A new passalid fossil (Insecta: Coleoptera) from the Santana Formation (Crato member, Lower Cretaceous), Araripe Basin, NE Brazil: Paleoecological and paleobiogeographic implications
Graphical abstract
Introduction
The superfamily Scarabaeoidea includes at least 28,000 described species (Browne and Scholtz, 1995; Lawrence et al., 2011; Bai et al., 2012), comprising one of the most diverse groups of polyphagous beetles. The 14 extinct and extant families include two families of ichnofossils (Bouchard et al., 2011), of which Passalidae Leach, 1815 can be recognized by its body outline (elongated and dorsoventrally depressed), prothorax separated from the elytra by a distinct narrowing, and mentum deeply emarginated apically (Schuster, 2002).
The family Passalidae comprises 1000 species, with widely differing richness between continents. Its members occur in many environments, including dry desert, wetlands, intertropical forest zones, and tropical forests; most species inhabit tropical forests (Reyes-Castillo, 1970; Boucher, 2006). The monophyly of the family is strongly supported by phylogenetic studies (see Scholtz and Grebennikov, 2005), and the group includes two subfamilies and seven tribes belonging to the clade Scarabaeoidea, close to Lucanidae and Diphyllostomatidae, which would be the earliest radiation in the superfamily (Scholtz and Chown, 1995; Boucher, 2006; Bai et al., 2012). However, Smith et al. (2006) placed Passalidae close to Geotrupidae + Pleocomidae, which would be the earliest radiation within Scarabaeoidea. This topology was questioned by Ahrens et al. (2014), who found Passalidae closer to Bolboceratidae. Furthermore, the hypothesis that the Passalidae are highly modified scarabs was recently questioned by McKenna et al. (2015), after the inclusion of Pleocomidae.
Passalid species can be used in ecological studies as potential environmental indicators because they are primary decomposers, i.e., adults and larvae live in and feed on decaying logs (Schuster, 1985; Schuster et al., 2000; Boucher, 2006; Mattos and Mermudes, 2014). The ethology of passalids is peculiar within the order Coleoptera: adults and immatures live in family groups with sub-social behavior, communicating by sounds, and the adults care for their offspring (Ohaus, 1900; Heymons, 1929; Reyes-Castillo, 1975; Schuster, 1975, 1983; Reyes-Castillo and Halffter, 1984).
The Passalidae comprises two subfamilies. The more-diverse subfamily Passalinae includes five tribes: Passalini Leach, 1815 (6 genera, 164 species), Proculini Kaup, 1868 (19 genera, 214 species), Solenocyclini Kaup, 1871 (9 genera, 35 species); Leptaulacini Kaup, 1871 (2 genera, 18 species), and Macrolinini Kaup, 1871 (22 genera, 46 species). The subfamily Aulacocyclinae is less diverse, with two tribes, Aulacocyclini (3 genera, 30 species) and Ceracupini (2 genera, 5 species), according to Hincks andand Dibb (1935; 1958), Boucher (2006), and Boucher et al. (2017). A third aulacocycline tribe, Ceracyclini (2 genera, 6 species) was proposed by Boucher et al. (2017), who transferred four species of Cylindrocaulus from Aulacocyclini.
Passalid fossils are extremely rare compared to members of other families in Scarabaeoidea. Up to the present, the approximately 231 described representatives of the superfamily included only four species of passalids (Krell, 2000, 2006; Bai et al., 2012; Boucher et al., 2017). The reason for this rarity is that passalid individuals inhabit the inner parts of fallen trunks, rendering them difficult to access when mineralized inside the logs (Cockerell, 1927; Reyes-Castillo, 1977; Boucher, 2006). Recently, Boucher et al. (2016) proposed a new family, Passalopalpidae, very close to Passalidae, represented only by species from the Cretaceous amber of Myanmar (earliest Cenomanian, 100 Ma).
The first description of a fossil species of Passalidae was contributed by Cockerell (1927), with Passalus indormitus from the Inferior Division of the John Davis Series of the late Oligocene (about 25 Ma), Crooked River Basin in Oregon, USA. Subsequently, the specimen was systematically reinterpreted twice by Reyes-Castillo (1970; 1977); in the last interpretation, Reyes-Castillo considered this species to be a probable member of the tribe Passalini. The second known member of Passalidae, Serrulus sinicus, was described by Hong (1983, 1985), and is still considered incertae sedis in the family; this species is from Shandong province (situated south of Beijing) of the Miocene Shanwang Formation (about 16 Ma). Later, Zhang (1989) identified this specimen as a member of Macrolinus Kaup (1868). Boucher (2006) considered this specimen, as described by Zhang, to be a member of Passalidae (Passalinae) because of its dorsoventrally flattened body, and placed it in Macrolinini Kaup (1871).
Subsequently, Boucher et al. (2017) described two sympatric fossil passalid species from the Cretaceous amber from the Hukawng Valley, Myanmar (about 100 Ma), Ceracyclus lotus and Ceracyclus jirouxi. These genera belong to Ceracyclini, as does Cylindrocaulus.
A fossil specimen of Passalidae from the Santana Formation, Araripe Basin, Brazil is described here, as a new genus and species of Passalinae. This specimen is the first record for the family in a Crato Litho-stratigraphic unit, and the oldest fossil of the family Passalidae, from the Aptian (about 112–114 Ma, Lower Cretaceous).
Section snippets
Geological setting
The Araripe Basin (Fig. 1, Fig. 2) is located in northeastern Brazil, central part of the Borborema Plateau, and occupies parts of the states of Piauí, Pernambuco, and Ceará. The basin covers an area larger than 8000 Km2 (Portela et al., 2014) and contains an elongate geomorphological feature oriented E-W, the Araripe Plateau.
The Araripe Basin is the largest inland basin of northeastern Brazil, with the most complex geological history (Assine, 2007), and consists of Paleozoic and Mesozoic
Repository
The specimen (MN 7732-I) described in this study is deposited at the paleoinvertebrate collection of the Departamento de Geologia e Paleontologia of the Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MN/UFRJ). In spite of the fire that destroyed most of the main museum complex in September 2018, this specimen resisted the fire and was completely recovered on 13 November 2018 by Sandro Scheffler. The most serious problem with the specimen is the presence of a
Systematic paleontology
Insecta Linnaeus, 1758
Order Coleoptera Linnaeus, 1758
Suborder Polyphaga Emery, 1886
Superfamily Scarabaeoidea Latreille, 1802
Family Passalidae Leach, 1815
Subfamily Passalinae Leach, 1815
Tribe Protopassalini trib. nov.
Type genus: Protopassalus gen. nov. described herein.
Diagnosis. Clypeus narrow and transverse (Fig. 3C, D), distinct at middle of anterior margin of head, exposed dorso-anterior with labrum, this developed and rectangular. Evidence of frontal ridges (Fig. 3C). Eyes subrounded,
Justification of placement in Passalidae
The assignment to Passalidae was based on clear indications, comprising the body size of 12 mm, subcylindrical body, and the following characteristics of the subfamily Passalinae. Head: 1) transverse frons; 2) deep depression on side of each eye; 3) asymmetric elevations on the frons; 4) anterior margin of the head evidently straight; 5) labrum developed; 6) clypeus exposed antero-dorsally; 7) evidence of frontal ridges; 8) central tubercle present, but vestigial, probably weathered; 9)
Concluding remarks
The fossil material is preserved three-dimensionally and provides the first view of anatomical structures by original tissues replaced by iron minerals, including the conspicuous characteristics of: clypeus narrow and transverse, exposed dorso-anteriorly; labrum developed, rectangular with subrounded eyes, anteriorly not emarginate, lacking ocular canthus; prothorax with anterior sides subrounded, feebly narrowed at posterior third, posterior margin sinuous; and elytra with row of coarse
Acknowledgments
JRMM was supported through the following grants: CNPq 306105/2016-0 and 311679/2019-6, Faperj 110.040/2014, 211.522/2016, and PROTAX CNPQ/CAPES 440479/2015-0. SMS was supported through a grant from Faperj E-26/200.110/2019 and CNPQ/MCTIC/CAPES/FNDCT - PROANTAR 442765/2018-5. The authors thank Dionizio Angelo de Moura Jr. for help in preparing the specimen. We thank Marcelo de Araújo Carvalho (Federal University of Rio de Janeiro), Luiz Felipe Lima da Silveira (Western Carolina University),
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Cretaceous beetles of the Jinju Formation (Coleoptera): An overview of the Jinju Formation, its coleopteran diversity, and past and future research
2024, Journal of Asia-Pacific EntomologyA palaeoecological analysis of the Cretaceous (Aptian) insect fauna of the Crato Formation, Brazil
2024, Palaeogeography, Palaeoclimatology, PalaeoecologyAptian ostracods from the Santana Group, Araripe Basin, Brazil
2022, Revue de MicropaleontologieCitation Excerpt :spp. 201–218 and up to current date, several of these have been assigned to the genera Damonella Anderson, 1966 and Pattersoncypris Bate, 1972. The Araripe Basin in northeast Brazil (Fig. 1) is known by the well preserved and abundant Cretaceous fossils of the Santana Group, recording such macro- and microfossils, as dinosaurs and pterosaurs (Kellner and Campos, 2002; Sayão et al., 2020a,b; Bantim et al., 2021), foraminifers and ostracods (Melo et al., 2020; Araripe et al., 2021), plants (Lima et al., 2012; Mohr et al., 2007; Sucerquia et al., 2015), terrestrial and marine palynomorphs (Arai and Assine, 2020; Goldberg et al., 2019; Teixeira et al., 2017), insects (Silva and Arruda, 1976; Dos Santos et al., 2021) and fishes (Brito and Yabumoto, 2011) among other groups. The Santana Group consists of, the Barbalha, Crato, Ipubi and Romualdo formations from base to top (Assine et al., 2014), with the Crato and Romualdo formations corresponding to two Cretaceous Konservat-Laggerstätten.
The role of microbial mats in the exquisite preservation of Aptian insect fossils from the Crato Lagerstätte, Brazil
2022, Cretaceous ResearchCitation Excerpt :However, Catto et al. (2016), Warren et al. (2017) and Varejão et al. (2019) presented several lines of evidence in favor of microbial influence on the origin of the laminated limestones, suggesting that the metabolic activity of these microorganisms played a significant role in their formation. The insect fossils recorded in the lithographic limestones of the Crato Formation have been generally analyzed as far as their taxonomy and paleobiology is concerned, mainly due to the preservation of external and internal features, including muscles, organs and other soft tissues (Barling et al., 2015, 2020, 2021; Osés et al., 2016; Bezerra et al., 2018, 2020, 2021; Dias et al., 2019; Dias and Carvalho, 2020; Moura-Júnior et al., 2020; Nel and Pouillon, 2020; Pouillon and Nel, 2020; Prado et al., 2020; Ribeiro et al., 2021b; Santos et al., 2021). Experimental studies of Iniesto et al. (2020) on the decay of insect larvae and its relationship with microbial mats allowed a comparative analysis between textural aspects of the larvae carcasses and the insect fossils’ microfabric from the Crato Formation.
Paranoikidae Zamboni, Martins-Neto & Popov, 2002, a junior synonym of Belostomatidae Leach, 1815 (Hemiptera: Heteroptera): Redescription of a giant water bug from Crato Formation, Lower Cretaceous of Brazil
2021, Cretaceous ResearchCitation Excerpt :The putative two great lineages may have appeared quickly, about 130 Ma ago, when the Gondwana breakup had already started but the continents were still very close to one another, allowing migrations through narrow seas. This hypothesis, which has to be thoroughly tested by means of formal phylogenetic and biogeographic analyses, seems to demonstrate that the distribution of Lethocerus does not reflect the major vicariant events that occurred in the Cretaceous, which are apparently related to the distribution of groups that have limited dispersal capacity (see Santos et al., 2021 for Coleoptera, Passalidae, and Boyer et al., 2007 and Boyer and Giribet, 2007 for Opiliones, Petallidae). The large size and great dispersal capacity of Lethocerus species, combined with their generalist habits and great resistance to stressful environments, may be connected to the morphological stability in deep time (see below).
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He is a coauthor that passed away during the preparation of the manuscript.