Plohophorini Glyptodonts (Xenarthra, Cingulata) From the Late Neogene of Northwestern Argentina. Insight Into Their Diversity, Evolutionary History, and Paleobiogeography

Plohophorini Glyptodonts (Xenarthra, Cingulata) From the Late Neogene of Northwestern Argentina. 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Insight Into Their Diversity, Evolutionary History, and Paleobiogeography Alizia Núñez-Blasco, Alfredo E. Zurita, Ricardo Bonini, Angel R. Miño-Boilini, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3914918/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 17 Aug, 2024 Read the published version in Journal of Mammalian Evolution → Version 1 posted 8 You are reading this latest preprint version Abstract Northwestern Argentina (NWA) contains, together with the Pampean region (PR), one of the most complete late Neogene continental sequences, in which a great diversity of palaeofauna was recognized, among which glyptodonts stand out. Recent evidence suggests that the Late Miocene was a period of extra-Patagonian diversification in southern South America for glyptodonts, perhaps stimulated by the expansion of C4 grasses and open environments (known as “Edad de las Planicies Australes”). Here we focus on one of the most poorly known glyptodonts of NWA, the Plohophorini, from the Villavil-Quillay basin (Catamarca Province). Our results show that, like other clades (e.g., Doedicurini), a single species can be recognized, Stromaphorus ameghini (Ameghino, 1889 ; ex Moreno, 1882 ), whose stratigraphic record spans from the latest Miocene to the Pliocene (ca. 7.14–3.3 Ma; Messinian-Zanclean). Cladistic analysis confirms the status of natural group of the tribe Plohophorini within Hoplophorinae (“austral clade”), in which S. ameghini appears as the sister species of the Pampean species S. trouessarti (Moreno, 1888 ) nov. comb. The oldest precise records of S. ameghini (ca. 7.14 Ma) provide a minimum age for the Plohophorini lineage. The evidence suggests that the diversity of glyptodonts from the late Neogene of NWA is composed of endemic species, different from those of the PR, although both areas share the same genera, as observed in other mammalian clades such as Hegetotheriidae and Dasypodidae. Finally, the cladistic analysis reveals, in a broader context, that the spine-like structure observed in the caudal tube of some genera (ie, Nopachtus , Propanochthus , and Panochthus ) is a homologous structure rather than a convergence as usually interpreted. On the contrary, the similar appearance of the ornamentation pattern represented by the multiplication of peripheral figures in the carapaces of the genera Stromaphorus and Nopachtus is, in fact, a convergence. Glyptodontidae Stromaphorus Phlyctaenopyga Late Miocene-Pliocene diversity evolutionary history Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction Xenarthra (Pilosa and Cingulata) is traditionally regarded as one of the main clades within Placentalia (O’Leary et al. 2013); however, they can be considered today as a relictual clade, compared to the high diversity of the fossil record along Cenozoic (McKenna and Bell 1997 ; Vizcaíno and Bargo 2014 ). Within this remarkable diversity, one of the most enigmatic clades is represented by Glyptodontidae (Late Eocene-Late Pleistocene), composed of medium (ca. 60–260 kg) to giant (ca. 2300 kg) armored herbivores (Vizcaíno et al. 2011 ; Soibelzon et al. 2012 ; Quiñones et al. 2020 ; Zamorano et al. 2021 ) and phylogenetically linked to another fossil enigmatic clade, Pampatheriidae (see Gaudin and Wible 2006 ; Gaudin and Lyon 2017 ; Fernicola et al. 2018 ). More recently, another proposal based on molecular evidence suggests that glyptodonts are in fact a subfamily (Glyptodontinae) within the "armadillos" Chlamyphoridae (Delsuc et al. 2016 ; Mitchell et al. 2016 ). In this contribution we focus exclusively on morphological aspects, and therefore we use the traditional classification, considering Glyptodontidae Gray, 1869; Pampatheriidae Paula Couto, 1954 and Dasypodidae Gray, 1821 as families at the same taxonomic level. This is also consistent with most recent contributions (e.g., Núñez-Blasco et al. 2021, 2022; Quiñones et al. 2023 ; Cuadrelli et al. 2023a , b ; Christen et al. 2023 ) using Glyptodontidae at the family level. In this framework, the analysis of the evolutionary history of glyptodonts since the Early and Middle Miocene (a moment when materials are enough informative; see Gaudin and Croft 2015 ) suggests that two large clades with different characteristics can be recognized, one of northern origin called Glyptodontinae, and other of southern origin (“austral clade”) that we propose here to name Hoplophorinae Huxley, 1864 (see, among others, Cuadrelli et al. 2020 ; Quiñones et al. 2020 , 2023 ; Nuñez-Blasco et al. 2021a, 2022 ; Barasoain et al. 2022a ). While Glyptodontinae dispersed to Central and North America during the Great American Biotic Interchange (GABI) (Carlini and Zurita 2010 ; Zurita et al. 2011 ; Gillette et al. 2016 ), Hoplophorinae, despite being more diverse, had a more restricted geographical distribution, limited mainly to high and middle latitudes of South America (Núñez-Blasco et al. 2020 ; Barasoain et al. 2022a ). Within Hoplophorinae, it is essential to analyze the evolution of glyptodonts in southern South America since the Late Miocene (ca. 9 Ma; see Barasoain et al. 2022a ) to understand their diversity and evolutionary history in the latest Miocene and Plio-Pleistocene, when terminal species became extinct in the latest Pleistocene together with the remaining megafauna (Prates and Pérez 2021 ; Prates et al. 2022 ; Carlini et al. 2022 ). The Northwestern region of Argentina (NWA) is an area with a remarkable diversity of glyptodonts (see Cabrera 1944 ; Zurita 2007a , b ; Nuñez Blasco et al. 2021a, 2022). It contains, together with the Pampean region (PR) of Argentina, one of the most complete Late Miocene-Pleistocene continental sequences of South America (Esteban et al. 2014 ; Bonini et al. 2016 , 2017 , 2021 ). In this context, a comprehensive revision of the diversity of glyptodonts from NWA (ie, taxonomic, phylogenetic, and chronostratigraphic) is currently carrying out (see Núñez-Blasco et al. 2021b , c ). These studies, including a detailed comparison with those taxa from the PR (Núñez-Blasco et al. 2022), have provided new information highlighting that, as observed in the late Neogene of the PR (see Zurita et al. 2016a , b , 2017 ), the diversity of glyptodonts seems to be more restricted than previously known. Plohophorini Castellanos, 1932 , is currently one of the most problematic traditional glyptodont tribes from a taxonomic, systematic, and phylogenetic standpoint. Within this tribe, the genus Phlyctaenopyga contains a species endemic to the NOA, Ph. ameghini (Ameghino, 1889 ), in addition to a late Neogene species in the RP, Ph. trouessarti (Moreno, 1888 ). The tribe Plohophorini also encompasses Stromaphorus compressidens (Moreno and Mercerat, 1891 ), another species interpreted as endemic to the NWA. Both taxa have been extensively investigated and described by numerous authors since the 19th century (e.g. Moreno 1882 , 1888 ; Moreno and Mercerat, 1891 ; Ameghino 1888a , b , 1889 , 1891a , b , 1902 , 1904 ; Lydekker 1894 ; Rovereto 1914 ; Cabrera 1939 , 1944 ; Castellanos 1925 , 1940 ; Zamorano et al. 2011 ; Zamorano 2012 ; Toriño and Perea 2018 ). In this scenario, the last revision of the glyptodont diversity from NWA was carried out by Cabrera ( 1944 ), more than seventy years ago, without phylogenetic and stratigraphic context and without taking into account taphonomic aspects. New field works carried out during the last years at NWA plus the revision of materials from different institutions (see Materials and Methods) yielded new and more complete specimens that allow us to perform, for the first time, a comprehensive revision of this particular assemblage of glyptodonts. In addition, chrono-stratigraphic and geological framework of these fossiliferous sequences from NWA were greatly improved in the last decades (see, among others see among others Bonini 2014 ; Bonini et al. 2016 , 2017 , 2021 ; Bossi y Muruaga 2009; Bossi et al. 1987 , 2001 ; Butler et al. 1984 ; Esteban et al. 2014 ; Georgieff et al. 2017 ; Latorre et al. 1997 ; Marshall y Patterson 1981; Marshall et al. 1979 ; Muruaga 2001a , b ;ñez-Blasco et al. 2020 ; Riggs y Patterson 1939; Sasso 1997 ; Hynek et al. 2012 ), a situation that allowed a detailed chronostratigraphic determination of the levels that contain these faunas. The present paper is the continuation of an exhaustive review of the diversity of late Neogene glyptodonts from NWA (see Núñez-Blasco et al. 2020 , 2021a , b , c ), based on the pioneering work of Cabrera ( 1944 ). We also perform the most comprehensive phylogenetic analysis of glyptodonts to date to evaluate the position of the accepted valid species of Plohophorini from NWA, highlighting the fact that this tribe (including species from PR and NWA of Argentina, and Uruguay) appears as a natural group in our analysis. Finally, this work complements those carried out by our team on the late Neogene glyptodont associations of the PR of Argentina (see Quiñones et al. 2023 ) allowing a more certain evolutionary scenario for these elusive and enigmatic mammals. Materials and methods Institutional abbreviations . AMNH , American Museum of Natural History, New York, USA; CC-MUFCA , Colección “Dr. Alfredo Castellanos”, Museo Universitario “Florentino y Carlos Ameghino”. Facultad de Ciencias Exactas, Ingeniería y Agrimensura - Universidad Nacional de Rosario. Rosario. Santa Fe, Argentina; FACENA , Facultad de Ciencias Exactas y Naturales y de Agrimensura, Corrientes Capital, Argentina; FC-CVF , Colección de vertebrados fósiles, Facultad de Ciencias, Universidad de la República, Montevideo, Uruguay; FMNH-P , Field Museum of Natural History, Paleontological collection, Chicago, Illinois, USA; GCF , Grupo Conservacionista de Fósiles, Museo Paleontologico ‘Fray Manuel de Torres,’ San Pedro, Buenos Aires, Argentina; IGM , Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, México; MACN , Sección Paleontología Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina; MCA , Museo Municipal de Ciencias Naturales ‘Carlos Ameghino,’ Mercedes, Buenos Aires, Argentina; MCH P , Sección Paleontología, Museo Arqueológico Condor Huasi, Belén, Catamarca, Argentina; MHNC , Museo de Historia Natural de Cochabamba ‘Alcide d’Orbigny’, Cochabamba, Bolivia; MLP-PV , División Paleontología Vertebrados, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Buenos Aires, Argentina; MMC , Museo Municipal de Colonia ‘Dr. Bautista Rebuffo’, Colonia de Sacramento, Uruguay; MMP , Museo Municipal de Ciencias Naturales ‘Lorenzo Scaglia’ Mar del Plata, Argentina; MSM , Arizona Museum of Natural History (formerly Mesa Southwest Museum), Arizona, USA; PVL , Colección de Paleozoología de la Facultad de Ciencias Naturales e Instituto ‘Miguel Lillo’, San Miguel de Tucumán, Argentina; PVSJ , Instituto y Museo de Ciencias Naturales, Universidad Nacional de San Juan, San Juan (San Juan Province, Argentina); Pz-Ctes , Colecciones Paleontológicas de la Universidad Nacional del Nordeste “Rafael Herbst”, Corrientes, Argentina; SGO PV , vertebrate paleontology collections, Museo Nacional de Historia Natural, Santiago, Chile; UCMP , University of California, Museum of Paleontology, Berkeley, California, USA; UFMG , Museo de la Universidade Federal de Minas Gerais, Belo Horizonte, Brazil. Anatomical abbreviations . bc , bony crest; dpm , descending process of the maxillae; fm , foramen magnum; if , infraorbital foramina; Mf , mf , upper and lower molariforms; no , nasal opening; oc , occipital condyle; on , orbital notch; sc , sagittal crest; za , zygomatic arch. Other abbreviations . FAD , First Appearance Datum; FOCP , Faldeo Occidental Cerro Pampa; LAD , Last Appearance Datum; Ma , Megaannun, (millon years ago); PCQ , Puerta de Corral Quemado; SFN , San Fernando Norte; SMV , Santa María Valley; VQB , Villavil-Quillay Basin. The most relevant materials for this study are housed in the paleontological collection of vertebrates from the Museo de La Plata (“Cabrera collection”) and the Paleontological collection from the Field Museum of Natural History (Riggs collection), under the following acronyms: skulls, MLP-PV 29-X-8-1, MLP-PV 29-X-10-1, FMNH-P14396; mandibles, MLP-PV 16–138, MLP-PV 29-X-8-1; carapaces MLP-PV 16–101, MLP-PV 29-X-10-2, FMNH-P14439; caudal tubes, MLP-PV 29-X-8-9, MLP-PV 29-X-10-1, FMNH-P14414, FMNH-P14532; (see Online Resource 1 for the nomenclatural history of Phlyctaenopyga ameghini (Ameghino, 1889 ) and Stromaphorus compressidens (Moreno y Mercerat, 1891); and Online Resource 2 for a list of all materials assigned to the species Stromaphorus ameghini (Ameghino, 1889 ; ex Moreno, 1882 ). A comparative study was carried out to obtain a morphological characterization of the specimens, including cranial and postcranial elements. The results are shown in the anatomical descriptions. For direct comparison, specimens referred to the following genera were used: Glyptodon Owen, 1839; Kelenkura Barasoain, Zurita, Croft, Montalvo, Contreras, Miño-Boilini and Tomassini, 2022; Plohophorus Ameghino, 1887; Pseudoplohophorus Castellanos, 1926 ; Cranithlastus Arias, Alonso and Malanca, 1978; Stromaphoropsis Kraglievich, 1932; Eleutherocercus Koken, 1888; Doedicurus Burmeister, 1874; Neosclerocalyptus Paula Couto, 1957; Eosclerocalyptus Ameghino, 1919; Hoplophorus Lund, 1839; Nopachtus Ameghino, 1888; Propanochthus Castellanos, 1925 ; Panochthus Burmeister, 1866. Additionally, specimens from the literature were included in the analysis. All measurements referred to in the text are listed in Online Resource 3 (table 1: skull; table 2: Holotype MLP-PV 16–101; table 3: carapace, and table 4: caudal tube); they were taken through a manual "Vernier" caliper, with an error range of 0.5mm and in some cases using ImageJ (1.53e) software. The linear dimensions are expressed in millimeters (mm). For descriptions and comparisons of the dorsal carapace, osteoderms and caudal tube, we follow the proposal of Zamorano et al. ( 2011 ), Porpino et al. ( 2014 ) and Toriño ( 2015 ). The numbering assigned to the figures of the ornamentation of caudal tubes (see Fig. 7 ) follows the proposal of Toriño ( 2015 ), and Toriño and Perea ( 2018 ). For chronological purposes, we follow the International Chronostratigraphic Chart v2022/02 (Cohen et al. 2020 ). Cladistic analysis . The phylogenetic study is based on a mixed matrix (see Online Resource 4 and Online Resource 5), consisting of a total of 96 characters, of which 95 are conventional (descriptive morphological characters) and one of geometric morphometry, to analyze a total of 35 taxa. The descriptive morphological characters are 68 binary and 27 multistate. This includes 40 skull characters, 4 autopodium characters, 26 carapace characters and 25 caudal armor characters. The information considered in this analysis, used both for the codification of states and landmark placement for geometric morphometry, was recorded via direct observation of the specimens and from photographs taken by the authors. The taxon-character matrix was constructed with Mesquite version 3.4 (Maddison and Maddison 2018 ) and to build the geometric morphometric character, photographs were compiled into .tps files using tpsUtil software, version 1.70 (Rohlf 2016 ), and landmarks were digitalized with tpsDig2 version 2.16 (Rohlf 2010 ); this matrix was analyzed via “Traditional search“ under the criterion of maximum parsimony, using TNT version 1.5 (Goloboff and Catalano, 2016 ), the algorithm used was TBR (10 trees to save per replication). It is worth noting that all characters were unordered and were weighted equally (1.0). Clade support was assessed via Absolute and Relative Bremer support, retaining suboptimal trees by 3 steps; see Bremer ( 1994 ); Goloboff and Farris ( 2001 ). In addition, a Jackknife analysis was carried out via “Traditional search” (TBR algorithm) with 100 replicates and 36 removal probability. Finally, the consensus tree obtained from the two most parsimonious trees (MPTs) was calibrated to obtain estimated divergence times and potential ghost ranges. The calibration followed the ‘equal’ method (with a root of 1 million-years) implemented in the strap package (see Bell and Lloyd, 2014 ) for R software version 4.3.0 (R Core team 2023 ). The resulting time-scaled tree was plotted against the International Chronostratigraphic Chart (Cohen et al. 2020 ) for visualization purposes. The R commands included in the appendices of Bell and Lloyd ( 2014 ) and Toriño et al. ( 2021 ) were adapted for these procedures. The input data for the analysis with R were the strict consensus tree exported in parenthetical notation from TNT as a .tre file, and a txt file including a list of taxa with their first and last record (see supplementary files). The list of biochrons for each taxon as well as the bibliographical references from which this information was taken can be found in the Online Resource 6 of the present work. The ingroup includes the following taxa: Boreostemma venezolensis Simpson, 1947; B. acostae (Villarroel 1983); Glyptodon jatunkhirkhi Cuadrelli, Zurita, Toriño, Miño-Boilini, Perea, Luna, Gillette and Medina, 2020; G. munizi Ameghino, 1881; G. reticulatus Owen, 1845; Glyptotherium texanum Osborn, 1903; Gl . cylindricum (Brown, 1912); Propalaehoplophorus australis Ameghino, 1887; Eucinepeltus petesatus Ameghino,1891; Cochlops muricatus Ameghino, 1889 ; Palaehoplophoroides rothi Scillato-Yané and Carlini, 1998; Palaehoplophorus meridionalis Ameghino, 1904 ; Kelenkura castroi Barasoain, Zurita, Croft, Montalvo, Contreras, Miño-Boilini and Tomassini, 2022; Plohophorus figuratus Ameghino, 1887; P. avellaneda Quiñones, Cuadrelli, De los Reyes, Luna, Poiré and Zurita, 2023; Pseudoplohophorus absolutus Perea, 2005; Ps. benvenutii (Castellanos, 1954); Stromaphorus ameghini (Ameghino, 1889 ); S. trouessarti (Moreno, 1888 ); Eleutherocercus solidus (Rovereto, 1914 ); E. antiquus (Ameghino, 1887); Doedicurus clavicaudatus (Owen, 1847); Neosclerocalyptus gouldi Zurita, Carlini and Scillato-Yané, 2008; N. ornatus (Owen, 1845); N. paseudornatus (Ameghino, 1889 ); N. paskoensis (Zurita, 2002); Nopachtus coagmentatus Ameghino, 1888; Hoplophorus euphractus Lund, 1839; Propanochthus bullifer (Burmeister, 1874); Panochthus intermedius Lydekker, 1895; P. tuberculatus (Owen, 1845) and Parapropalaehoplophorus septentrionalis Croft, Flynn and Wyss, 2007 . The extant dasypodid Euphractus sexcintus Linnaeus, 1758, and the pampathere Pampatherium humdboltii (Lund, 1839) were used as outgroups to root the tree. Geometric morphometrics . For the geometric morphometric character (Fig. 2 ) two-dimensional coordinates of 29 landmarks were digitalized from photographs of mandibles in lateral view of sixteen glyptodonts ( Eucinepeltus petesatus MACN 4760; Propalaehoplophorus australis MLP-PV 16 − 15; Pseudoplohophorus absolutus FC-CVF475-595; Glyptotherium texanum MSM 4818; Gl. cylindricum IGM 9563; Boreostemma acostae UCMP 38039; Glyptodon munizi GCF 10; G. reticulatus MCA 2015; Doedicurus clavicaudatus MACN 2762; Eleutherocercus solidus FMNH-P14437; Panochthus intermedius MHNC 13491; P. tuberculatus MLP-PV 16–29; Neosclerocalyptus ornatus MLP-PV 16–28; N. paskoensis MACN 18107; N. gouldi MCA 2010 and Parapropalaehoplophorus septentrionalis SGO PV 4165), one dasypodid ( Euphractus sexcintus FACENA 183) and one pampatheriid ( Pampatherium humboldtii MLP-PV 81-X-30-1); This method had previously been tested on Cingulate mandibles, with promising results in phylogenetics (see Nuñez-Blasco et al. 2021b). In this context, of the 29 landmarks, 5 are type I (Ladmarks 1, 10, 17, 18 and 19); 2 are type II (Ladmarks 2 and 3). Besides, there are 3 sets of semi-landmarks used to define the rest of the mandible morphology (green set, blue set and orange set). The green and blue sets (Fig. 2 : a, c) were defined based on the 65º angle formed between landmarks 3, 19 and 10, subdividing it into 7 portions. The orange set (Fig. 2 : b) was defined based on the 110º angle formed by landmarks 1, 3 and 19, subdividing it into 10 portions. Geographic and stratigraphic context The study area is in the Catamarca province, Northwestern Argentina, within the Northwestern Pampean Ranges geological province (Caminos, 1979 ). The Holotype materials of “ Phlyctaenopyga ” ameghini and “ Stromaphorus compressidens ” come from the Santa María Valley (SMV), while most of the fossils assigned to these two species come from the Villavil-Quillay basin (VQB). The SMV (26°53’S, 66°05’W) is a tectonic depression of more than 100 km long by 20 to 30 km wide, flanked to the east by the Cumbres Calchaquíes and Aconquija Ranges, and to the west by the Quilmes Ranges (Bossi et al. 2001 ). A great part of the paleontological materials from SMV was exhumed from the Late Miocene/Pliocene levels cropping out at Andalhuala de Arriba and Tiopunco localities (Catamarca and Tucumán provinces, respectively). In turn, the VQB (27°15’ S/66°54’ W) is a tecto-sedimentary basin included in the Hualfin valley, limited to the northwest by the Sierra de Altohuasi, to the southwest by the Cerro Durazno, to the northeast by the Sierra de Hualfín, to the east by the Complejo Volcánico Farallón Negro, and the southeast by the Cerro Pampa-Belén Range (Fig. 1 ). The materials found in the VQB come from the Andalhuala, and Chiquimil formations cropping out at Puerta de Corral Quemado (PCQ), San Fernando Norte (SFN), and Faldeo Occidental del Cerro Pampa (FOCP). The Neogene lithostratigraphic units recognized in these areas have been included in the Santa María Group, which is composed (from base to top) of Las Arcas, Chiquimil, Andalhuala, and Corral Quemado formations, ranging from ca. 10 to 3 Ma, being the Andalhuala Formation the most extended in both areas (Bossi et al. 1987 ; Muruaga 2001a , b ; Bossi and Muruaga 2009 ; Bonini et al. 2017 ). The Andalhuala Formation in the VQB was deposited between ca. 7.14 to 3.66 Ma (Latorre et al. 1997 ), corresponding to the fossiliferous beds that yielded the studied materials (Riggs and Patterson 1939 ; Marshall and Patterson 1981 ). The historical specimens collected in the SMV lack provenance data, except for the reference of “ Estratos Araucanos ” ( sensu Rovereto 1914 ) and “ Araucanense medio ” ( sensu Castellanos 1948a , b , 1969 ), mainly assigned to the upper section of Chiquimil and Andalhuala formations in the modern lithostratigraphic framework. Lithologically, the Andalhuala Formation in SMV is broadly characterized by sandstones and conglomerates poorly sorted, polymict and matrix-supported clasts, with scarce participation of pelites (Georgieff 1998 ). In both areas, SMV and VQB, this unit is composed of facies of brownish, reddish, and greyish sandstones with tabular, lenticular, trough cross-stratification, ripple marks, and cuneiform cross-stratification, often interbedded with tabular massive siltstones and several tuff beds. Moreover, the Andalhuala Formation shows several secondary features that evidence the subaerial exposure and water table fluctuations, which agree with the inferred permanent fluvial subenvironment passing to an eolian system with dunes (Bonini et al. 2017 , 2021 ). Although the literature mentions the same stratigraphic formations in VQB and SMV, it should be noted that the units that form the Santa María Group sometimes show important diachronies between those cropping out in both basins (see Spagnuolo et al. 2010 , 2013 , 2015 ; Georgieff et al. 2012a , b , 2017 ; Georgieff and Díaz 2014 ; Bonini 2014 ; Bonini et al. 2017 , 2021 ). In general, the inferred palaeoenvironments are shallow lacustrine (but also continental sabkha in SMV; Ibañez, 2001; Esteban et al. 2019 ) and sandy-gravel braided fluvial for the Chiquimil and Andalhuala formations at both sites, but the dating of the tuffaceous strata reveals that their deposition were older in VQB. A clear example is the Chiquimil Formation (see Fig. 1 ), which presents an age of 7.14 Ma for its top at East of PCQ, while in the SMV an age of ca. 6.88 was obtained from the top of the underlying unit, Las Arcas Formation (Georgieff et al. 2014; 2017 ); this shows that the deposition of the Chiquimil Formation at SMV was younger, and therefore the lithostratigraphic correlation between both areas cannot be established by that criteria. Likewise, the determination of the ages of the specimens analyzed in the present work has been carried out taking into account the absolute ages (geochronology) and not the formation (lithostratigraphy) from which they originate. Systematic Paleontology Superorder Xenarthra Cope, 1889 Order Cingulata Illiger, 1811 Family Glyptodontidae Gray, 1869 Tribe Plohophorini Castellanos, 1932 (nom. transl. Hoffstetter, 1958 ) Genus Stromaphorus Castellanos, 1925 (= Phlyctaenopyg a Cabrera, 1944 new synonymy) Species : Stromaphorus ameghini ( Ameghino, 1889 ; ex Moreno, 1882 ) Other species Stromaphorus trouessarti (Moreno, 1888 ) nov. comb. Stromaphorus ameghini ( Ameghino, 1889 ; ex Moreno, 1882 ) = Hoplophorus ameghinii Moreno, 1882 , p. 120 [nom. nud.]; Plohophorus ameghini Ameghino, 1889 , p. 825, plates LXIX, Figs. 19 and 20 and LXXXII, Figs. 5 and 6 ; Neuryurus compressidens Moreno and Mercerat, 1891 , p. 224; Plohophorus philippii Moreno and Mercerat, 1891 , p. 225; Plohophorus ameghinii Ameghino, 1902 , p. 2; Plohophorus ameghinoi Ameghino, 1904 , p. 288; Stromaphorus ameghinoi (Ameghino, 1904 ) Castellanos, 1925 , p. 96 (not 1940, p. 27) [n. comb.]; Stromaphorus philippii (Moreno and Mercerat, 1891 ), Cabrera 1939 [n. comb.]; Stromaphorus ameghinoi (Ameghino, 1904 ) Castellanos, 1940 , p. 27 (not 1925, p. 96); Urotherium compressidens Castellanos 1940 , p. 273; Stromaphorus compressidens (Moreno and Mercerat, 1891 ), Cabrera 1944 , p. 30 [n. comb.]. Phlyctaenopyga ameghini (Ameghino, 1889 ), Cabrera 1944 , p. 42 [n. gen, n. comb.]. Holotype MLP-PV 16–101, small fragment of carapace and some isolated osteoderms. Santa María Department, Catamarca Province, Argentina. Geographic and Stratigraphic Occurrence : Catamarca Province: Santa María Valley, Puerta de Corral Quemado, Corral Quemado and San Fernando; Chiquimil Formation, Andalhuala Formation and Corral Quemado Formation, Late Miocene-Pliocene (Cabrera 1944 ; Marshall and Patterson 1981 ; Bonini 2014 ; Zamorano et al. 2011 ). Tucumán Province: Tiopunco (Cabrera 1944 ; Zamorano et al. 2011 ). La Rioja Province: El Degolladito; Salicas Formation, Late Miocene (Brandoni and González-Ruiz 2020). Córdoba Province: Arroyo Los Chiflones, Villa Cura Brochero; Brochero Formation, Miocene-Pliocene (Cruz 2011 , 2013 ). Materials analysed in this work, geographic and stratigraphic provenance and age: FMNH-P14396 , deformed skull, Ampajango, Catamarca Province, Argentina; Chiquimil Formation, Stratigraphic level XI from Stahlecker in Marshall and Patterson ( 1981 , Appendix II) between 6.88 to 6.02 Ma, Messinian (Late Miocene); FMNH-P14414 , incomplete and deformed skull, dorsal carapace fragment, femur, two caudal rings, distal portion of caudal tube, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, Stratigraphic level 17 from Marshall and Patterson ( 1981 ), age ca. 6.9 to 6.3Ma, Messinian (Late Miocene); FMNH-P14439 , complete carapace, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, Stratigraphic level 26 from Marshall y Patterson (1981); age ca. 4.9 to 4.4 Ma, Zanclean (Early Pliocene); FMNH-P14532 , proximal portion of caudal tube, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; Andalhuala Formation/Corral Quemado Formation, Stratigraphic levels 15–32 from Marshall and Patterson 1981 ; age ca. 7.14 to 3.66 Ma, Miocene-Pliocene; FMNH-P15771 , small fragments of carapace, some from the anterior region and some from the posterior region, Tio Punco, Tafi, Tucumán Province, Argentina; unknown stratigraphic level, Pliocene, [originally classified as Glyptodontidae indet. in Marshall and Patterson 1981 ]; MCH-P39 , small fragment of carapace, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma, Zanclean (Early Pliocene); MCH-P174 , isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma, Zanclean (Early Pliocene). New studied specimen; MCH-P176 , isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma, Zanclean stage, Pliocene. New studied specimen; MCH-P177 , isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma., Zanclean (Early Pliocene). New studied specimen; MCH-P247 , isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, age ca. 4.72 to 4.79 Ma, Zanclean (Early Pliocene). New studied specimen; MCH-P323 , isolated osteoderm, Faldeo Occidental Cerro Pampa (San Fernando Sur), Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately below the tuff dated in 5.59 Ma, Messinian (Late Miocene). New studied specimen; MCH-P326 , isolated osteoderms, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, age ca. 3.6 to 4.72 Ma, Zanclean (Early Pliocene). New studied specimen; MCH-P328 , isolated osteoderms, fragments of carapace, fragments of caudal rings and fragments of caudal tube, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, age ca. 3.6 to 4.72 Ma, Zanclean (Early Pliocene). New studied specimen; MCH-P364 , isolated osteoderm, Western Slopes of Cerro Pampa. Chiquimil Formation, age more than 7.14 Ma; Messinian-Tortonian (Late Miocene). New studied specimen; MCH-P365 , 21 isolated osteoderms, Western Slopes of Cerro Pampa, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, above tuff dated in 5.59 Ma, Miocene-Pliocene. New studied specimen; MLP-PV 16–138 , incomplete and deformed left horizontal ramus, holotype of “ Stromaphorus compressidens ” ( Neuryurus compressidens Moreno and Mercerat, 1891 ), Bajo de Andalhualá, Santa María Department, Catamarca Province, Argentina; Corral Quemado Formation; MLP-PV 29-X-8-1 , laterally deformed skull, mandible and carapace fragments; referred to “Stromaphorus compressidens” by Cabrera ( 1944 ). Barranca del Palito Parado, campo del Jarillar, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; “Araucaniano”; MLP-PV 29-X-8-9 , caudal tube; referred to “Stromaphorus compressidens” by Cabrera ( 1944 ). Campo de los Cálibas, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; “Araucaniano”; MLP-PV 29-X-10-1 , incomplete skull, cephalic shield, carapace fragments, parts of the sacral and first caudal vertebrae, caudal tube and parts of the caudal rings; referred to “ Phlyctaenopyga ” ameghini by Cabrera ( 1944 ). Loma de la Greda, San Fernando, Belén Department, Catamarca Province, Argentina; “Araucaniano”; MLP-PV 29-X-10-2 , carapace deformed by pressure, parts of the first caudal ring, tube and pelvis; referred to “ Phlyctaenopyga ” ameghini by Cabrera ( 1944 ). Loma de la Greda, San Fernando, Belén Department, Catamarca Province, Argentina; “Araucaniano”. See Online Resource 2 for a complete list of materials historically referring to “ Phlyctaenopyga ” ameghini (Ameghino, 1889 ) and “ Stromaphorus compressidens ” (Moreno and Mercerat, 1891 ), and it is reassignment to Stromaphorus ameghini (Ameghino, 1889 ; ex Moreno, 1882 ). Emended diagnosis : Medium sized glyptodont (see Online Resource 3), larger than Pseudoplohophoru s but smaller than the Pleistocene genera Panochthus, Glyptodon , and Doedicurus . Lateral skull profile similar to Plohophorus and Pseudoplohophorus . Nasal region dorsoventrally reduced; nasofrontal area inclined 152º anteroventrally in front of the orbital notches; subelliptical orbital notch, with the main axis inclined 110° counterclockwise to the dorsoventral axis; inverted subtrapezoidal nasal aperture with both lateral margins convex; subtriangular rostral area in front of the orbital notches; without postorbital bar; zygomatic arches more elongated anteroposteriorly than those of other Plohophorini (eg., Plohophorus figuratus ); palate general morphology straight, as in Plohophorus and different from Pseudoplohophorus ; Mf1 and Mf2 with triangular morphology, Mf4-8 with very marked trilobulation, Mf4 coincides with the infraorbital foramen. The dorsal carapace is sub-rectangular with variation in the ornamentation depending on the sector. Anterior portion: rounded and flat central figure, surrounded by a single row of 10–12 small rounded peripheral figures. Middle portion: subcircular and slightly convex central figure, surrounded by a first row of 11 to 15 peripheral figures and a second row which is always incomplete; it differs clearly from Nopachtus , Panochthus and Propanochthus in which the second row is always complete. Postero-dorsal region (pelvic region): round and very convex central figure, surrounded by a first row of 12–21 peripheral figures and a second one, complete, between 21–27; this is the only area of the carapace in which the second row of peripheral figures is complete. Caudal margin with large convex central figures preceded anteriorly by 3–5 rows of peripheral figures, one row on the sides of the central figure and a free border on the posterior margin. In dorsal and ventral view, the caudal tube has parallel lateral borders along its entire length instead of tapering distally, as the caudal tubes of some genera (e.g. Pseudoplohophorus and Plohophorus ) and almost circular in transverse outline. Simple rosette pattern, with a smooth-surfaced central figure surrounded by a single complete row of shared peripheral figures. At the apex there are two large smooth terminal figures (T) which, in the more distal portion, are in contact. It has four lateral figures (numbered I, II, III and IV, see Fig. 7 ) that reduce in size towards the proximal portion. Figure I is characteristically convex, unlike figures II, III and IV, which are flatter. Between the lateral and marginal figures there is a row of small peripheral figures. Description and comparisons Cranium In lateral view (Fig. 3 : a, MLP-PV 29-X-8-1; e, FMNH-P14396 and i, MLP-PV 29-X-10-1) the dorsal profile of the skull is similar to Plohophorus figuratus (MLP-PV 16–153), Pseudoplohophorus absolutus (FC-CVF 475 / 595) and Ps. benvenutii (CC-MUFCA 1388). The naso-frontal area has an anteroventral tilt relative to the parieto-occipital area, forming an angle of 152º situated in front of the orbital notches, in contrast to Doedicurus , Eleutherocercus and Panochthus , in which both areas form a slightly tighter angle of ca. 140º, located posterior to or at the level of the orbital notch. The nasal region is dorsoventrally reduced (for measurements see Online Resource 3, table 1), a common feature among the Plohophorini, and different from the rest of the Glyptodontidae. The orbital notch is subelliptic, with the main axis inclined 110º in an anti-clockwise direction relative to the dorsoventral axis. In the ventral margin of the orbital notch a subtle bony crest is observed, similar to P. figuratus, Ps. absolutus , Ps. benvenutii and Eosclerocalyptus proximus (CC-MUFCA 703), but less pronounced than in Eleutherocercus and Neosclerocalyptu s. Like most glyptodonts, Stromaphorus ameghini lacks a post-orbital bar, unlike Eleutherocercus solidus (FMNH-P14437), E. antiquus (MACN 2894) and Panochthus tuberculatus (MLP-PV 16–29), in which it is present and well developed. On the other hand, the dorsal margin of the zygomatic arch is slightly concave, unlike P. figuratus , in which this concavity is greater and Ps. absolutus and Ps. benvenutii , which has a rather straight margin. In turn, the ventral margin of the zygomatic arch is straight and descends directly to join the descending process. In general terms, this whole structure is rather robust. In frontal view, (Fig. 3 : b, MLP-PV 29-X-8-1; f, FMNH-P14396 and j, MLP-PV 29-X-10-1) the nasal opening is sub-trapezoidal inverted, with its dorsal margin wider than the palate, and with both lateral margins convex, as in P. figuratus , Ps. absolutus , Ps. benvenutii , Kelenkura castroi (PVSJ-366), and Eo. proximus and bears a certain resemblance to E. antiquus . The descending processes are broken in all specimens. However, the infraorbital foramina are preserved; they are subcircular and small, different from Glyptodon munizi (GCF 10) and G. reticulatus (MCA 2015), in which they are circular and larger (see Zurita et al. 2013 ; Cuadrelli et al. 2019 ) (for measurements see Online Resource 3, table 1). In dorsal view, (Fig. 3 : c, MLP-PV 29-X-8-1, g, FMNH-P14396 and k, MLP-PV 29-X-10-1), the general morphology of the skull is very similar to P. figuratus and to a lesser degree to Ps. absolutus and Ps. benvenutii . The specimen MLP-PV 29-X-10-1 is largely reconstructed, especially in the whole area of the cranial vault; therefore, it is not reliable to describe the morphology of this region in this fossil. The rostral area in front of the orbital notches is subtriangular in outline, as is that of P. figuratus, Ps. absolutus , and Ps. benvenutii , but different from E. antiquus, E. solidus, Neosclerocalyptus paskoensis (MACN-Pv 18107), N. ornatus (MLP-PV 16–18), Glyptodon munizi and G. reticulatus in which the morphology is sub-rectangular. The orbital notches are open posteriorly due to the lack of post-orbital bar, while the maximum diameter of the fronto-parietal region coincides with this region; further back, there is a conspicuous post-orbital narrowing, also found in other species, such as P. figuratus , Eo. proximus , D. clavicaudatus or P. tuberculatus . The zygomatic arches are more elongated antero-posteriorly than in the other Plohophorini but not as much as K. castroi ; in turn, these more rounded forms differ from the quadrangular forms of E. solidus and E. antiquus . In all three specimens it can be observed a short and poorly developed sagittal crest, that starts at the nuchal crests and extends up to the contact with the fronto-parietal suture (for measurements see Online Resource 3, table 1). In occlusal view, (Fig. 3 : d, MLP-PV 29-X-8-1, h, FMNH-P14396 and l, MLP-PV 29-X-10-1), the general morphology of the palate is quite straight, like P. figuratus , and different from Ps. benvenutii and Ps. absolutus , which shows a relatively marked expansion in the posterior region of the palate (for measurements see Online Resource 3, table 1). One of the most remarkable features of this view is the morphology of the first molariforms. FMNH-P14396 is the only specimen in which the molariforms Mf1 and Mf2 are clearly visible, and they are triangular; in P. figuratus , instead, they are markedly circular. The Mf3 is not preserved. The Mf4 has a markedly trilobed morphology. From Mf4 onwards all molariforms (Mf4-8) have very pronounced trilobulation. It should be noted that Mf8 is small compared to the preceding molariforms. The main axes of Mf4-Mf8 are parallel to the longitudinal axis of the dental series, a characteristic shared with P. figuratus . In this view the Mf4 coincides with the infraorbital foramen; this situation differs from P. figuratus , Ps. absolutus and Ps. benvenutii in which the infraorbital foramen coincides with the Mf3. Mandible Descriptions are mainly based on MLP-PV 16–138 (holotype of Neuryurus compressidens= “ Stromaphorus compressidens ”) which includes incomplete left horizontal ramus, and MLP-PV 29-X-8-1, represented by two hemi-mandibles (left and right), Fig. 4 . In lateral view, the mandible looks quite gracile compared to those of other groups such as Doedicurus , Eleutherocercus, Neosclerocalyptu s and Panochthus . The antero-posterior diameter of the ascending ramus is shorter than the total length of the tooth series from mf1 to mf6. Like in Ps. absolutus , the ascending ramus is inclined forwards, drawing an angle of 70° between its anterior margin and the alveolar margin; this situation differs from the opening observed in Neosclerocalyptus , Doedicurus , Eleutherocercus , and Glyptodon , in which the angle is narrower of 65º. At the posterior border of the ascending ramus, the posterior-lower margin shows a rounded angular process, somewhat less accentuated than in Ps. absolutus. The horizontal ramus of the mandibular body reaches the maximum height at the level of the mf5 and its ventral margin is slightly convex, as in Ps. absolutus and Eo. proximus . Molariforms. The mf1 is absent in all specimens, only molariforms mf2-mf8 are preserved. The mf2 is broken at the lingual posterior margin, but its morphology is triangular or kidney-shaped with the posterior portion narrowed laterally, this morphology is very different from the rest of the Glyptodontidae, and is repeated in the first molariforms of the upper series of the skulls MLP-PV 29-X-8-1 and FMNH-P14396 (See Fig. 3 : d, h). The pattern of mf3 tends to trilobation, with a greater development of the lobules on the labial margin; while mf4-mf8 are clearly trilobated, very similar to Ps. absolutus and Eo. proximus , especially the anterior lobe of mf6-mf8, which is square with a small, curved incision on the labial margin of mf8. Dorsal carapace MLP-PV 29-X-10-2 and FMNH-P14439 are complete dorsal carapaces (Fig. 5 ), allowing the full description of the carapace morphology of Stromaphorus , including its ornamentation pattern variation. The carapace is composed of approximately 35–43 transverse bands of osteoderms, which are not always complete, especially in the anterior region. In lateral view the dorsal carapace is sub-rectangular in outline, short and globular, unlike Neosclerocalyptus , which shows an almost completely straight dorsal profile. In both specimens (MLP-PV 29-X-10-2 and FMNH-P14439) the most convex region is at the middle, showing some similarity with the genera Plohophorus , Pseudoplohophorus , Eosclerocalyptus , and Nopachtus ; it contrasts with Panochthus and Doedicurus in which the convexity is at the anterior region; and Glyptodon , with its greatest convexity in the posterior region; and Eleutherocercus , which is dome-shaped at the dorsal-pelvic region. The exposed surface of the osteoderms that forms the carapace shows a "rosette" pattern, although its morphology varies in different regions of the carapace. In the anterior region (lateral and central) the osteoderms have on their exposed surface a central figure surrounded by a single row of peripheral figures (simple rosette pattern). In the middle and posterior regions this pattern becomes more complex by the addition of incomplete rows of peripheral figures and accessory figures, all of them small and rounded, similar to the pattern observed in the genera Plohophorus , Pseudoplohophorus , Stromaphoropsis and Cranithlastus . Although this pattern is very similar to that of the genera Nopachtus , Panochthus and Propanochthus , it differs because in these latter taxa the rows of peripheral figurines are always complete and with a polygonal morphology. Thus, Nopachtus has up to 2 complete rows, while Panochthus and Propanochthus have between 3 to 7 complete rows per osteoderm, showing in some species a pattern of ornamentation mostly reticular in the dorsal region. In the antero-dorsal region of the carapace (Fig. 5 : a1, a4), the osteoderms in the area near the cephalic notch are pentagonal to hexagonal, composed of a large central figure surrounded by a single row of 10 to 12 very small rounded peripheral figures and occasional accessory figures; this reduction in size is especially noticeable in the osteoderms forming the cephalic notch, where in some cases only the central figure is observed. The smallest osteoderms of the entire carapace are found on the expanded antero-lateral margin (for measurements see Online Resource 3, table 3), with an irregular pentagonal morphology that gradually changes to rectangular towards the ventral edge. These osteoderms have a central figure surrounded by a single row of very small peripheral figures, unlike Neosclerocalyptus in which no peripheral figures are observed (Quiñones et al. 2020 ). In the mid-dorsal region of the carapace (Fig. 5 : a2, a5), the osteoderms are typically pentagonal or hexagonal in outline, with a subcircular, slightly convex central figure surrounded by a single, complete row of 11 to 15 peripheral figures; in addition, a second incomplete row of peripheral figures is observed at the anterior and posterior margins, often extending into adjacent osteoderms. Towards the lateral margins of the carapace, osteoderms become increasingly rectangular in shape and exhibit a gradual reduction in size. In these osteoderms, the central figure possesses a single complete row of peripheral figures, with remnants of a second incomplete row present along the anterior margin. This specific pattern of multiplication of peripheral figures distinguishes S. ameghini from Nopachtus , which exhibits up to two complete rows, and from Propanochtus and Panochthus , which can develop 5–7 complete rows surrounding the central figure. In the first osteoderms of the postero-dorsal region (pelvic region) Stromaphorus ameghini (MLP-PV 29-X-10-2) has in the first row a total of 12 to 21 and the second between 21 to 27, in contrast to Panochthus (e.g., P. tuberculatus (Owen, 1845) and P. greslebini Castellanos, 1942 in which a significant increase in the number of peripheral figures results in a completely reticular pattern). In the postero-dorsal region of the carapace (Fig. 5 : a3, a6), each of the osteoderms consists of a large rounded and bulging central figure, approximately 30 mm by 32. 4 mm, similar to Stromaphorus trouessarti nov. comb. (Moreno, 1888 ) more convex than in Nopachtus coagmentatus Ameghino, 1888, but less than in Nopachtus cabrerai (Zamorano, Scillato-Yané, Gonzalez-Ruiz, Zurita 2011). It should be noted that in this sector of the carapace, the main axis of the central figure is inclined towards the posterior region of the osteoderm, similar to S. trouessarti nov. comb. but different from N. cabrerai in which it is perfectly vertical. In these osteoderms the central figure is surrounded by a complete row of 12 to 21 peripheral figures; while on its posterior margin, near the caudal notch, a second row of incomplete figures occurs, unlike in Nopachtus coagmentatus , which has two complete rows of peripheral figures and N. cabrerai with the second row complete but shared with the adjoining osteoderms. On the other hand, the osteoderms that constitute the caudal margin present the central figure with an anterior margin composed of 3 to 5 rows of peripheral figurines (see Fig. 6 ), a row towards the sides of the central figure and a free edge towards the posterior margin. In this posterior region of the carapace, the osteoderms are aligned in orderly bands, counting between 8 and 10 bands of osteoderms. Caudal tube Overall, the caudal tube (Fig. 7 ) of Stromaphorus ameghini (MLP-PV 29-X-10-1; MLP-PV 29-X-8-9; FMNH-P14414 and FMNH-P14532), both in morphology and rosette ornamentation, is somewhat similar to those of Pseudoplohophorus (FC-CVF 475 / 595 and MMC 888), Plohophorus figuratus (MLP-PV 98-XI-21-1), Eosclerocalyptus proximus (PVL 375) and Neosclerocalyptus (MMP 4300; MACN 13084 and PZ-Ctes 5879); however, it differs from the tubes of Hoplophorus euphractus (UFMG 1235), Nopachtus coagmentatus (MLP-PV 16–122), Propanochthus bullifer (MANC-Pv 1761), Panochthus (MLP-PV 16–29; AMNH 11243; MHNC-13491 and MANC-Pv 5130), Eleutherocercus solidus (FMNH-P14437; MLP-PV 16–25; MLP-PV 29-X-10-21 and MACN 2893), and Doedicurus clavicaudatus (MLP-PV 16–23). The caudal tube of S. ameghini is almost circular in transverse outline. In dorsal and ventral views, it has a relatively constant diameter towards the distal portion, resulting in an almost semi-rectangular morphology (for measurements see Online Resource 3, table 4). It differs from Pseudoplohophorus and Plohophorus whose tubes narrow markedly towards the distal region, acquiring a conical morphology; from Eosclerocalyptus and Neosclerocalyptus , which have narrower and more gracile tubes; from Hoplophorus euphractus whose tube has a robust cylindrical morphology, but is slightly dorso-ventrally flattened from the middle to the terminal portion; from Propanochthus , Panochthus and Eleutherocercus which are dorso-ventrally flattened even more than in Hoplophorus and Nopachtus ; and Doedicurus in which the distal end of its caudal tube is similar to a club. In dorsal view (Fig. 7 : a, c), the ornamentation of the caudal tube is composed of a simple rosette pattern, in which each osteoderm has a smooth-surfaced central figure, surrounded by a single complete row of peripheral figures, which in turn are shared with the adjoining osteoderms. The peripheral figures are rounded (for measurements see Online Resource 3, table 4), being smaller than those observed in Plohophorus. The morphology of the central figures of the proximal portion varies from circular to subcircular, they are large, and arranged in transverse rows. However, towards the distal portion, the central figures become gradually smaller. In the middle portion, the central figures are clearly oval, while those of the more distal portion revert to a more circular shape, all of them surrounded by a single series of peripheral figures shared with the contiguous central figures. At the apex there are two large smooth terminal figures (T) bordering this end of the tube to the sides, the lateral margins of these figures, in the more distal portion, are in contact, as in Plohophorus and Pseudoplohophorus , In turn they differ from Hoplophorus euphractus and Nopachtus with characteristic large lateral spines, from Propanochthus and Panochthus with very rugose depressed elliptical lateral figures, raised in the middle, from Eleutherocercus and Doedicurus with very rugose elliptical lateral and apical figures, but completely concave. T figures are preceded by 4 smooth lateral figures (I, II, III and IV) which will be explained in detail in the lateral view. Between the T figures in their anterior region, there is a characteristic rounded figure called posterior figure (P), which is also present in other genera such as Pseudoplohophorus (see Toriño and Perea 2018 ). Towards proximal regions of the tube, before P, there is symmetry between the left (L) and right (R) marginal figures (see Fig. 7 : a, c), thus up to 10 pairs of marginal figures have been identified. Between the marginal figures, there are other central figures arranged in rows, the number of which increases towards the proximal region of the tube. It should be noted that the arrangement and number of these central figures are slightly variable between the different individuals of the species. In ventral view, the ornamentation of the caudal tube is very similar to the dorsal side, although the central figures are somewhat larger and flatter. These figures, particularly at the terminal portion, come into contact without peripheral figures between them. An inverse relationship exists between the size of the central figures and the surrounding peripheral figures. As the central figures expand, the peripheral figures become significantly smaller (may even disappear), unlike the larger peripheral figures of the dorsal region. In lateral view (Fig. 7 : b, d) there are large terminal figures (T) in the distal portion, with a highly convex surface, as well as four lateral figures (numbered I, II, III and IV in distal-proximal sense, see Fig. 7 ) which gradually reduce in size towards the proximal portion. Figure I is convex, unlike figures II, III and IV, which are flatter. Between the lateral and marginal figures, there is a row of small peripheral figures. Finally, the number of lateral figures varies from 3 to 4, even in the same individual. This is the case of MLP-PV 29-X-10-1 whose right side has the four figures described above (I, II, III and IV) and whose left side has only three of them (I, II and III). Additional observations Regarding S. trouessarti nov. comb., a redescription and comparative diagnosis of the species can be found in Zamorano et al. ( 2011 ). Phylogenetic analysis The analysis resulted in two MPTs, only differing in the location of both species of “Palaehoplophorini” ( Palaehoplophoroides rothi and Palaehoplophorus meridionalis ). In turn, the strict consensus shows RI = 0.933 and CI = 0.845; length = 174.020 steps (Fig. 8 ). It demonstrates the monophyly of the family Glyptodontidae mainly supported by dental and mandibular synapomorphies, such as the presence of flat occlusal surfaces on all molariforms [25:1], trilobulation [30:0;34:1] and loss of premaxillary teeth [26:0]; but also by exoskeletal features such as the loss of mobile bands [49:1]. As mentioned above (see Introduction) we prefer the use of Glyptodontidae at family level, in agreement with most of the recent contributions, and taking into account the noticeable divergent morphology of glyptodonts when compared to the remaining Cingulata diversity (see Machado et al. 2022 ). Consistent with the observations of Croft et al. ( 2007 ), the enigmatic Parapropalaehoplophorus septentrionalis is the sister species of remaining diversity of Glyptodontidae. Among the most characteristic mandibular morphology stands out as particularly distinctive. Given that this anatomical feature was the subject of a geometric morphometric analysis (Character 0), it will be discussed separately at the conclusion of this section. The MPT topology shows that the remaining diversity of Glyptodontidae is divided into two major clades, Glyptodontinae and Hoplophorinae (=”Austral clade” of Barasoian et al. 2022a; Quiñones et al. 2023 ); these two clades have already been reported in previous works (see Cuadrelli et al. 2020 ; Quiñones et al. 2020 ; Nuñez-Blasco et al. 2021c; Barasoian et al. 2022a). Glyptodontinae is mainly supported by endoskeletal synapomorphies, some from the carapace and some from the caudal armour (Node A). The topology of this node is in agreement with Zurita et al. ( 2013 ) and Cuadrelli et al. ( 2020 ). The second clade, Hoplophorinae, is supported by four exoskeletal synapomorphies (increase in size of cephalic shield osteoderms [35:0], acquisition of peripheral figures in caudal rings osteoderms [71:1], [72:1], and caudal armour composed of caudal rings and absence of terminal tubercle [73:0]). Some Burdigalian (Early-Middle Miocene) taxa ( Propaleoplohophorus australi s, Cochlops muricatu s and Eucinepeltus petesatus ) are located at the base of this second radiation forming a monophyletic group, supported by a reduction in the number of molariforms with trilobulation [27:3], rings representing 80% of total length of the caudal armour [75:1]; the species Pro. australi s and C. muricatu s are grouped together, supported by ambiguous synapomorphies related to the cephalic shield (acquisition of peripheral figures [36:1], [38:1]); E. petesatus is located as the sister species of this group (but see Barasoain et al. 2022a for another interpretation), the three taxa are forming the tribe Propalaehoplophorini (= Propalaehoplophorinae sensu Ameghino, 1891c ). It is interesting to analyze at this point the character 74; this describes the morphology of the caudal armor composed of caudal rings and a caudal tube, which are present in all members of the Hoplophorinae with the exception of Eucinepeltus petesatus and Propaleoplohophorus australis . These two basal species have neither terminal tubercle (as in the case of Glyptodontinae) nor a fully developed caudal tube, but have another structure that could be intermediate; therefore, in characters 73 and 74, they have been coded as absent [73:0; 74:0]. The “Palaehoplophorini” ( Palaehoplophoroides rothi and Palaehoplophorus meridionals ) forms a polytomy with the remaining extra-patagonian diversity, this politomy is supported by exoskeletal synapomorphies (similar ornamentation between dorsal and caudal armors [70:1], presence of caudal tube [74:1 ambiguous] and rings representing up to 60% or less of total length of the caudal armour [75:2]). In turn, the Late Miocene (ca. 10 − 9 Ma) species Kelenkura castroi , appears as the sister taxon of the Neogene and Quaternary remaining glyptodonts, in accordance with the proposal of Barasoain et al. ( 2022a ). This relationship is supported by synapomorphies related to the caudal tube (acquisition of central figures and rosette pattern in the dorsal region [80:1], [84:1] the latter subsequently modified in Doedicurus , Panochthus and Propanochthus ; and acquisition of lateral figure s [86:1], it must be noted that this state changes throughout the evolution of the clade). On the other hand, the clade clustering the late Neogene and Quaternary glyptodonts is strongly supported by six exoskeletal synapomorphies (acquisition of more peripheral figures in the posterior region of the carapace [64:2]), caudal tube without visible rings [76:4], [78:1], caudal tube with a rosette pattern with peripheral figures well-developed and completely surrounding the central figures [82:2], acquisition of five to seven large lateral figures [88:1], loss of large lateral figures at the apex of the caudal tube [93:0]), and two endoskeletal synapomorphies (both related to the degree of development of the third trochanter [43:1 ambiguous], [44:1 ambiguous]). The first subdivision (and the subject of this analysis) is formed by the monophyletic group that clusters the species Pseudoplohophorus absolutus , Ps. benvenuti , Plohophorus figuratus, Stromaphorus ameghini and Stromaphorus trouessarti nov. comb. comprising the tribe Plohophorini (Node B), it is supported by one dental synapomorphy (increasing of size of Mf1 [29:0]); and three related to the carapace ornamentation (multiplication of peripheral figures of dorsal osteoderms [56:1] and peripheral carapace figures with circular morphology [60:1], [61,0]). Describing in more detail the internal relationships of the species forming the tribe Plohophorini, the Pampean species Plohophorus figuratus appears as sister taxon to Plohophorus avellaneda (ornamentation of the proximal-ventral region of the caudal tube [95:0]); and these in turn as sister group of Stromaphorus ameghini + Stromaphorus trouessarti nov. comb. (supported by four carapace synapomorphies (loss of peripheral figures in osteorderms of the cephalic notch [52:0]; surface of the osteoderms of the posterior region of the carapace becoming blistered [62:2]; and increase in the number of peripheral figures in the posterior region of the carapace and caudal notch [64:4], [66:2]), thus confirming the condition of natural group of the genus Stromaphorus . The relationship between Stromaphorus and Plohophorus is supported by two synapomorphies (position of the infraorbitary foramen [15:2], and increase in the number of rows of peripheral figures in the posterior region of the carapace [63:2]); in turn, they form the sister group of the late Neogene Uruguayan taxa Pseudoplohophorus absolutus and Ps. benvenuti , ( supported by the morphology of the palate [24:1 ambiguous synapomorphy] ) . Plohophorini is recovered as the sister group of the remaining members of this second clade (Doedicurini + Hoplophorini + Neosclerocalyptini), wich is supported by a cranial synapomorphy (position of the infraorbitary foramen [15:1]), plus another from the caudal armour (acquisition of polygonal ornamentation in the caudal tube [85:1]) (but see Fernicola and Porpino 2012 for another proposal). The following node (Node C) encompasses the tribe Doedicurini (= Doedicurinae sensu Trouessart, 1897 ) supported by three cranial synapomorphies (angle between posterior margin of the orbital notch and the palate plane near to 90º [20:1], morphology of the palate acquiring a similar transverse expansion at their posterior and anterior margins [24:2 ambiguous], and increase in the antero-posterior diameter of Mf1, with oval morphology [29:1]); seven carapace synapomorphies (most of them related to the loss of ornamentation [52:3], [54:0], [62:3], [63:0], [64:0], [66:4], one related to the acquisition of interdigitation of the margin of its osteoderms [57:2]); and five caudal armour synapomorphies (transverse outline of distal third of caudal tube becoming dorsoventrally flattened [77:1 ambiguous], cylindrical-conical caudal tube composed of ankylosed osteoderms acquiring large lateral depressions (concave insertion structures) [86:3], [88:2], [89:1], [91:1]). Doedicurini appears as the sister group of Eosclerocalyptus proximus , forming a clade supported by two cranial synapomorphies (labiolingual trilobation evident from Mf2 [27:1 ambiguous], anteroposterior diameter of Mf1 less than 50% of the anteroposterior diameter of Mf2 [29:2 ambiguous]); two carapace synapomorphies (increasing in the number of polygonal peripheral figures in the carapace [61:2], the central figure occupies less than 50% of the total surface area of the osteoderms near the caudal notch [68:0 ambiguous]), plus another from the caudal armour (peripheral figures surround the entire central figure in the central lateral zone of the caudal tube [87:3]); and the group Neosclerocalyptini + Hoplophorini, supported cranial synapomorphies (acquisition of pneumatization of the rostral area of the skull [4:1], acquisition of peripheral figures in cephalic shield [36:1 ambiguous], acquisition of peripheral rows in the cephalic shield [38:1 ambiguous]). The tribe Neosclerocalyptini (Node D) is supported by six cranial synapomorphies (acquisition of ossified nasal cartilages [5:1] and [8:1], acquisition of a “V” groove separating the modified nasal area from the rest of the skull [9:1], ventral edge of the orbital notch coinciding with the 50% of the dorso-ventral diameter of the ossified nasal cartilages [12:1], distance between the infraorbitary foramen and the labial margin of the Mf3, equal or larger than the antero-posterior diameter of the Mf1-Mf2 [14:1], acquisition of cephalic shield with sub-angular outline [37:1]); and three carapace synapomorphies (loss of convexity of carapace, in lateral view [47:0] and [48:3], acquisition of antero-lateral expansion of the carapace [50:1]). Neosclerocalyptini is the sister group of the tribe Hoplophorini (Node E) grouped by a total of five synapomorphies, of which the most relevant have been those of the caudal armour (acquisition of sigmoid contour in the anterolateral border of the nasal openings [21:1], transverse outline of distal third of caudal tube dorsoventrally flattened [77:1 ambiguous], acquisition of convex insertion structures in the lateral figures of the caudal tube [86:2], increasing rows of peripheral figures around the entire central lateral figure [87:4], acquisition of depressions with convex central area in the terminal zone of the caudal tube [90:1]), this latter (Hoplophorini) formed by the species Hoplophorus euphractus , Nopachtus coagmentatus , Propanochtus bullifer , Panochthus tuberculatus , and P. intermedius . In turn, the geometric morphometry applied to the mandibles in lateral view has been included as an additional character (Character 0) in the phylogenetic matrix, resulting in a mixed matrix (conventional characters + geometric morphometry characters). Character 0 has played a crucial role in establishing Pa. septentrionalis as the sister taxon to Glyptodontinae + Hoplophorinae. This is primarily attributed to the unique 90º angle formed by the horizontal and ascending mandibular rami. This morphology represents a clearly plesiomorphic condition, intermediate between that of Pampatheriidae or Dasypodidae (greater than 90º) and the more derived Glyptodontidae (less than 90º). On the other hand, it also contributed for grouping the Propaleoplohophorini by their morphological affinity. Finally, the geometric morphometry has increased the support of some nodes when calculating the Bremer, absolute Bremer, and Jackknifing indices (Fig. 8 ) in comparison with the values obtained in trees of similar topology from previous analyses (see Cuadrelli et al. 2020 ; Quiñones et al. 2020 ; Nuñez-Blasco et al. 2021c, 2022 ; Barasoain et al. 2022a ). The most interesting nodes, positively affected, were those that group Plohophorini (Node B), Doedicurini (Node C), Neosclerocalyptini (Node D), and Hoplophorini (Node E). The support values obtained for Node B (Plohophorini) are for the first time notably high, Bremer index of 100, absolute Bremer index of "3?" (calculated with 3-step suboptimality) and Jackknifing of 87. Finally, the comparison between the morphotypes representative of each of these subsets, also increased the support of the node that groups them (Doedicurini + Hoplophorini) resulting in a Bremer index value of 40, absolute Bremer index of 2 (calculated with 3-step suboptimality) and Jackknifing of 63. Local geographical and stratigraphic distribution The holotypes of “ Phlyctaenopyga ” ameghini MLP-PV 16–101 ( Plohophorus ameghini , Ameghino, 1889 ) and “ Stromaphorus compressidens ” MLP-PV 16–138 ( Neuryurus compressidens , Moreno and Mercerat, 1891 ) come from the "Araucanian" levels of the “Bajo de Andalhuala” in the southern part of the SMV, probably from the upper levels of the Chiquimil Formation or from the Andalhuala Formation (i.e. between 6.88 to 4.85–3.4 Ma; Bonini et al. 2021 ). The skull FMNH-P14396, from Ampajango locality (Santa María Department, Catamarca Province) originaly referred to S. compressidens , comes from XI level (by Marshall and Patterson 1981 , Appendix II), and according to the correlation proposed by other authors, such as Georgieff et al. ( 2017 ) and Bonini et al. ( 2021 ), this level is between 6.88 to 6.02 Ma. The specimen FMNH-P14367, a carapace here reassigned to a juvenile of S. ameghini from Loma Rica locality (Andalhuala, Santa María Department, Catamarca Province), cames from XVII level (Marshall and Patterson 1981 ), and according to Georgieff et al. ( 2017 ) and Bonini et al. ( 2021 ), this level is above 6.02Ma. The materials MLP-PV 16–134, MLP-PV 16–135, MLP-PV 19–136 (Holotype of Plohophorus philippii Moreno and Mercerat, 1891 ), and MLP-PV 16–137 referred to “ S. compressidens ”, come from north ridge of Loma Rica (Andalhuala, Santa María Department, Catamarca Province), based on the stratigraphic levels outcropping in that area, their age could be slightly lower than 6.02Ma (see Fig. 1 ). Regarding the Villavil-Quillay Basin (located ca. 78 km at southwest from the Santa Maria Valley), there are several localities where numerous fossils have been found, belonging to both species (“ Phlyctaenopyga ” ameghini and “ Stromaphorus compressidens ”). The most remarkable localities are Puerta de Corral Quemado, Belén Department (“ Ph. ” ameghini : MLP-PV 29-X-8-2; MLP-PV 29-X-8-7; MLP-PV 29-X-10-6; MLP-PV 29-X-10-10; FMNH-P14439; FMNH-P14532; FMNH-P14414; FMNH-P14447; CCPCQ 03-DPA-Pv264 and “ S. compressidens ”: MLP-PV 29-X-8-1; MLP-PV 29-X-8-9; MLP-PV 29-X-10-54; FMNH-P14494; FMNH-P14520; FMNH-P14414; FMNH-P14494; FMNH-P14520) and Corral Quemado, Belén Department (“ Ph. ” ameghini : MLP-PV 31-XI-12-7). In turn, the second most fossiliferous locality for these species is San Fernando (North and South), Belén Department (“ Ph. ” ameghini : MLP-PV 29-X-10-1; MLP-PV 29-X-10-2; MLP-PV 29-X-10-3; MLP-PV 29-X-10-5; MLP-PV 29-X-10-40; MCH-P39; MCH-P176; MCH-P177; MCH-P247; MCH-P323, and “ S. compressidens ”: MLP-PV 29-X-10-8; MLP-PV 29-X-10-51; MCH-P174; MCH-P328) (see Cabrera 1944 ; Marshall and Paterson 1981; Bonini 2014 ). It should be noted that the remains referring to both taxa not only appear indistinctly in these localities, but also in the same time interval, specifically in the Andalhuala and Chiquimil formations, in strata dated between 7.14 +/- 0.05 and 3.66 +/- 0.05 Ma. In adittion, we also report at least two records located above the tuff dated at 3.66 (FMNH-P14447) and below that dated at 7.14 (MCH-P364), but without greater stratigraphic precision. Discussion Taxomomy and nomenclature Glyptodonts were very probably the most fascinating and unusual xenarthrans that ever existed. Recent findings of glyptodonts from late Neogene sediments of Ecuador, which are morphologically distant from those known of La Venta (Colombia) and Anzoategui (Venezuela) (see Zurita et al. 2013 ) in northern South America, and from those of southern South America (see González-Ruiz 2010 ; Barasoian et al. 2022a), reveal that much more studies are necessary to improve our knowledge about their evolutionary history (Román-Carrión et al. 2022 ). As mentioned above, the tribe Plohophorini ( sensu Castellanos 1932 ; Hoffstetter 1958 ) is one of the most enigmatic groups within the southern clade Hoplophorinae, and particularly those taxa from the late Neogene of the NWA region were, before this contribution, poorly known from the taxonomic, stratigraphic and phylogenetic viewpoint. In the Villavil-Quillay Basin and Santa María Valley two species were recognized, “ Phlyctaenopyga ” ameghini (Ameghino, 1889 ) and “ Stromaphorus compressidens ” (Moreno and Mercerat, 1891 ), both sharing the same stratigraphic and geographic provenance (see Cabrera 1944 ; this work). Two issues can be inferred from geographical and stratigraphic provenance data (see Fig. 1 ): (1) the remains are quite numerous, so they were common taxa in this area; (2) two different, but at the same time so similar species coexisting at the same time in the same area would have resulted in high interspecific competition, due to direct rivalry for the same resources. From an ecological perspective, the co-ocurrence of two different species is hardly feasible, as it was observed in Pleistocene genera (see Cuadrelli et al., 2020 ). In this regard, a careful morphologic and taxonomic analysis made in this contribution revealed that, actually, both names must be considered as synonyms, being Stromaphorus ameghini (Ameghino, 1889 ; ex Moreno, 1882 ) the only species of Plohophorini present in the late Neogene sequences of NWA. Following the ICZN 2000, Art. 23.1 and 23.3, the genus Stromaphorus Castellanos, 1925 , has priority over Phlyctaenopyg a Cabrera, 1944 , while the specific epithet “ ameghini ” has priority over “ compressidens ” (see Online Resource 1). Besides, the comparative study with “ Phlyctaenopyga ” trouessarti from the Monte Hermoso Formation (Pliocene) revealed the presence of some shared characters between both species at the level of the dorsal carapace (see Description and comparisons) that suggests the inclusion of this Pampean species in the genus Stromaphorus , as S. trouessarti (Moreno, 1888 ) nov. comb. In consequence, the genus Stromaphorus now encompasses two species, S. ameghini and S. trouessarti nov. comb. In summary, given the noticeable anatomical similarities between both species already discussed (ie., “ Ph. ” ameghini and “ S. compressidens ”), and taking into account that no spatial discontinuities are observed (through their geographical provenance) nor temporal discontinuities (through their stratigraphic provenance as well as the dating of the tuffaceous levels), following the criteria of Simpson (1951) and Castro et al. (2013), the evidence supports the existence of a single species, Stromaphorus ameghini , among Plohophorini glyptodonts of NWA. Phylogeny As observed in previous analyses (e.g., Barasoian et al. 2022a; Quiñones et al. 2023 ), two large clades can be observed. One of northern origin, Glyptodontinae, and the other object of this study, Hoplophorinae, which includes the remaining diversity. Within the southern lineage Hoplophorinae, cladistic results show that the tribe Plohophorini ( sensu Castellanos 1932 (nom. transl. Hoffstetter 1958 ) appears as the sister group of Doedicurini + Hoplophorini + Neosclerocaliptini, forming part of the extra-Patagonian radiation starting with Kelenkura castroi (see Barasoain et al. 2022a ). The result of our analysis shows the condition of natural group of Plohophorini (Fig. 8 , Node B), a result that agrees with Quiñones et al. ( 2023 ); this condition is supported by several cranial and postcranial synapomorphies. Regrettably, we were no able to include all the species traditionally interpreted as belonging to Plohophorini (see Hofstetter 1958; Paula Couto, 1979 ), since many of them are recognized on the basis of very fragmentary type material within a strict typological taxonomic criterion and lacking precise stratigraphic provenance (Toriño and Perea 2018 ). Besides this, one interesting point is the distribution of the different species in this so far poorly known tribe. First, as observed in the topology of the MPT (see Fig. 8 ), the monophyly of the genera Stromaphorus ( S. ameghini + S. trouessarti nov. comb. ), Plohophorus ( P. figuratus + P. avellaneda ) and Pseudoplohophorus ( Ps. absolutus + Ps. benvenuti ) is confirmed and supported by several cranial and postcranial synapomorphies (see Online Resource 4 and Online Resource 5). Moreover, Plohophorus figuratus + P. avellaneda appears as the sister group of the clade clustering S. ameghini + S. trouessarti nov. comb. , condition supported by a couple of cranial and exoskeletal synampomorhies. Beyond the phylogeny of the Plohophorini, another interesting point refers to the location of the enigmatic species Nopachtus coagmentatus , here interpreted as the sister and earliest divergent taxa of the lineage containing Hoplophorus , Propanochthus bullifer + Panochthus spp. The phylogenetic position of N. coagmentatus has been controversial since Ameghino ( 1889 ) in a pre-cladistic time. More recently, Zamorano and Brandoni ( 2013 ) placed this species close to the genera Plophophorus, Pseudoplohophorus, Propanochtus , “ Phlyctaenopyga ”, and Stromaphorus . However, and as mentioned above, our results show that N. coagmentatus is in fact the sister species within the lineage including Hoplophorus , Propanochthus , and Panochthus spp. (Hoplophorini). This suggests that the multiplication of peripheral figures of this lineage and that of the Plohophorini here analyzed (i.e., Pseudoplohophoru s, Plohophorus and Stromaphorus ) is actually a homoplasy; in this sense our results coincide with those presented by Fernicola and Porpino ( 2012 ), in which these authors already postulated a possible homoplasy between the rosette patterns ornamenting the dorsal carapace of these groups. This hypothesis is also supported by the evident morphological similitude in the caudal tubes of Nopachtus, Propanochthu s, Hoplophorus , and Panochthus , which in turn are very different from those of Pseudoplohophorus , Plohophorus and Stromaphorus . Also, the spine-like structure observed in the caudal tubes of the lineage Nopachtus coagmentatus, Hoplophorus euphractus, Propanochthus bullifer , and Panochthus spp. is a homologous structure rather than a convergence as usually interpreted. Finally, it is noteworthy the position of the Early Miocene glyptodont P. septentrionalis as the earliest divergent member of Glyptodontidae, supported by geometric morphometric character (Character 0). This result is consistent with the analysis of Croft et al. ( 2007 ) that recovered this species as the sister taxon of all remaining glyptodonts. Paleobiogeography, palaeoenvironments, stratigraphy, and First Appearance Datum (FAD) of Plohophorini In southern South America (ca 48° to 21° S), where records of Neogene glyptodonts are quite frequent (see Gaudin and Croft 2015 ; Zurita et al. 2016a ), a recent revision of the diversity of the Late Miocene [Chasicoan lapse, Tortonian (ca. 10 − 9 Ma; see Prevosti et al. 2021 )] in central Argentina revealed a much more restricted diversity than previously supposed, with the presence of one species, Kelenkura castroi (Barasoain et al., 2022). Following the palaeophytogeographic scheme proposed by Barreda et al. ( 2007 ; Fig. 2.2.), the Proto-Spinal/Steppe province was mainly characterized during the Late Miocene ( ca . 10 − 8 Ma) by the presence of low xerophytic shrubs or trees and mostly halophytic herbaceous vegetation in open environments, consistent with the progressive aridization and open biomes, and a shift toward the consumption of C4 plants in several herbivorous clades (Domingo et al. 2020 ; Sanz-Pérez et al. 2023 ). The presence of a single species, Kelenkura castroi , in central Argentina represents the first extra-Patagonian radiation within Hoplophorinae at the onset of the so-called "Age of the Southern Plains", an event that took place between 10 − 3 Ma and that was a direct consequence of the pulse of drying and cooling observed at the end of the Miocene Optimum Climatic (see Zachos et al. 2001 ; Westerhold et al. 2020 ; Domingo et al. 2020 ; Candela et al. 2021 ). The subsequent period of diversification of Hoplophorinae may have been influenced by the progressive development of open biomes, as suggested by previous studies (see Zurita et al. 2016a ; Barasoain et al. 2022a ; Domingo et al. 2020 ). The FAD of S. ameghini in the NOA is not clear, since depending on the criterion it can be established at ca. 9 or 7 Ma. The material MCH-P364 assigned to this species was exhumed from lacustrine sediments considered as lateral facies changes of the upper section of the El Áspero Member in transition to El Jarillal Member (Chiquimil Formation) outcropping in the FOCP, San Fernando, Catamarca (Moyano, 2003 ; Muruaga et al. 2003 ; Armella and Bonini, 2020 ). The intrusive andesitic lacolith defined as the El Áspero Member in Villavil locality was dated in 9.14 ± 0.09 Ma (Sasso, 1997 ), and the lacustrine facies in the FOCP have been interpreted as part of damming of a similar age volcanic event (see Bossi et al. 1999 ; Moyano, 2003 ; Georgieff et al. 2017 ). Likewise, the upper limit of the Jarillal Member has been dated in 7.14 ± 0.02 (Latorre et al. 1997 ). If this is correct, it implies that records of Plohophorini in the NWA are older than those of the PR, where the Chasicoan glyptodonts (ca. 9 − 8) are limited to K. castroi , since the presence of supposed Plohophorini (see Bondesio et al. 1980 ) was discarded by Barasoain et al. ( 2022a ). According to the available evidence, no records of Plohophorini are present in the PR until the Montehermosan lapse, ca. 5.3 Ma (see Tomassini et al. 2013 ; Prevosti et al. 2021 ). A similar situation occurs with Doedicurini glyptodonts, which have their oldest records in the NWA (see Núñez-Blasco et al. 2020 , 2021c , 2022). As mentioned above, the subsequent evolutionary history of the Plohophorini glyptodonts from NWA here analyzed mostly includes the latest Miocene (ca. 7.14 Ma) to the Late Pliocene (ca. 3.66 Ma) interval, during the Messsinian and Zanclean lapse, characterized by an evident expansion of C4-dominated grasslands in southern South America in ca . 8–7 Ma (Hynek et al. 2012 ; Domingo et al. 2020 ). From a stratigraphic viewpoint, the materials here analyzed mostly come from the Andalhuala Formation, ranging from 7.14.to 3.66 in VQB (see Esteban et al. 2014 ; Georgieff et al. 2017 ; Bonini et al. 2021 ), except one particular fossil (FMNH-P14447) that comes from the Corral Quemado Formation and thus could be younger than 3.66 Ma. However, its imprecise stratigraphical provenance does not allow us to infer its real age. It must be noted that the ghost range predicted for this group in the calibrated phylogeny extends its origin down to ca. 10 Ma (Fig. 8 ). As mentioned above, there is an interesting temporal overlap between the expansion of C4 grassland and the diversification of Hoplophorinae, which deserves further study (see also Núñez-Blasco et al. 2020 , 2021c ). Regarding the Plohophorini recorded in Argentina and here analyzed, S. ameghini, S. trouessarti nov. comb. , and P. figuratus lived during the latest Miocene-Pliocene under different paleoenvironmental conditions. This is because S. trouessarti nov. comb. and P. figuratus were geographically restricted to the Proto-Espinal/Steppe province, while the distribution of S. ameghini coincides with the Neotropical province, this latter, and according to Barreda et al. ( 2007 ;Fig. 2.2), characterized by vegetation like the current Chaqueña Province (xerophytic forests, associated with palm groves, savannahs and halophytic shrub-steppe, and hydrophytic communities linked to watercourses.). It is important to emphasize that a similar paleobiogeographic pattern was already observed by Núñez-Blasco et al. ( 2021c , 2022) when analyzing another clade of glyptodonts, Doedicurini, present in the late Neogene sediments of the NW and PR of Argentina. The genus Eleutherocercus contains two species, E. antiquus (Zanclean-middle Piacencian; ca 5.33–2.53 Ma) from the PR of Argentina, and E. solidus (middle Messinian-middle Piacencian; ca 7.14-3Ma) from NWA (Zurita et al. 2016a ; Núñez-Blasco et al. 2020 , 2021c ). Moreover, even though they are currently under study, a similar geographic pattern could be inferred for another genus present in the late Neogene of NW and PR of Argentina, Eosclerocalyptus . This clusters two species, E. proximus , restricted to NWA (Catamarca Province) and E. lineatus , in the PR (Buenos Aires Province) (Zurita and Tomassini 2006 ; Zurita 2007a , b ). Something similar happens with the distribution of Nopachtus Ameghino, 1888, represented by the species N. coagmentatus (Ameghino, 1888) in NWA (Catamarca Province, see Zamorano et al. 2017 )d rdoba Province, and N. cabrerai Zamorano et al. ( 2011 ) in the PR (Buenos Aires Province). Consequently, during the Late Miocene and Pliocene, NWA (Neotropical province sensu Barreda et al. 2007 ) and the PR (Proto-Spinal/Steppe province sensu Barreda et al. 2007 ) shared the same genera ( Eleutherocercus , Stromaphorus, Nopachtus , and Eosclerocalyptus ) but different species. More precisely, within the Neotropical province, some differences between NWA and NEA (North Eastern Argentina) seem possible, since glyptodonts from the “Mesopotamiense” (Late Miocene) include another species of Eleutherocercus ( E. paranensis ), some putative species of “Palaehoplophorini” and other very poorly characterized taxa, mostly limited to isolated osteoderms and/or fragmented portions of the dorsal carapace (Scillato-Yané et al. 2013 ). Perhaps, the closed and forested biomes and tropical/subtropical conditions inferred for the Late Miocene of NEA (see Schmidt et al. 2020 ) stimulated this difference between NW and NE regions. However, further studies with more complete materials from the Ituzaingó Formation are necessary in order to infer the taxonomic diversity of glyptodonts in NEA. A similar distribution of different species of Tremacyllus (Hegetotheriidae, Notoungulata) in NWA and PR was observed. Armella et al. (2022) interpreted these differences as related to paleoenvironmental and paleo-phytogeographic features. These patterns could be interpreted as disjunctive distribution following the criteria of Morrone and Escalante ( 2009 ) in which a barrier (rivers, mountains, vegetation, paleoecological differences) could have interrupted gene flow between different populations and derived in the allopatric speciation of these populations. In the case of the glyptodonts and taking into account the Neogene geologic history of NWA, these barriers could be related to tectonic and/or climatic-environmental changes. During the Late Miocene, NWA experienced a series of tectonic uplifts, which resulted in the elevation of the mountain chain of Northwestern Pampean and Aconquija Ranges (Bossi et al. 2001 ; Georgieff et al. 2017 ), which probably developed a physical barrier for the distribution of the different groups of animals. In addition, by this time, one of the most important environmental changes is recorded in NWA, where plants with C4 photosynthetic pathway became dominant (Latorre et al. 1997 ; Hynek et al. 2012 ; Sanz-Pérez et al. 2023 ), perhaps in response to global temperatures gradually decreasing through the Late Miocene to the Early Pliocene (6 Ma) (Prevosti and Forasiepi 2018 ). So, two scenarios, and probably not the only ones, can be considered. In this respect, the endemic diversity of glyptodonts observed in NWA is also recorded in other related clades of cingulates, the “armadillos” Chlamyphoridae, in which at least two species ( Paleuphractus argentinus (Moreno and Mercerat, 1891 ) Kraglievich, 1934 and Paraeuphractus prominens (Moreno and Mercerat, 1891 ) Scillato-Yane, 1975 are considered as endemic of this area (see Barasoain et al. 2022b ). As mentioned above, this particular pattern observed in armadillos and glyptodonts could be explained by the tectonic activity of the uplift of the Andean mountains, resulting in a more temperate climate during the Late Miocene and Pliocene lapse, because the cordillera acted as a barrier for the Atlantic wet winds (Starck and Anzótegui 2001 ; Barasoain et al. 2022b ). Moreover, palaeobotanical evidence from the late Neogene sequence of NWA indicates warmer conditions with a marked seasonality (Anzótegui et al. 2017 ; Candela et al. 2021 ). Besides, the stratigraphic distribution of the Plohophorini glyptodonts of PR (ie. S. trouessarti nov. comb. and P. figuratus ) from the Monte Hermoso Formation (ca. 4.7–3.7 Ma; see Prevosti et al. 2021 ) mostly coincides with a warm and humid pulse recorded during the Pliocene (ca. 4.7–3.1 Ma) (Haug et al. 2001 ; Domingo et al. 2020 ; Prevosti et al. 2021 ), and a similar scenario can be inferred for both species of Pseudoplohophorus from the Camacho Formation (ca. 7.2-6 Ma) in Uruguay (Del Río et al. 2018 ; Aumond et al. 2021 ; Perea et al. 2020 ). In the PR, the records of Plohophorini ( Plohophorus figuratus ) continue through the Chapadmalalan (Zanclean and Piacenzian, ca. 3.7–3.04 Ma; see Zurita et al. 2016a ; Prevosti et al. 2021 ) and Marplatan ( P. avellaneda ) (Piacenzian, ca 2.8 Ma) ages under mostly wet and warm conditions. According to the fossil evidence, records belonging to this tribe are among the most frequent in glyptodonts, proving that the populations were healthy in that period (see Zurita et al. 2016a ). The last records of Plohophorini in the PR, at ca. 2.53 Ma, (see Quiñones et al. 2023 ) and at ca. 3.66 Ma in NWA (this work) coincides with a global cooling generated by the expansion of ice in Antarctica at ~ 3.0 and 2.7 Ma (see Domingo et al. 2020 ; Prevosti et al. 2021 ). In addition, these last records chronologically overlap with the arrival, from lower and middle latitude, of the other large clade of Glyptodontidae, the Glyptodontinae (Cuadrelli, 2020). Conclusions The results of the present contribution indicate that: (1) The names “ Stromaphorus compressidens ” (Moreno and Mercerat, 1891 ) and “ Phlyctaenopyga ” ameghini (Ameghino, 1889 ) refer to the same species; therefore, a synonymy is established with Stromaphorus ameghini (Ameghino, 1889 ; ex Moreno, 1882 ) representing the only Plohophorini glyptodont in the late Neogene sequence of NWA. (2) The cladistic analysis confirms that the genera Pseudoplohophorus , Plohophorus and Stromaphorus are monophyletic, being Plohophorini a well-supported clade within Hoplophorinae. (3) The location of the enigmatic species Nopachtus coagmentatus within Hoplophorini lineage indicates that the spine-like structure of the caudal tube must be interpreted as a homologous structure rather than a convergence. On the contrary, the multiplication of peripheral figures in both lineages (i.e., Hoplophorini and Plohophorini) is a homoplasy. (4) From a stratigraphic point of view, the FAD of S. ameghini is not precise since it could be established between ca.7 or 9 Ma; however, regardless of the discrepancy in this numerical range, a minimum age is established for the Plohophorini lineage, being the records of NWA older than those of the PR, where the diversity achieved by this clade was larger. The certain stratigraphic distribution of S. ameghini in the NWA is ca. 7.14–3.66 Ma, while the distribution of Plohophorini (ie, Plohophorus spp. and S. trouessarti nov. comb. ) in the PR is ca. 4.7–2.53 Ma. (5) The same genera of glyptodonts have different species in NWA and PR; this differentiation is probably related to the different palaeophytogeographic pattern. S. trouessarti nov. comb. , P. figuratus , E. antiquus , Eo. lineatus and N. cabrerai were geographically restricted to the Proto-Espinal/Steppe province. In turn, the distribution of the species S. ameghini , E. solidus , Eo. proximus and N. coagmentatus coincides with the Neotropical province. Furthermore, this separation could have been accentuated by the presence of a geographical barrier (e.g. course of a large river, mountainous elevation, etc.). (6) The last record of Plohophorini at ca. 2.53 Ma in the PR coincides with a global cooling event very close to the end of the Pliocene and the beginning of the Pleistocene, while those of the NWA are somewhat older, in ca. 3.66 Ma. Declarations Acknowledgements We thank Susana Bargo, Martín L. de los Reyes and Marcelo Reguero (Museo de La Plata, La Plata, Argentina), Laura Cruz (Museo Argentino de Ciencias Naturales Bernardino Rivadavia, MACN, Buenos Aires, Argentina), William Simpson and Keneth Angielezyk (Field Museum of Natural History FMNH, Chicago, USA) and the Museo “Condor Huasi” Belén Catamarca, Argentina, for granting us access to their collections; and Dirección de Antropología de Catamarca for permitting work in this province. We also thank Gabriela Schmidt (CONICET-Prov. ER-UADER, Entre Ríos, Argentina), Johana Baez and Juan Manuel Robledo (CECOAL-CONICET-UNNE, Corrientes Capital, Argentina), Federico Degrange and Ivana Tapia (CICTERRA-CONICET-UNC, Córdoba, Argentina), Lucia M. Ibáñez and Matias A. Armella (Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, Tucumán, Argentina), María Carolina Madozzo Jaén (INSUGEO-CONICET-UNT, Tucumán, Argentina), for their help during fieldwork. Special mention to Marcos Roig for his help during the palaeobiogeographical analysis, Nicolás Bauzá with phylogenetic morphometric analysis, Daniel Perea for the photographs provided, and Alvaro Mones and Cristina Luisa Scioscia for his help with nomenclatural issues. Author Contributions Statement Núñez-Blasco A performed the geometric morphometric analysis and applied it to the phylogenetic analysis, and also prepared all the online resource; Núñez-Blasco A., Zurita A. and Toriño P. performed the phylogenetic analysis and the chronological calibration of the tree with R; Núñez-Blasco A. and Zurita A. E. made the taxonomical and systematic assignment; Núñez-Blasco A., Zurita A. E., Bonini R., Miño-Boilini A. R. and Toriño P. wrote the introduction and materials and methods texts and Historical background Núñez-Blasco A., Zurita A. E., Quiñones S. I. and Zamorano M. wrote the anatomical descriptions; Nuñez-Blasco A. and Bonini R. prepared figures 1-8. Núñez-Blasco A., Zurita A. E., Bonini R., Miño-Boilini A. R., Toriño P. and Zamorano M. wrote the discussion and conclusion texts; Georgieff S. analysed the stratigraphy and relative dating of the fossils. All authors reviewed the manuscript. Funding This research was partially funded by PICT 2019-03412, PICT 2018 003380, PI Q002/21 (SGCyT-UNNE), and SNI_2020_1_1010231 ANII (P.T.). Data Availability All data generated and analyzed during this study are included in this published article and its supplementary iles. Disclosure statement No potential conflict of interest was reported by the authors. References Ameghino F (1888a) Rápidas diagnosis de algunos mamíferos fósiles nuevos de la República Argentina . Pablo E. Coni e Hijos, Buenos Aires. 17pp. Ameghino F (1888b) Lista de las especies de mamíferos fósiles del mioceno superior de Monte Hermoso, hasta ahora conocidas . Pablo E. Coni e Hijos, Buenos Aires. 21pp. Ameghino F (1889) Contribuciones al conocimiento de los mamíferos fósiles de la República Argentina, Actas Acad Nac de Cienc en Córdoba VI, Buenos Aires. pp 850, lams. LVI and LXXXVII. Ameghino F (1891a) Sobre algunos restos de mamíferos fósiles, recogidos por el Señor Manuel B. 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Zurita","email":"","orcid":"","institution":"Centro de Ecología Aplicada del Litoral (CECOAL-CONICET)","correspondingAuthor":false,"prefix":"","firstName":"Alfredo","middleName":"E.","lastName":"Zurita","suffix":""},{"id":270374269,"identity":"df4ec98c-46e1-4daa-b9c5-bfd2e252c9b6","order_by":2,"name":"Ricardo Bonini","email":"","orcid":"","institution":"Instituto de investigaciones Arqueológicas y Paleontológicas del Cuaternario Pampeano (INCUAPA-CONICET), Facultad de Ciencias Sociales, Universidad Nacional del Centro de la Provincia de Buenos Aires","correspondingAuthor":false,"prefix":"","firstName":"Ricardo","middleName":"","lastName":"Bonini","suffix":""},{"id":270374270,"identity":"8e9b4e6e-38f1-422a-bfac-a357a705f73b","order_by":3,"name":"Angel R. 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Quiñones","email":"","orcid":"","institution":"Centro de Ecología Aplicada del Litoral (CECOAL-CONICET)","correspondingAuthor":false,"prefix":"","firstName":"Sofia","middleName":"I.","lastName":"Quiñones","suffix":""},{"id":270374272,"identity":"4ea1aff4-f6fd-47c8-b6aa-dd7352ca7adb","order_by":5,"name":"Pablo Toriño","email":"","orcid":"","institution":"Instituto de Ciencias Geológicas, Facultad de Ciencias, Iguá 4225, 11400","correspondingAuthor":false,"prefix":"","firstName":"Pablo","middleName":"","lastName":"Toriño","suffix":""},{"id":270374273,"identity":"78af6e25-d19f-4707-aa6a-b54157adf283","order_by":6,"name":"Martín Zamorano","email":"","orcid":"","institution":"División Paleontología de Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata. Paseo del Bosque s/n. Código postal 1900. La Plata. CONICET.","correspondingAuthor":false,"prefix":"","firstName":"Martín","middleName":"","lastName":"Zamorano","suffix":""},{"id":270374274,"identity":"4b8f42f4-cf58-4c22-a817-b0037920887c","order_by":7,"name":"Sergio M. Georgieff","email":"","orcid":"","institution":"IESGLO, Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, Miguel Lillo 205, T4000JFE y CONICET, CCT NOA Sur, San Miguel de Tucumán","correspondingAuthor":false,"prefix":"","firstName":"Sergio","middleName":"M.","lastName":"Georgieff","suffix":""}],"badges":[],"createdAt":"2024-01-31 19:31:50","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3914918/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3914918/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10914-024-09726-3","type":"published","date":"2024-08-17T15:56:58+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":50548482,"identity":"5ae30bcf-637f-4731-b1ab-397d492095de","added_by":"auto","created_at":"2024-02-02 10:03:03","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":11558057,"visible":true,"origin":"","legend":"\u003cp\u003eGeographical map and stratigraphic distribution of fossils historically referred to “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e” and “\u003cem\u003ePhlyctaenopyga\u003c/em\u003e”\u003cem\u003e ameghini\u003c/em\u003e. The stratigraphic profiles represent the general series for each basin, based on information from Marshall and Patterson \u003cstrong\u003e(\u003c/strong\u003e1981\u003cstrong\u003e)\u003c/strong\u003e. Roman numerals correspond to the numbering used by Stahlecker for the Chiquimil section and Arabic numerals for the Puerta de Corral Quemado section. The sedimentation rate ages was taken from Esteban et al (2014); 6,88 Ma date from Spagnuolo et al (2015);\u003cstrong\u003e \u003c/strong\u003elithostratigraphic assignments on the basis of Marshall y Patterson (1981), Muruaga (2001a), Hynek et al (2012), Georgieff et al (2017), Bonini et al (2020).\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/77c78e0504bd01deb6f39725.png"},{"id":50548475,"identity":"d0e41c4e-aa51-4fd1-b3f4-9b9d6cfe6d85","added_by":"auto","created_at":"2024-02-02 10:03:03","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1128723,"visible":true,"origin":"","legend":"\u003cp\u003eVisual representation of semilandmarks: a, Graphic representation of the green semilandmarks; b, Graphic representation of the orange semilandmarks; c, Graphic representation of the blue semilandmarks; d, Landmarks of the mandible. 1, most anterior part of the mandible; 2, most anterior part of the cheek teeth at the alveolar level; 3, crossing point between the ascending and the horizontal rami of the mandible; 4-9, green semilandmarks; 10, upper extreme of the coronoid process; 11-16, blue semilandmarks; 17, anterior point of the condylar apophysis; 18, highest midpoint of the condylar apophysis; 19, posterior point of the condylar apophysis; 20-28, orange semilandmarks; e, Landmarks with links.\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/4bfd4a8665e674b8972a9e91.png"},{"id":50548485,"identity":"5faf3089-0bf9-4872-857a-eb8d91358856","added_by":"auto","created_at":"2024-02-02 10:03:04","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":29263220,"visible":true,"origin":"","legend":"\u003cp\u003eSkulls of \u003cem\u003eStromaphorus ameghini\u003c/em\u003e MLP-PV 29-X-8-1, referred to “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e” by Cabrera (1944): a, lateral view (inverted); b, frontal view; c, dorsal view; d, occlusal view. FMNH-P14396, referred to \u003cem\u003eStromaphorus? \u003c/em\u003esp. by Marshall and Patterson (1981): e, lateral view; f, frontal view; g, dorsal view; h, occlusal view. MLP-PV 29-X-10-1, referred to \u003cem\u003ePhlyctaenopyga ameghini\u003c/em\u003e by Cabrera (1944): i, lateral view; j, frontal view; k, dorsal view; l, occlusal view. Grey striped areas indicate artificially reconstructed parts. Scale bar 100mm.\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/8eafcabfe58948ef493a2a7e.png"},{"id":50548720,"identity":"7ab41e3c-6ea0-4a9c-9cd6-e2e58c7bf58e","added_by":"auto","created_at":"2024-02-02 10:11:03","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":24556170,"visible":true,"origin":"","legend":"\u003cp\u003eMandible of \u003cem\u003eStromaphorus ameghini\u003c/em\u003e; ap, angular process; ms, mandibular symphysis. a, MLP-PV 16-138, left hemimandible, Holotype of \u003cem\u003eNeuyurus compressidens\u003c/em\u003e (\u003cem\u003eStromaphorus compressidens\u003c/em\u003e): 1, Labial lateral view; 2, Lingual lateral view; 3, Oclussal view. b, MLP-PV 29-X-8-1, left hemimandible: 1, Labial lateral view; 2, Lingual lateral view; 3, Occlusal view. c, MLP-PV 29-X-8-1, right hemimandible: 1, Labial lateral view; 2, Lingual lateral view; 3, Oclussal view. Scale bar 100mm.\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/96baa16ae07da34cf980fc0a.png"},{"id":50548487,"identity":"142fe9be-8e57-4281-8a6c-9b327484df3d","added_by":"auto","created_at":"2024-02-02 10:03:04","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":40090369,"visible":true,"origin":"","legend":"\u003cp\u003eCarapace and osteoderms of \u003cem\u003eStromaphorus ameghini\u003c/em\u003eFMNH-P14439: a, General carapace morphology in lateral view; 1, Upper anterior region; 2, Upper middle region; 3, Upper posterior region; 4, Lower anterior region; 5, Lower middle region; 6, Lower posterior region. Arrow points to the anterior region. b, Holotype of \u003cem\u003ePlohophorus ameghini \u003c/em\u003e(MLP-PV 16-101); 1, detail of some osteoderms from the holotype. Scale bar 100mm.\u003c/p\u003e","description":"","filename":"Fig5.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/203797f1bfcb8ddbf7899a30.png"},{"id":50548721,"identity":"9eea7803-934b-4304-9caf-621f976d6cba","added_by":"auto","created_at":"2024-02-02 10:11:04","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":19030905,"visible":true,"origin":"","legend":"\u003cp\u003eDetail of posterior region osteoderms of \u003cem\u003eStromaphorus ameghini \u003c/em\u003e. a, FMNH-P14439; b, MLP-PV 29-X-10-2. Scale bar 100mm.\u003c/p\u003e","description":"","filename":"Fig6.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/ef32365daafc8431ada69cc3.png"},{"id":50548488,"identity":"6a8482da-de63-411c-9c5a-fe8a49d07f81","added_by":"auto","created_at":"2024-02-02 10:03:04","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":24156677,"visible":true,"origin":"","legend":"\u003cp\u003eCaudal tube of \u003cem\u003eStromaphorus ameghini\u003c/em\u003e MLP-PV 29-X-10-1, originally referred to “\u003cem\u003ePhlyctaenopyga\u003c/em\u003e”\u003cem\u003e ameghini \u003c/em\u003eby Cabrera (1944): a, Dorsal view, Photography and diagram; b, Lateral left view, Photography and diagram. MLP-PV 29-X-8-9, originally referred to “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e” by Cabrera (1944): c, Dorsal view, Photography and diagram; d, Lateral right view (inverted) Photography and diagram. The terminal (T) and lateral figures (I, II, III, and IV) are shown in yellow. In blue the marginal figures are marked, left side (L) and right side (R). In white the central figures. In black the broken parts. In grey the parts that are not clearly visible. Scale bar 100mm.\u003c/p\u003e","description":"","filename":"Fig7.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/ddc21c957e9cac4d4e4270da.png"},{"id":50548486,"identity":"c4e7d163-bdb9-44ac-95e7-22517a62f406","added_by":"auto","created_at":"2024-02-02 10:03:04","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":8100972,"visible":true,"origin":"","legend":"\u003cp\u003eStrict consensus from two most parsimonious tree\u003cstrong\u003es\u003c/strong\u003e (MPT\u003cstrong\u003es\u003c/strong\u003e) resulting from the cladistic analysis of Glyptodontidae based on TNT parsimony analysis of 95 characters from 33 taxa (tree length 174.020 steps; Consistency Index 0.845; Retention Index 0.933). The number above each node represents Jackknife values, numbers under each node show relative Bremer support (right) and absolute Bremer support (left). Node A: Glyptodontinae; Node B: Plohophorini; Node C: Doedicurini; Node D: Neosclerocalyptini; Node E: Hoplophorini. The tree has been calibrated following the ‘equal’ criterion using the \u003cem\u003estrap\u003c/em\u003e package in \u003cem\u003eR\u003c/em\u003e (root length: 1 million-years; see main text for further details). Chronostratigraphic calibration of the phlylogenetic relationships is compared to benthic foraminifera δ\u003csup\u003e18\u003c/sup\u003eO values from Westerhold et al. (2020) to visualize the evolution of the Plohophorini clade in the context of paleoenvironment and climate changes throughout the Neogene in north-western Argentina. Abbreviations: LOW, Late Oligocene Warming; MMCO, Middle Miocene Climatic Optimun; LMC, Late Miocene Cooling; PCO, Pliocene Climatic Optimun; Exp. C4 plants, expansion of C4 plants; VQB, Villavil-Quillay Basin.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e","description":"","filename":"Fig8.png","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/a12ee1b81e0206a435ac9cf9.png"},{"id":63073566,"identity":"a5bfba64-ea81-44c2-979f-987b2ee89444","added_by":"auto","created_at":"2024-08-22 20:25:03","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":293530794,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/d172162e-0918-4a14-bd77-f92ce99bd03a.pdf"},{"id":50548477,"identity":"fe836d15-35a9-45a3-880f-b34979cf520e","added_by":"auto","created_at":"2024-02-02 10:03:03","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":16253,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource1.docx","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/3a68dcc3d65a0199e96719b5.docx"},{"id":50548717,"identity":"cdfdc58a-3ab5-4f69-9a82-cdd2347f84d3","added_by":"auto","created_at":"2024-02-02 10:11:03","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":22463,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource2.docx","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/3c7da35332b98f10822a692b.docx"},{"id":50548718,"identity":"486ff52d-d302-49b5-b01c-f41ac53867ca","added_by":"auto","created_at":"2024-02-02 10:11:03","extension":"docx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":24670,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource3.docx","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/9ab0b08844de96047fc37087.docx"},{"id":50549259,"identity":"10971cc1-ff63-4e44-b7f9-ea366627b20e","added_by":"auto","created_at":"2024-02-02 10:19:03","extension":"docx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":20529,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource4.docx","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/49366219280fc29a052e41b7.docx"},{"id":50548483,"identity":"19876166-9d2d-4fac-a2c0-650cedd119bd","added_by":"auto","created_at":"2024-02-02 10:03:04","extension":"docx","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":79760,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource5.docx","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/3e09c9ffba461ff879dcfee6.docx"},{"id":50548480,"identity":"b9878c9c-0a6d-44a6-8a82-68e661297d6d","added_by":"auto","created_at":"2024-02-02 10:03:03","extension":"docx","order_by":6,"title":"","display":"","copyAsset":false,"role":"supplement","size":27279,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource6.docx","url":"https://assets-eu.researchsquare.com/files/rs-3914918/v1/f7753d967c09f4a71adc26e2.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"\u003cp\u003ePlohophorini Glyptodonts (Xenarthra, Cingulata) From the Late Neogene of Northwestern Argentina. Insight Into Their Diversity, Evolutionary History, and Paleobiogeography\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eXenarthra (Pilosa and Cingulata) is traditionally regarded as one of the main clades within Placentalia (O\u0026rsquo;Leary et al. 2013); however, they can be considered today as a relictual clade, compared to the high diversity of the fossil record along Cenozoic (McKenna and Bell \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Vizca\u0026iacute;no and Bargo \u003cspan citationid=\"CR126\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Within this remarkable diversity, one of the most enigmatic clades is represented by Glyptodontidae (Late Eocene-Late Pleistocene), composed of medium (ca. 60\u0026ndash;260 kg) to giant (ca. 2300 kg) armored herbivores (Vizca\u0026iacute;no et al. \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Soibelzon et al. \u003cspan citationid=\"CR116\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Zamorano et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and phylogenetically linked to another fossil enigmatic clade, Pampatheriidae (see Gaudin and Wible \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Gaudin and Lyon \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Fernicola et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). More recently, another proposal based on molecular evidence suggests that glyptodonts are in fact a subfamily (Glyptodontinae) within the \"armadillos\" Chlamyphoridae (Delsuc et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Mitchell et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). In this contribution we focus exclusively on morphological aspects, and therefore we use the traditional classification, considering Glyptodontidae Gray, 1869; Pampatheriidae Paula Couto, 1954 and Dasypodidae Gray, 1821 as families at the same taxonomic level. This is also consistent with most recent contributions (e.g., N\u0026uacute;\u0026ntilde;ez-Blasco et al. 2021, 2022; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Cuadrelli et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2023a\u003c/span\u003e, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003eb\u003c/span\u003e; Christen et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) using Glyptodontidae at the family level.\u003c/p\u003e \u003cp\u003eIn this framework, the analysis of the evolutionary history of glyptodonts since the Early and Middle Miocene (a moment when materials are enough informative; see Gaudin and Croft \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) suggests that two large clades with different characteristics can be recognized, one of northern origin called Glyptodontinae, and other of southern origin (\u0026ldquo;austral clade\u0026rdquo;) that we propose here to name Hoplophorinae Huxley, 1864 (see, among others, Cuadrelli et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Nu\u0026ntilde;ez-Blasco et al. 2021a, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e). While Glyptodontinae dispersed to Central and North America during the Great American Biotic Interchange (GABI) (Carlini and Zurita \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Zurita et al. \u003cspan citationid=\"CR138\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Gillette et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), Hoplophorinae, despite being more diverse, had a more restricted geographical distribution, limited mainly to high and middle latitudes of South America (N\u0026uacute;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e). Within Hoplophorinae, it is essential to analyze the evolution of glyptodonts in southern South America since the Late Miocene (ca. 9 Ma; see Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e) to understand their diversity and evolutionary history in the latest Miocene and Plio-Pleistocene, when terminal species became extinct in the latest Pleistocene together with the remaining megafauna (Prates and P\u0026eacute;rez \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Prates et al. \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Carlini et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Northwestern region of Argentina (NWA) is an area with a remarkable diversity of glyptodonts (see Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e; Zurita \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2007a\u003c/span\u003e,\u003cspan citationid=\"CR137\" class=\"CitationRef\"\u003eb\u003c/span\u003e; Nu\u0026ntilde;ez Blasco et al. 2021a, 2022). It contains, together with the Pampean region (PR) of Argentina, one of the most complete Late Miocene-Pleistocene continental sequences of South America (Esteban et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Bonini et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In this context, a comprehensive revision of the diversity of glyptodonts from NWA (ie, taxonomic, phylogenetic, and chronostratigraphic) is currently carrying out (see N\u0026uacute;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e, \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003ec\u003c/span\u003e). These studies, including a detailed comparison with those taxa from the PR (N\u0026uacute;\u0026ntilde;ez-Blasco et al. 2022), have provided new information highlighting that, as observed in the late Neogene of the PR (see Zurita et al. \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e, \u003cspan citationid=\"CR141\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR142\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), the diversity of glyptodonts seems to be more restricted than previously known.\u003c/p\u003e \u003cp\u003ePlohophorini Castellanos, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1932\u003c/span\u003e, is currently one of the most problematic traditional glyptodont tribes from a taxonomic, systematic, and phylogenetic standpoint. Within this tribe, the genus \u003cem\u003ePhlyctaenopyga\u003c/em\u003e contains a species endemic to the NOA, \u003cem\u003ePh. ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e), in addition to a late Neogene species in the RP, \u003cem\u003ePh. trouessarti\u003c/em\u003e (Moreno, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e). The tribe Plohophorini also encompasses \u003cem\u003eStromaphorus compressidens\u003c/em\u003e (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), another species interpreted as endemic to the NWA. Both taxa have been extensively investigated and described by numerous authors since the 19th century (e.g. Moreno \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e; Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e; Ameghino \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1888a\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1891a\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1902\u003c/span\u003e, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1904\u003c/span\u003e; Lydekker \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e1894\u003c/span\u003e; Rovereto \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e1914\u003c/span\u003e; Cabrera \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1939\u003c/span\u003e, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e; Castellanos \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1925\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1940\u003c/span\u003e; Zamorano et al. \u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Zamorano \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Tori\u0026ntilde;o and Perea \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this scenario, the last revision of the glyptodont diversity from NWA was carried out by Cabrera (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e), more than seventy years ago, without phylogenetic and stratigraphic context and without taking into account taphonomic aspects. New field works carried out during the last years at NWA plus the revision of materials from different institutions (see Materials and Methods) yielded new and more complete specimens that allow us to perform, for the first time, a comprehensive revision of this particular assemblage of glyptodonts. In addition, chrono-stratigraphic and geological framework of these fossiliferous sequences from NWA were greatly improved in the last decades (see, among others see among others Bonini \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Bonini et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Bossi y Muruaga 2009; Bossi et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1987\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Butler et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Esteban et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Georgieff et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Latorre et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Marshall y Patterson 1981; Marshall et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Muruaga \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2001a\u003c/span\u003e, \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003eb\u003c/span\u003e;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Riggs y Patterson 1939; Sasso \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Hynek et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), a situation that allowed a detailed chronostratigraphic determination of the levels that contain these faunas.\u003c/p\u003e \u003cp\u003eThe present paper is the continuation of an exhaustive review of the diversity of late Neogene glyptodonts from NWA (see N\u0026uacute;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e, \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003ec\u003c/span\u003e), based on the pioneering work of Cabrera (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e). We also perform the most comprehensive phylogenetic analysis of glyptodonts to date to evaluate the position of the accepted valid species of Plohophorini from NWA, highlighting the fact that this tribe (including species from PR and NWA of Argentina, and Uruguay) appears as a natural group in our analysis. Finally, this work complements those carried out by our team on the late Neogene glyptodont associations of the PR of Argentina (see Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) allowing a more certain evolutionary scenario for these elusive and enigmatic mammals.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e \u003cb\u003eInstitutional abbreviations\u003c/b\u003e.\u003c/p\u003e \u003cp\u003e \u003cb\u003eAMNH\u003c/b\u003e, American Museum of Natural History, New York, USA; \u003cb\u003eCC-MUFCA\u003c/b\u003e, Colección “Dr. Alfredo Castellanos”, Museo Universitario “Florentino y Carlos Ameghino”. Facultad de Ciencias Exactas, Ingeniería y Agrimensura - Universidad Nacional de Rosario. Rosario. Santa Fe, Argentina; \u003cb\u003eFACENA\u003c/b\u003e, Facultad de Ciencias Exactas y Naturales y de Agrimensura, Corrientes Capital, Argentina; \u003cb\u003eFC-CVF\u003c/b\u003e, Colección de vertebrados fósiles, Facultad de Ciencias, Universidad de la República, Montevideo, Uruguay; \u003cb\u003eFMNH-P\u003c/b\u003e, Field Museum of Natural History, Paleontological collection, Chicago, Illinois, USA; \u003cb\u003eGCF\u003c/b\u003e, Grupo Conservacionista de Fósiles, Museo Paleontologico ‘Fray Manuel de Torres,’ San Pedro, Buenos Aires, Argentina; \u003cb\u003eIGM\u003c/b\u003e, Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, México; \u003cb\u003eMACN\u003c/b\u003e, Sección Paleontología Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina; \u003cb\u003eMCA\u003c/b\u003e, Museo Municipal de Ciencias Naturales ‘Carlos Ameghino,’ Mercedes, Buenos Aires, Argentina; \u003cb\u003eMCH P\u003c/b\u003e, Sección Paleontología, Museo Arqueológico Condor Huasi, Belén, Catamarca, Argentina; \u003cb\u003eMHNC\u003c/b\u003e, Museo de Historia Natural de Cochabamba ‘Alcide d’Orbigny’, Cochabamba, Bolivia; \u003cb\u003eMLP-PV\u003c/b\u003e, División Paleontología Vertebrados, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Buenos Aires, Argentina; \u003cb\u003eMMC\u003c/b\u003e, Museo Municipal de Colonia ‘Dr. Bautista Rebuffo’, Colonia de Sacramento, Uruguay; \u003cb\u003eMMP\u003c/b\u003e, Museo Municipal de Ciencias Naturales ‘Lorenzo Scaglia’ Mar del Plata, Argentina; \u003cb\u003eMSM\u003c/b\u003e, Arizona Museum of Natural History (formerly Mesa Southwest Museum), Arizona, USA; \u003cb\u003ePVL\u003c/b\u003e, Colección de Paleozoología de la Facultad de Ciencias Naturales e Instituto ‘Miguel Lillo’, San Miguel de Tucumán, Argentina; \u003cb\u003ePVSJ\u003c/b\u003e, Instituto y Museo de Ciencias Naturales, Universidad Nacional de San Juan, San Juan (San Juan Province, Argentina); \u003cb\u003ePz-Ctes\u003c/b\u003e, Colecciones Paleontológicas de la Universidad Nacional del Nordeste “Rafael Herbst”, Corrientes, Argentina; \u003cb\u003eSGO PV\u003c/b\u003e, vertebrate paleontology collections, Museo Nacional de Historia Natural, Santiago, Chile; \u003cb\u003eUCMP\u003c/b\u003e, University of California, Museum of Paleontology, Berkeley, California, USA; \u003cb\u003eUFMG\u003c/b\u003e, Museo de la Universidade Federal de Minas Gerais, Belo Horizonte, Brazil.\u003c/p\u003e \u003cp\u003e \u003cb\u003eAnatomical abbreviations\u003c/b\u003e.\u003c/p\u003e \u003cp\u003e \u003cb\u003ebc\u003c/b\u003e, bony crest; \u003cb\u003edpm\u003c/b\u003e, descending process of the maxillae; \u003cb\u003efm\u003c/b\u003e, foramen magnum; \u003cb\u003eif\u003c/b\u003e, infraorbital foramina; \u003cb\u003eMf\u003c/b\u003e, \u003cb\u003emf\u003c/b\u003e, upper and lower molariforms; \u003cb\u003eno\u003c/b\u003e, nasal opening; \u003cb\u003eoc\u003c/b\u003e, occipital condyle; \u003cb\u003eon\u003c/b\u003e, orbital notch; \u003cb\u003esc\u003c/b\u003e, sagittal crest; \u003cb\u003eza\u003c/b\u003e, zygomatic arch.\u003c/p\u003e \u003cp\u003e \u003cb\u003eOther abbreviations\u003c/b\u003e.\u003c/p\u003e \u003cp\u003e \u003cb\u003eFAD\u003c/b\u003e, First Appearance Datum; \u003cb\u003eFOCP\u003c/b\u003e, Faldeo Occidental Cerro Pampa; \u003cb\u003eLAD\u003c/b\u003e, Last Appearance Datum; \u003cb\u003eMa\u003c/b\u003e, Megaannun, (millon years ago); \u003cb\u003ePCQ\u003c/b\u003e, Puerta de Corral Quemado; \u003cb\u003eSFN\u003c/b\u003e, San Fernando Norte; \u003cb\u003eSMV\u003c/b\u003e, Santa María Valley; \u003cb\u003eVQB\u003c/b\u003e, Villavil-Quillay Basin.\u003c/p\u003e \u003cp\u003eThe most relevant materials for this study are housed in the paleontological collection of vertebrates from the Museo de La Plata (“Cabrera collection”) and the Paleontological collection from the Field Museum of Natural History (Riggs collection), under the following acronyms: skulls, MLP-PV 29-X-8-1, MLP-PV 29-X-10-1, FMNH-P14396; mandibles, MLP-PV 16–138, MLP-PV 29-X-8-1; carapaces MLP-PV 16–101, MLP-PV 29-X-10-2, FMNH-P14439; caudal tubes, MLP-PV 29-X-8-9, MLP-PV 29-X-10-1, FMNH-P14414, FMNH-P14532; (see Online Resource 1 for the nomenclatural history of \u003cem\u003ePhlyctaenopyga ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e) and \u003cem\u003eStromaphorus compressidens\u003c/em\u003e (Moreno y Mercerat, 1891); and Online Resource 2 for a list of all materials assigned to the species \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; ex Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA comparative study was carried out to obtain a morphological characterization of the specimens, including cranial and postcranial elements. The results are shown in the anatomical descriptions. For direct comparison, specimens referred to the following genera were used: \u003cem\u003eGlyptodon\u003c/em\u003e Owen, 1839; \u003cem\u003eKelenkura\u003c/em\u003e Barasoain, Zurita, Croft, Montalvo, Contreras, Miño-Boilini and Tomassini, 2022; \u003cem\u003ePlohophorus\u003c/em\u003e Ameghino, 1887; \u003cem\u003ePseudoplohophorus\u003c/em\u003e Castellanos, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1926\u003c/span\u003e; \u003cem\u003eCranithlastus\u003c/em\u003e Arias, Alonso and Malanca, 1978; \u003cem\u003eStromaphoropsis\u003c/em\u003e Kraglievich, 1932; \u003cem\u003eEleutherocercus\u003c/em\u003e Koken, 1888; \u003cem\u003eDoedicurus\u003c/em\u003e Burmeister, 1874; \u003cem\u003eNeosclerocalyptus\u003c/em\u003e Paula Couto, 1957; \u003cem\u003eEosclerocalyptus\u003c/em\u003e Ameghino, 1919; \u003cem\u003eHoplophorus\u003c/em\u003e Lund, 1839; \u003cem\u003eNopachtus\u003c/em\u003e Ameghino, 1888; \u003cem\u003ePropanochthus\u003c/em\u003e Castellanos, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1925\u003c/span\u003e; \u003cem\u003ePanochthus\u003c/em\u003e Burmeister, 1866. Additionally, specimens from the literature were included in the analysis.\u003c/p\u003e \u003cp\u003eAll measurements referred to in the text are listed in Online Resource 3 (table 1: skull; table 2: Holotype MLP-PV 16–101; table 3: carapace, and table 4: caudal tube); they were taken through a manual \"Vernier\" caliper, with an error range of 0.5mm and in some cases using ImageJ (1.53e) software. The linear dimensions are expressed in millimeters (mm). For descriptions and comparisons of the dorsal carapace, osteoderms and caudal tube, we follow the proposal of Zamorano et al. (\u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), Porpino et al. (\u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and Toriño (\u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The numbering assigned to the figures of the ornamentation of caudal tubes (see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e) follows the proposal of Toriño (\u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), and Toriño and Perea (\u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). For chronological purposes, we follow the International Chronostratigraphic Chart v2022/02 (Cohen et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eCladistic analysis\u003c/b\u003e.\u003c/p\u003e \u003cp\u003eThe phylogenetic study is based on a mixed matrix (see Online Resource 4 and Online Resource 5), consisting of a total of 96 characters, of which 95 are conventional (descriptive morphological characters) and one of geometric morphometry, to analyze a total of 35 taxa. The descriptive morphological characters are 68 binary and 27 multistate. This includes 40 skull characters, 4 autopodium characters, 26 carapace characters and 25 caudal armor characters. The information considered in this analysis, used both for the codification of states and landmark placement for geometric morphometry, was recorded via direct observation of the specimens and from photographs taken by the authors.\u003c/p\u003e \u003cp\u003eThe taxon-character matrix was constructed with Mesquite version 3.4 (Maddison and Maddison \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and to build the geometric morphometric character, photographs were compiled into .tps files using tpsUtil software, version 1.70 (Rohlf \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), and landmarks were digitalized with tpsDig2 version 2.16 (Rohlf \u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e2010\u003c/span\u003e); this matrix was analyzed via “Traditional search“ under the criterion of maximum parsimony, using TNT version 1.5 (Goloboff and Catalano, \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), the algorithm used was TBR (10 trees to save per replication). It is worth noting that all characters were unordered and were weighted equally (1.0). Clade support was assessed via Absolute and Relative Bremer support, retaining suboptimal trees by 3 steps; see Bremer (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1994\u003c/span\u003e); Goloboff and Farris (\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). In addition, a Jackknife analysis was carried out via “Traditional search” (TBR algorithm) with 100 replicates and 36 removal probability. Finally, the consensus tree obtained from the two most parsimonious trees (MPTs) was calibrated to obtain estimated divergence times and potential ghost ranges. The calibration followed the ‘equal’ method (with a root of 1\u0026nbsp;million-years) implemented in the strap package (see Bell and Lloyd, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) for R software version 4.3.0 (R Core team \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The resulting time-scaled tree was plotted against the International Chronostratigraphic Chart (Cohen et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) for visualization purposes. The R commands included in the appendices of Bell and Lloyd (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and Toriño et al. (\u003cspan citationid=\"CR124\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) were adapted for these procedures. The input data for the analysis with R were the strict consensus tree exported in parenthetical notation from TNT as a .tre file, and a txt file including a list of taxa with their first and last record (see supplementary files). The list of biochrons for each taxon as well as the bibliographical references from which this information was taken can be found in the Online Resource 6 of the present work.\u003c/p\u003e \u003cp\u003eThe ingroup includes the following taxa: \u003cem\u003eBoreostemma venezolensis\u003c/em\u003e Simpson, 1947; \u003cem\u003eB. acostae\u003c/em\u003e (Villarroel 1983); \u003cem\u003eGlyptodon jatunkhirkhi\u003c/em\u003e Cuadrelli, Zurita, Toriño, Miño-Boilini, Perea, Luna, Gillette and Medina, 2020; \u003cem\u003eG. munizi\u003c/em\u003e Ameghino, 1881; \u003cem\u003eG. reticulatus\u003c/em\u003e Owen, 1845; \u003cem\u003eGlyptotherium texanum\u003c/em\u003e Osborn, 1903; \u003cem\u003eGl\u003c/em\u003e. \u003cem\u003ecylindricum\u003c/em\u003e (Brown, 1912); \u003cem\u003ePropalaehoplophorus australis\u003c/em\u003e Ameghino, 1887; \u003cem\u003eEucinepeltus petesatus\u003c/em\u003e Ameghino,1891; \u003cem\u003eCochlops muricatus\u003c/em\u003e Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; \u003cem\u003ePalaehoplophoroides rothi\u003c/em\u003e Scillato-Yané and Carlini, 1998; \u003cem\u003ePalaehoplophorus meridionalis\u003c/em\u003e Ameghino, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1904\u003c/span\u003e; \u003cem\u003eKelenkura castroi\u003c/em\u003e Barasoain, Zurita, Croft, Montalvo, Contreras, Miño-Boilini and Tomassini, 2022; \u003cem\u003ePlohophorus figuratus\u003c/em\u003e Ameghino, 1887; \u003cem\u003eP. avellaneda\u003c/em\u003e Quiñones, Cuadrelli, De los Reyes, Luna, Poiré and Zurita, 2023; \u003cem\u003ePseudoplohophorus absolutus\u003c/em\u003e Perea, 2005; \u003cem\u003ePs. benvenutii\u003c/em\u003e (Castellanos, 1954); \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e); \u003cem\u003eS. trouessarti\u003c/em\u003e (Moreno, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e); \u003cem\u003eEleutherocercus solidus\u003c/em\u003e (Rovereto, \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e1914\u003c/span\u003e); \u003cem\u003eE. antiquus\u003c/em\u003e (Ameghino, 1887); \u003cem\u003eDoedicurus clavicaudatus\u003c/em\u003e (Owen, 1847); \u003cem\u003eNeosclerocalyptus gouldi\u003c/em\u003e Zurita, Carlini and Scillato-Yané, 2008; \u003cem\u003eN. ornatus\u003c/em\u003e (Owen, 1845); \u003cem\u003eN. paseudornatus\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e); \u003cem\u003eN. paskoensis\u003c/em\u003e (Zurita, 2002); \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e Ameghino, 1888; \u003cem\u003eHoplophorus euphractus\u003c/em\u003e Lund, 1839; \u003cem\u003ePropanochthus bullifer\u003c/em\u003e (Burmeister, 1874); \u003cem\u003ePanochthus intermedius\u003c/em\u003e Lydekker, 1895; \u003cem\u003eP. tuberculatus\u003c/em\u003e (Owen, 1845) and \u003cem\u003eParapropalaehoplophorus septentrionalis\u003c/em\u003e Croft, Flynn and Wyss, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2007\u003c/span\u003e. The extant dasypodid \u003cem\u003eEuphractus sexcintus\u003c/em\u003e Linnaeus, 1758, and the pampathere \u003cem\u003ePampatherium humdboltii\u003c/em\u003e (Lund, 1839) were used as outgroups to root the tree.\u003c/p\u003e \u003cp\u003e \u003cb\u003eGeometric morphometrics\u003c/b\u003e.\u003c/p\u003e \u003cp\u003eFor the geometric morphometric character (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) two-dimensional coordinates of 29 landmarks were digitalized from photographs of mandibles in lateral view of sixteen glyptodonts (\u003cem\u003eEucinepeltus petesatus\u003c/em\u003e MACN 4760; \u003cem\u003ePropalaehoplophorus australis\u003c/em\u003e MLP-PV 16 − 15; \u003cem\u003ePseudoplohophorus absolutus\u003c/em\u003e FC-CVF475-595; \u003cem\u003eGlyptotherium texanum\u003c/em\u003e MSM 4818; \u003cem\u003eGl. cylindricum\u003c/em\u003e IGM 9563; \u003cem\u003eBoreostemma acostae\u003c/em\u003e UCMP 38039; \u003cem\u003eGlyptodon munizi\u003c/em\u003e GCF 10; \u003cem\u003eG. reticulatus\u003c/em\u003e MCA 2015; \u003cem\u003eDoedicurus clavicaudatus\u003c/em\u003e MACN 2762; \u003cem\u003eEleutherocercus solidus\u003c/em\u003e FMNH-P14437; \u003cem\u003ePanochthus intermedius\u003c/em\u003e MHNC 13491; \u003cem\u003eP. tuberculatus\u003c/em\u003e MLP-PV 16–29; \u003cem\u003eNeosclerocalyptus ornatus\u003c/em\u003e MLP-PV 16–28; \u003cem\u003eN. paskoensis\u003c/em\u003e MACN 18107; \u003cem\u003eN. gouldi\u003c/em\u003e MCA 2010 and \u003cem\u003eParapropalaehoplophorus septentrionalis\u003c/em\u003e SGO PV 4165), one dasypodid (\u003cem\u003eEuphractus sexcintus\u003c/em\u003e FACENA 183) and one pampatheriid (\u003cem\u003ePampatherium humboldtii\u003c/em\u003e MLP-PV 81-X-30-1); This method had previously been tested on Cingulate mandibles, with promising results in phylogenetics (see Nuñez-Blasco et al. 2021b). In this context, of the 29 landmarks, 5 are type I (Ladmarks 1, 10, 17, 18 and 19); 2 are type II (Ladmarks 2 and 3). Besides, there are 3 sets of semi-landmarks used to define the rest of the mandible morphology (green set, blue set and orange set). The green and blue sets (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e: a, c) were defined based on the 65º angle formed between landmarks 3, 19 and 10, subdividing it into 7 portions. The orange set (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e: b) was defined based on the 110º angle formed by landmarks 1, 3 and 19, subdividing it into 10 portions.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eGeographic and stratigraphic context\u003c/h2\u003e \u003cp\u003eThe study area is in the Catamarca province, Northwestern Argentina, within the Northwestern Pampean Ranges geological province (Caminos, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1979\u003c/span\u003e). The Holotype materials of “\u003cem\u003ePhlyctaenopyga\u003c/em\u003e” \u003cem\u003eameghini\u003c/em\u003e and “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e” come from the Santa María Valley (SMV), while most of the fossils assigned to these two species come from the Villavil-Quillay basin (VQB).\u003c/p\u003e \u003cp\u003eThe SMV (26°53’S, 66°05’W) is a tectonic depression of more than 100 km long by 20 to 30 km wide, flanked to the east by the Cumbres Calchaquíes and Aconquija Ranges, and to the west by the Quilmes Ranges (Bossi et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). A great part of the paleontological materials from SMV was exhumed from the Late Miocene/Pliocene levels cropping out at Andalhuala de Arriba and Tiopunco localities (Catamarca and Tucumán provinces, respectively). In turn, the VQB (27°15’ S/66°54’ W) is a tecto-sedimentary basin included in the Hualfin valley, limited to the northwest by the Sierra de Altohuasi, to the southwest by the Cerro Durazno, to the northeast by the Sierra de Hualfín, to the east by the Complejo Volcánico Farallón Negro, and the southeast by the Cerro Pampa-Belén Range (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The materials found in the VQB come from the Andalhuala, and Chiquimil formations cropping out at Puerta de Corral Quemado (PCQ), San Fernando Norte (SFN), and Faldeo Occidental del Cerro Pampa (FOCP).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe Neogene lithostratigraphic units recognized in these areas have been included in the Santa María Group, which is composed (from base to top) of Las Arcas, Chiquimil, Andalhuala, and Corral Quemado formations, ranging from \u003cem\u003eca.\u003c/em\u003e 10 to 3 Ma, being the Andalhuala Formation the most extended in both areas (Bossi et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Muruaga \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2001a\u003c/span\u003e, \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003eb\u003c/span\u003e; Bossi and Muruaga \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Bonini et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The Andalhuala Formation in the VQB was deposited between ca. 7.14 to 3.66 Ma (Latorre et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), corresponding to the fossiliferous beds that yielded the studied materials (Riggs and Patterson \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e1939\u003c/span\u003e; Marshall and Patterson \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e). The historical specimens collected in the SMV lack provenance data, except for the reference of “\u003cem\u003eEstratos Araucanos\u003c/em\u003e” (\u003cem\u003esensu\u003c/em\u003e Rovereto \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e1914\u003c/span\u003e) and “\u003cem\u003eAraucanense medio\u003c/em\u003e” (\u003cem\u003esensu\u003c/em\u003e Castellanos \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1948a\u003c/span\u003e, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1969\u003c/span\u003e), mainly assigned to the upper section of Chiquimil and Andalhuala formations in the modern lithostratigraphic framework. Lithologically, the Andalhuala Formation in SMV is broadly characterized by sandstones and conglomerates poorly sorted, polymict and matrix-supported clasts, with scarce participation of pelites (Georgieff \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). In both areas, SMV and VQB, this unit is composed of facies of brownish, reddish, and greyish sandstones with tabular, lenticular, trough cross-stratification, ripple marks, and cuneiform cross-stratification, often interbedded with tabular massive siltstones and several tuff beds. Moreover, the Andalhuala Formation shows several secondary features that evidence the subaerial exposure and water table fluctuations, which agree with the inferred permanent fluvial subenvironment passing to an eolian system with dunes (Bonini et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough the literature mentions the same stratigraphic formations in VQB and SMV, it should be noted that the units that form the Santa María Group sometimes show important diachronies between those cropping out in both basins (see Spagnuolo et al. \u003cspan citationid=\"CR117\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, \u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e2013\u003c/span\u003e, \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Georgieff et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2012a\u003c/span\u003e, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Georgieff and Díaz \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Bonini \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Bonini et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In general, the inferred palaeoenvironments are shallow lacustrine (but also continental sabkha in SMV; Ibañez, 2001; Esteban et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and sandy-gravel braided fluvial for the Chiquimil and Andalhuala formations at both sites, but the dating of the tuffaceous strata reveals that their deposition were older in VQB. A clear example is the Chiquimil Formation (see Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e), which presents an age of 7.14 Ma for its top at East of PCQ, while in the SMV an age of ca. 6.88 was obtained from the top of the underlying unit, Las Arcas Formation (Georgieff et al. 2014; \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e); this shows that the deposition of the Chiquimil Formation at SMV was younger, and therefore the lithostratigraphic correlation between both areas cannot be established by that criteria. Likewise, the determination of the ages of the specimens analyzed in the present work has been carried out taking into account the absolute ages (geochronology) and not the formation (lithostratigraphy) from which they originate.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eSystematic Paleontology\u003c/h2\u003e \u003cp\u003eSuperorder \u003cb\u003eXenarthra\u003c/b\u003e Cope, 1889\u003c/p\u003e \u003cp\u003eOrder \u003cb\u003eCingulata\u003c/b\u003e Illiger, 1811\u003c/p\u003e \u003cp\u003eFamily \u003cb\u003eGlyptodontidae\u003c/b\u003e Gray, 1869\u003c/p\u003e \u003cp\u003eTribe \u003cb\u003ePlohophorini\u003c/b\u003e Castellanos, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1932\u003c/span\u003e (nom. transl. Hoffstetter, \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1958\u003c/span\u003e)\u003c/p\u003e \u003cp\u003eGenus \u003cb\u003eStromaphorus\u003c/b\u003e Castellanos, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1925\u003c/span\u003e\u003c/p\u003e \u003cp\u003e(= \u003cem\u003ePhlyctaenopyg\u003c/em\u003ea Cabrera, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e new synonymy)\u003c/p\u003e \u003cp\u003e \u003cb\u003eSpecies\u003c/b\u003e: \u003cb\u003eStromaphorus ameghini\u003c/b\u003e \u003cb\u003e(\u003c/b\u003eAmeghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; \u003cb\u003eex\u003c/b\u003e Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e\u003cb\u003e)\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eOther species\u003c/strong\u003e \u003c/p\u003e\u003cp\u003e \u003cem\u003eStromaphorus trouessarti\u003c/em\u003e (Moreno, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e) \u003cem\u003enov. comb.\u003c/em\u003e\u003c/p\u003e \u003cp\u003e\u003c/p\u003e \u003cp\u003e \u003cb\u003eStromaphorus ameghini\u003c/b\u003e \u003cb\u003e(\u003c/b\u003eAmeghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; \u003cb\u003eex\u003c/b\u003e Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e\u003cb\u003e)\u003c/b\u003e\u003c/p\u003e \u003cp\u003e= \u003cem\u003eHoplophorus ameghinii\u003c/em\u003e Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e, p. 120 [nom. nud.]; \u003cem\u003ePlohophorus ameghini\u003c/em\u003e Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e, p. 825, plates LXIX, Figs.\u0026nbsp;19 and 20 and LXXXII, Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e and \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e; \u003cem\u003eNeuryurus compressidens\u003c/em\u003e Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e, p. 224; \u003cem\u003ePlohophorus philippii\u003c/em\u003e Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e, p. 225; \u003cem\u003ePlohophorus ameghinii\u003c/em\u003e Ameghino, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1902\u003c/span\u003e, p. 2; \u003cem\u003ePlohophorus ameghinoi\u003c/em\u003e Ameghino, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1904\u003c/span\u003e, p. 288; \u003cem\u003eStromaphorus ameghinoi\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1904\u003c/span\u003e) Castellanos, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1925\u003c/span\u003e, p. 96 (not 1940, p. 27) [n. comb.]; \u003cem\u003eStromaphorus philippii\u003c/em\u003e (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), Cabrera \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1939\u003c/span\u003e [n. comb.]; \u003cem\u003eStromaphorus ameghinoi\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1904\u003c/span\u003e) Castellanos, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1940\u003c/span\u003e, p. 27 (not 1925, p. 96); \u003cem\u003eUrotherium compressidens\u003c/em\u003e Castellanos \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1940\u003c/span\u003e, p. 273; \u003cem\u003eStromaphorus compressidens\u003c/em\u003e (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e, p. 30 [n. comb.]. \u003cem\u003ePhlyctaenopyga ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e), Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e, p. 42 [n. gen, n. comb.].\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHolotype\u003c/strong\u003e \u003c/p\u003e\u003cp\u003eMLP-PV 16–101, small fragment of carapace and some isolated osteoderms. Santa María Department, Catamarca Province, Argentina.\u003c/p\u003e \u003cp\u003e\u003c/p\u003e \u003cp\u003e \u003cb\u003eGeographic and Stratigraphic Occurrence\u003c/b\u003e: Catamarca Province: Santa María Valley, Puerta de Corral Quemado, Corral Quemado and San Fernando; Chiquimil Formation, Andalhuala Formation and Corral Quemado Formation, Late Miocene-Pliocene (Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e; Marshall and Patterson \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Bonini \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zamorano et al. \u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Tucumán Province: Tiopunco (Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e; Zamorano et al. \u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). La Rioja Province: El Degolladito; Salicas Formation, Late Miocene (Brandoni and González-Ruiz 2020). Córdoba Province: Arroyo Los Chiflones, Villa Cura Brochero; Brochero Formation, Miocene-Pliocene (Cruz \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eMaterials analysed in this work, geographic and stratigraphic provenance and age: FMNH-P14396\u003c/b\u003e, deformed skull, Ampajango, Catamarca Province, Argentina; Chiquimil Formation, Stratigraphic level XI from Stahlecker in Marshall and Patterson (\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e, Appendix II) between 6.88 to 6.02 Ma, Messinian (Late Miocene); \u003cb\u003eFMNH-P14414\u003c/b\u003e, incomplete and deformed skull, dorsal carapace fragment, femur, two caudal rings, distal portion of caudal tube, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, Stratigraphic level 17 from Marshall and Patterson (\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e), age ca. 6.9 to 6.3Ma, Messinian (Late Miocene); \u003cb\u003eFMNH-P14439\u003c/b\u003e, complete carapace, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, Stratigraphic level 26 from Marshall y Patterson (1981); age ca. 4.9 to 4.4 Ma, Zanclean (Early Pliocene); \u003cb\u003eFMNH-P14532\u003c/b\u003e, proximal portion of caudal tube, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; Andalhuala Formation/Corral Quemado Formation, Stratigraphic levels 15–32 from Marshall and Patterson \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; age ca. 7.14 to 3.66 Ma, Miocene-Pliocene; \u003cb\u003eFMNH-P15771\u003c/b\u003e, small fragments of carapace, some from the anterior region and some from the posterior region, Tio Punco, Tafi, Tucumán Province, Argentina; unknown stratigraphic level, Pliocene, [originally classified as Glyptodontidae indet. in Marshall and Patterson \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e]; \u003cb\u003eMCH-P39\u003c/b\u003e, small fragment of carapace, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma, Zanclean (Early Pliocene); \u003cb\u003eMCH-P174\u003c/b\u003e, isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma, Zanclean (Early Pliocene). New studied specimen; \u003cb\u003eMCH-P176\u003c/b\u003e, isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma, Zanclean stage, Pliocene. New studied specimen; \u003cb\u003eMCH-P177\u003c/b\u003e, isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately above the tuff dated in ca. 4.72 Ma., Zanclean (Early Pliocene). New studied specimen; \u003cb\u003eMCH-P247\u003c/b\u003e, isolated osteoderm, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, age ca. 4.72 to 4.79 Ma, Zanclean (Early Pliocene). New studied specimen; \u003cb\u003eMCH-P323\u003c/b\u003e, isolated osteoderm, Faldeo Occidental Cerro Pampa (San Fernando Sur), Belén Department, Catamarca Province, Argentina; Andalhuala Formation, immediately below the tuff dated in 5.59 Ma, Messinian (Late Miocene). New studied specimen; \u003cb\u003eMCH-P326\u003c/b\u003e, isolated osteoderms, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, age ca. 3.6 to 4.72 Ma, Zanclean (Early Pliocene). New studied specimen; \u003cb\u003eMCH-P328\u003c/b\u003e, isolated osteoderms, fragments of carapace, fragments of caudal rings and fragments of caudal tube, San Fernando Norte, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, age ca. 3.6 to 4.72 Ma, Zanclean (Early Pliocene). New studied specimen; \u003cb\u003eMCH-P364\u003c/b\u003e, isolated osteoderm, Western Slopes of Cerro Pampa. Chiquimil Formation, age more than 7.14 Ma; Messinian-Tortonian (Late Miocene). New studied specimen; \u003cb\u003eMCH-P365\u003c/b\u003e, 21 isolated osteoderms, Western Slopes of Cerro Pampa, Belén Department, Catamarca Province, Argentina; Andalhuala Formation, above tuff dated in 5.59 Ma, Miocene-Pliocene. New studied specimen; \u003cb\u003eMLP-PV 16–138\u003c/b\u003e, incomplete and deformed left horizontal ramus, holotype of “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e” (\u003cem\u003eNeuryurus compressidens\u003c/em\u003e Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), Bajo de Andalhualá, Santa María Department, Catamarca Province, Argentina; Corral Quemado Formation; \u003cb\u003eMLP-PV 29-X-8-1\u003c/b\u003e, laterally deformed skull, mandible and carapace fragments; referred to \u003cem\u003e“Stromaphorus compressidens”\u003c/em\u003e by Cabrera (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e). Barranca del Palito Parado, campo del Jarillar, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; “Araucaniano”; \u003cb\u003eMLP-PV 29-X-8-9\u003c/b\u003e, caudal tube; referred to \u003cem\u003e“Stromaphorus compressidens”\u003c/em\u003e by Cabrera (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e). Campo de los Cálibas, Puerta de Corral Quemado, Belén Department, Catamarca Province, Argentina; “Araucaniano”; \u003cb\u003eMLP-PV 29-X-10-1\u003c/b\u003e, incomplete skull, cephalic shield, carapace fragments, parts of the sacral and first caudal vertebrae, caudal tube and parts of the caudal rings; referred to “\u003cem\u003ePhlyctaenopyga\u003c/em\u003e” \u003cem\u003eameghini\u003c/em\u003e by Cabrera (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e). Loma de la Greda, San Fernando, Belén Department, Catamarca Province, Argentina; “Araucaniano”; \u003cb\u003eMLP-PV 29-X-10-2\u003c/b\u003e, carapace deformed by pressure, parts of the first caudal ring, tube and pelvis; referred to “\u003cem\u003ePhlyctaenopyga\u003c/em\u003e” \u003cem\u003eameghini\u003c/em\u003e by Cabrera (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e). Loma de la Greda, San Fernando, Belén Department, Catamarca Province, Argentina; “Araucaniano”. See Online Resource 2 for a complete list of materials historically referring to “\u003cem\u003ePhlyctaenopyga\u003c/em\u003e” \u003cem\u003eameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e) and “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e” (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), and it is reassignment to \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; ex Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eEmended diagnosis\u003c/b\u003e: Medium sized glyptodont (see Online Resource 3), larger than \u003cem\u003ePseudoplohophoru\u003c/em\u003es but smaller than the Pleistocene genera \u003cem\u003ePanochthus, Glyptodon\u003c/em\u003e, and \u003cem\u003eDoedicurus\u003c/em\u003e. Lateral skull profile similar to \u003cem\u003ePlohophorus\u003c/em\u003e and \u003cem\u003ePseudoplohophorus\u003c/em\u003e. Nasal region dorsoventrally reduced; nasofrontal area inclined 152º anteroventrally in front of the orbital notches; subelliptical orbital notch, with the main axis inclined 110° counterclockwise to the dorsoventral axis; inverted subtrapezoidal nasal aperture with both lateral margins convex; subtriangular rostral area in front of the orbital notches; without postorbital bar; zygomatic arches more elongated anteroposteriorly than those of other Plohophorini (eg., \u003cem\u003ePlohophorus figuratus\u003c/em\u003e); palate general morphology straight, as in \u003cem\u003ePlohophorus\u003c/em\u003e and different from \u003cem\u003ePseudoplohophorus\u003c/em\u003e; Mf1 and Mf2 with triangular morphology, Mf4-8 with very marked trilobulation, Mf4 coincides with the infraorbital foramen. The dorsal carapace is sub-rectangular with variation in the ornamentation depending on the sector. Anterior portion: rounded and flat central figure, surrounded by a single row of 10–12 small rounded peripheral figures. Middle portion: subcircular and slightly convex central figure, surrounded by a first row of 11 to 15 peripheral figures and a second row which is always incomplete; it differs clearly from \u003cem\u003eNopachtus\u003c/em\u003e, \u003cem\u003ePanochthus\u003c/em\u003e and \u003cem\u003ePropanochthus\u003c/em\u003e in which the second row is always complete. Postero-dorsal region (pelvic region): round and very convex central figure, surrounded by a first row of 12–21 peripheral figures and a second one, complete, between 21–27; this is the only area of the carapace in which the second row of peripheral figures is complete. Caudal margin with large convex central figures preceded anteriorly by 3–5 rows of peripheral figures, one row on the sides of the central figure and a free border on the posterior margin. In dorsal and ventral view, the caudal tube has parallel lateral borders along its entire length instead of tapering distally, as the caudal tubes of some genera (e.g. \u003cem\u003ePseudoplohophorus\u003c/em\u003e and \u003cem\u003ePlohophorus\u003c/em\u003e) and almost circular in transverse outline. Simple rosette pattern, with a smooth-surfaced central figure surrounded by a single complete row of shared peripheral figures. At the apex there are two large smooth terminal figures (T) which, in the more distal portion, are in contact. It has four lateral figures (numbered I, II, III and IV, see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e) that reduce in size towards the proximal portion. Figure I is characteristically convex, unlike figures II, III and IV, which are flatter. Between the lateral and marginal figures there is a row of small peripheral figures.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003cdiv id=\"Sec6\" class=\"Section3\"\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e "},{"header":"Description and comparisons","content":"\u003ch2\u003eCranium\u003c/h2\u003e\u003cp\u003eIn lateral view (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e: a, MLP-PV 29-X-8-1; e, FMNH-P14396 and i, MLP-PV 29-X-10-1) the dorsal profile of the skull is similar to \u003cem\u003ePlohophorus figuratus\u003c/em\u003e (MLP-PV 16–153), \u003cem\u003ePseudoplohophorus absolutus\u003c/em\u003e (FC-CVF 475\u003cb\u003e/\u003c/b\u003e595) and \u003cem\u003ePs. benvenutii\u003c/em\u003e (CC-MUFCA 1388). The naso-frontal area has an anteroventral tilt relative to the parieto-occipital area, forming an angle of 152º situated in front of the orbital notches, in contrast to \u003cem\u003eDoedicurus\u003c/em\u003e, \u003cem\u003eEleutherocercus\u003c/em\u003e and \u003cem\u003ePanochthus\u003c/em\u003e, in which both areas form a slightly tighter angle of ca. 140º, located posterior to or at the level of the orbital notch. The nasal region is dorsoventrally reduced (for measurements see Online Resource 3, table 1), a common feature among the Plohophorini, and different from the rest of the Glyptodontidae. The orbital notch is subelliptic, with the main axis inclined 110º in an anti-clockwise direction relative to the dorsoventral axis. In the ventral margin of the orbital notch a subtle bony crest is observed, similar to \u003cem\u003eP. figuratus, Ps. absolutus\u003c/em\u003e, \u003cem\u003ePs. benvenutii\u003c/em\u003e and \u003cem\u003eEosclerocalyptus proximus\u003c/em\u003e (CC-MUFCA 703), but less pronounced than in \u003cem\u003eEleutherocercus\u003c/em\u003e and \u003cem\u003eNeosclerocalyptu\u003c/em\u003es. Like most glyptodonts, \u003cem\u003eStromaphorus ameghini\u003c/em\u003e lacks a post-orbital bar, unlike \u003cem\u003eEleutherocercus solidus\u003c/em\u003e (FMNH-P14437), \u003cem\u003eE. antiquus\u003c/em\u003e (MACN 2894) and \u003cem\u003ePanochthus tuberculatus\u003c/em\u003e (MLP-PV 16–29), in which it is present and well developed. On the other hand, the dorsal margin of the zygomatic arch is slightly concave, unlike \u003cem\u003eP. figuratus\u003c/em\u003e, in which this concavity is greater and \u003cem\u003ePs. absolutus\u003c/em\u003e and \u003cem\u003ePs. benvenutii\u003c/em\u003e, which has a rather straight margin. In turn, the ventral margin of the zygomatic arch is straight and descends directly to join the descending process. In general terms, this whole structure is rather robust.\u003c/p\u003e\u003cp\u003eIn frontal view, (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e: b, MLP-PV 29-X-8-1; f, FMNH-P14396 and j, MLP-PV 29-X-10-1) the nasal opening is sub-trapezoidal inverted, with its dorsal margin wider than the palate, and with both lateral margins convex, as in \u003cem\u003eP. figuratus\u003c/em\u003e, \u003cem\u003ePs. absolutus\u003c/em\u003e, \u003cem\u003ePs. benvenutii\u003c/em\u003e, \u003cem\u003eKelenkura castroi\u003c/em\u003e (PVSJ-366), and \u003cem\u003eEo. proximus\u003c/em\u003e and bears a certain resemblance to \u003cem\u003eE. antiquus\u003c/em\u003e. The descending processes are broken in all specimens. However, the infraorbital foramina are preserved; they are subcircular and small, different from \u003cem\u003eGlyptodon munizi\u003c/em\u003e (GCF 10) and \u003cem\u003eG. reticulatus\u003c/em\u003e (MCA 2015), in which they are circular and larger (see Zurita et al. \u003cspan citationid=\"CR139\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Cuadrelli et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) (for measurements see Online Resource 3, table 1).\u003c/p\u003e\u003cp\u003eIn dorsal view, (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e: c, MLP-PV 29-X-8-1, g, FMNH-P14396 and k, MLP-PV 29-X-10-1), the general morphology of the skull is very similar to \u003cem\u003eP. figuratus\u003c/em\u003e and to a lesser degree to \u003cem\u003ePs. absolutus\u003c/em\u003e and \u003cem\u003ePs. benvenutii\u003c/em\u003e. The specimen MLP-PV 29-X-10-1 is largely reconstructed, especially in the whole area of the cranial vault; therefore, it is not reliable to describe the morphology of this region in this fossil. The rostral area in front of the orbital notches is subtriangular in outline, as is that of \u003cem\u003eP. figuratus, Ps. absolutus\u003c/em\u003e, and \u003cem\u003ePs. benvenutii\u003c/em\u003e, but different from \u003cem\u003eE. antiquus, E. solidus, Neosclerocalyptus paskoensis\u003c/em\u003e (MACN-Pv 18107), \u003cem\u003eN. ornatus\u003c/em\u003e (MLP-PV 16–18), \u003cem\u003eGlyptodon munizi\u003c/em\u003e and \u003cem\u003eG. reticulatus\u003c/em\u003e in which the morphology is sub-rectangular. The orbital notches are open posteriorly due to the lack of post-orbital bar, while the maximum diameter of the fronto-parietal region coincides with this region; further back, there is a conspicuous post-orbital narrowing, also found in other species, such as \u003cem\u003eP. figuratus\u003c/em\u003e, \u003cem\u003eEo. proximus\u003c/em\u003e, \u003cem\u003eD. clavicaudatus\u003c/em\u003e or \u003cem\u003eP. tuberculatus\u003c/em\u003e. The zygomatic arches are more elongated antero-posteriorly than in the other Plohophorini but not as much as \u003cem\u003eK. castroi\u003c/em\u003e; in turn, these more rounded forms differ from the quadrangular forms of \u003cem\u003eE. solidus\u003c/em\u003e and \u003cem\u003eE. antiquus\u003c/em\u003e. In all three specimens it can be observed a short and poorly developed sagittal crest, that starts at the nuchal crests and extends up to the contact with the fronto-parietal suture (for measurements see Online Resource 3, table 1).\u003c/p\u003e\u003cp\u003eIn occlusal view, (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e: d, MLP-PV 29-X-8-1, h, FMNH-P14396 and l, MLP-PV 29-X-10-1), the general morphology of the palate is quite straight, like \u003cem\u003eP. figuratus\u003c/em\u003e, and different from \u003cem\u003ePs. benvenutii\u003c/em\u003e and \u003cem\u003ePs. absolutus\u003c/em\u003e, which shows a relatively marked expansion in the posterior region of the palate (for measurements see Online Resource 3, table 1). One of the most remarkable features of this view is the morphology of the first molariforms. FMNH-P14396 is the only specimen in which the molariforms Mf1 and Mf2 are clearly visible, and they are triangular; in \u003cem\u003eP. figuratus\u003c/em\u003e, instead, they are markedly circular. The Mf3 is not preserved. The Mf4 has a markedly trilobed morphology. From Mf4 onwards all molariforms (Mf4-8) have very pronounced trilobulation. It should be noted that Mf8 is small compared to the preceding molariforms. The main axes of Mf4-Mf8 are parallel to the longitudinal axis of the dental series, a characteristic shared with \u003cem\u003eP. figuratus\u003c/em\u003e. In this view the Mf4 coincides with the infraorbital foramen; this situation differs from \u003cem\u003eP. figuratus\u003c/em\u003e, \u003cem\u003ePs. absolutus\u003c/em\u003e and \u003cem\u003ePs. benvenutii\u003c/em\u003e in which the infraorbital foramen coincides with the Mf3.\u003c/p\u003e\u003ch2\u003eMandible\u003c/h2\u003e\u003cp\u003eDescriptions are mainly based on MLP-PV 16–138 (holotype of \u003cem\u003eNeuryurus compressidens=\u003c/em\u003e “\u003cem\u003eStromaphorus compressidens\u003c/em\u003e”) which includes incomplete left horizontal ramus, and MLP-PV 29-X-8-1, represented by two hemi-mandibles (left and right), Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e. In lateral view, the mandible looks quite gracile compared to those of other groups such as \u003cem\u003eDoedicurus\u003c/em\u003e, \u003cem\u003eEleutherocercus, Neosclerocalyptu\u003c/em\u003es and \u003cem\u003ePanochthus\u003c/em\u003e. The antero-posterior diameter of the ascending ramus is shorter than the total length of the tooth series from mf1 to mf6. Like in \u003cem\u003ePs. absolutus\u003c/em\u003e, the ascending ramus is inclined forwards, drawing an angle of 70° between its anterior margin and the alveolar margin; this situation differs from the opening observed in \u003cem\u003eNeosclerocalyptus\u003c/em\u003e, \u003cem\u003eDoedicurus\u003c/em\u003e, \u003cem\u003eEleutherocercus\u003c/em\u003e, and \u003cem\u003eGlyptodon\u003c/em\u003e, in which the angle is narrower of 65º. At the posterior border of the ascending ramus, the posterior-lower margin shows a rounded angular process, somewhat less accentuated than in \u003cem\u003ePs. absolutus.\u003c/em\u003e The horizontal ramus of the mandibular body reaches the maximum height at the level of the mf5 and its ventral margin is slightly convex, as in \u003cem\u003ePs. absolutus\u003c/em\u003e and \u003cem\u003eEo. proximus\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eMolariforms. The mf1 is absent in all specimens, only molariforms mf2-mf8 are preserved. The mf2 is broken at the lingual posterior margin, but its morphology is triangular or kidney-shaped with the posterior portion narrowed laterally, this morphology is very different from the rest of the Glyptodontidae, and is repeated in the first molariforms of the upper series of the skulls MLP-PV 29-X-8-1 and FMNH-P14396 (See Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e: d, h). The pattern of mf3 tends to trilobation, with a greater development of the lobules on the labial margin; while mf4-mf8 are clearly trilobated, very similar to \u003cem\u003ePs. absolutus\u003c/em\u003e and \u003cem\u003eEo. proximus\u003c/em\u003e, especially the anterior lobe of mf6-mf8, which is square with a small, curved incision on the labial margin of mf8.\u003c/p\u003e\u003ch2\u003eDorsal carapace\u003c/h2\u003e\u003cp\u003eMLP-PV 29-X-10-2 and FMNH-P14439 are complete dorsal carapaces (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e), allowing the full description of the carapace morphology of \u003cem\u003eStromaphorus\u003c/em\u003e, including its ornamentation pattern variation. The carapace is composed of approximately 35–43 transverse bands of osteoderms, which are not always complete, especially in the anterior region.\u003c/p\u003e\u003cp\u003eIn lateral view the dorsal carapace is sub-rectangular in outline, short and globular, unlike \u003cem\u003eNeosclerocalyptus\u003c/em\u003e, which shows an almost completely straight dorsal profile. In both specimens (MLP-PV 29-X-10-2 and FMNH-P14439) the most convex region is at the middle, showing some similarity with the genera \u003cem\u003ePlohophorus\u003c/em\u003e, \u003cem\u003ePseudoplohophorus\u003c/em\u003e, \u003cem\u003eEosclerocalyptus\u003c/em\u003e, and \u003cem\u003eNopachtus\u003c/em\u003e; it contrasts with \u003cem\u003ePanochthus\u003c/em\u003e and \u003cem\u003eDoedicurus\u003c/em\u003e in which the convexity is at the anterior region; and \u003cem\u003eGlyptodon\u003c/em\u003e, with its greatest convexity in the posterior region; and \u003cem\u003eEleutherocercus\u003c/em\u003e, which is dome-shaped at the dorsal-pelvic region. The exposed surface of the osteoderms that forms the carapace shows a \"rosette\" pattern, although its morphology varies in different regions of the carapace. In the anterior region (lateral and central) the osteoderms have on their exposed surface a central figure surrounded by a single row of peripheral figures (simple rosette pattern). In the middle and posterior regions this pattern becomes more complex by the addition of incomplete rows of peripheral figures and accessory figures, all of them small and rounded, similar to the pattern observed in the genera \u003cem\u003ePlohophorus\u003c/em\u003e, \u003cem\u003ePseudoplohophorus\u003c/em\u003e, \u003cem\u003eStromaphoropsis\u003c/em\u003e and \u003cem\u003eCranithlastus\u003c/em\u003e. Although this pattern is very similar to that of the genera \u003cem\u003eNopachtus\u003c/em\u003e, \u003cem\u003ePanochthus\u003c/em\u003e and \u003cem\u003ePropanochthus\u003c/em\u003e, it differs because in these latter taxa the rows of peripheral figurines are always complete and with a polygonal morphology. Thus, \u003cem\u003eNopachtus\u003c/em\u003e has up to 2 complete rows, while \u003cem\u003ePanochthus\u003c/em\u003e and \u003cem\u003ePropanochthus\u003c/em\u003e have between 3 to 7 complete rows per osteoderm, showing in some species a pattern of ornamentation mostly reticular in the dorsal region.\u003c/p\u003e\u003cp\u003eIn the antero-dorsal region of the carapace (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e: a1, a4), the osteoderms in the area near the cephalic notch are pentagonal to hexagonal, composed of a large central figure surrounded by a single row of 10 to 12 very small rounded peripheral figures and occasional accessory figures; this reduction in size is especially noticeable in the osteoderms forming the cephalic notch, where in some cases only the central figure is observed. The smallest osteoderms of the entire carapace are found on the expanded antero-lateral margin (for measurements see Online Resource 3, table 3), with an irregular pentagonal morphology that gradually changes to rectangular towards the ventral edge. These osteoderms have a central figure surrounded by a single row of very small peripheral figures, unlike \u003cem\u003eNeosclerocalyptus\u003c/em\u003e in which no peripheral figures are observed (Quiñones et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn the mid-dorsal region of the carapace (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e: a2, a5), the osteoderms are typically pentagonal or hexagonal in outline, with a subcircular, slightly convex central figure surrounded by a single, complete row of 11 to 15 peripheral figures; in addition, a second incomplete row of peripheral figures is observed at the anterior and posterior margins, often extending into adjacent osteoderms. Towards the lateral margins of the carapace, osteoderms become increasingly rectangular in shape and exhibit a gradual reduction in size. In these osteoderms, the central figure possesses a single complete row of peripheral figures, with remnants of a second incomplete row present along the anterior margin. This specific pattern of multiplication of peripheral figures distinguishes \u003cem\u003eS. ameghini\u003c/em\u003e from \u003cem\u003eNopachtus\u003c/em\u003e, which exhibits up to two complete rows, and from \u003cem\u003ePropanochtus\u003c/em\u003e and \u003cem\u003ePanochthus\u003c/em\u003e, which can develop 5–7 complete rows surrounding the central figure.\u003c/p\u003e\u003cp\u003eIn the first osteoderms of the postero-dorsal region (pelvic region) \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (MLP-PV 29-X-10-2) has in the first row a total of 12 to 21 and the second between 21 to 27, in contrast to \u003cem\u003ePanochthus\u003c/em\u003e (e.g., \u003cem\u003eP. tuberculatus\u003c/em\u003e (Owen, 1845) and \u003cem\u003eP. greslebini\u003c/em\u003e Castellanos, 1942 in which a significant increase in the number of peripheral figures results in a completely reticular pattern). In the postero-dorsal region of the carapace (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e: a3, a6), each of the osteoderms consists of a large rounded and bulging central figure, approximately 30 mm by 32. 4 mm, similar to \u003cem\u003eStromaphorus trouessarti nov. comb.\u003c/em\u003e (Moreno, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e) more convex than in \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e Ameghino, 1888, but less than in \u003cem\u003eNopachtus cabrerai\u003c/em\u003e (Zamorano, Scillato-Yané, Gonzalez-Ruiz, Zurita 2011). It should be noted that in this sector of the carapace, the main axis of the central figure is inclined towards the posterior region of the osteoderm, similar to \u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e but different from \u003cem\u003eN. cabrerai\u003c/em\u003e in which it is perfectly vertical. In these osteoderms the central figure is surrounded by a complete row of 12 to 21 peripheral figures; while on its posterior margin, near the caudal notch, a second row of incomplete figures occurs, unlike in \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e, which has two complete rows of peripheral figures and \u003cem\u003eN. cabrerai\u003c/em\u003e with the second row complete but shared with the adjoining osteoderms. On the other hand, the osteoderms that constitute the caudal margin present the central figure with an anterior margin composed of 3 to 5 rows of peripheral figurines (see Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e), a row towards the sides of the central figure and a free edge towards the posterior margin. In this posterior region of the carapace, the osteoderms are aligned in orderly bands, counting between 8 and 10 bands of osteoderms.\u003c/p\u003e\u003ch2\u003eCaudal tube\u003c/h2\u003e\u003cp\u003eOverall, the caudal tube (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e) of \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (MLP-PV 29-X-10-1; MLP-PV 29-X-8-9; FMNH-P14414 and FMNH-P14532), both in morphology and rosette ornamentation, is somewhat similar to those of \u003cem\u003ePseudoplohophorus\u003c/em\u003e (FC-CVF 475\u003cb\u003e/\u003c/b\u003e595 and MMC 888), \u003cem\u003ePlohophorus figuratus\u003c/em\u003e (MLP-PV 98-XI-21-1), \u003cem\u003eEosclerocalyptus proximus\u003c/em\u003e (PVL 375) and \u003cem\u003eNeosclerocalyptus\u003c/em\u003e (MMP 4300; MACN 13084 and PZ-Ctes 5879); however, it differs from the tubes of \u003cem\u003eHoplophorus euphractus\u003c/em\u003e (UFMG 1235), \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e (MLP-PV 16–122), \u003cem\u003ePropanochthus bullifer\u003c/em\u003e (MANC-Pv 1761), \u003cem\u003ePanochthus\u003c/em\u003e (MLP-PV 16–29; AMNH 11243; MHNC-13491 and MANC-Pv 5130), \u003cem\u003eEleutherocercus solidus\u003c/em\u003e (FMNH-P14437; MLP-PV 16–25; MLP-PV 29-X-10-21 and MACN 2893), and \u003cem\u003eDoedicurus clavicaudatus\u003c/em\u003e (MLP-PV 16–23).\u003c/p\u003e\u003cp\u003eThe caudal tube of \u003cem\u003eS. ameghini\u003c/em\u003e is almost circular in transverse outline. In dorsal and ventral views, it has a relatively constant diameter towards the distal portion, resulting in an almost semi-rectangular morphology (for measurements see Online Resource 3, table 4). It differs from \u003cem\u003ePseudoplohophorus\u003c/em\u003e and \u003cem\u003ePlohophorus\u003c/em\u003e whose tubes narrow markedly towards the distal region, acquiring a conical morphology; from \u003cem\u003eEosclerocalyptus\u003c/em\u003e and \u003cem\u003eNeosclerocalyptus\u003c/em\u003e, which have narrower and more gracile tubes; from \u003cem\u003eHoplophorus euphractus\u003c/em\u003e whose tube has a robust cylindrical morphology, but is slightly dorso-ventrally flattened from the middle to the terminal portion; from \u003cem\u003ePropanochthus\u003c/em\u003e, \u003cem\u003ePanochthus\u003c/em\u003e and \u003cem\u003eEleutherocercus\u003c/em\u003e which are dorso-ventrally flattened even more than in \u003cem\u003eHoplophorus\u003c/em\u003e and \u003cem\u003eNopachtus\u003c/em\u003e; and \u003cem\u003eDoedicurus\u003c/em\u003e in which the distal end of its caudal tube is similar to a club.\u003c/p\u003e\u003cp\u003eIn dorsal view (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e: a, c), the ornamentation of the caudal tube is composed of a simple rosette pattern, in which each osteoderm has a smooth-surfaced central figure, surrounded by a single complete row of peripheral figures, which in turn are shared with the adjoining osteoderms. The peripheral figures are rounded (for measurements see Online Resource 3, table 4), being smaller than those observed in \u003cem\u003ePlohophorus.\u003c/em\u003e The morphology of the central figures of the proximal portion varies from circular to subcircular, they are large, and arranged in transverse rows. However, towards the distal portion, the central figures become gradually smaller. In the middle portion, the central figures are clearly oval, while those of the more distal portion revert to a more circular shape, all of them surrounded by a single series of peripheral figures shared with the contiguous central figures.\u003c/p\u003e\u003cp\u003eAt the apex there are two large smooth terminal figures (T) bordering this end of the tube to the sides, the lateral margins of these figures, in the more distal portion, are in contact, as in \u003cem\u003ePlohophorus\u003c/em\u003e and \u003cem\u003ePseudoplohophorus\u003c/em\u003e, In turn they differ from \u003cem\u003eHoplophorus euphractus\u003c/em\u003e and \u003cem\u003eNopachtus\u003c/em\u003e with characteristic large lateral spines, from \u003cem\u003ePropanochthus\u003c/em\u003e and \u003cem\u003ePanochthus\u003c/em\u003e with very rugose depressed elliptical lateral figures, raised in the middle, from \u003cem\u003eEleutherocercus\u003c/em\u003e and \u003cem\u003eDoedicurus\u003c/em\u003e with very rugose elliptical lateral and apical figures, but completely concave. T figures are preceded by 4 smooth lateral figures (I, II, III and IV) which will be explained in detail in the lateral view. Between the T figures in their anterior region, there is a characteristic rounded figure called posterior figure (P), which is also present in other genera such as \u003cem\u003ePseudoplohophorus\u003c/em\u003e (see Toriño and Perea \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Towards proximal regions of the tube, before P, there is symmetry between the left (L) and right (R) marginal figures (see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e: a, c), thus up to 10 pairs of marginal figures have been identified. Between the marginal figures, there are other central figures arranged in rows, the number of which increases towards the proximal region of the tube. It should be noted that the arrangement and number of these central figures are slightly variable between the different individuals of the species.\u003c/p\u003e\u003cp\u003eIn ventral view, the ornamentation of the caudal tube is very similar to the dorsal side, although the central figures are somewhat larger and flatter. These figures, particularly at the terminal portion, come into contact without peripheral figures between them. An inverse relationship exists between the size of the central figures and the surrounding peripheral figures. As the central figures expand, the peripheral figures become significantly smaller (may even disappear), unlike the larger peripheral figures of the dorsal region.\u003c/p\u003e\u003cp\u003eIn lateral view (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e: b, d) there are large terminal figures (T) in the distal portion, with a highly convex surface, as well as four lateral figures (numbered I, II, III and IV in distal-proximal sense, see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e7\u003c/span\u003e) which gradually reduce in size towards the proximal portion. Figure I is convex, unlike figures II, III and IV, which are flatter. Between the lateral and marginal figures, there is a row of small peripheral figures. Finally, the number of lateral figures varies from 3 to 4, even in the same individual. This is the case of MLP-PV 29-X-10-1 whose right side has the four figures described above (I, II, III and IV) and whose left side has only three of them (I, II and III).\u003c/p\u003e\n\u003ch3\u003eAdditional observations\u003c/h3\u003e\n\u003cp\u003eRegarding \u003cem\u003eS. trouessarti\u003c/em\u003e nov. comb., a redescription and comparative diagnosis of the species can be found in Zamorano et al. (\u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003ePhylogenetic analysis\u003c/h2\u003e \u003cp\u003eThe analysis resulted in two MPTs, only differing in the location of both species of \u0026ldquo;Palaehoplophorini\u0026rdquo; (\u003cem\u003ePalaehoplophoroides rothi\u003c/em\u003e and \u003cem\u003ePalaehoplophorus meridionalis\u003c/em\u003e). In turn, the strict consensus shows RI\u0026thinsp;=\u0026thinsp;0.933 and CI\u0026thinsp;=\u0026thinsp;0.845; length\u0026thinsp;=\u0026thinsp;174.020 steps (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). It demonstrates the monophyly of the family Glyptodontidae mainly supported by dental and mandibular synapomorphies, such as the presence of flat occlusal surfaces on all molariforms [25:1], trilobulation [30:0;34:1] and loss of premaxillary teeth [26:0]; but also by exoskeletal features such as the loss of mobile bands [49:1]. As mentioned above (see Introduction) we prefer the use of Glyptodontidae at family level, in agreement with most of the recent contributions, and taking into account the noticeable divergent morphology of glyptodonts when compared to the remaining Cingulata diversity (see Machado et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Consistent with the observations of Croft et al. (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), the enigmatic \u003cem\u003eParapropalaehoplophorus septentrionalis\u003c/em\u003e is the sister species of remaining diversity of Glyptodontidae. Among the most characteristic mandibular morphology stands out as particularly distinctive. Given that this anatomical feature was the subject of a geometric morphometric analysis (Character 0), it will be discussed separately at the conclusion of this section.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe MPT topology shows that the remaining diversity of Glyptodontidae is divided into two major clades, Glyptodontinae and Hoplophorinae (=\u0026rdquo;Austral clade\u0026rdquo; of Barasoian et al. 2022a; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e); these two clades have already been reported in previous works (see Cuadrelli et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Nu\u0026ntilde;ez-Blasco et al. 2021c; Barasoian et al. 2022a). Glyptodontinae is mainly supported by endoskeletal synapomorphies, some from the carapace and some from the caudal armour (Node A). The topology of this node is in agreement with Zurita et al. (\u003cspan citationid=\"CR139\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and Cuadrelli et al. (\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe second clade, Hoplophorinae, is supported by four exoskeletal synapomorphies (increase in size of cephalic shield osteoderms [35:0], acquisition of peripheral figures in caudal rings osteoderms [71:1], [72:1], and caudal armour composed of caudal rings and absence of terminal tubercle [73:0]). Some Burdigalian (Early-Middle Miocene) taxa (\u003cem\u003ePropaleoplohophorus australi\u003c/em\u003es, \u003cem\u003eCochlops muricatu\u003c/em\u003es and \u003cem\u003eEucinepeltus petesatus\u003c/em\u003e) are located at the base of this second radiation forming a monophyletic group, supported by a reduction in the number of molariforms with trilobulation [27:3], rings representing 80% of total length of the caudal armour [75:1]; the species \u003cem\u003ePro. australi\u003c/em\u003es and \u003cem\u003eC. muricatu\u003c/em\u003es are grouped together, supported by ambiguous synapomorphies related to the cephalic shield (acquisition of peripheral figures [36:1], [38:1]); \u003cem\u003eE. petesatus\u003c/em\u003e is located as the sister species of this group (but see Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e for another interpretation), the three taxa are forming the tribe Propalaehoplophorini (=\u0026thinsp;Propalaehoplophorinae \u003cem\u003esensu\u003c/em\u003e Ameghino, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1891c\u003c/span\u003e). It is interesting to analyze at this point the character 74; this describes the morphology of the caudal armor composed of caudal rings and a caudal tube, which are present in all members of the Hoplophorinae with the exception of \u003cem\u003eEucinepeltus petesatus\u003c/em\u003e and \u003cem\u003ePropaleoplohophorus australis\u003c/em\u003e. These two basal species have neither terminal tubercle (as in the case of Glyptodontinae) nor a fully developed caudal tube, but have another structure that could be intermediate; therefore, in characters 73 and 74, they have been coded as absent [73:0; 74:0].\u003c/p\u003e \u003cp\u003eThe \u0026ldquo;Palaehoplophorini\u0026rdquo; (\u003cem\u003ePalaehoplophoroides rothi\u003c/em\u003e and \u003cem\u003ePalaehoplophorus meridionals\u003c/em\u003e) forms a polytomy with the remaining extra-patagonian diversity, this politomy is supported by exoskeletal synapomorphies (similar ornamentation between dorsal and caudal armors [70:1], presence of caudal tube [74:1 ambiguous] and rings representing up to 60% or less of total length of the caudal armour [75:2]). In turn, the Late Miocene (ca. 10\u0026thinsp;\u0026minus;\u0026thinsp;9 Ma) species \u003cem\u003eKelenkura castroi\u003c/em\u003e, appears as the sister taxon of the Neogene and Quaternary remaining glyptodonts, in accordance with the proposal of Barasoain et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e). This relationship is supported by synapomorphies related to the caudal tube (acquisition of central figures and rosette pattern in the dorsal region [80:1], [84:1] the latter subsequently modified in \u003cem\u003eDoedicurus\u003c/em\u003e, \u003cem\u003ePanochthus\u003c/em\u003e and \u003cem\u003ePropanochthus\u003c/em\u003e; and acquisition of lateral figure\u003cb\u003es\u003c/b\u003e [86:1], it must be noted that this state changes throughout the evolution of the clade).\u003c/p\u003e \u003cp\u003eOn the other hand, the clade clustering the late Neogene and Quaternary glyptodonts is strongly supported by six exoskeletal synapomorphies (acquisition of more peripheral figures in the posterior region of the carapace [64:2]), caudal tube without visible rings [76:4], [78:1], caudal tube with a rosette pattern with peripheral figures well-developed and completely surrounding the central figures [82:2], acquisition of five to seven large lateral figures [88:1], loss of large lateral figures at the apex of the caudal tube [93:0]), and two endoskeletal synapomorphies (both related to the degree of development of the third trochanter [43:1 ambiguous], [44:1 ambiguous]).\u003c/p\u003e \u003cp\u003eThe first subdivision (and the subject of this analysis) is formed by the monophyletic group that clusters the species \u003cem\u003ePseudoplohophorus absolutus\u003c/em\u003e, \u003cem\u003ePs. benvenuti\u003c/em\u003e, \u003cem\u003ePlohophorus figuratus, Stromaphorus ameghini\u003c/em\u003e and \u003cem\u003eStromaphorus trouessarti nov. comb.\u003c/em\u003e comprising the tribe Plohophorini (Node B), it is supported by one dental synapomorphy (increasing of size of Mf1 [29:0]); and three related to the carapace ornamentation (multiplication of peripheral figures of dorsal osteoderms [56:1] and peripheral carapace figures with circular morphology [60:1], [61,0]).\u003c/p\u003e \u003cp\u003eDescribing in more detail the internal relationships of the species forming the tribe Plohophorini, the Pampean species \u003cem\u003ePlohophorus figuratus\u003c/em\u003e appears as sister taxon to \u003cem\u003ePlohophorus avellaneda\u003c/em\u003e (ornamentation of the proximal-ventral region of the caudal tube [95:0]); and these in turn as sister group of \u003cem\u003eStromaphorus ameghini\u0026thinsp;+\u0026thinsp;Stromaphorus trouessarti nov. comb.\u003c/em\u003e (supported by four carapace synapomorphies (loss of peripheral figures in osteorderms of the cephalic notch [52:0]; surface of the osteoderms of the posterior region of the carapace becoming blistered [62:2]; and increase in the number of peripheral figures in the posterior region of the carapace and caudal notch [64:4], [66:2]), thus confirming the condition of natural group of the genus \u003cem\u003eStromaphorus\u003c/em\u003e. The relationship between \u003cem\u003eStromaphorus\u003c/em\u003e and \u003cem\u003ePlohophorus\u003c/em\u003e is supported by two synapomorphies (position of the infraorbitary foramen [15:2], and increase in the number of rows of peripheral figures in the posterior region of the carapace [63:2]); in turn, they form the sister group of the late Neogene Uruguayan taxa \u003cem\u003ePseudoplohophorus absolutus\u003c/em\u003e and \u003cem\u003ePs. benvenuti\u003c/em\u003e, \u003cb\u003e(\u003c/b\u003esupported by the morphology of the palate [24:1 ambiguous synapomorphy]\u003cb\u003e)\u003c/b\u003e.\u003c/p\u003e \u003cp\u003ePlohophorini is recovered as the sister group of the remaining members of this second clade (Doedicurini\u0026thinsp;+\u0026thinsp;Hoplophorini\u0026thinsp;+\u0026thinsp;Neosclerocalyptini), wich is supported by a cranial synapomorphy (position of the infraorbitary foramen [15:1]), plus another from the caudal armour (acquisition of polygonal ornamentation in the caudal tube [85:1]) (but see Fernicola and Porpino \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e for another proposal). The following node (Node C) encompasses the tribe Doedicurini (=\u0026thinsp;Doedicurinae \u003cem\u003esensu\u003c/em\u003e Trouessart, \u003cspan citationid=\"CR125\" class=\"CitationRef\"\u003e1897\u003c/span\u003e) supported by three cranial synapomorphies (angle between posterior margin of the orbital notch and the palate plane near to 90\u0026ordm; [20:1], morphology of the palate acquiring a similar transverse expansion at their posterior and anterior margins [24:2 ambiguous], and increase in the antero-posterior diameter of Mf1, with oval morphology [29:1]); seven carapace synapomorphies (most of them related to the loss of ornamentation [52:3], [54:0], [62:3], [63:0], [64:0], [66:4], one related to the acquisition of interdigitation of the margin of its osteoderms [57:2]); and five caudal armour synapomorphies (transverse outline of distal third of caudal tube becoming dorsoventrally flattened [77:1 ambiguous], cylindrical-conical caudal tube composed of ankylosed osteoderms acquiring large lateral depressions (concave insertion structures) [86:3], [88:2], [89:1], [91:1]).\u003c/p\u003e \u003cp\u003eDoedicurini appears as the sister group of \u003cem\u003eEosclerocalyptus proximus\u003c/em\u003e, forming a clade supported by two cranial synapomorphies (labiolingual trilobation evident from Mf2 [27:1 ambiguous], anteroposterior diameter of Mf1 less than 50% of the anteroposterior diameter of Mf2 [29:2 ambiguous]); two carapace synapomorphies (increasing in the number of polygonal peripheral figures in the carapace [61:2], the central figure occupies less than 50% of the total surface area of the osteoderms near the caudal notch [68:0 ambiguous]), plus another from the caudal armour (peripheral figures surround the entire central figure in the central lateral zone of the caudal tube [87:3]); and the group Neosclerocalyptini\u0026thinsp;+\u0026thinsp;Hoplophorini, supported cranial synapomorphies (acquisition of pneumatization of the rostral area of the skull [4:1], acquisition of peripheral figures in cephalic shield [36:1 ambiguous], acquisition of peripheral rows in the cephalic shield [38:1 ambiguous]).\u003c/p\u003e \u003cp\u003eThe tribe Neosclerocalyptini (Node D) is supported by six cranial synapomorphies (acquisition of ossified nasal cartilages [5:1] and [8:1], acquisition of a \u0026ldquo;V\u0026rdquo; groove separating the modified nasal area from the rest of the skull [9:1], ventral edge of the orbital notch coinciding with the 50% of the dorso-ventral diameter of the ossified nasal cartilages [12:1], distance between the infraorbitary foramen and the labial margin of the Mf3, equal or larger than the antero-posterior diameter of the Mf1-Mf2 [14:1], acquisition of cephalic shield with sub-angular outline [37:1]); and three carapace synapomorphies (loss of convexity of carapace, in lateral view [47:0] and [48:3], acquisition of antero-lateral expansion of the carapace [50:1]). Neosclerocalyptini is the sister group of the tribe Hoplophorini (Node E) grouped by a total of five synapomorphies, of which the most relevant have been those of the caudal armour (acquisition of sigmoid contour in the anterolateral border of the nasal openings [21:1], transverse outline of distal third of caudal tube dorsoventrally flattened [77:1 ambiguous], acquisition of convex insertion structures in the lateral figures of the caudal tube [86:2], increasing rows of peripheral figures around the entire central lateral figure [87:4], acquisition of depressions with convex central area in the terminal zone of the caudal tube [90:1]), this latter (Hoplophorini) formed by the species \u003cem\u003eHoplophorus euphractus\u003c/em\u003e, \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e, \u003cem\u003ePropanochtus bullifer\u003c/em\u003e, \u003cem\u003ePanochthus tuberculatus\u003c/em\u003e, and \u003cem\u003eP. intermedius\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eIn turn, the geometric morphometry applied to the mandibles in lateral view has been included as an additional character (Character 0) in the phylogenetic matrix, resulting in a mixed matrix (conventional characters\u0026thinsp;+\u0026thinsp;geometric morphometry characters). Character 0 has played a crucial role in establishing \u003cem\u003ePa. septentrionalis\u003c/em\u003e as the sister taxon to Glyptodontinae\u0026thinsp;+\u0026thinsp;Hoplophorinae. This is primarily attributed to the unique 90\u0026ordm; angle formed by the horizontal and ascending mandibular rami. This morphology represents a clearly plesiomorphic condition, intermediate between that of Pampatheriidae or Dasypodidae (greater than 90\u0026ordm;) and the more derived Glyptodontidae (less than 90\u0026ordm;). On the other hand, it also contributed for grouping the Propaleoplohophorini by their morphological affinity.\u003c/p\u003e \u003cp\u003eFinally, the geometric morphometry has increased the support of some nodes when calculating the Bremer, absolute Bremer, and Jackknifing indices (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e) in comparison with the values obtained in trees of similar topology from previous analyses (see Cuadrelli et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Nu\u0026ntilde;ez-Blasco et al. 2021c, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e). The most interesting nodes, positively affected, were those that group Plohophorini (Node B), Doedicurini (Node C), Neosclerocalyptini (Node D), and Hoplophorini (Node E). The support values obtained for Node B (Plohophorini) are for the first time notably high, Bremer index of 100, absolute Bremer index of \"3?\" (calculated with 3-step suboptimality) and Jackknifing of 87. Finally, the comparison between the morphotypes representative of each of these subsets, also increased the support of the node that groups them (Doedicurini\u0026thinsp;+\u0026thinsp;Hoplophorini) resulting in a Bremer index value of 40, absolute Bremer index of 2 (calculated with 3-step suboptimality) and Jackknifing of 63.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eLocal geographical and stratigraphic distribution\u003c/h2\u003e \u003cp\u003eThe holotypes of \u0026ldquo;\u003cem\u003ePhlyctaenopyga\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e MLP-PV 16\u0026ndash;101 (\u003cem\u003ePlohophorus ameghini\u003c/em\u003e, Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e) and \u0026ldquo;\u003cem\u003eStromaphorus compressidens\u003c/em\u003e\u0026rdquo; MLP-PV 16\u0026ndash;138 (\u003cem\u003eNeuryurus compressidens\u003c/em\u003e, Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e) come from the \"Araucanian\" levels of the \u0026ldquo;Bajo de Andalhuala\u0026rdquo; in the southern part of the SMV, probably from the upper levels of the Chiquimil Formation or from the Andalhuala Formation (i.e. between 6.88 to 4.85\u0026ndash;3.4 Ma; Bonini et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The skull FMNH-P14396, from Ampajango locality (Santa Mar\u0026iacute;a Department, Catamarca Province) originaly referred to \u003cem\u003eS. compressidens\u003c/em\u003e, comes from XI level (by Marshall and Patterson \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e, Appendix II), and according to the correlation proposed by other authors, such as Georgieff et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and Bonini et al. (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), this level is between 6.88 to 6.02 Ma. The specimen FMNH-P14367, a carapace here reassigned to a juvenile of \u003cem\u003eS. ameghini\u003c/em\u003e from Loma Rica locality (Andalhuala, Santa Mar\u0026iacute;a Department, Catamarca Province), cames from XVII level (Marshall and Patterson \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1981\u003c/span\u003e), and according to Georgieff et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and Bonini et al. (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), this level is above 6.02Ma. The materials MLP-PV 16\u0026ndash;134, MLP-PV 16\u0026ndash;135, MLP-PV 19\u0026ndash;136 (Holotype of \u003cem\u003ePlohophorus philippii\u003c/em\u003e Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), and MLP-PV 16\u0026ndash;137 referred to \u0026ldquo;\u003cem\u003eS. compressidens\u003c/em\u003e\u0026rdquo;, come from north ridge of Loma Rica (Andalhuala, Santa Mar\u0026iacute;a Department, Catamarca Province), based on the stratigraphic levels outcropping in that area, their age could be slightly lower than 6.02Ma (see Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding the Villavil-Quillay Basin (located ca. 78 km at southwest from the Santa Maria Valley), there are several localities where numerous fossils have been found, belonging to both species (\u0026ldquo;\u003cem\u003ePhlyctaenopyga\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e and \u0026ldquo;\u003cem\u003eStromaphorus compressidens\u003c/em\u003e\u0026rdquo;). The most remarkable localities are Puerta de Corral Quemado, Bel\u0026eacute;n Department (\u0026ldquo;\u003cem\u003ePh.\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e: MLP-PV 29-X-8-2; MLP-PV 29-X-8-7; MLP-PV 29-X-10-6; MLP-PV 29-X-10-10; FMNH-P14439; FMNH-P14532; FMNH-P14414; FMNH-P14447; CCPCQ 03-DPA-Pv264 and \u0026ldquo;\u003cem\u003eS. compressidens\u003c/em\u003e\u0026rdquo;: MLP-PV 29-X-8-1; MLP-PV 29-X-8-9; MLP-PV 29-X-10-54; FMNH-P14494; FMNH-P14520; FMNH-P14414; FMNH-P14494; FMNH-P14520) and Corral Quemado, Bel\u0026eacute;n Department (\u0026ldquo;\u003cem\u003ePh.\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e: MLP-PV 31-XI-12-7). In turn, the second most fossiliferous locality for these species is San Fernando (North and South), Bel\u0026eacute;n Department (\u0026ldquo;\u003cem\u003ePh.\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e: MLP-PV 29-X-10-1; MLP-PV 29-X-10-2; MLP-PV 29-X-10-3; MLP-PV 29-X-10-5; MLP-PV 29-X-10-40; MCH-P39; MCH-P176; MCH-P177; MCH-P247; MCH-P323, and \u0026ldquo;\u003cem\u003eS. compressidens\u003c/em\u003e\u0026rdquo;: MLP-PV 29-X-10-8; MLP-PV 29-X-10-51; MCH-P174; MCH-P328) (see Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e; Marshall and Paterson 1981; Bonini \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). It should be noted that the remains referring to both taxa not only appear indistinctly in these localities, but also in the same time interval, specifically in the Andalhuala and Chiquimil formations, in strata dated between 7.14 +/- 0.05 and 3.66 +/- 0.05 Ma. In adittion, we also report at least two records located above the tuff dated at 3.66 (FMNH-P14447) and below that dated at 7.14 (MCH-P364), but without greater stratigraphic precision.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eTaxomomy and nomenclature\u003c/h2\u003e \u003cp\u003eGlyptodonts were very probably the most fascinating and unusual xenarthrans that ever existed. Recent findings of glyptodonts from late Neogene sediments of Ecuador, which are morphologically distant from those known of La Venta (Colombia) and Anzoategui (Venezuela) (see Zurita et al. \u003cspan citationid=\"CR139\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) in northern South America, and from those of southern South America (see Gonz\u0026aacute;lez-Ruiz \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Barasoian et al. 2022a), reveal that much more studies are necessary to improve our knowledge about their evolutionary history (Rom\u0026aacute;n-Carri\u0026oacute;n et al. \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). As mentioned above, the tribe Plohophorini (\u003cem\u003esensu\u003c/em\u003e Castellanos \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1932\u003c/span\u003e; Hoffstetter \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1958\u003c/span\u003e) is one of the most enigmatic groups within the southern clade Hoplophorinae, and particularly those taxa from the late Neogene of the NWA region were, before this contribution, poorly known from the taxonomic, stratigraphic and phylogenetic viewpoint. In the Villavil-Quillay Basin and Santa Mar\u0026iacute;a Valley two species were recognized, \u0026ldquo;\u003cem\u003ePhlyctaenopyga\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e) and \u0026ldquo;\u003cem\u003eStromaphorus compressidens\u003c/em\u003e\u0026rdquo; (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e), both sharing the same stratigraphic and geographic provenance (see Cabrera \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e; this work). Two issues can be inferred from geographical and stratigraphic provenance data (see Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e1\u003c/span\u003e): (1) the remains are quite numerous, so they were common taxa in this area; (2) two different, but at the same time so similar species coexisting at the same time in the same area would have resulted in high interspecific competition, due to direct rivalry for the same resources. From an ecological perspective, the co-ocurrence of two different species is hardly feasible, as it was observed in Pleistocene genera (see Cuadrelli et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this regard, a careful morphologic and taxonomic analysis made in this contribution revealed that, actually, both names must be considered as synonyms, being \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; ex Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e) the only species of Plohophorini present in the late Neogene sequences of NWA. Following the ICZN 2000, Art. 23.1 and 23.3, the genus \u003cem\u003eStromaphorus\u003c/em\u003e Castellanos, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1925\u003c/span\u003e, has priority over \u003cem\u003ePhlyctaenopyg\u003c/em\u003ea Cabrera, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1944\u003c/span\u003e, while the specific epithet \u0026ldquo;\u003cem\u003eameghini\u003c/em\u003e\u0026rdquo; has priority over \u0026ldquo;\u003cem\u003ecompressidens\u003c/em\u003e\u0026rdquo; (see Online Resource 1). Besides, the comparative study with \u0026ldquo;\u003cem\u003ePhlyctaenopyga\u003c/em\u003e\u0026rdquo; \u003cem\u003etrouessarti\u003c/em\u003e from the Monte Hermoso Formation (Pliocene) revealed the presence of some shared characters between both species at the level of the dorsal carapace (see Description and comparisons) that suggests the inclusion of this Pampean species in the genus \u003cem\u003eStromaphorus\u003c/em\u003e, as \u003cem\u003eS. trouessarti\u003c/em\u003e (Moreno, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e) \u003cem\u003enov. comb.\u003c/em\u003e In consequence, the genus \u003cem\u003eStromaphorus\u003c/em\u003e now encompasses two species, \u003cem\u003eS. ameghini\u003c/em\u003e and \u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e\u003c/p\u003e \u003cp\u003eIn summary, given the noticeable anatomical similarities between both species already discussed (ie., \u0026ldquo;\u003cem\u003ePh.\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e and \u0026ldquo;\u003cem\u003eS. compressidens\u003c/em\u003e\u0026rdquo;), and taking into account that no spatial discontinuities are observed (through their geographical provenance) nor temporal discontinuities (through their stratigraphic provenance as well as the dating of the tuffaceous levels), following the criteria of Simpson (1951) and Castro et al. (2013), the evidence supports the existence of a single species, \u003cem\u003eStromaphorus ameghini\u003c/em\u003e, among Plohophorini glyptodonts of NWA.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003ePhylogeny\u003c/h2\u003e \u003cp\u003eAs observed in previous analyses (e.g., Barasoian et al. 2022a; Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), two large clades can be observed. One of northern origin, Glyptodontinae, and the other object of this study, Hoplophorinae, which includes the remaining diversity.\u003c/p\u003e \u003cp\u003eWithin the southern lineage Hoplophorinae, cladistic results show that the tribe Plohophorini (\u003cem\u003esensu\u003c/em\u003e Castellanos \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1932\u003c/span\u003e (nom. transl. Hoffstetter \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1958\u003c/span\u003e) appears as the sister group of Doedicurini\u0026thinsp;+\u0026thinsp;Hoplophorini\u0026thinsp;+\u0026thinsp;Neosclerocaliptini, forming part of the extra-Patagonian radiation starting with \u003cem\u003eKelenkura castroi\u003c/em\u003e (see Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e). The result of our analysis shows the condition of natural group of Plohophorini (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e, Node B), a result that agrees with Qui\u0026ntilde;ones et al. (\u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e); this condition is supported by several cranial and postcranial synapomorphies. Regrettably, we were no able to include all the species traditionally interpreted as belonging to Plohophorini (see Hofstetter 1958; Paula Couto, \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e1979\u003c/span\u003e), since many of them are recognized on the basis of very fragmentary type material within a strict typological taxonomic criterion and lacking precise stratigraphic provenance (Tori\u0026ntilde;o and Perea \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Besides this, one interesting point is the distribution of the different species in this so far poorly known tribe. First, as observed in the topology of the MPT (see Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e), the monophyly of the genera \u003cem\u003eStromaphorus\u003c/em\u003e (\u003cem\u003eS. ameghini\u0026thinsp;+\u0026thinsp;S. trouessarti nov. comb.\u003c/em\u003e), \u003cem\u003ePlohophorus\u003c/em\u003e (\u003cem\u003eP. figuratus\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eP. avellaneda\u003c/em\u003e) and \u003cem\u003ePseudoplohophorus\u003c/em\u003e (\u003cem\u003ePs. absolutus\u0026thinsp;+\u0026thinsp;Ps. benvenuti\u003c/em\u003e) is confirmed and supported by several cranial and postcranial synapomorphies (see Online Resource 4 and Online Resource 5). Moreover, \u003cem\u003ePlohophorus figuratus\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eP. avellaneda\u003c/em\u003e appears as the sister group of the clade clustering \u003cem\u003eS. ameghini\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e, condition supported by a couple of cranial and exoskeletal synampomorhies.\u003c/p\u003e \u003cp\u003eBeyond the phylogeny of the Plohophorini, another interesting point refers to the location of the enigmatic species \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e, here interpreted as the sister and earliest divergent taxa of the lineage containing \u003cem\u003eHoplophorus\u003c/em\u003e, \u003cem\u003ePropanochthus bullifer\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003ePanochthus\u003c/em\u003e spp. The phylogenetic position of \u003cem\u003eN. coagmentatus\u003c/em\u003e has been controversial since Ameghino (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e) in a pre-cladistic time. More recently, Zamorano and Brandoni (\u003cspan citationid=\"CR131\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) placed this species close to the genera \u003cem\u003ePlophophorus, Pseudoplohophorus, Propanochtus\u003c/em\u003e, \u0026ldquo;\u003cem\u003ePhlyctaenopyga\u003c/em\u003e\u0026rdquo;, and \u003cem\u003eStromaphorus\u003c/em\u003e. However, and as mentioned above, our results show that \u003cem\u003eN. coagmentatus\u003c/em\u003e is in fact the sister species within the lineage including \u003cem\u003eHoplophorus\u003c/em\u003e, \u003cem\u003ePropanochthus\u003c/em\u003e, and \u003cem\u003ePanochthus\u003c/em\u003e spp. (Hoplophorini). This suggests that the multiplication of peripheral figures of this lineage and that of the Plohophorini here analyzed (i.e., \u003cem\u003ePseudoplohophoru\u003c/em\u003es, \u003cem\u003ePlohophorus\u003c/em\u003e and \u003cem\u003eStromaphorus\u003c/em\u003e) is actually a homoplasy; in this sense our results coincide with those presented by Fernicola and Porpino (\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), in which these authors already postulated a possible homoplasy between the rosette patterns ornamenting the dorsal carapace of these groups. This hypothesis is also supported by the evident morphological similitude in the caudal tubes of \u003cem\u003eNopachtus, Propanochthu\u003c/em\u003es, \u003cem\u003eHoplophorus\u003c/em\u003e, and \u003cem\u003ePanochthus\u003c/em\u003e, which in turn are very different from those of \u003cem\u003ePseudoplohophorus\u003c/em\u003e, \u003cem\u003ePlohophorus\u003c/em\u003e and \u003cem\u003eStromaphorus\u003c/em\u003e. Also, the spine-like structure observed in the caudal tubes of the lineage \u003cem\u003eNopachtus coagmentatus, Hoplophorus euphractus, Propanochthus bullifer\u003c/em\u003e, and \u003cem\u003ePanochthus\u003c/em\u003e spp. is a homologous structure rather than a convergence as usually interpreted.\u003c/p\u003e \u003cp\u003eFinally, it is noteworthy the position of the Early Miocene glyptodont \u003cem\u003eP. septentrionalis\u003c/em\u003e as the earliest divergent member of Glyptodontidae, supported by geometric morphometric character (Character 0). This result is consistent with the analysis of Croft et al. (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) that recovered this species as the sister taxon of all remaining glyptodonts.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003ePaleobiogeography, palaeoenvironments, stratigraphy, and First Appearance Datum (FAD) of Plohophorini\u003c/h2\u003e \u003cp\u003eIn southern South America (ca 48\u0026deg; to 21\u0026deg; S), where records of Neogene glyptodonts are quite frequent (see Gaudin and Croft \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Zurita et al. \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e), \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003ea\u003c/span\u003e recent revision of the diversity of the Late Miocene [Chasicoan lapse, Tortonian (ca. 10\u0026thinsp;\u0026minus;\u0026thinsp;9 Ma; see Prevosti et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e)] in central Argentina revealed a much more restricted diversity than previously supposed, with the presence of one species, \u003cem\u003eKelenkura castroi\u003c/em\u003e (Barasoain et al., 2022). Following the palaeophytogeographic scheme proposed by Barreda et al. (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Fig.\u0026nbsp;2.2.), the Proto-Spinal/Steppe province was mainly characterized during the Late Miocene (\u003cem\u003eca\u003c/em\u003e. 10\u0026thinsp;\u0026minus;\u0026thinsp;8 Ma) by the presence of low xerophytic shrubs or trees and mostly halophytic herbaceous vegetation in open environments, consistent with the progressive aridization and open biomes, and a shift toward the consumption of C4 plants in several herbivorous clades (Domingo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sanz-P\u0026eacute;rez et al. \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The presence of a single species, \u003cem\u003eKelenkura castroi\u003c/em\u003e, in central Argentina represents the first extra-Patagonian radiation within Hoplophorinae at the onset of the so-called \"Age of the Southern Plains\", an event that took place between 10\u0026thinsp;\u0026minus;\u0026thinsp;3 Ma and that was a direct consequence of the pulse of drying and cooling observed at the end of the Miocene Optimum Climatic (see Zachos et al. \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Westerhold et al. \u003cspan citationid=\"CR128\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Domingo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Candela et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The subsequent period of diversification of Hoplophorinae may have been influenced by the progressive development of open biomes, as suggested by previous studies (see Zurita et al. \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e; Barasoain et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e; Domingo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe FAD of \u003cem\u003eS. ameghini\u003c/em\u003e in the NOA is not clear, since depending on the criterion it can be established at ca. 9 or 7 Ma. The material MCH-P364 assigned to this species was exhumed from lacustrine sediments considered as lateral facies changes of the upper section of the El \u0026Aacute;spero Member in transition to El Jarillal Member (Chiquimil Formation) outcropping in the FOCP, San Fernando, Catamarca (Moyano, \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Muruaga et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Armella and Bonini, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The intrusive andesitic lacolith defined as the El \u0026Aacute;spero Member in Villavil locality was dated in 9.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 Ma (Sasso, \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), and the lacustrine facies in the FOCP have been interpreted as part of damming of a similar age volcanic event (see Bossi et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Moyano, \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Georgieff et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Likewise, the upper limit of the Jarillal Member has been dated in 7.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 (Latorre et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). If this is correct, it implies that records of Plohophorini in the NWA are older than those of the PR, where the Chasicoan glyptodonts (ca. 9\u0026thinsp;\u0026minus;\u0026thinsp;8) are limited to \u003cem\u003eK. castroi\u003c/em\u003e, since the presence of supposed Plohophorini (see Bondesio et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1980\u003c/span\u003e) was discarded by Barasoain et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e). According to the available evidence, no records of Plohophorini are present in the PR until the Montehermosan lapse, ca. 5.3 Ma (see Tomassini et al. \u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Prevosti et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). A similar situation occurs with Doedicurini glyptodonts, which have their oldest records in the NWA (see N\u0026uacute;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e, 2022). As mentioned above, the subsequent evolutionary history of the Plohophorini glyptodonts from NWA here analyzed mostly includes the latest Miocene (ca. 7.14 Ma) to the Late Pliocene (ca. 3.66 Ma) interval, during the Messsinian and Zanclean lapse, characterized by an evident expansion of C4-dominated grasslands in southern South America in \u003cem\u003eca\u003c/em\u003e. 8\u0026ndash;7 Ma (Hynek et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Domingo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). From a stratigraphic viewpoint, the materials here analyzed mostly come from the Andalhuala Formation, ranging from 7.14.to 3.66 in VQB (see Esteban et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Georgieff et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Bonini et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), except one particular fossil (FMNH-P14447) that comes from the Corral Quemado Formation and thus could be younger than 3.66 Ma. However, its imprecise stratigraphical provenance does not allow us to infer its real age. It must be noted that the ghost range predicted for this group in the calibrated phylogeny extends its origin down to \u003cem\u003eca.\u003c/em\u003e 10 Ma (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). As mentioned above, there is an interesting temporal overlap between the expansion of C4 grassland and the diversification of Hoplophorinae, which deserves further study (see also N\u0026uacute;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding the Plohophorini recorded in Argentina and here analyzed, \u003cem\u003eS. ameghini, S. trouessarti nov. comb.\u003c/em\u003e, and \u003cem\u003eP. figuratus\u003c/em\u003e lived during the latest Miocene-Pliocene under different paleoenvironmental conditions. This is because \u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e and \u003cem\u003eP. figuratus\u003c/em\u003e were geographically restricted to the Proto-Espinal/Steppe province, while the distribution of \u003cem\u003eS. ameghini\u003c/em\u003e coincides with the Neotropical province, this latter, and according to Barreda et al. (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e;Fig.\u0026nbsp;2.2), characterized by vegetation like the current Chaque\u0026ntilde;a Province (xerophytic forests, associated with palm groves, savannahs and halophytic shrub-steppe, and hydrophytic communities linked to watercourses.). It is important to emphasize that a similar paleobiogeographic pattern was already observed by N\u0026uacute;\u0026ntilde;ez-Blasco et al. (\u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e, 2022) when analyzing another clade of glyptodonts, Doedicurini, present in the late Neogene sediments of the NW and PR of Argentina. The genus \u003cem\u003eEleutherocercus\u003c/em\u003e contains two species, \u003cem\u003eE. antiquus\u003c/em\u003e (Zanclean-middle Piacencian; ca 5.33\u0026ndash;2.53 Ma) from the PR of Argentina, and \u003cem\u003eE. solidus\u003c/em\u003e (middle Messinian-middle Piacencian; ca 7.14-3Ma) from NWA (Zurita et al. \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e; N\u0026uacute;\u0026ntilde;ez-Blasco et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2021c\u003c/span\u003e). Moreover, even though they are currently under study, a similar geographic pattern could be inferred for another genus present in the late Neogene of NW and PR of Argentina, \u003cem\u003eEosclerocalyptus\u003c/em\u003e. This clusters two species, \u003cem\u003eE. proximus\u003c/em\u003e, restricted to NWA (Catamarca Province) and \u003cem\u003eE. lineatus\u003c/em\u003e, in the PR (Buenos Aires Province) (Zurita and Tomassini \u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Zurita \u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e2007a\u003c/span\u003e,\u003cspan citationid=\"CR137\" class=\"CitationRef\"\u003eb\u003c/span\u003e). Something similar happens with the distribution of \u003cem\u003eNopachtus\u003c/em\u003e Ameghino, 1888, represented by the species \u003cem\u003eN. coagmentatus\u003c/em\u003e (Ameghino, 1888) in NWA (Catamarca Province, see Zamorano et al. \u003cspan citationid=\"CR133\" class=\"CitationRef\"\u003e2017\u003c/span\u003e)d rdoba Province, and \u003cem\u003eN. cabrerai\u003c/em\u003e Zamorano et al. (\u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) in the PR (Buenos Aires Province).\u003c/p\u003e \u003cp\u003eConsequently, during the Late Miocene and Pliocene, NWA (Neotropical province \u003cem\u003esensu\u003c/em\u003e Barreda et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) and the PR (Proto-Spinal/Steppe province \u003cem\u003esensu\u003c/em\u003e Barreda et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) shared the same genera (\u003cem\u003eEleutherocercus\u003c/em\u003e, \u003cem\u003eStromaphorus, Nopachtus\u003c/em\u003e, and \u003cem\u003eEosclerocalyptus\u003c/em\u003e) but different species. More precisely, within the Neotropical province, some differences between NWA and NEA (North Eastern Argentina) seem possible, since glyptodonts from the \u0026ldquo;Mesopotamiense\u0026rdquo; (Late Miocene) include another species of \u003cem\u003eEleutherocercus\u003c/em\u003e (\u003cem\u003eE. paranensis\u003c/em\u003e), some putative species of \u0026ldquo;Palaehoplophorini\u0026rdquo; and other very poorly characterized taxa, mostly limited to isolated osteoderms and/or fragmented portions of the dorsal carapace (Scillato-Yan\u0026eacute; et al. \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Perhaps, the closed and forested biomes and tropical/subtropical conditions inferred for the Late Miocene of NEA (see Schmidt et al. \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) stimulated this difference between NW and NE regions. However, further studies with more complete materials from the Ituzaing\u0026oacute; Formation are necessary in order to infer the taxonomic diversity of glyptodonts in NEA.\u003c/p\u003e \u003cp\u003eA similar distribution of different species of \u003cem\u003eTremacyllus\u003c/em\u003e (Hegetotheriidae, Notoungulata) in NWA and PR was observed. Armella et al. (2022) interpreted these differences as related to paleoenvironmental and paleo-phytogeographic features. These patterns could be interpreted as disjunctive distribution following the criteria of Morrone and Escalante (\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) in which a barrier (rivers, mountains, vegetation, paleoecological differences) could have interrupted gene flow between different populations and derived in the allopatric speciation of these populations. In the case of the glyptodonts and taking into account the Neogene geologic history of NWA, these barriers could be related to tectonic and/or climatic-environmental changes. During the Late Miocene, NWA experienced a series of tectonic uplifts, which resulted in the elevation of the mountain chain of Northwestern Pampean and Aconquija Ranges (Bossi et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Georgieff et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), which probably developed a physical barrier for the distribution of the different groups of animals. In addition, by this time, one of the most important environmental changes is recorded in NWA, where plants with C4 photosynthetic pathway became dominant (Latorre et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Hynek et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Sanz-P\u0026eacute;rez et al. \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), perhaps in response to global temperatures gradually decreasing through the Late Miocene to the Early Pliocene (6 Ma) (Prevosti and Forasiepi \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). So, two scenarios, and probably not the only ones, can be considered.\u003c/p\u003e \u003cp\u003eIn this respect, the endemic diversity of glyptodonts observed in NWA is also recorded in other related clades of cingulates, the \u0026ldquo;armadillos\u0026rdquo; Chlamyphoridae, in which at least two species (\u003cem\u003ePaleuphractus argentinus\u003c/em\u003e (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e) Kraglievich, 1934 and \u003cem\u003eParaeuphractus prominens\u003c/em\u003e (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e) Scillato-Yane, 1975 are considered as endemic of this area (see Barasoain et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e). As mentioned above, this particular pattern observed in armadillos and glyptodonts could be explained by the tectonic activity of the uplift of the Andean mountains, resulting in a more temperate climate during the Late Miocene and Pliocene lapse, because the cordillera acted as a barrier for the Atlantic wet winds (Starck and Anz\u0026oacute;tegui \u003cspan citationid=\"CR120\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Barasoain et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e). Moreover, palaeobotanical evidence from the late Neogene sequence of NWA indicates warmer conditions with a marked seasonality (Anz\u0026oacute;tegui et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Candela et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBesides, the stratigraphic distribution of the Plohophorini glyptodonts of PR (ie. \u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e and \u003cem\u003eP. figuratus\u003c/em\u003e) from the Monte Hermoso Formation (ca. 4.7\u0026ndash;3.7 Ma; see Prevosti et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) mostly coincides with a warm and humid pulse recorded during the Pliocene (ca. 4.7\u0026ndash;3.1 Ma) (Haug et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Domingo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Prevosti et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), and a similar scenario can be inferred for both species of \u003cem\u003ePseudoplohophorus\u003c/em\u003e from the Camacho Formation (ca. 7.2-6 Ma) in Uruguay (Del R\u0026iacute;o et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Aumond et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Perea et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In the PR, the records of Plohophorini (\u003cem\u003ePlohophorus figuratus\u003c/em\u003e) continue through the Chapadmalalan (Zanclean and Piacenzian, ca. 3.7\u0026ndash;3.04 Ma; see Zurita et al. \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e; Prevosti et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and Marplatan (\u003cem\u003eP. avellaneda\u003c/em\u003e) (Piacenzian, ca 2.8 Ma) ages under mostly wet and warm conditions. According to the fossil evidence, records belonging to this tribe are among the most frequent in glyptodonts, proving that the populations were healthy in that period (see Zurita et al. \u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e). The last records of Plohophorini in the PR, at ca. 2.53 Ma, (see Qui\u0026ntilde;ones et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and at ca. 3.66 Ma in NWA (this work) coincides with a global cooling generated by the expansion of ice in Antarctica at ~\u0026thinsp;3.0 and 2.7 Ma (see Domingo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Prevosti et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In addition, these last records chronologically overlap with the arrival, from lower and middle latitude, of the other large clade of Glyptodontidae, the Glyptodontinae (Cuadrelli, 2020).\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe results of the present contribution indicate that: (1) The names \u0026ldquo;\u003cem\u003eStromaphorus compressidens\u003c/em\u003e\u0026rdquo; (Moreno and Mercerat, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1891\u003c/span\u003e) and \u0026ldquo;\u003cem\u003ePhlyctaenopyga\u003c/em\u003e\u0026rdquo; \u003cem\u003eameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e) refer to the same species; therefore, a synonymy is established with \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; ex Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e) representing the only Plohophorini glyptodont in the late Neogene sequence of NWA. (2) The cladistic analysis confirms that the genera \u003cem\u003ePseudoplohophorus\u003c/em\u003e, \u003cem\u003ePlohophorus\u003c/em\u003e and \u003cem\u003eStromaphorus\u003c/em\u003e are monophyletic, being Plohophorini a well-supported clade within Hoplophorinae. (3) The location of the enigmatic species \u003cem\u003eNopachtus coagmentatus\u003c/em\u003e within Hoplophorini lineage indicates that the spine-like structure of the caudal tube must be interpreted as a homologous structure rather than a convergence. On the contrary, the multiplication of peripheral figures in both lineages (i.e., Hoplophorini and Plohophorini) is a homoplasy. (4) From a stratigraphic point of view, the FAD of \u003cem\u003eS. ameghini\u003c/em\u003e is not precise since it could be established between ca.7 or 9 Ma; however, regardless of the discrepancy in this numerical range, a minimum age is established for the Plohophorini lineage, being the records of NWA older than those of the PR, where the diversity achieved by this clade was larger. The certain stratigraphic distribution of \u003cem\u003eS. ameghini\u003c/em\u003e in the NWA is ca. 7.14\u0026ndash;3.66 Ma, while the distribution of Plohophorini (ie, \u003cem\u003ePlohophorus\u003c/em\u003e spp. and \u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e) in the PR is ca. 4.7\u0026ndash;2.53 Ma. (5) The same genera of glyptodonts have different species in NWA and PR; this differentiation is probably related to the different palaeophytogeographic pattern. \u003cem\u003eS. trouessarti nov. comb.\u003c/em\u003e, \u003cem\u003eP. figuratus\u003c/em\u003e, \u003cem\u003eE. antiquus\u003c/em\u003e, \u003cem\u003eEo. lineatus\u003c/em\u003e and \u003cem\u003eN. cabrerai\u003c/em\u003e were geographically restricted to the Proto-Espinal/Steppe province. In turn, the distribution of the species \u003cem\u003eS. ameghini\u003c/em\u003e, \u003cem\u003eE. solidus\u003c/em\u003e, \u003cem\u003eEo. proximus\u003c/em\u003e and \u003cem\u003eN. coagmentatus\u003c/em\u003e coincides with the Neotropical province. Furthermore, this separation could have been accentuated by the presence of a geographical barrier (e.g. course of a large river, mountainous elevation, etc.). (6) The last record of Plohophorini at ca. 2.53 Ma in the PR coincides with a global cooling event very close to the end of the Pliocene and the beginning of the Pleistocene, while those of the NWA are somewhat older, in ca. 3.66 Ma.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Susana Bargo, Mart\u0026iacute;n L. de los Reyes and Marcelo Reguero (Museo de La Plata, La Plata, Argentina), Laura Cruz (Museo Argentino de Ciencias Naturales Bernardino Rivadavia, MACN, Buenos Aires, Argentina), William Simpson and Keneth Angielezyk (Field Museum of Natural History FMNH, Chicago, USA) and the Museo \u0026ldquo;Condor Huasi\u0026rdquo; Bel\u0026eacute;n Catamarca, Argentina, for granting us access to their collections; and Direcci\u0026oacute;n de Antropolog\u0026iacute;a de Catamarca for permitting work in this province. We also thank Gabriela Schmidt (CONICET-Prov. ER-UADER, Entre R\u0026iacute;os, Argentina), Johana Baez and Juan Manuel Robledo (CECOAL-CONICET-UNNE, Corrientes Capital, Argentina), Federico Degrange and Ivana Tapia (CICTERRA-CONICET-UNC, C\u0026oacute;rdoba, Argentina), Lucia M. Ib\u0026aacute;\u0026ntilde;ez and Matias A. Armella (Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucum\u0026aacute;n, Tucum\u0026aacute;n, Argentina), Mar\u0026iacute;a Carolina Madozzo Ja\u0026eacute;n (INSUGEO-CONICET-UNT, Tucum\u0026aacute;n, Argentina), for their help during fieldwork. Special mention to Marcos Roig for his help during the palaeobiogeographical analysis, Nicol\u0026aacute;s Bauz\u0026aacute; with phylogenetic morphometric analysis, Daniel Perea for the photographs provided, and Alvaro Mones and Cristina Luisa Scioscia for his help with nomenclatural issues. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eN\u0026uacute;\u0026ntilde;ez-Blasco A performed the geometric morphometric analysis and applied it to the phylogenetic analysis, and also prepared all the\u0026nbsp;online resource; N\u0026uacute;\u0026ntilde;ez-Blasco A., Zurita A. and Tori\u0026ntilde;o P. performed the phylogenetic analysis and the chronological calibration of the tree with R; N\u0026uacute;\u0026ntilde;ez-Blasco A. and Zurita A. E. made the taxonomical and systematic assignment; N\u0026uacute;\u0026ntilde;ez-Blasco A., Zurita A. E., Bonini R., Mi\u0026ntilde;o-Boilini A. R. and Tori\u0026ntilde;o P. wrote the introduction and\u0026nbsp;materials and methods\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003etexts and\u0026nbsp;Historical background\u0026nbsp;N\u0026uacute;\u0026ntilde;ez-Blasco A., Zurita A. E., Qui\u0026ntilde;ones S. I. and Zamorano M. wrote the anatomical descriptions; Nu\u0026ntilde;ez-Blasco A. and Bonini R. prepared figures 1-8.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eN\u0026uacute;\u0026ntilde;ez-Blasco A., Zurita A. E., Bonini R., Mi\u0026ntilde;o-Boilini A. R., Tori\u0026ntilde;o P. and Zamorano M. wrote the discussion and conclusion texts; Georgieff S. analysed the stratigraphy and relative dating of the fossils. All authors reviewed the manuscript.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was partially funded by\u0026nbsp;PICT 2019-03412, PICT 2018 003380, PI Q002/21 (SGCyT-UNNE), and SNI_2020_1_1010231 ANII (P.T.).\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data generated and analyzed during this study are included in this published article and its supplementary iles.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDisclosure statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNo potential conflict of interest was reported by the authors.\u0026nbsp;\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAmeghino F (1888a) \u003cem\u003eR\u0026aacute;pidas diagnosis de algunos mam\u0026iacute;feros f\u0026oacute;siles nuevos de la Rep\u0026uacute;blica Argentina\u003c/em\u003e. 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Hist Biol 29(8):1076-1088.\u003cu\u003e \u003c/u\u003ehttps://doi.org/10.1080/08912963.2016.1278443\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-mammalian-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jomm","sideBox":"Learn more about [Journal of Mammalian Evolution](http://link.springer.com/journal/10914)","snPcode":"10914","submissionUrl":"https://submission.nature.com/new-submission/10914/3","title":"Journal of Mammalian Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Glyptodontidae, Stromaphorus, Phlyctaenopyga, Late Miocene-Pliocene, diversity, evolutionary history","lastPublishedDoi":"10.21203/rs.3.rs-3914918/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3914918/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eNorthwestern Argentina (NWA) contains, together with the Pampean region (PR), one of the most complete late Neogene continental sequences, in which a great diversity of palaeofauna was recognized, among which glyptodonts stand out. Recent evidence suggests that the Late Miocene was a period of extra-Patagonian diversification in southern South America for glyptodonts, perhaps stimulated by the expansion of C4 grasses and open environments (known as \u0026ldquo;Edad de las Planicies Australes\u0026rdquo;). Here we focus on one of the most poorly known glyptodonts of NWA, the Plohophorini, from the Villavil-Quillay basin (Catamarca Province). Our results show that, like other clades (e.g., Doedicurini), a single species can be recognized, \u003cem\u003eStromaphorus ameghini\u003c/em\u003e (Ameghino, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1889\u003c/span\u003e; ex Moreno, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1882\u003c/span\u003e), whose stratigraphic record spans from the latest Miocene to the Pliocene (ca. 7.14\u0026ndash;3.3 Ma; Messinian-Zanclean). Cladistic analysis confirms the status of natural group of the tribe Plohophorini within Hoplophorinae (\u0026ldquo;austral clade\u0026rdquo;), in which \u003cem\u003eS. ameghini\u003c/em\u003e appears as the sister species of the Pampean species \u003cem\u003eS. trouessarti\u003c/em\u003e (Moreno, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1888\u003c/span\u003e) nov. comb. The oldest precise records of \u003cem\u003eS. ameghini\u003c/em\u003e (ca. 7.14 Ma) provide a minimum age for the Plohophorini lineage. The evidence suggests that the diversity of glyptodonts from the late Neogene of NWA is composed of endemic species, different from those of the PR, although both areas share the same genera, as observed in other mammalian clades such as Hegetotheriidae and Dasypodidae. Finally, the cladistic analysis reveals, in a broader context, that the spine-like structure observed in the caudal tube of some genera (ie, \u003cem\u003eNopachtus\u003c/em\u003e, \u003cem\u003ePropanochthus\u003c/em\u003e, and \u003cem\u003ePanochthus\u003c/em\u003e) is a homologous structure rather than a convergence as usually interpreted. On the contrary, the similar appearance of the ornamentation pattern represented by the multiplication of peripheral figures in the carapaces of the genera \u003cem\u003eStromaphorus\u003c/em\u003e and \u003cem\u003eNopachtus\u003c/em\u003e is, in fact, a convergence.\u003c/p\u003e","manuscriptTitle":"Plohophorini Glyptodonts (Xenarthra, Cingulata) From the Late Neogene of Northwestern Argentina. Insight Into Their Diversity, Evolutionary History, and Paleobiogeography","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-02-02 10:02:58","doi":"10.21203/rs.3.rs-3914918/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-04-02T13:06:17+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-03-19T22:30:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"494afc47-a3a0-460d-843c-84b8b2826a67","date":"2024-02-09T12:50:53+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"5e295169-12e8-499f-9204-98684a0b80be","date":"2024-02-05T14:25:29+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-02-05T14:00:48+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-02-01T03:41:37+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-02-01T03:41:37+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Mammalian Evolution","date":"2024-01-31T19:21:40+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-mammalian-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jomm","sideBox":"Learn more about [Journal of Mammalian Evolution](http://link.springer.com/journal/10914)","snPcode":"10914","submissionUrl":"https://submission.nature.com/new-submission/10914/3","title":"Journal of Mammalian Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"12b53575-f31c-4fcf-afef-a59d517b9516","owner":[],"postedDate":"February 2nd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-08-22T19:24:17+00:00","versionOfRecord":{"articleIdentity":"rs-3914918","link":"https://doi.org/10.1007/s10914-024-09726-3","journal":{"identity":"journal-of-mammalian-evolution","isVorOnly":false,"title":"Journal of Mammalian Evolution"},"publishedOn":"2024-08-17 15:56:58","publishedOnDateReadable":"August 17th, 2024"},"versionCreatedAt":"2024-02-02 10:02:58","video":"","vorDoi":"10.1007/s10914-024-09726-3","vorDoiUrl":"https://doi.org/10.1007/s10914-024-09726-3","workflowStages":[]},"version":"v1","identity":"rs-3914918","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3914918","identity":"rs-3914918","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}