diff --git a/data/03/82/87/038287CBFFC2FF8F79475229E85FB08A.xml b/data/03/82/87/038287CBFFC2FF8F79475229E85FB08A.xml new file mode 100644 index 00000000000..fe43d7fe639 --- /dev/null +++ b/data/03/82/87/038287CBFFC2FF8F79475229E85FB08A.xml @@ -0,0 +1,18741 @@ + + + +Woznikella triradiata n. gen., n. sp. - a new kannemeyeriiform dicynodont from the Late Triassic of northern Pangea and the global distribution of Triassic dicynodonts + + + +Author + +Szczygielski, Tomasz +Institute of Paleobiology, Polish Academy of sciences, Twarda 51 / 55, 00 - 818 Warsaw (Poland) +szczygielski@twarda.pan.pl + + + +Author + +Sulej, Tomasz +Institute of Paleobiology, Polish Academy of sciences, Twarda 51 / 55, 00 - 818 Warsaw (Poland) +sulej@twarda.pan.pl + +text + + +Comptes Rendus Palevol + + +2023 + +2023-05-16 + + +22 + + +16 + + +279 +406 + + + + +http://dx.doi.org/10.5852/cr-palevol2023v22a16 + +journal article +305805 +10.5852/cr-palevol2023v22a16 +96b937a1-56d3-4b29-93b2-5b859c159da7 +1777-571X +14260551 +urn:lsid:zoobank.org:pub:5BEBCC73-819E-47AE-9610-1E50B1AD6B60 + + + + + + +Woznikella triradiata + +n. gen., n. sp. + + + + + +( +Figs 1-16 +) + + + +urn:lsid:zoobank.org:act: +DACADFD8-C682-4872-82C0-7CB34025868D + + + + + + + +HOLOTYPE + +. — +ZPAL V. 34/1 +, partial skeleton + +. + + + + +DIAGNOSIS +. — A medium-sized stahleckeriid-related dicynodont with triradiate, Y-shaped branching pattern of premaxillary ridges, paired medial ridge on the mental surface of the dentary, slender scapula, wakly-angled scapular spine, well-demarcated acromion process projecting well beyond the border of the scapula, and coracoid foramen completely enclosed by the procoracoid, characterized by an autapomorphy: acromion directed anterodorsally rather than anteroventrally or anteriorly. Differs from non-kannemeyeriiform dicynodonts and + +Dinodontosaurus tener + +in presence of a distinct supinator process. Differs from most shansiodontids and stahleckeriids in the gracile scapular blade withsignificant terminal flaring of the scapula. Differs from more derived stahleckeriines in having a less pronounced supinator process. Differs from + +Ischigualastia jenseni +Cox, 1962 + +and + +Jachaleria candelariensis + +Araujo & Gonzaga, +1980 + + +in scapula not contributing to the coracoid foramen. Differs from + +Stahleckeria potens +von Huene, 1935 + +, + +Placerias hesternus +Lucas, 1904 + +, + +Eubrachiosaurus browni + +, and + +Zambiasaurus submersus + +Cox, +1969 + + +in having a weakly developed scapular spine. + + + + + +DERIVATIO +NOMINIS +. — +Woznikella + +in reference to the name of the +type +locality, Woźniki; + +triradiata + +in reference to the triradiate branching pattern of the premaxillary ridges. + + + +TYPE + + +LOCALITY + +. — Woźniki, southern +Poland +. + + + + + +TYPE + + +HORIZON + +. — The Patoka Member of the Grabowa Formation, Carnian ( + +Sulej +et al. +2011 + +, see + +Szulc +et al. +2015b + +for the arguments for the Norian age of the locality; see Discussion for the rationale for its Carnian age). + + +REFERRED +SPECIMEN +. — +SMNS +91416, a fragmentary mandible from the Carnian of Markt Obernzenn ( +Bavaria +, +Germany +), Stuttgart Formation (Shilfsandstein). + + + + +DISTRIBUTION +. — Carnian (and?Norian) of +Poland +and +Germany +. + + + + +DESCRIPTION + + +Skull and mandible + + +The preserved parts of the skull and mandible ( +Figs 1-6 +) lack pronounced rugosities, which may be either taxonomic or related to the young age of the individual. Based on the shape of the beak (both of the premaxilla and dentary) and the width of the preserved skull table elements, at least preorbitally the skull was apparently relatively narrow ( +Fig. 7 +). Unlike in, e.g., + +Kannemeyeria aganosteus +Kammerer & Ordoñez, 2021 + +, + +Kannemeyeria simocephalus + +, + +Kannemeyeria lophorhinus +Renaut, Damiani, Yates & Hancox, 2003 + +, + +Lystrosaurus +spp. + +(with the possible exception of + +L. youngi + +), + +Rechnisaurus cristarhynchus +Chowdhury, 1970 + +, + +Sangusaurus edentatus +Cox, 1969 + +, + +Sangusaurus parringtonii +Cruickshank, 1986a + +, + +Shaanbeikannemeyeria xilougouensis +Cheng, 1980 + +, + +Shansiodon wangi +Yeh, 1959 + +, + +Shansiodon wuhsiangensis +Yeh, 1959 + +, + +Stahleckeria potens + +, + +Sungeodon kimkraemerae +Maisch & Matzke, 2014 + +, + +Tetragonias njalilus +( +von Huene, 1942 +) + +, + +Ufudocyclops mukanelai + +, + +Vinceria andina +Bonaparte, 1969 + +, + +Wadiasaurus indicus +Chowdhury, 1970 + +, and + +Zambiasaurus submersus +Cox, 1969 + +there is no conspicuous median ridge on the skull ( +Weithofer 1888 +; +Broom 1899 +; +Haughton 1915 +; +Pearson 1924a +; +Case 1934 +; +Yuan & Young 1934b +; +Young 1935 +; +von Huene 1942 +; +Camp 1956 +; +Yeh 1959 +; +Sun 1964 +; +Cruickshank 1967 +; +Chowdhury 1970 +; +Crozier 1970 +; +Cluver 1971 +; +Kalandadze 1975 +; +Cheng 1980 +; +Cruickshank 1986a +; +Bandyopadhyay 1989 +; +Pickford 1995 +; +Schwanke-Peruzzo & Araújo-Barbarena 1995 +; +Maisch 2001 +; + +Renaut +et al. +2003 + +; + +Surkov +et al. +2005 + +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +; + +Grine +et al. +2006 + +; +Domnanovich & Marsicano 2012 +; + +Angielczyk +et al. +2014 + +; +Maisch & Matzke 2014 +; + +Angielczyk +et al. +2017 + +; + +Kammerer +et al. +2019 + +; +Kammerer & Ordoñez 2021 +). Note that the presence and degree of development of the median ridge is ontogeny-dependent at least in some dicynodont taxa (e.g., +Kammerer & Ordoñez 2021 +). + + + +FIG +. 1. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1/2: +A -E +, premaxillae in dorsal ( +A +), lateral right ( +B +), ventral ( +C +), lateral left ( +D +), and anterior ( +E +) view. Scale bar: 5 cm. + + + +Comparison of overall proportions of the mandiblular bones shows that the mandible of + +Woznikella triradiata + +n. gen., n. sp. +was proportionally significantly longer relative to its height than that of; e.g., + +Angonisaurus cruickshanki + +, + +Ischigualastia jenseni + +, + +Lystrosaurus +spp. + +, + +Myosaurus gracilis +Haughton, 1917 + +, + +Sangusaurus parringtonii + +, and + +Stahleckeria potens + +, and more reminiscent to that of, e.g., + +Kannemeyeria simocephalus + +, + +Sinokannemeyeria +spp. + +, or + +Wadiasaurus indicus + +(see, e.g., +Cox & Li 1983 +; +Pearson 1924a +; +von Huene 1935 +, +1936 +; +Young 1935 +; +Broom 1937 +; +Janensch 1952 +; +Camp 1956 +; +Sun 1963 +, +1964 +; +Cox 1965 +; +Cluver 1971 +, +1974 +; +Keyser 1974 +; +Cruickshank 1986a +; +Bandyopadhyay 1988 +; +Renaut 2000 +; + +Hancox +et al. +2013 + +; + +Angielczyk +et al. +2017 + +). + + +Premaxilla + + +The premaxilla ( +ZPAL +V +. 34/1/2; +Fig. 1 +) is fused and well preserved. Only the right side and the tip of the left side of the dorsal process as well as a small fragment of the posteriormost tip are broken. The anterior part is skewed to the left, but it is unclear whether this deformation is taphonomical or if it occurred during the life of the animal. The anterior tip in dorsoventral aspect is blunt (squared off), rather than sharply pointed as in, e.g., + +Kannemeyeria lophorhinus + +, apparently + +Moghreberia nmachouensis +Dutuit, 1980 + +, + +Placerias hesternus + +, “ + +Putillosaurus sennikovi +” +Surkov, 2005 + +, and + +Ufudocyclops mukanelai + +(see +Camp & Welles 1956 +; +Cox 1965 +; +Dutuit 1980 +, +1988 +; +Renaut 2000 +; +Surkov 2005 +; + +Angielczyk +et al. +2014 + +; + +Kammerer +et al. +2019 + +). Nonetheless, it appears to be narrower than in + +Dinodontosaurus brevirostris +Cox, 1968 + +, + +Dinodontosaurus tener +( +von Huene, 1935 +) + +, + +Dolichuranus primaevus +Keyser, 1973 + +, + +Jachaleria candelariensis + +, + +Rechnisaurus cristarhynchus + +, + +Sinokannemeyeria sanchuanheensis +Cheng, 1980 + +, and + +Stahleckeria potens + +, in which the bone is about as wide as it is long in dorsal view and shows nearly no rostral tapering ( + +Tupi +Caldas +1936 + +; +Cox 1965 +, +1968 +; +Chowdhury 1970 +; +Keyser 1973 +; +Araújo & Gonzaga 1980 +; +Cheng 1980 +; +Bandyopadhyay 1989 +; +Maisch 2001 +; +Vega-Dias & Schultz 2004 +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +; +Kammerer & Ordoñez 2021 +). Its edge and anterodorsal surface are slightly concave medially, so the beak is M-shaped in dorsoventral aspect, similar to + +Ischigualastia jenseni + +, + +Jachaleria candelariensis + +, + +Parakannemeyeria chengi +Liu, 2004 + +, + +Parakannemeyeria ningwuensis +Sun, 1963 + +, + +Parakannemeyeria youngi +Sun, 1963 + +, + +Rhadiodromus mariae +Surkov, 2003 + +, + +Uralokannemeyeria vjuschkovi +Danilov, 1971 + +, and + +Stahleckeria potens + +(see +Sun 1963 +; +Cox 1965 +; +Danilov 1971 +; +Araújo & Gonzaga 1980 +; +Maisch 2001 +; +Surkov 2003 +; +Liu 2004 +; +Vega-Dias & Schultz 2004 +; +Kammerer & Ordoñez 2021 +). In contrast to, e.g., + +Dinodontosaurus brevirostris + +, + +Dinodontosaurus tener + +, + +Ischigualastia jenseni + +(pers. obs.: PVSJ 545), + +Jachaleria candelariensis + +, + +Lystrosaurus +spp. + +(with the possible exception of + +L. georgi + +, + +L. hedini +Young, 1935 + +, and + +L. youngi + +), + +Parakannemeyeria ningwuensis + +, and + +Sinokannemeyeria yingchiaoensis +Sun, 1963 + +, there is no pronounced notch in the median part of the tomial edge (unless the tips of the beak of +ZPAL +V +. 34/1/2 are damaged; +Yuan & Young 1934b +; +Young 1935 +; +Sun 1963 +, +1964 +; +Cluver 1971 +; +Kalandadze 1975 +; +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 2004 +; +Surkov 2005 +; +Morato 2006 +; + +Grine +et al. +2006 + +; +Kammerer & Ordoñez 2021 +). + + +In lateral view ( +Fig. 1C +), accounting for deformation, the labial (tomial) edge of the premaxilla was apparently directed anteroventrally, roughly perpendicular to the anterodorsally facing surface of the nasal process. This differs from at least some specimens of + +Tetragonias njalilus + +, in which the labial edge of the beak was set at an obtuse angle to its anteroventrally facing surface and thus the tip of the beak was slightly recessed ( +Cruickshank 1967 +; +Kammerer & Ordoñez 2021 +). The lateral surface of the anterior half is coarsely rugose, indicative of a keratinous rhamphotheca ( +Fig 1B, D, E +). The premaxilla forms the anterior edge of the naris. The anterior outline of the narial fossa is straight in the upper part and reaches very close to the ventral edge of the premaxilla, similar to, e.g., + +Acratophorus argentinensis +( +Bonaparte, 1965 +) + +, + +Kannemeyeria simocephalus + +, + +Parakannemeyeria dolichocephala +Sun, 1960 + +, + +Parakannemeyeria ningwuensis + +, + +Parakannemeyeria youngi + +, “ + +Putillosaurus sennikovi + +”, + +Rechnisaurus cristarhynchus + +, + +Repelinosaurus robustus +Olivier, Battail, Bourquin, Rossignol, Steyer & Jalil, 2019 + +, + +Rhadiodromus mariae + +, + +Rhinodicynodon gracile +Kalanadze, 1970 + +, + +Rabidosaurus cristatus +Kalanadze, 1970 + +, the South African + +Shansiodon +sp. + +, + +Sinokannemeyeria pearsoni +Young, 1937 + +, + +Sinokannemeyeria sanchuanheensis + +, + +Ufudocyclops mukanelai + +, + +Vinceria andina + +, + +Wadiasaurus indicus + +, and + +Xiyukannemeyeria brevirostris +( +Sun, 1978 +) + +(see +Haughton 1915 +; +Pearson 1924a +; +Case 1934 +; +Watson 1948 +; +Sun 1960 +, +1963 +; +Cruickshank 1965 +; + +Bonaparte 1966 +a, 1967 + +; +Chowdhury 1970 +; +Kalandadze 1970 +; +Cheng 1980 +; +Bandyopadhyay 1988 +, +1989 +; +Renaut 2000 +; +Renaut & Hancox 2001 +; +Liu & Li 2003 +; +Surkov 2003 +, +2005 +; +Domnanovich & Marsicano 2012 +; + +Hancox +et al. +2013 + +; + +Kammerer +et al. +2019 + +; + +Olivier +et al. +2019 + +; +Kammerer & Ordoñez 2021 +). The premaxilla differs dramatically in proportions from the anteroposteriorly short and vertically elongated premaxillae of + +Lystrosaurus +spp. + +( +Yuan & Young 1934b +; +Young 1935 +; +Sun 1964 +; e.g., +Cluver 1971 +; +Colbert 1974 +; +Kalandadze 1975 +; + +Surkov +et al. +2005 + +; + +Grine +et al. +2006 + +). On the ventral (palatal) surface of the premaxilla, each of the anteriorly pronounced tips of the beak projects rearward to form a predominantly posteriorly and gently medially directed, rugose ridge ( +Fig. 1C +). Anteriorly, these ridges are separated by the median palatal groove, but merge at approximately 2/3 of the premaxilla’s length into a single, higher, posteriorly directed median (posterior) palatal ridge, resulting in a Y-shaped structure, similar to most Triassic dicynodonts with the exception of + +Kombuisia frerensis +Hotton, 1974 + +and + +Myosaurus gracilis + +, which lack the anterior ridges ( +Cluver 1974 +; +Hotton 1974 +; +Fröbisch 2007 +). There are no lateral anterior ridges comparable to those of + +Kombuisia frerensis + +(see +Hotton 1974 +). The anterior ridges in + +Woznikella triradiata + +n. gen., n. sp. +are more widely separated than in, e.g., + +Jachaleria canderlariensis + +and + +Kannemeyeria lophorhinus + +( +Crozier 1970 +; +Araújo & Gonzaga 1980 +; +Pickford 1995 +; +Renaut 2000 +; +Vega-Dias & Schultz 2004 +; + +Angielczyk +et al. +2014 + +). Unlike in, e.g., + +Acratophorus argentinensis + +, + +Angonisaurus cruickshanki +Cox & Li, 1983 + +, + +Counillonia superoculis +Olivier, Battail, Bourquin, Rossignol, Steyer & Jalil, 2019 + +, “ + +Cristonasus koltaeviensis + +”, + +Dolichuranus primaevus + +, + +Kannemeyeria simocephalus + +, + +Kannemeyeria lophorhinus + +, “ + +Kannemeyeria +” +latirostris +Crozier, 1970 + +, + +Jachaleria candelariensis + +, + +Parakannemeyeria dolichocephala + +, “ + +Putillosaurus sennikovi + +”, + +Rechnisaurus cristarhynchus + +, + +Repelinosaurus robustus + +, + +Sangusaurus edentatus + +, + +Sangusaurus parringtonii + +, + +Sinokannemeyeria yingchiaoensis + +, + +Ufudocyclops mukanelai + +, + +Uralokannemeyeria vjuschkovi + +, and + +Zambiasaurus submersus + +, the ridges diverge anteriorly rather than being parallel ( +Pearson 1924a +; +Case 1934 +; +Toerien 1953 +; +Sun 1960 +, +1963 +; +Bonaparte 1966a +; +Cox 1969 +; +Chowdhury 1970 +; +Crozier 1970 +; +Danilov 1971 +; +Keyser 1973 +; +Araújo & Gonzaga 1980 +; +Cox & Li 1983 +; +Bandyopadhyay 1989 +; +Pickford 1995 +; +Schwanke-Peruzzo & Araújo-Barbarena 1995 +; +Surkov 1999a +; +Renaut 2000 +; +Vega-Dias & Schultz 2004 +; +Surkov 2005 +; + +Damiani +et al. +2007 + +; + +Hancox +et al. +2013 + +; + +Angielczyk +et al. +2014 + +, +2017 +; + +Kammerer +et al. +2019 + +; + +Olivier +et al. +2019 + +; +Kammerer & Ordoñez 2021 +). They nearly reach the anterolateral corners of the beak, unlike, e.g., in + +Acratophorus argentinensis + +, + +Dinodontosaurus +spp. + +, + +Jachaleria candelariensis + +, + +Lystrosaurus +spp. + +, + +Rechnisaurus cristarhynchus + +, + +Shaanbeikannemeyeria xilougouensis + +, + +Stahleckeria potens + +, + +Vinceria andina + +, and + +Wadiasaurus indicus + +, in which the snout stretches out further laterally ( +von Huene 1935 +; +Toerien 1953 +; +Cox 1965 +, +1968 +; +Bonaparte 1966a +; +Chowdhury 1970 +; +Cluver 1971 +; +Cheng 1980 +; +Araújo & Gonzaga 1980 +; +Bandyopadhyay 1988 +, +1989 +; +Renaut 2000 +; +Renaut & Hancox 2001 +; +Vega-Dias & Schultz 2004 +; +Morato 2006 +; +Domnanovich & Marsicano 2012 +; + +Abdala +et al. +2013 + +; +Kammerer & Ordoñez 2021 +). The three ridges are continuous with each other and the point of their branching is well demarcated and ventrally convex, in contrast to + +Jachaleria candelariensis + +, + +Kannemeyeria lophorhinus + +, + +Rabidosaurus cristatus + +(pers. obs.), and + +Rhinodicynodon gracile + +, in which the posterior (median) ridge is flanked by the anterior ones but not connected to them ( +Crozier 1970 +; +Araújo & Gonzaga 1980 +; +Pickford 1995 +; +Renaut 2000 +; +Vega-Dias & Schultz 2004 +; + +Angielczyk +et al. +2014 + +; +Kammerer & Ordoñez 2021 +) or, e.g., + +Acratophorus argentinensis + +, + +Angonisaurus cruickshanki + +, + +Counillonia superoculis + +, + +Dolichuranus primaevus + +, “ + +Kannemeyeria +” +latirostris + +, + +Lystrosaurus +spp. + +, + +Parakannemeyeria dolichocephala + +, + +Placerias hesternus + +, + +Rechnisaurus cristarhynchus + +, + +Repelinosaurus robustus + +, + +Sangusaurus parringtonii + +, + +Shaanbeikannemeyeria xilougouensis + +, the South African + +Shansiodon +sp. + +, + +Sinokannemeyeria baidaoyuensis +Liu, 2015 + +, + +Ufudocyclops mukanelai + +, + +Vinceria andina + +, + +Wadiasaurus indicus + +, and + +Zambiasarus submersus + +, in which the ridges are separate and the branching point is indistinct ( +Camp 1956 +; +Camp & Welles 1956 +; +Sun 1960 +; +Cox 1965 +, +1969 +; +Crozier 1970 +; +Chowdhury 1970 +; +Cluver 1971 +; +Keyser 1973 +; +Cheng 1980 +; +Cox & Li 1983 +; +Bandyopadhyay 1988 +, +1989 +; +Renaut 2000 +; +Renaut & Hancox 2001 +; + +Damiani +et al. +2007 + +; +Domnanovich & Marsicano 2012 +; + +Hancox +et al. +2013 + +; + +Angielczyk +et al. +2014 + +, +2017 +; +Liu 2015 +; + +Kammerer +et al. +2019 + +; + +Olivier +et al. +2019 + +; +Kammerer & Ordoñez 2021 +). In + +Kannemeyeria simocephalus + +, both of the latter morphologies can be observed, but the lateral and median palatal ridges never seem to form a well-defined connection ( +Pearson 1924a +; +Case 1934 +; +Toerien 1953 +; +Renaut 2000 +). This character is usually poorly presented in published figures, but apparently the only Triassic dicynodonts with such a well-defined triradiate branching pattern of the premaxillary ridges comparable to that of + +Woznikella triradiata + +n. gen., n. sp. +Are + +Dinodontosaurus brevirostris + +and + +Dinodontosaurus tener + +(see +Cox 1965 +, +1968 +; +Morato 2006 +; +Kammerer & Ordoñez 2021 +), + +Ischigualastia jenseni + +(see +Kammerer & Ordoñez 2021 +; pers. obs.: PVSJ 545), + +Kannemeyeria aganosteus + +(see +Kammerer & Ordoñez 2021 +), + +Rhadiodromus mariae + +(see +Surkov 2003 +), + +Tetragonias njalilus + +( +von Huene 1942 +; +Cruickshank 1967 +; + +Hancox +et al. +2013 + +), + +Uralokannemeyeria vjuschkovi + +(see +Danilov 1971 +), and the specimens illustrated by +Camp (1956) +as a juvenile + +Kannemeyeria +“ +vanhoepeni + +” and by +Cruickshank (1965) +as a juvenile + +Kannemeyeria +“ +latifrons + +” +Broom 1899 +. In the latter two cases this is based only on the interpretative drawings – this morphology is not clearly described, and no photographs of these specimens were published, so the accuracy of the drawings cannot be verified. In + +Ufudocyclops mukanelai + +, the anterior and posterior ridges do not form a connection, but the surface between them is slightly convex ( + +Hancox +et al. +2013 + +; + +Kammerer +et al. +2019 + +). As preserved, the anteriormost parts of these ridges are slightly deeper than the lateral labial edge, and thus visible in lateral view, in contrast to, e.g., + +Placerias hesternus + +(see +Camp & Welles 1956 +; +Cox 1965 +). The area for the connection with the palatines and vomers is very poorly preserved, so it is not possible to establish the anterior extent of those bones. The dorsal surface of the palatal plate is convex and a single longitudinal ridge spans along its entire midline ( +Fig. 1A +). Large parts of the surfaces on either side of this ridge probably served as facets for the maxillae. These facets are outlined with gentle, anterodorsally aligned, posterodorsally convex grooves, suggesting that the maxillae nearly reached anteriorly the front of the narial fossa, spanning along approximately posterior two thirds of the premaxillae length ( +Fig. 1B, D +). This is relatively further than in, e.g., “ + +Nasoplanites danilovi + +”, + +Placerias hesternus + +, and “ + +Putillosaurus sennikovi + +”, but comparable to + +Kannemeyeria simocephalus + +(see +Pearson 1924a +; +Camp & Welles 1956 +; +Surkov 1999a +; +Renaut 2000 +; +Surkov 2005 +). Both premaxillae are damaged in a way that obscures the connections with the septomaxillae. + + + +Nasal + + + +The right nasal ( +ZPAL +V +. 34/1/42; +Fig. 2 +A-D) is preserved almost completely. It consists of two parts, lateral and dorsal. The bone is slightly deformed due to compaction, which is clearly visible in the anteromedial part, which is concave dorsally (not corresponding with the alignment of the preserved part of the dorsal process of the premaxilla, thus unlikely to represent nasal depressions; compare to, e.g., +Chowdhury 1970 +; +Bandyopadhyay 1989 +; +Kammerer & Ordoñez 2021 +). The deformation also likely affected the angle between the lateral and dorsal plates ( +Fig. 2B +). Unlike in, e.g., “ + +Planitorostris pechoriensis + +” (see +Surkov 1999b +), the lateral part has a coarsely rugose surface (although the development of the rugosity is mild in comparison with many Kannemeyeriiform taxa) with numerous large vascular openings. It suggests that + +Woznikella triradiata + +n. gen., n. sp. +had either a sensitive tip of the snout or keratinous covering in that area (e.g., +Keyser & Cruickshank 1979 +; +Surkov 2003 +, +2006 +; + +Morato +et al. +2005 + +; +Morato 2006 +; + +Benoit +et al. +2018 + +). The lateral surface of the nasal is roughly even with the lateral edge of the dorsal table and is vertical and mostly flat, without prominent lateral depression ( +Fig. 2D +), unlike in, e.g., some specimens of + +Dinodontosaurus brevirostris + +and + +Dinodontosaurus tener + +, + +Rhadiodromus klimovi +( +Efremov, 1938 +) + +, + +Rhadiodromus mariae + +, + +Rhinodicynodon gracile + +, + +Sinokannemeyeria yingchiaoensis + +, and + +Xiyukannemeyeria brevirostris + +( +Sun 1963 +, 1978; +Cox 1965 +, +1968 +; +Vjuschkov 1969 +; +Kalandadze 1970 +; +Surkov 2003 +; +Morato 2006 +). This may imply the absence of a postnarial depression (although the lack of maxilla makes it uncertain). The ventral edge of the lateral part is sinusoidal. Two ventral processes, which formed the sutural connection with the lacrimal, have distinct ridges on the lateral surfaces. A shallow depression, which is probably the area for the suture with the dorsal process of the premaxilla, is visible on the dorsal surface ( +Fig. 2A +). The edge of this sutural field is directed posteromedially and it spans for about half the length of the dorsal plate of the nasal. Aside from that, there are no pronounced grooves comparable with those described in, e.g., “ + +Calleonasus furvus + +” or “ + +Elatosaurus facetus + +” (see +Kalandadze & Sennikov 1985 +; +Surkov 1999a +). The contact with the septomaxilla was apparently very small, only at the edge of the naris, due to most of the ventral edge of the nasal being occupied by the lacrimal. The contact with the prefrontal is slightly broken and difficult to interpret, but at least part of the facet for the prefrontal is preserved in the posterior third of the bone. As preserved, the medial portions of the nasals and frontals are separated by a diamond-shaped empty space, which was filled either by a short, pointy median anterior process of the frontal, or by a supernumerary internasal bone ( +Fig. 7A, B +), as in + +Jachaleria +spp. + +or some individuals of + +Lystrosaurus +spp. + +( +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 2004 +; + +Jasinoski +et al. +2014 + +; +Kammerer & Ordoñez 2021 +). The nasal roofed the naris, and its anterior edge formed the thick posterodprsal edge of the nostril ( +Fig. 2B, C +), similar to, e.g., + +Dolichuranus primaevus + +, + +Acratophorus argentinensis + +, + +Kannemeyeria lophorhinus + +, + +Kannemeyeria simocephalus + +, + +Parakannemeyeria dolichocephala + +, + +Parakannemeyeria ningwuensis + +, + +Rabidosaurus cristatus + +, the South African + +Shansiodon +sp. + +, + +Sinokannemeyeria pearsoni +, +Sinokannemeyeria yingchiaoensis + +, + +Sungeodon kimkraemerae + +, + +Ufudocyclops mukanelai + +, + +Uralokannemeyeria vjuschkovi + +, and + +Vinceria andina + +( +Pearson 1924a +; +Case 1934 +; +Sun 1960 +, +1963 +; + +Bonaparte 1966 +a, 1967 + +, +1969 +; +Kalandadze 1970 +; +Danilov 1971 +; +Keyser 1973 +; +Pickford 1995 +; +Renaut 2000 +; +Maisch 2001 +; +Renaut & Hancox 2001 +; + +Damiani +et al. +2007 + +; +Domnanovich & Marsicano 2012 +; + +Hancox +et al. +2013 + +; +Maisch & Matzke 2014 +; + +Angielczyk +et al. +2014 + +; + +Kammerer +et al. +2019 + +; +Kammerer & Ordoñez 2021 +). + + + +FIG +. 2. — + +Woznikella triradiata + +n. gen., n. sp. +: +A -D +, ZPAL V. 34/1/42, right nasal in dorsal ( +A +), ventral ( +B +), anterior ( +C +), and lateral ( +D +) view; +E -H +, ZPAL V. 34/1/80, right lacrimal in lateral ( +E +), medial ( +F +), posterior ( +G +), and ventral ( +H +) view. Scale bar: 5 cm. + + + + +Lacrimal + + + +The right lacrimal ( +ZPAL +V +. 34/1/80; +Fig. 2 +E-H)) is almost complete, only a small triangular fragment is missing in the middle of the posterior edge. The bone is subtriangular, approximately as high as it is long, and thus unusually equidimensional compared to other dicynodonts. The lateral surface is flat and the sutural surfaces are prominent ( +Fig. 2E +). The ventral edge is the most massive, with a triangular surface for the contact with the maxilla and, possibly the jugal and/ or palatine ( +Fig. 2H +). Unlike, e.g., + +Placerias hesternus + +, there is no conspicuous jugal process ( +Camp & Welles 1956 +). The anteroventral orbit margin is formed by a thin lappet of bone. The suture for the prefrontal is inverted V-shaped. Internally, a ridge is visible in the posterior part of the bone ( +Fig. 2F +). This ridge is most distinctive in the ventral part, where it floors and encases anteriorly a pocket-like lacrimal pit, which opens posteriorly as a large lacrimal foramen ( +Fig. 2G +). The ridge continues faintly along the posterior edge towards the dorsal part of the bone. The anterior opening of the lacrimal duct is located on the internal surface, in the anterior part of the bone, and is directed anterodorsally. The specimen articulates well with the preserved nasal ( +ZPAL +V +. 34/1/42) and based on that articulation it appears that the lacrimal nearly reached the naris, preventing or nearly preventing the contact between the nasal and the maxilla – the very posterior edge of the naris was formed by another bone, but it is currently impossible to establish whether this was maxilla or septomaxilla ( +Fig. 7 +). The anterior extent of the lacrimal, which is usually covered by the more superficial bones, is unknown in many other Triassic dicynodonts, but it is similar to that of + +Woznikella triradiata + +n. gen., n. sp. +at least in + +Ischigualastia jenseni + +, + +Jachaleria candelariensis + +, + +Kannemeyeria simocephalus + +, + +Parakannemeyeria dolichocephala + +, + +Parakannemeyeria shenmuensis +Cheng, 1980 + +, + +Parakannemeyeria youngi + +, + +Placerias hesternus + +, + +Rhadiodromus mariae + +, + +Shaanbeikannemeyeria xilougouensis + +, possibly + +Shansiodon wangi + +, the South African + +Shansiodon +sp. + +, + +Stahleckeria potens + +, + +Sinokannemeyeria baidaoyuensis + +, + +Sinokannemeyeria pearsoni + +, + +Sinokannemeyeria sanchuanheensis + +, + +Sungeodon kimkraemerae + +, and + +Vinceria andina + +, in which the lacrimal reaches or nearly reaches the nostril ( +Pearson 1924a +; +von Huene 1935 +; +Camp 1956 +; +Camp & Welles 1956 +; +Yeh 1959 +; +Sun 1960 +, +1963 +; +Cox 1965 +; +Bonaparte 1969 +; +Cheng 1980 +; +Araújo & Gonzaga 1980 +; +Hancox 1998 +; +Renaut 2000 +; +Renaut & Hancox 2001 +; +Surkov 2003 +; +Vega-Dias & Schultz 2004 +; +Domnanovich & Marsicano 2012 +; + +Abdala +et al. +2013 + +; + +Hancox +et al. +2013 + +; +Maisch & Matzke 2014 +; +Liu 2015 +; +Kammerer & Ordoñez 2021 +). In contrast to + +Kannemeyeria simocephalus + +, + +Parakannemeyeria youngi + +(and, likely, the other species of that genus), + +Placerias hesternus + +, and + +Stahleckeria potens + +, however, except for a narrow margin along the dorsoanterior and anterior border, which shows characteristics of a squamous suture, the lacrimal in + +Woznikella triradiata + +n. gen., n. sp. +was almost entirely exposed externally ( +Pearson 1924a +; +von Huene 1935 +; +Camp 1956 +; +Sun 1963 +; +Renaut 2000 +; +Maisch 2001 +; + +Abdala +et al. +2013 + +; +Kammerer & Ordoñez 2021 +). In + +Dinodontosaurus brevirostris + +and + +Dinodontosaurus tener + +the morphology of the lacrimal is variable ( +Cox 1965 +, +1968 +; +Morato 2006 +). In the specimens MCZ 1628, MCZ 3454, DGM 309, and the “ + +Chanaria platyceps + +” +holotype +(UNLaR 14) the lacrimal has a large exposure, unlike in the specimens MCZ 1670 and MCZ 1687 of the same species. However, in MCZ 1628 it is subrectangular and still significantly separated from the nostril, which is located in the anterior third of the narial fossa, whereas in MCZ 3454, DGM 309, and UNLaR 14 it reaches further rostrally ( +Cox 1965 +, +1968 +). In + +Dinodontosaurus brevirostris +MCZ + +3454 and likely UNLaR 14, as well as in + +Dinodontosaurus tener +DGM + +309, + +Parakannemeyeria youngi + +, + +Sinokannemeyeria baidaoyuensis + +, and + +Sinokannemeyeria pearsoni + +the lacrimal meets the septomaxilla anteriorly, separating the maxilla from the nasal, but the septomaxilla seems to cover about third of this distance ( +Sun 1963 +; +Cox 1968 +; +Liu 2015 +), thus more than in + +Woznikella triradiata + +n. gen., n. sp. +In + +Dinodontosaurus tener + +specimen DGM 530R the separation is incomplete and the lacrimal is relatively small, probably due to relatively short preorbital length of that skull ( +Cox 1968 +). Variable lacrimals, generally large but not separating the nasals from the maxillae, were observed in the series of specimens described by +Morato (2006) +. Some age-related variation was also described in + +Kannemeyeria simocephalus + +by +Renaut (2000) +, with larger individuals having proportionally smaller exposure of the lacrimal, but in no case was the anterior part of the lacrimal externally exposed +in vivo +in that species. In + +Jachaleria candelariensis + +the lacrimal, as exposed laterally, is much lower dorsoventrally and more strap-like ( +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 2004 +). The nasal is also completely or almost completely separated from the maxilla by the lacrimal and septomaxilla in + +Lystrosaurus +spp. + +and the South African + +Shansiodon +sp. + +, but due to the shortening of the preorbital part of the skull, the lacrimal remains relatively short (e.g., +Sun 1964 +; +Cluver 1971 +; +Li 1988 +; + +Hancox +et al. +2013 + +). + + + +FIG +. 3. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V.34/1/1: +A -E +, right frontal in dorsal ( +A +), ventral ( +B +), medial ( +C +), anterior ( +D +), and lateral ( +E +) view.Scale bar: 5 cm. + + + + +Frontal + + + +The right frontal ( +ZPAL +V +. 34/1/1; +Fig. 3 +) is almost complete, only the top of the dorsal (preparietal) process, a small fragment of the medial edge, and the anterior tip are broken. The frontal plate is nearly as wide as long and its anterior edge is bowed. The frontal pair thus was fan-shaped in dorsal view and apparently lacked prominent and narrow lateral processes or conspicuous embayments for the nasal or prefrontals, similar to, e.g., + +Dolichuranus primaevus + +, + +Lystrosaurus +spp. + +(with the exception of + +L. curvatus +( +Owen, 1876 +) + +and + +L. hedini + +), the South African + +Shansiodon +sp. + +, + +Sinokannemeyeria sanchuanheensis + +, and + +Tetragonias njalilus + +, but different from, e.g., + +Acratophorus argentinensis + +, + +Dinodontosaurus +spp. + +, + +Elephantosaurus jachimovitschi +Vjuschkov, 1969 + +, + +Ischigualastia jenseni + +, + +Jachaleria candelariensis + +, “ + +Kannemeyeria +” +latirostris + +, + +Moghreberia nmachouensis + +, + +Parakannemeyeria dolichocephala + +, + +Parakannemeyeria youngi + +, “ + +Parvobestiola bashkiriensis + +”, + +Placerias hesternus + +, + +Rechnisaurus cristarhynchus + +, + +Rhadiodromus mariae + +, + +Rhinodicynodon gracile + +, + +Shaanbeikannemeyeria xilougouensis + +, + +Sinokannemeyeria yingchiaoensis + +, + +Stahleckeria potens + +, + +Ufudocyclops mukanelai + +, and + +Vinceria andina + +( +von Huene 1935 +, +1942 +; +Young 1935 +; +Camp & Welles 1956 +; +Sun 1960 +, +1963 +; +Cox 1965 +, +1968 +; +Bonaparte 1966a +; +Cruickshank 1967 +; +Vjuschkov 1969 +; +Crozier 1970 +; +Kalandadze 1970 +; +Chowdhury 1970 +; +Cluver 1971 +; +Cheng 1980 +; +Araújo & Gonzaga 1980 +; +Dutuit 1988 +; +Bandyopadhyay 1989 +; +Lucas & Harris 1996 +; + +Surkov 1999 +a, 2003 + +; +Maisch 2001 +; +Vega-Dias & Schultz 2004 +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +; + +Damiani +et al. +2007 + +; +Domnanovich & Marsicano 2012 +; + +Abdala +et al. +2013 + +; + +Hancox +et al. +2013 + +; +Kammerer 2018 +; + +Kammerer +et al. +2019 + +; +Kammerer & Ordoñez 2021 +). The anterior median process is broken but, considering that the posterior edge of the nasal appears to be complete, it likely either was relatively short and ended in a point, or a supernumerary internasal bone was present, as in + +Jachaleria +spp. + +and some individuals of + +Lystrosaurus +spp. + +( +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 2004 +; + +Jasinoski +et al. +2014 + +; +Kammerer & Ordoñez 2021 +). It must be noted, however, that the layout of the anterior frontal suture exhibits some variability, even bilaterally within a single individual (e.g., +Camp 1956 +; +Cox 1965 +; +Renaut 2000 +; +Kammerer & Ordoñez 2021 +). The frontal is proportionally much wider than in the narrow-roofed skulls of + +Counillonia superoculis + +, + +Myosaurus gracilis + +, + +Kombuisia frerensis + +, + +Kombuisia antarctica +Fröbisch, Angielczyk & Sidor, 2010 + +, and + +Repelinosaurus robustus + +(see +Haughton 1917 +; +Cluver 1974 +; +Hotton 1974 +; +Hammer & Cosgriff 1981 +; +Fröbisch 2007 +; + +Olivier +et al. +2019 + +). It is much longer than the anteroposteriorly shortened (at least, as exposed externally) frontal of + +Sangusaurus parringtonii + +(see + +Angielczyk +et al. +2017 + +; +Kammerer & Ordoñez 2021 +). + + +The most distinctive feature of the frontal is the strongly elevated dorsal process similar to that of + +Kannemeyeria simocephalus + +(e.g., +Weithofer 1888 +; +Broom 1937 +; +Watson 1948 +; +Kammerer & Ordoñez 2021 +). On the anterior and lateral surfaces of this process, the area for sutural contact with the postorbital is visible. This area continues throughout the posterodorsal edge of the bone, reaching its lateral edge, where it ends in a point. + + +The lateral edge of the frontal contributed to the margin of the orbit, and in dorsoventral aspect is slightly oblique to the long axis of the skull (turned anteromedially). The contribution is restricted, similar as in, e.g., + +Dolichuranus primaevus + +and + +Rechnisaurus cristarhynchus + +, but larger than in at least some specimens of + +Dinodontosaurus +spp. + +, + +Ischigualastia jenseni + +, “ + +Kannemeyeria +” +latirostris + + +, +Stahleckeria potens + +, and + +Uralokannemeyeria vjuschkovi + +, and smaller than in most specimens of + +Kannemeyeria simocephalus + +, + +Kombuisia +spp. + +, and + +Myosaurus gracilis + +(see +Haughton 1917 +; +Pearson 1924a +; +Case 1934 +; +von Huene 1935 +; +Camp 1956 +; +Cox 1965 +, +1968 +; +Crozier 1970 +; +Danilov 1971 +; +Cluver 1974 +; +Hammer & Cosgriff 1981 +; +Bandyopadhyay 1989 +; +Renaut 2000 +; +Maisch 2001 +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +; + +Damiani +et al. +2007 + +; +Fröbisch 2007 +; + +Fröbisch +et al. +2010 + +; + +Abdala +et al. +2013 + +; +Kammerer & Ordoñez 2021 +). This differs from + +Jachaleria candelariensis + +, in which the frontal is excluded from the edge of the orbit by the prefrontal and postorbital ( +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 2004 +). In anterior view, the part of the skull roof formed by the frontal has its medial part noticeably more elevated dorsally than the lateral one, so this part of the skull was convex, similar to, e.g., + +Ischigualastia jenseni + +(see +Cox 1965 +; +Kammerer & Ordoñez 2021 +). + + + +FIG +. 4. — + +Woznikella triradiata + +n. gen.,n. sp. +: +A -E +, ZPAL V.34/1/3:dentaries in dorsal ( +A +), ventral ( +B +), lateral left ( +C +), anterior ( +D +), and lateral right ( +E +) view; +F -J +, SMNS 91416: dentaries in dorsal ( +F +), ventral ( +G +), lateral left ( +H +), anterior ( +I +), and lateral right ( +J +) view. Scale bar: 5 cm. + + + +The pineal foramen is located in the posteromedial corner of the bone. In that aspect, + +Woznikella triradiata + +n. gen., n. sp. +differs from + +Kombuisia frerensis + +, in which the pineal foramen is absent ( +Hotton 1974 +; +Fröbisch 2007 +). The area of the suture with the left frontal is damaged, but it appears that the preparietal bone was absent. The internal side is also poorly preserved, and a large fragment of sediment was left unprepared to avoid destroying the specimen, obscuring most of the internal structures. Still, a deep fissure is visible in the center of the internal surface of the bone. + + + +TABLE +1. — Occurrences of Triassic dicynodonts. Note that the list is not comprehensive and likely numerous other mentions exist, particularly in local, nonEnglish literature. The Country column includes all mentions deemed significant (i.e., not merely listing taxa from original literature, but providing mentions, descriptions, and/or images of new specimens,original data or reinterpretations of morphology, confirming the geographical and/or temporal presence of taxa, confirming or providing novel insights into their validity, or at least presenting them in a novel biostratigraphic context). For simplicity, the Formation column includes only the references providing the most recent and most precise subdivisions either verbatim or in a form unambiguously recognizable without extensive research of local geological literature. The historically used + +Lystrosaurus +Cope, 1870a + +AZ was recently redefined and renamed to + +Lystrosaurus declivis +( +Owen, 1859 +) AZ ( +Botha & Smith 2020 +) + +. Because of the presence of + +Lystrosaurus +spp. + +also in the underlying + +Lystrosaurus maccaigi +( +Seeley,1898 +) + +- + +Moschorhinus +Broom, 1920 +SZ + +of + +Daptocephalus +Hoepen, 1934 AZ + +(former + +Dicynodon +AZ + +), the meaning of the historical + +Lystrosaurus +AZ + +is somewhat ambiguous and may be dependent on the author (see Table 2). Therefore, the name is used here in parentheses, implying that although in most cases it was probably synonymous with + +Lystrosaurus declivis +AZ + +, it can potentially include the top of the + +Daptocephalus +AZ. If + +the occurrence is not explicitly mentioned in the context of a changed naming scheme, the papers allowing referral (e.g., naming both the old formation and the new formation) are provided. The synonymy of Chinese + +Lystrosaurus +spp. + +follows +Camp & Liu (2011) +. Age includes selected single most recent original study with preference towards radiometric dates, unless those are unavailable, and a sound disagreement exists between several studies. Note that the bottom age (252.24 ± 0.11 Ma) of the + +Lystrosaurus declivis +AZ + +obtained by + +Gastaldo +et al. +(2020) + +is barely about 0.3 Ma older than the currently recognized Triassic/Permian boundary (251.902 ± 0.024 Ma). See + +Botha +et al. +(2020) + +for the entirely Triassic estimation of the age of that assemblage and +Botha & Smith (2020) +for discussion. Additional noteworthy contributions, not providing precise specific or geographic information include: +Broom 1900b +; +Toerien 1955 +; +Watson 1960 +; +Cruickshank 1964 +; +Anderson & Cruickshank 1978 +; +Kemp 1982 +; +Hotton 1986 +; + +King 1990 +b, 1991 + +; +King & Cluver 1991 +; +Smith 1995 +; +King & Jenkins 1997 +; +Cox 1998 +; +Surkov 1998b +; + +Neveling +et al. +1999 + +; +Smith & Ward 2001 +; +Angielczyk & Kurkin 2003 +; +Ray & Chinsamy 2003 +; + +Neveling +et al. +2005 + +; +Whitney & Sidor 2020 +; +Kammerer 2021 +; + +Liu +et al. +2021 + +; + +Whitney +et al. +2021 + +. Abbreviations: +AZ +, assemblage zone; +SZ +, subzone. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Taxon + +Country + +Formation + +Age +
+ +Acratophorus +argentinensis + +( +Bonaparte, 1965 +) + +Argentina ( +Bonaparte 1965 +, + +1966 +a, 1967 + +, +1970 +, +1978 +, +1981 +; +Cruickshank 1970 +; +Keyser & Cruickshank 1979 +; +Brink 1986 +; +King 1988 +; +Renaut 2000 +; +Renaut & Hancox 2001 +; +Domnanovich & Marsicano 2012 +; +Mancuso & Irmis 2019 +; + +Ordoñez +et al. +2019 + +; +Kammerer & Ordoñez 2021 +; + +Escobar +et al. +2023 + +) + +Río Seco de la Quebrada ( +Domnanovich & Marsicano 2012 +; +Mancuso & Irmis 2019 +) Quebrada de los Fósiles ( +Bonaparte 1981 +; +Domnanovich & Marsicano 2012 +) + +Carnian ( + +Ottone +et al. +2014 + +)?Ladinian-Carnian ( + +Ottone +et al. +2014 + +) +
+ +Angonisaurus +cruickshanki + +Cox & Li, 1983 + +Tanzania ( +Cox & Li 1983 +; +King 1988 +; +Hancox 1998 +; +Surkov & Benton 2004 +; + +Kammerer +et al. +2017 + +, +2019 +) + +Manda ( +Cox & Li 1983 +; +King 1988 +; +Hancox 1998 +; +Surkov & Benton 2004 +; + +Kammerer +et al. +2017 + +, +2019 +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Anomodontia +indet. + +Antarctica ( + +Cosgriff +et al. +1978 + +) + +Lower Fremouw ( + +Cosgriff +et al. +1978 + +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+South Africa ( +Watson 1960 +) + + +Cynognathus +AZ ( +Watson 1960 +) + + +Anisian-Carnian ( + +Hancox +et al. +2020 + +) +
+Tanzania ( +Haughton 1932 +; + +Attridge +et al. +1964 + +) + +Manda ( + +Attridge +et al. +1964 + +; +Cruickshank 1965 +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+“ + +Azarifeneria barrati + +” +Dutuit, 1989a + +Morocco ( +Dutuit 1989a +; +Gauffre 1993 +) + +Argana t5 ( +Dutuit 1989b +; +Gauffre 1993 +) + +Carnian/?Norian ( + +Kammerer +et al. +2012 + +) +
+“ + +Azarifeneria robustus + +” +Dutuit, 1989b + +Morocco ( +Dutuit 1989b +; +Gauffre 1993 +) + +Argana t5 ( +Dutuit 1989b +; +Gauffre 1993 +) + +Carnian/?Norian ( + +Kammerer +et al. +2012 + +) +
+ +Counillonia +superoculis + +Olivier, Battail, Bourquin, Rossignol, Steyer & Jalil, 2019 + +Laos ( + +Olivier +et al. +2019 + +) + +Purple Claystone ( + +Olivier +et al. +2019 + +) + +?Late Permian ( +Liu 2020 +)/ Early Triassic ( + +Olivier +et al +. 2019 + +) +
Dicynodontia indet. +Antarctica ( + +Hammer +et al. +2004 + +) + +Lashly C ( + +Hammer +et al. +2004 + +) + +Carnian ( + +Hammer +et al. +2004 + +) +
+Argentina ( + +Rogers +et al. +2001 + +) + +Chañares ( + +Rogers +et al. +2001 + +) + +Late Ladinian-early Carnian ( + +Ezcurra +et al. +2017 + +) +
+Australia ( +King 1983 +, +1988 +; +Thulborn 1983a +, b, +1990 +) + +Arcadia ( +King 1983 +; +Thulborn 1983a +, b, +1990 +) + +Early Triassic ( +Northwood 1997 +) +
+Brazil ( +von Huene 1935 +; +Holz & Schultz 1998 +; + +Da Rosa +et al. +2004 + +, +2005 +; + +Martinelli +et al. +2005 + +, +2017 +; + +Pavantino +et al. +2020 + +) + +Santa Maria + +Dinodontosaurus +AZ + +( + +Martinelli +et al. +2017 + +; + +Pavantino +et al. +2020 + +) + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/Carnian ( + +Ordoñez +et al. +2020 + +) +
+Canada ( +Sues & Olsen 2015 +) + +Wolfville ( +Sues & Olsen 2015 +) + +Carnian ( +Sues & Olsen 2015 +) +
+China ( +Young 1935 +, +1937 +, +1939 +, +1946 +) + +Not given ( +Young 1935 +, +1939 +) + +?Late Permian-Induan ( + +Tong +et al. +2018 + +) +
+Ermaying ( +Young 1937 +) + +Anisian ( + +Liu +et al. +2017 + +) +
+India ( +Bandyopadhyay 1988 +; + +Kutty +et al. +1988 + +; +Kutty & Sengupta 1989 +; +Bandyopadhyay & Ray 2020 +) + +Bhimaram ( + +Kutty +et al. +1988 + +; Kutty & Sengupta 1989; +Bandyopadhyay & Ray 2020 +) +Late Anisian-Ladinian (Bandyopadhyay & Ray 2020)
+Lower Maleri ( +Bandyopadhyay 1988 +; +Kutty & Sengupta 1989 +) + +Carnian ( +Bandyopadhyay & Ray 2020 +) +
+Japan ( + +Jinnouchi +et al. +2018 + +) + +Momonoki ( + +Jinnouchi +et al. +2018 + +) + +Carnian ( + +Jinnouchi +et al. +2018 + +) +
+Mozambique ( + +Araújo +et al. +2020 + +) + +Fubué ( + +Araújo +et al. +2020 + +) + +Early Triassic ( + +Araújo +et al. +2020 + +) +
+Namibia ( +Smith & Swart 2002 +; +Abdala & Smith 2009 +; + +Abdala +et al. +2013 + +) + +Omingonde ( +Smith & Swart 2002 +; +Abdala & Smith 2009 +; + +Abdala +et al. +2013 + +) + +Anisian-Ladinian ( + +Wynd +et al. +2018 + +; + +Zieger +et al. +2020 + +) +
+Russia ( + +Shishkin +et al. +1995 + +; + +Tverdokhlebov +et al. +2002 + +) + +Bukobay ( + +Shishkin +et al. +1995 + +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+Donguz ( + +Shishkin +et al. +1995 + +; + +Tverdokhlebov +et al. +2002 + +) + +Anisian ( +Ivakhnenko 2008 +) +
+South Africa ( +Nicolas & Rubidge 2010 +; + +Bordy +et al. +2020 + +) + + +Cynognathus +AZ ( +Nicolas & Rubidge 2010 +) + + +Anisian-Carnian ( + +Hancox +et al. +2020 + +) +
+Lower Elliott + +Scalenodontoides +AZ + +( + +Bordy +et al. +2020 + +; + +Viglietti +et al. +2020b + +) + +Norian-Rhaetian ( + +Bordy +et al. +2020 + +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Taxon + +Country + +Formation + +Age +
+“ + +Lystrosaurus +AZ + +” ( +Nicolas & Rubidge 2010 +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+Tanzania ( +Charig 1956 +; +Nesbitt & Butler 2013 +; + +Nesbitt +et al. +2013 + +) + +Manda ( +Charig 1956 +; +Nesbitt & Butler 2013 +; + +Nesbitt +et al. +2013 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+United States ( + +Sues +et al. +2003 + +; + +Nesbitt +et al. +2006 + +) + +Upper Moenkopi ( + +Nesbitt +et al. +2006 + +) + +Late Anisian ( + +Haque +et al. +2021 + +) +
+Newark Supergroup, Lithofacies Association II ( + +Sues +et al. +2003 + +) + +Late Carnian/early Norian ( + +Sues +et al. +2003 + +) +
+Zambia ( +Kemp 1975 +; + +Peecook +et al. +2018 + +; + +Whitney +et al. +2019 + +) + +Ntawere ( +Kemp 1975 +; + +Peecook +et al. +2018 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Dicynodontia indet. (“ + +Austrobrachyops + + +jenseni + +” +Colbert & Cosgriff, 1974 +) + +Antarctica ( +Colbert & Cosgriff 1974 +; +Warren & Marsicano 2000 +) + +Lower Fremouw ( +Colbert & Cosgriff 1974 +; +Warren & Marsicano 2000 +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+Dicynodontia indet. (“cf. +Dinodontosaurus +sp.”) + +Brazil ( + +Raugust +et al. +2013 + +; + +Martinelli +et al. +2017 + +; + +Schultz +et al. +2020 + +) + +?Santa Maria + +Santacruzodon +AZ + +( + +Raugust +et al. +2013 + +; + +Martinelli +et al. +2017 + +; + +Schultz +et al. +2020 + +) + +Carnian ( + +Philipp +et al. +2018 + +) +
+Dicynodontia indet. (“ + +Fukangolepis +barbaros + +” +Yang, 1978 +) + +China ( +Yang 1978 +; +Lucas & Hunt 1993b +) + +Upper Karamay ( +Yang 1978 +; +Lucas & Hunt 1993b +) + +Ladinian ( + +Tong +et al. +2018 + +) +
+Dicynodontia indet. (“ + +Dicynodon +sp. + +”) + +India ( +Lydekker 1879 +) + +Panchet ( +Lydekker 1879 +) + +Early Triassic ( +Prasad & Pundir 2020 +) +
+Dicynodontia indet. (“ + +Lystrosaurus +sp. + +”) + +Tanzania ( +Boonstra 1953 +; +Cruickshank 1967 +) + +Manda ( +Cruickshank 1965 +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Dicynodontia indet. (“ + +Ruhuhuungulasaurus +croucheri + +” +Larkin, 1994 +/ “ + +Shansiodon +sp. + +”) + +Tanzania ( +Larkin 1994 +; +Surkov & Benton 2004 +, +2008 +; + +Angielczyk +et al. +2014 + +; + +Kammerer +et al. +2017 + +) + +Manda ( +Larkin 1994 +; +Surkov & Benton 2004 +, +2008 +; + +Kammerer +et al. +2017 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Dicynodontia indet. (“ +Brachybrachium +breviceps +” +Williston, 1904 +) + +United States ( +Williston 1904 +; +von Huene 1911 +, +1935 +; + +Camp & Welles 1956 + +; +King 1988 +; + +Huber +et al. +1993 + +; +Lucas & Hunt 1993a +; +Long & Murry 1995 +; +Lucas 1998 +; +Lucas & Heckert 2002 +; + +Kammerer +et al. +2013 + +) + +Popo Agie ( +Williston 1904 +; +von Huene 1911 +; +1935 +; +Camp & Welles 1956 +; +King 1988 +; + +Huber +et al. +1993 + +; +Lucas & Hunt 1993a +; +Long & Murry 1995 +; +Lucas 1998 +; +Lucas & Heckert 2002 +; + +Kammerer +et al. +2013 + +) + +Carnian ( + +Hartman +et al. +2015 + +) +
+?Dicynodontia indet. (“aff. + +Dinodontosaurus + +sp.”) + +Germany ( +Lucas 2007 +) + +Upper Muschelkalk ( +Lucas 2007 +) +Anisian/Ladinian (German Stratigraphic Commission 2016)
+?Dicynodontia indet. (“cf. + +Placerias +sp. + +”) + +France ( +Broili 1921 +; +Schmidt 1928 +; +von Huene 1935 +; +Camp & Welles 1956 +; +Lucas & Wild 1995 +; + +Maisch +et al. +2009 + +) + +Upper Muschelkalk ( +Broili 1921 +; +Schmidt 1928 +; +von Huene 1935 +; +Camp & Welles 1956 +; +Lucas & Wild 1995 +; + +Maisch +et al. +2009) + +Anisian/Ladinian (German Stratigraphic Commission 2016)
Dicynodontoidea indet. +Australia ( + +Rozefelds +et al. +2011 + +) + +Upper Parmeer Supergroup ( + +Rozefelds +et al. +2011 + +) + +Late Permian-Early Triassic ( + +Rozefelds +et al. +2011 + +) +
+Dicynodontoidea indet. (“ + +Putillosaurus +sennikovi + +” +Surkov, 2005 +) + +Russia ( + +Shishkin +et al. +2000 + +; +Angielczyk & Kurkin 2003 +; +Surkov 2005 +; +Ivakhnenko 2008 +) + +Donskaya Luka ( +Surkov 2005 +; +Ivakhnenko 2008 +) + +Olenekian ( +Ivakhnenko 2008 +) +
+ +Dinodontosaurus +brevirostris + +Cox, 1968 + +Argentina ( +Cox 1968 +; +Bonaparte 1970 +, +1978 +, +1997 +; +Keyser 1974 +; +Keyser & Cruickshank 1979 +; +Brink 1988 +; +King 1988 +; + +Morato +et al. +2006 + +; + +Ordoñez +et al. +2019 + +; + +Escobar +et al. +2021 + +, +2023 +; +Kammerer & Ordoñez 2021 +) + +Chañares ( +Cox 1968 +; +Bonaparte 1970 +, +1978 +, +1997 +; +Keyser 1974 +; +Brink 1988 +; +King 1988 +; + +Ordoñez +et al. +2019 + +; + +Escobar +et al. +2021 + +, +2023 +; +Kammerer & Ordoñez 2021 +) + +Late Ladinian-early Carnian ( + +Ezcurra +et al. +2017 + +) +
+?Brazil ( +Cox 1968 +; + +Escobar +et al. +2023 + +) + +Santa Maria ( +Cox 1968 +) + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/early Carnian ( + +Ordoñez +et al. +2020 + +) +
+ +Dinodontosaurus tener + +( +von Huene, 1935 +) + +Brazil ( +von Huene 1935 +; +Tupi Caldas 1936 +; +Romer 1943 +; +Cox 1965 +; +Bonaparte 1970 +, +1978 +; +Keyser 1974 +; +Keyser & Cruickshank 1979 +; +Brink 1986 +, +1988 +; +King 1988 +; +Walter 1989 +; +Lucas & Harris 1996 +; +Lucas 2002 +; +Schwanke & Melo 2002 +; + +Vega-Dias +et al. +2004 + +; + +Da Rosa +et al. +2004 + +; +Morato 2006 +; + +Morato +et al. +2006 + +, +2008 +; + +Langer +et al. +2007 + +; + +Kammerer +et al. +2011 + +; + +de Oliveira Bueno +et al. +2011 + +; + +Abdala +et al. +2013 + +; +Dassie 2014 +; + +Martinelli +et al. +2017 + +; +Da Silveira 2017 +; +Kammerer 2018 +; +Mancuso & Irmis 2019 +; + +Ordoñez +et al. +2019 + +, +2020 +; + +Schultz +et al. +2020 + +; + +Escobar +et al. +2021 + +, +2023 +; +Kammerer & Ordoñez 2021 +; + +Knaus +et al. +2021 + +; +Maisch 2021 +) + +Santa Maria + +Dinodontosaurus +AZ + +( +Lucas 2002 +; + +Langer +et al. +2007 + +; + +de Oliveira Bueno +et al. +2011 + +; + +Abdala +et al. +2013 + +; +Dassie 2014 +; +Da Silveira 2017 +; + +Martinelli +et al. +2017 + +; Mancuso & Irmis 2019; + +Ordoñez +et al. +2020 + +; + +Schultz +et al. +2020 + +; +Kammerer & Ordoñez 2021 +; +Maisch 2021 +) + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/ early Carnian ( + +Ordoñez +et al. +2020 + +) +
+ +Dinodontosaurus +sp. + + +Brazil ( + +Da Rosa +et al. +2005 + +; + +Martinelli +et al. +2005 + +, +2016 +, +2017 +; + +Reichel +et al. +2009 + +; +Mainardes Dutra 2015 +; + +Ugalde +et al. +2018 + +; + +Corecco +et al. +2020 + +, +2021 +; + +Pavantino +et al. +2020 + +) + +Santa Maria + +Dinodontosaurus +AZ + +(Martinelli +et al. +2016, 2017; + +Ugalde +et al. +2018 + +; + +Corecco +et al. +2020 + +, +2021 +; + +Pavantino +et al. +2020) + + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/Carnian ( + +Ordoñez +et al. +2020 + +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
TaxonCountryFormation +Age +
+ +Dinodontosaurus +sp. + +(“ +Dicynodon turpior +” +von Huene, 1935 +, “ + +Dicynodon +cf. +turior + +”) + +Brazil ( +von Huene 1935 +, +1944 +,; +Cox 1965 +; +Bonaparte 1978 +; + +Brink 1986 + +; +King 1988 +; +Lucas 1993a +; +Lucas & Harris 1996 +; + +Langer +et al. +2007 + +; + +Kammerer +et al. +2011 + +; Kammerer & Ordoñez 2021) + +Santa Maria + +Dinodontosaurus +AZ + +( +Cox 1965 +; +King 1988 +; +Lucas 1993a +; +Lucas & Harris 1996 +; + +Langer +et al. +2007 + +; + +Kammerer +et al. + +2011; +Kammerer & Ordoñez 2021 +) + + + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/Carnian ( + +Ordoñez +et al. +2020 + +) +
+cf. + +Dinodontosaurus +sp. + + +Argentina ( + +Ezcurra +et al. +2017 + +) + +Chañares + +Tarjadia +AZ ( + +Ezcurra +et al. +2017 + +) + + +Ladinian/Carnian ( + +Ezcurra +et al. +2017 + +) +
+ +Dolichuranus primaevus + +Keyser, 1973 + +Namibia ( +Keyser 1973 +; +Cooper 1980 +; +Brink 1986 +; +King 1988 +; + +Damiani +et al. +2007 + +; +Govender & Yates 2009 +) + +Omingonde ( +Keyser 1973 +, +1974 +; +Cooper 1980 +; +Brink 1986 +; +King 1988 +; +Damiani et al. 2007 +) + +Anisian-Ladinian ( + +Wynd +et al. +2018 + +; + +Zieger +et al. +2020 + +) +
+cf. + +Dolichuranus +primaevus + +Keyser, 1973 + +Namibia ( +Govender & Yates 2009 +) + +Omingonde ( +Govender & Yates 2009 +) + +Anisian-Ladinian ( + +Wynd +et al. +2018 + +; + +Zieger +et al. +2020 + +) +
+ +Dolichuranus +sp. + + +Tanzania ( + +Kammerer +et al. +2017 + +; + +Peecook +et al. +2018 + +, + +Smith +et al. +2018 + +, + +Wynd +et al. +2018 + +) + +Manda ( + +Kammerer +et al. +2017 + +, + +Peecook +et al. +2018 + +, + +Wynd +et al. +2018 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ +Elephantosaurus +jachimovitschi + +Vjuschkov, 1969 + +Russia ( +Vjuschkov 1969 +; +Keyser & Cruickshank 1979 +; +King 1988 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; Battail & +Surkov 2000 +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Bukobay ( + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; Ivakhnenko 2008; + +Kammerer +et al. +2013 + +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+ +Eubrachiosaurus browni + +Williston, 1904 + +United States ( +Williston 1904 +; +Cross & Howe 1905 +; +von Huene 1911 +, +1926a +, +1935 +; +Pearson 1924a +; +Camp & Welles 1956 +; +King 1988 +; + +Huber +et al. +1993 + +; +Long & Murry 1995 +; +Lucas 1998 +; +Lucas & Heckert 2002 +; + +Kammerer +et al. +2013 + +) + +Popo Agie ( +Williston 1904 +; +von Huene 1911 +, +1926a +, +1935 +; +Camp & Welles 1956 +; +King 1988 +; + +Huber +et al. +1993 + +; +Long & Murry 1995 +; +Lucas 1998 +; +Lucas & Heckert 2002 +; + +Kammerer +et al. +2013 + +) + +Carnian ( + +Hartman +et al. +2015 + +) +
+ +Ischigualastia jenseni + +Cox, 1962 + +Argentina ( +Cox 1962 +, +1965 +; +Bonaparte 1970 +, +1978 +, +1997 +; +Keyser 1974 +; +Keyser & Cruickshank 1979 +; +King 1988 +; +Vega-Dias & Schwanke 2004a +; + +Martínez +et al. +2012 + +; + +Kammerer +et al. +2017 + +; +Kammerer 2018 +; + +Ordoñez +et al. +2019 + +; + +Wynd +et al. +2020 + +; +Maisch 2021 +; +Kammerer & Ordoñez 2021 +) + +Ischigualasto B1 ( + +Martínez +et al. +2012 + +) + +Late Carnian ( + +Martínez +et al. +2011 + +) +
+cf. + +Ischigualastia jenseni + +Cox, 1962 + +India ( +Edler 2000 +) + +Pipariya ( +Edler 2000 +) + +Carnian ( + +Chatterjee +et al. +2017 + +) +
+cf. + +Ischigualastia +sp. + + +India ( +Bandyopadhyay 1988 +; +Kutty & Sengupta 1989 +; + +Novas +et al. +2010 + +; +Bandyopadhyay & Ray 2020 +) + +Upper Maleri ( +Bandyopadhyay 1988 +; +Kutty & Sengupta 1989 +; + +Novas +et al. +2010 + +; Bandyopadhyay & Ray 2020) + +Norian ( +Bandyopadhyay & Ray 2020 +) +
+ +Jachaleria +candelariensis + +Araújo & Gonzaga, 1980 + +Brazil ( +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 1999 +, +2003 +, +2004 +, +2007 +; +Schultz & Vega-Dias 2003 +; VegaDias & Schwanke 2004b, 2005; + +Vega-Dias +et al. +2004 + +; + +Morato +et al. +2005 + +; + +Langer +et al. +2007 + +; +Francischini Filho 2014 +; +Kammerer 2018 +; + +Corecco +et al. +2020 + +, +2021 +; + +Schultz +et al. +2020 + +; + +Wynd +et al. +2020 + +; +Maisch 2021 +) + +Caturrita ( +Lucas 1993a +; Schultz & Vega-Dias 2003; +Vega-Dias & Schultz 2003 +, +2007 +; +Vega-Dias & Schwanke 2004b +, +2005 +; + +Langer +et al. +2007 + +; +Francischini Filho 2014 +; +Kammerer 2018 +; +Kammerer & Ordoñez 2021 +; +Maisch 2021 +) + +Norian ( + +Langer +et al. +2018 + +) +
+cf. + +Jachaleria +canderlariensis + +Araújo & Gonzaga, 1980 + +Brazil ( + +Martinelli +et al. +2020 + +) + +Caturrita ( + +Martinelli +et al. +2020 + +) + +Norian ( + +Langer +et al. +2018 + +) +
+ +Jachaleria colorata + +Bonaparte, 1970 + +Argentina ( +Bonaparte 1966b +, +1970 +, +1978 +, +1997 +; +Keyser 1974 +; +King 1988 +; +Vega-Dias & Schwanke 2004b +, +2005 +; + +Martínez +et al. +2012 + +; +Francischini Filho 2014 +; +Kammerer 2018 +; + +Ordoñez +et al. +2019 + +) + +Ischigualasto B3 ( + +Martínez +et al. +2012 + +) + +Norian ( + +Currie +et al. +2009 + +) +
+Lower Los Colorados ( + +Bonaparte 1966 +a, 1970 + +, +1978 +; +Araújo & Gonzaga 1980 +; Vega- Dias & Schwanke 2005; + +Martínez +et al. +2012 + +; +Francischini Filho 2014 +; +Kammerer 2018 +) + +Norian ( + +Currie +et al. +2009 + +) +
+ +Kannemeyeria +aganosteus + +Kammerer & Ordoñez, 2021 + +Argentina ( +Bonaparte 1981 +; +DeFauw 1993 +; +FrÖbisch 2009 +; +Kammerer & Ordoñez 2021 +) + +Quebrada de los Fósiles ( +Bonaparte 1981 +; +DeFauw 1993 +) + +?Ladinian-Carnian ( + +Ottone +et al. +2014 + +) +
+Río Seco de la Quebrada ( +Kammerer & Ordoñez 2021 +) + +Carnian ( + +Ottone +et al. +2014 + +) +
+“ + +Kannemeyeria + +” + +latirostris + +Crozier, 1970 + +Zambia ( +Brink 1963 +, +1986 +; +Drysdall & Kitching 1963 +; Kitching 1963; +Crozier 1970 +; +Cruickshank 1970 +; +Keyser & Cruickshank 1979 +; +King 1988 +; + +Angielczyk +et al. +2014 + +; + +Peecook +et al. +2018 + +) + +Ntawere ( +Brink 1963 +, +1986 +; +Drysdall & Kitching 1963 +; +Kitching 1963 +; +Crozier 1970 +; +Cruickshank 1970 +; +Keyser & Cruickshank 1979 +; +King 1988 +; + +Angielczyk +et al. +2014 + +; + +Peecook +et al. +2018 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ +Kannemeyeria +lophorhinus + +Renaut, Damiani, Yates & Hancox, 2003 + +Namibia ( +Keyser 1973 +; +Brink 1986 +; +King 1988 +; Bandyopadhyay 1989; +Pickford 1995 +; +Renaut 2000 +; + +Renaut +et al. +2003 + +; +Govender & Yates 2009 +) + +Omingonde ( +Keyser 1973 +; +Brink 1986 +; +King 1988 +; +Bandyopadhyay 1989 +; +Pickford 1995 +; +Renaut 2000 +; + +Renaut +et al. +2003 + +; Govender & Yates 2009) + +Anisian-Ladinian ( + +Wynd +et al. +2018 + +; + +Zieger +et al. +2020 + +) +
+?Tanzania ( +Cox 1991 +; +Renaut 2000 +) + +Manda ( +Cox 1991 +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Zambia ( +Brink 1963 +, +1986 +; +Drysdall & Kitching 1963 +; +Crozier 1970 +; +Keyser & Cruickshank 1979 +; +King 1988 +; +Bandyopadhyay 1989 +; +Renaut 2000 +; + +Renaut +et al. +2003 + +; + +Angielczyk +et al. +2014 + +; + +Peecook +et al. +2018 + +) + +Ntawere ( +Brink 1963 +, +1986 +; Drysdall & Kitching 1963; +Kitching 1963 +; +Crozier 1970 +; Keyser & Cruickshank 1979; +King 1988 +; Bandyopadhyay 1989; +Renaut 2000 +; + +Renaut +et al. +2003 + +; + +Angielczyk +et al. +2014 + +; + +Peecook +et al. +2018 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Taxon +Country +Formation + +Age +
+cf. + +Kannemeyeria + + +lophorhinus + + +Namibia ( +Govender & Yates 2009 +) + +Omingonde ( +Govender & Yates 2009 +) + +Anisian-Ladinian ( + +Wynd +et al. +2018 + +; + +Zieger +et al. +2020 + +) +
+ +Kannemeyeria +simocephalus + +( +Weithofer, 1888 +) + +South Africa ( +Weithofer 1888 +; +Broom 1899 +, +1909 +, +1913a +, +1915 +, +1923 +, +1932 +, +1937 +; +Seeley 1904 +, +1908 +; +Jaekel 1911 +; +Watson 1912a +, +1917 +, +1948 +; +Haughton 1915 +, +1917 +, +1924 +, +1963 +; +Pearson 1924a +, b; +von Huene 1925 +; +Case 1934 +; +Courtenay-Latimer 1948 +; +Toerien 1951 +, +1953 +, +1955 +; Haughton & Brink 1954; +Camp 1956 +; +Cruickshank 1970 +, +1975 +; +Anonymous 1972 +; +Keyser 1974 +; +Kitching 1977 +; +Keyser & Cruickshank 1979 +; +Brink 1986 +; +King 1988 +; +Hancox 1998 +; +MacRae 1999 +; +Renaut 2000 +; Nesbitt & Angielczyk 2002; +Damiani & Kitching 2003 +; +Neveling 2004 +; Surkov & Benton 2004; +Govender 2005 +; +Ray 2006 +; Nicolas 2007; +Jinnah & Rubidge 2007 +; +Damiani 2008 +; FrÖbisch & Reisz 2008; + +Govender +et al. +2008 + +; +Botha-Brink & Angielczyk 2010 +; +Nicolas & Rubidge 2010 +; + +Kammerer +et al. +2011 + +, +2017 +; + +Smith +et al. +2012 + +, +2020b +; + +Butler +et al. +2016 + +; +Kammerer 2018 +; + +Hancox +et al. +2020 + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Viglietti +et al. +2022 + +) + + +Cynognathus +AZ +Trirachodon-Kannemeyeria + +SZ ( + +Hancox +et al. +2020 + +; + +Smith +et al. +2020b + +; + +Viglietti +et al. +2022 + +) + +Early Anisian ( + +Hancox +et al. +2020 + +) +
+Tanzania ( +Haughton 1932 +; +Stockley 1932 +; +Boonstra 1953 +; + +Attridge +et al. +1964 + +; +Cruickshank 1965 +; Keyser & Cruickshank 1979; + +Kammerer +et al. +2017 + +) + +Manda ( +Stockley 1932 +; + +Attridge +et al. +1964 + +; +Cruickshank 1965 +; +Keyser & Cruickshank 1979 +; + +Kammerer +et al. +2017 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Kannemeyeriidae +indet. + +Argentina ( +Romer 1966 +; +Cox 1968 +; + +Ezcurra +et al. +2015 + +) + +Tarjados Formation ( +Cox 1968 +; + +Ezcurra +et al. +2015 + +) + +Ladinian ( + +Ezcurra +et al. +2017 + +) +
+China ( +Sun 1963 +, +1973 +; +Young 1964 +; +Liu 1973 +, +2015 +; +Cheng 1980 +) + +Upper Ermaying ( +Sun 1963 +; +Cheng 1980 +; +Liu 2015 +) + +Anisian ( + +Liu +et al. +2017 + +) +
+Lower Karamay ( +Liu 1973 +; +Sun 1973 +; +Liu & Li 2003 +) + +Anisian ( + +Tong +et al. +2018 + +) +
+Tongchuan I ( +Liu 2015 +) + +Anisian ( + +Tong +et al. +2018 + +) +
+India ( +Bandyopadhyay 1988 +; +Mukherjee & Sengupta 1998 +; +Bandyopadhyay & Sengupta 1999 +; +Bandyopadhyay & Ray 2020 +) + +Denwa ( +Bandyopadhyay 1988 +; +Mukherjee & Sengupta 1998 +; +Bandyopadhyay & Sengupta 1999 +; +Bandyopadhyay & Ray 2020 +) + +Anisian or younger ( + +Peecook +et al. +2018 + +) +
+Madagascar ( + +Flynn +et al. +1999 + +) + +Isalo II ( + +Flynn +et al. +1999 + +) + +Carnian ( + +Flynn +et al. +1999 + +) +
+Russia ( + +Ivakhnenko +et al. +1997 + +) + +Bukobay ( + +Ivakhnenko +et al. +1997 + +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+South Africa ( +Warren 1998 +) + +Cynognathus +AZ ( +Warren 1998 +) + +Anisian-Carnian ( + +Hancox + +et al. +2020) + + +
+United States ( + +Small +et al. +2022 + +) + +Tecovas ( + +Small +et al. +2022 + +) + + +Norian ( + +Small +et al. +2022 + +) + +
+Kannemeyeriiformes +indet. + +Antarctica ( + +Hammer +et al. +1987 + +; +Hammer 1990 +, +1995 +; + +Sidor +et al. +2014 + +; + +Smith +et al. +2020a + +) + +Upper Fremouw ( + +Hammer +et al. +1987 + +; +Hammer 1990 +, +1995 +; + +Sidor +et al. +2014 + +; + +Smith +et al. +2020a + +) + +Late Anisian-Ladinian ( + +Elliot +et al. +2017 + +) +
+Argentina ( +Zavattieri & Arcucci 2007 +; +Kammerer & Ordoñez 2021 +) + +Cerro de las Cabras ( +Zavattieri & Arcucci 2007 +; +Kammerer & Ordoñez 2021 +) + +Late Anisian-early Ladinian ( + +Cariglino +et al. +2016 + +) +
+Kazakhstan ( + +Tverdokhlebov +et al. +2020 + +) + +Inder ( + +Tverdokhlebov +et al. +2020 + +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+Russia ( + +Tverdokhlebov +et al. +2020 + +) + +Bukobay ( + +Tverdokhlebov +et al. +2020 + +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+South Africa ( + +Hancox +et al. +2013 + +) + + +Cynognathus +AZ +Cricodon-Ufudocyclops +SZ + +( + +Hancox +et al. +2013 + +, +2020 +) + +Late Anisian-Carnian ( + +Hancox +et al. +2020 + +) +
+United States ( +Nesbitt & Angielczyk 2002 +; +Lehman & Chatterjee 2005 +; +Mueller & Chatterjee 2007 +; +Martz 2008 +; + +Martz +et al. +2013 + +; + +Small +et al. +2022 + +) + +Cooper Canyon ( +Lehman & Chatterjee 2005 +; +Martz 2008 +; + +Martz +et al. +2013 + +) Upper Moenkopi ( +Nesbitt & Angielczyk 2002 +) Tecovas ( + +Small +et al. +2022 + +) + +Norian ( + +Martz +et al. +2013 + +) +
+Late Anisian ( + +Haque +et al. +2021 + +) +
+Norian ( + +Small +et al. +2022 + +) +
+Zambia ( + +Angielczyk +et al. +2014 + +) + +Ntawere ( + +Angielczyk +et al. +2014 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+Kannemeyeriiformes +indet. (“ + +Calleonasus + + +furvus + +” +Kalandadze & Sennikov, 1985 +) + +Russia ( +Kalandadze & Sennikov 1985 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; + +Surkov 1999 +a, 2003 + +; +Battail & Surkov 2000 +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+Kannemeyeriiformes +indet. (“ + +Cristonasus + +koltaeviensis +” +Surkov, 1999a +) + +Russia ( + +Surkov 1999 +a, 2003 + +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+Kannemeyeriiformes +indet. (“ + +Edaxosaurus + + +edentatus + +” Kalandadze & Sennikov, 1985) + +Russia ( +Kalandadze & Sennikov 1985 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; +Surkov 2003 +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+Kannemeyeriiformes +indet. (“ + +Elatosaurus + + +facetus + +” +Kalandadze & Sennikov, 1985 +) + +Russia ( +Kalandadze & Sennikov 1985 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; + +Surkov 1999 +a, 2003 + +; +Battail & Surkov 2000 +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Bukobay ( +Kalandadze & Sennikov 1985 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; +Ivakhnenko 2008 +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Taxon + +Country + +Formation + +Age +
+Kannemeyeriiformes +indet. (“ + +Nasoplanites +danilovi + +” +Surkov, 1999a +) + +Russia ( + +Surkov 1999 +a, 2003 + +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+Kannemeyeriiformes +indet. (“ + +Parvobestiola + +bashkiriensis +” +Surkov, 1999a +) + +Russia ( + +Surkov 1999 +a, 2003 + +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+Kannemeyeriiformes +indet. (“ + +Planitorostris + + +pechoriensis + +” +Surkov, 1999b +) + +Russia ( +Surkov 1999b +; +Ivakhnenko 2008 +; + +Kammerer +et al. +2013 + +) + +Bukobay ( +Surkov 1999b +; + +Shishkin +et al. +2000 + +) + +Ladinian ( + +Tverdokhlebov +et al. +2020 + +) +
+ +Kombuisia antarctica + +FrÖbisch, Angielczyk & Sidor, 2010 + +Antarctica ( +DeFauw 1989 +; + +FrÖbisch +et al. +2010 + +) + +Lower Fremouw ( +DeFauw 1989 +; + +FrÖbisch +et al. +2010 + +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+ +Kombuisia frerensis + +Hotton, 1974 + +South Africa ( +Hotton 1974 +; +Keyser 1974 +; +Kitching 1977 +; +Keyser & Cruickshank 1979 +; +Brink 1986 +; +King 1988 +; +FrÖbisch 2007 +; +Nicolas & Rubidge 2010 +; + +Smith +et al. +2020b + +, +2012 +; + +Hancox +et al. +2020 + +; + +Viglietti +et al. +2022 + +) + + +Cynognathus +AZ +Trirachodon-Kannemeyeria + +SZ ( + +Hancox +et al. +2020 + +; + +Smith +et al. +2020b + +; + +Viglietti +et al. +2022 + +) + +Early Anisian ( + +Hancox +et al. +2020 + +) +
+ +Lisowicia bojani +Sulej + +& Niedźwiedzki, 2018 + +Poland ( + +Dzik +et al. +2008a + +, b; +Vogel 2008 +; Sulej & Niedźwiedzki 2009, 2019; + +Niedźwiedzki +et al. +2011 + +; +Racki & Lucas 2020 +; +Romano & Manucci 2021 +) + +Grabowa ( + +Szulc +et al. +2015b + +; +Racki & Lucas 2020 +) + +Norian/Rhaetian (Kowal-Linka +et al. +2019) +
+cf. + +Lisowicia bojani + +Sulej & Niedźwiedzki, 2018 + +Poland ( + +Budziszewska-Karwowska +et al. +2010 + +; +Sadlok & Wawrzyniak 2013 +; + +Sulej +et al. +2019 + +; +Racki & Lucas 2020 +) + +Grabowa ( + +Szulc +et al. +2015b + +; +Racki & Lucas 2020 +) + +Norian ( + +Szulc +et al. +2015b + +) +
+ +Lystrosaurus curvatus + +( +Owen, 1876 +) + +Antarctica ( +Colbert 1972 +, +1973 +, +1974 +, +1975 +; +Cosgriff & Hammer 1981 +; +Kulik & Sidor 2023 +) + +Lower Fremouw ( +Colbert 1974 +; +Cosgriff & Hammer 1981 +; +Kulik & Sidor 2023 +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+South Africa ( +Owen 1876 +; +Seeley 1898 +; +Broom 1903 +, +1907 +, +1909 +, +1932 +, +1940 +; +Van Hoepen 1915 +, +1916 +; Haughton 1924; +von Huene 1925 +; +Brink 1951 +, +1986 +; Haughton & Brink 1954; +Barry 1968 +; +Kitching 1968 +, +1977 +; +Cluver 1971 +; +Colbert 1974 +, +1975 +; +Keyser & Cruickshank 1979 +; + +Cosgriff +et al. +1982 + +; +DeFauw 1986 +; +King 1988 +; +Surkov & Benton 2004 +; +Ray 2005 +; +Smith & Botha 2005 +; + +Grine +et al. +2006 + +; +Botha & Smith 2006 +, +2007 +, +2020 +; + +Jasinoski +et al. +2009 + +, +2010 +, +2014 +; +Camp 2010 +; + +Kammerer +et al. +2011 + +; +Jasinoski & Chinsamy-Turan 2012 +; + +Smith +et al. +2012 + +, +2020b +; +Smith & Botha-Brink 2014 +; + +Botha-Brink +et al. +2014 + +, +2016 +; + +Rubidge +et al. +2016 + +; + +Viglietti +et al. +2016 + +; +Botha-Brink 2017 +; + +Benoit +et al. +2018 + +; +Botha 2020 +; Gastaldo +et al. +2020; +Modesto 2020 +; +Viglietti 2020 +; + +Botha +et al. +2020 + +; + +Liu +et al. +2022 + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Viglietti +et al. +2022 + +; +Kulik & Sidor 2023 +) + + +Daptocephalus +AZ + +, + +Lystrosaurus + +maccaigi-Moschorhinus +SZ + + +( + +Smith +et al. +2020b + +; +Viglietti 2020 +; + +Liu +et al. +2022 + +; + +Viglietti +et al. +2022 + +) + +Late Permian ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus declivis +AZ + +( +Botha & Smith 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; +Viglietti 2020 +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus + +cf. + +curvatus + +( +Owen, 1876 +) + +India ( +Gupta & Das 2011 +; +Bandyopadhyay & Ray 2020 +; +Kulik & Sidor 2023 +) + +Panchet ( +Gupta & Das 2011 +; Bandyopadhyay & Ray 2020; +Kulik & Sidor 2023 +) + +Early Triassic ( +Prasad & Pundir 2020 +) +
+South Africa ( +Kitching 1977 +; +DeFauw 1986 +) + +“ + +Lystrosaurus +AZ + +” ( +Kitching 1977 +; +DeFauw 1986 +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus declivis + +( +Owen, 1859 +) + +South Africa ( +Owen 1859 +, 1860, +1862 +, +1876 +; +Broom 1903 +, +1909 +, +1915 +, +1932 +; +Watson 1912b +; +Van Hoepen 1916 +; +Haughton 1924 +; +von Huene 1925 +, +1931 +; +Brink 1951 +, +1986 +; +Haughton & Brink 1954 +; +Toerien 1954 +; +Barry 1968 +; +Kitching 1968 +, +1977 +; +Cluver 1971 +; +Colbert 1974 +; Keyser & Cruickshank 1979; + +Cosgriff +et al. +1982 + +; +DeFauw 1986 +; +King 1988 +; + +Thackeray +et al. +1998 + +; +Angielczyk 2002 +; + +Retallack +et al. +2003 + +; +Neveling 2004 +; +Smith & Botha 2005 +; + +Grine +et al. +2006 + +; +Botha & Smith 2006 +, +2007 +, +2020 +; + +Jasinoski +et al. +2009 + +, +2014 +; +Camp 2010 +; + +Modesto +et al. +2010 + +; Botha-Brink & Angielczyk 2010; + +Kammerer +et al. +2011 + +; + +Viglietti +et al. +2013 + +; + +Botha-Brink +et al. +2016 + +; +Smith & Botha-Brink 2014 +; Botha-Brink +et al. +2014; + +Rubidge +et al. +2016 + +; + +Benoit +et al. +2018 + +; +Thackeray 2018 +, +2019 +; + +Gastaldo +et al. +2019 + +, +2020 +; +Botha 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; +Viglietti 2020 +; + +Botha +et al. +2020 + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Smith +et al. +2022 + +; + +Viglietti +et al. +2022 + +; +Kulik & Sidor 2023 +) + + +Lystrosaurus declivis +AZ + +( +Botha 2020 +; +Botha & Smith 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; +Viglietti 2020 +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Smith +et al. +2022 + +; + +Viglietti +et al. +2022 + +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus +cf. +declivis + +( +Owen, 1859 +) + +India ( +Gupta & Das 2011 +; +Bandyopadhyay & Ray 2020 +; +Kulik & Sidor 2023 +) + +Panchet ( +Gupta & Das 2011 +; Bandyopadhyay & Ray 2020; +Kulik & Sidor 2023 +) + +Early Triassic ( +Prasad & Pundir 2020 +) +
+South Africa ( +Broom 1915 +; +von Huene 1931 +) + +“ + +Lystrosaurus +AZ + +” ( +Broom 1915 +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus georgi + +Kalandadze, 1975 + +Russia ( +Kalandadze 1974 +, +1975 +; +Colbert 1982 +; +Lozovskii 1983 +; +King 1988 +; +Ochev 1992 +; + +Ivakhnenko +et al. +1997 + +; Battail & +Surkov 2000 +; + +Surkov +et al. +2005 + +; +Ivakhnenko 2008 +) + +Vokhmian ( + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; + +Surkov +et al. +2005 + +; +Ivakhnenko 2008 +) + +Induan ( +Ivakhnenko 2008 +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Taxon +Country +Formation + +Age +
+ +Lystrosaurus hedini + +Young, 1935 + +China ( +Yuan & Young 1934b +; +Young 1935 +, +1939 +, +1946 +; +Brink 1951 +, +1986 +, +1988 +; +Liu 1973 +; +Sun 1973 +; +Colbert 1974 +, +1982 +; +Yuhe 1983 +; +Cheng 1986 +; +King 1988 +; +Li 1988 +; +Lucas 2001 +; + +Liu +et al. +2002 + +; +Li & Sun 2008 +; +Camp 2010 +; +Camp & Liu 2011 +; +Li 2015 +; +Modesto 2020 +; +Kulik & Sidor 2023 +) + +Jiucaiyuan ( +Yuhe 1983 +; +Li 1988 +; +Lucas 2001 +; + +Liu +et al. +2002 + +; +Li & Sun 2008 +) Upper Guodikeng ( + +Liu +et al. +2002 + +; +Li & Sun 2008 +) + +Induan ( + +Tong +et al. +2018 + +) Induan ( + +Tong +et al. +2018 + +) +
+Mongolia ( +Gubin & Sinitza 1993 +; +Kulik & Sidor 2023 +) + +Upper Noyan Somon ( +Gubin & Sinitza 1993 +) + +Early Triassic ( +Gubin & Sinitza 1993 +) +
+ +Lystrosaurus +cf. +hedini + +Young, 1935 + +China ( +Young 1939 +; + +Marilao +et al. +2020 + +) + +Jiucaiyuan ( + +Marilao +et al. +2020 + +) + +Induan ( + +Tong +et al. +2018 + +) +
+Not given ( +Young 1939 +) + +?Induan ( + +Tong +et al. +2018 + +) +
+ +Lystrosaurus maccaigi + +( +Seeley, 1898 +) + +Antarctica ( +Cosgriff & Hammer 1979 +; + +Cosgriff +et al. +1982 + +; +Kulik & Sidor 2023 +) + +Lower Fremouw ( +Cosgriff & Hammer 1979 +; + +Cosgriff +et al. +1982 + +; +Kulik & Sidor 2023 +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+South Africa ( +Seeley 1898 +; +Broom 1909 +, +1932 +, +1903 +; +Van Hoepen 1915 +; +Haughton 1924 +; +von Huene 1925 +; +Brink 1951 +; +Toerien 1953 +; +Haughton & Brink 1954 +; Kitching 1968, 1977; +Cluver 1971 +; +Colbert 1974 +; +Keyser & Cruickshank 1979 +; + +Cosgriff +et al. +1982 + +; +King 1988 +; +Ray 2005 +; +Smith & Botha 2005 +; +Botha & Smith 2006 +; + +Grine +et al. +2006 + +; +Botha & Smith 2007 +, +2020 +; + +Jasinoski +et al. +2010 + +, +2014 +; +Botha-Brink & Angielczyk 2010 +; +Jasinoski & Chinsamy-Turan 2012 +; +Smith & Botha-Brink 2014 +; Botha-Brink +et al. +2014, 2016; + +Rubidge +et al. +2016 + +; + +Viglietti +et al. +2016 + +; + +Gastaldo +et al. +2019 + +, +2020 +; +Botha 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; +Viglietti 2020 +; + +Botha +et al. +2020 + +; + +Liu +et al. +2022 + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Viglietti +et al. +2022 + +; +Kulik & Sidor 2023 +) + + +Daptocephalus +AZ + +, + +Lystrosaurus +maccaigi- + + +Moschorhinus +SZ + +( + +Smith +et al. +2020b + +; +Viglietti 2020 +; + +Liu +et al. +2022 + +; + +Viglietti +et al. +2022 + +) + +Late Permian ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus declivis +AZ + +( +Botha 2020 +; +Botha & Smith 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; +Viglietti 2020 +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus murrayi + +( +Huxley, 1859 +) + +Antarctica ( +Colbert 1970a +, +1971a +, +1972 +, +1973 +, 1974, 1975; + +Cosgriff +et al. +1978 + +; +Retallack & Hammer 1998 +; +Kulik & Sidor 2023 +) + +Lower Fremouw ( +Colbert 1974 +; + +Cosgriff +et al. +1978 + +; +Retallack & Hammer 1998 +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+India ( +Huxley 1865 +; +Lydekker 1877 +, +1879 +, +1889 +; +Cotter 1918 +; +Das Gupta 1922 +; +von Huene 1935 +; +Brink 1951 +, +1988 +; +Robinson 1958 +; +Tripathi & Puri 1961 +; Sahani & Tripathi 1962; +Tripathi & Satsangi 1963 +; + +Colbert 1971 +a, 1973 + +, +1974 +; + +Cosgriff +et al. +1982 + +; +King 1988 +; +Ray 2005 +, +2006 +; + +Ray +et al. +2005 + +, +2009a +, b; + +Kammerer +et al. +2011 + +; Bandyopadhyay & Ray 2020; +Kulik & Sidor 2023 +) + +Panchet ( +Huxley 1865 +; +Lydekker 1877 +, Early Triassic (Prasad & 1879, 1889; +Cotter 1918 +; +Das Gupta 1922 +; Pundir 2020) +von Huene 1935 +; +Robinson 1958 +; +Tripathi & Puri 1961 +; +Sahani & Tripathi 1962 +; +Tripathi & Satsangi 1963 +; + +Colbert 1971 +a, 1974 + +; + +Cosgriff +et al. +1982 + +; +Brink 1988 +; +King 1988 +; +Ray 2005 +, +2006 +; + +Ray +et al. +2005 + +, +2009a +, b; + +Kammerer +et al. +2011 + +; +Bandyopadhyay & Ray 2020 +) +
+South Africa ( +Huxley 1859 +; +Owen 1860b +; +Cope 1870a +; +Seeley 1889 +; +Broom 1902 +, +1909 +, +1932 +, +1941 +; +Van Hoepen 1916 +; +Haughton 1917 +, +1924 +; +von Huene 1925 +; Arambourg 1943; +Brink 1951 +, +1986 +; +Janensch 1952 +; Haughton & Brink 1954; +Camp 1956 +, +2010 +; +Cruickshank 1967 +; +Barry 1968 +; +Kitching 1968 +, +1977 +; +Cluver 1971 +; + +Colbert 1971 +a, 1974 + +; +Aulie 1974 +; +Keyser & Cruickshank 1979 +; + +Cosgriff +et al. +1982 + +; +DeFauw 1986 +; +King 1988 +; +Chinsamy & Rubidge 1993 +; + +Thackeray +et al. +1998 + +; +Ray 2005 +, +2006 +; + +Ray +et al. +2005 + +, +2009a +; +Smith & Botha 2005 +; + +Grine +et al. +2006 + +; +Botha & Smith 2006 +, +2007 +, +2020 +; + +Jasinoski +et al. +2010 + +, +2014 +; + +Modesto +et al. +2010 + +; +Botha-Brink & Angielczyk 2010 +; + +Kammerer +et al. +2011 + +; +Jasinoski & Chinsamy-Turan 2012 +; + +Botha-Brink +et al. +2016 + +; +Smith & Botha-Brink 2014 +; Botha-Brink +et al. +2014; + +Rubidge +et al. +2016 + +; + +Gastaldo +et al. +2017 + +, +2019 +; +Thackeray 2018 +, +2019 +; +Botha 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; + +Botha +et al. +2020 + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Smith +et al. +2022 + +; + +Viglietti +et al. +2022 + +; +Kulik & Sidor 2023 +) + + +Lystrosaurus declivis +AZ + +( +Botha 2020 +; +Botha & Smith 2020 +; +Modesto 2020 +; + +Smith +et al. +2020b + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Smith +et al. +2022 + +; + +Viglietti +et al. +2022 + +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus + +cf. + +murrayi + +( +Huxley, 1859 +) + +South Africa ( +Broom 1900a +, +1915 +) + +?“ + +Lystrosaurus +AZ + +” ( +Broom 1915 +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus youngi + +Sun, 1964 + +China ( +Sun 1964 +, +1973 +; +Liu 1973 +; +Colbert 1974 +, +1982 +; +Yuhe 1983 +; +Li 1988 +; +Lucas 2001 +; +Li & Sun 2008 +; +Camp 2010 +; +Camp & Liu 2011 +; +Li 2015 +; +Kulik & Sidor 2023 +) + +Jiucaiyuan ( +Yuhe 1983 +; +Li 1988 +; +Lucas 2001 +; + +Liu +et al. +2002 + +; +Li & Sun 2008 +) + +Induan ( + +Tong +et al. +2018 + +) +
+ +Lystrosaurus +cf. +youngi + +Sun, 1964 + +China ( + +Liu +et al. +2002 + +; +Li 2015 +) + +Upper Guodikeng ( + +Liu +et al. +2002 + +) + +Induan ( + +Tong +et al. +2018 + +) +
+ +Lystrosaurus +sp. + +(“ + +Dicynodon +strigops + +” +Broom, 1913b +) + +South Africa ( +Broom 1913b +, +1915 +; +Haughton 1917 +, +1924 +; +von Huene 1925 +; +Brink 1951 +; +Haughton & Brink 1954 +; +Cluver 1971 +; +Colbert 1974 +; + +Kammerer +et al. +2011 + +) + +“ + +Lystrosaurus +AZ + +” ( +Haughton 1917 +; +von Huene 1925 +; +Cluver 1971 +; +Colbert 1974 +; + +Kammerer +et al. +2011 + +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus +sp. + +(“ + +Lystrosaurus +weidenreichi + +” +Young, 1939 +) + +China ( +Young 1939 +, +1946 +; +Brink 1951 +; +Cluver 1971 +; +Colbert 1974 +, +1982 +; +King 1988 +; +Li 1988 +; +Lucas 2001 +; + +Liu +et al. +2002 + +; +Li & Sun 2008 +; +Camp & Liu 2011 +) + +Jiucaiyuan ( +Li 1988 +; +Lucas 2001 +; +Li & Sun 2008 +) + +Induan ( + +Tong +et al. +2018 + +) +
+ +Lystrosaurus +sp. + + +Antarctica ( +Colbert 1970a +, b, +1971b +, +1974 +; + +Elliot +et al. +1970 + +) + +Lower Fremouw ( + +Elliot +et al. +1970 + +; +Colbert 1974 +)Olenekian ( + +Elliot +et al. +2017 + +) +
+Australia ( +Northwood 1997 +) + +Arcadia ( +Northwood 1997 +) + +Early Triassic ( +Northwood 1997 +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
TaxonCountryFormation +Age +
+China ( +Young 1939 +, +1946 +; + +Liu +et al. +2002 + +; +Maisch & Matzke 2014 +; +Li 2015 +; + +Han +et al. +2021 + +; + +Kulik +et al. +2021 + +) + +Upper Guodikeng ( + +Liu +et al. +2002 + +; +Li 2015 +) + +Induan ( + +Tong +et al. +2018 + +) +
+Jiucaiyuan ( +Maisch & Matzke 2014 +; + +Han +et al. +2021 + +; + +Kulik +et al. +2021 + +) + +Induan ( + +Tong +et al. +2018 + +) +
+Not given ( +Young 1939 +) + +?Induan ( + +Tong +et al. +2018 + +) +
+India ( +Bandyopadhyay & Ray 2020 +) + +Kamthi ( +Bandyopadhyay & Ray 2020 +) + +Early Triassic ( +Prasad & Pundir 2020 +) +
+South Africa ( +Broom 1903 +, +1908a +, b; +Watson 1913 +; +Brink 1951 +; +Janensch 1952 +; +Haughton 1963 +; Crompton & Hotton 1967; +Cruickshank 1968 +; +Cluver 1971 +; +DeFauw 1986 +; +Groenewald 1991 +; +Cox 1998 +; +MacRae 1999 +; Damiani & Welman 2001; + +Damiani +et al. +2003 + +; +Neveling 2004 +; +Surkov & Benton 2004 +; + +Grine +et al. +2006 + +; + +Jasinoski +et al. +2010 + +, +2014 +; +Modesto & Botha-Brink 2010 +; +Jasinoski & Chinsamy-Turan 2012 +; +Angielczyk & Rubidge 2013 +; + +Botha-Brink +et al. +2014 + +; + +Gastaldo +et al. +2017 + +, +2019 +; Olivier +et al. +2017; +Krummeck & Bordy 2018 +; + +Whitney +et al. +2019 + +; + +Botha +et al. +2020 + +; + +Knaus +et al. +2021 + +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +) + +“ + +Lystrosaurus +AZ + +” ( +Cluver 1971 +; +DeFauw 1986 +; +Groenewald 1991 +; +MacRae 1999 +; +Damiani & Welman 2001 +; + +Damiani +et al. +2003 + +; +Neveling 2004 +; Surkov & Benton 2004; + +Grine +et al. +2006 + +; +Modesto & Botha-Brink 2010 +; + +Botha-Brink +et al. +2014 + +; + +Olivier +et al. +2017 + +; + +Gastaldo +et al. +2017 + +; +Krummeck & Bordy 2018 +; + +Whitney +et al. +2019 + +; + +Botha +et al. +2020 + +; + +Knaus +et al. +2021 + +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Lystrosaurus declivis +AZ + +( +Olroyd 2022 +; +Olroyd & Sidor 2022 +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+? + +Lystrosaurus +sp. + +(“ +Dicynodon seeleyi +” + +Broili, 1908 + +) + +South Africa ( +Broili 1908 +; + +Kammerer +et al. +2011 + +) + +“? + +Lystrosaurus +AZ + +” ( + +Kammerer +et al. +2011 + +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+? + +Lystrosaurus +sp. + + +China ( +Yin & Peng 2000 +) Russia ( +Ochev 1992 +; + +Shishkin +et al. +2000 + +) + +Shaofanggou ( +Yin & Peng 2000 +) Vokhmian ( +Ochev 1992 +; + +Shishkin +et al. +2000 + +) + +Olenekian ( + +Tong +et al. +2018 + +) Induan ( +Ivakhnenko 2008 +) +
+cf. + +Lystrosaurus +sp. + + +Mozambique ( + +Araújo +et al. +2020 + +) + +Fubué ( + +Araújo +et al. +2020 + +) + +Early Triassic ( + +Araújo +et al. +2020 + +) +
+ +Moghreberia +nmachouensis + +Dutuit, 1980 + +Morocco ( +Dutuit 1980 +, +1988 +; +Bandyopadhyay 1988 +; +King 1988 +; +Gauffre 1993 +; + +Olivier +et al. +2017 + +; +Kammerer 2018 +) + +Argana t5 ( +Dutuit 1988 +, +1989b +; +Gauffre 1993 +) + +Carnian/?Norian ( + +Kammerer +et al. +2012 + +) +
+ +Myosaurus gracilis + +Haughton, 1917 + +Antarctica ( +Cosgriff & Hammer 1979 +; +Hammer & Cosgriff 1981 +; +DeFauw 1989 +) + +Lower Fremouw ( +Cosgriff & Hammer 1979 +; +Hammer & Cosgriff 1981 +; +DeFauw 1989 +) + +Olenekian ( + +Elliot +et al. +2017 + +) +
+South Africa ( +Haughton 1917 +, +1924 +; +von Huene 1925 +; +Broom 1932 +; +Haughton & Brink 1954 +; +Kitching 1968 +, +1977 +; +Keyser 1974 +; +Cluver 1974 +; +Keyser & Cruickshank 1979 +; +Hammer & Cosgriff 1981 +; +Brink 1986 +; +King 1988 +; +Botha & Smith 2006 +; +Surkov 2006 +; +Botha & Smith 2020 +; +Nicolas & Rubidge 2010 +; + +Smith +et al. +2012 + +, +2020b +; + +Benoit +et al. +2018 + +; + +Viglietti +et al. +2022 + +) + + +Lystrosaurus declivis +AZ + +( +Botha & Smith 2020 +; + +Smith +et al. +2020b + +; + +Viglietti +et al. +2022 + +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ +Parakannemeyeria + + +chengi +Liu, 2004 + + +China ( +Liu 2004 +; +Li & Sun 2008 +; +Li 2015 +) + +Lower Karamay ( +Liu 2004 +; +Li 2015 +) + +Anisian ( + +Tong +et al. +2018 + +) +
+ +Parakannemeyeria +dolichocephala + +Sun, 1960 + +China ( +Sun 1960 +, +1963 +; +Young 1964 +; Keyser & Cruickshank Upper Ermaying ( +Sun 1963 +; +Brink 1986 +; King 1979; +Brink 1986 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) 1988; +Li & Sun 2008 +; +Li 2015 +) + +Late Anisian ( +Liu et al. 2017 +) +
+ +Parakannemeyeria +ningwuensis + +Sun, 1963 + +China ( +Sun 1963 +; +Young 1964 +; +Keyser & Cruickshank 1979 +; +Cheng 1980 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +; +Liu 2022 +) + +Ermaying ( +Sun 1963 +; +Cheng 1980 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +; +Liu 2022 +) + +Anisian ( + +Liu +et al. +2017 + +) +
+ +Parakannemeyeria +shenmuensis + +Cheng, 1980 + +China ( +Cheng 1980 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) + +Upper Ermaying ( +Cheng 1980 +; +King 1988 +; +Li & Sun 2008 +) + +Late Anisian ( +Liu et al. 2017 +) +
+ +Parakannemeyeria + + +youngi +Sun, 1963 + + +China ( +Sun 1963 +; +Young 1963 +, +1964 +; Keyser & Cruickshank Upper Ermaying ( +Sun 1963 +; +Brink 1988 +; King 1979; +Yuhe 1983 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; 1988; +Li & Sun 2008 +; +Li 2015 +) +Li 2015 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ +Parakannemeyeria + + +cf. +youngi +Sun, 1963 + + +China ( +Sun 1963 +; +Young 1964 +) + +Upper Ermaying ( +Sun 1963 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ +Parakannemeyeria +sp. + + +China ( +Sun 1963 +; +Young 1964 +; +Cheng 1980 +; +Liu 2015 +; +Liu 2022 +) + +Upper Ermaying ( +Sun 1963 +; +Cheng 1980 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+Lower Ermaying ( +Liu 2022 +) + +Early Anisian ( + +Liu +et al. +2017 + +) +
+Tongchuan I ( +Liu 2015 +) + +Late Anisian ( + +Tong +et al. +2018 + +) +
+ +Pentasaurus goggai + +Kammerer, 2018 + +South Africa ( +Kammerer 2018 +; + +Viglietti +et al. +2020b + +) + +Lower Elliott + +Scalenodontoides +AZ + +( + +Viglietti +et al. +2020b + +) + +Norian-Rhaetian ( + +Bordy +et al. +2020 + +) +
+ +Placerias hesternus + +Lucas, 1904 + +United States ( +Lucas 1904 +; +Cross & Howe 1905 +; +Cross 1908 +; +Darton 1910 +; +von Huene 1911 +, +1926a +, +1935 +, +1936 +; +Broom 1914 +; +Case 1915 +; +Gregory 1917 +; +Pearson 1924a +; +Camp 1956 +; +Camp & Welles 1956 +; +Cox 1965 +; +Baird & Patterson 1968 +; +Keyser 1974 +; +Keyser & Cruickshank 1979 +; +Rowe 1979 +; +Walter 1985 +; +Brink 1986 +, +1988 +; +King 1988 +; + +Olsen +et al. +1989 + +; + +Huber +et al. +1993 + +; +Long & Murry 1995 +; +Lucas 1998 +; +Olsen & Huber 1998 +; + +Fiorillo +et al. +2000 + +; +Heckert & Lucas 2003 +; + +Green +et al. +2010 + +; +Parker & Martz 2011 +; + +Kammerer +et al. +2013 + +; +Kammerer 2018 +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +) + +Bluewater Creek ( +Lucas 1998 +; Lucas & Heckert 2002; +Heckert & Lucas 2003 +) + +Norian ( + +Ramezani +et al. +2014 + +) +
+Cooper Canyon ( +Lehman & Chatterjee 2005 +) + +Norian ( + +Martz +et al. +2013 + +) +
+Pekin ( +Baird & Patterson 1968 +; + +Olsen +et al. +1989 + +; + +Huber +et al. +1993 + +; +Lucas & Hunt 1993a +; +Lucas 1998 +; +Olsen & Huber 1998 +) + +Carnian ( + +Heckert +et al. +2017 + +) +
+Petrified Forest ( +Murry & Long 1989 +; +Long & Murry 1995 +; +Lucas 1998 +; + +Fiorillo +et al. +2000 + +; +Lucas & Heckert 2002 +; + +Green +et al. +2010 + +) + +Norian ( + +Ramezani +et al. +2014 + +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
TaxonCountryFormation +Age +
+ +Placerias +sp. + + +United States ( +Murry & Long 1989 +; +Heckert 1997 +; Heckert & +Lucas 2003 +) + +Bluewater Creek ( +Heckert 1997 +; +Heckert & Lucas 2003 +) + +Norian ( + +Ramezani +et al. +2014 + +) +
+Petrified Forest ( +Murry & Long 1989 +) + +Norian ( + +Ramezani +et al. +2014 + +) +
+? + +Placerias +sp. + + +United States ( +Long & Murry 1995 +; +Parker & Martz 2011 +) + +Petrified Forest ( +Long & Murry 1995 +; +Parker & Martz 2011 +) + +Norian ( + +Ramezani +et al. +2014 + +) +
+ +Rabidosaurus cristatus + +Kalanadze, 1970 + +Russia ( +Kalandadze 1970 +; +Keyser & Cruickshank 1979 +; +King 1988 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; +Ivakhnenko 2008 +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+ +Rechnisaurus +cristarhynchus + +Chowdhury, 1970 + +India ( + +Jain +et al. +1964 + +; +Chowdhury 1970 +; Keyser & Cruickshank 1979; +Brink 1986 +; +King 1988 +; +Bandyopadhyay 1989 +; +Bandyopadhyay & Ray 2020 +) + +Yerrapalli ( + +Jain +et al. +1964 + +; +Chowdhury 1970 +; +Keyser & Cruickshank 1979 +; +Brink 1986 +; +King 1988 +; +Bandyopadhyay 1989 +; Bandyopadhyay & Ray 2020) + +Anisian or younger ( + +Ottone +et al. +2014 + +) +
+Tanzania ( +Cox 1991 +; +Renaut 2000 +; +Surkov & Benton 2004 +; + +Kammerer +et al. +2017 + +) + +Manda ( +Cox 1991 +; +Surkov & Benton 2004 +; + +Kammerer +et al. +2017 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ +Repelinosaurus +robustus + +Olivier, Battail, Bourquin, Rossignol, Steyer & Jalil, 2019 + +Laos ( + +Olivier +et al. +2019 + +) + +Purple Claystone ( + +Olivier +et al. +2019 + +) + +?Late Permian ( +Liu 2020 +)/ Early Triassic +
+( + +Olivier +et al. +2019 + +) +
+ +Rhadiodromus klimovi + +( +Efremov, 1938 +) + +Russia ( +Efremov 1938 +, +1940 +, +1951 +; +Vjuschkov 1969 +; Kalan-dadze 1970; +King 1988 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; +Ivakhnenko 2008 +) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+ +Rhadiodromus mariae + +Surkov, 2003 + +Russia ( +Surkov 2003 +; +Surkov & Benton 2004 +; +Ivakhnenko 2008 +) + +Upper Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+ +Rhinodicynodon gracile + +Kalanadze, 1970 + +Russia ( +Kalandadze 1970 +; +Keyser & Cruickshank 1979 +; +King 1988 +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Surkov 1998a +, b; +Battail & Surkov 2000 +; +Ivakhnenko 2008 +; + +Hancox +et al. +2013 + +) + +Upper Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+ +Sangusaurus edentatus + +Cox, 1969 + +Zambia ( +Cox 1969 +; +King 1988 +; + +Angielczyk +et al. +2014 + +, +2017 +; + +Peecook +et al. +2018 + +) + +Ntawere ( +Cox 1969 +; +King 1988 +; + +Angielczyk +et al. +2014 + +, +2017 +; + +Peecook +et al. +2018 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ +Sangusaurus +parringtonii + +Cruickshank, 1986a + +Tanzania ( +Cruickshank 1986a +, b; +Surkov & Benton 2004 +; + +Angielczyk +et al. +2017 + +; + +Kammerer +et al. +2017 + +) + +Manda ( +Cruickshank 1986a +, b; Surkov & Benton 2004; + +Angielczyk +et al. +2017 + +; + +Kammerer +et al. +2017 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ +Sangusaurus +sp. + +(“ +Ischigualastia +sp.”/” +Stahleckeria + +potens + +”) + +Brazil ( +Schwanke-Peruzzo 1990 +; + +Schwanke-Peruzzo & +Araújo-Barbarena 1995 + +; +Lucas 2002 +; + +Langer +et al. +2007 + +; + +Dassie 2014 + +; +Kammerer & Ordoñez 2021 +) + +Santa Maria ( +Schwanke-Peruzzo 1990 +; +Schwanke-Peruzzo & Araújo-Barbarena 1995 +; +Lucas 2002 +; + +Langer +et al. +2007 + +; Dassie 2014; +Kammerer & Ordoñez 2021 +) + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/Carnian ( + +Ordoñez +et al. +2020 + +) +
+ +Shaanbeikannemeyeria + +xilougouensis +Cheng, 1980 + +China ( +Cheng 1980 +; +Li 1980 +; +King 1988 +; +Li & Sun 2008 +; + +Li 2015 + +; +Liu 2022 +; + +Escobar +et al. +2023 + +) + +Lower Ermaying ( +Cheng 1980 +; +Li 1980 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +; +Liu 2022 +) + +Early Anisian ( + +Liu +et al. +2017 + +) +
+Upper Heshanggou ( +Liu 2022 +) + +Early Anisian ( + +Liu +et al. +2017 + +) +
+ +Shansiodon wangi +Yeh, 1959 + + +China ( +Yeh 1959 +; +Sun 1963 +; +Young 1964 +; +Keyser & Cruickshank 1979 +; +Cheng 1980 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; + +Kammerer +et al. +2013 + +; +Li 2015 +) + +Upper Ermaying ( +Yeh 1959 +; +Sun 1963 +; +Cheng 1980 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ +Shanisodon +wuhsiangensis + +Yeh, 1959 + +China ( +Yeh 1959 +; +Keyser & Cruickshank 1979 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; + +Hancox +et al. +2013 + +; +Li 2015 +) + +Upper Ermaying ( +Yeh 1959 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; + +Hancox +et al. +2013 + +; +Li 2015 +) + +Late Anisian ( +Liu et al. 2017 +) +
+ +Shansiodon +sp. + + +China ( +Yeh 1959 +; +Sun 1963 +; +Young 1964 +; +Cheng 1980 +; +King 1988 +) + +Upper Ermaying ( +Yeh 1959 +; +Sun 1963 +; +Cheng 1980 +; +King 1988 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+South Africa ( +Hancox 1998 +; +Nicolas & Rubidge 2010 +; + + +Smith +et al. +2012 + + +, +2020b +; + +Hancox +et al. +2013 + +, +2020 +; + +Viglietti +et al. +2022 + +; + +Escobar +et al. +2023 + +) + +Cynognathus +AZ +Cricodon-Ufudocyclops +SZ ( + +Hancox +et al. +2020 + +; + +Smith +et al. +2020b + +; + +Viglietti +et al. +2022 + +) + +Late Anisian-Carnian ( + +Hancox +et al. +2020 + +) +
+? + +Shansiodon +sp. + + +China ( +Sun 1963 +; +Young 1964 +) + +Upper Ermaying ( +Sun 1963 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ +Sinokannemeyeria +baidaoyuensis + +Liu, 2015 + +China ( +Liu 2015 +) + +Tongchuan I ( +Liu 2015 +) + +Late Anisian ( +Tong et al. 2018 +) +
+ +Sinokannemeyeria +pearsoni + +Young, 1937 + +China ( +Young 1937 +, +1946 +, +1964 +; +Sun 1963 +; +Keyser & Cruickshank 1979 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) + +Upper Ermaying ( +Sun 1963 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) + +Late Anisian ( +Liu et al. 2017 +) +
+cf. + +Sinokannemeyeria + + +pearsoni +Young, 1937 + + +China ( +Sun 1963 +; +Young 1964 +) + +Upper Ermaying ( +Sun 1963 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ +Sinokannemeyeria + +sanchuanheensis +Cheng, 1980 + +China ( +Cheng 1980 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) + +Upper Ermaying ( +Cheng 1980 +; +King 1988 +; + +Li & +Sun 2008 + +; +Li 2015 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ +Sinokannemeyeria + +yingchiaoensis +Sun, 1963 + +China ( +Sun 1963 +; +Keyser & Cruickshank 1979 +; +Brink 1988 +; + +King 1988 + +; +Li & Sun 2008 +; +Li 2015 +) + +Upper Ermaying ( +Sun 1963 +; +Brink 1988 +; +King 1988 +; +Li & Sun 2008 +; +Li 2015 +) + +Late Anisian ( + +Liu +et al. +2017 + +) +
+ + + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
TaxonCountryFormationAge
+ +Sinokannemeyeria +sp. + + +China ( +Sun 1963 +; +Cheng 1980 +; +King 1988 +) + +Upper Ermaying ( +Sun 1963 +; +Cheng 1980 +; +King 1988 +) + +Late Anisian ( + +Liu +et al. +2017) + +
+ +Stahleckeria potens + +von Huene, 1935 + +Brazil ( +von Huene 1935 +, +1936 +, +1949 +; +Romer & Price 1944 +; +Camp 1956 +; +Bonaparte 1970 +, +1978 +; Keyser & Cruickshank 1979; +King 1988 +; +Brink 1988 +; +Walter 1989 +; +Lucas 1993a +, +2002 +; +Maisch 2001 +, +2021 +; +Surkov & Benton 2004 +; +Vega-Dias & Schwanke 2004a +; + +Vega-Dias +et al. +2005 + +; + +Langer +et al. +2007 + +; + +Abdala +et al. +2013 + +; +Vega & Maisch 2014 +; +Dassie 2014 +; + +Kammerer +et al. +2017 + +; Kammerer 2018; +Mancuso & Irmis 2019 +; + +Ordoñez +et al. +2019 + +; + +Schultz +et al. +2020 + +; +Kammerer & Ordoñez 2021 +) + +Santa Maria + +Dinodontosaurus +AZ + +( +Lucas 2002 +; + +Langer +et al. +2007 + +; + +Abdala +et al. +2013 + +; +Dassie 2014 +; + +Martinelli +et al. +2017 + +; Mancuso & Irmis 2019; + +Schultz +et al. +2020 + +; +Maisch 2021 +) + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/Carnian ( + +Ordoñez +et al. +2020 + +) +
+Namibia ( + +Abdala +et al. +2013 + +; +Mancuso & Irmis 2019 +; +Kammerer & Ordoñez 2021 +; +Maisch 2021 +; +Olroyd 2022 +; +Olroyd & Sidor 2022 +; + +Preuschoft +et al. +2022 + +) + +Omingonde ( + +Abdala +et al. +2013 + +; +Mancuso & Irmis 2019 +; +Kammerer & Ordoñez 2021 +; +Maisch 2021 +) + +Anisian-Ladinian ( + +Wynd +et al. +2018 + +; + +Zieger +et al. +2020 + +) +
+ +Stahleckeria +sp. + + +Argentina ( +Mancuso & Irmis 2019 +; + +Escobar +et al. +2021 + +; +Kammerer & Ordoñez 2021 +) + +Chañares +Massetognathus-Chanaresuchus +AZ ( +Mancuso & Irmis 2019 +; + +Escobar +et al. +2021 + +) + +Early Carnian ( + +Ezcurra +et al. +2017 + +) +
+Stahleckeriidae +indet. (“ + +Angonisaurus +sp. + +”) + +Antarctica ( + +Sidor +et al. +2014 + +; + +Kammerer +et al. +2019 + +) + +Upper Fremouw ( + +Sidor +et al. +2014 + +; Kammerer +et al. +2019) + +Late Anisian-Ladinian ( + +Elliot +et al. +2017 + +) +
+Stahleckeriidae +indet. + +Brazil ( +Maisch 2021 +) + +Santa Maria + +Dinodontosaurus +AZ + +( +Maisch 2021 +) + +Ladinian-Carnian ( + +Philipp +et al. +2018 + +)/Carnian +
+( + +Ordoñez +et al. +2020 + +) +
+India ( +Bandyopadhyay 1988 +; +Bandyopadhyay & Sengupta 1999 +; +Bandyopadhyay & Ray 2020 +) + +Denwa ( +Bandyopadhyay 1988 +; Bandyopadhyay & Sengupta 1999; +Bandyopadhyay & Ray 2020 +) + +Anisian or younger ( + +Peecook +et al. +2018 + +) +
+South Africa ( +Watson 1917 +; +Hancox & Rubidge 1994 +; +Govender 2005 +) + +Cynognathus +AZ +Trirachodon-Kannemeyeria +SZ +( +Govender 2005 +; + +Hancox +et al. +2020 + +) + +Early Anisian ( + +Hancox +et al. +2020 + +) +
+Tanzania ( + +Kammerer +et al. +2017 + +) + +Manda ( + +Kammerer +et al. +2017 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
Stahleckeriinae indet. +Argentina ( +Cox 1968 +; +Bonaparte 1997 +; + +Escobar +et al. +2021 + +) + +Chañares +Tarjadia +AZ ( + +Escobar +et al. +2021 + +) + +Ladinian +/Carnian ( + +Ezcurra +et al. +2017 + +) +
+United States ( +Lucas & Hunt 1993a +; +Long & Murry 1995 +; + +Green +et al. +2005 + +; +Green 2012 +; + +Kammerer +et al. +2013 + +) + +Pekin ( + +Green +et al. +2005 + +; +Green 2012 +; Kammerer +et al. +2013) + +Carnian ( + +Heckert +et al. +2017 + +) +
+Santa Rosa ( +Lucas & Hunt 1993a +; +Long & Murry 1995 +; + +Kammerer +et al. +2013 + +) + +Carnian ( + +Hartman +et al. +2015 + +) +
+ +Sungeodon +kimkraemerae + +Maisch & Matzke, 2014 + +China ( +Maisch & Matzke 2014 +) + +Jiucaiyuan ( +Maisch & Matzke 2014 +) + +Induan ( + +Tong +et al. +2018 + +) +
+ +Tetragonias njalilus + +( +von Huene, 1942 +) + +Tanzania ( +von Huene 1942 +; +Haughton & Brink 1954 +; +Cruickshank 1964 +, +1967 +; +Bonaparte 1966a +; +Keyser & Cruickshank 1979 +; +King 1988 +; +Surkov & Benton 2004 +; +FrÖbisch 2006 +; + +Kammerer +et al. +2011 + +, +2017 +; + +Hancox +et al. +2013 + +; + +Preuschoft +et al. +2022 + +) + +Manda ( +von Huene 1942 +; +Haughton & Brink 1954 +; +Bonaparte 1966b +; +Cruickshank 1967 +; +Keyser & Cruickshank 1979 +; +King 1988 +; +Surkov & Benton 2004 +; +FrÖbisch 2006 +; + +Kammerer +et al. +2011 + +, +2017 +; + +Hancox +et al. +2013 + +; + +Preuschoft +et al. +2022 + +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ +Tetragonias +sp. + + +Argentina ( +Bonaparte 1981 +; +Domnanovich & Marsicano 2012 +; +Kammerer & Ordoñez 2021 +) + +Quebrada de los Fósiles ( +Bonaparte 1981 +; +Domnanovich & Marsicano 2012 +; Kammerer & Ordoñez 2021) + +?Ladinian-Carnian ( + +Ottone +et al. +2014 + +) +
+ +Ufudocyclops + +mukanelai +Kammerer, Viglietti, Hancox, Butler & Choiniere, 2019 + +South Africa ( +Hancox & Rubidge 1996 +; +Hancox 1998 +; + +Nicolas & Rubidge 2010 + +; + +Smith +et al. +2012 + +, +2020b +; + +Hancox +et al. +2013 + +, +2020 +; + +Kammerer +et al. +2019 + +; + +Viglietti +et al. +2022 + +) + +Cynognathus +AZ +Cricodon-Ufudocyclops +SZ ( + +Hancox +et al. +2020 + +; + +Smith +et al. +2020b + +; + +Viglietti +et al. +2022 + +) + +Late Anisian-Carnian +( + +Hancox +et al. +2020 + +) +
+ +Uralokannemeyeria + +vjuschkovi +Danilov, 1971 + +Russia ( +Danilov 1971 +, +1973 +; +Keyser & Cruickshank 1979 +; + +King 1988 + +; + +Shishkin +et al. +1995 + +; + +Ivakhnenko +et al. +1997 + +; +Battail & Surkov 2000 +; +Surkov & Benton 2004 +; Ivakhnenko 2008) + +Lower Donguz ( +Surkov 2003 +) + +Anisian ( +Ivakhnenko 2008 +) +
+ +Vinceria andina + +Bonaparte, 1969 + +Argentina ( +Bonaparte 1969 +, +1970 +, +1978 +; +Keyser 1974 +; +Keyser & Cruickshank 1979 +; +King 1988 +; +Domnanovich & Marsicano 2006a +, +2012 +; + +Hancox +et al. +2013 + +; + +Ordoñez +et al. +2019 + +; +Kammerer & Ordoñez 2021 +) + +Cerro de las Cabras ( +Bonaparte 1969 +; +King 1988 +; +Domnanovich & Marsicano 2006a +, +2012 +; +Kammerer & Ordoñez 2021 +) + +Late Anisian-early Ladinian ( + +Cariglino +et al. +2016 + +) +
+ +Wadiasaurus indicus + +Chowdhury, 1970 + +India ( + +Jain +et al. +1964 + +; +Chowdhury 1970 +; Keyser & Cruickshank 1979; +Bandyopadhyay 1988 +; +King 1988 +; +Ray 2006 +; + +Ray +et al. +2009 +a, b, 2010 + +; +Bandyopadhyay & Ray 2020 +) + +Yerrapalli ( + +Jain +et al. +1964 + +; +Chowdhury 1970 +; +Keyser & Cruickshank 1979 +; Bandyopadhyay 1988; +King 1988 +; +Ray 2006 +; + +Ray +et al. +2009 +a, b, 2010 + +; +Bandyopadhyay & Ray 2020 +) + +Anisian or younger ( + +Ottone +et al. +2014 + +) +
+ +Woznikella triradiata + +n. gen., n. sp. + +Germany ( +Schoch 2012 +) + +Stuttgart ( +Schoch 2012 +) + +Carnian ( +Schoch 2012 +) +
+Poland ( + +Sulej +et al. +2011 + +; +Racki & Lucas 2020 +) + +Grabowa ( + +Szulc +et al. +2015b + +; +Racki & Lucas 2020 +) + +Carnian ( + +Sulej +et al. +2011 + +)/ Norian (( + +Szulc +et al. +2015b + +) +
+ + + + + +rostris +( +Sun, 1978 +) + + + + + + + +
+ +Xiyukannemeyeria +brevi- + + +China ( +Sun 1978 +; +King 1988 +; +Liu & Li 2003 +; +Li & Sun 2008 +) +
+Lower Karamay ( +Liu & Li 2003 +) + +Anisian ( + +Tong +et al. +2018 + +) +
+ + +TABLE +1. — Continuation. + + + + + + + + + + + + + + + + + + +
+Taxon + +Country + +Formation + +Age +
+ +Zambiasaurus +submersus + +Cox, 1969 + +Zambia ( + +Attridge +et al. +1964 + +; +Cox 1969 +; +King 1988 +; + +Angielczyk +et al. +2014 + +; +Kammerer 2018 +; + +Peecook +et al. +2018 + +; +Li 2015 +) + +Ntawere ( + +Attridge +et al. +1964 + +; +Cox 1969 +; +Keyser & Cruickshank 1979 +; +King 1988 +; + +Angielczyk +et al. +2014 + +; +Kammerer 2018 +; + +Peecook +et al. +2018 + +; +Li 2015 +) + +Anisian or younger ( + +Wynd +et al. +2018 + +) +
+ + +Dentary + + +The dentaries ( +ZPAL +V +. 34/1/3; +Fig. 4 +) are fused together at the symphysis. Each dentary is a massive yet narrow bone with a dorsally projected and dorsolaterally pointed anterodorsal tip, triangular in cross-section. Between the tips there is a narrow notch ( +Fig. 4D +), like in + +Angonisaurus cruickshanki + +and + +Kannemeyeria simocephalus + +(see +Pearson 1924a +; +Cox & Li 1983 +; +Renaut 2000 +; + +Hancox +et al. +2013 + +). + + +In lateral view the dentary is gently hooked ( +Fig. 4D +), similarly to that of + +Angonisaurus cruickshanki + +, + +Dolichuranus primaevus + +, + +Kannemeyeria simocephalus + +, + +Kannemeyeria lophorhinus + +, “ + +Kannemeyeria +” +latirostris + +, + +Kombuisia frerensis + +, + +Kombuisia antarctica + +, + +Lystrosaurus +spp. + +, + +Myosaurus gracilis + +, + +Parakannemeyeria dolichocephala + +, + +Rhinodicynodon gracile + +, + +Sangusaurus parringtonii + +, + +Tetragonias njalilus + +, + +Wadiasaurus indicus + +, and + +Xiyukannemeyeria brevirostris + +(see +Broom 1923 +, +1937 +; +Pearson 1924a +; +von Huene 1942 +; +Sun 1960 +; +Cruickshank 1967 +, +1986a +; +Crozier 1970 +; +Kalandadze 1970 +; +Cluver 1971 +, +1974 +; +Keyser 1973 +; +Hotton 1974 +; +Hammer & Cosgriff 1981 +; +Cox & Li 1983 +; +Bandyopadhyay 1988 +; +Renaut 2000 +; +Liu & Li 2003 +; + +Damiani +et al. +2007 + +; +Fröbisch 2007 +; + +Fröbisch +et al. +2010 + +; + +Hancox +et al. +2013 + +; + +Angielczyk +et al. +2014 + +, +2017 +; +Kammerer 2018 +) and unlike the non-hookeed or minimally hooked mandibles of, e.g., + +Dinodontosaurus + +s pp., + +Ischigualastia jenseni + +, + +Parakannemeyeria ningwuensis + +, + +Sinokannemeyeria yingchiaoensis + +, + +Stahleckeria potens + +, and + +Vinceria andina + +(see +von Huene 1935 +; +Romer & Price 1944 +; +Camp 1956 +; +Sun 1963 +; +Cox 1965 +, +1968 +; +Lucas 2002 +; +Domnanovich & Marsicano 2012 +; + +Abdala +et al. +2013 + +; +Kammerer 2018 +; +Kammerer & Ordoñez 2021 +; + +Escobar +et al. +2023 + +). Unlike in + +Dinodontosaurus brevirostris + +, the labial (tomial) edge is continuous with the posterior part of the bone and directed either dorsally or (in the anteriormost part) posterodorsally, but never predominantly anterodorsally ( +Cox 1968 +; +Kammerer & Ordoñez 2021 +; + +Escobar +et al. +2023 + +). The anterior portion of the lateral dentary surface exhibits a coarsely rugose texture indicating the presence of a rhamphotheca-covered beak. The rugosities extend ventrally, towards the mentum, and posteriorly. The posteroventral limit of the rugose area is concave. From the rostral corner of that concavity, rostrodorsally, crossing the rugose surface towards the dorsal edge of the dentaries, paired, anastomosing vascular grooves are visible. Just anterior to them, shallow, gently expressed lateral grooves similar to those of + +Dolichuranus primaevus + +, + +Kannemeyeria simocephalus + +, + +Kannemeyeria lophorhinus + +, + +Parakannemeyeria ningwuensis + +, + +Sangusaurus parringtonii + +, and + +Sinokannemeyeria yingchiaoensis + +span from the lateral surface of each of the tips of the beak posteroventrally towards the ventral edge of the bone ( +Camp 1956 +; +Sun 1963 +; +Cruickshank 1986a +; +Renaut 2000 +; + +Damiani +et al. +2007 + +; + +Angielczyk +et al. +2017 + +). Like in + +Kannemeyeria simocephalus + +, + +Kannemeyeria lophorhinus + +, and + +Sangusaurus parringtonii + +, the grooves do not follow the ventral curvature of the dentary, but are straight ( +Camp 1956 +; +Renaut 2000 +; + +Angielczyk +et al. +2017 + +). Paired, more defined grooves coupled with paired ridges are also present laterally on the dentaries of at least the South African + +Lystrosaurus +spp. + +( +Cluver 1971 +). The lateral dentary shelf ( +Cluver 1971 +) is not developed, but the lateral surfaces of the posterior part of the dentary above the mandibular fenestra show a laterally symmetrical, rounded, indistinct broadening at the level of the split of the dorsal (tomial), medial (lingual), and lateral (labial) laminae of the dentary ramus, at the roof of the Meckelian canal. This differs significantly from the exceptionally pronounced dentary shelf of + +Pentasaurus goggai +Kammerer, 2018 + +(see +Kammerer 2018 +). + + +The symphysis in dorsal view is relatively long and narrow but the mandible broadens ventrally ( +Fig. 4A +), similar to the mandibles of, e.g., + +Kannemeyeria simocephalus + +, + +Sinokannemeyeria yingchiaoensis + +, and + +Tetragonias njalilus + +(see +Pearson 1924a +; +Sun 1963 +; +Cruickshank 1967 +). It bears a single wide, deep groove in the junction between the left and right rami, as is typical for dicynodonts.The dorsal surface of each ramus shows two parallel longitudinal ridges separated by a marked groove (posterior dentary sulcus posteriorly and dentary table anteriorly; see discussion in +Angielczyk & Rubidge 2013 +) along its length, with the lingual one higher than the labial around the midlength, and diminishing gradually posteriorly and sharply anteriorly. The anterior end of each lingual ridge is associated with a rounded, mediolaterally compressed peg. Anterior to that peg each lingual ridge continues rostrally, but becomes less prominent and gently diverges laterally, making the median trough slightly wider frontally and eventually ending in the lateral point of the beak. This morphology is different than in + +Dinodontosaurus tener + +as figured by +von Huene (1935) +and later by +Lucas & Harris (1996) +, in which the symphyseal part is shorter, the lingual ridges nearly meet anteriorly, and the terminal part of the mandible appears wider and more rounded, but (aside from rostral broadening of the median groove) it is generally similar to that of + +Kannemeyeria simocephalus + +and + +Tetragonias njalilus + +(see +Pearson 1924a +; +Cruickshank 1967 +; +Renaut 2000 +). The lingual ridge in the latter, however, is concave dorsally, lower than the labial, and the anterior pegs appear slightly larger, so they are visible laterally ( +Cruickshank 1967 +). Anteroposteriorly short lingual ridges taller than the labial ridges are also present in + +Angonisaurus cruickshanki + +(“rugosities” of +Cox & Li 1983 +; + +Hancox +et al. +2013 + +). In + +Stahleckeria potens + +the lingual ridge is also taller than the labial ( +von Huene 1935 +; +Camp 1956 +; + +Abdala +et al. +2013 + +; +Kammerer 2018 +). The morphology of + +Woznikella triradiata + +n. gen., n. sp. +is also different than in + +Placerias hesternus + +( +Camp & Welles 1956 +; +Kammerer 2018 +; pers. obs.) in which the labial ridge is consistently taller than the lingual and straight. In the latter species the rostral ends of the lingual ridges are visible in lateral view ( +Camp & Welles 1956 +), but in + +Woznikella triradiata + +n. gen., n. sp. +they are obscured by the very high dorsal labial edges of the rami. The anatomy of the scoop-shaped dorsal surfaces of the symphyseal areas of + +Kombuisia frerensis + +and + +Myosaurus gracilis + +is much simpler and the median groove in these species is wider ( +Cluver 1974 +; +Hotton 1974 +; +Hammer & Cosgriff 1981 +; +Fröbisch 2007 +). + + + +FIG +. 5. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V.34/1/5: +A -D +, right angular in lateral ( +A +), dorsal ( +B +), ventral ( +C +), and medial ( +D +) view. Scale bar: 5 cm. + + + +In dorsoventral aspect, the dentaries have roughly parallel lateral edges in their anterior part and strongly diverge posteriorly. They are much more slender than the dentaries of + +Dolichuranus primaevus + +, + +Kombuisia +spp. + +, + +Lystrosaurus +spp. + +, + +Myosaurus gracilis + +, + +Pentasaurus goggai + +, + +Shaanbeikannemeyeria xilougouensis + +, + +Shansiodon wangi + +, + +Shansiodon wuhsiangensis + +, + +Sinokannemeyeria baidaoyuensis + +, + +Vinceria andina + +, and + +Xiyukannemeyeria brevirostris + +(see +Young 1935 +; +Yeh 1959 +; +Cluver 1971 +, +1974 +; +Keyser 1973 +; +Hotton 1974 +; +Li 1980 +; +Hammer & Cosgriff 1981 +; +Liu & Li 2003 +; + +Damiani +et al. +2007 + +; +Fröbisch 2007 +; + +Fröbisch +et al. +2010 + +; +Domnanovich & Marsicano 2012 +; +Liu 2015 +; +Kammerer 2018 +). + + +The ventral part is complete only in the right ramus and the mentum is strongly bowed, more so than in, e.g., + +Dinodontosaurus brevirostris + +or + +Shaanbeikannemeyeria xilougouensis + +( +Cox 1968 +; +Kammerer & Ordoñez 2021 +; +Liu 2022 +; + +Escobar +et al. +2023 + +). It is, however, strikingly different from the top-heavy, rostrally vertical dentaries of + +Sungeodon kimkraemerae + +and + +Vinceria andina + +, in which the rostral and the posteroventral surfaces are set at an acute angle ( +Domnanovich & Marsicano 2012 +; +Maisch & Matzke 2014 +). The lateral bone lamella of the dentary in + +Woznikella triradiata + +n. gen., n. sp. +becomes thinner ventrally, forming a large surface for a suture with the splenial (not preserved) and angular. On the anteroventral (mental) surface of the dentaries two sharply defined, parallel, longitudinal grooves are separated by a wide ridge with a narrow but distinct third groove running medially along the symphysis ( +Fig. 4B, D +). A ridge-like sagittal structure is also present in + +Kannemeyeria simocephalus + +, + +Parakannemeyeria ningwuensis + +, + +Pentasaurus goggai + +, + +Placerias hesternus + +, + +Sinokannemeyeria sanchuanheensis + +, + +Sungeodon kimkraemerae + +, + +Tetragonias njalilus + +, and + +Wadiasaurus indicus + +( +Camp & Welles 1956 +; +Sun 1963 +; +Cruickshank 1967 +; +Cheng 1980 +; +Bandyopadhyay 1988 +; +Renaut 2000 +; +Maisch & Matzke 2014 +; +Kammerer 2018 +). In + +Tetragonias njalilus + +, however, it projects its own tip from the anterior edge of the beak ( +Cruickshank 1967 +). The tips of the beak bear narrow, longitudinal grooves close to their lateral edges. + + +The Meckelian canal exposed on the ventral surface of the dorsal part of the disarticulated dentary takes the form of a longitudinal groove which is widest anteriorly and narrows posteriorly ( +Fig. 4B +). This groove ends in a point in contrast to + +Placerias hesternus + +, in which it forms a vertical edge ( +Camp & Welles 1956 +and pers. obs.). In most dicynodonts at least part of this groove accommodates an anterior process of the surangular. + + +The posteriormost part of the dentary is formed by a thin blade, which contacted the surangular. Its ventral edge contributed to the mandibular fenestra. Below that, a plate-like process for the angular was present, but only its base is preserved on the right side of +ZPAL +V +. 34/1/3. Given the imprint on the left angular ( +ZPAL +V +. 34/1/81), this process was relatively large, about one fourth to one third the length of the dentary, very different than, e.g., the unusually small process of + +Dolichuranus primaevus + +(see + +Damiani +et al. +2007 + +). Unlike + +Myosaurus gracilis + +and + +Tetragonias njalilus + +, the dentary was apparently level with the more posterior parts of the mandible, not set at an angle ( +von Huene 1942 +; +Cruickshank 1967 +; +Cluver 1974 +). + + + +FIG +. 6. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1/4: +A -E +, right posterior mandibular bone complex in dorsal ( +A +), lateral ( +B +), medial ( +C +), ventral ( +D +), and posterior ( +E +) view. Scale bar: 5 cm. + + + +ZPAL +V +. 34/1/3 is notably similar to +SMNS +91416, the partial mandible from the Stuttgart Formation in +Bavaria +( +Germany +) described by +Schoch (2012) +. The similarities include: general shape in lateral view (hooked, with bowed mental surface); presence of a median notch; presence and shape of anterodorsal tips; rugosity of the surface; shape of the ramphotheca, with distinctly concave posteroventral edge; position and shape of a lateral groove; position and shape of lateral vascular grooves; presence of a symphyseal ridge on the rostroventral (mental) surface of the mandible flanked by lateral grooves and with a sagittal groove (less distinct than in +ZPAL +V +. 34/1/3, possibly due to difference in ontogenetic age); presence of what appears to be a lingual ridge higher than the labial edge. Based on the published photographs, +SMNS +91416 appears to be wider than +ZPAL +V +. 34/1/3, but this, at least to some extent, is caused by the foreshortening due to the perspective and slightly different scale of the panels captioned as the dorsal and ventral view. Because of the morphological similarity, geographical and temporal proximity, and absence of other dicynodonts in the European Carnian, we therefore tentatively refer +SMNS +91416 to + +Woznikella triradiata + +n. gen., n. sp. + + + +FIG +. 7. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1: +A -F +, reconstruction of the skull ( +A +, +C +, +E +) and mandible ( +B +, +D +, +F +) in anterior ( +A +, +B +), dorsal ( +C +, +D +), and lateral left ( +E +, +F +) view. Scale bar: 5 cm. + + + + +Angular + + + +Only the posterior portion of the left angular ( +ZPAL +V +. 34/1/81) is preserved, and its posterodorsal edge is incomplete. The right angular ( +ZPAL +V +. 34/1/5; +Fig. 5 +) is almost complete. The ventral and dorsal edges are deeply concave in lateromedial aspect. The reflected lamina is very well preserved in +ZPAL +V +. 34/1/5, even though it is a thin plate. Its medial surface is slightly concave, whereas the lateral surface is convex. Medial to the reflected lamina is the main body of the angular, and between the two there is a deep notch, the anterior limit of which is exposed in lateral view. In lateromedial aspect, the main body protrudes dorsally above the reflected lamina ( +Fig. 4A, D +). Unfortunately, the dorsal edge of the angular is broken in both specimens so its exact height remains unknown – the probable outline of the articular surface on the labial side of the surangular suggests that approximately +5 mm +are missing. On the medial surface of the main body a long, shallow groove is visible ( +Fig. 4D +). Its edges are pronounced and mostly parallel in its posterior part, and anteriorly it becomes very wide. It is gently sinuous, with its anterior end turned gently dorsally, the middle part approximately parallel to the long axis of the bone, and the posterior end downturned. In the posterior and middle part, it is situated closer to the dorsal than to the ventral edge of the bone, whereas anteriorly its ventral edge comes closer to the ventral edge of the bone while the dorsal edge diminishes, leaving the groove open dorsally. An additional short ridge is present above the middle part of the groove, close to the half of the angular. These structures probably form the area of connection with the surangular. + + +A large, clearly demarcated, depressed and mostly flat area for the suture with the dentary is visible on the lateral surface in the anterior part of the angular ( +Fig. 4A, B +). Its posterior end is located around the mid-length of the angular, its dorsal edge quickly merges with the dorsal edge of the angular, and its ventral edge is gently bowed, concave dorsally, and reaches the ventral edge of the angular close to its anterior tip. + + + +FIG +. 8. — + +Woznikella triradiata + +n. gen., n. sp. +, cervical vertebrae: +A -F +, ZPAL V.34/1/27:left part of the neural arch of the atlas in anterior ( +A +), lateral ( +B +), posterior ( +C +), medial ( +D +), dorsal ( +E +), and ventral ( +F +) view; +G -J +, ZPAL V. 34/1/30: cervical neural arch in anterior ( +G +), lateral left ( +H +), posterior ( +I +), and dorsal ( +J +) view; +K -M +, ZPAL V. 34/1/66: cervical vertebral centrum in anterior ( +K +), lateral right ( +L +), and dorsal ( +M +) view; +N -P +, ZPAL V. 34/1/77: posterior cervical vertebral centrum in anterior ( +N +), lateral right ( +O +), and dorsal ( +P +) view. Scale bar: 5 cm. + + + + +Surangular + + + +The right surangular is fused with the articular and prearticular ( +ZPAL +V +. 34/1/4; +Fig. 6 +). As preserved, its dorsal edge forms a triangular coronoid eminence. The boundary of that structure is damaged, but what appears to be remnants of its natural edge suggest that the general shape is not be significantly altered. The surangular is similar to the homologous bone of, e.g., + +Angonisaurus cruickshanki + +and + +Lisowicia bojani + +(see +Cox& Li 1983 +; +Sulej & Niedźwiedzki 2019 +). The surangular has a completely flat lingual surface. A distinct ridge on the labial surface begins in front of the coronoid eminence and posteriorly it continues onto the labial surface of the articular. A narrow ridge and a groove beneath it are close to the ventral edge of the labial side, but the anteroventral part of the bone is broken. This structure was probably the area of contact with the angular. + + + +Prearticular + + + +The right prearticular is fused with the articular and surangular ( +ZPAL +V +. 34/1/4; +Fig. 6 +). It is a thin blade of roughly rectangular shape. Its dorsal edge is curved, so the highest point is in the middle of the bone, in front of the coronoid eminence of the surangular. + + + +Articular + + + +The right articular is fused with the surangular and prearticular ( +ZPAL +V +. 34/1/4; +Fig. 6 +) and is broadened posterodorsally into a triple-edged condyle. The central ridge is the longest and highest, the inner ridge is the shortest. The retroarticular (digastric) process is very short, as in, e.g., + +Ischigulastia +jenseni + +, most + +Shaanbeikannemeyeria xilougouensis + +, and + +Wadiasaurus indicus + +, but it is longer than in + +Vinceria andina + +(see +Cox 1965 +; +Bandyopadhyay 1988 +; +Domnanovich & Marsicano 2012 +; +Liu 2022 +). Note, however, that this character shows some variability, even bilaterally within a single individual ( +Camp 1956 +). The fourth, labialmost ridge, which is located outside of the condyle, begins on the surangular like in + +Stachleckeria +potens + +and ends on the retroarticular (digastric) process like in + +Kannemeyeria simocephalus + +(see +Camp 1956 +). + + + +Vertebral column + + + +All centra and neural arches are preserved separately ( +Figs 8 +; +9 +A-M), with the exception of +ZPAL +V +. 34/1/12, a posterior dorsal vertebra ( +Fig. 8 +N-Q). Although the elements were mostly found scattered, their morphology and documented location within the association allow the establishment of their approximate relative positions within the sequence. + + + +Neural arch of the atlas + + + +Both halves of the neural arch of the atlas are preserved. The left part of the atlas neural arch is complete ( +ZPAL +V +. 34/1/27; +Fig. 8 +A-F). In lateral view, it is composed of an almost flat lateral process and an elongated posterodorsal process. The posterodorsal process is slightly asymmetric and located entirely to the left of the midline of the neural canal, most likely acting functionally as a postzygapophysis. The postzygapophyseal surface is almost oval with nearly straight medial edge. Above the postzygapophysis, a distinct ridge is present on the dorsomedial side of the process. At the base of the posterodorsal process, a large prezygapophysis is very distinct with a deep notch in its posterior margin. On the anterior portion (main body) there is a flat facet for the exoccipital and a concave, rounded facet for the centrum ventromedially. The lateral process is directed predominantly posterodorsally, its long axis nearly parallel to the posterodorsal process. It is very high and its lateral surface forms a large, oval, flat area, with a gentle but clear, laterally directed lip along its dorsal edge. The overall morphology is nearly identical to that of the neural arch of the atlas of + +Kannemeyeria simocephalus + +(see +Pearson 1924b +; +Govender 2005 +; + +Govender +et al. +2008 + +) but more elongated posterodorsally than in + +Wadiasarus indicus + +(see +Bandyopadhyay 1988 +). + + +The right part of the atlas neural arch ( +ZPAL +V +. 34/1/69) is preserved in the same bone accumulation but has a slightly different state of preservation and shape due to deformation. It is broken into at least three pieces, slightly distorted, and the surface of the bone is reddish and in a bad condition. Both processes, lateral and posterodorsal, are eroded. The lateral process is misshapen and only the base of the dorsal process is preserved. Both modes of preservation are congruent with the state of the rest of the skeleton (see the Material section). + + + +Cervical vertebrae + + + +Five cervical vertebral centra ( +ZPAL +V +. 34/1/66, +ZPAL +V +. 34/1/69, +ZPAL +V +. 34/1/70, +ZPAL +V +. 34/1/76, and +ZPAL +V +. 34/1/77; +Fig. 8 +K-P) and three cervical neural arches ( +ZPAL +V +. 34/1/14, and +ZPAL +V +. 34/1/30; +Fig. 8 +G-J) are preserved and are mostly complete ( +ZPAL +V +. 34/1/14 has its neural process deformed and broken, +ZPAL +V +. 34/1/70 is incomplete laterally on its left side, +ZPAL +V +. 34/1/76 has its posteroventral part broken off). Based on their morphology, the centra can be arranged in a following sequence (anterior to posterior): +ZPAL +V +. 34/1/66, +ZPAL +V +. 34/1/70, +ZPAL +V +. 34/1/69, +ZPAL +V +. 34/1/77, +ZPAL +V +. 34/1/76. The neural arches cannot be attributed to any particular centrum, but +ZPAL +V +. 34/1/14 is larger and has a longer neural process than +ZPAL +V +. 34/1/30, indicating that it is more posterior. + + +The centra are wide and rounded, subpentagonal in their anteroposterior aspect. Anterior and posterior surfaces are concave (weakly amphicoelous) with pronounced notochordal pits in their centers. The anterior centra are anteroposteriorly short, but gradually increase in length posteriorly, so +ZPAL +V +. 34/1/76 is about double the length of +ZPAL +V +. 34/1/66. The two anteriormost centra ( +ZPAL +V +. 34/1/66 and +ZPAL +V +. 34/1/70) have small, knob-like parapophyses located around the mid-height of the centrum ( +Fig. 8 +K-M), but the parapophyses of the more posterior centra increase in size and extend further dorsally, in +ZPAL +V +. 34/1/77 and +ZPAL +V +. 34/1/76 reaching around half of the centrum’s length and nearly reaching the articular surfaces for the neural arch pedicles ( +Fig. 8 +N-P; compare with +Camp & Welles 1956 +; +Sun 1963 +; +Bandyopadhyay 1988 +; +Surkov 1998a +; +Vega-Dias & Schultz 2004 +). The dorsal surfaces are wide, and the surfaces for articulation with the neural arches change from wider than long in the anterior cervical vertebral centra to almost rectangular in the posterior, and gently slope ventrolaterally. The centra show little to no keeling ventrally. Ventrally and laterally, particularly above the parapophyses, the cortices of the three anteriormost preserved centra are perforated by vascular canals. + + +The neural arch +ZPAL +V +. 34/1/30 has a very low and anteroposteriorly short neural process, with an additional articulation surface on its anterior edge ( +Fig. 8 +G-J). The apex of the neural process is subtriangular in cross-section, with sharpened anterior and posterolateral edges, and the posterior surface is concave. In anteroposterior aspect, the process is slightly expanded laterally near the apex ( +Fig. 8G, I +). In lateral view, the anterior edge of the process is gently concave and slanted posterodorsally, with the posterior edge subvertical ( +Fig. 8H +). There is a narrow, dorsoventrally aligned rugose band on the posterior surface, likely an attachment point for muscles or ligaments ( +Fig. 8I +). The neural process is much higher, although plastically deformed and dorsally incomplete, in +ZPAL +V +. 34/1/14. It is, furthermore, shifted posteriorly, nearly completely beyond the posterior limit of the transverse processes. The zygapophyses are nearly horizontal. The prezygapophyses are oval, and the postzygapophyses are ovoid (long axes directed posterolaterally) and concave ventrally.In lateral view, the dorsal (non-articular) surfaces of the postzygapophyses of +ZPAL +V +. 34/1/30 are continuous with and parallel to the dorsal (articular) surfaces of the prezygapophyses, and they gently slope anteroventrally. A distinctive vertical ridge is present above each postzygapophysis. In +ZPAL +V +. 34/1/14, the articular surfaces of both the pre- and postzygapophyses are roughly level with each other, but aligned more ventromedially, and the postzygapophyses are more elongate than in +ZPAL +V +. 34/1/30, with their long axes directed more posteriorly. In both specimens, the prezygapophyses and postzygapophyses are separated mesially along their entire length (the prezygapophyses wider than the postzygapophyses), but in +ZPAL +V +. 34/1/14 the latter are closer to each other than in +ZPAL +V +. 34/1/30. The diapophyses are present beneath the prezygapophyses and are relatively small dorsoventrally, but the transverse processes extend horizontally just above the neurocentral suture, with their ventrolateral surfaces convex (compare with +Young 1937 +; +Sun 1963 +; +Cheng 1980 +; +Bandyopadhyay 1988 +; +Vega-Dias & Schultz 2004 +; +Liu 2015 +). + + + +FIG +. 9. — + +Woznikella triradiata + +n. gen., n. sp. +, dorsal vertebrae: +A -D +, ZPAL V. 34/1/31: anterior dorsal neural arch in anterior ( +A +), lateral right ( +B +), posterior ( +C +), and dorsal ( +D +) view; +E -F +, ZPAL V. 34/1/35:middle dorsal neural arch in anterior ( +E +) and lateral right ( +F +) view; +G -J +, ZPAL V.34/1/33: more posterior middle dorsal neural arch in anterior ( +G +), lateral left ( +H +), posterior ( +I +), and dorsal ( +J +) view; +K -M +, ZPAL V. 34/1/11: dorsal vertebral centrum in anterior ( +K +), lateral left ( +L +), and dorsal ( +M +) view; +N -Q +, ZPAL V. 34/1/12: posterior dorsal vertebra in anterior ( +N +), lateral left ( +O +), posterior ( +P +), and dorsal ( +Q +) view. Scale bar: 5 cm. + + + + +TABLE +2. — Remains historically considered to belong to post-Permian Dicynodontia or confused with them,but in fact of Permian age and/or not being dicynodonts. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Original description + +Country + +Comments +
+“ + +Colossoemys + +macrococcygeana +” Rodrigues, +1892 +/“cf. + +Dicynodon turpior + +” +von Huene, 1935 +unnumbered specimen ( +von Huene 1944 +) + +Peru ( +Rodrigues 1892 +; von Huene 1944; +Camp & Welles 1956 +) + +A bone fragment described as an ilium of a Miocene chelid turtle (one of several syntypes described and figured by +Rodrigues 1892 +; see + +Ferreira +et al. +2006 + +for more information about the history of the syntype material), later reinterpreted as a partial humerus of a Late Triassic dicynodont ( +von Huene 1944 +; +Camp & Welles 1956 +), now lost; probably part of a xenarthran humerus ( +Price 1956 +; Paula-Couto 1960; + +de Lapparent de Broin +et al. +1993 + +; + +Ferreira +et al. +2006 + +) +
+ +Daptocephalus leoniceps + +( +Owen, 1876 +) AMNHFARB 5598, RC 96, SAM-PK-7849 ( +Broom 1921 +, +1948 +, Haughton 1924, +von Huene 1925 +, +Haughton & Brink 1954 +, +Cruickshank 1967 +) + +South Africa ( +Owen 1876 +; Broom 1921, 1948; +Haughton 1924 +; +von Huene 1925 +; Haughton & Brink 1954; +Cruickshank 1967 +; Kitching 1977; + +Kammerer +et al. +2011 + +) + +Three skulls originally described as “ + +Dicynodon osborni +” Broom + +, 1921, “ + +Dicynodon leontocephalus +” +Broom, 1948 + +, and “ + +Dicynodon + +watsoni +” +Broom, 1921 +(all subsequently synonymized with + +Daptocephalus leoniceps + +; see + +Kammerer +et al. +2011 + +), and originally thought to originate from the Triassic “ + +Lystrosaurus + +AZ” ( +Broom 1921 +, +1948 +; +Haughton 1924 +; +von Huene 1925 +; +Haughton & Brink 1954 +); independently, +Cruickshank (1967) +, citing personal communication with Crompton, noted the presence of + +Daptocephalus leoniceps + +in the “ + +Lystrosaurus +AZ + +”; most likely, however, all occurrences of + +Daptocephalus leoniceps + +are Permian and the confusion arose due to the presence of + +Lystrosaurus +spp. + +in the latest Permian + +Lystrosaurus +maccaigi-Moschorhinus + +SZ of the + +Daptocephalus +AZ + +( +Kitching 1977 +; + +Kammerer +et al. +2011 + +; +Viglietti 2020 +) +
+“ +Dicynodon incisivum +” +Repelin, 1923 +unnumbered specimen + +Laos ( +Repelin 1923 +; Yuan & Young 1934b; +von Huene 1935 +; Piveteau 1938; +Colbert 1982 +; +Brink 1988 +; +King 1988 +; +Battail 2009 +; +FrÖbisch 2009 +; + +Kammerer +et al. +2011 + +) + + + +Nomen +dubium + +, specimen lost; originally believed to be Triassic; + +probably Permian ( +Yuan & Young 1934b +; +Colbert 1982 +; Battail 2009; +FrÖbisch 2009 +; + +Kammerer +et al. +2011 + +) +
+“ +Dicynodon ingens +” +Broom, 1908b +unnumbered specimens + +South Africa ( +Broom 1908b +, +1909 +; +Haughton 1924 +; Kammerer +et al. +2011) + + + +Nomen +dubium + +, several cranial and postcranial specimens, now lost; + + +noted by +Broom (1909) +as possibly originating from the Triassic + +“ + +Lystrosaurus +AZ + +”; likely Permian ( +Haughton 1924 +; + +Kammerer +et al. +2011 + +) +
+“ +Dicynodon rosmarus +” +Cope, 1870b +unnumbered specimens + +United States ( +Cope 1870a +, b, c; +von Huene 1926b +, +1935 +; Kammerer +et al. +2013) + + + +Nomen +dubium + +, two teeth from the Upper Triassic of the New + +Oxford Formation described as dicynodont tusks, now lost; probably archosaur teeth, possibly phytosaur ( +von Huene 1926b +; + +Kammerer +et al. +2013 + +) +
+Dicynodontia indet. QMF990 (former QM F15.990) ( +Thulborn & Turner 2003 +) + +Australia ( +Thulborn & Turner 2003 +; +Knutsen & Oerlemans 2020 +) + +Bone fragments, one of which was described as a dicynodont maxilla from the Lower Cretaceous of the Allaru Formation; probably a post-Cretaceous diprotodontid mammal ( +Knutsen & Oerlemans 2020 +) +
+Dicynodontia indet. YPM VPPU 020750 ( +Baird & Olsen 1983 +) + +Canada ( +Baird & Olsen 1983 +; Olsen +et al. +1989; +Lucas & Hunt 1993a +; +Nesbitt & Angielczyk 2002 +; Sues & Olsen 2015) + +Waethered skull fragment from the Carnian Wolfville Formation, initially mentioned as an indeterminate dicynodont; reinterpreded as a paracrocodylomorph pseudosuchian by +Sues & Olsen (2015) +
+“aff. + +Dinodontosaurus +sp. + +” SMNS 56891 ( +Lucas & Wild 1995 +) + +Germany ( +Lucas & Wild 1995 +; + +Maisch +et al. +2009 + +) + +Humerus from the Middle Triassic of the Muschelkalk; probably a temnospondyl ( + +Maisch +et al. +2009 + +) +
+Geikia elginensis +Newton, 1893 +GSM 90998-91015 + +Scotland ( +Newton 1893 +; von Huene 1913; + +Kammerer +et al. +2011 + +) + +A series of sandstone fragments with natural molds of a skull, mandible, and limb fragments from the Cuttie’s Hillock Sandstone +in Scotland, originally considered to of Triassic age ( +Judd 1885 +; +Newton 1893 +); subsequent biostratigraphic correlations suggest a Permian age for that assemblage (e.g., +von Huene 1913 +; +Rowe 1980 +; + +Kammerer +et al. +2011 + +) +
+Gordonia traquairi +Newton, 1893 +GSE 11703 and several other specimens + +Scotland ( +Judd 1885 +; +Newton 1893 +; +von Huene 1913 +; Kammerer +et al. +2011) + + +Several specimens described as + +Gordonia traquairi +Newton, 1893 + +, + +Gordonia duffiana +Newton, 1893 + +, + +Gordonia huxleyana +Newton, + + +1893, and + +Gordonia juddiana +Newton, 1893 + +, which were later synonymized ( +King 1988 +; + +Kammerer +et al. +2011 + +), all coming from the Cuttie’s Hillock Sandstone in Scotland, originally considered to be of Triassic age ( +Judd 1885 +; +Newton 1893 +); subsequent biostratigraphic correlations suggest a Permian age for that assemblage (e.g., +von Huene 1913 +; +Rowe 1980 +; + +Kammerer +et al. +2011 + +) +
+ +Herpetochirus brachycnemus + +Seeley, 1985 NHMUK R2588 + +South Africa ( +Seeley 1895 +; +von Huene 1925 +; +Kammerer 2009 +) + +Fragmentary postcranial skeleton from South Africa, noted by +von Huene (1925) +as a dicynodont from the Triassic “ + +Lystrosaurus +AZ + +”; probably Permian therocephalian ( +Seeley 1895 +; +Kammerer 2009 +) +
+ + + +Table 2 +. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + +
+Original description + +Country + +Comments +
+“ + +Ischigualastia boreni + +” +Edler, 2000 +TTU P-9427 + +United States ( +Edler 2000 +; Nesbitt & Bone fragment from the Norian of the Cooper Canyon Formation, +Angielczyk 2002 +; +Martz 2008 +) interpreted as a preorbital part of a stahleckeriid skull; its dicynodont affinity was subsequently questioned ( +Nesbitt & Angielczyk 2002 +); probably posterior part of a pseudosuchian mandible ( +Martz 2008 +) +
+“ + +Lystrosaurus +sp. + +” MCN PV1872 and MCN PV1873 ( +Schwanke & Kellner 1999 +, +Langer & Lavina 2000 +) + +Brazil ( +Schwanke & Kellner 1999 +; +Langer & Lavina 2000 +; Dias-da-Silva +et al. +2017) + +Two specimens from the Lower Triassic of the Sanga do Cabral Formation, described as lystrosaurid stapes; probably procolophonid sacral ribs ( + +Dias-da-Silva +et al. +2017 + +) +
+ +Theromus leptonotus + +Seeley, 1985 NHMUK R2585, NHMUK R2587 + +South Africa ( +Seeley 1895 +; Watson 1911; +von Huene 1925 +; Kammerer 2009) + +Fragmentary postcranial skeleton from South Africa, noted by +von Huene (1925) +as a dicynodont from the Triassic “ + +Lystrosaurus + +AZ”; in fact either a therocephalian ( +Seeley 1895 +; +Watson 1911 +) or non-dicynodont anomodont ( +Kammerer 2009 +), probably + +Permian ( +Seeley 1895 +; +Watson 1911 +) + +
+ + + +Dorsal vertebrae + + + +There are six isolated neural arches ( +ZPAL +V +. 34/1/13, +ZPAL +V +. 34/1/31, +ZPAL +V +. 34/1/33, +ZPAL +V +. 34/1/34, +ZPAL +V +. 34/1/35, and +ZPAL +V +. 34/1/36; +Fig. 9 +A-J), four isolated vertebral centra ( +ZPAL +V +. 34/1/11, +ZPAL +V +. 34/1/67, +ZPAL +V +. 34/1/68, and +ZPAL +V +. 34/1/71; +Fig. 9 +K-M), and a single mostly complete vertebra ( +ZPAL +V +. 34/1/12; +Fig. 8 +N-Q) that can be classified as belonging to the dorsal series. Based on recovery documentation, out of the isolated elements, the centrum +ZPAL +V +. 34/1/71 and the neural arch +ZPAL +V +. 34/1/36 were found in near articulation, and +ZPAL +V +. 34/1/34 was found right behind them. The centrum +ZPAL +V +. 34/1/67 was found below the neural arch +ZPAL +V +. 34/1/31. The shapes and sizes of the neurocentral articulation facets as well as the state of preservation suggest that the centra +ZPAL +V +. 34/1/68 and +ZPAL +V +. 34/1/11 ( +Fig. 9 +K-M) may belong to the neural arches +ZPAL +V +. 34/1/34 ( +Fig. 9 +K-M) and +ZPAL +V +. 34/1/33 ( +Fig. 9 +G-J), respectively, and that they come from neighboring segments. Based on those data and gradual changes of morphology, the elements may be hypothesized to form the following relative sequence (anterior to posterior): +ZPAL +V +. 34/1/31, +ZPAL +V +. 34/1/67, +ZPAL +V +. 34/1/35, +ZPAL +V +. 34/1/71 + +ZPAL +V +. 34/1/36, +ZPAL +V +. 34/1/68 + +ZPAL +V +. 34/1/34, +ZPAL +V +. 34/1/11 + +ZPAL +V +. 34/1/33, +ZPAL +V +. 34/1/13, +ZPAL +V +. 34/1/12. However, this sequence should be treated with caution, particularly when it comes to the vertebral centra, due to the lesser number of morphological details observable on them. + + +The anterior part of the dorsal vertebral column is represented by a single neural arch, +ZPAL +V +. 34/1/31 ( +Fig. 9 +A-D). Its anterior position within the dorsal section of the vertebral column is indicated by its extensive, wide synapophyses spanning from the tips of the transverse processes, ventromedially towards the neurocentral suture, and apparently continuing onto the centrum (compare with +Pearson 1924b +; +Young 1937 +; +Sun 1963 +; +Cox 1965 +; +Cruickshank 1967 +; +Bandyopadhyay 1988 +). The transverse processes are wing-like, massive, and similar to those of the cervical neural arches, but gently raised laterally. Their ventrolateral surfaces are nearly straight, and their edges are angular. The dorsal edges of the transverse processes face anterodorsally and form distinct lateromedial ridges anteriorly and posterodorsally. The synapophyses are wider dorsally than ventrally and skewed posterodorsally in lateral view. In contrast to the cervical neural arches, the prezygapophyses of +ZPAL +V +. 34/1/31 have subrectangular outlines and are aligned dorsolaterally rather than subhorizontally; the angle between their anterior edges is approximately 90° in the medial part, but their lateral parts gently turn dorsally. The postzygapophyses are ovoid, shorter than in the cervical neural arch +ZPAL +V +. 34/1/14, and set at a more acute angle than the prezygapophyses (about 60°). Contralaterally, both the pre- and postzygapophyses are separated only by a narrow notch. The neural process is high and anteroposteriorly long with a rounded and gently anteroposteriorly expanded dorsal end. It is directed gently posteriorly. The neural process apex is fusiform in cross-section and only slightly expanded laterally. At the base of the neural process there is a distinct vertical ridge anteriorly, reaching the notch between the prezygapophyses, and a vertically expanded, rhomboid depression posteriorly, limited by the postzygapophyses ventrolaterally and by low, rounded ridges dorsolaterally. The anterior edge of the base of the neural process reaches anteriorly past the level of the anterior edge of the synapophyses. + + +ZPAL +V +. 34/1/35 represents a slightly more posterior neural arch ( +Fig. 9E, F +). In contrast to +ZPAL +V +. 34/1/31, the dorsolaterally directed transverse process in this specimen is free from the synapophysis and is plate-like. Only a low ridge continues from the dorsal end of the synapophysis onto the posteroventral surface of the transverse process. The dorsal extent of the synapophyses is below the dorsolateral edges of the prezygapophyses, which is lower than in +ZPAL +V +. 34/1/31 but higher than in succeeding vertebrae. It is well defined in anteroposterior aspect and clearly separated from the transverse process, as in the remaining, more posterior neural arches. The dorsal part of the synapophysis is rounded and anteroposteriorly wider than the ventral part. The prezygapophyses are set at a more acute angle than in +ZPAL +V +. 34/1/31 (about 60°), the postzygapophyses are set at a greater angle (about 80°) and the dorsal process is directed more posterodorsally. The morphology is reminiscent of that figured and described by +Pearson (1924b) +, +Govender (2005) +, and + +Govender +et al. +(2008) + +for the mid-dorsal vertebrae of + +Kannemeyeria simocephalus + +. + + +The succeeding neural arches show gradual changes in morphology: 1) the synapophyses recede farther ventrally compared to +ZPAL +V +. 34/1/35 and develop a conspicuous dorsal tip, thus attaining a subtriangular shape with gently bowed anterodorsal and posterodorsal edges; 2) the prezygapophyses and postzygapophyses are ovoid and become more horizontal; 3) the dorsal processes become inclined even more posterodorsally and their bases shift more posteriorly relative to the anterior extent of the synapophyses; 4) the apices of the dorsal processes expand more laterally, particularly in the anterior part, so they become teardrop-shaped in cross-section, and develop subtle medial grooves anteriorly; and 5) the depressions in the posterior bases of the neural processes become deeper and more conspicuous. The posteriormost preserved vertebra, +ZPAL +V +. 34/1/12, presents the most extreme morphology ( +Fig. 9 +N-Q). The synapophyses in this specimen are particularly low but anteroposteriorly wide, the prezygapophyses are short but the postzygapophyseal part is extended posteriorly due to the posterior shift of the neural process. The outline of the articular surface of the postzygapophyses, nonetheless, takes up only the posterior half of that area. The transverse processes are more horizontal, both in the lateral and anteroposterior aspect. The posterodorsal alignment of the neural process is particularly pronounced. In lateral view, the process presents subtle sigmoidal curvature, with the midsection directed more posteriorly than the base and the apex. The base of the process presents a strong ridge, which unlike in +ZPAL +V +. 34/1/31 does not extend to the notch between the prezygapophyses, but ends at the level of the base of the transverse processes.The apex of the neural process is expanded also in the middle part, thus presenting a pentagonal cross-section. The groove in its anterior face is subtle, but more conspicuous than in the preceding neural arches. + + +The dorsal vertebral centra are weakly amphicoelous with distinct notochordal pits in the anterior and posterior facets, and are ovoid in anterioposterior aspect. They are comparable in length to the posterior cervical vertebral centra but slightly narrower dorsally, so the facets for the neural arches are longer than wide. Some of them (particularly +ZPAL +V +. 34/1/67 and +ZPAL +V +. 34/1/12) bear a rounded ventral keel. The central parts of synapophyses are located in the anterodorsal corners of the centra. As in the case of their neural parts, they become smaller posteriorly. Most of them present convex posteroventral edges, similar to those in the posterior cervical vertebrae, with the exception of +ZPAL +V +. 34/1/12, in which this edge is straight and the synapophysis has a rhomboid outline. + + +The observed morphological anteroposterior progression is consistent with the changes observed previously in other +Kannemeyeriiformes +(compare with +Pearson 1924b +; +Young 1937 +; +Sun 1963 +; +Cruickshank 1967 +; +Bandyopadhyay 1988 +). It is furthermore consistent with the shape and size of the ribs and their articular regions (see below). According to + +Vega-Dias +et al. +(2004) + +the vertebrae of Triassic dicynodonts have no diagnostic value at the family or generic level. However, it must be noted that the relative scarcity of published data on dicynodont vertebrae (in many cases, only selected vertebrae were figured and their precise location within the column was not determined), has made meaningful comparisons difficult, and variation in these elements may be more extensive than currently recognized. In contrast to + +Jachaleria candelariensis + +, the dorsal vertebral centra of + +Woznikella triradiata + +n. gen., n. sp. +are significantly longer and the transverse processes are more pronounced ( +Araújo & Gonzaga 1980 +; +Vega-Dias & Schultz 2004 +; + +Martinelli +et al. +2020 + +). They are also narrower relative to height, particularly in the ventral region, than in, e.g., + +Dinodontosaurus tener + +, + +Ischigualastia jenseni + +(at least for part of the dorsal vertebral column), + +Jachaleria colorata +Bonaparte, 1970 + +, + +Kannemeyeria simocephalus + +, + +Lisowicia bojani + +, + +Pentasaurus goggai + +, + +Rhinodicynodon gracile + +, + +Sangusaurus parringtonii + +, + +Sinokannemeyeria pearsoni + +, and + +Stahleckeria potens + +(see +von Huene 1935 +; +Young 1937 +; +Romer & Price 1944 +; +Cox 1965 +; +Araújo & Gonzaga 1980 +; +Surkov 1998a +; +Vega-Dias & Schultz 2004 +; +Morato 2006 +; + +Angielczyk +et al. +2017 + +; +Kammerer 2018 +; +Sulej & Niedźwiedzki 2019 +; + +Martinelli +et al. +2020 + +), being more in line with, e.g., + +Parakannemeyeria shenmuensis + +, + +Placerias hesternus + +, + +Shansiodon wangi + +, + +Sinokannemeyeria yingchiaoensis + +, + +Wadiasaurus indicus + +, or + +Zambiasaurus submersus + +(e.g., +Camp & Welles 1956 +; +Yeh 1959 +; +Sun 1963 +; +Cox 1969 +; +Cheng 1980 +; +Bandyopadhyay 1988 +). The shape of the centra, however, aside from taphonomic factors, may potentially be impacted by ontogeny (e.g., +von Huene 1935 +). If the assignment of the neural arches to the vertebral centra is correct, the posterior ends of their pedicels could project posteriorly in the mid-dorsal vertebrae, forming protuberances consistent with the morphology seen in at least some other +Kannemeyeriiformes +( +von Huene 1935 +; +Araújo & Gonzaga 1980 +; +Bandyopadhyay 1988 +; +Vega-Dias & Schultz 2004 +; + +Martinelli +et al. +2020 + +). + + + +Ribs + + + +Ribs are represented by some almost complete specimens and many fragments of varying sizes. Among them several morphotypes may be distinguished, which generally agree with the morphologies presented for + +Jachaleria candelariensis + +by +Vega-Dias & Schultz (2004) +, + +Kannemeyeria simocephalus + +by +Pearson (1924b) +, and + +Sinokannemeyeria yingchiaoensis + +by +Sun (1963) +, and correspond well with the articulation facets observed on the vertebrae (see above). + + + +FIG +. 10. — + +Woznikella triradiata + +n. gen., n. sp. +, ribs: +A +, +B +, ZPAL V. 34/1/82 in anterior ( +A +) and posterior ( +B +) view; +C +, +D +, ZPAL V. 39/1/20 in anterior ( +C +) and posterior ( +D +) view; +E +, +F +, ZPAL V. 34/1/208 in anterior ( +E +) and posterior ( +F +) view; +G +, +H +, ZPAL V. 34/1/17 in anterior ( +G +) and posterior ( +H +) view; +I +, +J +, ZPAL V. 34/1/26 in?anterior ( +I +) and?posterior ( +J +) view; +K -Q +, closeups of the articular surfaces: +K +, ZPAL V. 34/1/16; +L +, ZPAL V. 34/1/82; +M +, ZPAL V. 34 /1/15; +N +, ZPAL V. 34/1/20; +O +, ZPAL V. 34/1/208; +P +, ZPAL V. 34/1/17; +Q +, ZPAL V. 34/1/26. Scale bars: A-J, 5 cm; K-N, O-Q, 1 cm. + + + +ZPAL +V +. 34/1/26 is a very short ( +14 cm +along the parietal edge), anteroposteriorly flattened rib with a distinct curvature and conspicuous decrease of the visceroparietal width along its length ( +Fig. 10I, J, Q +). The proximal part in anteroposterior aspect is gently flared, slightly sloped anteromedially in proximal view, and exceptionally flat (29 × +8 mm +). Unlike the remaining ribs, it does not form a defined articular surface and its edge is uneven, so it is likely that this rib formed a sutural connection with the vertebra rather than a movable articulation. As preserved, the specimen terminates in a blunt 4 × +6 mm +large tip. Possibly this is a break, but it seems unlikely that the rib continued much further in life. Given its different morphology, this specimen is tentatively identified as a cervical rib.Ribs with anteroposteriorly narrow and visceroparietally wide shafts are common and attain the largest sizes, possibly coming from the widest, anterior section of the ribcage. The best-preserved specimen in this category is +ZPAL +V +. 34/1/82 ( +Fig. 10A, B, L +). It is a very long rib with a complete head and tubercle, but with a broken distal end. The rib head is set on a short neck and is relatively small compared to the expanded tubercle, the visceroparietal size of which nearly equals the visceroparietal width of the shaft. This amounts to the proximalmost part of the rib being almost two times the size of the shaft visceroparietal width. The articular surface for the diapophysis forms an inverted pear-shape in proximal view and is much larger than the oval surface for the parapophysis. Both facets are separated by a constriction giving the proximal part of the rib an inverted figure 8 shape in articular view. The posterior edges of the articular surfaces protrude in a ridge-like fashion, as does the dorsal edge of the facet for the diapophysis. The specimen is broken in several places and its distal part is crushed and distorted, but it appears to originally have a relatively straight shaft with nearly parallel edges. The specimen +ZPAL +V +. 34/1/16 presents overall the same morphology but the edges of the articular facets also form conspicuous ridges anteriorly ( +Fig. 10K +). The shaft of this specimen is gently sinuous in visceroparietal aspect, but it is not clear whether this waviness was present in vivo, or if it was acquired postmortem because of distortion. In the specimen +ZPAL +V +. 34/1/83, which represents the same morphotype, the curvature of the shaft is slightly more pronounced, likely as a result of a smaller circumference of the rib cage at its level. In +ZPAL +V +. 34/1/15, the head and tubercle merge in a single, elongated subvertical articular surface ( +Fig. 10M +). More posterior, visceroparietally wide ribs with a smaller single, vertical or nearly vertical articular surface lying in the same plane as the shaft and a curved proximal region ( +ZPAL +V +. 34/1/20, +ZPAL +V +. 34/1/21, +ZPAL +V +. 34/1/85) can reach similar visceroparietal width as the previous +type +, but were likely shorter, considering their curvature.In +ZPAL +V +. 34/1/20 ( +Fig. 10C, D, N +) and +ZPAL +V +. 34/1/85, the articular surface is sinuous in anteroposterior aspect, convex in its ventral part, and recessed in the dorsal part. In +ZPAL +V +. 34/1/20 it is oblique, directed ventromedially in anteroposterior aspect, and dorsally projected into a small process, whereas in +ZPAL +V +. 34/1/85 the dorsal part is nearly horizontal in anteroposterior aspect. In both specimens, the articular surfaces are slightly sloped anteroventrally in proximal view. The dorsal and ventral parts are continuous but in proximal view there is a very subtle constriction in the middle of the articular surface, giving it a peanut-shaped outline and suggesting that this area is formed by merging of the head and tubercle. In +ZPAL +V +. 34/1/21, the articular surface in proximal view is vertical and convex, inverted comma-shaped, with the thinner end directed posterodorsally and the anterodorsal edge convex. The specimen is visceroparietally narrower than +ZPAL +V +. 34/1/20 and +ZPAL +V +. 34/1/85. In all specimens of this +type +, the shaft is anteroposteriorly flattened in the proximal part, the anterior and posterior surfaces are gently concave just distal to the curve, and more distally the shaft becomes oval in cross-section and visceroparietally narrower. + + +The distal (ventral) ends of the ribs belonging to the previous two morphotypes are completely preserved in +ZPAL +V +. 34/1/18 and +ZPAL +V +. 34/1/20. Based on these specimens, it may be established that the oval cross-section, which is present around the midlength of the ribs, becomes flattened again and increases in visceroparietal width distally. The ribs terminate with swollen and rounded ends, ovoid in distal view. + + +Strongly curved ribs with narrow and long shafts likely come from the posteriormost part of the rib cage. This kind of rib is represented by the specimens +ZPAL +V +. 34/1/17 ( +Fig. 10G, H, P +) and +ZPAL +V +. 34/1/23. The articular surface is only preserved in +ZPAL +V +. 34/1/17. It is horizontal, single, and elongated (the anteroposterior width about twice that of the shaft) with a straight ventral edge and a convex dorsal edge. In dorsovental aspect, the anteriormost point of the articular surface is protruding. The neck expands dorsally towards a smooth dorsal curve, which constitutes the visceroparietally widest part of the shaft. The shaft just distal to the curve becomes subtriangular in cross-section, with a rounded visceral and posterior edge, an angular anterodorsal edge, and a concave posterovisceral surface, forming along the dorsolateral part of the rib a small, posteriorly directed lappet of bone. More distally, the rib becomes oval in cross-section, with the longer axis directed visceroparietally. + + + +Scapula + + + +The nearly complete, right scapula ( +ZPAL +V +. 34/1/7; +Fig. 11 +A-D) only lacks some minor fragments of its ventral and posterior edge and the posterodorsal tip of the blade. It is roughly hourglass-shaped in lateromedial aspect, with the anterior edge more concave than the posterior, and the minimal breadth just below the midlength, above the acromion ( +Fig. 11A, B +). It is gently bowed and twisted in anteroposterior view, with its dorsal and ventral ends turned medially and deflected anteromedially ( +Fig. 11C, D +). The scapular spine is damaged but seems to be only weakly developed and the prespinal surface is flat ( +Fig. 11C +) – in this aspect it differs from dicynodonts such as + +Eubrachiosaurus browni + +, + +Placerias hesternus + +, + +Stahleckeria potens + +, the indeterminate stahleckeriines (MCZ 3459 and CRILAR-Pv 82) from the Chañares Formation, + +Rhinodicynodon gracile + +, + +Tetragonias njalilus + +, or even + +Zambiasaurus submersus + +, which have a more pronounced spine ( +von Huene 1935 +; +Romer & Price 1944 +; +Camp & Welles 1956 +; +Cruickshank 1967 +; +Cox 1968 +, +1969 +; +Surkov 1998a +; + +Kammerer +et al. +2013 + +; + +Escobar +et al. +2021 + +). The acromion is conspicuous, mediolaterally flattened, hooked medially, protrudes well past the anterior margin of the scapula, and has a blunt, thickened tip forming medially a small tubercle ( +Fig. 11B +) similar to that of + +Stahleckeria potens + +and + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”) figured by +von Huene (1935) +. The presence of a well-developed acromion differentiates + +Woznikella triradiata + +n. gen., n. sp. +from cf. + +Dolichuranus primaevus + +and + +Ischigualastia jenseni + +, which lack this process ( +Cox 1965 +; +Govender & Yates 2009 +; + +Escobar +et al. +2021 + +). Nonetheless, the acromion is not as large, relative to the rest of the scapula, as in, e.g., + +Acratophorus argentinensis + +, + +Dinodontosaurus tener + +, + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”), + +Lystrosaurus +spp. + +, + +Rhinodicynodon gracile + +, or + +Shansiodon wangi + +(see +Broom 1908a +; +von Huene 1935 +; +Young 1935 +; +Yeh 1959 +; +Cox 1965 +; +Bonaparte 1966a +; +DeFauw 1986 +; +Surkov 1998a +; +Morato 2006 +; +Ray 2006 +; + +Kammerer +et al. +2013 + +; + +Escobar +et al. +2021 + +). Unlike in most Triassic dicynodonts (e.g., +Pearson 1924b +; +von Huene 1935 +; +Broom 1937 +; +Romer & Price 1944 +; +Yeh 1959 +; +Sun 1960 +, +1963 +; +Bonaparte 1966a +; +Cruickshank 1967 +; +Colbert 1974 +; +Araújo & Gonzaga 1980 +; +Cox & Li 1983 +; +Bandyopadhyay 1988 +; +Lucas & Harris 1996 +; +Surkov 1998a +; +Govender 2005 +; + +Surkov +et al. +2005 + +; +Ray 2006 +; + +Govender +et al. +2008 + +; + +Kammerer +et al. +2013 + +; + +Escobar +et al. +2021 + +), it is directed anterodorsally rather than anteriorly, laterally, or antero- or ventrolaterally. The acromion had been broken off during recovery or preparation, causing a wide crack laterally, but its medial and ventral surfaces indicate that the pieces were well fitted together, and that no misalignment occurred. Other than the direction of the acromion, the scapula mostly resembles the narrow and gently flared “kannemeyeriid-like” ( +sensu + +Kammerer +et al. +2013 + +; note that the morphologies are not restricted to the clades after which they are named) scapulae of, e.g., + +Kannemeyeria simocephalus + +, + +Parakannemeyeria youngi + +, + +Rhinodicynodon gracile + +, + +Sinokannemeyeria yingchiaoensis + +, + +Wadiasaurus indicus + +, the unnamed stahleckeriine (MCZ 3459) from the Chañares Formation, the morphotype B stahleckeriid of +Govender (2005) +, and possibly + +Eubrachiosaurus browni + +in its general shape and proportions ( +Watson 1917 +; +Pearson 1924b +; +Broom 1937 +; +Sun 1963 +; +Cox 1968 +; +Bandyopadhyay 1988 +; +Surkov 1998a +; +Govender 2005 +; +Ray 2006 +; + +Govender +et al. +2008 + +; + +Kammerer +et al. +2013 + +). +Govender (2005) +and + +Govender +et al. +(2008) + +noted that + +Kannemeyeria simocephalus + +had relatively wide scapular blade and that the narrow-bladed scapulae previously attributed to that species by +Pearson (1924b) +differ significantly, resembling more +Govender’s (2005) +morphotypeB stahleckeriid, the scapula of which, however, also fits better the “kannemeyeriid-like +type +” +sensu + +Kammerer +et al. +(2013) + +. +ZPAL +V +. 34/1/7 differs from the typical “shansiodontid-” and “stahleckeriid-like” ( +sensu + +Kammerer +et al. +2013 + +) scapulae of + +Acratophorus argentinensis + +, + +Dinodontosaurus tener + +, + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”), + +Ischigualastia jenseni + +, + +Jachaleria candelariensis + +, cf. + +Kannemeyeria lophorhinus + +, + +Shansiodon wangi +, +Tetragonias njalilus + +, and + +Stahleckeria potens + +, which are less gracile, have a more expanded scapular blade, and less expanded point of contact with the coracoids ( +von Huene 1935 +; +Romer & Price 1944 +; +Yeh 1959 +; +Cox 1965 +; +Bonaparte 1966a +; +Cruickshank 1967 +; +Araújo & Gonzaga 1980 +; +Lucas 2002 +; +Vega-Dias & Schultz 2004 +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +; +Govender & Yates 2009 +). The scapulae attributed to + +Lystrosaurus +spp. + +show some variability, but generally seem to be stouter as well (e.g., +Huxley 1865 +; +Broom 1908a +; +Young 1935 +; +Colbert 1974 +; +DeFauw 1986 +; +Ray 2006 +). The scapula of + +Zambiasaurus submersus + +is similar in its slenderness, but less flared at both ends ( +Cox 1969 +; +Govender 2005 +; + +Kammerer +et al. +2013 + +), whereas the scapula of + +Placerias hesternus + +is less constricted ( +Camp & Welles 1956 +; + +Kammerer +et al. +2013 + +). A lesser degree of flaring around the glenoid area also differentiates it from the scapula of + +Angonisaurus cruickshanki + +( +Cox & Li 1983 +; +Govender 2005 +). The scapula of + +Lisowicia bojani + +is less gracile as well, and has a significantly more expanded ventral part ( +Sulej & Niedźwiedzki 2019 +). The scapula of + +Woznikella triradiata + +n. gen., n. sp. +is thickest at the glenoid, the articular surface of which in the scapular part is gently concave, rhomboid with rounded corners and set at an angle of ~100° relative to the anteroventral facet for the coracoids. This articular surface is approximately half the length of the gently convex facet for the coracoids. The attachment site for the triceps muscle is present as a rugose field on the posterolateral edge of the scapula above the glenoid ( +Fig. 11A, D +). There is no trace of a +m. teres major +scar (e.g., +Vega-Dias & Schultz 2004 +), but it might have occupied the unpreserved posterior tip of the scapula. Along most of the visceral surface of the scapula, a gentle but clear striation is present but there is no tubercle at the base of the acromion like that described by +Govender (2005) +and + +Govender +et al. +(2008) + +in + +Kannemeyeria simocephalus + +or + +Tetragonias njalilus + +. +Govender (2005) +described a fossa near the same area in her morphotype B stahleckeriid, and a gentle depression seems to be also present at the anteromedially surface of +ZPAL +V +. 34/1/7, right above the acromion, but it is smaller and it is difficult to establish whether the two are homologous. + + + +FIG +. 11. — + +Woznikella triradiata + +n. gen., n. sp. +, scapulocoracoid: +A -D +, ZPAL V. 34/1/7, right scapula in lateral ( +A +), medial ( +B +), anterior ( +C +), and posterior ( +D +) view; +E +, +F +, ZPAL V. 34/1/107, right procoracoid in lateral ( +E +) and medial ( +F +) view. Scale bar: 5 cm. + + + + +FIG +. 12. — + +Woznikella triradiata + +n. gen., n. sp. +, clavicles: +A -D +, ZPAL V. 34/1/75, left clavicle in posterior ( +A +), anteroventral ( +B +), anterodorsal ( +C +), and posterodorsal ( +D +) view; +E -H +, ZPAL V. 34/1/74, right clavicle in posteroventral ( +E +), anteroventral ( +F +), anterodorsal ( +G +), and posterodorsal ( +H +) view. Scale bar: 5 cm. + + + + +FIG +. 13. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1/9,?left ulna: +A +, +B +, proximal part in dorsal (anterior) ( +A +), and ventral (posterior) ( +B +) view; +C -G +, distal part in dorsal (anterior) ( +C +), medial ( +D +), lateral ( +E +), ventral (posterior) ( +F +), and distal ( +G +) view. Scale bar: 5 cm. + + + + +Procoracoid + + + +The right procoracoid ( +ZPAL +V +. 34/1/107; +Fig. 11E, F +) is preserved almost in its entirety but with its anterior end damaged. The facet for the scapula lacks sutural characteristics and the ventral edge is rounded and porous, but it is unclear whether this state is caused by weathering or if the bone was still finished in cartilage at the time of the animal’s death. The posterior suture with the coracoid is nearly straight and closed – it is recognizable as a raised, rugose ridge on both the visceral and external surface of the specimen. The procoracoid foramen is fully enclosed by the procoracoid, although it is very close to the contact with the scapula, especially on the visceral side. As a result, its walls are directed slightly posteroventrally. The procoracoid is very gently sigmoid in anterior view, with its ventral part thin and faintly directed medially. Its external surface is otherwise nearly flat. Viscerally, however, there is a pronounced thickening forming a rounded shelf below the procoracoid foramen and in the posterodorsal part of the bone, behind the foramen and close to the glenoid. There is no indication that the procoracoid itself contributed to the articular surface of the glenoid, unlike, e.g., + +Tetragonias njalilus + +(see +Cruickshank 1967 +). + + + +Coracoid + + + +Only a minute part of the anterior edge of the right coracoid is preserved connected by a closed, nearly straight suture to the right procoracoid ( +ZPAL +V +. 34/1/107; +Fig. 11E, F +). Nothing can be said about that bone other than that the preserved part thickens dorsally, towards the glenoid area. + + + +Clavicle + + + +Both clavicles are preserved (right +ZPAL +V +. 34/1/74, left +ZPAL +V +. 34/1/75; +Fig. 12 +) but the medial ends of both are broken near the area for contact with the interclavicle. Each clavicle is twisted; the lateral part is vertical while the medial end is horizontal. The lateral part is gently bent posteriorly. The lateral end with the area of contact with the acromion process of scapula is well preserved and well developed. Its anterior surface is convex, whereas the posterior one is concave in ventral part and its dorsal part forms a massive ridge along the dorsal edge of the lateral half of the bone. The ventral edge of the lateral end forms a ridge along the posterior margin of the rest of the clavicle. Unlike in + +Sinokannemeyeria yingchiaoensis + +, the clavicles of + +Woznikella triradiata + +n. gen., n. sp. +lack finger-like processes at the scapular ends ( +Sun 1963 +). This may be taxonomic or may result from their incomplete ossification due to young ontogenetic age in the individual. + + + +Humerus + + + +The right humerus ( +ZPAL +V +. 34/1/6; +Fig. 13 +) is preserved in two parts, lacks the anterior portion of the proximal end and the midshaft, and most of the ventral surfaces of the preserved fragments are heavily damaged. The articular surface of the proximal end is slightly raised and subtriangular in dorsal aspect, but still rather inconspicuous ( +Fig. 13A, C, F +), at least in part due to taphonomic factors, but possibly also due to immature ontogenetic age of the individual – the well-preserved fragments of its surface are porous, indicative of a well-developed cartilaginous finish. As in + +Kannemeyeria simocephalus + +, its distal corner terminates in a gentle ridge ( +Govender 2005 +; + +Govender +et al. +2008 + +). Similarly, the medial process (insertion of +m. subcoracoscapularis +) is short, lacks any rounded tip, lies nearly in line with the posterior (medial) surface of the shaft, and takes form of a relatively short, raised lip, continuous along the proximal and posterior edge of the proximal humeral end, rather than a distinct projection. In that respect, it is similar to, e.g., cf. + +Dolichuranus primaevus + +, + +Lisowicia bojani + +, + +Parakannemeyeria shenmuensis + +, + +Parakannemeyeria youngi + +, + +Sinokannemeyeria yingchiaoensis + +, the morphotype B stahleckeriid of +Govender (2005) +, possibly + +Eubrachiosaurus browni + +and – to some extent – + +Angonisaurus cruickshanki + +, + +Kannemeyeria simocephalus + +, and + +Sinokannemeyeria pearsoni + +, but different from, e.g., + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”), + +Dinodontosaurus tener + +, at least some representatives of + +Lystrosaurus + +and + +Placerias hesternus + +, + +Shansiodon wangi + +, + +Stahleckeria potens + +, + +Wadiasaurus indicus + +, + +Xiyukannemeyeria brevirostris + +, and + +Zambiasaurus submersus + +(see +Williston 1904 +; +Watson 1917 +; +Pearson 1924b +; +von Huene 1935 +; +Young 1937 +; +Camp & Welles 1956 +; +Yeh 1959 +; +Sun 1963 +, 1978; +Cox 1965 +, +1969 +; +Cheng 1980 +; +Bandyopadhyay 1988 +; +Lucas & Harris 1996 +; +Lucas 2002 +; +Govender 2005 +; +Ray 2006 +; + +Dzik +et al. +2008a + +; + +Govender +et al. +2008 + +; +Govender & Yates 2009 +; + +Kammerer +et al. +2013 + +; +Sulej & Niedźwiedzki 2019 +). A similar morphology is also present in the specimen MCZ 3118 of + +Ischigualastia jenseni + +as pictured by +Cox (1965) +, but the specimen PVL 3807 has a more pronounced rounded medial process ( + +Kammerer +et al. +2013 + +). The proximal end thickens in that area, but the significant damage of the ventral surface obscures details of the morphology. The deltopectoral crest is not preserved, so its exact size and shape cannot be determined, but based on the curvature of the proximal edge of the proximal end, at least proximally it did not reach much further anteriorly (laterally) from the proximal condyle than the medial process reaches posteriorly (medially). The bicipital fossa in the preserved part seems to be relatively shallow and gently concave, but the ventral surface is significantly damaged ( +Fig. 13B, F +). The attachment of +m. latissimus dorsi +is inconspicuous, taking form of a rugose, longitudinally extended area ( +Fig. 13C +). The distal end is relatively well preserved in dorsal aspect, presenting a marked smaller entepi- and larger ectepicondyle with rugosities and porous distal surfaces indicative of a cartilaginous cap ( +Fig. 13A, B, D, E, G +). The distal end is slightly narrower than the proximal end. Its outline in this aspect is triangular and its anterior (lateral) and posterior (medial) expansion is relatively gentle and symmetrical, similar to that in + +Kannemeyeria simocephalus + +, some specimens of + +Lystrosaurus +spp. + +(except + +L. georgi + +, + +L. hedini + +, ‘ + +Lystrosaurus weidenreichi +’ +Young, 1939 + +, and some specimens from +South Africa +and +Antarctica +; see +Young 1935 +, +1939 +; +Colbert 1974 +; +DeFauw 1986 +; + +Surkov +et al. +2005 + +), and + +Zambiasaurus submersus + +, but unlike, e.g., cf. + +Acratophorus argentinensis + +, + +Dolichuranus primaevus + +, + +Dinodontosaurus tener + +, + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”), + +Eubrachiosaurus browni + +, + +Ischigualastia jenseni + +, + +Parakannemeyeria dolichocephala + +, + +Parakannemeyeria ningwuensis + +, + +Pentasaurus goggai + +, some (but apparently not all – see +Camp & Welles 1956 +) specimens of + +Placerias hesternus + +, + +Rhinodicynodon gracile + +, + +Sangusaurus parringtonii + +, + +Shansiodon wangi + +, + +Sinokannemeyeria baidaoyuensis + +, + +Sinokannemeyeria pearsoni + +, + +Sinokannemeyeria sanchuanheensis + +, + +Sinokannemeyeria yingchiaoensis + +, + +Stahleckeria potens + +, + +Wadiasaurus indicus + +, or + +Xiyukannemeyeria brevirostris + +( +Williston 1904 +; +Watson 1917 +; +Pearson 1924b +; +von Huene 1935 +; +Young 1937 +; +Camp 1956 +; +Yeh 1959 +; +Sun 1960 +, +1963 +, 1978; +Tripathi & Puri 1961 +; +Cox 1965 +, +1969 +; + +Bonaparte 1966 +a, 1967 + +; +Colbert 1974 +; +Cheng 1980 +; +Bandyopadhyay 1988 +; +Lucas & Harris 1996 +; +Lucas 1998 +, +2002 +; +Surkov 1998a +; + +Surkov +et al. +2005 + +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +; +Ray 2006 +; + +Govender +et al. +2008 + +; +Govender & Yates 2009 +; + +Kammerer +et al. +2013 + +; + +Angielczyk +et al. +2014 + +, +2017 +; +Liu 2015 +; +Kammerer 2018 +; +Kammerer & Ordoñez 2021 +). It should be noted, however, that the proportions, symmetry, and shape of the distal humeral end are in +Kannemeyeriiformes +subject to change during ontogeny and intraspecific variability ( +Cox 1969 +; + +Kammerer +et al. +2013 + +; + +Angielczyk +et al. +2014 + +). The supinator process is easily noticeable, proximodistally elongated and rugose, but rather low ( +Fig. 13B, E +). This morphology differs from non-kannemeyeriiform dicynodonts and + +Dinodontosaurus tener + +which lack the supinator process ( +Morato 2006 +; +Kammerer 2018 +), but also stahleckeriines ( + +Eubrachiosaurus browni + +, + +Ischigualastia jenseni + +, + +Stahleckeria potens + +) and + +Lisowicia bojani + +, which have a very pronounced, tab-like supinator process ( +Williston 1904 +; +Cox 1965 +; + +Vega-Dias +et al. +2005 + +; + +Dzik +et al. +2008a + +; + +Kammerer +et al. +2013 + +; +Kammerer 2018 +; +Sulej & Niedźwiedzki 2019 +). In that respect, it resembles + +Pentasaurus goggai + +and + +Zambiasaurus submersus + +the most ( +Cox 1969 +; + +Angielczyk +et al. +2014 + +; +Kammerer 2018 +). The intercondylar groove is well-defined both distally and dorsally. The olecranon fossa is well expressed distally but fades towards the base of the distal end, its broadly concave and its anterior (lateral) and posterior (medial) edges are rounded. The trochlea is readily distinguishable but relatively low and restricted in extent. In ventral aspect, most of the surface except for the anterior (lateral) portion is destroyed. Most of the capitulum and the edges of the entepicondylar foramen are not preserved. Distal to the supinator process, on the ventral surface of the bone, a single, oval puncture is present, likely a bite mark ( +Fig. 13B +). The incompleteness of the bone makes it impossible to establish the exact inclination (angle) between the proximal and distal end, but it seems to have been relatively low. + + + +FIG +. 14. — + +Woznikella triradiata + +n. gen.,n. sp. +, ZPAL V.34/1/6, right humerus: +A +, dorsal view; +B +, ventral view; +C +, proximal part in posterior (medial) view; +D +, +E +, distal part in posterior (medial) ( +D +), and anterior (lateral) ( +E +) view; +F +, proximal view; +G +, distal view. Scale bar: 5 cm. + + + + +FIG +. 15. — + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1/10, right radius: +A -E +, proximal part in dorsal (anterior) ( +A +), lateral ( +B +), ventral (posterior) ( +C +), medial ( +D +), and proximal ( +E +) view; +F -J +, distal part in dorsal (anterior) ( +F +), lateral ( +G +), ventral (posterior) ( +H +), medial ( +I +), and distal ( +J +) view. Scale bar: 5 cm. + + + + +Ulna + + + +The probable left ulna ( +ZPAL +V +. 34/1/9; +Fig. 14 +) is mostly complete (despite the lack of the olecranon process), but its proximal end is severely damaged and distorted, making identification difficult. As preserved, the proximal end is wide, roughly triangular in anteroposterior (dorsoventral) view ( +Fig. 14A, B +). In anterior (dorsal) aspect, a clearly marked radial notch and a gently bowed outline of what seems to be the sigmoid notch are visible ( +Fig. 14A +). Although the lack of olecranon may be an effect of damage or immaturity of the individual, the whole edge of the sigmoid process is sharp, unlike the very rounded edges in the specimens of the (juvenile or worn) +holotype +of + +Shaanbeikannnemeyeria “buerdongia + +” figured by +Li (1980) +or juvenile + +Stahleckeria potens + +figured by +Lucas (2002) +and + +Vega-Dias +et al. +(2005) + +. It particularly differs from the ulnae of, e.g., + +Ischigualastia jenseni + +, + +Lisowicia bojani + +, + +Parakannemeyeria youngi + +, + +Placerias hesternus + +, + +Shansiodon wangi + +, + +Sinokannemeyeria yingchiaoensis + +, + +Stahleckeria potens + +, + +Wadiasaurus indicus + +, and the unidentified kannemeyeriid from the Baidaoyu locality (IVPP +V +19365) which have nearly vertical sigmoid notches ( +von Huene 1935 +; +Romer & Price 1944 +; +Camp & Welles 1956 +; +Yeh 1959 +; +Sun 1963 +; +Cox 1965 +; +Bandyopadhyay 1988 +; +Lucas 1993a +; +Ray 2006 +; +Liu 2015 +; +Sulej & Niedźwiedzki 2019 +). Despite the damage, the curvature of the bone is clearly visible, as the proximal end is gently turned and expanded medially. The distal end of the bone is slightly expanded, fusiform in distal view, ends bluntly and is rather featureless ( +Fig. 14 +C-G), in a manner resembling, e.g., juvenile + +Dinodontosaurus tener + +, + +Dolichuranus primaevus + +, + +Jachaleria candelariensis + +, + +Kannemeyeria simocephalus + +, + +Lystrosaurus +spp. + +, + +Parakannemeyeria youngi + +, and + +Zambiasaurus submersus + +(see +Young 1935 +; +Sun 1963 +; +Cox 1969 +; +Colbert 1974 +; +Vega-Dias & Schultz 2004 +; +Govender 2005 +; +Morato 2006 +; + +Govender +et al. +2008 + +; +Govender & Yates 2009 +). Unlike, e.g., + +Dinodontosaurus tener + +as figured by +Cox (1965) +, it lacks a pronounced medial process for contact with the radius. Right above the distal end, in anterior (dorsal) aspect there is a gentle depression, likely accommodating the distal end of the radius, but possibly exaggerated by crushing ( +Fig. 14 +C-E). Despite the crushing, the ulna is significantly more gracile than in the juvenile + +Dinodontosaurus tener + +figured by +Morato (2006) +, + +Lisowicia bojani + +figured by +Sulej & Niedźwiedzki (2019) +, and in + +Stahleckeria potens + +as figured by +Lucas (2002) +and + +Vega-Dias +et al. +(2005) + +. The diagnostic value of this character is unclear, however, as the ulna of the latter was described by + +Vega-Dias +et al. +(2005) + +as diagenetically swollen. + + + +Radius + + + +The proximal end of the right radius ( +ZPAL +V +. 34/1/10. +Fig. 15 +) is crushed, and the proximal part of the shaft and the distal end are missing as well. Despite crushing, the proximal articular surface is subovoid with the lateral portion gently directed ventrally (posteriorly) in a comma-shaped fashion ( +Fig. 15E +).The edge of this portion is gently raised and rounded, surrounding a subtle depression. Medially, this depression is limited by a comparably gentle convexity, comprising most of the articular surface. The raised edges of the lateral portion do not merge smoothly with the convexity, but instead they end medially as low but noticeable bumps, separated from the convexity by shallow fossae.This separation may, however, be an artifact of preservation or preparation, or an effect of incomplete ossification. In the dorsal (anterior) and ventral (posterior) view the edges of the proximal articular surface are sinuous, with the lateral part of the edge higher than the medial part in the anterior (dorsal) view and the medial part of the edge being higher than the lateral part in the posterior (ventral) view. Due to this sinuousness, the articular convexity is exposed in the anterior (dorsal) view ( +Fig. 15A +). This shape may be impacted by crushing, but other dicynodonts (e.g., + +Jachaleria candelariensis + +) show a similar morphology (e.g., +Vega-Dias & Schultz 2004 +). The shaft is relatively slender and as preserved the bone shows only moderate expansion at each end. Distally, the specimen is slightly flattened anteroposteriorly (dorsoventrally) and ends in a sediment-filled concavity, either as a result of taphonomical processes or incomplete ossification due to immaturity of the individual ( +Fig. 15J +). There is no evidence of pronounced distal expansions, such as in + +Stahleckeria potens + +or + +Dinodontosaurus tener + +radii ( +Romer & Price 1944 +; +Lucas 2002 +; + +Vega-Dias +et al. +2005 + +; +Morato 2006 +). Along the anterolateral edge of the bone an elongated muscle scar is present, directed slightly proximomedially. The damage and incompleteness of the specimen preclude observation of further meaningful features.In general outline, the bone resembles the radius of + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”), + +Dinodontosaurus tener + +, at least some species of + +Lystrosaurus + +, + +Shaanbeikannemeyeria xilougouensis + +, + +Sinokannemeyeria baidaoyuensis + +, + +Sinokannemeyeria yingchiaoensis + +, + +Xiyukannemeyeria brevirostris + +, as well as + +Parakannemeyeria +sp. + +, + +Jachaleria candelariensis + +, or + +Wadiasaurus indicus + +, but is not as robust as the latter three ( +von Huene 1935 +; +Young 1935 +; +Sun 1963 +, 1978; +Cox 1965 +; +Colbert 1974 +; +Li 1980 +; +Bandyopadhyay 1988 +; +Lucas & Harris 1996 +; +Vega-Dias & Schultz 2004 +; +Ray 2006 +; +Liu 2015 +; +Liu 2022 +). It differs from the radii of, e.g., + +Ischigualastria +jenseni + +, + +Stahleckeria potens + +, and + +Wadiasaurus indicus + +, which are much less gracile ( +von Huene 1935 +; +Romer & Price 1944 +; +Cox 1965 +; +Lucas 2002 +; + +Vega-Dias +et al. +2005 + +; +Ray 2006 +). + + + +Femur + + + +The left femur ( +ZPAL +V +. 34/1/8; +Fig. 16 +) is preserved in three pieces:the proximal end ( +Fig. 16 +A-E), part of the shaft ( +Fig.16 +FI), and the medial condyle ( +Fig.16 +J-M).The articular surface of the proximal head is nearly circular in proximal view ( +Fig. 16B +) and gently turned anteromedially ( +Fig. 16A +), although still directed mostly proximally. Unlike, e.g., + +Ischigualastia jenseni + +, + +Placerias hesternus + +, + +Rhinodicynodon gracile + +, + +Shansiodon wangi + +, or + +Sinokannemeyeria pearsoni + +, there is no clear neck separating it from the rest of the proximal end ( +Young 1937 +; +Camp & Welles 1956 +; +Yeh 1959 +; +Cox 1965 +; +Surkov 1998a +; + +Kammerer +et al. +2013 + +). Still, the proximal end is more geometrically complex than in most Permian dicynodonts and + +Lystrosaurus +spp. + +(e.g., +Young 1935 +; +Colbert 1974 +; +Yuhe 1983 +; +DeFauw 1986 +; + +Surkov +et al. +2005 + +; +Ray 2006 +). The greater trochanter is roughly half the size of the articular head, both in the anteroposterior ( +Fig. 16A, D +) and in the proximal aspect ( +Fig. 16E +), but it is clearly demarcated, unlike in e.g., + +Dinodontosaurus tener + +, + +Dolichuranus primaevus + +, + +Kannemeyeria simocephalus + +, the specimen figured by +Pearson (1924b) +as + +K. simocephalus + +(but not attributable to that species according to +Govender 2005 +and +Govender & Yates 2009 +), + +Shaanbeikannemeyeria xilougouensis + +, or the unnamed stahleckeriids from the Manda Beds (NMT RB463) and the +Pekin +Formation (NCSM 21719) ( +Pearson 1924b +; +Govender 2005 +; +Morato 2006 +; +Govender & Yates 2009 +; +Green 2012 +; + +Kammerer +et al. +2013 + +, +2017 +; +Liu 2022 +). In proximal aspect it is positioned approximately at the level of the middle of the articular head ( +Fig. 16E +), unlike, e.g., + +Dinodontosaurus tener + +and + +Sangusaurus parringtonii + +, in which the greater trochanter is aligned with the posterior limit of the articular head ( +Morato 2006 +; + +Angielczyk +et al. +2017 + +), but similar to + +Lisowicia bojani + +, + +Tetragonias njalilus + +, the unnamed Manda Beds specimens ( +Cruickshank 1967 +; + +Kammerer +et al. +2017 + +; +Sulej & Niedźwiedzki 2019 +), and + +Placerias hesternus + +(pers. obs.), although in the latter four the major trochanter is less bulbous. The outline of the proximal end in anteroposterior aspect resembles + +Tetragonias njalilus + +specimen UMCZ T754 figured by + +Kammerer +et al. +(2013 + +; but much less so the specimen figured by +Cruickshank 1967 +), and the unnamed Manda Beds stahleckeriid (NMT RB463) both in anteroposterior and mediolateral aspects ( + +Kammerer +et al. +2017 + +). Laterally, the greater trochanter projects a tubercle, narrower than its main body and limited in proximodistal span ( +Fig. 16C +). The third trochanter is not preserved and there is no indication of a pronounced trochanteric crest spanning between the greater and third trochanter, at least in the proximal part of the bone. Unlike most dicynodonts, the mediolateral diameter of the bone starts to clearly decrease just below the tip of the trochanter major ( +Fig. 16 +N-I +′ +). This is unlike, e.g., + +Dolichuranus primaevus + +, + +Dinodontosaurus tener + +, + +Dinodontosaurus +sp. + +(“ + +Dicynodon turpior + +”), + +Lisowicia bojani + +, at least some species of + +Lystrosaurus + +, + +Parakannemeyeria youngi + +, + +Rhinodicynodon gracile + +, + +Sangusaurus parringtonii + +, + +Shaanbeikannemeyeria xilougouensis + +, + +Shansiodon wangi + +, + +Sinokannemeyeria pearsoni + +, + +Stahleckeria potens + +, + +Wadiasaurus indicus + +, the unnamed Denwa Formation specimens, the unnamed +Pekin +Formation specimen (NCSM 21719), the unnamed stahleckeriid from the Santa Rosa Formation (NMMNH P-13001), and the morphotype B stahleckeriid of +Govender (2005) +but similar to, e.g., young + +Kannemeyeria simocephalus + +and possibly + +Parakannemeyeria dolichocephala + +(see +von Huene 1935 +; +Young 1935 +, +1937 +; +Yeh 1959 +; +Sun 1963 +; +Cox 1965 +; +Li 1980 +; +Yuhe 1983 +; +Cruickshank 1986b +; +DeFauw 1986 +; +Bandyopadhyay 1988 +; +Lucas & Harris 1996 +; +Surkov 1998a +; +Bandyopadhyay & Sengupta 1999 +; +Govender 2005 +; +Morato 2006 +; +Ray 2006 +; + +Dzik +et al. +2008a + +; + +Govender +et al. +2008 + +; +Govender & Yates 2009 +; +Green 2012 +; + +Kammerer +et al. +2013 + +; + +Angielczyk +et al. +2017 + +; +Sulej & Niedźwiedzki 2019 +). A clear trochanteric crest is present in the + +Ischigualastia jenseni + +specimen figured by +Cox (1965) +, but in the specimen PVL 3807 the morphology is more reminiscent of +ZPAL +V +. 34/1/8 ( + +Kammerer +et al. +2013 + +).Some dicynodonts,such as + +Acratophorus argentinensis + +, + +Placerias hesternus + +(pers. obs.), and the Manda Beds stahleckeriid (NMT RB463), exhibit a trochanteric crest that originates proximally wide and inconspicuous in height, but becomes thinner and/or more pronounced distally ( + +Bonaparte 1966 +a, 1967 + +, + +Kammerer +et al. +2017 + +), whereas + +Tetragonias njalilus + +has a more distinct separation of the major and third trochanter ( +Fröbisch 2006 +; + +Kammerer +et al. +2017 + +); such morphologies cannot be ruled out for +ZPAL +V +. 34/1/8 due to its incompleteness. Juvenile + +Zambiasaurus submersus + +has a rather shallow trochanteric crest, but the greater trochanter is apparently narrower and less bulbous than in +ZPAL +V +. 34/1/8 – the morphology in adults is unfortunately unknown ( +Cox 1969 +). + + + +FIG +. 16. — Kannemeyeriiform femora: +A -M +, + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1/8, left femur: +A -E +, proximal part in anterior ( +A +), medial ( +B +), lateral ( +C +), posterior ( +D +), and proximal ( +E +) view; +F -I +, shaft in anterior ( +F +), medial ( +G +), lateral ( +H +), and posterior ( +I +) view; +J -M +, medial condyle in anterior ( +J +), medial ( +K +), posterior ( +L +), and distal ( +M +) view; +N -I′ +, outlines of femora (not to scale): +N +, “ +Ruhuhuungulasaurus croucheri +” NHMUK R12710 (after +Kammerer et al. 2017 +); +O +, + +Rhinodicynodon gracile +Kalandadze, 1970 +PIN 1579 + +/50 (after +Surkov 1998a +); +P +, + +Tetragonias njalilus +( +von Huene, 1942 +) GPIT + +/RE/7110 (after + +Kammerer +et al. +2017 + +); +Q +, + +Dinodontosaurus tener +( +von Huene,1935 +) UFRGS + +/PV0113T (after +Morato 2006 +); +R +, + +Acratophorus argentinensis +Kammerer & Ordoñez, 2021 +PVL + +3465 ( +Bonaparte 1966a +); +S +, + +Wadiasaurus indicus +Chowdhury, 1970 +ISI R + +172/1 (after +Bandyopadhyay 1988 +); +T +, + +Sinokannemeyeria pearsoni +Young, 1937 + +(number not given, after +Young 1937 +); +U +, +Parakannemyeria + +youngi +Sun, 1963 +IVPP V + +972 (after +Sun 1963 +); +V +, + +Shaanbeikannemeyeria xilougouensis +Cheng, 1980 +IVPP V + +6033 (after +Li 1980 +); +W +, young + +Kannemeyeria simocephalus +( +Weithofer, 1888 +) UMZC T + +757 (after +Kammerer et al. 2017 +); +X +, + +Kannemeyeria simocephalus +( +Weithofer, 1888 +) NHMUK R + +3740 (after +Kammerer et al. 2017 +); +Y +, + +Woznikella triradiata + +n. gen., n. sp. +, ZPAL V. 34/1/8; +Z +, + +Stahleckeria potens +von Huene, 1935 +GPIT + +unnumbered (after +Kammerer et al. 2017 +); +A′ +, + +Sangusaurus parringtonii +Cruickshank,1986a + +UMZC T1225 (after +Kammerer et al.2017 +); +B′ +, + +Ischigualastia jenseni +Cox, 1962 +PVL + +3807 (after +Kammerer et al. 2017 +); +C′ +, + +Ischigualastia jenseni +MCZ + +3120 (after +Cox 1965 +); +D′ +, stahleckeriid NMT RB463 (after +Kammerer et al.2017 +); +E′ +, stahleckeriid NMMNH P-13001 (after + +Kammerer +et al. +2013 + +); +F′ +, morphotype B stahleckeriid BP/1/3518 (after +Govender 2005 +); +G′ +, + +Zambiasaurus submersus +Cox,1969 +NHMUK R + +9123 (after +Angielczyk et al. 2014 +); +H′ +, + +Lisowicia bojani +Sulej & Niedźwiedzki, 2019 +ZPAL V. + +33/74 (after +Sulej & Niedźwiedzki 2019 +); +I′ +, + +Placerias hesternus +Lucas, 1904 +UCMP + +3294 (after + +Kammerer +et al. +2013 + +). Indicated is an approximate angle between the dorsal (just lateral to the head) and lateral (immediately behind the curve) surfaces of the greater trochanter ( +green +if acute, +red +if obtuse) as well as the arrangement between the proximal medial (just distal to the head) and lateral edges of the bone: +green +if clearly converging (tangents meet along or just distal to the bone length, the bone decreases in diameter in its proximal part), +orange +if subparallel (tangents never meet or meet far beyond the distal end of the bone, the bone keeps roughly constant diameter in its proximal part), +red +if clearly diverging (tangents never meet, the bone increases in diameter in its proximal part). Scale bar: 5 cm. + + + + +FIG +. 17. — Phylogeny of +Kannemeyeriiformes +: +A +, strict consensus tree based on the analysis with all taxa included; +B +, majority rule (50%) consensus tree based on the analysis with + +Ufudocyclops mukanelai +Kammerer,Viglietti,Hancox,Butler & Choiniere, 2019 + +inactivated (the topology used for paleobiogeographic analyses; strict consensus differs only in the presence of an unresolved polytomy of + +Eubrachiosaurus browni +Williston, 1904 + +, + +Sangusaurus parringtonii +Cruickshank, 1986a + +, and + +Sangusaurus +sp. + +). Numbers next to nodes indicate bootstrap values above 50. Lighter parts of stratigraphic ranges indicate uncertainty of the ages of the Manda, Ntawere, and Yerrapalli formations (see text for discussion). + + + + +FIG +. 18. — Paleobiogeographic analysis of dicynodonts.Ancestral states estimated based on BioGeoBEARS analysis (DEC+J).Note that region names are based on the known dicynodont-yielding localities included in the regions and thus do not accurately reflect their potential geographic extent.Triassic taxa indicated in +bold +. + + + + +TABLE +3. — Occurrences of Triassic tracks possibly produced by dicynodonts.Note that the list is not comprehensive and likely numerous other mentions exist, particularly in local, non-English literature. The Country column includes all mentions deemed significant (i.e., not merely listing taxa from original literature, but providing mentions, descriptions, and/or images of new specimens, original data or reinterpretations of morphology, confirming the geographical and/or temporal presence of taxa, confirming or providing novel insights into their validity, or at least presenting them in a novel biostratigraphic context). For simplicity, the Formation column includes only the references providing the most recent and most precise subdivisions either verbatim or in a form unambiguously recognizable without extensive research of local geological literature. The historically used + +Lystrosaurus +Cope, 1870 +AZ + +was recently redefined and renamed to + +Lystrosaurus declivis +Brink, 1951 +AZ ( +Botha & Smith 2020 +) + +. Because of the presence of + +Lystrosaurus +spp. + +also in the underlying + +Lystrosaurus maccaigi +( +Seeley, 1898 +) + +- + +Moschorhinus +Broom, 1920 +SZ + +of +Daptocephalus Hoepen, 1934 AZ +(former +Dicynodon AZ +), the meaning of the historical + +Lystrosaurus +AZ + +is somewhat ambiguous and may be dependent on the author (see Table 2). Therefore, the name is used here in parentheses, implying that although in most cases it was probably synonymous with + +Lystrosaurus declivis +AZ + +, it can potentially include the top of the +Daptocephalus AZ. If +the occurrence is not explicitly mentioned in the context of a changed naming scheme, the papers allowing referral (e.g., naming both the old formation and the new formation) are provided. Age includes selected single most recent original study with preference towards radiometric dates, unless those are unavailable, and a sound disagreement exists between several studies. Note that the bottom age (252.24 ± 0.11 Ma) of the + +Lystrosaurus declivis +AZ + +obtained by + +Gastaldo +et al. +(2020) + +is barely about 0.3 Ma older than the currently recognized Triassic/Permian boundary (251.902 ± 0.024 Ma). See + +Botha +et al. +(2020) + +for the entirely Triassic estimation of the age of that assemblage and +Botha & Smith (2020) +for discussion. Abbreviation: +AZ +, assemblage zone. At least some + +Dicynodontipus + +ispp. could be produced by cynodonts (e.g. + +Marchetti +et al. +2019 + +) + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
TaxonCountryFormation +Age +
?Dicynodont tracks +Poland ( + +Sulej +et al. +2011 + +; Sadlok & Wawrzyniak 2013) + +Grabowa ( +Sadlok & Wawrzyniak 2013 +; ( + +Szulc +et al. +2015b + +; +Racki & Lucas 2020 +) + +Carnian ( + +Sulej +et al. +2011 + +)/ Norian (( + +Szulc +et al. +2015b + +) +
+” + +Dicynodontipus + +” +bellambiensis +Retallack, 1996 + +Australia ( +Retallack 1996 +; + +Klein +et al. +2015 + +; + +Francischini +et al. +2018 + +; Díaz-Martínez +et al. +2019; +Klein & Lucas 2021 +) + +Coal Cliff Sandstone ( +Retallack 1996 +; + +Francischini +et al. +2018 + +; Díaz-Martínez +et al. +2019) + +Induan ( +Retallack 1996 +) +
+ +Dicynodontipus geinitzi + +( +Hornstein, 1876 +) + +England ( +Demathieu & Haubold 1971 +; +Haubold 1974 +, +1984 +) + +Helsby Sandstone (Demathieu & +Haubold 1971 +; +Pollard 1981 +) + +Late Olenekian-Anisian ( + +Medici +et al. +2019 + +) +
+Germany ( +Hornstein 1876 +; Rühle von +Lilienstern 1944 +; Demathieu & Haubold 1971; +Haubold 1971 +, +1974 +, +1984 +, +Demathieu & Fichter 1989 +; Francischini + +et al. +2018 + +; + +Díaz-Martínez +et al. +2019 + +; +Klein & Lucas 2021 +) +Middle Buntsandstein (Demathieu & Fichter 1989) +Early Triassic (German Stratigraphic +Commission 2016 +) +
+ +Dicynodontipus + +isp. + +Argentina ( +Casamiquela 1964 +, +1975 +; +Haubold 1971 +, +1974 +, +1984 +; Leonardi & Oliveira 1990; +Leonardi 1994 +; Marsicano & Barredo 2004; Melchor & de Valais 2006; +Domnanovich & Marsicano 2006b +; +Calvo 2007 +; + +Domnanovich +et al. +2008 + +; + +Francischini +et al. +2018 + +; + +Citton +et al. +2018a + +, b, 2019, 2021; + +Lagnaoui +et al. +2019 + +; + +Díaz-Martínez +et al. +2019 + +; De Valais +et al. +2020) + +Cerro de Las Cabras (Melchor & de Valais 2006; + +Francischini +et al. +2018 + +; + +Díaz-Martínez +et al. +2019 + +; Lagnaoui +et al. +2019) + +Late Anisian-early Ladinian ( + +Cariglino +et al. +2016 + +) +
+Corral de Piedra (Marsicano & Barredo 2004; +Melchor & de Valais 2006 +; + +Díaz-Martínez +et al. +2019 + +) +Late Carnian (Marsicano & Barredo 2004)
+Sierra de Las Higueras (Leonardi 1994; +Melchor & de Valais 2006 +; + +Francischini +et al. +2018 + +) + +Ladinian (Francischini +et al. +2018) +
+Los Menucos Complex (Melchor & de Valais 2006; Domnanovich & Marsicano 2006b; +Calvo 2007 +; + +Domnanovich +et al. +2008 + +; Citton +et al. + +2018 +a, b, 2019 + +, +2021 +; Francischini + +et al. +2018 + +; + +Díaz-Martínez +et al. +2019 + +; + +De Valais +et al. +2020 + +) + +Late Permian-Early Triassic ( +Falco 2019 +) +
+Brazil ( + +Francischini +et al. +2018 + +; + +Díaz-Martínez +et al. +2019 + +) + +Pirambója ( + +Francischini +et al. +2018 + +; + +Díaz-Martínez +et al. +2019 + +) + +Late Permian-Induan ( + +Francischini +et al. +2018 + +) +
+Catalonia ( + +Valdiserri +et al. +2009 + +; + +Díaz-Martínez +et al. +2019 + +) + +Lower Buntsandstein (Valdiserri +et al. +2009) + +Early Triassic (German Stratigraphic +Commission 2016 +) +
+England ( +Haubold 1971 +; +Pollard 1981 +; + +Francischini +et al. +2018 + +) + +Helsby Sandstone ( +Pollard 1981 +; + +Francischini +et al. +2018 + +) + +Late Olenekian-Anisian ( + +Medici +et al. +2019 + +) +
+Germany ( +Demathieu & Haubold 1971 +; +Haubold 1984 +; +Klein & Lucas 2018 +; + +Díaz-Martínez +et al. +2019 + +) + +Buntsandstein (Demathieu & Haubold 1971; +Haubold 1984 +) + +Olenekian-early Anisian (German Stratigraphic +Commission 2016 +) +
Middle Muschelkalk (Klein & Lucas 2018) +Anisian (German Stratigraphic +Commission 2016 +) +
+cf. + +Dicynodontipus + +isp. + +South Africa ( + +Díaz-Martínez +et al. +2019 + +; + +Marchetti +et al. +2019 + +; +Botha & Smith 2020 +) + + +Lystrosaurus declivis +AZ + +(Botha & +Smith 2020 +) + +Latest Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+? + +Dicynodontipus + +isp. + +Germany ( +Demathieu & Haubold 1971 +) + +Lower and Middle Buntsandstein ( +Demathieu & Haubold 1971 +) + +Early Triassic (German Stratigraphic +Commission 2016 +) +
+ +Dolomitipes accordii + +(Ceoloni, Conti, Mariotti & Nicosia, 1986 + +South Africa ( +Watson 1960 +; +Haubold 1971 +; +Smith & Botha-Brink 2014 +; Marchetti +et al. +) 2019; +Klein & Lucas 2021 +) + +“ + +Lystrosaurus +AZ + +” ( +Watson 1960 +; +Haubold 1971 +; + +Marchetti +et al. +2019 + +; +Klein & Lucas 2021 +) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
+ + + +Table 3 +. — Continuation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Taxon + +Country + +Formation + +Age +
+ +Dolomitipes + +isp. + +South Africa ( +Smith & Botha-Brink 2014 +; + +Marchetti +et al. +2019 + +) + +“ + +Lystrosaurus +AZ + +” (Marchetti +et al. +2019) + +Late Permian-Early Triassic ( + +Gastaldo +et al. +2020 + +) +
?Lystrosaurid tracks +Antarctica ( + +MacDonald +et al. +1992 + +) + +Middle Fremouw (MacDonald +et al. +1992) + +Early Anisian (Elliot +et al. +2017) +
+ +Pentasauropus + +argentinae +Lagnaoui, Melchor, Bellosi, Villegas & Espinoza, 2019 + +Argentina ( +Marsicano & Barredo 2004 +; +Porchetti & Nicosia 2007 +; Citton et al. 2018a; + +Díaz-Martínez +et al. +2019 + +; Lagnaoui +et al. +2019; +Klein & Lucas 2021 +) + +Cerro de las Cabras (Díaz-Martínez +et al. +2019; + +Lagnaoui +et al. +2019 + +) + +Late Anisian-early Ladinian ( + +Cariglino +et al. +2016 + +) +
+Corral de Piedra (Díaz-Martínez +et al. +2019; + +Lagnaoui +et al. +2019 + +) + +Ladinian (Barredo +et al. +2012) +
+Los Menucos Complex (Domnanovich +et al. +2008; + +Citton +et al. +2018a + +; Díaz-Martínez +et al. +2019; Lagnaoui +et al. +2019 +) + +Late Anisian-?Rhaetian ( + +Díaz-Martínez +et al. +2019 + +) +
+ +Pentasauropus + +incredibilis +Ellenberger, 1972 + +Lesotho ( +Ellenberger 1955 +, +1970 +, +1972 +; +Ellenberger & Ellenberger 1958 +, +1960 +; + +Ellenberger +et al. +1970 + +; +Haubold 1971 +, +1974 +, +1984 +; +Olsen & Galton 1984 +; +Porchetti & Nicosia 2007 +; Bordy +et al. +2017 +; +Kammerer 2018 +; Díaz-Martínez +et al. +2019 +; +Viglietti 2020 +; +Klein & Lucas 2021 +) + +Lower Elliot + +Scalenodontoides +AZ + +( + +Viglietti +et al. +2020b + +) +Norian-Rhaetian (Bordy et al. 2020)
+ +Pentasauropus + +isp. + +Argentina ( + +Domnanovich +et al. +2008 + +; Citton +et al. +2018 +a, b, 2019 +; Díaz-Martínez et al. 2019; + +Lagnaoui +et al. +2019 + +; De Valais +et al. +2020) + +Los Menucos Complex (Domnanovich +et al. +2008; + +Citton +et al. +2018a + +, b, 2019; + +Díaz-Martínez +et al. +2019 + +; + +Lagnaoui +et al. +2019 + +; De Valais +et al. +2020 +) + +Late Anisian-?Rhaetian ( + +Díaz-Martínez +et al. +2019 + +) +
+United States ( +Lockley & Hunt 1995 +; + +Lockley +et al. +1996 + +; + +Gaston +et al. +2003 + +; +Hunt & Lucas 2007 +; Porchetti & Nicosia 2007; + +Hunt-Foster +et al. +2016 + +; Díaz-Martínez +et al. +2019) + +Rock Point ( + +Lockley +et al. +1996 + +; + +Gaston +et al. +2003 + +; Hunt & Lucas +2007 +; + +Hunt-Foster +et al. +2016 + +; + +Díaz-Martínez +et al. +2019 + +) + +Rhaetian (Ramezani +et al. +2014) +
+cf. + +Pentasauropus + +isp. + +United States ( +Olsen & Galton 1984 +; +Conrad & Lockley 1986 +; Porchetti & Nicosia 2007; + +Díaz-Martínez +et al. +2019 + +) + +Chinle Group ( +Conrad & Lockley 1986 +; +Porchetti & Nicosia 2007 +) + +Norian-Rhaetian (Ramezani +et al. +2014) +
+Gettysburg ( +Olsen & Galton 1984 +; + +Díaz-Martínez +et al. +2019 + +) + +Norian ( +Gee & Jasinski 2021 +) +
?Therapsid tracks +Brazil ( +Leonardi 1994 +; Francischini +et al. +2018) +Lower Sanga do Cabral (Leonardi 1994) +Early Triassic (Dias-da-Silva +et al. +2017) +
+United States ( + +Conrad +et al. +1987 + +; Lockley & Hunt 1995; +Hunt & Lucas 2007 +) + +Sloan Canyon ( + +Conrad +et al. +1987 + +; +Hunt & Lucas 2007 +) +Rhaetian (Hunt & Lucas 2007)
Therapsid tracks +France ( +Ellenberger 1965 +; + +Courel +et al. +1968 + +) + +Not given ( +Ellenberger 1965 +; Courel +et al. +1968) + +Triassic ( + +Courel +et al. +1968 + +) +
+Italy ( + +Avanzini +et al. +2011 + +) + +Not given ( + +Avanzini +et al. +2011 + +) + +Anisian ( + +Avanzini +et al. +2011 + +) +
+Poland ( +Klein & Niedźwiedzki 2012 +; + +Francischini +et al. +2018 + +) + +Wióry ( +Klein & Niedźwiedzki 2012 +; + +Francischini +et al. +2018 + +) +Olenekian (Klein & Niedźwiedzki 2012)
+ +Therapsipus cumminsi + +Hunt, Santucci, Lockley & Olson, 1993 + +United States ( + +Hunt +et al. +1993 + +; Lockley & Hunt 1995; +Nesbitt & Angielczyk 2002 +; +Klein & Lucas 2010 +, +2021 +) + +Moenkopi Holbrook Member (Hunt +et al. +1993; +Lockley & Hunt 1995 +; +Nesbitt & Angielczyk 2002 +; Klein & +Lucas 2010 +, +2021 +) + +Late Anisian (Haque +et al. +2021) +
+ +Therapsipus + +isp. + +United States ( +Hunt & Lucas 1993 +; Lucas +et al. +2003; +Klein & Lucas 2010 +) + +Moenkopi Anton Chico Member ( +Hunt & Lucas 1993 +; Lucas +et al. +2003; +Klein & Lucas 2010 +) + +Late Anisian (Haque +et al. +2021) +
+cf. + +Therapsipus + +isp. + +France ( + +Courel +et al. +1968 + +; + +Hunt +et al. +1993 + +) + +Not given ( + +Courel +et al. +1968 + +; Hunt +et al. +1993) + +Early Triassic (Courel +et al. +1968) +
+Poland ( +Klein & Niedźwiedzki 2012 +; Klein & +Lucas 2021 +) + +Wióry ( +Klein & Niedźwiedzki 2012 +; +Klein & Lucas 2021 +) +Olenekian (Klein & Niedźwiedzki 2012)
+ + +The base of the proximal end is flattened anteroposteriorly, as is the preserved part of the shaft ( +Fig. 16 +F-I). The mediolateral diameter of these parts is similar, unlike in + +Lystrosaurus +spp. + +, + +Sangusaurus parringtonii + +, + +Shansiodon wangi + +, or + +Wadiasaurus indicus + +in which the femur exhibits a clear constriction along the diaphysis ( +Young 1935 +; +Yeh 1959 +; +Yuhe 1983 +; +Cruickshank 1986b +; +DeFauw 1986 +; +Bandyopadhyay 1988 +; +Ray 2006 +; + +Angielczyk +et al. +2017 + +). The medial condyle of the distal end ( +Fig. 16 +J-M) terminates at a relatively sharp angle in the anterior aspect and is relatively narrow and elongated anteroposteriorly in ventral view ( +Fig. 16M +), similar as in the unnamed Manda Beds stahleckeriid ( + +Kammerer +et al. +2017 + +). Its surface is rugose. The preserved fragment indicates a clear separation of the condyles both in the anteroposterior and in the distal aspect. + + + + \ No newline at end of file diff --git a/data/03/82/87/038287CBFFC2FFA7792751CEEB3DB54F.xml b/data/03/82/87/038287CBFFC2FFA7792751CEEB3DB54F.xml new file mode 100644 index 00000000000..1100518cbbc --- /dev/null +++ b/data/03/82/87/038287CBFFC2FFA7792751CEEB3DB54F.xml @@ -0,0 +1,92 @@ + + + +Woznikella triradiata n. gen., n. sp. - a new kannemeyeriiform dicynodont from the Late Triassic of northern Pangea and the global distribution of Triassic dicynodonts + + + +Author + +Szczygielski, Tomasz +Institute of Paleobiology, Polish Academy of sciences, Twarda 51 / 55, 00 - 818 Warsaw (Poland) +szczygielski@twarda.pan.pl + + + +Author + +Sulej, Tomasz +Institute of Paleobiology, Polish Academy of sciences, Twarda 51 / 55, 00 - 818 Warsaw (Poland) +sulej@twarda.pan.pl + +text + + +Comptes Rendus Palevol + + +2023 + +2023-05-16 + + +22 + + +16 + + +279 +406 + + + + +http://dx.doi.org/10.5852/cr-palevol2023v22a16 + +journal article +305805 +10.5852/cr-palevol2023v22a16 +96b937a1-56d3-4b29-93b2-5b859c159da7 +1777-571X +14260551 +urn:lsid:zoobank.org:pub:5BEBCC73-819E-47AE-9610-1E50B1AD6B60 + + + + + + +Woznikella + +n. gen. + + + + + + +urn:lsid:zoobank.org:act: +39A734F4-1403-43C1-9D99-1DB3F1901478 + + + + + + + +TYPE + + + +SPECIES + +. — + +Woznikella triradiata + +n. sp. + + + + \ No newline at end of file diff --git a/data/03/B0/DE/03B0DE22FFA0322F8FDCF9F4FAACA2A0.xml b/data/03/B0/DE/03B0DE22FFA0322F8FDCF9F4FAACA2A0.xml new file mode 100644 index 00000000000..8c061ebc299 --- /dev/null +++ b/data/03/B0/DE/03B0DE22FFA0322F8FDCF9F4FAACA2A0.xml @@ -0,0 +1,621 @@ + + + +Insect (Hexapoda) diversity in the oceanic archipelago of Fernando de Noronha, Brazil: Lauxaniidae (Diptera) + + + +Author + +Soares, Matheus M. M. +Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil + + + +Author + +Gaimari, Stephen D. +Dipterists Society, P. O. Box 231113, Sacramento, California 95823, USA + + + +Author + +Corrêa-Neto, José De J. +Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil + + + +Author + +Limeira-De-Oliveira, Francisco +Universidade Estadual do Maranhão, Laboratório de Estudos dos Invertebrados, Praça Duque de Caixas, Morro do Alecrim, s / n, 65604 - 380 Caxias, MA, Brazil + + + +Author + +Rafael, José A. +Instituto Nacional de Pesquisas da Amazônia, INPA, Coordenação de Biodiversidade, Manaus, Amazonas, Brazil + +text + + +Zootaxa + + +2024 + +2024-11-07 + + +5537 + + +1 + + +1 +23 + + + + +http://dx.doi.org/10.11646/zootaxa.5537.1.1 + +journal article +305806 +10.11646/zootaxa.5537.1.1 +5944fc67-97a0-4a74-87b1-12f5c691bd58 +1175-5326 +14239256 +91F512AE-5BAC-458D-992D-DF4EAF055183 + + + + + + + +Poecilominettia octovittata +( +Williston, 1896 +) + + + + + + + +( +Figs. 7–8 +) + + + + + + + +Sapromyza octovittata +Williston, 1896: 382 + + +. + + + + + + +Type locality: +Saint Vincent +( +West Indies +). +Lectotype +female +(designated by + +Gaimari & Silva +2020 + +), Natural History Museum, London, United Kingdom (examined +SDG +). + + + + + + +Poecilominettia +sp. 2 + +( + +Rafael +et al. +2020 + +). + + + + +Diagnosis. +Easily differentiated from the other congeneric species by the distinct pattern of spots on the thorax, wing and abdomen ( +Fig. 7A, B +); femora yellow, tibiae with a ventral brown spot near base ( +Fig. 7A, B +); anterior margin of wing dark brown ( +Fig. 7F +); surstylus ovoid, 1.5X longer than wide at widest point, with 2 pointed processes, 1 at middle of inner edge and 1 at apex ( +Fig. 8A, B +). + + + + +FIGURE 7. + +Poecilominettia octovittata +( +Williston, 1896 +) + +. +A, C–H +male (INPA), +B +(living female from municipality of Olha d’Água das Flores, state of Alagoas), +I, J +female (INPA). +A, B. +Habitus, lateral view; +C, D. +Head anterior and lateral views, respectively; +E. +Head and thorax, dorsal view; +F. +Wing; +G. +Male terminalia, posterior view; +H. +Male abdomen, dorsal view; +I–J. +Female abdomen, dorsal and ventral views respectively. Abbreviations: afrs = anterior fronto-orbital seta; dc str = dorsocentral stripe; ep spt = spot on epandrium; fc spt = facial spot; kpt s = katepisternal seta; flgm 1 = first flagellomere; ot = ocellar triangle; sal str = supra-alar stripe. Photograph +B +provided by Alenilson Rodrigues. + + + + +FIGURE 8. + +Poecilominettia octovittata +( +Williston, 1896 +) + +. +A–D +male (INPA), +E–H +female (INPA). +A, B. +Epandrium in posterior and lateral views, respectively; +C, D. +Internal appendages in lateral and anterior views, respectively; +E–G +. Female terminalia in lateral, dorsal and ventral views, respectively; +H. +Spermathecae. Abbreviations: bp = basiphallus; cerc = cercus; dp = distiphallus; e = epiproct; epand = epandrium; h = hypoproct; hypd = hypandrium; ej apod = ejaculatory apodeme; te = teeth of distiphallus; st = sternite; tg = tergite; pgt = postgonite; sur = surstylus. + + + + +Redescription. +Head +( +Fig. 7A–E +). Mostly yellowish, except frons with a narrow central dark brown median stripe extended from ocellar triangle to edge of lunule, subparallel to tapered anteriorly; face with an ovoid, median dark brown spot at lower margin; and with dark brown spot in area of gena and bottom of parafacial. Head 1.6X higher than long, 1.3X wider than high, 1.1X wider than scutum; eye 1.4X higher than long; gena height (directly below eye) 0.2X eye height. Vertex silvery-yellow pruinose, rounded; inner vertical seta long and strong, about 1.5X longer than outer vertical seta, with distance between them about 1/3 that from inner vertical seta to central vertex. Ocellar triangle dark brown, small, slightly raised, placed slightly in front of vertex, at same level as inner vertical seta; ocelli subequal in size and arranged in small equilateral triangle, covered with few tiny setulae. Ocellar setae proclinate and divergent, weak. Postocellar setae cruciate, slightly longer than ocellar seta. Two rows of short postocular setulae. Median occipital sclerite with silvery pruinosity and supracervical setae above occipital foramen. Frons 1.2X wider than long, frons flat, curved evenly into facial plane; with 2 strong reclinate fronto-orbital setae, posterior seta slightly longer than anterior seta, located near midpoint between inner vertical seta and anterior seta; anterior seta closer to lunule than to posterior seta; frons setulose at anterior 1/2, wholly bronzy pruinose except for central brown stripe. Lunule low, straight. Antenna mostly orangish-yellow, except scape and arista dark brown; scape with few short dorsal setae; pedicel with 1 long dorsal seta at middle length, with crown of short setae at apex and 3–4 long ventral setae; first flagellomere about 1.6X longer than high, and about 2X longer than scape and pedicel combined, with rounded apex; arista inserted dorsobasally on first flagellomere, covered with short and sparse rays. Gena with dense row of strong setae along ventral edge, extended onto lower parafacial; postgena with short and strong setae. Face about 1.2X wider than long, face and parafacial silvery pruinose, except parafacial darkened ventrally. Clypeus orange, narrow. Maxillary palpus and labellum brown, ventral surface of maxillary palpus with rows of long setae. + + +Thorax +( +Fig. 7A, B, E +). Scutum slightly arched, about 1.2X longer than wide; scutellum about 1/3 as long as scutum, with width at base about 1.75X greater than length; scutum orangish-yellow, with dorsocentral and supra-alar brown stripes, pale yellow on postpronotum through notopleuron; scutellum pale yellow with dark brown lateral stripes as extended through dorsocentral stripes of scutum; pleural region mostly orangish-yellow, except anterodorsal quadrant of anepisternum, upper margin of katepisternum, lower margin of anepimeron, and meron dark brown. Chaetotaxy: 0+3 dorsocentral setae, anterior seta slightly shorter than middle seta, located close to transverse suture, middle seta located at midpoint between posterior seta and anterior seta; prescutellar acrostichal seta present, weak; 1 postpronotal seta; 2 notopleural setae, in anterior and posterior corners, anterior seta slightly longer than posterior; 1 postsutural intra-alar seta present; 1 presutural and 1 postsutural supra-alar setae; 2 postalar setae, in anterior and posterior corners; 7–8 irregular rows of acrostichal setulae between dorsocentral setal rows, row of setulae along dorsocentral area present, and setulose outside dorsocentral row; proepisternal seta present, strong; anepisternum with anepisternal seta along posterior edge, otherwise setulose in posterior 2/3; anepimeron bare; 2 katepisternal setae present, both strong, posterior one slightly stronger, katepisternum otherwise setulose through middle, and with group of long ventral setae, close to the coxa; 2 pairs scutellar setae, anterior setae subparallel, posterior setae cruciate. +Legs +( +Fig. 7A, B +). Pale yellow, except all tibiae with a ventral brown spot near base. Fore femur with posteroventral row of long and strong setae, about 1.5X longer than width of femur, posterior and posterodorsal rows of setae from base to apex, shorter and thinner than posteroventral row of setae; ctenidium absent. Mid femur with anterior row of short and strong setae in apical 1/2, and with 1 short curved posteroventral preapical seta. Hind femur with anteroventral row of short setae in apical 1/3 and 1 anterodorsal long preapical seta. All tibiae with preapical dorsal seta, strongest on mid tibia; mid tibia with 1 longer preapical ventral seta, as long as dorsal seta; hind tibia with 1 short apical ventral spur. +Wing +( +Fig. 7A, F +). Mostly hyaline, except dark brown in anterior 1/3 (through cells sc, r +1 +, and most of r +2+3 +) and curved around distal part of wing to the tip of vein M +1 +, on crossveins rm and dm-m, and with a spot at the midpoint of the distal part of vein M +1 +. Veins dark brown, except yellow in basal veins and sections of veins R +4+5 +and M +1 +( +Fig. 7F +). Length +3.5 mm +; 2.5X longer than high. Cell dm about 4X longer than crossvein dm-m. Crossvein r-m located at midpoint of cell dm. Vein R +4+5 +ending at wing tip, subparallel with veins R +2+3 +and M +1 +. Crossvein dm-m straight, located slightly basal to midpoint between crossvein r-m and tip of vein M +1 +. Vein M +4 +about 0.2X as long as crossvein dm-m. Vein CuA+CuP short, about 1/2 length of vein A +1 +. + + +Abdomen +( +Fig. 7G, H +). Tapered gradually after segment 4; mostly yellow, except for lateral margins of syntergite 1+2 and tergites 3–6 dark brown forming a narrow stripe, paired semicircular brown spots along posterolateral margins of syntergite 1+2 and tergites 3–6, and central brown macula on tergites 3–6 forming a narrow stripe. Tergites covered with short setulae, longest and strongest along posterior edges of syntergite 1+2 and tergites 3–6. Sternites pale yellow to white, except lateral margins of sternites 3–5 dark brown, sternite 1 bare, sternites 2–5 setulose, with slightly longer setae at lateral margins; sternites 2–5 each slightly wider than long, subequal in width sternite to sternite; sternite 6 a membranous transverse strip. +Male genitalia +( +Figs. 7G +, +8A–D +). Pale yellow, epandrium dorsally with rectangular dark brown spot at middle. Syntergosternite 7+8 simple, bare, transversely saddle-shaped. Epandrium simple, saddle-shaped; mostly bare, with median row of short setae and with row of long setae at posterior edge, 2X longer than widest point. Surstylus large, leaf-shaped in lateral view, 1.5X longer than wide at widest point, with 2 medially-pointed processes, inner process at middle of inner edge and apical process at apex, with sparse short setulae in posterior 1/2, close to the inner process and with 2 short inner setae above the process. Subepandrial sclerite narrow, about 5X longer than wide. Cercus short, ovoid, heavily setose. Ejaculatory apodeme long and narrow, weak sclerotized. Phallus mostly tubular, widened towards apex, about 2X longer than wide; apex of distiphallus with short, sclerotized teeth, 2 apical long and pointed teeth, apparently articulated with distiphallus, 1 short and stout tooth at base of distiphallus. Hypandrium a narrow transverse band. Postgonite short, rounded and weak sclerotized, with small seta. + + +Female: +Similar to male. Sternite 6 pale yellow to white, except lateral margin dark brown (as in sternites 3–5) ( +Fig. 7K +). +Female terminalia +( +Figs. 7I, J +, +8E–H +). Tergite 7 not fused to sternite 7. Tergite 8 narrow, somewhat saddle-shaped, weakly sclerotized, with posterior row of setae. Sternite 8 somewhat quadrangular, lateral edges curved, heavily setose, dark brown, except for median rectangular yellow area. Epiproct narrow, with row of strong setae. Hypoproct simple, semicircular, heavily setulose. Spermathecae round, with configuration 1+2, paired spermathecae; surface smooth. + + + + +Material examined. + + +BRAZIL +, + +Pernambuco +, +Fernando de Noronha +: +Capim-Açu +, +03°51′17″S +– +32°26′26″W +, + +09–24.vi.2019 + +, +large Malaise trap +, +J.A. Rafael +, +F. Limeira-de-Oliveira +& +L.C. Castro +( +7 ♂ +, +36 ♀ +, +INPA +; +1 ♂ +, +3 ♀ +, +AMNH +; +1 ♂ +, +3 ♀ +, +CSCA +; +1 ♂ +, +3 ♀ +, +USNM +; +1 ♂ +, +3 ♀ +, +SDG +) + +; + +same data, except + +08–27.x.2019 + +( +1 ♀ +, dissected, +INPA +) + +; + +same data, except +03°51′30″S +– +32°25′50″W +, + +01–09.vi.2019 + +, varredura [sweeping], +J.A. Rafael +, +F. Limeira-de-Oliveira +& +D.M.M. Mendes +( +9 ♀ +, +INPA +) + +; + +Sueste Mangue +, +03°51′30″S +– +32°25′50″W +, + +01–09.vi.2019 + +, armadilha luminosa [ +light trap +] ( +3 ♂ +, +7 ♀ +, +INPA +) + +; + +same data, except + +20–27.ii.2020 + +, +Malaise trap +, +J.A. Rafael +, +F. Limeira-de-Oliveira +& +P.C. Grossi +( +1 ♂ +, +4 ♀ +, +INPA +) + +; + +Trilha Sancho +, +03°51′30″S +– +32°25′50″W +, + +01–09.vi.2019 + +, +Malaise trap +, +J.A. Rafael +, +F. Limeira-de-Oliveira +& +D.M.M. Mendes +( +11 ♂ +, 2 dissected, +17 ♀ +, 2 dissected, +INPA +; +1 ♂ +, +1 ♀ +, +AMNH +; +1 ♂ +, +1 ♀ +, +CSCA +; +1 ♂ +, +1 ♀ +, +USNM +; +1 ♂ +, +1 ♀ +, +SDG +) + +; + +same data, +except CDC-UV light trap +( +2 ♂ +, +3 ♀ +, +INPA +) + +; + +same data, except Caracas ( +1 ♀ +, +INPA +) + +. Additionally, 16,422 specimens from nearly all localities and dates in ethanol. + + + + +FIGURE 9. +Monthly averages of temperature, humidity and rainfall in the Fernando de Noronha archipelago, Pernambuco, Brazil. + + + + +FIGURE 10. +Pearson correlation between weather variables in the Fernando de Noronha archipelago, Pernambuco, Brazil. + + + + +FIGURE 11. +Seasonal occurence of + +Pachyopella flavida +( +Wiedemann, 1824 +) + +, + +Poecilominettia erebus +Soares & Gaimari + +, + +sp. nov. + +and + +Poecilominettia octovittata +( +Williston, 1896 +) + +, collected with interception traps in the Fernando de Noronha archipelago, Pernambuco, Brasil. + + + + +FIGURE 12. +Abundance of (a–b) + +Pachyopella flavida +( +Wiedemann, 1824 +) + +and (c–d) + +Poecilominettia octovittata +( +Williston, 1896 +) + +in relation to relative air humidity (%) and precipitation (mm) in the Fernando de Noronha Archipelago, Pernambuco, Brazil. + + + + +Remarks. + +Poecilominettia octovittata + +is easily recognized by the patterns of stripes and spots on the head and body, and by the wing pattern of being darkened anteriorly, on the crossveins, and in the distal part of vein M +1 +, as well as the distinctive characteristics of the male genitalia (particularly the shape and structure of the surstylus and the phallus) and female terminalia (particularly the paired darkened patches on sternite 8). The species was described from +Saint Vincent +(West Indies), and later recorded from +Brazil +( +Gaimari & Silva, 2020 +). This is by far the most abundant lauxaniid in Fernando de Noronha, after nine months we collected 16,548 specimens in all traps, representing 87,6% of the specimens collected, with +13,416 in +Capim-açu, +2,769 in +Sancho and +363 in +other localities of the archipelago. It is worth noting that +Fig. 7B +is a living specimen photographed in Olho d’Água das +Flores +, in the state of Alagoas, being the first record of this species from the state (https://www.inaturalist.org/ observations/140062464). + + +Discussion. +The genus + +Poecilominettia + +comprises 81 described species, all found in the Neotropics except for three strictly Nearctic species, and with 56 of the Neotropical species found in +Panama +, and 10 recorded in +Brazil +( +Gaimari & Silva 2020 +). The revisionary work of +Broadhead (1989) +represents the most comprehensive treatment of the genus, with a key to species and illustrations for most species. +Broadhead (1989) +also discusses the biology of the species of this genus, finding that adults feed on fungal hyphae and fungal spores, based on examination of food particles found in the pseudo-tracheal canals and gut contents. She also discusses the idea that although their larval habits are not known, they are generally thought to develop within the tissues of fall, decaying leaves. Some species of the genus are also known to feed in decaying matter in birds’ nests ( +Miller & Foote 1975 +, +Miller 1977 +). Further, +Miller & Foote (1976) +provide details of the morphology of immatures for a species of + +Poecilominettia + +. + + + + \ No newline at end of file diff --git a/data/03/B0/DE/03B0DE22FFA432348FDCFA40FDE2A6CC.xml b/data/03/B0/DE/03B0DE22FFA432348FDCFA40FDE2A6CC.xml index 28d2b655e18..60ac93d0021 100644 --- a/data/03/B0/DE/03B0DE22FFA432348FDCFA40FDE2A6CC.xml +++ b/data/03/B0/DE/03B0DE22FFA432348FDCFA40FDE2A6CC.xml @@ -1,73 +1,75 @@ - - - -Insect (Hexapoda) diversity in the oceanic archipelago of Fernando de Noronha, Brazil: Lauxaniidae (Diptera) + + + +Insect (Hexapoda) diversity in the oceanic archipelago of Fernando de Noronha, Brazil: Lauxaniidae (Diptera) - - -Author + + +Author -Soares, Matheus M. M. -Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil +Soares, Matheus M. M. +Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil - - -Author + + +Author -Gaimari, Stephen D. -Dipterists Society, P. O. Box 231113, Sacramento, California 95823, USA +Gaimari, Stephen D. +Dipterists Society, P. O. Box 231113, Sacramento, California 95823, USA - - -Author + + +Author -Corrêa-Neto, José De J. -Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil +Corrêa-Neto, José De J. +Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil - - -Author + + +Author -Limeira-De-Oliveira, Francisco -Universidade Estadual do Maranhão, Laboratório de Estudos dos Invertebrados, Praça Duque de Caixas, Morro do Alecrim, s / n, 65604 - 380 Caxias, MA, Brazil +Limeira-De-Oliveira, Francisco +Universidade Estadual do Maranhão, Laboratório de Estudos dos Invertebrados, Praça Duque de Caixas, Morro do Alecrim, s / n, 65604 - 380 Caxias, MA, Brazil - - -Author + + +Author -Rafael, José A. -Instituto Nacional de Pesquisas da Amazônia, INPA, Coordenação de Biodiversidade, Manaus, Amazonas, Brazil +Rafael, José A. +Instituto Nacional de Pesquisas da Amazônia, INPA, Coordenação de Biodiversidade, Manaus, Amazonas, Brazil -text - - -Zootaxa +text + + +Zootaxa - -2024 - -2024-11-07 + +2024 + +2024-11-07 - -5537 + +5537 - -1 + +1 - -1 -23 + +1 +23 - -http://dx.doi.org/10.11646/zootaxa.5537.1.1 + +http://dx.doi.org/10.11646/zootaxa.5537.1.1 -journal article -10.11646/zootaxa.5537.1.1 -1175-5326 -14239256 -91F512AE-5BAC-458D-992D-DF4EAF055183 +journal article +305806 +10.11646/zootaxa.5537.1.1 +5944fc67-97a0-4a74-87b1-12f5c691bd58 +1175-5326 +14239256 +91F512AE-5BAC-458D-992D-DF4EAF055183 @@ -83,7 +85,7 @@ Soares & Gaimari - + ( Figs. 5 @@ -91,6 +93,8 @@ Soares & Gaimari 6 ) + + Poecilominettia @@ -266,7 +270,7 @@ Similar to male ( Type material examined. - + Holotype ♂ ( @@ -276,14 +280,14 @@ Similar to male ( , PE [ Pernambuco ], -Fernando -de / -Noronha +Fernando de / Noronha , 3°51′17″S –/ 32°26′26″W -, Capim-Açu”, “ +, +Capim-Açu +”, “ 11–27.xi.2019 @@ -291,12 +295,14 @@ de / Malaise trap / G[grande], J.A. Rafael -, F. Limeira- / de-Oliveira, +, +F. Limeira- / de-Oliveira +, L.C. Castro ” . - + PARATYPES . @@ -318,12 +324,12 @@ de / , large Malaise trap -, J.A. -Rafael -, F. -Limeira-de-Oliveira -& L.C. -Castro +, +J.A. Rafael +, +F. Limeira-de-Oliveira +& +L.C. Castro ( 8 ♀ , @@ -361,13 +367,13 @@ same data, except , 5 ♀ , -AMNH +AMNH ; 1 ♂ , 8 ♀ , -MNRJ +MNRJ ) ; @@ -381,13 +387,13 @@ same data, except , 5 ♀ , -CSCA +CSCA ; 1 ♂ , 13 ♀ , -MZUSP +MZUSP ) ; @@ -399,11 +405,11 @@ same data, except ( 10 ♀ , -AMNH +AMNH ; 10 ♀ , -CSCA +CSCA ; 10 ♀ , @@ -411,21 +417,21 @@ same data, except ; 19 ♀ , -INPA +INPA ; 10 ♀ , -MNRJ +MNRJ ; 10 ♀ , -MZUSP +MZUSP ; 1 ♂ , 5 ♀ , -USNM +USNM ) ; @@ -434,14 +440,12 @@ same data, except 11–27.xi.2019 -[same data as -holotype -] ( +[same data as holotype] ( 5 ♂ , 2 dissected, 3 ♀ , 2 dissected, -INPA +INPA ) ; @@ -453,11 +457,11 @@ same data, except ( 17 ♀ , -INPA +INPA ) ; - + Sancho , 03º51′17″S @@ -478,7 +482,7 @@ same data, except ( 21 ♀ , -INPA +INPA ; 5 ♀ , @@ -486,7 +490,7 @@ same data, except ) ; - + same data, except 24.vi–8.vii.2019 @@ -496,15 +500,15 @@ same data, except ( 5 ♀ , -AMNH +AMNH ; 3 ♀ , -CSCA +CSCA ; 3 ♀ , -USNM +USNM ; 3 ♀ , @@ -520,7 +524,7 @@ same data, except ( 1 ♀ , -INPA +INPA ) ; @@ -532,15 +536,15 @@ same data, except ( 4 ♀ , -AMNH +AMNH ; 5 ♀ , -CSCA +CSCA ; 4 ♀ , -USNM +USNM ; 4 ♀ , @@ -556,16 +560,16 @@ same data, except ( 19 ♀ , -INPA +INPA ; 5 ♀ , -USNM +USNM ) ; - -Trilha Sancho + +Trilha Sancho , 3°51′30″S – @@ -578,24 +582,24 @@ same data, except Malaise trap , J.A. Rafael -, F. -Limeira-de-Oliveira +, +F. Limeira-de-Oliveira & D.M.M. Mendes [=same data as holotype] ( 27 ♀ , -INPA +INPA ) ; - + same data, except armadilha luminosa [ light trap ] ( 1 ♀ , -INPA +INPA ) ; @@ -607,25 +611,25 @@ same data, except ( 1 ♂ , dissected, -INPA +INPA ) ; - + same data, except 24.vi–08.vii.2019 -, J.A. -Rafael -, F. -Limeira-de-Oliveira +, +J.A. Rafael +, +F. Limeira-de-Oliveira & L.C. Castro ( 1 ♀ , dissected, -INPA +INPA ) ; @@ -637,25 +641,25 @@ same data, except ( 1 ♂ , dissected, -INPA +INPA ) ; - + same data, except 11–27.ii.2020 -, J.A. -Rafael -, F. -Limeira-de-Oliveira +, +J.A. Rafael +, +F. Limeira-de-Oliveira & P.C. Grossi ( 2 ♀ , -INPA +INPA ) . diff --git a/data/03/B0/DE/03B0DE22FFAC32388FDCFA11FA12A4EE.xml b/data/03/B0/DE/03B0DE22FFAC32388FDCFA11FA12A4EE.xml new file mode 100644 index 00000000000..d5d5301b48f --- /dev/null +++ b/data/03/B0/DE/03B0DE22FFAC32388FDCFA11FA12A4EE.xml @@ -0,0 +1,226 @@ + + + +Insect (Hexapoda) diversity in the oceanic archipelago of Fernando de Noronha, Brazil: Lauxaniidae (Diptera) + + + +Author + +Soares, Matheus M. M. +Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil + + + +Author + +Gaimari, Stephen D. +Dipterists Society, P. O. Box 231113, Sacramento, California 95823, USA + + + +Author + +Corrêa-Neto, José De J. +Graduate Program in Entomology (PPG-Ent), Instituto Nacional de Pesquisas da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis, CEP 69067 - 375, Manaus, Amazonas, Brazil + + + +Author + +Limeira-De-Oliveira, Francisco +Universidade Estadual do Maranhão, Laboratório de Estudos dos Invertebrados, Praça Duque de Caixas, Morro do Alecrim, s / n, 65604 - 380 Caxias, MA, Brazil + + + +Author + +Rafael, José A. +Instituto Nacional de Pesquisas da Amazônia, INPA, Coordenação de Biodiversidade, Manaus, Amazonas, Brazil + +text + + +Zootaxa + + +2024 + +2024-11-07 + + +5537 + + +1 + + +1 +23 + + + + +http://dx.doi.org/10.11646/zootaxa.5537.1.1 + +journal article +305806 +10.11646/zootaxa.5537.1.1 +5944fc67-97a0-4a74-87b1-12f5c691bd58 +1175-5326 +14239256 +91F512AE-5BAC-458D-992D-DF4EAF055183 + + + + + + +Key to +Lauxaniidae +from the oceanic Archipelago of Fernando de Noronha + + + + + + + + +1 +Anterior fronto-orbital seta reclinate ( +Figs. 5C, D +, +7C, D +); ocellar triangle not strongly contrasting with remainder of frons ( +Figs. 5E +, +7E +); postsutural intra-alar seta present ( +Fig. 5E +); 2 katepisternal setae ( +Figs. 5A +, +7A +); epandrium with dorsal rectangular brown spot ( +Figs. 5G +, +6A +, +7G +, +8A +)....................................................................... +2 + + + + +- Anterior fronto-orbital seta inclinate ( +Figs. 1C +, +3C +); ocellar triangle surrounded by dark brown patch contrasting with otherwise orange frons ( +Figs. 1E +, +3C +); postsutural intra-alar seta absent ( +Figs. 1E +, +3E +); 1 katepisternal seta ( +Figs. 1A +, +3B +); epandrium homogeneously yellow to pale yellow ( +Figs. 1G +, +2A–C +, +3G +, +4A, B +)............................................. +3 + + + + + + +2 +Face yellow with an oval dark brown spot at middle of lower margin ( +Fig. 7C +); thorax with dorsocentral and supra-alar brown stripes ( +Fig. 7E +); anterior margin of wing dark brown ( +Fig. 7F +).............. + +Poecilominettia octovittata +( +Williston, 1896 +) + + + + + +- +Face homogeneously brownish ( +Fig. 5C +); thorax homogenously brownish ( +Fig. 5E +); wing wholly hyaline ( +Fig. 5F +)................................................................... + +Poecilominettia erebus +Soares & Gaimari + +, + +sp. nov. + + + + + + + +3 +First-flagellomere elongated, about 8X longer than high ( +Fig. 3A, B, D +); face yellow with median rounded dark brown spot ( +Fig. 3C +); surstylus elongated, about 4X longer than wide, somewhat finger-shaped ( +Fig. 4A, B +)........................................................................................... + +Pachyopella flavida +( +Wiedemann, 1824 +) + + + + + +- +First flagellomere short, about 2.5X longer than high ( +Fig. 1D +); face homogeneously yellow ( +Fig. 1C +); surstylus somewhat oval, 1.5X longer than wide at widest point, with pointed apex ( +Fig. 2A–C +)... + +Camptoprosopella equatorialis +Shewell, 1939 + + + + + + + \ No newline at end of file diff --git a/data/5E/0B/9C/5E0B9CDA5EBB5422A4E7C3B6A82F2FC7.xml b/data/5E/0B/9C/5E0B9CDA5EBB5422A4E7C3B6A82F2FC7.xml new file mode 100644 index 00000000000..99337988652 --- /dev/null +++ b/data/5E/0B/9C/5E0B9CDA5EBB5422A4E7C3B6A82F2FC7.xml @@ -0,0 +1,520 @@ + + + +The new genus Purpurata (Lepidoptera, Crambidae, Spilomelinae), with descriptions of two new species from China + + + +Author + +Xue, Xin-Lei +https://orcid.org/0009-0000-5697-0872 +College of Plant Protection, Southwest University, Chongqing 400715, China & Yibin Academy of Southwest University, Yibin, Sichuan 644000, China + + + +Author + +Lu, Xiao-Qiang +https://orcid.org/0009-0009-3696-7778 +College of Plant Protection, Southwest University, Chongqing 400715, China & Guangyuan Center for Disease Control and Prevention, Guangyuan, Sichuan 628040, China + + + +Author + +Du, Xi-Cui +0000-0002-7796-7303 +College of Plant Protection, Southwest University, Chongqing 400715, China & Yibin Academy of Southwest University, Yibin, Sichuan 644000, China + +text + + +ZooKeys + + +2024 + +2024-12-02 + + +1219 + + +175 +194 + + + +journal article +10.3897/zookeys.1219.131102 +A7CEF41B-6FE9-48DF-87BD-7474CD8AEAB7 + + + + + +Purpurata lurida + +sp. nov. + + + + +Figs 10 +, +17 +, +17 a – c +, +18 + + + + +Diagnosis. + + +This species is similar to + +P. obfuscalis + +in appearance and genitalia, but can be distinguished by the body color paler than the latter (Fig. +10 +); uncus short and broad triangular (Fig. +17 a +); valva shorter and broader than the latter, with distal 1 / 3 narrowed very gradually and rounded apically (Fig. +17 +). In + +P. obfuscalis + +, the uncus is semicircular (Fig. +12 a +); the distal 1 / 3 of the valva is narrowed gradually and pointed apically (Fig. +12 +). + + + + +Type material. + + + + +Holotype + +: +♂ +pinned, with genitalia in a separate slide. + +China + +• + +Guizhou Prov. + +, +Kuankuoshui Nature Reserve +, alt. + +1500 m + +, + +15 August 2010 + +, +Xi-Cui Du +leg., genitalia slide number: XLJ 14084 + +. + + +Paratypes + +: pinned, some with genitalia in separate slides. + +China + + +• + + +Hubei Prov. + +, +2 ♂♂ +, +Hejiaping Town +, +Changyang County +, alt. + +800 m + +, + +18 June 2018 + +, +Xiao-Qiang Lu +& +Xi-Cui Du +leg. + +• + +2 ♂♂ +, +Xingdou Mountain +, +Maoba Town +, +Enshi +, alt. + +780 m + +, + +30 July 2012 + +, +Jun Zhang +& +Xiao-Bing Fu +leg. + +• + + +Chongqing +Municipality + +, +2 ♂♂ +, +Jinyun Mountain +, alt. + +550 m + +, 22, + +July 2010 + +, +Li Kang +& Xing-Fu +Fu +leg., genitalia slide number: XLJ 14103 + +• + +2 ♂♂ +, +1 ♀ +, +Jinyun Mountain +, + +29 July 2010 + +, +Xi-Cui Du +& +Chao-Wei Bi +leg., genitalia slide number: XLJ 14104 +♀ + +• + +1 ♂ +, +Jinyun Mountain +, + +30 July 2012 + +, +Li-Jun Xu +& Jian-Bo +Cao +leg. + +• + +2 ♂♂ +, +Jinyin Mountain +, +Qianjiang District +, alt. + +1100 m + +, + +26 July 2012 + +, +Li-Jun Xu +& +Jun-Zhang +leg. + +• + + +Yunnan Prov. + +, +1 ♂ +, +Daxichang village +, +Malipo County +, alt. + +1465 m + +, + +5 June 2015 + +, +Man-Fei Tao +leg. + +• + +2 ♀♀ +, +Xishuangbanna Tropical Botanical Garden +, alt. + +659 m + +, + +28 May 2015 + +, +Man-Fei Tao +leg., genitalia slide number: LXQ 18316 + +• + + +Hainan Prov. + +, +1 ♂ +, +Diaoluo Mountain +vocational village, alt. + +500 m + +, + +23 May 2014 + +, +Li-Jun Xu +& +Dan Xu +leg. + +• + + +Zhejiang Prov. + +, +3 ♂♂ +, +Tianmu Mountain +, alt. + +400 m + +, 24, 25, + +30 July 2011 + +, +Xi-Cui Du +& +Xiao-Bing Fu +leg. + + + + + + + +Genitalia of + +Purpurata +species + +12, 13 + +P. obfuscalis + +12 +male, genitalia slide no. XXL 23275 +13 +female, genitalia slide no. XLJ 14221 +14 + +P. iopasalis + +, male, genitalia slide no. XXL 23277 +15, 16 + +P. directa + +sp. nov. +15 +male, holotype, genitalia slide no. LXQ 18315 +16 +female, paratype, genitalia slide no. XLJ 14147 +12 a, 14 a, 15 a +gnathos +12 b, 14 b, 15 b +fibula +12 c, 14 c, 15 c +cornuti and posterior protrusion of phallus. Scale bars: 1.0 mm ( +12–16 +); 0.1 mm ( +12 a – c, 14 a – c, 15 a – c +). + + + + + + + +Genitalia of + +Purpurata +species + +17, 18 + +P. lurida + +sp. nov. +17 +male, holotype, genitalia slide no. XLJ 14084 +18 +female, paratype, genitalia slide no. LXQ 18316 +19, 20 + +P. plagiatalis + +19 +male, genitalia slide no. HGQ 13237 +20 +female, genitalia slide no. XLJ 14204 +17 a, 19 a +gnathos +17 b, 19 b +fibula +17 c, 19 c +cornuti and posterior protrusion of phallus. Scale bars: 1.0 mm ( +17–20 +); 0.1 mm ( +17 a – c, 19 a – c +). + + + + + +Description. + + +Habitus +(Fig. +10 +). Forewing length 11.0–13.5 mm, wingspan 24.0–30.0 mm. Frons and vertex yellowish brown. Labial palpus with 1 +st +segment yellowish white ventrally, the remainder yellowish brown. Maxillary palpus brown. Antenna yellowish brown, with ventral cilia ~ 1 / +3 in +length of diameter of flagellomere in male. Patagium and tegula yellow. Thorax yellow dorsally, white ventrally. Legs yellowish white, distal end of front tibia black. Wings pale yellow, with purple-brown lines and patches. Forewing with three small spots at base, another spot near basal dorsum; antemedial line slightly wavy, accompanied by a large elliptical pale patch inside; orbicular stigma a dark brown dot; discoidal stigma reniform, yellow centrally; postmedial line slightly obliquely inward from costa, dentate and excurved between M +2 +and CuA +2 +, then incurved to discoidal stigma below and sinuous to inner margin; an irregular large patch near apex beyond anterior postmedial line, another misty patch near tornus below CuA +2 +; a line of spots along marginal line. Hindwing with discoidal stigma a short oblique stripe; postmedial line same as forewing before CuA +2 +, not apparent afterwards; an irregular large patch near apex beyond anterior postmedial line; a band below discoidal stigma, accompanied by another irregular misty wide band extended to tornus. Cilia of fore and hind wings yellowish white. Abdomen yellow dorsally, white ventrally; 1 +st +and 2 +nd +tergites with pale black spots laterally and 7 +th +tergites black posteriorly in male. + + +Male genitalia +(Fig. +17 +). Uncus short and broad triangular. Gnathos vestigial to a narrow band and bearing a few short setae (Fig. +17 a +). Valva broad tongue-shaped, with distal 1 / 3 narrowed very gradually, and rounded apically; costa arched medially and bearing a cluster of long setae; fibula a short lamina, setose basally (Fig. +17 b +). Saccus oval. Juxta a broad plate. Phallus posteriorly with a finger-like protruding sclerite, with a thick, needle-like cornutus and a brush-like cornutus composed of a spine cluster (Fig. +17 c +). + + +Female genitalia +(Fig. +18 +). Apophyses anteriores ~ 2 × length of apophyses posteriores. Antrum broad, ductus seminalis originating from antrum. Ductus bursae ~ 2 × length of corpus bursae. Corpus bursae nearly oval, with a round signum. + + + + +Distribution. + + +China +( +Chongqing +, +Guizhou +, +Yunnan +, +Hainan +, +Hubei +, +Zhejiang +). + + + + +Etymology. + + +The species name +lurida +is derived from the Latin word +luridus +, an adjective, meaning pale yellow, indicating the pale wing color. + + + + \ No newline at end of file diff --git a/data/7E/0D/92/7E0D929D4AEA576BB1D944A98979533A.xml b/data/7E/0D/92/7E0D929D4AEA576BB1D944A98979533A.xml new file mode 100644 index 00000000000..0d534348a0b --- /dev/null +++ b/data/7E/0D/92/7E0D929D4AEA576BB1D944A98979533A.xml @@ -0,0 +1,481 @@ + + + +Rubus tingzhouensis (Rosaceae), a new species from Fujian, China + + + +Author + +Chen, Ming +https://orcid.org/0009-0001-9894-8339 +Fujian Maternity and Child Health Hospital, College of Clinical Medicine for Obstetrics & Gynecology and Pediatrics, Fujian Medical University, Fuzhou 350001, China + + + +Author + +Lin, Gui-Chan +https://orcid.org/0009-0001-7911-358X +College of Pharmacy, Fujian University of Traditional Chinese Medicine, Fuzhou 350122, China & Fujian Health College, Fuzhou, Fujian 350101, China + + + +Author + +Wang, Tao +https://orcid.org/0009-0001-5749-4174 +Xiamen Gaoxin School, Xiamen 361000, China + + + +Author + +Zhuang, Yi-Xue +0000-0002-0295-9405 +College of Pharmacy, Fujian University of Traditional Chinese Medicine, Fuzhou 350122, China + + + +Author + +Yao, Yi-Xin +0000-0001-6948-6975 +State Key Laboratory of Quality Research in Chinese Medicine Institute of Chinese Medical Sciences, University of Macau, Macau 519000, China + + + +Author + +Yang, Cheng-Zi +0000-0002-5527-7934 +College of Pharmacy, Fujian University of Traditional Chinese Medicine, Fuzhou 350122, China + + + +Author + +Qin, Yuan +0000-0003-4713-6151 +Fujian Provincial Key Laboratory of Haixia Applied Plant Systems Biology, Center for Genomics and Biotechnology, College of Life Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China + + + +Author + +Lin, Yan-Xiang +https://orcid.org/0009-0008-6971-792X +College of Pharmacy, Fujian University of Traditional Chinese Medicine, Fuzhou 350122, China + + + +Author + +An, Chang +0000-0002-7657-7936 +Fujian Provincial Key Laboratory of Haixia Applied Plant Systems Biology, Center for Genomics and Biotechnology, College of Life Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China + +text + + +PhytoKeys + + +2024 + +2024-12-02 + + +249 + + +251 +267 + + + +journal article +10.3897/phytokeys.249.138951 + + + + + +Rubus tingzhouensis +C. An & G. C. Lin + +sp. nov. + + + + +Figs 3 +, +4 + + + + + +Type +. + + + + +China +• +Fujian +: +Longyan City +, +Changting County +, +Xuancheng Town +, +Xiashe Village +, + +25 ° 24 ' 06 " N +, +116 ° 22 ' 34 " E + +, forests on mountain slopes, alt. ca. + +351 m + +, + +18 April 2024 + +, + +C +. An & +G +. +C +. Lin. 240418 + +( +holotype +: + +IBSC + +[barcode 1021457!]; +isotypes +: + +PE + +[barcode 02468523!, 02468524!]) + +. + + + + + + + +Rubus tingzhouensis +C. An & G. C. Lin +A + +plant in natural habitat +B +leaf (adaxial surface) +C +leaf (abaxial surface) +D, E, F +flower and inflorescence +G +androecium +H +stamen +I +stigma +J +petal +K +stipule. + + + + + +Diagnosis. + + +This species is similar to + +R. swinhoei + +in its growth habit, with ovate to oblong-lanceolate leaf blades and botryoid inflorescences that may be terminal or axillary. However, + +R. tingzhouensis + +can be distinguished by its dense indumentum of long, reddish-purple stipitate glands, soft setae and light yellow short trichomes on the plant surface. It also has scattered epidermal prickles, adding to its distinct appearance. Additionally, it has deeply laciniate stipules measuring +1–1.5 cm +in length, which are significantly more divided than those of related species, making them a key distinguishing feature. + + + + + + +Illustration of + +Rubus tingzhouensis +C. An & G. C. Lin +A + +habit +B +petal +C +longitudinal section of flower +D +free lobes of calyx +E +stamen +F +pistil +G +flower +H +stipule +I +bract +J +margin and trichome of leaf +K +trichome of abaxial leaf +L +section of leafed stem. Drawn by Yunxiao Liu. + + + + + +Description. + + + +Vines, lianas and shrubs. +Stems + +cylindrical, greyish-brown, with dense, reddish-purple long glandular hairs, soft bristles, short yellowish hairs and sparse prickles, apically rooting. +Leaves +simple; blades ovate to oblong-lanceolate, herbaceous, 8–16 × +3.5–6 cm +, apex acuminate to acute, base cordate; adaxially flat, hirsutullous with long, purple stipitate glands along veins, abaxially densely yellowish-brown tomentose and pubescent, with long soft hairs along mid-ribs; principal veins sparsely retrorsely aculeolate, margin unevenly serrate to doubly serrate, apex acuminate to acute, lateral veins 9 to 10 pairs; petiole +1.5–2 cm +, with dense, long, purplish-red glandular hairs and soft bristles; stipules caducous, free, pinnatipartite, lobes narrowly elliptic or lanceolate, densely covered with long glandular hairs and tomentose-villous, +1–1.5 cm +. +Inflorescences +terminal or axillary, short botryoid, 5–10 flowered; involucral bracts +6–9 mm +, lobed, lobes linear or lanceolate, villous, rachis and pedicels with dense reddish-purple long glandular hairs and soft bristles; peduncle +5–10 cm +, pedicels +1.5–2 cm +; +Flowers +2–2.5 cm +. Sepals ovate-lanceolate, +5–6 mm +, apex acuminate to caudate, outer sepals usually 2 - or 3 - laciniate; abaxially densely greyish-white pubescent, purplish-red long glandular hairs and soft bristles, adaxially densely greyish-white pubescent. Petals white, broadly ovate to oblong, 5–6 × +4–5 mm +, base barely clawed, slightly shorter than sepals. Stamens many; filaments linear, lower part slightly broader; anthers with few long hairs. Carpels many, style longer than stamens, glabrous. + + + + +Phenology. + +Flowering in March to May, fruits have not been seen yet. + + + +Etymology. + + +The specific epithet + +‘ +tingzhouensis + +’ refers to the ancient region Tingzhou (汀州) in south-western +Fujian +, +China +, where this species was discovered. The Chinese name, “ 红毛木莓 ” (hong mao mu mei), reflects the plant’s dense covering of reddish-purple long hairs. + + + + +Distribution and ecology. + + +Currently, this species is only found in Changting County and Shanghang County, +Fujian Province +, +China +(Fig. +5 +). It is sporadically distributed in the understorey of the primary forest in mountain valleys at an altitude of + +300– +400 m. + +The habitat features a thick layer of dead branches and leaf litter, as well as a substantial amount of humus, supporting vigorous growth of herbaceous, shrubby and woody plants. Associated species include + +Vernicia montana +Lour. + +and + +Pterocarya stenoptera + +C +. D. +C +. in the tree layer; + +R. corchorifolius + +L +. f., + +Buddleja lindleyana +Fortune + +, + +Diplospora dubia +(Lindl.) Masam. + +, + +Callicarpa kochiana +Makino + +, + +Ilex pubescens +Hook. & Arn. + +, + +Itea omeiensis +C. K. Schneid. + +, + +C. formosana +R. Br. + +, + +Clerodendrum cyrtophyllum +Turcz. + +, + +Loropetalum chinense +(R. Br.) Oliv. + +, + +Mallotus apelta +(Lour.) Müll. Arg. + +, + +Trema tomentosa +(Roxb.) H. Hara + +, + +Phyllanthus glaucus +Wall. ex Müll. Arg. + +, + +Melastoma malabathricum + +L +. and + +R. reflexus +Ker Gawl. + +in the shrub layer; + +Lysimachia alfredii +Hance + +, + +Senecio scandens +Buch. + +- Ham. ex D. Don, + +Blechnopsis orientalis + +( +L +.) +C +. Presl and + +Dicranopteris pedata +(Houtt.) Nakaike + +in the herbaceous layer. + + + + + + +Geographical distribution of + +R. tingzhouensis + +in China (red star). + + + + + +Conservation assessment. + + + +R. tingzhouensis + +is documented in limited populations distributed within sparse forests on mountain slopes in Shanghang, Changting and Wuping County, +Fujian Province +or thrives along stream margins and under mixed forests. Notably, one population in Changting County is adjacent to a scenic locale, heightening its susceptibility to considerable anthropogenic disturbance. Furthermore, most of these populations are located outside designated conservation zones, making them vulnerable to ongoing exploitation of woodland resources by local residents. Thus, this newly-recognised species is assigned a preliminary status of Vulnerable ( + +VU + +D 2) according to the +IUCN +Red List Categories and Criteria ( +Standards and Group 2006 +; +Betts et al. 2020 +), reflecting a population with a severely limited occupancy range (typically less than +20 km +2 +) or few locations (typically five or fewer). + + + + \ No newline at end of file diff --git a/data/BB/7A/03/BB7A032FC1246B1BFF4D28CFFED5F876.xml b/data/BB/7A/03/BB7A032FC1246B1BFF4D28CFFED5F876.xml new file mode 100644 index 00000000000..d6e307e1b10 --- /dev/null +++ b/data/BB/7A/03/BB7A032FC1246B1BFF4D28CFFED5F876.xml @@ -0,0 +1,358 @@ + + + +First records of the treehopper Trichaetipyga infantilis (Ball, 1937) in Ecuador with the description of its nymph and notes of natural history, and morphology + + + +Author + +Montalvo-Salazar, Jorge L. +0009-0003-1221-0304 +Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Instituto de Biodiversidad Tropical IBIOTROP, Laboratorio de Zoología Terrestre, Museo de Zoología, Quito 170901, Ecuador. +jorgemontalvo2000@gmail.com + + + +Author + +Rueda-Rodríguez, María P. +0009-0002-8190-7239 +Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Instituto de Biodiversidad Tropical IBIOTROP, Laboratorio de Zoología Terrestre, Museo de Zoología, Quito 170901, Ecuador. & Tandayapa Cloud Forest Station, Colegio de Ciencias Biológicas y Ambientales, Universidad San Francisco de Quito USFQ, P. O. Box 17 - 1200 - 841, Quito, Ecuador. +mariapazrr03@gmail.com + + + +Author + +Torres Arizaga, David M. +0009-0004-9485-9963 +Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Instituto de Biodiversidad Tropical IBIOTROP, Laboratorio de Zoología Terrestre, Museo de Zoología, Quito 170901, Ecuador. +davidtorresarizaga@gmail.com + +text + + +Zootaxa + + +2024 + +2024-11-25 + + +5541 + + +1 + + +89 +94 + + + + +http://dx.doi.org/10.11646/zootaxa.5541.1.6 + +journal article +305807 +10.11646/zootaxa.5541.1.6 +1dd6b56f-af7d-4a68-a743-78d20ddde04b +1175-5326 +14240846 + + + + + + +Trichaetipyga infantilis +( +Ball, 1937 +) + + + + + + +( +Figures 1–3 +) + + + + +Distribution: +Mexico +, +Costa Rica +, +Venezuela +, +Colombia +, and +Ecuador +( +New Record +) ( +Fig. 4 +). We report here the southern-most records for the species, from the northern Amazonian lowlands and foothills of northwestern Ecuadorian Andes, around +620 km +and +760 km +. The species frequently inhabits lowlands below +700 m +of elevation, however, Flórez-V. +et al. +(2015) have recorded the species from> +1800 m +of elevation in the Colombian Andes. + + + + + +Material examined: One nymph, +two males +and +three females +( +ZSFQ-i7928 +, +9059,11131 +, +17972 +) from +Nueva Loja +( +Sucumbíos +) [ +0.102022ºN +76.887272ºW +, + +300 m + +of elevation]; + + +one male +and +one female +(ZSFQ-i17711, 17712) from +Santo Domingo +( +Santo Domingo de los Tsáchilas +) [ +0.302838ºS +79.155478ºW +, + +500 m + +of elevation] + +. + + + + +FIGURE 1. + +Trichaetipyga infantilis + +habitus. A. male in lateral view. Female: B. frontal view, C. lateral view, D. dorsal view. Nymph: E. frontal view, F. lateral view. + + + + +FIGURE 2. + +Trichaetipyga infantilis + +female. Abdomen: A. lateral view, B. ventral view. C. gonoplac in lateral view. First valvula: D. lateral view, E. close-up of anterior portion. Second valvula, F. lateral view, G. close-up of anterior portion, H. closeup of apex. + + + + +Female genitalia description. +Pygofer surface smooth and almost glabrous except for small setae on ventral surface around aperture ( +Figs 2A, B +). Gonoplac apical half broad, gradually swollen, apex shortly rounded; dorsal margin straight, ventral margin more sclerotized, convex, and with macrosetae ( +Fig. 2C +). First valvula curved upward, narrowing towards apex; ramus extending almost throughout valvula, dorsal sculptured area restricted in distal half with dense fine oblique integumental lines gradually increasing from the dorsal margin to ramus, ventral sculpted area restricted at apex with subperpendicular irregular integumental lines widely separated ( +Fig. 2D +); apex acute and dorsal margin straight ( +Fig. 2E +). Second valvula more sclerotized than first, curved upward, tapering apically, apex acutely rounded ( +Fig. 2F +); apical fifth of dorsal margin irregularly serrate ( +Fig. 2G +), apex finely serrate ( +Fig. 2H +). + + +Nymph description +( +Figs 1E, F +, +3A +). Coloration. Pale yellowish with reddish bands or patches on the head, thorax, wing pad, legs, and abdomen. +Overall body. +Laterally compressed with long scoli with stalked chalazae on every segment; dense chalazae on head, between scoli, legs, and ventral margins of thoracic and abdominal tergites. +Head. +One pair of scoli longer than head height and obliquely directed forward; covered with large and simple chalazae. +Prothorax +. Pre- and postmetopidum each with pair of dorsal scoli, longer than head scoli and obliquely directed forward; pronotum with triangular posterior apex slightly curved upwards, hidden between mesothoracic scoli in lateral view and not surpassing mesonotum. +Mesothorax +. One pair of dorsal scoli obliquely directed forwards and as long as postmetopidum scoli; enlarged chalazae setae on ventral margin of forewing pad. +Metathorax +. Dorsally with paired scoli. +Legs +. Chalazae present from coxa to tarsus, enlarged on tibiae. +Abdomen +. Terga III–VII laterally glabrous with dorsal, subparallel scoli of subequal lengths, 8 or more x their basal widths; tergum IX scoli half-length of anterior tergal scoli and located apically; terga III–VIII ventrolateral margins each with 2 pairs of enlarged chalazae and dense small chalazae on ventral margin; segment IX in lateral view equal to combined lengths of segments V–VII, dorsal margin with 2 parallel rows of enlarged chalazae. + + + +FIGURE 3. + +Trichaetipyga infantilis + +in its natural environment. A. nymph on stalk base of + +Paspalum +sp. + +, B. adult on spike of + +Paspalum +sp. + +, C. adult on a leaf of + +Ipomoea +sp. + + + + +Few +Ceresini +nymphs have been described, including only some species from the genera + +Stictocephala + +, + +Spissistilus +, +Tortistilus +, +Stictolobus +, + +and + +Ceresa + +. The nymphs of those genera and + +Trichaetipyga infantilis + +have dorsal spine-like scoli and stalked chalazae on the head, pronotum, and abdomen ( + +Quisenberry +et al. +1978 + +; + +Grosso +et al. +2014 + +; +Wheeler & Rothschild 2020 +). + + +Natural history. +This species inhabits pastures at the borders of water bodies or forests and is abundant in highly disturbed areas. Adults and nymphs are mainly solitary although, on one occasion, +three adult +individuals shared the same plant but did not feed on it. The nymph stayed in the stalk base of their host plants while adults usually in the high portion of the plant ( +Figs 3A–C +). This behavior has been described in more detail for other Ceresine species, in which the emerged nymphs move to the base of the plant and gradually climb up while they grow and become more active ( + +Grosso +et al. +2014 + +; + +Khan +et al. +2022 + +). + + +Adults and the nymph were not attended by ants or other hymenopterans. We recorded + +T. infantilis + +on both dicots and monocots ( +Table 1 +), although it was more frequently found in + +Paspalum +sp. + +( +Poaceae +), which we suspect to be the main host plant because the nymphs have been only seen on that plant species and it was the most frequently seen plant association of the adults. +Wheeler & Rothschild (2020) +found the nymphs of another Ceresine, + +Stictolobus minutus +(Funkhouser) + +, are specialized on a single species of +Poaceae +. Other Ceresine species have been observed to feed on grasses but mostly are recorded as polyphagous on dicots and monocots (Flórez-V. +et al. +2015). + + + + +Remarks. +Kopp & Yonke (1979) +argued that + +Trichaetipyga infantilis + +varies in the relative development of the suprahumeral horns (see their Figs 43–48). The +syntypes +of this species ( +United States +National Museum of Natural History, available at http://n +2t +.net/ark:/65665/3a2cc0b1f-e932-4087-9157-d76bd425ce21), have short suprahumeral horns, with those of the female slightly more pronounced than the male. In contrast, in all our speicmens the suprahumeral horns are strongly developed and acute in females and males ( +Figs 1A–D +), as is the specimen illustrated by Flórez-V. +et al. +(2015) from +Colombia +(see Figs their 36A, 37D). Further studies might reveal that + +T. infantilis + +represents more than one cryptic species. + + + + \ No newline at end of file diff --git a/data/DC/BC/6F/DCBC6FA6131B5DB69B62E8E56D69D3FB.xml b/data/DC/BC/6F/DCBC6FA6131B5DB69B62E8E56D69D3FB.xml new file mode 100644 index 00000000000..31214df9402 --- /dev/null +++ b/data/DC/BC/6F/DCBC6FA6131B5DB69B62E8E56D69D3FB.xml @@ -0,0 +1,587 @@ + + + +The new genus Purpurata (Lepidoptera, Crambidae, Spilomelinae), with descriptions of two new species from China + + + +Author + +Xue, Xin-Lei +https://orcid.org/0009-0000-5697-0872 +College of Plant Protection, Southwest University, Chongqing 400715, China & Yibin Academy of Southwest University, Yibin, Sichuan 644000, China + + + +Author + +Lu, Xiao-Qiang +https://orcid.org/0009-0009-3696-7778 +College of Plant Protection, Southwest University, Chongqing 400715, China & Guangyuan Center for Disease Control and Prevention, Guangyuan, Sichuan 628040, China + + + +Author + +Du, Xi-Cui +0000-0002-7796-7303 +College of Plant Protection, Southwest University, Chongqing 400715, China & Yibin Academy of Southwest University, Yibin, Sichuan 644000, China + +text + + +ZooKeys + + +2024 + +2024-12-02 + + +1219 + + +175 +194 + + + +journal article +10.3897/zookeys.1219.131102 +A7CEF41B-6FE9-48DF-87BD-7474CD8AEAB7 + + + + + +Purpurata directa + +sp. nov. + + + + +Figs 2 +, +8 +, +9 +, +15 +, +15 a – c +, +16 + + + + + + + +Patania iopasalis + + +: + +Xu and Du 2016: 132 + +, figs 5–8. +Type +locality: +India +(misidentification). + + + + + + + +Diagnosis. + + +This species is very similar to + +P. iopasalis + +in appearance, but can be distinguished by ventral cilia ~ 1 / 3 length of flagellomere diameter in male, antemedial line of forewing relatively straight and slightly inclined outward, postmedial line of fore and hind wings smooth linear between M +2 +and CuA +2 +(Figs +8 +, +9 +); uncus semicircular, gnathos a transverse lamina, and valva broader than the latter in male genitalia (Fig. +15 +); a signum present in female genitalia (Fig. +16 +). In + +P. iopasalis + +, the ventral cilia are approximately equal in length of flagellomere diameter in male, the antemedial line is slightly wavy, the postmedial line of fore and hind wings punctiform between M +2 +and CuA +2 +(Fig. +7 +); the uncus is nearly triangular, and the gnathos is finger-like in the male genitalia (Fig. +14 a +); and the signum is absent in the female genitalia. + + + + +Type material. + + + + +Holotype + +: +♂ +pinned, with genitalia in a separate slide. + +China + +• + +Hunan Prov. + +, +Huping Mountain +, +Shimen County +, alt. + +350 m + +, + +6 June 2017 + +, +Jian-Yue Qiu +& +Hao Xu +leg., genitalia slide number: LXQ 18315 + +. + + +Paratypes + +: pinned, some with genitalia in the separate slides, respectively. + +China + + +• + +6 ♂♂ +, same data as holotype + +• + + +Hubei Prov. + +, +1 ♂ +, +Hejiaping Town +, +Changyang County +, alt. + +800 m + +, + +18 May 2018 + +, +Xian-Qiang Lu +& +Xi-Cui Du +leg. + +• + + +Chongqing +Municipality + +, +2 ♀♀ +, +Daheba +, +Jinfo Mountain +, alt. + +600 m + +, 14, + +17 July 2017 + +, +Shi-Man Bu +leg + +• + + +Sichuan Prov. + +, +2 ♂♂ +, +Emei Mountain +, alt. + +863 m + +, + +17 July 2011 + +, Jian-Bo +Cao +leg. + +• + + +Guizhou Prov. + +, +3 ♂♂ +, +Baishao +, +Kuankuoshui +, alt. + +800 m + +, + +10 August 2010 + +, +Xi-Cui Du +leg., genitalia slide number: HGQ 13018, HGQ 13019 + +• + +3 ♂♂ +, +1 ♀ +, +Wengang village +, +Libo County +, alt. + +1345 m + +, + +20 July 2015 + +, +Ji-Ping Wan +leg. + +• + + +Yunnan Prov. + +, +1 ♂ +, +Baihualing Village +, +Baoshan City +, alt. + +1520 m + +, + +13 July 2007 + +, +Dan-Dan Zhang +leg. + +• + +1 ♂ +, +Daxichang Village +, +Malipo County +, alt. + +1465 m + +, + +7 August 2007 + +, +Man-Fei Tao +leg. + +• + + +Hainan Prov. + +, +1 ♂ +, +Bawangling National Forest +Park, + +11 June 2010 + +, +Li Kang +leg., genitalia slide number: XLJ 14105 + +• + +3 ♂♂ +, +1 ♀ +, +Wuzhi Mountain +, alt. + +795 m + +, 18, 19, + +21 May 2014 + +, +Li-Jun Xu +& +Dan Xu +leg., genitalia slide number: XD 15024 +♂ +, XLJ 14146 +♂ +, XLJ 14147 +♀ + +• + +1 ♂ +, +Diaoluo Mountain +, alt. + +500 m + +, + +25 May 2014 + +, +Li-Jun Xu +& +Dan Xu +leg., genitalia slide number: XD 15049 + +• + + +Guangxi Zhuang +Autonomous Region + +, +3 ♂♂ +, +Nonggang National Nature Reserve +, +Longzhou County +, alt. + +188 m + +, 25, 27, + +30 July 2011 + +, +Gui-Qing He +leg., genitalia slide number: XLJ 14083 (being identified as + +Patania iopasalis + +by +Xu and Du (2016) +), XLJ 13199 + +• + +3 ♂♂ +, +Nonggang National Nature Reserve +, +Longzhou County +, alt. + +188 m + +, + +2 August 2011 + +, +Gui-Qing He +leg., genitalia slide number: XLJ 13164 + +• + +5 ♂♂ +, +Mulun Nature Reserve +, alt. + +288 m + +, + +21 July 2015 + +, +Dan Xu +leg. + +• + + +Guangdong Prov. + +, +5 ♂♂ +, +Renhua County +, +Danxia Mountain +, alt. + +408 m + +, + +31 May 2018 + +, +Feng-Xia He +leg. + + + + + +Description. + + +Habitus +(Figs +2 +, +8 +, +9 +). Forewing length 9.0–13.5 mm, wingspan 21.0–30.0 mm. Frons and vertex yellow. Labial palpus yellowish white, with distal part of 2 +nd +segment brown. Maxillary palpus yellowish white basally, brown near distal end. Antenna yellowish brown, with brown spots on scape, ventral cilia ~ 1 / +3 in +length of diameter of flagellomere in male. Patagium and tegula yellow, with brown patches. Thorax yellowish brown dorsally, white ventrally. Legs pale yellow, front coxa and middle tibia with outer sides black at the base and distal end, front tibia with distal end black. Wings yellow, with purple-brown lines and patches. Forewing with three small spots at base, another spot near basal dorsum; antemedial line relatively straight, slightly inclined outward, accompanied by a large elliptical pale patch inside; orbicular stigma a dark brown dot; discoidal stigma reniform, yellow centrally, connected to postmedial line posteriorly; postmedial line slightly obliquely inward from costa, straightly excurved between M +2 +and CuA +2 +, then incurved to discoidal stigma below and sinuous to inner margin; an irregular large patch between anterior postmedial line and terminal margin, another irregular large patch below discoidal stigma and extended to tornus; a line of small spots along terminal margin. Hindwing with discoidal stigma a short oblique stripe; postmedial line same as forewing before CuA +2 +, not apparent afterwards; an irregular large patch near apex beyond anterior postmedial line; a band below discoidal stigma slightly inclined towards tornus, accompanied by another irregular misty wide band extended to tornus. Cilia of fore and hind wings purple-brown, white basally. Abdomen yellow dorsally, white ventrally; 1 +st +and 2 +nd +tergites with black spots laterally; 7 +th +and 8 +th +tergites black posteriorly in male; terminally black in female. + + +Male genitalia +(Fig. +15 +). Uncus semicircular. Gnathos a transverse lamina, with posterior end arc-shaped and setose (Fig. +15 a +). Valva broad tongue-shaped, with long setae along distal costa; sacculus a narrowed band; fibula a triangular lamina, setose basally (Fig. +15 b +). Saccus cylindrical, with rounded end. Juxta a narrowed plate. Phallus posteriorly with an oval protruding sclerite, with a thick, needle-like cornutus and a brush-like cornutus composed of a spine cluster (Fig. +15 c +). + + +Female genitalia +(Fig. +16 +). Apophyses anteriores ~ 2 × length of apophyses posteriores. Ductus seminalis originating from antrum. Ductus bursae ~ 2 × length of corpus bursae. Corpus bursae nearly rounded, with a short transverse bar-like signum. + + + + +Distribution. + + +China +( +Chongqing +, +Sichuan +, +Guizhou +, +Yunnan +, +Hainan +, +Hubei +, +Hunan +, +Guangdong +, +Guangxi +). + + + + +Etymology. + + +The species name +directa +is derived from the Latin word +directus +, an adjective, meaning straight, indicating the antemedial line of forewing relatively straight. + + + + \ No newline at end of file diff --git a/data/E4/B9/80/E4B98048D516566BA014B8652ABF5E82.xml b/data/E4/B9/80/E4B98048D516566BA014B8652ABF5E82.xml new file mode 100644 index 00000000000..d01c17a8adc --- /dev/null +++ b/data/E4/B9/80/E4B98048D516566BA014B8652ABF5E82.xml @@ -0,0 +1,147 @@ + + + +The new genus Purpurata (Lepidoptera, Crambidae, Spilomelinae), with descriptions of two new species from China + + + +Author + +Xue, Xin-Lei +https://orcid.org/0009-0000-5697-0872 +College of Plant Protection, Southwest University, Chongqing 400715, China & Yibin Academy of Southwest University, Yibin, Sichuan 644000, China + + + +Author + +Lu, Xiao-Qiang +https://orcid.org/0009-0009-3696-7778 +College of Plant Protection, Southwest University, Chongqing 400715, China & Guangyuan Center for Disease Control and Prevention, Guangyuan, Sichuan 628040, China + + + +Author + +Du, Xi-Cui +0000-0002-7796-7303 +College of Plant Protection, Southwest University, Chongqing 400715, China & Yibin Academy of Southwest University, Yibin, Sichuan 644000, China + +text + + +ZooKeys + + +2024 + +2024-12-02 + + +1219 + + +175 +194 + + + +journal article +10.3897/zookeys.1219.131102 +A7CEF41B-6FE9-48DF-87BD-7474CD8AEAB7 + + + + + +Purpurata shompen +( +Singh & Ahmad, 2022 +) + +comb. nov. + + + + + + + +Patania shompen + +Singh & Ahmad in + +Singh et al. 2022: 14 + +, figs 1–2, 5–7. +Type +locality: +India +(Great Nicobar Island). + + + + + + + + +Diagnosis. + + +This species is similar to + +P. iopasalis + +, but can be distinguished by the postmedial line of forewing punctiform between M +2 +and CuA +2 +, and the postmedial line of hindwing linear between M +2 +and CuA +2 +( +Singh et al. 2022 +: fig. 1); phallus posteriorly with a rectangular protruding sclerite in male genitalia ( +Singh et al. 2022 +: fig. 7). In + +P. iopasalis + +, the postmedial line of the fore and hind wings are punctiform between M +2 +and CuA +2 +(Fig. +7 +); and phallus posteriorly with a nail head-like protruding sclerite in male genitalia (Fig. +14 c +). + + + + +Distribution. + + +India +(Great Nicobar Island) ( +Singh et al. 2022 +). + + + + +Remarks. + + +This species is not found in +China +. The diagnosis is summarized based on the description and images of habitus and genitalia by +Singh et al. (2022) +. + + + + \ No newline at end of file