doi:10.4072/rbp.2023.2.01

A NEW NOMINAL GENUS FOR “PRESTOSUCHUS” CHINIQUENSIS 
HUENE, 1938 (TRIASSIC OF SOUTHERN BRAZIL): HUENESUCHUS,

GENUS NOVUS ET COMBINATIO NOVA

EDIO-ERNST KISCHLAT 
Divisão de Bacias Sedimentares (DIBASE), Superintendência de Porto Alegre (SUREG-PA), 

Serviço Geológico do Brasil (SGB/CPRM). Rua Banco da Província, 105, 90.840-030, Porto Alegre, 
Rio Grande do Sul, Brazil. edio.kischlat@sgb.gov.br

ABSTRACT – The nominal genus Prestosuchus Huene was originally proposed comprising two nominal species, but without a valid 
indication of the type-species. According to the International Code of Zoological Nomenclature, this indication is essential for proposals 
after the year 1930. Consequently, both nominal species, although valid, have a very uncomfortable situation. Therefore, a new nominal 
genus, Huenesuchus, is here proposed to correct this nomenclatural situation to be used in the new combination Huenesuchus chiniquensis. 
In addition, it is noted that two class-group names that have been used lately in the literature are previously occupied. The first, Suchia Krebs, 
is previously occupied by Simpson. The second, Loricata Merrem, is previously occupied by Schumacher. Therefore, two substitute names 
are here proposed: Holosuchia for the first and Loricatosuchia for the second.  

Keywords: Archosauria, Prestosuchus, Huenesuchus, Santa Maria Formation, Triassic.

RESUMO – O gênero nominal Prestosuchus Huene foi originalmente proposto compreendendo duas espécies nominais, mas sem indicação 
válida de espécie-tipo. De acordo com o Código Internacional de Nomenclatura Zoológica, esta indicação é essencial para propostas após o 
ano de 1930. Como consequência, ambas as espécies nominais, embora válidas, possuem uma situação bastante incômoda. Portanto, é aqui 
proposto um novo gênero nominal, Huenesuchus, para corrigir esta situação nomenclatural, a ser utilizado na nova combinação Huenesuchus 
chiniquensis. Além disso, nota-se que dois nomes de grupo-classe que ultimamente têm sido utilizados na literatura estão previamente ocupados. 
O primeiro, Suchia Krebs, está ocupado pelo de Simpson. O segundo, Loricata Merrem, está ocupado pelo de Schumacher. Portanto, são 
aqui propostos dois nomes substitutos: Holosuchia para o primeiro e Loricatosuchia para o segundo.

Palavras-chave: Archosauria, Prestosuchus, Huenesuchus, Formação Santa Maria, Triássico.

69

INTRODUCTION

Crocodylotarsian archosaurs of the Brazilian Triassic have 
been the subject of several doctoral/master’s theses (Mattar, 
1985; Azevedo, 1991; Kischlat, 2003; Mastrantonio, 2010; 
França, 2011; Lacerda, 2012; Raugust, 2014; Roberto-da-
Silva, 2017; Santos, 2017) and articles (Mattar, 1987, 1989; 
Azevedo, 1995; Kischlat, 2002; França et al., 2011, 2013; 
Mastrantonio et al., 2013, 2019; Roberto-da-Silva et al., 
2016, 2020; Lacerda et al., 2015, 2016, 2018; Desojo et al., 
2020) and Prestosuchus chiniquensis Huene, 1938a, has been 
at the center of discussion in the presence of new specimens 
belonging to this taxon. However, the formal availability 
of this nominal genus has not been investigated until now. 
The application of the International Code of Zoological 
Nomenclature (I.C.Z.N., 1999) shows that Prestosuchus 
Huene, 1938a, is an unavailable genus name. The goal of 

this paper is to discuss the nomenclatural validity of the 
nominal genus Prestosuchus Huene, 1938a, as well as some 
higher order names in the Archosaurian lineage towards the 
current crocodilians. 

MATERIAL AND METHODS

The anatomical terms follow that proposed by the 
anatomical committees (Baumel et al., 1993; I.C.V.G.A.N., 
2012; F.I.P.A.T., 2019) in place of traditional descriptive ones. 
Concerning archosaurs, the avian nomenclature (Baumel et 
al., 1993) occupies a central position. For a comprehensive 
understanding of general terminology, I suggest Collin (2007).

Throughout the body of this paper, the articles of the 
International Code of Zoological Nomenclature (I.C.Z.N., 
1999, hereinafter referred to as the Code) are precisely cited 
when and where they are relevant. I propose to use the symbol 

Revista Brasileira de Paleontologia, 26(2):69–96, Abril/Junho 2023
A Journal of the Brazilian Society of Paleontology

https://orcid.org/0000-0001-9400-7956


70 Revista Brasileira de Paleontologia, 26(1), 2023

“₢” representing “character”. This symbol was once used 
for the old Brazilian currency (cruzeiro) and nowadays, this 
currency is obsolete. Now, this symbol can have a utility.
Institucional Abbreviations: CPEZ ,  Coleção de 
Paleontologia do Museu Paleontológico e Arqueológico 
Walter Ilha, São Pedro do Sul, Rio Grande do Sul, 
Brazil; SNSB-BSPG, Staatliche Naturwissenschaftliche 
Sammlungen Bayerns (SNSB), Bayerische Staatssammlung 
für Paläontologie und Geologie (BSPG), München, Germany; 
UFRGS-PV, Laboratório de Paleovertebrados, Universidade 
Federal do Rio Grande do Sul, Porto Alegre, Rio Grande do 
Sul, Brazil; ULBRA-PVT, Coleção de Paleovertebrados, 
Universidade Luterana do Brasil, Canoas, Rio Grande do 
Sul, Brazil.

BACKGROUND: FRIEDRICH VON HUENE 
AND HIS WORK IN BRAZIL

Huene’s (1935–42) book, Die Fossilen Reptilien des 
Sudamerikanischen Gondwanalandes, is the start point of 
the knowledge of the Brazilian Triassic Paleontology. This 
work has a Portuguese translation published in 1990 by Carlos 
Burguer Júnior and revised by M.C. Barberena (see Huene, 
1990). Huene’s (1935–42) book was published in four parts 
(Lieferüngen). The first one (dealing with Anomodontia) 
was published on December 1st, 1935. The second one 
(Cynodontia) was published on October 26th, 1936, and 
the third and fourth parts (Pseudosuchia and Saurischia, 
Rhynchosauridae, and the final section – Schlußabschnitt) 
were published in Spring (Frühjahr) of 1942 (Huene, 1942: 
v, 1990:7). 

There is a previous report (Huene, 1933a:130) in the 
journal Forschungen und Fortschritte where he introduced 
three new binomina Stahleckeria potens, Belesodon 
magnificus, and Prestosuchus chiniquensis. Concerning the 
binomen Prestosuchus chiniquensis, it is a nomen nudum 
(I.C.Z.N., 1999:111) because it was not described nor defined, 
directly or indirectly by a bibliographic reference (arts. 13.1.1 
& 13.1.2).

There is also a Referate (report) authored by himself 
published in 1938a in the journal Neues Jahrbuch für 
Mineralogie, Geologie und Paläontologie. In his 1938 
Referate, he proposed (as a redundant nomenclatural 
act) the new binomina of the first two parts of his book 
(Huene, 1935 and 1936) indicating them as new (n. g. and/
or n. sp.) and there are no problems concerning priority 
with those already published, but the new binomina of 
the yet unpublished third part were published in advance. 
So, the year of publication of the binomina Prestosuchus 
chiniquensis, Prestosuchus loricatus, Rhadinosuchus gracilis, 
Hoplitosaurus raui, Rauisuchus tiradentes, Procerosuchus 
celer, and Spondylosoma absconditum is 1938a, and not 1942. 
Concerning the fourth part on Rhynchosauridae there were no 
new binomina, and all of them were already published earlier 
in Huene (1926, 1929). 

THE CODE’S “N. G., N. SP.” PAST RULE FOR 
ORIGINAL TYPE DESIGNATION

Charles W. Stiles proposed (“Opinion 7”) to the 
International Commission on Zoological Nomenclature 
(I.C.Z.N., 1910:10) that “if an author publishes a new genus 
and marks one of the species ‘n. g., n. sp.’, but does not 
otherwise specifically designate the genotype, such citation 
(‘n. g., n. sp.’) is to be construed under Art. 30a as type by 
the original designation” (his italics). It was approved, but the 
Commission noted that “this method of designating the type 
species does not […] represent the best method to adopt; on 
the contrary” and “urges all authors to state definitely that a 
certain species is type, regardless of the number of species 
placed in the genus”.

The “Opinion 7” was expressly included in the 1913 
Code’s version approved in the Ninth International Congress 
of Zoology at Monaco (I.C.Z.N., 1914:902) in Art. 30a 
(“When in the original publication of a genus, one of the 
species is definitely designated as type, this species shall 
be accepted as type, regardless of any other considerations. 
(Type by original Designation)”). In later Code versions, 
this phrasing of Art. 30a continued to be the same (I.C.Z.N., 
1958:xix), but in the 1927 Tenth International Congress of 
Zoology at Budapest, an amendment was included in Art. 
25 calling for a “definite unambiguous designation of the 
type species” (I.C.Z.N., 1945:143) and the “Opinion 7” 
only “remains valid and binding as respects generic names 
published in the period from 1st January 1758 up to, and 
including, 31st December 1930, but it is no longer applicable 
as respects any generic name published after that date” 
(I.C.Z.N., 1945:144). This subject was also discussed again 
at the Thirteenth International Congress of Zoology in Paris 
in 1948 (I.C.Z.N., 1950:152–153).

In the end, the formula “gen. n., sp. n.” (genus novus, 
species nova), was expressly disabled as an original 
designation for type-species after 1930 in the First Edition 
of the Code [I.C.Z.N., 1961:67, Art. 68(a)(i)], and repeated 
in the later three editions [I.C.Z.N. 1964:67, Art. 68(a)(i), 
1985:129, Art. 68(b)(i), 1999:70, Art. 68.2.1].

HUENESUCHUS NEW NOMINAL GENUS

Huene (1938a, 1942) introduced the nominal genus 
Prestosuchus including two nominal species: P. chiniquensis 
and P. loricatus. He (1938a:146, 1942:161) used the 
expression “n. g. n. sp.” applied to the first of his two species 
(P. chiniquensis) and, for the second (P. loricatus), he used the 
expression “n. sp.” (Huene, 1938a:147, 1942:185). So, Kuhn 
(1961a:87) indicated P. chiniquensis as the “genotypus” (cf. 
I.C.Z.N., 1912:45, footnote), followed by Krebs (1976:75, 
“Typusart”) and Barberena (1978:63, “Type-Species”).

As quoted above, the Code (Art. 13.3) asks that “every 
new genus-group name published after 1930 [...] must [...] be 



Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 71

accompanied by the fixation of a type species in the original 
publication [...]” and the expressions “gen. n., sp. n.” are 
only valid as original designation if applied before 1931 (Art. 
68.2.1). It should also be noted that any later type fixation 
likewise only applies to a taxon “established before 1931” 
(Art. 69). As a consequence, neither Kuhn’s (1961a:87), nor 
Krebs’ (1976:75), nor Barberena’s (1978:63) type indications 
can be accepted as subsequent type designations.

The absence of a valid original type-species fixation 
challenges the availability of Prestosuchus as introduced by 
Huene (1938a, 1942). It is not a matter of invalidity of a name, 
but a matter of availability, and Prestosuchus Huene, 1938a 
(or 1942) is, today, a nomen nudum (I.C.Z.N., 1999:111).

As a nomen nudum, Prestosuchus is “not an available 
name, and therefore the same name may be made available 
later for the same or a different concept” with new authorship 
and date (I.C.Z.N., 1999:111).

The possibility of understanding a past proposal of a 
new homonymous substitutive name with the same concept 
would point again to Kuhn (1961a), but, in this case, it had to 
“be expressly proposed as a new replacement name (nomen 
novum)” (Arts. 13.3, & 67.8), which is not the case in Kuhn 
(1961a), nor in any other superfluous type-species indications 
of Krebs (1976) and Barberena (1978). On the other hand, a 
formal reintroduction of the new homonymous Prestosuchus 
could also be done here, with the present date and authorship, 
but it will raise ambiguity and confusion in the literature.

Reintroducing Prestosuchus as a new genus will bring 
some problems because many authors/readers will not 
understand these nomenclatural meanders in changing, in 
practice, only the authorship. Prestosuchus is the type-genus 
of the family-group name (Prestosuchidae) and the eponym 
of the class-group name (Prestosuchia). An unusual situation 
will happen with Prestosuchus Huene, 1838 (nomen nudum), 
and the new homonym because both would be also synonyms 
in having the same species name chiniquensis associated. Two 
names being, at the same time, synonyms, and homonyms of 
each other will be very confusing. And this would also happen 
with the coordinated names Prestosuchidae and Prestosuchia.

Nomenclature is “a system of names, and provisions for 
their formation and use” (I.C.Z.N., 1999:111). On the other 
hand, Taxonomy is “the theory and practice of classifying 
organisms” (I.C.Z.N., 1999:119). Therefore, Taxonomy 
concerns the discovery/recognition of different taxa and their 
interrelationships, and Nomenclature refers to rules for a name 
to be applied to each discovered/recognized taxon.

Currently, there is already ambiguity and taxonomic 
confusion in applying the binomen Prestosuchus chiniquensis 
in several recent phylogenetic analyses and this subject should 
be properly raised. Of special interest are the huge dataset and 
the phylogenetic analysis presented by Ezcurra (2015, 2016) 
that was iterated by several authors (see below). His coding 
sequence for “Prestosuchus chiniquensis” does not reflect, nor 
include the type-material and, as consequence, this binomen 
is applied to a different taxon. Today nearly almost all the 
information in the literature, except for the description of 
the type-material of P. chiniquensis in Desojo et al. (2020), 

refers to this new taxon named preliminarily by Kischlat 
& Barberena (in Kischlat, 2002) as “Karamuru vorax”. In 
sum, the binomen Prestosuchus chiniquensis is currently 
taxonomically corrupted and this should be properly fixed.

Evaluating the nomenclatural fault discussed above and 
the current taxonomic corruption, is, therefore, necessary 
to propose a new generic name to be used in a binomen 
with the nominal species chiniquensis that nowadays only 
comprises the type-material. Here I propose the new genus 
name Huenesuchus gen. nov., to be used in the new binomen 
combination Huenesuchus chiniquensis (Huene, 1938a).

Concerning the species-group name [Prestosuchus] 
loricatus originally associated with Prestosuchus Huene, 
1838 (nomen nudum), it was included in its nominal genus 
Abaporu by Kischlat (2002:501).

SYSTEMATIC PALEONTOLOGY

ARCHOSAURIA Cope, 1870

Definition. “All the descendants of the most recent common 
ancestor of extant birds and crocodiles” (Gauthier, 1986:8). 
Recently, Gauthier & Padian (2019:1187) proposed an 
updated definition as the crown clade containing Alligator 
Cuvier, 1807, and Compsognathus Wagner, 1859, but here 
I continue with my previous definition (Kischlat, 2002:276) 
as the crown clade containing Crocodilus Cuvier, 1807, and 
Megalosaurus Buckland, 1824 (see Appendix 1).

CROCODYLOTARSI Benton & Clark, 1988

Definition. “Crocodiles and archosaurs closer to crocodiles 
than to birds” (Gauthier & Padian, 1985:189). Here I continue 
with my previous definition (Kischlat, 2002:277) as the 
stem containing Crocodylus Laurenti, 1768, but not Vultur 
Linnæus, 1758 (see Appendix 1).

HOLOSUCHIA (nomen substitutum)

Definition. Node, the least inclusive clade containing 
Aetosaurus Fraas, 1877, Rauisuchus Huene, 1938a, 
Huenesuchus gen. nov., and Crocodylus Laurenti, 1768 (updated 
and simplified from Nesbitt, 2011:195) (see Appendix 1). 
Suchia Krebs, 1974, is previously occupied by Suchia 
Simpson, 1937 (Crocodiliformes). Therefore, I propose 
the substitute class-group name Holosuchia [cf. Art. 1.2.2, 
“Articles (…10.6…) also regulate names of taxa at ranks 
above the family group” which deals with the invalidity of 
junior homonyms].

LORICATOSUCHIA (nomen substitutum)

Definition. Stem, the most inclusive clade containing 
Crocodylus Laurenti, 1768, but not Poposaurus Mehl, 
1915, Ornithosuchus Newton, 1893, and Aetosaurus Fraas, 
1877 (simplified from Nesbitt, 2011:200) (see Appendix 1). 
Loricata Merrem, 1820, is previously occupied by Loricata 



72 Revista Brasileira de Paleontologia, 26(1), 2023

Schumacher, 1817 (Mollusca). Therefore, I propose the 
substitute class-group name Loricatosuchia [cf. Art. 1.2.2, 
“Articles (…10.6…) also regulate names of taxa at ranks 
above the family group” which deals with the invalidity of 
junior homonyms].

HUENESUCHIA new class-group name

Definition. Stem, the most inclusive clade containing 
Huenesuchus, gen. nov., but not Crocodylus Laurenti, 1768 
(see Appendix 1).
Eponym. Huenesuchus gen. nov.
Nomenclatural note. As Prestosuchus Huene, 1938a (also 
in 1942) is an unavailable name (nomen nudum, I.C.Z.N., 
1999:111), the obscure class-group name Prestosuchia 
Parrish, 1993 (p. 308; also in Parrish, 1994:204) lost its 
formal eponym. Therefore, I propose Huenesuchia as a new 
class-group name (cf. analogy to Art. 37.2, which deals with 
family-group names). 

HUENESUCHIDAE new family-group name
urn:lsid:zoobank.org:act:AD764DC2-B33C-48BF-AF41-

DD2746A1D83D

Definition. Node, necessarily including Huenesuchus gen. 
nov. and, today, Stagonosuchus Huene, 1938b; to be formally 
defined in the future when greater diversity is known.
Type-genus. Huenesuchus gen. nov.
Diagnosis. Today, following the present phylogenetic 
analysis (see ahead), huenesuchids include Huenesuchus, 
gen. nov., and Stagonosuchus Huene, 1938b; which share the 
following unambiguous apomorphies: Ischium with Margo 
ventralis with an abrupt change in angle between Extremitas 
adacetabularis and Scapus (₢296.1); Fibula in its Extremitas 
distalis, with a fossa on its Facies medialis (₢422.1).
Nomenclatural note. As Prestosuchus Huene, 1938a (also 
in 1942) is an unavailable name (nomen nudum, I.C.Z.N., 
1999:111), the family-group name Prestosuchidae Romer, 
1966 (p. 368), turns also to be unavailable (Art. 37.2).

Huenesuchus gen. nov.
urn:lsid:zoobank.org:act:E5B41FEA-CF59-4955-9C70-

5B72A3CD0792

Type-species. Huenesuchus chiniquensis (Huene, 1938a), 
comb. nov.
 
Derivatio nominis. In honor of Friedrich von Huene, who 
unfortunately proposed the genus-group name Prestosuchus 
in 1938a (repeated in 1942) using a notation for fixation of 
the type-species that would be suppressed shortly soon, being 
only valid before the year 1931 (cf. Art. 68.2.1); and -suchus 
(Gr. σοῦχος, soûchos: crocodile; Bailly, 2020:2124), the name 
used by ancient Egyptians from Arsinoë (today el-Fayyūm) 
for crocodiles (Strábonos [Στράβωνoσ], ex Jones, 1967:106). 
Included species. Only the type-species.

Lectotype. SNSB-BSPG AS XXV 1~3/5~6/8~12/28~35, 
designated by Krebs (1976), comprising a complete Symphysis 
mandibulæ with most of the left Ramus mandibulæ preserved 
with some teeth, right Os premaxillare also with some teeth, 
and an incomplete postcranial skeleton. Desojo et al. (2020:8) 
indicated the number of the lectotype as “1–3/5–11/28–41/49”, 
but this is not correct because they included #7, #36~#41, and 
#49, and excluded #12, mixing specimens that are not part 
of the type-material (see Table 1). According to the SNSB-
BSPG catalog (see Supplementary file 1) there are sequential 
numbers that were here abbreviated as 1~3/5~6/8~12/28~35.
Paralectotype. An incomplete sacrum and right ilium, part of 
the last truncal vertebra, and some sacral osteoderms (SNSB-
BSPG AS XXV 7), recently described by Desojo et al. (2020).
Diagnosis. Symphysis mandibulæ with both splenials dorsally 
reaching well rostrally the dental tooth I and, in ventral view, 
with an extensive Sutura interspleniales reaching caudally 
well the level of the dental tooth V, possibly reaching the 
dental tooth VI. Scapula with the Acromion not developed, 
with a notch (Incisura acromialis) cutting the margin of the 
Os coracoidale, forming an uncination in this bone; Margo 
medialis of Os coracoidale convexly curved; Corpus humeri 
probably relatively thin. Absence of Crista supraacetabularis 
ilii; Ischium with an abrupt change in angle between 
Extremitas adacetabularis and Scapus; Femur with the 
Condylus medialis (Norma distalis) tapering to a point on the 
medial portion; Fibula with a fossa (“lunate fossa”) distally, 
on Facies medialis; Calcaneus with a short and wide Tuber. 
The paralectotype shows the transition point of the double 
row of paramedian pair of osteoderms to only a sagittal pair 
on the first caudal vertebra. (All updated from both Kischlat, 
2002, and Desojo et al., 2020).
Nomenclatural note. Prestosuchus Huene, 1938a (also 
in 1942) is an unavailable name (nomen nudum, I.C.Z.N., 
1999:111), so Huenesuchus is not a nomen novum because 
it does not replace any available name (Art. 12.2.3). But, in 
practice, it works as such because the present type-species 
(i.e., Huenesuchus chiniquensis comb. nov.) has the same 
name-bearing specimen as for the previously proposed type-
species (i.e., “Prestosuchus” chiniquensis). As an unavailable 
name, the nominal genus Prestosuchus Huene, 1938a (nomen 
nudum), should be always cited under quotation marks when 
associated with chiniquensis in the binomen “Prestosuchus” 
chiniquensis.
Stratigraphic procedence. Pinheiro-Chiniquá Sequence, 
Dinodontosaurus Assemblage Zone, Santa Maria Formation, 
Ladinian/Carnian (Middle/Upper Triassic) (Schultz et al., 
2020:5). The correct name for the locality in the Municipality 
of Candelária (State of Rio Grande do Sul, Brazil), is Pinheiro 
(29°47’30”S 52°44’25”W), in singular, and not the plural 
“Pinheiros”, as expressed by several previous authors (e.g., 
Barberena, 1978:63; Azevedo, 1991:2; Schultz et al., 2020:6). 
The Brazilian Code of Stratigraphic Nomenclature (Petri 
et al., 1986:382) asks for the “immutability of consecrated 
names”, but the name “Pinheiros” was formally listed twice 
in the Stratigraphic Lexicon of Brazil (Branco, 1984:317; 



Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 73

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.



74 Revista Brasileira de Paleontologia, 26(1), 2023

Sp
ec

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. C
on

t.



Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 75

Hasui & Baptista, 1984:317). Thus, it is convenient to rise 
this issue for proper evaluation elsewhere. 
Type locality. “Weg Sanga”, Chiniquá (29°40’1”S 
54°22’1”W), Municipality of São Pedro do Sul, west of 
Santa Maria City, Rio Grande do Sul, Brazil.

DISCUSSION

Huene (1938a:146) introduced the binomen “Prestosuchus” 
chiniquensis assembling two specimens from different 
localities as the hypodigm (vide Simpson, 1940:418, 
1945:30, 1961:185, for this concept). Later, Krebs (1976:76, 
repeated in Kischlat & Barberena, 1999:53) designated 
the first more complete specimen (SNSB-BSPG AS XXV 
1~3/5~6/8~12/28~35), that which has the mandible, as the 
lectotype. Consequently, the second specimen (SNSB-BSPG 
AS XXV 7) turned out to be the paralectotype (Art. 74.1.3). An 
extraneous third specimen (from Sanga Pascoal, at Pinheiro, 
UFRGS-PV 156T), described in Barberena (1978), was once 
considered as the “type” (Azevedo, 1995:61, “tipo”), but this 
apparent designation is invalid because this specimen is not 
part of the original type-material (cf. Art. 74.2; Kischlat & 
Barberena, 1999).

The lectotype has been cited with the number “1–
3/5–11/28–41/49” (Nesbitt, 2009:53, 2011:33; Desojo 
et al., 2020:8), but following the SNSB-BSPG Catalog 
(Supplementary file 1) this is not correct and should be 
properly fixed. This Catalog was kindly provided by Dr. Peter 
Wellnhofer when I visited the Bayerische Staatsammlung in 
München (November 1997), and it uses the acronym “1933 
L” (instead of “AS XXV”) as I have used it before (Kischlat, 
2002).

The specimen SNSB-BSPG AS XXV 7 concerns 
the paralectotype of Huenesuchus chiniquensis, and the 
specimens SNSB-BSPG AS XXV 36~41 were only originally 
referred to H. chiniquensis with doubts (Huene, 1942:185, 
1990:196; “Prestosuchus chiniquensis (?)”). These are not 
part of the type-series (cf. Art. 72.4.1). Desojo et al. (2020:8) 
also did not expressly include specimen #12 in their full 
number “1–3/5–11/28–41/49”, but they described it as part 
of the holotype.

Desojo et al. (2020:23, fig. 8.3) figured a proximal rib 
fragment as being the SNSB-BSPG AS XXV 49. But this same 
specimen was figured in Huene (1942, 1990, pl. 20, fig. 1) and 
the SNSB-BSPG Catalog lists it as #8. In this Catalog, the true 
#49 is indicated as part of a dorsal rib (“dorsalrippenstück”) 
belonging to “Prestosuchus loricatus”, but I didn’t find its 
description in Huene’s (1942:185–191) text. On the other 
hand, the description of three rib fragments (#19 and #21a~b) 
is present (Huene, 1942:188, “Dorsalrippen”).

Huene (1942:186) hypothesized that the type-material 
of Prestosuchus loricatus  (SNSB-BSPG AS XXV 
13~24/26~27/43~48) belongs to a single individual (“einem 
einzigen Individuum herrühren”). Thus, all material should 
be called the holotype and not as syntypes, and only syntypes 
could become lectotype and paralectotype(s) (cf. Art. 73.2.2). 
Any restriction of this holotype should follow Art. 73.1.5 (“If 

a subsequent author finds that a holotype [...] is not derived 
from an individual animal, the extraneous components may, 
by appropriate citation, be excluded from the holotype”), and 
Huene (1942:186) expressly indicated that in case of doubt, 
the species name should be attached to the presacral vertebral 
remains (“Im Zweifelsfall soll der Speciesname an den 
Praesacralwirbelresten hängen”). The composite nature of the 
holotype is still to be demonstrated (although very probable), 
but if something is Prestosuchus loricatus, the two presacral 
vertebrae (#13a~b) are the core. Huene (1942:190–191) 
also referred to additional material (SNSB-BSPG AS XXV 
4/25/42) in doubts (“Prestosuchus loricatus (?)”) and these 
specimens are not part of the type-series (cf. Art. 72.4.1). I 
didn’t unequivocally find the citation of specimens #46a~l in 
Huene’s (1942:185–191) text (“Ein isoliert Dornfortsatz”?, 
p. 189).

Desojo et al. (2020:6) indicated the full number of the 
“lectotype” (or “holotype”, p. 9) of Prestosuchus loricatus 
as “13–24/26–27/44–48”, excluding specimen #43, which 
was included in Huene (1942:189). Concerning the purported 
“paralectotype”, they did not cite expressly any specimen, but 
according to their dataset it is the additional material (SNSB-
BSPG AS XXV 4/25/42) originally referred with doubts to 
Prestosuchus loricatus. Aside from the fault in not following 
Art. 72.4.1 (exclusion of doubt specimens from type-series) 
and Art. 73.2.2 (only syntypes could become lecto- and 
paralectotypes), these lectotype/paralectotype indications 
were not done as an express statement (cf. Art. 74.7) and they 
are not valid (although repeated in Nesbitt et al., 2020:38; 
Tolchard et al., 2021:597; Butler et al., 2022:4).

The first mention of the lectotype of Prestosuchus 
chiniquensis concerns a short note made by Huene (1929:54) 
briefly describing a sketch of the mandible (SNSB-BSPG AS 
XXV 1) sent to him from Brazil and supposedly identified as 
a probable belodont (“wahrscheinlich Parasuchier”). Later, 
Huene (1942:164) gave more information about the material 
assembled under this binomen explaining that the mandible 
and some bones were collected in 1923 by Vicentino Prestes 
de Almeida (on whom the name “Prestosuchus” was based; 
Beltrão, 1965:20). In his trip to southern Brazil in 1928 
(vide Huene, 1930), after some searching, he collected the 
remaining material. It came from the locality called “Sanga 
am Wege” or “Weg Sanga” (Huene & Stahlecker, 1931:40, 
1968:35; Huene, 1942:161, 1990:171; “Sanga da Estrada” 
in Portuguese), at the locality of Chiniquá (29°40’1”S 
54°22’1”W), Municipality of São Pedro do Sul.

The paralectotype of Huenesuchus chiniquensis (SNSB-
BSPG AS XXV 7) came from an upper layer from a near 
locality called “Sanga des Theotonio Beles Xavier” (Huene 
& Stahlecker, 1931:38–39, 1968:34, “Sanga Béles”; Huene, 
1942:183). “Cynodontier Sanga” is another name for this 
locality (Huene, 1935:93, 1942:325, 1990:103, 341, “Sanga 
dos Cinodontes” in Portuguese) and it was figured in Huene & 
Stahlecker (1931:38, 1968:34, fig. 17) and Huene (1942:327, 
1990:344, fig. 66). The paralectotype was also considered a 
supposed belodont (“Parasuchier”) in Huene & Stahlecker 
(1931:39, 1968:34).



76 Revista Brasileira de Paleontologia, 26(1), 2023

Recently, Desojo et al. (2020:6–7) redescribed all the 
type-series of Huenesuchus chiniquensis and accepted their 
conspecifity like Huene (1938a, 1942) originally hypothesized. 
This approach should be accepted as a hypothesis to be 
tested in the future when better and more complete material 
comes to hand. The assumption of this hypothesis implies 
that both lectotype and paralectotype specimens are closely 
related, representing the same taxon, and all the remaining 
available specimens described elsewhere (Barberena, 1978; 
Azevedo, 1991, 1995; Kischlat, 2002, 2003; Mastrantonio, 
2010; França, 2011; França et al., 2011, 2013; Lacerda, 2012; 
Mastrantonio et al., 2013, 2019; Raugust, 2014; Roberto-da-
Silva et al., 2016, 2020; Lacerda et al., 2016; Roberto-da-
Silva, 2017; Damke et al., 2022) should be compared to them, 
chiefly to the lectotype. In this way, the specimen from Sanga 
Pascoal (UFRGS-PV 156T; Barberena, 1978; Azevedo, 1991, 
1995) was recognized as being another taxon (Kischlat, 2002, 
2003). On the other hand, Decuriasuchus (França, 2011:50; 
França et al., 2011:391, 2013:474) was mainly compared to 
the specimen from Sanga Pascoal (UFRGS-PV 156T) that 
was then identified as H. chiniquensis.

The lectotype of Huenesuchus chiniquensis was 
individually scored in Parrish’s (1993), Nesbitt’s (2009, 
2011), and França’s (2011) datasets. The paralectotype 
was individually scored in França (2011) and Desojo et al. 
(2020). The combined type-series (lectotype+paralectotype), 
as originally proposed by Huene (1938a), was scored in 
Desojo et al. (2020). And the lectotype and the Sanga Pascoal 
specimen (UFRGS-PV 156T) were combined and scored 
in the datasets of Benton & Walker (2002), Benton (2004), 
Nesbitt (2009, 2011), and Brusatte et al. (2010).

Benton’s (2004) dataset was later iterated by Li et al. 
(2006). Nesbitt’s (2011) dataset was later iterated by several 
authors (Butler et al., 2011, 2014, 2018, 2022; Nesbitt et al., 
2011, 2013a, c, 2014, 2017, 2018a, b, 2020; Li et al., 2012, 
2016; Langer & Ferigolo, 2013; Lecuona, 2013; Nesbitt & 
Butler, 2013; Sues & Schoch, 2013; Baczko et al., 2014, 
2020; Raugust, 2014; Sookias et al., 2014a, b; Lautenschlager 
& Rauhut, 2015; Zanno et al., 2015; Cabreira et al., 2016; 
Lacerda et al., 2016, 2018; Lecuona et al., 2016; Lessner et 
al., 2016; Niedźwiedzki et al., 2016; Roberto˗da˗Silva et 
al., 2016, 2020; Nesbitt & Desojo, 2017; Roberto˗da˗Silva, 
2017; Stocker et al., 2017; Müller et al., 2018; Sarıgül et al., 
2018; Garcia et al., 2019, 2021; Barrett et al., 2020; Desojo et 
al., 2020; Kammerer et al., 2020; Marsh et al., 2020; Baron, 
2021; DallaVecchia, 2021; Tolchard et al., 2021; Parker et 
al., 2021; Damke et al., 2022; ) and many of them including 
modifications/corrections and inserting new taxa. With few 
exceptions (e.g., Li et al., 2012, 2016; Desojo et al., 2020) 
the combined score of the lectotype plus the Sanga Pascoal 
specimen (UFRGS-PV 156T) was the standard sequence used 
in their phylogenetic analyses for “Prestosuchus chiniquensis” 
terminal. This was also the case in Brusatte’s et al. (2010) 
dataset, with the combined lectotype plus UFRGS-PV 156T 
as the only available sequence and it was later iterated by 
several authors (Mastrantonio, 2010; Benton & Walker, 
2011; Butler et al., 2011; França et al., 2011; Lacerda, 2012; 

Lautenschlager & Rauhut, 2015; Nesbitt et al., 2014) also 
with modifications/corrections and insertions of new taxa.

Finally the binomen “Prestosuchus” chiniquensis was 
used for scoring an assemblage of specimens not including 
the lectotype (i.e., the name-bearing specimen) by Ezcurra 
(2015:183, tab. 5.1; 2016:110, tab. 1), which was also later 
iterated by several authors (Ezcurra et al., 2017, 2019, 2020b, 
2021a, b, c, d, 2022; Nesbitt et al., 2017, 2018b; Sengupta 
et al., 2017, 2022; Stoker et al., 2017; Ezcurra & Butler, 
2018; Oliveira et al., 2018; Spiekman, 2018; Butler et al., 
2019; Peecook et al., 2019; Baczko et al., 2020; Barrett et 
al., 2020; Bennett, 2020; Foffa et al., 2020, 2022; Maidment 
et al., 2020; Müller et al., 2020; Scheyer et al., 2020; Sues et 
al., 2020, 2021; Troteyn & Ezcurra, 2020; Wynd et al., 2020; 
DallaVecchia, 2021; Parker et al., 2021; Ezcurra & Sues, 
2022; Kellner et al., 2022; Martínez et al., 2022; Müller & 
Garcia, 2022; Pretto et al., 2022; Sengupta & Bandyopadhyay, 
2022; Simão-Oliveira et al., 2022; Chen & Liu, 2023). 
Therefore, these scorings don’t reflect, nor include the type-
material and, as consequence, they are not representative of 
H. chiniquensis.

As noted above, Kischlat (2002, 2003) considered the 
Sanga Pascoal specimen (UFRGS-PV 156T) as a different 
taxon from Huenesuchus chiniquensis and the binomen 
Karamuru vorax Kischlat & Barberena in Kischlat, 2002 (cf. 
Code’s Recommendation 51E) was preliminarily proposed 
for this new taxon.

Two new nearly complete specimens were discovered 
after, the first in March 2003 (UFRGS-PV 629T; Mastrantonio 
et al., 2009) and the second in May 2010 (ULBRA-PVT 2810; 
Cabreira et al., 2011). For me, at each time of discovery, 
both specimens showed marked differences in mandibular 
morphology with that of the lectotype of Huenesuchus 
chiniquensis, and many similarities with the mandible of 
the Sanga Pascoal specimen (UFRGS-PV 156T). But, as the 
specimen from Sanga Pascoal was previously identified as 
“Prestosuchus chiniquensis” (Barberena, 1978; Azevedo, 
1991, 1995), an obvious conclusion quickly came, and my 
previous hypothesis was then rejected by all involved. In the 
literature this rejection was mainly because of nomenclatural 
problems rather than the recognition of osteological 
differences to understand that it represents a new taxon 
(Mastrantonio, 2010:43; França, 2011:20; Lacerda, 2012:29; 
Raugust, 2014:1.72; Lacerda et al., 2016:3; Roberto-da-Silva, 
2017:40; Roberto-da-Silva et al., 2020:993; Desojo et al., 
2020:3). In short, these authors confused Nomenclature with 
Taxonomy and did not conclude what, to me, was obvious 
(i.e., the presence of two different taxa whatever the names 
available). Although the binomen “Karamuru vorax” (but see 
Abdala et al., 2009:84) is not available (cf. Arts. 16.1 & 16.4), 
its formal unavailability does not preclude the discovery/
recognition of a new taxon, even if no name was formally 
proposed. Unfortunately, after 2003 I could not continue my 
studies on the subject and, with the discovery of these two 
new specimens (UFRGS-PV 629T and ULBRA-PVT 281), I 
waited for the conclusions of those involved. Now I feel quite 
comfortable going back and defending my earlier hypothesis.



Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 77

Differences in jaw morphology between the lectotype and 
the new taxon called “Karamuru” are easier to observe using 
specimen UFRGS-PV 629T than with the others (UFRGS-PV 
156T and ULBRA-PVT 281) because in these the mandibles 
are tightly adpressed to the skulls.

The actual length of the lectotype mandible is unknown 
because the caudal part is missing, but the preserved part has 
49.0 cm (Desojo et al., 2020:11). Otherwise, the complete 
hemimandible of the specimen UFRGS-PV 629T has 47.19 
cm in length (Mastrantonio, 2010:128). Thus, in linear 
dimensions, the lectotype has a larger mandible and it would 
represent a larger individual. For comparisons, both mandibles 
were normalized with the same dimension from the rostral 
tip of the symphysis to the caudal tip of the ventrocaudal 
process of the dental bone (Supplementary file 2). The 

actual dimension of the lectotype mandible is approximately 
150% larger than that of UFRGS-PV 629T, so its apparent 
slenderness does not correspond to a juvenile condition of the 
lectotype. Thus, the relative height is shorter and the Ramus 
mandibulae is slenderer. There is a lesser curvature in the 
Margo oralis and the Eminentia rostralis is more developed 
under tooth III. The rostral teeth (II–IV) are relatively 
procumbent when compared to the highly recumbent ones 
in UFRGS-PV 629T.

These differences should be added to the most important one 
in the morphology of mandibular symphysis of the lectotype 
with both splenials dorsally reaching well rostrally the dental 
tooth I (Figure 1C) (cf. Huene, 1942:161, “Die Splenialia 
reichen bis an die vordere Spitze der Symphyse”, 1990:171) 
and, in ventral view, with the Sutura interspleniales reaching 

Figure 1. Mandible of the lectotype of Huenesuchus chiniquensis (SNSB-BSPG AS XXV 1). A, lateral view. B, dorsal view. C-D, detail of the symphysis 
in laterorostral and ventral view, respectively. All the pictures from the author except (D) which is modified from Desojo et al. (2020). Scale bars = 50 mm.  



78 Revista Brasileira de Paleontologia, 26(1), 2023

Figure 2. Drawline showing the main differences in scapulocoracoids between the (A) lectotype of Huenesuchus chiniquensis (SNSB-BSPG AS XXV 12) 
and the (B) new taxon (UFRGS-PV 629T, reversed). Abbreviations: Cav. glen., glenoid cavity (Cavitas glenoidale); Inc. acrom., acromial notch (Incisura 
acromialis); Fac. artic. clav., articular surface for clavicle (Facies articularis clavicularis); Synd. coracoscap., coracoscapular syndesmosis (Syndesmosis 
coracoscapularis). [A, modified from Parrish (1993), B, modified from Mastrantonio (2010)]. Scale bars = 50 mm.

caudally well the level of the dental tooth V (Figure 1D), 
possibly reaching the dental tooth VI (cf. Desojo et al., 
2020:17, fig. 5.1–2; cf. Raugust, 2014:44, fig. 21ed). These 
differences are certainly connected to the mode of prey, and 
both were carnivorous archosaurs. 

Concerning the morphology of an elongated mandibular 
symphysis of the lectotype, the ventral extension of the 
intersplenial suture draws attention to the condition found 
in Mambawakale (Butler et al., 2022:15, figs. 10–11), which 
shows, basically, the same morphology of the mandible of 
H. chiniquensis, with an elongated Symphysis mandibula 
extending up to dental tooth VIII with a clear and elongated 
Sutura interspleniales rostrally from, at least, tooth II to 
caudally reaching tooth VIII.

Concerning the Scapula (Figures 2A and 3A) of the 
lectotype, the Acromion is not developed, and there is a 
notch (Incisura acromialis) cutting the margin of the Os 
coracoidale, forming an uncination in this bone. In the new 
taxon, the Acromion is well developed and the Incisura 
acromialis is absent (the scapulocoracoid is only preserved in 
UFRGS-PV 629T and ULBRA-PVT 281; Figures 2B and 3B).

Desojo et al. (2020:35) also gave several characters 
found in the lectotype Huenesuchus chiniquensis (and 
scored in their dataset) that differentiate it from the specimen 
UFRGS-PV 629T: absence of Crista supraacetabularis ilii 
(₢265.0); Ischium with an abrupt change in angle between 
the Extremitas adacetabularis and the Scapus (₢296.1); 
Femur with the Condylus medialis (Norma distalis) tapering 
to a point on the medial portion (₢320.0); and, Fibula with a 
fossa (“lunate fossa”) distally on Facies medialis (₢422.1). 
All these characters are not preserved in UFRGS-PV 156T, 

but only in UFRGS-PV 629T (₢320 also in ULBRA-PVT 
281) and they show a different condition (cf. Mastrantonio, 
2010; Roberto-da-Silva et al., 2016; Roberto-da-Silva, 2017) 
from that found in the lectotype of H. chiniquensis (cf. Desojo 
et al., 2020).

On the other hand, the paralectotype shows the transition 
point between the double row of paramedian pair of 
osteoderms to only a double sagittal row on the first caudal 
vertebra (Figure 4). This condition is also found in 
Decuriasuchus quartacolonia (França, 2011:142, fig. 35A), 
Postosuchus alisonae (Peyer et al., 2008:373, fig. 6E), and, 
apparently, also in Ticinosuchus ferox (Krebs, 1965:114). 
Other taxa, such as Saurosuchus galilei (Sill, 1974:349, fig. 
10) and Fasolasuchus tenax (Bonaparte, 1981:74, fig. 20) 
also show symmetric osteoderms supposed as from the tail, 
but they are not preserved articulated. The crocodylomorph 
Dromicosuchus grallator has a double row of paired 
paramedian osteoderms on the first two caudal vertebrae 
(Sues et al., 2003:330). The condition found in the specimen 
UFRGS 629T is that double symmetric sagittal osteoderms 
are found on both sacral vertebrae and the transition point 
is between the Vertebra truncale ultima (= truncale XVII, 
= presacrale XXV) and Vertebra sacrale I (Mastrantonio, 
2010:159, figs. 84A and 87C; Raugust, 2014:2.84, fig. 41A).

 The dataset of Desojo et al. (2020) is an iteration of 
that of Nesbitt (2019, 2011) plus more characters and taxa. 
The existence of intersplenial suture is not evaluated and 
their ₢160 only concerns its absence or presence rostrally 
(plesiomorphy) and presence relative to one-third of the lower 
jaw (apomorphy). This same wording was used by Ezcurra 
(2015:₢251, 2016:₢265) and later iterations. The presence 



Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 79

Figure 3. Scapulocoracoids of several Brazilian Triassic specimens (left side) normalized with the approximately horizontal coracoscapular syndesmosis. 
A, lectotype of Huenesuchus chiniquensis (SNSB-BSPG AS XXV 12) comprising both claviculae, interclavicula, both coracoids, and left scapula (with 
incomplete dorsal Lamina). B, complete scapula and coracoid of the new taxon (UFRGS-PV 629T). C, holotype of Procerosuchus celer (SNSB-BSPG AS 
XXV 134) with the coracoid and the fragmentary scapula (only Corpus scapulae preserved). D, Baum Sanga specimen (CPEZ 239b, reversed as left) with 
the coracoid badly preserved attached with only part of the scapular body. [All the pictures from the author except B which is modified from Mastrantonio 
(2010)]. Scale bars = 50 mm.

of intersplenial suture in the condition of Huenesuchus (and 
Mambawakale) was only evaluated in two other expanded 
iterations of Nesbitt’s (2009, 2011) dataset (Lecuona, 
2013:₢413.1; Lecuona et al., 2016:₢413.1), but the terminal 
“Prestosuchus chiniquensis” was coded as “?”. Another 
dataset concerning only later crocodylomorphs (Leardi et 
al., 2017:₢83.1) also uses this approach. Evaluating all these 
datasets (and there are differences in codings that should 

be investigated), the presence of intersplenial suture was 
coded for Euparkeria, belodonts, aetosaurs, ornithosuchians, 
Yonghesuchus as well in several crocodylomorphs.

Kischlat (2002:291, 2003:263) noted that the condition 
in UFRGS-PV 156T was different from the lectotype of 
Huenesuchus chiniquensis, but the full medial view of the 
symphysis was precluded for more accurate observations. 
Otherwise, the condition in UFRGS-PV 629T (Mastrantonio, 



80 Revista Brasileira de Paleontologia, 26(1), 2023

2010:135, fig. 72A; Mastrantonio et al., 2019:189, fig. 17) 
shows that there isn’t any clear Facies articularis dentalis 
dentale (dental articular face for the other dental bone). So, 
the contact between both dental bones was not strong, and 
this suggests that the symphysis was formed by ligaments in 
a syndesmosis (cf. Holliday et al., 2010:1351; cf. Holliday & 
Nesbitt, 2013:556). The condition in ULBRA-PVT 281 is very 
similar with the dentals (= dentaries) only attached to each 
other in a probable syndesmotic symphysis (Roberto-da-Silva 
et al., 2016:983, fig. 5; Roberto-da-Silva, 2017:148, fig. 4). 

In the classification of symphysis types, the specimen 
UFRGS-PV 156T was considered by Holliday & Nesbitt 
(2013:564, tab. 1) as from Class I, characterized by flat 
symphyseal plates, conspicuous smooth region, and equally 
high and deep joint. Otherwise, Huenesuchus chiniquensis, 
with an extensive symphysis, although formed largely by the 
splenials, appears to be near Class II symphysis, a condition 
also indicated for Saurosuchus (Alcober, 2000:315, fig. 10A; 
Holliday & Nesbitt, 2013:565, fig. 7c).

I noted previously (Kischlat, 2002:290) that the holotype 
of Procerosuchus celer (SNSB-BSPG AS XXV 131~139), 
could represent a juvenile of Huenesuchus chiniquensis 

(endorsed by Desojo & Rauhut, 2008:19; and Baczko et al., 
2019:14). But I was wrong because there is no evidence that 
the holotype of P. celer could be an ontogenetic miniature of 
the lectotype of H. chiniquensis.

Both the holotype of Procerosuchus celer and the 
lectotype of Huenesuchus chiniquensis were found in 
the same locality (“Weg-Sanga”; Huene & Stahlecker, 
1931:40, 1968:35; Huene, 1942:217, 1990:228; “Sanga da 
Estrada” in Portuguese), but in different places (“Grabung 
37” in the former, “Grabung 34” in the latter; Huene, 
1942:161, 1990:171). The type-series of the dicynodont 
Dinodontosaurus turpior (Huene, 1935:76, 1990:85) also 
came from the Weg-Sanga (“Grabung 38”). 

Three remarkable details are present in the holotype 
of Procerosuchus celer. As noted by Huene (1938a:148, 
1942:220, 1990:230) the morphology of the humerus shaft is 
exceptionally thin, resembling the condition of the additional 
material recently described (Lacerda, 2012:98, f. 53; Lacerda 
et al., 2016:30, fig. 19) as Huenesuchus chiniquensis. This 
latter specimen (now CPEZ 239b) I personally partially 
collected in 1994 (but I didn’t find it) in the Baum Sanga 
locality (Huene & Stahlecker, 1931:39, 1968:35; Beltrão, 

Figure 4. Dorsal view of the paralectotype of Huenesuchus chiniquensis (SNSB-BSPG AS XXV 7) comprising the dorsal margin of the right (dexter) iliac 
blade, both right sacral ribs, and several osteoderms (Osteoderma paramedianum), the right double paramedian set of the first sacral vertebra (Vertebra sacrale 
I), both sides of double paramedian set of the second sacral vertebra (Vertebra sacrale II), and the sagittal double set (Osteoderma sagittale) of the first caudal 
vertebra (Vertebral caudale I). Scale bar = 100 mm. 



Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 81

1968:86), mixed with the holotype of Archaeopelta 
arborensis Desojo et al., 2011. This is the same locality 
where the dicynodont Stahleckeria potens and the aphanosaur 
Spondylosoma absconditum were found (Huene, 1933a:129, 
1933b:129; 1935:1, 1942:326; 1990:11, 342).

The condition of the humerus in the lectotype of 
Huenesuchus chiniquensis could not be fully observed 
because the shaft is not preserved, but it is suggestive of being 
relatively thin (pers. obs.; Raugust, 2014:104, fig. 52D–E; 
Desojo et al., 2020:27, fig. 10.4–6).

The second detail is the presence of a notable Incisura 
acromialis with also a uncinated margin of the coracoid at 
the joint between coracoid and scapula (Huene, 1942:219, 
1990:230, pl. 29.1), which is also found in the lectotype of 
Huenesuchus chiniquensis (pers. obs.; Huene, 1942:171, pl. 
21.1, 1990:182; Raugust, 2014:99, fig. 49, “ent”; Desojo et 
al., 2020:23, fig. 9, “oi”), but in Procerosuchus celer the 
acromion is well developed (i.e., both margins of scapula 
and coracoid have similar cranial limits; Figure 3C). An 
Incisura acromialis could be potentially present, although the 
uncination is not clearly preserved, in the additional material 
from Baum Sanga (CPEZ 239b; Lacerda, 2012:96, fig. 97; 
Lacerda et al., 2016:29, fig. 18A, “?coracoid foramen”; 
Figure 3D). The third detail concerns the medial margin of 
the coracoids. It is nearly straight in the holotype of P. celer 
and curved in the lectotype of H. chiniquensis (Figures 2A, 
3A, C). In sum, there are notable morphological differences 
between H. chiniquensis and P. celer, and, as consequence, 
they represent different taxa, although sharing the slenderness 
of humeri and the presence of acromial uncinated incisure.

As cited before, critical parts of the lectotype of 
Huenesuchus chiniquensis (mandible and trunk plus fore/
hindlimbs) were collected on two different occasions. Mixed 
material is commonly found in Santa Maria Formation, 
therefore the mixed condition of the lectotype cannot be ruled 
out. In this hypothetical case, the binomen H.chiniquensis 
(Huene, 1938a) should be restricted to the mandible (SNSB-
BSPG AS XXV 1) with all the remaining parts having this 
mandible as the core of the binomen (cf. Art. 73.1.5).

PRELIMINARY PHYLOGENETIC 
ANALYSIS

Kischlat (2002, 2003) hypothesized that the Sanga Pascoal 
specimen (UFRGS-PV 156T) was taxonomically different 
from Huenesuchus chiniquensis. To preliminary test the 
interrelationships of H. chiniquensis and this possible new 
taxon, I used Desojo’s et al. (2020) modified dataset. This 
one was initially taken from Nesbitt (2011), modified by 
Butler et al. (2014), and later by Nesbitt & Desojo (2017). 
As Desojo et al. (2020) did not indicate which characters 
are ordered, I followed Nesbitt & Desojo (2017) ordering 
characters 32, 52, 121, 137, 139, 156, 168, 188, 223, 247, 
258, 269, 271, 291, 297, 328, 356, 399, and 413, with the 
following modifications: (1) exclusion of the lectotype 
and paralectotype of Huenesuchus chiniquensis, but using 

Huene’s (1938a) original hypodigm, expressly endorsed by 
the authors, and using the combined sequence; (2) exclusion 
of Vale Verde specimen (UFRGS-PV 152T); (3) exclusion 
the Baum Sanga specimen (CPEZ 239b); (4) exclusion of 
the “paralectotype” (sic!, probably SNSB-BSPG AS XXV 
4/25/42) of “Prestosuchus” loricatus; (5) exclusion of 
the individual sequences of Pseudolagosuchus major and 
Lewisuchus admixtus, but using their combined sequence, 
reflecting a new hypodigm comprising both binomina 
(Ezcurra et al., 2020a; Agnolín et al., 2021); (6) substitution of 
the sequence of Rauisuchus tiradentes in the characters 1–412 
by the updated sequence given by Lautenschlager & Rauhut 
(2015), but changing the ₢75(3→2); (7) substitution of the 
sequence of Parringtonia gracilis in the characters 1–412 
by the updated sequence given by Nesbitt et al. (2018a); (8) 
inclusion of the sequence of Teratosaurus suevicus given by 
Lessner et al. (2016) completing characters 413–422 with 
“?”; and, the most important, (9) the combined sequence 
of both the lectotype and paralectotype of Huenesuchus 
chiniquensis, given by Desojo et al. (2020:47), fails in the 
scores for ₢412(?→0), ₢416(?→1), ₢417(?→1), ₢418(?→1), 
₢419(?→1), ₢420(?→0), ₢421(?→1) and ₢422(?→1), but all 
were scored in the lectotype. Except for these modifications, 
I did not modify/correct any other score. This will be done in 
an upcoming paper dealing with the proposition of the new 
taxon using UFRGS-PV 156T as the name-bearing specimen 
(cf. Kischlat, 2002, 2003).

Mesosuchus browni and Prolacerta broomi were used 
as outgroups, as originally proposed by Nesbitt (2009:403, 
2011:185). Instead, he did not implement this composite 
outgroup because he used the software TNT 1.0 (Goloboff 
et al., 2003, 2008) for his phylogenetic analyses. So far as I 
know, TNT software does not work with a composite outgroup 
group like PAUP 4.0, but this is possible as recently shown 
(Goloboff, 2022:154).

The matrix with 91 taxa (two as outgroup) was analyzed 
under equally weighted parsimony using P.A.U.P. 4.0β10 
(Swofford, 2002) and 4.0α (Swofford & Bell, 2017) and 
the results were the same. I used a heuristic search and 100 
replicates, using a batch procedure (see Appendix 2). This 
resulted in 79,380 maximum parsimony trees (MPTs). When 
multistate taxa are interpreted as a variable (polymorphisms 
and uncertainty, see Swofford & Bell, 2017:105 for 
differences), the length has 1,476 steps (CI = 0.348; RI = 
0.754); when interpreted as all uncertainties the length has 
1,456 steps (CI = 0.339; RI = 0.754). The strict consensus 
tree has a good resolution with a polytomy only at basal 
Avemetatarsalia and Crocodylomorpha+Teratosauria clades 
(Figure 5). The majority rule tree (50%) shows these clades 
with a better resolution inside (see Supplementary file 3). Such 
results are not different from the results achieved by Nesbitt 
(2009, 2011) and the later iterations. 

The result shows the original hypodigm of Huenesuchus 
chiniquensis nested with Stagonosuchus nyassicus, and, 
higher in the next clade, the specimen UFRGS-PV 156T 
nested with Luperosuchus fractus and Saurosuchus galilei.



82 Revista Brasileira de Paleontologia, 26(1), 2023

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Kischlat – A new nominal genus for “Prestosuchus” chiniquensis Huene, 1938 83

This preliminary result shows that the specimen UFRGS-
PV 156T is a different taxon from Huenesuchus chiniquensis 
contradicting the opinion of many researchers and supporting 
my previous hypothesis (Kischlat, 2002, 2003). Huenesuchus 
chiniquensis is related to Stagonosuchus nyassicus, a result 
virtually achieved by Desojo et al. (2020). But UFRGS-PV 
156T is closer to Luperosuchus fractus and Saurosuchus 
galilei, as well as all the remaining teratosaurians and 
crocodylomorphs.

FINAL REMARKS

Unfortunately, the universally used nominal genus 
Prestosuchus Huene, 1938a (nomen nudum), is not 
available in the light of the Code. As such, the specific name 
chiniquensis, part of the original Huene’s (1938a, 1942) 
binomen “Prestosuchus” chiniquensis, turned out to be an 
orphan. This awkward nomenclatural situation was fixed here.

Huenesuchus chiniquensis was a taxon that reached a 
large size with a mandible exceeding 50 cm and with a huge 
femur measuring nearly 45 centimeters. “‘Prestosuchus’ 
chiniquensis” has been used as the iconic taxon representative 
for Brazilian Triassic, but the specimen usually figured 
(UFRGS-PV 156T) is not Huenesuchus chiniquensis but 
represents a new taxon, as noted before (Kischlat, 2002, 2003).

Past errors in the literature were corrected here and I hope 
that this present contribution can provide a new nomenclatural 
and taxonomic understanding of the hypercanivorous 
crocodylotarsans archosaurs of the Triassic fauna of Rio 
Grande do Sul (Brazil). The choice of honoring Friedrich von 
Huene, and his seminal work, introducing the new nominal 
genus Huenesuchus, should be considered as a tribute that 
I could not avoid doing, literally calling this already long-
known taxon paronomastically as “Huene’s suchus from 
Chiniquá”.

ACKNOWLEDGEMENTS

I am especially grateful to my former advisor (deceased), 
Dr. Mario C. Barberena, who from our first conversation 
in 1994 alerted me about the nomenclatural problems of 
Prestosuchus and the large skull described by him in 1978 
(“I stuck it in Prestosuchus, but it’s not Prestosuchus!”). 
I wish to thank many colleagues that helped me on my trip to 
Germany (land of my grandfather Ernst-Wilhelm Kischlat) 
in late 1997: D.J. Gower (British Museum, Natural History, 
London), E.S. Gafiney (American Museum of Natural History, 
New York), M.W. Maisch (Institut und Museum für Geologie 
und Paläontologie, Eberhard-Karls-Universität, Tübingen), P. 
Wellnhofer (Bayerische Staatssammlung für Paläontologie 
und Geologie, München), R. Wild and R. Schoch (Staatliches 
Museum für Naturkunde, Stuttgart). Several papers were 
downloaded from the Wikipaleo community and R. Guzman-
Gutierrez (Universidad Humanista de las Américas, Monterey, 
Mexico) helped me many times. B.L. Horn (project manager 
here in Serviço Geológico do Brasil, Porto Alegre) and the 

anonymous reviewers that improved my text, as well the 
always kind A.M. Ribeiro (Museu de Ciências Naturais, 
Porto Alegre). A special thanks to F.D. Ross, who helped me 
on numerous occasions in the first decade of the 2000s, when 
I had a really troubled time. He gave me the strength to get 
here. Thank you all.

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