Brazilian Miocene crabs I. Taxonomic review of Cyclocancer tuberculatus Beurlen, 1958 and Hepatella amazonica Beurlen, 1958 (Pancrustacea, Decapoda, Brachyura)

Abstract

In describing Cyclocancer to accommodate his new species C. tuberculatus, Beurlen highlighted that his new genus was a cancroid intermediate in form between Atelecyclus and Cancer, but with enough characters to be placed in a new genus. Examination of C. tuberculatus type material revealed that Cyclocancer is a junior synonym of Hepatus. Therefore, C. tuberculatus is herein transferred into Hepatus as H. beurleni nomen novum because H. tuberculatus (Beurlen, 1958) is a junior homonym of H. tuberculatusSaussure (1858), a junior synonym of Hepatus pudibundus (Herbst, 1785). Miohepatus gen. nov., comb. nov. is proposed here to accommodate Hepatella amazonica Beurlen from the Pirabas Formation, Neogene, Pará, Brazil, based on new morphological evidence from the type material.

Keywords
Fossil decapods; Pirabas Formation; Systematics; Aethridae; Aethroidea

INTRODUCTION

Karl Beurlen (1901-1985) was a German geologist and paleontologist who worked in Brazil from 1950 to 1969, invited by the Departamento Nacional de Produção Mineral (DNPM), currently Agência Nacional de Mineração (ANM). During this period, he described several fossil crustacean species, with special reference to the decapods. One of his important works was in the Pirabas Formation, north region of Brazil, where he described several Miocene species (Beurlen, 1958) in the first work describing decapod crustaceans from that formation (Aguilera & Páes, 2012). Although with undeniable importance, this work needs revision, mostly in the descriptions and updated images, with new photos of the species studied.

The revaluation of the species described by Beurlen revealed that Hepatella amazonicaBeurlen, 1958, does not belong to Hepatella Smith, in Verril (1869) and cannot be assigned to an existing genus. Therefore, the new genus Miohepatus is described here with Miohepatus amazonicus as a new combination. Also, morphological evidence based on the type material from the Pirabas Formation revealed that CyclocancerBeurlen, 1958, is a junior synonym of Hepatus Latreille, in Bosc, 1801. Cyclocancer tuberculatusBeurlen, 1958 is transferred herein into Hepatus as H. beurleni nomen novum.

Geological setting

The Pirabas Formation (Maury, 1925) consists of mixed siliciclastic-carbonate deposits associated with shallow-water coastal settings (grainstone and consolidated packstone, stratified wackestone to packstone, and laminated mudstone) (Rossetti et al., 2013; Aguilera et al., 2022). It is a young Cenozoic formation, deposited on the Brazilian equatorial platform of late early (~16 Ma) to late middle (13-12 Ma to younger) Miocene Pirabas Formation (Gomes et al., 2023) (Fig. 1). The Pirabas Formation is representative of a variety of shallow water paleoenvironments, including surf zones, tidal flats, coastal lagoons, and tidal inlet environments (Góes et al., 1990; Rossetti & Góes, 2004; Rossetti et al., 2013; Borges, 2016; Aguilera et al., 2020; Lima et al., 2020). The Pirabas Formation has been divided into two groups (α, β) and three main facies (α1, α2, and β) (Aguilera et al., 2022). The studied outcrops of Ponta da Fazenda in the Ilha de Fortaleza and Colônia Pedro Teixeira in Capanema are included in facies α1 (characterized by echinoderm-bryozoan packstone to rudstone rich in mollusks) that suggest an outer inner platform exposed to oceanic influence.

Figure 1
(A, B) Location map of Pirabas Formation, Pará, Brazil. (A) Early to middle Miocene paleogeographic reconstruction of Pirabas Formation (23-13 Ma). (B) Pirabas Formation at Colônia Pedro Teixeira and Fazenda outcrops. Paleogeographic reconstruction and location map primarily based on Aguilera et al. (2016, 2022).

MATERIAL AND METHODS

The material studied is deposited in the invertebrate paleontology collections of the Museu de Ciências da Terra (MCTer/SGB-CPRM), and Museu Nacional da Universidade Federal do Rio de Janeiro (MNRJ). The Hepatus beurleni nomen nov. is a single fossil specimen, with part and counterpart (Fig. 2). The redescription of Miohepatus amazonicus gen. nov., comb. nov. is based on the single type specimen from the Pirabas Formation. Descriptions, drawings, and photographs were made using a stereomicroscope Nikon SMZ800N equipped with camera lucida and a Leica EZ4W, both with digital camera attached.

Figure 2
(A-E) Hepatus beurleni nomen nov.. (A) Hepatus beurleni original label as Cyclocancer tuberculatusBeurlen, 1958. (B) Holotype MNRJ-4617-I, part and counterpart. (C) Cyclocancer tuberculatusBeurlen, 1958 original photograph and illustration. (D) and (E) dorsal and frontal view. Scale: 20 mm.

The Brachyura classification of Ng et al. (2008) was mostly used in suprafamilial rankings. Infrafamilial rankings mostly follow De Grave et al. (2009) and Schweitzer et al. (2010). We also used the following abbreviations: cl, carapace length; cw, carapace width (taken at the widest point); coll., collector or collected by; † indicates taxa with recent and fossil representants; †† indicates taxa exclusively known from fossils; L/W, length/width ratio.

This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN.

RESULTS

Systematic Paleontology

Brachyura Latreille, 1802

Eubrachyura de Saint Laurent, 1980

Aethroidea Dana, 1851

Aethridae Dana, 1851

Type genus:Aethra Latreille, in Cuvier, 1816.

Other genera:ActaeomorphaMiers, 1877; Drachiella† Guinot in Serène & Soh, 1976; Eohepatella†† Beschin & De Angeli, 2017; Eriosachila†† Blow & Manning, 1996; Hepatella† Smith in Verril, 1869; Hepatiscus†† Bittner, 1875; Hepatus Latreille, in Bosc, 1801; IlerdapatiscusArtal & Van Bakel, 2018; Mainhepatiscus†† De Angeli & Beschin, 1999; Matutites†† Blow & Manning, 1996; Miohepatus gen. nov.; Osachila† Stimpson, 1871; Politohepatiscus zorziniBeschin, Busulini, Fornaciari, Papazzoni & Tessier, 2018; Prehepatus†† Rathbun, 1935; Priabonella†† Beschin, De Angeli, Checchi & Mietto, 2006; Pseudohepatiscus†† Blow & Manning, 1996; Sakaila Manning & Holthuis, 1981.

Remarks: Aethridae currently embraces fossil and extant members which share similarities proposed by Guinot (1966, 1967) and they are currently positioned in Aethroidea (Ng et al., 2008; De Grave et al., 2009). Fossil specimens generally are attributed to this family by the general carapace outline and ornamentation, which is octagonal to subquadrate, wider than long; antero- and posterolateral margins clearly demarcated; and anterolateral margins entire to cristate, may be expanded to form subclypeiform structure (Davie, 2002).

Genus Hepatus Latreille, in Bosc, 1801

Hepatus Latreille, in Bosc, 1801: 55, 56 [type species: Calappa angustataFabricius, 1798, by monotypy, junior synonym of Hepatus pudibundus (Herbst, 1785)].

HepatusLatreille, 1802: 22 [type species: Calappa angustataFabricius, 1798, by monotypy, junior synonym of Hepatus pudibundus (Herbst, 1785)].

HepatulusFowler, 1912: 590 [type species: Hepatus fasciatusLatreille, 1803; unnecessary replacement name for Hepatus Latreille, in Bosc, 1801].

HepatoidesBalss, 1957: 1612 [unnecessary replacement name for Hepatus Latreille, in Bosc, 1801].

CyclocancerBeurlen, 1958: 15 [type species Cyclocancer tuberculatusBeurlen, 1958, by monotypy].

Included species:Hepatus beurleni†† nomen nov.; H. biformis†† Collins & Todd in Todd & Collins, 2005; H. bottomsi†† Blow, 2003; H. chiliensis† H. Milne-Edwards, 1837; H. epheliticus (Linnaeus, 1763); H. gronoviiHolthuis, 1959; H. guraboensis†† Collins in Collins et al., 2009; H. kossmanniNeumann, 1878; H. lineatus† Rathbun, 1898; H. lineatinus†† Collins & Todd in Todd & Collins, 2005; H. nodosus†† Collins & Morris, 1976; H. pauli†† Collins, Garvie & Mellish, 2014; H. praecox†† Collins, Donovan & Dixon, 1996; H. pudibundus (Herbst, 1785); H. scaberHolthuis, 1959; H. spinimarginatus†† Feldmann, Schweitzer & Encinas, 2005.

Remarks: The authorship of Hepatus has been generally attributed to Latreille (1802: 22) (e.g.,Desmarest & Bosc, 1830; Holthuis, 1959; Glaessner, 1969; Ng et al., 2008; De Grave et al., 2009; Schweitzer et al., 2010). Hepatus was recently attributed to Bosc (1801: 55, 56) (see WoRMS, 2023). Bosc (1801: 29, 48) makes it clear that he used some results of Latreille’s manuscript - which was in preparation. More importantly, Bosc (1801: 29, 48) attributed authorship to Latreille (see Opinion 1133: 97 by C. Dupuis, Case 1879, BZN 1979: 97 for TalitrusBosc, 1802). According to the ICZN (1999: Art. 50.1.1) “if it is clear from the contents that some person other than an author of the work is alone responsible both for the name or act and for satisfying the criteria of availability other than actual publication, then that other person is the author of the name or act”. Thus, the authorship for Hepatus should therefore be Latreille, in Bosc (1801: 55, 56) (type species Calappa angustataFabricius, 1798, by monotypy). Desmarest & Bosc (1830: 42, 209), in the second edition of Bosc (1801, 1802) attributed the authorship of Hepatus to Latreille. Here we follow Dupuis (Opinion 1133: 97) (for more details see ICZN Case 1879; Bousfield & Holthuis, 1969; Dupuis, 1975; Holthuis, 1975).

Hepatus beurleni nomen nov.

(Fig. 2A-E)

https://zoobank.org/78AF63CA-91E9-499D-B3E8-ED1BE02E7A26

Cyclocancer tuberculatusBeurlen, 1958: 15 [type locality: Pirabas Formation, Brazil, Pará holotype (MNRJ-4617-I). - Schweitzer et al., 2010: 102; Távora et al., 2010b: 214; Luque et al., 2017: 12.

Type material: Holotype MNRJ-4617-I, part and counterpart, early-middle Miocene, Pirabas Formation, Brazil, Pará, Ilha de Fortaleza, Ponta da Fazenda, C.S. Ferreira e O.R. da Cunha coll.

Occurrence and age: Brazil, Pará, Ilha de Fortaleza, Ponta da Fazenda, Pirabas Formation, early-middle Miocene.

Etymology: The specific epithet of the new name honors Karl Beurlen, a prominent German paleontologist who worked in the Instituto Geológico Brasileiro and described this species.

Diagnose: Carapace ovoid, wider than long (cw 38 mm, cl 28 mm; W/L ratio 1.4). Rostrum narrow, orbits small. Anterior margins convex, strongly crenated, lobes extending to anterior posterolateral margins. Posterolateral margins short, slightly concave, upturned anteriorly, crenulated posteriorly, with a large knob-like lobe separating it from the posterior margin. Posterior margin narrow, straight, crenulated. Greatest width near midlength. Carapace with 8 protuberances on gastric, cardiac, and branchial regions; mesogastric region higher than the remaining carapace regions; protuberances surmounted with a cluster of small granules. Mesobranchial protuberance oblong, transversal separated from gastric and cardiac regions by a shallow groove.

Remarks:Cyclocancer tuberculatusBeurlen, 1958 is herein transferred into Hepatus as H. beurleni nomen nov., a new species name for H. tuberculatus (Beurlen, 1958) as it became a junior homonym of H. tuberculatusSaussure (1858), a junior synonym of Hepatus pudibundus (Herbst, 1785).

Janssen & Müller (1984) were the first to indicate that Cyclocancer tuberculatus is more related to Hepatus than to any other Cancroidea (superfamily where CyclocancerBeurlen, 1958 was originally included). Salva & Feldmann (2001) also questioned the position of Cyclocancer in Cancridae, although not citing its resemblance with Hepatus. Feldmann et al. (2005) mentioned that when the front of the carapace is missing, as is the case of the holotype of Hepatus beurleni nomen nov., it is difficult to distinguish Hepatus from the cancrid Metacarcinus A. Milne-Edwards, 1862; however, the presence of distinct elevations of the carapace in Hepatus makes possible to differentiate it from Metacarcinus.

Hepatus beurleninomen nov., has the diagnostic characters of Hepatus (as understood by Feldmann et al., 2005: 433), such as carapace much broader than long, broadly obovate, vaulted; crenated anterolateral margins; posterolateral margin concave; and posterior margin narrow.

Among the fossil species, H. lineatinus and H. biformis from the late Pliocene of Panama and Miocene-Pliocene of Panama and Costa Rica, respectively, differ from H. beurleni nomen nov. (characters within brackets) by having the carapace anterolateral margins tridenticulate (vs anterolateral margins with short lobes of about the same size). Hepatus guraboensis Collins, in Collins et al., 2009, from the Miocene of the Dominican Republic stands apart by having the anterolateral margins with short triangular spines increasing in size posteriorly (vs anterolateral margins strongly crenated). Hepatus nodosusCollins & Morris, 1976, from the Miocene of Trinidad differs by having the anterolateral margins divided into four blunt teeth (vs anterolateral margins strongly crenated, extending to anterior posterolateral margins). Hepatus bottomsiBlow, 2003, from the Pliocene of Virginia, United States, can be distinguished by the anterolateral margins of the carapace consisting of 40 blunt denticles situated from 12-13 or more bidentate or tridentate teeth (vs anterolateral margins consisting of simple lobes). Hepatus pauliCollins, Garvie & Mellish, 2014, from the Pleistocene of Texas, United States, differs by having 12-13 tridentate spines on the anterolateral margins (vs anterolateral margins consisting of simple lobes). Hepatus spinimarginatusFeldmann, Schweitzer & Encinas, 2005, from the Miocene of Chile differs by having the posterolateral margins rimmed by very finely beaded elevation (vs posterolateral margin strongly crenulated).

The following set of characters differentiate H. beurleni nomen nov., from all of its extant congeners: 1) posterior margin narrow, crenulated, almost straight; 2) posterolateral margin strongly crenulated with a large lobe between the posterolateral and posterior margins; and 3) anterolateral margin crenated, with short lobes of about the same size.

The holotype MNRJ-4617-I had been missing for decades and was considered lost, but was recently found together in the MCTer collection, probably due to an old forgotten loan, posteriorly returned to the institution of origin (Museu Nacional do Rio de Janeiro/MNRJ). This fact allowed the specimen to escape the fire that hit the Museu Nacional in 2018 and destroyed a large part of the paleontological collections, which gives it even greater importance.

Genus Miohepatus new genus

https://zoobank.org/76AE969B-27E0-4112-85A5-DDE9B2F5A503

Type species:Hepatella amazonicaBeurlen, 1958, by present designation. Gender masculine.

Included species:Miohepatus amazonicusgen. nov., comb. nov. (Beurlen, 1958) [Hepatella]; M. peruvianus gen. nov., comb. nov. (Rathbun, 1933) [Hepatella].

Diagnosis: As for the type species.

Occurrence: Brazil, Pará, Colônia Pedro Teixeira, Capanema, Pirabas Formation, early-middle Miocene.

Etymology: The generic name is a combination of the generic name Hepatus and Miocene in allusion to the age of the type species.

Remarks:Miohepatusgen. nov. is erected here to accommodate Hepatella amazonicaBeurlen, 1958 from the early-middle Miocene, Pirabas Formation, Pará, Brazil, originally assigned to the recent genus Hepatella Smith, in Verril, 1869 (type species Hepatella amica Smith, in Verrill, 1869) from the tropical eastern Pacific Beurlen (1958) wrongly justified the assignment of his species to Hepatella by: the 1) presence of a carapace with remarkable elevations in the gastric, cardiac and branchial regions; 2) prominent rostrum; and 3) concave anterolateral margins. However, he was correct in suggesting a close relationship between Hepatella amazonica and H. peruviana, a recent species from the east Pacific coast. Accordingly, Hepatella peruviana is transferred along with H. amazonica to Miohepatus gen. nov.

Miohepatusgen. nov., as revealed by both species, can be differentiated from Hepatella (characters for H. amica within brackets) by the: 1) carapace octagonal, slightly longer than wide, 1.1 width/length ratio (vs carapace subrectangular, remarkable longer than wide, 1.7 width/length ratio); 2) rostrum extending well beyond orbits (vs rostrum slightly exceeding orbits); and 3) posterolateral margins strongly concave (vs posterolateral margins nearly parallel).

The new genus differs from Hepatus in that the carapace dorsal surface is uneven, vaulted in the central region, with eight protuberances in the cardiac, gastric and branchial regions (vs carapace dorsal surface almost smooth in adults, convex, carapace regions poorly differentiated in Hepatus).

The rostrum bilobed with a median sulcus in Miohepatus gen. nov. is similar to that of OsachilaStimpson, 1871 (type species Osachila tuberosaStimpson, 1871), MainhepatiscusDe Angeli & Beschin, 1999 (type species M. zannatoiDe Angeli & Beschin, 1999), PolitohepatiscusBeschin, Busulini, Fornaciari, Papazzoni & Tessier, 2018 (type species P. zorziniBeschin, Busulini, Fornaciari, Papazzoni & Tessier, 2018), and PriabonellaBeschin, De Angeli, Checchi & Mietto, 2006 (type species P. violattiBeschin, De Angeli, Checchi & Mietto, 2006). However, Miohepatus gen. nov. can be separated from Osachila by having a posterolateral margin of carapace crenulated (vs posterolateral margin of carapace nearly straight, obtuse, rugose, and armed with two of three tuberculliform teeth in Osachila); in having a slightly convex posterior margin of the carapace (vs posterior margin of carapace bilobed in Mainhepatiscus); by the dorsal surface of the carapace ornamented with protuberances (vs carapace dorsal surface smooth in Politohepatiscus); and by having a carapace wider than long (vs carapace slightly longer than wide in Priabonella).

Drachiela Guinot, in Serène & Soh, 1976 [type species D. sculpta (Haswell, 1879)], EriosachilaBlow & Manning, 1996 [type species E. petitiBlow & Manning, 1996], IlerdapatiscusArtal & Van Bakel, 2018 [type species I. guardiae Artal & Van Bakel, 2018], MatutitesBlow & Manning, 1996 [type species M. anthonyaeBlow & Manning, 1996], PseudohepatiscusBlow & Manning, 1996 [type species P. marinoiBlow & Manning, 1996], all stand apart from Miohepatus gen. nov. by having the rostrum slightly exceeding the orbits (vs rostrum distinctly exceeding the orbits in Miohepatus gen. nov.).

The new genus differs from Aethra Latreille, in Cuvier, 1816 [type species A. scruposa (Linnaeus, 1764)] by having an octagonal carapace and concave posterolateral margin (vs carapace elliptical and posterolateral margin slightly convex in Aethra). It differs from ActaeomorphaMiers, 1877 [type species A. erosaMiers, 1877] in having three tubercles on the gastric region (vs two large elevations on the gastric region, behind each orbit in Actaeomorpha); and differs from SakailaManning & Holthuis, 1981 [type species S. africanaManning & Holthuis, 1981] by having a crenulated posterolateral margins and carapace dorsal surface with eight protuberances (vs posterolateral margins with eight distinct teeth and carapace dorsal surface with six major protuberances in Sakaila).

The new genus shares with HepatiscusBittner, 1875 (type species H. poverelliVía, 1959) the presence of eight protuberances on the carapace dorsal surface, bilobed rostrum, and concave posterolateral margins. However, Hepatiscus stands apart in having the anterolateral carapace margins with six lobes instead of the anterolateral carapace margins crenated, with 12 small lobes as in Miohepatus gen. nov. Miohepatus gen. nov. differs from EohepatellaBeschin & De Angeli, 2017 [type species E. planaBeschin & De Angeli, 2017] by having an octagonal carapace and the presence of eight protuberances on the carapace dorsal surface (vs oval carapace and six protuberances on the carapace dorsal surface in Eohepatella). PrehepatusRathbun, 1935 is known only by chelae (see Schweitzer et al., 2006).

Miohepatus amazonicus ( Beurlen, 1958 ) new combination

(Fig. 3A-E)

https://zoobank.org/DD1D6AF9-E23C-4B95-9F30-1C8A3EFC96A6

Figure 3
(A-E) Miohepatus amazonicum new genus, new combination. (A) Holotype MCT.I.4458. (B) Miohepatus amazonicus original label as Hepatella amazonicaBeurlen, 1958. (C) Hepatella amazonicaBeurlen, 1958 original photograph and illustration. (D) dorsal view; red arrow: rostrum; white arrows: concave posterolateral margins. (E) left frontolateral view; red arrows: bilobed rostrum; yellow arrow: left orbit; green arrow: left anterolateral margin. Scale: 10 mm.

Hepatella amazonicaBeurlen, 1958: 3, pl. I fig. 1, pl. III, fig. 1 [type locality: Pirabas Formation, Brazil, Pará; holotype in MCTer/SGB-CPRM]. - Collins et al., 2009: 95; Schweitzer et al., 2010: 85; Távora et al., 2010b: 212; Távora et al., 2010a: 50, 54; Aguilera & Páes, 2012: 35, tab. 1; Aguilera et al., 2013: 113; Luque et al., 2017: 8.

Hepatella amazonica - Vega et al., 2009: 56, pl. 1, figs. 15-18; Aguilera et al., 2010: 109, fig. 6.2.15-6.2.19 [not Hepatella amazonicaBeurlen, 1958].

Hepatella amazzonica - Beschin & De Angeli, 2017: 25 [misspelling].

Material examined: Holotype MCT-I 4458, early-middle Miocene, Pirabas Formation, Brazil, Pará, Capanema, Colônia Pedro Teixeira, F. Ackermann coll.

Occurrence and age: Brazil, Pará, Colônia Pedro Teixeira, Capanema, Pirabas Formation, early-middle Miocene.

Redescription: Carapace octagonal, wider than long (cw 18.5 mm, cl 15 mm; W/L ratio 1.2); greatest width at the posterolateral corner. Carapace dorsal surface uneven, vaulted centrally, concave marginally, anterior surface punctate, regions well defined. Cervical groove well defined. Rostrum partially preserved, bilobed, extending well beyond orbits; lobes separated medially by short cleft. Left orbit well preserved, small, circular, well visible in dorsal view. Carapace with eight distinct protuberances on cardiac, branchial, and gastric regions. Cardiac, mesobranchial and metagastric protuberances slightly higher than metabranchial and protogastric. Each protuberance bears a single large central granule, surrounded by some smaller ones. Anterolateral margins crenated, with 12 small rounded lobes. Posterolateral margins distinctly concave, arcuate, crenulated; shorter than anterolateral margins. Posterior margin narrow, crenulated, slightly convex.

Remarks:Miohepatus amazonicusgen. nov., comb. nov. is closely related to M. peruvianus gen. nov., comb. nov. However, they differ in the presence of a small protuberance on each metabranchial region in M. amazonicus gen. nov., comb. nov., whereas the metabranchial protuberance is absent in M. peruvianus.

Miohepatus amazonicusgen. nov., comb. nov. differs from Hepatus beurleni nomen nov., comb. nov. [characters for H. beurleni within brackets] by having the carapace octagonal (vs carapace ovoid) and the carapace W/L ratio of 1.2 (vs carapace W/L ratio of 1.4). In addition, H. beurleni has a large knob-like lobe separating the posterolateral margins from the posterior margin, which is absent in M. amazonicus.

Vega et al. (2009) mentioned the occurrence of M. amazonicusgen. nov., comb. nov. (as H. amazonica) from the lower Miocene of the Chiapas, Mexico. However, from the figures provided by Vega et al. (2009) it seems that their material is not attributable to M. amazonicus. The main differences are in the curvature of the posterolateral margin of the carapace, much more accentuated in M. amazonicus gen. nov., comb. nov. (Fig. 2) (vs almost straight in the specimen figured by Vega et al., 2009: pl. 1, figs. 15-18); and in the surface of the carapace with six main protuberances and two small protuberances on the metabranchial region in M. amazonicus gen. nov., comb. nov. (Fig. 2) (vs eight equally inflated elevations on the carapace in the material studied by Vega et al., 2009: pl. 1, figs. 15-18). Actually, the general carapace shape and the short bilobed rostrum of specimens from Mexico are similar to that of Eriosachila rather than to M. amazonicus gen. nov., comb. nov., although with quite different posterolateral margin (Vega et al., 2009; pl. 1). A reexamination of the material from Vega et al. (2009) is needed to clarify its taxonomic assignment.

ACKNOWLEDGEMENTS:

We would like to thank Sandro Scheffler for his assistance in the invertebrate paleontological collection of Museu Nacional (UFRJ). We kindly thank the anonymous reviewers and Associated Editor Marcos Tavares for their constructive comments to the manuscript.

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• FUNDING INFORMATION:

  This work is part of an effort from the Fundação Cearense de Apoio ao Desenvolvimento Científico e Tecnológico (FUNCAP) for the study of paleontology in the Araripe Basin (Edital 08/2021). RCS was financed by FAPERJ #E-26/210.294-2021. DL, AP, and WS were financed by FUNCAP with the following grants #PV1-0187-00019.01.00/21 to DL; #BP3-0139-00166.01.00/18, BP4-00172-00173.01.00/20 to APP; and #PV1-0187-00033.01.00/21 and #6647309/2017 to WS. Financiadora de Estudos e Projetos (FINEP) provided equipment. Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (grants #315185/2020-1 to WS and 404937/2018-7 to OA, and fellowships 304693/2021-9 to OA). Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) (fellowships E-26/201.035/2021 to OA). Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001.

Edited by

Edited by:

Marcos Domingos Siqueira Tavares

Publication Dates

• Publication in this collection
  21 Apr 2023
• Date of issue
  2023

History

• Received
  24 Nov 2022
• Accepted
  13 Mar 2023
• Published
  06 Apr 2023