The Crato Fossil Beds of Brazil: Window into an Ancient World
Abstract
This beautifully illustrated 2007 volume describes the entire flora and fauna of the famous Lower Cretaceous Crato Formation of Brazil - one of the world's most important fossil deposits, exhibiting exceptional preservation. A wide range of invertebrates and vertebrates are covered, including extended sections on pterosaurs and insects. Two chapters are devoted to plants. Many of the chapters include descriptions of new species and re-descriptions and appraisals of taxa published in obscure places, rendering them available to a wider audience. Fossil descriptions are supported by detailed explanations of the geological history of the deposit and its tectonic setting. Drawing on expertise from around the world and specimens from the most important museum collections, this book forms an essential reference for researchers and enthusiasts with an interest in Mesozoic fossils.
... The bond place run of the breastbone in spiders plays a relevant role in their travel feed and general physical coordination. Drawing from various sources here's an in-depth exploration of this aspect: Anatomical Connectivity: Muscles responsible for important physiological Roles originate from the prosoma and attach to specific regions of the sternum via tendons or apodemes (Martill et al., 2007;Minh et al., 2013). ...
... These attachment points provide a stable for muscle fibers facilitating robust and coordinated movements. Coordination of movement: bond to the breastbone allows for right and fast movements important for search web-constructing and apologetic behaviors (Martill et al., 2007) away catching and quiet these muscles spiders do adroit actions over aggregate appendages. Muscle Roleality: Sternum-attached muscles play decisive roles in extending and retracting legs manipulating prey and constructing silk strands (Menon, 2007). ...
... Muscle Roleality: Sternum-attached muscles play decisive roles in extending and retracting legs manipulating prey and constructing silk strands (Menon, 2007). This reality enables spiders to perform intricate tasks with Productivity and precision Biomechanical Productivity: The sternum's role as an attachment site improves biomechanical productivity by minimizing energy expenditure and maximizing muscular output (Martill et al., 2007). End propinquity to the eye of the lot reduces automatic disadvantages allowing spiders to get sound movements with nominal effort adjustations to ecologic niches: ...
This study investigates the coxosternal or ventral sternal region, in scorpions and spiders. We collected 10 specimens of Scorpions from provinces Baghdad in region Abo Kreab and Salahuddin provinces in Tikrit then 7 specimens of Spiders were collected from variant regions in provinces Baghdad and various genera and families were collected and processed for examination. The aim was to compare the morphology of the coxosternal between these two arachnid groups and analyze the underlying functional and evolutionary implications of any structural variations. So the result of this study on coxosternal region of scorpions in the Family: Buthidae like Androctonus crassicauda, is not wide the majority of members of this family have a triangular cut and the female genital cover is divided, coxosternal region in Orthochirus scrobiculosus do not extend forward and do not form lobes, edges of the scales grainy and it is finely grained the majority of individuals have a triangular cut, Mesobuthus eupeus the ventral side is shiny and wide with strong hairs and the sternum is triangular, coxosternal region in Family: Scorpionidae like Scorpio Maurus in the ventral side do not have lobes extending forward, the sternum is pentagonal. But in Family: Salticidae of spiders the coxosternal region in Hasarius adansoni, are oval shape sternum or variable in Thyene imperialis, and Evarcha seyun, are circle shape sternum or variable, coxosternal region in Neoscona subfusca, family Araniedae are heart-shaped or triangular. The expected results include identifying distinct coxosternal shapes in scorpions compared to spiders. These variations might be linked to specific Roles like locomotion and internal organ protection. The determined geomorphology gets to render Understandings into the evolutionary relationships betwixt scorpions and spiders. This comparative analysis is expected to Add to a better understanding of how the coxosternal plays a role in the biology of scorpions and spiders. The known variations get bid important information along the development and practical adjustations inside these arachnids
... The Crato Formation (Araripe Basin, northeastern Brazil; Assine et al., 2014;Fambrini et al., 2020;Varejão et al., 2021) is renowned worldwide because of the exceptionally preserved fossils that have been unearthed from a laminated limestone interval at the base of the unit (Martill et al., 2007;Heimhofer et al., 2010;Varejão et al., 2019a). This carbonate bed contains diverse fossils of terrestrial organisms that have undergone microbial entombment (Varejão et al., 2019a). ...
... This carbonate bed contains diverse fossils of terrestrial organisms that have undergone microbial entombment (Varejão et al., 2019a). This preserves delicate structures like fragile flowers and leaves, making it one of the most important Cretaceous Konservat-Lagerstätten (i.e., sites of exceptional preservation of soft tissues under unusual conditions) worldwide (Maisey, 1991;Heads et al., 2005;Martill et al., 2007;Ribeiro et al., 2021). ...
... The Aptian transitional phase comprises the rocks of the Santana Group, represented in ascending order by the Barbalha, Crato, Ipubi, and Romualdo Formations (Assine et al., 2014). Among them, the mixed carbonate-siliciclastic of the Crato Formation encompasses a Konservat-Lagerstätte (Crato Konservat-Lagerstätte, or CKL; see Ribeiro et al., 2021) in a laminated limestone deposit ≤8 m thick at the base of the unit (Grimaldi, 1990;Maisey, 1991;Martill et al., 2005Martill et al., , 2007Ribeiro et al., 2021;Varejão et al., 2019aVarejão et al., , 2021. ...
Premise:
The Lower Cretaceous Crato Konservat-Lagerstätte (CKL) preserves a rich flora including early angiosperms from northern Gondwana. From this area, the recently described fossil genus Santaniella was interpreted as a ranunculid (presumably Ranunculaceae). However, based on our examination of an additional specimen and a new phylogenetic analysis, we offer an alternative interpretation.
Methods:
The new fossil was collected from an active quarry for paving stones in the state of Ceará, north-eastern Brazil. We assessed support for alternative phylogenetic hypotheses using a combined analysis of morphological data and DNA sequence data using Bayesian inference. We used a consensus network to visualize the posterior distribution of trees and we used RoguePlot to illustrate the support for alternative positions on a scaffold tree.
Results:
The new material includes a flower-like structure not present in the original material and also includes follicles preserved at early stages of development. The flower-like structure is a compact terminal cluster of elliptical sterile laminar organs surrounding internal filamentous structures that occur on flexuous axes. Phylogenetic analyses did not support its placement among eudicots. Instead Santaniella appears to fall in the magnoliid clade.
Conclusions:
The presence of seeds in a marginal-linear placentation and enclosed in a follicle supports the fossil as an angiosperm. However, even though most characters are clearly recognizable, its combination of characters does not provide strong support for a close relationship to any extant order of flowering plants. Its position under the magnoliid clade is intriguing, and based on plicate carpels it is definitely a mesangiosperm. This article is protected by copyright. All rights reserved.
... A região é rica em diversidade geológica e com abundância de fósseis, bem preservados e diversificados, tornando-a o destino de geocientistas de todo o mundo (VIANA, 2001;ARAI;COIMBRA, 1990;BRUNO;HESSEL, 2006;CARVALHO;MELO, 2012;CARVALHO;SANTOS, 2005;MARTILL;BECHLY;LOVERIDGE, 2007;MOURA;BARRETO;BÁEZ, 2006;SANTOS, 1991;VIANA;NEUMANN, 2002). Por ser parte do Geopark Araripe, espera-se ampla visitação de turistas, inclusive do exterior, que proporcione o desenvolvimento local pela vertente do Geoturismo. ...
... A região é rica em diversidade geológica e com abundância de fósseis, bem preservados e diversificados, tornando-a o destino de geocientistas de todo o mundo (VIANA, 2001;ARAI;COIMBRA, 1990;BRUNO;HESSEL, 2006;CARVALHO;MELO, 2012;CARVALHO;SANTOS, 2005;MARTILL;BECHLY;LOVERIDGE, 2007;MOURA;BARRETO;BÁEZ, 2006;SANTOS, 1991;VIANA;NEUMANN, 2002). Por ser parte do Geopark Araripe, espera-se ampla visitação de turistas, inclusive do exterior, que proporcione o desenvolvimento local pela vertente do Geoturismo. ...
... A região é rica em diversidade geológica e com abundância de fósseis, bem preservados e diversificados, tornando-a o destino de geocientistas de todo o mundo (VIANA, 2001;ARAI;COIMBRA, 1990;BRUNO;HESSEL, 2006;CARVALHO;MELO, 2012;CARVALHO;SANTOS, 2005;MARTILL;BECHLY;LOVERIDGE, 2007;MOURA;BARRETO;BÁEZ, 2006;SANTOS, 1991;VIANA;NEUMANN, 2002). Por ser parte do Geopark Araripe, espera-se ampla visitação de turistas, inclusive do exterior, que proporcione o desenvolvimento local pela vertente do Geoturismo. ...
A revista Terr@ Plural tem a missão de publicar artigos científicos relacionados à área de Gestão do Território que contribuam com o desenvolvimento do conhecimento teórico e metodológico deste campo de saber. Além disso, visa estimular o debate acadêmico daqueles que atuam na temática e ampliar as relações com profissionais de outras regiões do Brasil e do exterior. A revista é uma publicação semestral e é composta pelas seções de artigos e resenhas.
... This lagerstätten type often displays plenty of fossils and could simultaneously represent a kondensat lagerstätte. The Early Cretaceous Crato Fossil Lagerstätte in northeastern Brazil provides both a lake deposit with numerous fossils of various groups (plants, insects, fishes, and reptiles) and fossils in excellent preservation (e.g., Martill et al. 2007a;Bezerra et al. 2020a). Since 2006, when the Araripe Geopark was established, the Pedra Cariri geosite refers to this lagerstätte and its exceptionally preserved fossils (Araripe UNESCO Global Geopark (Brazil) 2015). ...
... It is quite important to state that every geological unit (rock unit) represents a certain period that could be regarded in terms of geological imagination as a "moment" but representing several 10.000s or 100.000s of years. For the rock bed of the CFL, the Crato limestone, a depositional history of about 100.000 years was assumed (Martill et al. 2007a) but not confirmed by any reliable approach yet. Based on the nonoriented withdrawal of fossils from the limestone, no kind of succession or evolutionary adaptations of biota to environmental shifts within the Crato world has been documented so far, although some development in habitat architecture, landscape, and vegetation structure is conceivable. ...
... The basin is regarded as one of the world's most important fossil-bearing regions of the Mesozoic due to its extraordinary wealth of exceptionally well-preserved fossils from two Konservat-Lagerstätten (cf. Martill et al. 2007; see also the chapter "▶ The Crato Flora, a 115 Million Years Old Window into the Cretaceous World of Brazil" by Kunzmann et al., this volume). ...
... The Crato Formation mainly consists of lacustrine deposits which contain one of the Konservat-Lagerstätten of the basin. Laminated limestones of this formation contain a diverse and abundant fauna and flora (e.g., Martill et al. 2007; cf. different chapters in this book). ...
... During the deposition of the Crato Formation, there is a wide variety of continental and transitional environments, also with records of marine incursions (Neumann and Cabrera 2002a, b;Varejão et al. 2021a;Ribeiro et al. 2021). Within the lacustrine hypersaline succession (Varejão et al. 2021a), there are abundant and diverse exquisite preserved fossils that give the Lagerstätte status for the unit (Martill 2007;Selden and Nudds 2012). The Crato Biota, as referred by , is characterized by fungi, plants, arthropods, fish, frogs, lizards, turtles, pterosaurs, non-avian dinosaurs, and birds living in a wetland-type ecosystem influenced by climatic oscillations between wetter and drier periods (Ribeiro et al. 2021). ...
... During the Aptian, alkaline lakes were one of the main depositional environments in the Araripe Basin, representing the Lagerstätte succession of the Crato Formation. Although there are records of marine facies association (Varejão et al. 2021a, b) the dinosaur footprints of the Crato Formation are recognized in deposits from the margins of alkaline and hypersaline lakes, where other exceptionally preserved fossils ( Fig. 6.12) of vertebrates, invertebrates, plants, and even fungi are also found (Martill et al. 2007;Carvalho et al. 2021a;Dias et al. , 2023. The tracks of the Crato Formation occur in the Três Irmãos Quarry, Nova Olinda County, which is mainly composed of micritic limestone, with levels of marls and fine-grained siliciclastic beds (Neumann and Cabrera 2002a, b). ...
Fossil footprints are generally recognized by morphological features from the feet registered on the unconsolidated substrate; therefore, they can also be understood as biogenic primary deformation structures. The footprints can be found in the bedding surface and as cross-section deformations of the strata. Then, the substrate properties and the animal’s behavior allow for a wide range of track morphologies. The main modes of footprint preservation can thus be evaluated as the relationship between the substrate and the lower surface of the feet. In the Araripe Basin, both kinds of preservation are found in the Mauriti, Rio da Batateira, Crato, and Exu formations. In the Mauriti Formation, the footprints are imprints in the bedding surface of fine to coarse-grained sandstones. The cross-section footprints occur in the Rio da Batateira, Crato, and Exu formations. These footprints are produced by foot pressure on a depositional surface, and transmitted downward inside a bed or a bed set. In some cases, they are difficult to recognize, as they can be misinterpreted as load or liquefaction inorganic features related to sediment compaction, usually triggered by earthquakes and not to trampling by terrestrial vertebrates. They are in reality load casts produced by dinosaur trampling, allowing evaluation of substrate consistency besides the potential trackmaker identification. The temporal and environmental contexts of the dinosaur footprints from the Araripe Basin include the dinosaur trampling in fluvial sand bars, floodplains, deltas, and saline-alkaline lake borders.
... In contrast, less than 10% of the specimens collected in Cretaceous amber inclusions correspond to Chironomidae 36,58 . While chironomids (in particular larval stages) are typically rare or absent in amber deposits, as chances of being caught in the resinous secretions of the trees are scarcer in the aquatic environment, its absence in Cretaceous lithic sediments is also noteworthy and, as far as we are aware, only a partial wing of a possible chironomid has been so far reported from the Early Cretaceous of Brazil 59 . ...
... This may also be indicative that lepidopterans may have acquired an outstanding role as pollinators and probably nectarivores by the latest Cretaceous. Such condition is reminiscent to Cenozoic insect associations, in which lepidopterans constitute one of the most frequent and diversified clades of phytophagous insects 36,69,70 , contrasting with the Early Cretaceous times, when lepidopterans were numerically scarce [52][53][54]59 . ...
Insect faunas from the latest Cretaceous are poorly known worldwide. Particularly, in the Southern Hemisphere, there is a gap regarding insect assemblages in the Campanian-Maastrichtian interval. Here we present an insect assemblage from the Maastrichtian Chorrillo Formation, southern Argentina, represented by well-preserved and non-deformed, chitinous microscopic remains including head capsules, wings and scales. Identified clades include Chironomidae dipterans, Coelolepida lepidopterans, and Ephemeroptera. The assemblage taxonomically resembles those of Cenozoic age, rather than other Mesozoic assemblages, in being composed by diverse chironomids and lepidopterans. To the best of our knowledge, present discovery constitutes the first insect body fossils for the Maastrichtian in the Southern Hemisphere, thus filling the gap between well-known Early Cretaceous entomofaunas and those of Paleogene age. The presented evidence shows that modern clades of chironomids were already dominant and diversified by the end of the Cretaceous, in concert with the parallel radiation of aquatic angiosperms which became dominant in freshwater habitats. This exceptional finding encourages the active search of microscopic remains of fossil arthropods in other geological units, which could provide a unique way of enhancing our knowledge on the past diversity of the clade.
... The Crato Formation was deposited in a stratified lacustrine environment (Martill et al., 2007;Osés et al., 2016;Varejão et al., 2019;Barling et al., 2020) or perhaps a semi-arid wetland (Ribeiro et al., 2021), and yielded a diverse fos-sil assemblage (Martill et al., 2007). The preservation fidelity varies among different taxa but is extremely high for insects (Fig. 1A). ...
... The Crato Formation was deposited in a stratified lacustrine environment (Martill et al., 2007;Osés et al., 2016;Varejão et al., 2019;Barling et al., 2020) or perhaps a semi-arid wetland (Ribeiro et al., 2021), and yielded a diverse fos-sil assemblage (Martill et al., 2007). The preservation fidelity varies among different taxa but is extremely high for insects (Fig. 1A). ...
Prokaryotes, including bacteria, are a major component of both modern and ancient ecosystems. Although fossilized prokaryotes are commonly discovered in sedimentary rocks, it is rare to find them preserved in situ alongside macrofossils, particularly as pyritized cells in sites of exceptional fossil preservation. We examined prokaryotes preserved in the Lower Cretaceous Crato Formation of Brazil and demonstrate the widespread presence of spherical microorganisms preserved on the surface of Crato invertebrate fossils. These microorganisms were pyritized, covering decaying carcasses, 1.14 ± 0.01 μm in size, hollow with smooth surfaces, and can be found as aggregates resembling modern prokaryotes, particularly, coccoid bacterial colonies. It is likely that the observed microorganisms covered the carcasses before permissive conditions were established for pyritization, which must have been so rapid as to inhibit the autolysis of their delicate membranes. This is a new record of prokaryote fossils preserved in pyrite in association with macrofossils, which highlights the unique diagenetic and paleoenvironmental conditions of the Crato Formation that facilitated this mode of fossilization.
... Well exposures of the unit were characterised and sampled by the present study at the Três Irmãos Quarry (section 1BAr17, Fig. 2A), ''Cascatinha" (small waterfall) point at the Batateira River (outcrop 3BAr06, Fig. 2C), and Serra do Mãozinha hill (section 2BAr05, Fig. 2F), also the unit was completely recorded by the well-core 2-AR-SR-1A-CE (Fig. 2D) from 62.40 m depth down to the base. The basal strata of the Crato Formation constitute Konservat-Laggerstätte carbonates considered by Martill et al. (2007) as having been deposited in a protected lagoonal system, with probably fluctuating salinities and reducing conditions attested by the presence of sulphide minerals (Heimhofer and Martill, 2007). Intermittent connection of the Crato lagoon with the sea perhaps through restricted channels is suggested by the presence of certain marine fish taxa which could have entered the lagoon for reproduction and use it as a nursery (Brito, 2007;Brito and Yabumoto, 2011). ...
... Heterolithic facies represented by black and calcareous shale with limestone concretions and interstratified with siltstone, claystone and limestone levels occur over the basal coarse-grained facies of the Romualdo Formation (Assine et al., 2014). The black shales level bearing abundant early diagenetic carbonate concretions with well-preserved fossils, represents another Konservat-Lagerstätte (Maisey, 1991;Martill et al., 2007) of the Santana Group. As well as ostracods and other marine microfossils, a diverse macrofossil invertebrate and vertebrate fauna like echinoids, decapod crustaceans, molluscs, fishes, dinosaurs, pterosaurs are recovered (Kellner and Campos, 2002;Brito and Yabumoto, 2011;Prado et al., 2018aPrado et al., , 2018bPrado et al., , 2019Melo et al., 2020;Sayão et al., 2020;Araripe et al., 2022;Guzmán et al., 2022;Pereira et al., 2022;Tomé et al., 2022). ...
The Santana Group, consisting of the Barbalha, Crato, Ipubi and Romualdo formations, records the postrift sequence of the Gondwana break-up in the Araripe Basin. The post-rift phase of the South Atlantic rifting is dated as the Alagoas Stage, which corresponds to the ostracod biozone 011. A detailed ostracod
stratigraphical distribution along the Santana Group, allowed to standardise a common ostracod for the biozone name and a code, the Pattersoncypris micropapillosa Biozone (OST-011). Four different ostracod associations were recognized establishing the subzones: Pattersoncypris cucurves (OST-011.1), Pattersoncypris cucurves-Neuquenocypris berthoui (OST-011.2), Damonella grandiensis (OST-011.3) and Pattersoncypris crepata (OST-011.4). Planktic foraminifera were also recovered in different associations,
leading to identify two international biostratigraphic intervals and calibrate the ostracod subzones, the Early Aptian Leupoldina cabri Zone includes the OST-011.1 and OST-011.2 subzones, and the upper Late Aptian Hedbergella infracretacea–Microhedbergella miniglobularis composite zone correlated to the OST-011.4 subzone. Between these zones there is an interval without foraminiferal biostratigraphy resolution, the OST-011.3 subzone which is assigned to the lower Late Aptian. The palaeoenvironmental evolution of
the Santana Group was reconstituted from the integrated study of ostracods, planktic and benthic foraminifera. Through the Aptian, the Araripe Basin evolved from a transitional to a marine environment, recording estuarine, fluvial to bayhead deltas, a bay coastal watershed that reached better-marine conditions under extreme aridity and the full installation of an epeiric sea flooding the area.
... Type species: Cratovitisma oldreadi Bechly, 2007 Composition. Cratovitisma bechlyi Podstrelená in Podstrelená and Sendi (2018) ;Podstrelená and Sendi 2018). ...
... n., Pseudojantaropterix lebani and Lepidopterix vegrandis were reported (Podstrelená and Sendi 2018;Vršanský 2003a;Sendi et al. 2020a), while Compunctiotypus hudai was described from Jordanian amber (Kaddumi 2005). Cratovitisma is distributed in Brazil, Myanmar and Lebanon and Pseudojantaropterix is found in Lebanese and Jantaropterix in northern Myanmar amber (Bechly 2007) and New Jersey amber (Vršanský 2003a). On the other hand, Lepidopterix is only reported in Lebanese amber and Compunctiotypus only in Jordanian amber (Sendi et al. 2020a;Kaddumi 2005). ...
The earliest amber with numerous important paleobiological samples, and the only significant Lower Cretaceous one containing cockroaches, comes from the Lebanese regions Bouarij, Falougha, Hammana-Mdeirij, Jezzine, Mechmech, Qanat Bakish, Bqaatouta and Rihane (and 21 unstudied outcrops), and from Jordanian amber of Wadi Zerka and Syrian amber of Bloudane. Complex data (RTFI spectroscopy, palynology and entomofauna with diverse and abundant Nematocera) indicated that the amber originated in a subtropical, warm and humid gymnosperm-dominated forest in close proximity to the sea. 79 specimens of cockroaches s. s. (excluding Isoptera, Mantodea and possibly Chresmoda) were housed in 18 species (and three species incertae sedis), including Pravdupovediac neklam, P. maaloufi gen. et spp. n; Elisama globosa, Cratovitisma spinosa and Neoblattella nechapetomu spp. n. The former three belong to the dominant family Blattulidae Vishniakova, 1982 (n = 14), while Umenocoleidae Chen et Tan, 1973 were codominant (n = 10) in the present fossil record. Diversity and disparity of the preserved forms was moderately high and comparable to the Middle Jurassic and Lower Cretaceous sedimentary sites – i.e., higher than in Early Jurassic sites and lower than in Upper Jurassic Karabastau sedimentary rocks and Late Cretaceous North Myanmar amber. Scarce syninclusions in small amber pieces represent a coccid, psychodid, ceratopogonid, possibly a pathogenic fungus and plant trichomes. Palaeogeographically Lepidopterix Sendi, 2020; Cryptoblatta Sendi, 2019; Miniblattina Sendi, 2021; Pravdupovediac gen. n. and possibly Nymphoblatta Vršanský et Grimaldi, 2004 (Lebanese amber); Compunctiotypus Kaddumi, 2005 and Grandocularis Kaddumi, 2005 (Jordanian amber); and Anenev Vršanský, Oružinský, Sendi, Choufani, El-Halabi et Azar, 2019 (Syrian amber) are reported only in the herein studied amber, while the Jantaropterix/ Pseudojantaropterix-complex; Cratovitisma Bechly, 2007; Nymphoblatta Vršanský et Grimaldi, 2004 aff. Perlucipecta Wei et Ren, 2013 and aff. Sivis Vršanský, 2009; Pseudomantina Sendi et Vršanský, 2021; Elisama Giebel, 1856; Ocelloblattula Anisyutkin et Gorokhov, 2008, ?Rhipidoblatta Vishniakova, 1968; Balatronis Šmídová et Lei, 2017 and Neoblatella Shelford, 1911 (nowadays restricted to the Americas) were cosmopolitan with occurrences in both Laurasia and Gondwana. The low percentage (0.4% compared with 2.5–5% in North Myanmar amber) of cockroaches preserved in Lebanese amber could be due to lower viscosity of the amber-forming resin, and/or due to different paleoenvironments, suggested also for Taimyr amber with a moderate climate (constituting 0.3% cockroaches of all samples). Phylogenetically, several still living families, such as Ectobiidae Brunner von Wattenwyl, 1865 and Blattidae Latreille, 1810 have their oldest representatives within Lebanese amber, suggesting that the amber is advanced in evolutionary aspect among all Mesozoic ecosystems (all other families went extinct near Cretaceous–Paleogene boundary, K/Pg). Furthermore, Neoblattella is the earliest recorded still living genus. However, respective species from the present amber complex do not reveal any specific phylogenetical signal and weren’t measurably more basal when compared with their younger North Myanmar amber counterparts. A wide range of morphotypes with various ecological adaptations are documented, suggesting already established, sophisticated trophic relationships, similar as in today’s ecosystems. Nevertheless, given the abundance of blattulids and mesoblattids, most had probably a standard detritivorous feeding habit - the more advanced morphotypes known from Myanmar amber were absent. To summarise, the herein studied amber complex reveals origination of a moderately high diversity and disparity of forms (corresponding to the beginning of the Cretaceous Terrestrial Revolution), similar to Upper Jurassic assemblages, with a number of advanced forms, but lower than in burmite, which might be caused due to collection bias, shorter deposition time, milder tropical climate, or due to a different ecosystem (e.g. with less diversified angiosperms). The present amber reveals the earliest record of already triggered Mesozoic-to-Cenozoic ecosystem change within global forests.
... The Early Cretaceous Crato Formation is located in the Araripe Basin, which outcrops primarily along the slopes of the Chapada do Araripe plateau near the border of the Brazilian states of Ceará, Pernambuco, and Piauí. The Crato Formation is part of the Santana Group, and its lower part [often referred to as the Nova Olinda member, but see Ribeiro et al. (2021)] is now recognized as one of the most important Lagerstätten in the world, due to the combination of its interesting age and the exceptional preservation and diversity of its fossil fauna and flora (Fig. 2) (Martill et al., 2007). Traditionally, the sedimentary environment of the Crato Formation has been interpreted as a hypersaline lacustrine system (Varejão et al., 2019). ...
... Megaloptera classification follows , and Raphidioptera classification follows Engel (2002). The lower-level (genus and species) taxonomic and nomenclatural content of the catalog are generally based on Martill et al. (2007), Moura-Júnior et al. (2018), and the Neuropterida Species of the World . All original descriptions were re-checked. ...
... The pronotum shape is similar to the basal Cratovitisma Bechly, 2007 where the pronotum is centrally elevated and with a rounded posterior margin and flattened paranotalia, but less rounded (sharper) anterior margins. Among Cratovitisma, species are usually also much broader with shorter antenna. ...
... Although, Cratovitisma Bechly, 2007, Alienopterix Mlynský et al., 2018, Antophiloblatta Sendi in Sendi et al. (2020a, Perspicuus Koubová in Koubová and Mlynský (2020), Jantaropterix Vršanský et Grimaldi in Vršanský (2003), Nadveruzenie Vršanský, Hinkelman et Sendi in Vršanský et al. (2021), Vzrkadlenie Vršanský in Sendi et al. (2020a) were housed recently in the synonimised (and not revived formally) Cratovitismidae Bechly, 2007 according to Luo et al. (2021aLuo et al. ( , 2022 contrasting with Szabó et al. (2022) and Kováčová (2022Kováčová ( , 2023, they were used in the present network analysis, and grouped together (Fig. 14). Morphological boundaries between Umenocoleidae and the synonimised Cratovitismidae remain unclear as punctures of smaller size with sensilla also appear in taxa, such as Cratovitisma (see Fig. 2 in Podstrelená and Sendi 2018), and Trapezionotum (Fig. 5a, g). ...
Umenocoleidae is an enigmatic, extinct Mesozoic beetle-like roach family with cosmopolitan distribution erroneously attributed to various insect orders by past authors. Here unusual and well-preserved Archaeospinapteryx tartarensis gen. et sp. n., Trapezionotum vrsanskyi gen. et sp. n., Poikiloprosopon celiae gen. et sp. n., Vzrkadlenie karneri sp. n., and V. saint- germaini sp. n. are described from Cenomanian Burmese amber of the Hukawng Valley. Furthermore, Nigropterix angustata gen. et sp. n. and Umenotypus maculatus gen. et sp. n. are reported from Turonian sediments of Kzyl-Zhar, Kazakhstan. ?Classopolis pollen grains directly attached on the frons of the head of one specimen indiciate pollen transfer, as reported in other representatives of Umenocoleoidea. The herein studied material increases our knowledge of the diversity of this bizarre lineage and extends its temporal and paleogeographic distribution.
... The Early Cretaceous Crato Formation is located in the Araripe Basin, which outcrops primarily along the slopes of the Chapada do Araripe plateau near the border of the Brazilian states of Ceará, Pernambuco, and Piauí. The Crato Formation is part of the Santana Group, and its lower part [often referred to as the Nova Olinda member, but see Ribeiro et al. (2021)] is now recognized as one of the most important Lagerstätten in the world, due to the combination of its interesting age and the exceptional preservation and diversity of its fossil fauna and flora (Fig. 2) (Martill et al., 2007). Traditionally, the sedimentary environment of the Crato Formation has been interpreted as a hypersaline lacustrine system (Varejão et al., 2019). ...
... Megaloptera classification follows , and Raphidioptera classification follows Engel (2002). The lower-level (genus and species) taxonomic and nomenclatural content of the catalog are generally based on Martill et al. (2007), Moura-Júnior et al. (2018), and the Neuropterida Species of the World (Oswald, 2022). All original descriptions were re-checked. ...
The currently-known extinct Neuropterida fauna of Brazil comprises 18 families, 55 genera and 99 species (Neuroptera: 15 families, 49 genera, 89 species; Raphidioptera: 1 family, 2 genera, 5 species, + 1 genus incertae sedis with 2 species; Megaloptera: 1 family, 2 genera, 2 species; Order incertae sedis: 1 family, 1 genus, 1 species). All of these species have been described since 1980 and they collectively constitute the overwhelming majority of extinct Neuropterida presently known from South America. Other than one species of Permithonidae (Neuropterida incertae sedis) described from the Late Permian Irati Formation (Paraná Basin) of Rio Grande do Sul state, all extinct Brazilian Neuropterida taxa are known only from the Early Cretaceous Crato Formation (Araripe Basin) in Ceará and Pernambuco states. Of the 35 authors who have contributed to the descriptive literature of this fauna over the past 42 years, the work of Brazilian paleontologists Rafael Gioia Martins-Neto [1954-2010] and Maria Aparecida Vulcano-d'Andretta [1921-2018] - who together contributed to the description of 67 species - is especially notable. However, since their deaths, extinct Neuropterida have received little attention from Brazilian scientists and most later work on this fauna has been undertaken by workers outside Brazil. We provide this updated synthesis of the extinct Neuropterida of Brazil in order to centralize baseline knowledge for this fauna, to support efforts to document and conserve Brazilian fossil sites, and to encourage future work on the fauna, particularly by the next generation of Brazilian paleoentomologists.
Keywords:
Fossil; Crato formation; Irati formation; Taxonomy
... Recently, Warren et al. (2017) and Varejão et al. (2019Varejão et al. ( , 2021 suggested a shallow hypersaline coastal lacustrine system dominated by benthic microbial mats. The Crato Formation is divided into four members including, from bottom to top: the Nova Olinda, Caldas, Jamacaru and Casa de Pedra (Martill et al. 2007). The lowest Nova Olinda Member crops out in the northern and eastern flanks of the Araripe Basin, where the fossil insects from this study were found. ...
... The lowest Nova Olinda Member crops out in the northern and eastern flanks of the Araripe Basin, where the fossil insects from this study were found. The palaeoenvironmental conditions and exceptional preservation of fossils have made the Crato Formation well known worldwide as a Cretaceous fossil Lagerstätte (Martill et al. 2007). ...
Many studies have improved our understanding of the mode of preservation at the Crato fossil Lagerstätte. The high degree of preservation of the Crato mineralized insects is thought to be a consequence of the diffusion of ions through carcasses and envelopment by bacteria that, in turn, created microenvironmental conditions that led to mineralization, mainly pyritization. Pyritized insects have been oxidized by in situ weathering to more stable oxide/hydroxy minerals during Quaternary time. This transformation is essential to maintain the palaeontological information acquired during microbially induced pyritization in an oxidizing atmosphere. However, intense weathering can diminish or obscure the morphological fidelity, and little attention has been paid to the post-diagenetic processes experienced by these fossils. Here, we aim to determine the degree of alteration undergone by Crato pyritized insects using the following combination of analytical tools: scanning electron microscopy, energy dispersive X-ray spectroscopy, Fourier transform infrared and Raman spectroscopy. Our results show that well-preserved insects are preferentially replaced by haematite and poorly preserved fossils are replaced by goethite. In addition, we recorded three types of post-diagenetic alteration: insects with iron-oxide overgrowths; insects associated with black coatings, sometimes with the formation of dendrites; and insects preserved as an impression, where only the outline of the body remains. All of these alterations have the potential to distort or tarnish palaeontological information. Here, we measured the effects of such telodiagenetic alterations at macro and micro scales. Therefore, this taphonomic approach has wide applicability wherever fine-grained deposits bearing mineralized insects are found.
... Its origin is related to transcurrent tectonics occurred during the Early Cretaceous, controlled by Proterozoic lineaments and regional E-W faulting, related to the opening of the South Atlantic Ocean (Matos, 1992), particularly to the Brazilian Equatorial Margin's basins. Besides the high quantity and quality of its invertebrate and vertebrate macrofossils (e.g., Maisey, 1991;Martill et al., 2007;Carvalho et al., 2015aCarvalho et al., ,b, 2019cAgnolin et al., 2020;Iniesto et al., 2021;Kroth et al., 2021;Carvalho et al., 2023a;Carvalho, 2020, 2022;Dias et al., , 2023Santos et al., 2023), dinosaur footprints are commonly found in four lithostratigraphic units: Mauriti, Rio da Batateira, Crato, and Exu formations (cf. Carvalho et al., 1995Carvalho et al., , 2018Carvalho et al., , 2019aCarvalho et al., ,b, 2021aCarvalho et al., ,b, 2022Carvalho et al., , 2023b. ...
A NEW DINOSAUR TRACKSITE FROM THE ARARIPE BASIN (BRAZIL) AND THE PUTATIVE EARLY PALEOZOIC AGE FOR THE MAURITI FORMATION
Pp 97-102 In: Louis H.Taylor, Robert G. Raynolds, and Spencer G. Lucas, eds., 2024, Vertebrate Paleoichnology: A Tribute to Martin Lockley. New Mexico Museum of Natural History and Science Bulletin.
Footprints in the Araripe Basin (Northeastern Brazil) are previously known in the Mauriti Formation only from the Milagres ichnosite (Milagres County, Ceará State) on coarse-to fine-grained sandstones. Since this lithostratigraphic unit is considered and mapped as Silurian-Devonian (despite the fact that no macro-or microfossils established its age) the presence of such footprints shows a temporal inconsistency. The dinosaur footprints found in a new ichnosite (Mauriti County, Ceará State) reinforce the Mesozoic age for the Mauriti Formation, and due to the proximity of the nearby Rio do Peixe basins and to the similarity of their dinosaur footprints, a particularly Early Cretaceous age is suggested. This new ichnosite, herein named as Mauriti ichnosite, presents seven isolated footprints. There are four tridactyl, mesaxonic isolated footprints with pointed (?theropod) and rounded digits (?ornithopod). The other imprints are rounded depressions with no clear digit impressions, surrounded by displacement rims or presenting fluidization features of indeterminate trackmakers. The partial sandstone filling of the footprints is similar to the surrounding matrix. They range from 30-48 cm in length and 25-48 cm in width. The paleoenvironmental interpretation of the dinoturbation strata is fluvial braided. The trackmakers are small and large bipeds, despite some pointed digits induce to consider them as theropods related to those already known in the Araripe Basin's Cretaceous formations. This new tracksite confirms the need to revise the age of the Mauriti Formation and the involved paleogeography, establishing a new stratigraphic framework for the lower successions of the Araripe Basin.
... Since the fossil here presented has both sword-shaped elongate ovipositor valves, not as long as those of Carboniferous roachoids, and external short ootheca (Figs. 2D, E, S2B), three conclusions of importance for clarifying some aspects of dictyopteran evolution can be drawn: i. the appearance of the ootheca in cockroaches, and thus in general of maternal brood care in insects, dates back at least to the Middle Triassic, and not to Early Cretaceous 39,[41][42][43] or Jurassic 44 as previously thought; ii. the ootheca has evolved as a structure to protect and bear only a few eggs; and iii. the reduction in length of the ovipositor valves was not a prerequisite to producing the ootheca. The latter statement implies that the dramatic ovipositor reduction occurred after the ability to deposit eggs within this coriaceous structure was gained. ...
The Triassic represents a critical period for understanding the turnover of insect fauna from the Paleozoic to the Mesozoic following the end-Permian mass extinctions (EPME); however, fossil deposits from the Early-Middle Triassic are scarce. The exceptionally preserved 239 million-year-old fossil insect fauna recorded at Monte San Giorgio (Switzerland), including 248 fossils representing 15 major insect clades is presented here. Besides the exceptional features, including their small size and excellent preservation, the fossils have importance in the evolutionary history of the group. The taxonomic and ecological diversity recovered, including both freshwater (dragonflies and caddisflies) and terrestrial taxa (true bugs and wasps), demonstrates that complex environments sustained a paleocommunity dominated by monurans (thought not to have survived the EPME), midges, and beetles. Interestingly, a blattodean-like fossil bearing an external ootheca was also found, important for understanding Paleozoic roachoids to extant cockroaches’ transition and the evolution of maternal brood care. Moreover, the youngest and first complete specimen of †Permithonidae and the oldest sawfly fossils were discovered. Finally, round-shaped bodies, compatible with seminal capsules or lycophyte spores, were found on the abdomens of several midge-like individuals. If these are spores, non-seed-bearing plants could have been the first entomophilous plants rather than gymnosperms, as recently supposed. Altogether, these fossils contribute substantially to understanding insect evolution and Paleozoic-Mesozoic faunal turnover.
... Many cockroach fossils are adpressions (e.g. lee 2016 ;Bechly 2007;Vršanský 1999;Vršanský and Ansorge 2001;Kováčová et al. 2023;Káčerová and Azar 2023;Majtaník and Kotulová 2023;Barna et al. 2023), but the best preserved and most popular source of fossil cockroaches is amber (see Poinar 2023;anisyutkin and Perkovski 2023). amber inclusions of cockroaches have been documented from a few Mesozoic ambers, e.g. from France, the United States, Lebanon, Jordan, Myanmar, Syria, russia, and Hungary (anisyutkin and Gorochov 2008;Vršanský and Grimaldi 2003;Vršanský 2004Vršanský , 2008aVršanský ,b, 2009Vršanský , 2019Vršanský et al. 2019;Koubová & Mlynský 2020;Li and Huang 2023;Szabó et al. 2022d;. ...
Mesozoic amber cockroaches are rare compared to sedimentary imprints and are only known from Myanmar, Lebanon, France, Russia and USA. Perspicuus csincsii sp. n. from the Late Cretaceous (Santonian) ajkaite amber of Hungary with very long cerci, cup-like terminal palpomere of the maxillary palp and fully carinated legs, is clearly distinguished from all other Vitisminae species by the unique, black forewing with small light-coloured dots/stripes and the absence of pubescence on the wings (autapomorphies). The second cockroach described from the ajkaite adds important data to the distribution pattern of Vitisminae during the Late Cretaceous and further highlights the similarity of the ajkaite and burmite biotas.
... Due to the absence of true marine fossils, there is strong indication that the Crato Formation strata were deposited under lacustrine conditions (Neumann et al., 2003;Heimhofer et al., 2010;Varejão et al., 2021), though the presence of halite pseudomorphs in some sections suggests that the basin experienced salinity variations with increased arid and evaporitic conditions (Martill, 2007;Warren et al., 2017;Heimhofer et al., 2010;Storari et al., 2021). The layers of the Crato Formation have received several stratigraphic classifications to date. ...
The Crato Formation (Santana Group, Araripe Basin, Brazil) bears a high abundance of fossils in exceptional state of preservation, with insects from the order Orthoptera standing out. However, so far, few studies have explored their preserved inner organs in detail. Here, we provide the first detailed description of fossilized proventriculi from nine Grylloidea (Orthoptera: Ensifera) specimens of the Crato Formation. In all analyzed specimens, the external cuticle of the abdomen is cracked exposing the proventriculi, which are preserved as a tridimensional organ with a globular body and a tubular neck, similar to that of modern crickets. However, in the globular region of all fossils analyzed there are 9–12 rows of parallel divisions, differing from the modern crickets which have, more frequently, six. SEM images of two specimens revealed the exceptional preservation of internal median teeth, folds, and microvilli texture preserved in the organ.
... The Crato Formation consists of a thick succession of meter-scale limestone banks interbedded with equally thick beds of shales and sandstones (Assine et al., 2014). Within the Crato Formation, the CKL is restricted to an~8 m thick limestone interval at the lower part of the unit (Martill et al., 2007;Varejão et al., 2019Varejão et al., , 2021 (Fig. 1). Recent dating from the slightly younger Romualdo Formation (Barreto et al., 2022) found it to be 110.5 ± 7.4 Mya old, thus indicating an Aptian age for the Crato limestones. ...
The angiosperm Araripia florifera was originally described based on a fossil specimen including a few lobed leaves and floral buds. Although nothing was known of the internal structure of the flowers, based on the external similarity of the flower buds, it was compared with Calycanthaceae (Laurales). A new fossil from the type locality provides the first morphological evidence for the gynoecium of A. florifera. An androecium was not found, but new evidence is still needed to determine whether the flowers are unisexual. The gynoecium of this fossil species is syncarpous or monomerous, the ovary is superior, globose and distinctly stipitate, and the style is elongated and relatively thick. This new floral information disagrees with suggestions that Araripia belonged to the family Calycanthaceae, because this extant family is characterized by perigynous flowers, with apocarpous gynoecium and ovaries that are never stipitate. We also question the leaf arrangement, originally proposed as opposite, but alternate in this new fossil. We provide a phylogenetic hypothesis combining morphological and DNA sequence data using Bayesian inference. Our phylogenetic analyses indicate that A. florifera is more likely an extinct lineage in the stem group of Laurales, and is hereby placed in its own family Araripiaceae.
... In surface area, the Araripe Basin include the Araripe Plateau (Chapada do Araripe), up to 900 m highs and extending some 200 km from east to west (Maisey, 1991). The basin is filled with coarse and fine-grained clastic sediments and chemical deposits with many lithostratigraphic proposals (Beurlen, 1971;Machado et al., 1990;Ponte and Appi, 1990;Mabesoone et al., 2000;Heimhofer and Martill, 2007;Martill, 2007aMartill, , 2007bHeimhofer et al., 2010;Paula-Freitas and Borghi, 2011;Rios-Netto et al., 2012;Assine et al., 2014;Araripe et al., 2022). ...
... The Aptian Crato Member (Santana Formation) of the Araripe Basin is considered an excellent example of a mixed siliciclastic-carbonate paleolake, known chiefly for its outcrops of fine-grained laminated limestones bearing well-preserved fossiliferous content, which has received considerable attention over the last few decades (e.g., Grimaldi 1990;Carvalho and Viana 1993;Mohr and Eklund 2003;Makarkin and Menon 2005;Martill et al. 2007a;Báez et al. 2009;Figueiredo and Kellner 2009;Carvalho et al. 2019;Varejão et al. 2019;Ribeiro et al. 2021), being recognized as a Konservat-Lagerstätte. However, the main characteristics of the depositional environment of this lithostratigraphic unit are still a matter of debate. ...
There is wide recognition of lacustrine sediments as excellent archives of a basin's depositional history due to their high sensibility to environmental changes. Among them, microbial limestones are one of the most valuable tools for paleoenvironmental reconstruction, as the biological agents responsible for their genesis tend to respond to short-lived variations of the depositional setting creating specific precipitation patterns. We here document and investigate the sedimentary features of a specific sedimentary layer, remarkable by the extraordinary lateral continuity of its textural attributes over kilometer distances. This marker horizon occurs among the first carbonate layers of the Crato Member (Aptian, Araripe Basin, NE Brazil), commonly assigned as a paleolacustrine system. We build upon a multiscale comparative analysis (mesoscale, microscale, and chemical) to outline the main processes and paleoenvironmental settings that prompted this interval's widespread and laterally almost uniform depos ition. A lamination pattern identified in different well cores was scrutinized and compared, proving the striking lateral continuity of the layer and attesting that autochthonous biologically-induced mineralization was the primary mechanism of the formation of the microbialites. Compositional and stable isotope results also show similar trends throughout the well cores, where minor differences represent the influence of local processes. The studied interval encompassed a relatively swift transition of organic shales rich in ostracod valves to planar stromatolites, where both developed in the anoxic benthonic zone of a freshwater lake. The precipitation of the overlying finely laminated limestones is related to a change in the carbonate genetic mechanism as a response to a more stable lacustrine stratification. The widespread formation of microbialites preserving an almost identical textural pattern must be related to a regional event, constituting a rare example of a preserved ancient biostrome. Moreover, the investigation of this sedimentary layer can further contribute to determining the roles of different biotic and abiotic processes in microbialite precipitation over large areas.
... The Pseudofrenopsis specimens studied came from the limestone succession named Crato Formation (Fig. 1C), which comprises one of the most important Cretaceous fossil Lagerstätten of Gondwana (Martill et al., 2007). It consists of gray, dark brown to black shales and gray to brown laminated limestones and is interpreted as deposited in a continental lake complex during a phase of tectonic quiescence (Neumann et al., 2002). ...
This study is the first contribution towards the estimation of paleoatmospheric carbon dioxide concentration (pCO2) from the paleoequatorial late Aptian Crato Formation (Santana Group, Araripe Basin, NE Brazil) based on stomatal numbers. Cuticular and epidermal silicon replicas of Pseudofrenelopsis capillata Sucerquia, Bernardes-de-Oliveira & Mohr (Cheirolepidiaceae, Coniferales) were observed under scanning electron microscopy to analyze their stomatal frequency and distribution for paleoatmospheric and paleoenvironmental inferences. Stomatal counting yielded a mean stomatal density (SD) and stomatal index (SI) of 68.8 mm-2 and 6.7. Based on the mean SI value of four modern nearest living equivalent (NLE) conifer species, the stomatal ratio (SR) was calculated to be 1.4. The paleoatmospheric pCO2 range was estimated to be between 514.9 ppmv (Recent standardization) and 1029.8 ppmv (Carboniferous standardization). These values were found to be consistent with results from the literature based on other latitudes in the Early Cretaceous and were discussed in the light of the paleoenvironmental context that prevailed around the peri-equatorial Crato paleolake, including the occurrence of stomatal clustering in the leaves of P. capillata.
... Revisions of the geology and paleontology of the Crato Formation is found in Martill et al. (2007aMartill et al. ( , 2007b and Ribeiro et al. (2021). The age of the Crato Formation is considered to be upper Aptian (Early Cretaceous) (Pons et al., 1991;Heimhofer and Hochuli 2010;Varejão et al., 2021). ...
... Several basins located at the modern continental Atlantic margins of South America and Africa began to form during the Jurassic and Cretaceous as diachronous rifts, thus recording the onset of the Gondwana breakup (Matos, 1992;Darros de Matos, 1999;Mohriak, 2003). Some sedimentary basins of this rift system in Northeast Brazil (e.g., Araripe, Tucano basins) contain Cretaceous deposits, in which distinct fossil groups are exceptionally preserved (see Martill et al., 2007;Varejão et al., 2019;Souza et al., 2022) and are considered Konservat-Lagerst€ atten (sensu Seilacher et al., 1985). ...
This paper describes a new genus and species of hangingfly (Mecoptera: Bittacidae) from the Lower Cretaceous Crato Formation of Brazil. Petrulebittacus martinsnetoi gen. et sp. nov. represents the first described Mesozoic Mecoptera fossil from Brazil and the oldest known Bittacidae from the New World. Comparisons are made with other fossil and extant Bittacidae genera. This discovery adds to the growing evidence that Bittacidae diversity during the Cretaceous may have been higher than previously thought. This paper also highlights the importance of housing important fossil specimens in public collections to ensure proper scientific study and description.
The Crato Formation, an Early Cretaceous lacustrine Lagerstätte in Northeastern Brazil, is noteworthy for soft-tissue fossilization. This unit records an abundant and diverse palaeoentomofauna, which preserve external and internal elements, including organs and delicate tissues. Here, we report the oldest and only known fossilized insect ovary fragments, found in the laminated limestone of the Crato Formation, Araripe Basin. It is a female Baissogryllidae, an extinct lineage of true crickets (Grylloidea). Dissection of the extant cricket, Endecous ( Notendecous ) onthophagus (Berg, 1891), facilitated accurate morphological and morphometric comparisons, enhancing our interpretation of the fossil structures. Besides the ovaries, dozens of elliptical microelements, resembling fossilized "eggs," were found in the female abdomen. These structures are significantly smaller than the mature oocytes of E . ( N .) onthophagus or even early-stage oocytes. It remains open whether these microelements represent immature baissogryllid oocytes or if Cretaceous cricket eggs were inherently smaller. Alternatively, this could be a preservation artefact. Taphonomic signatures suggest that this ancient Grylloidea lived close to a lacustrine environment, likely using lake margins for oviposition. Despite the open wings indicating death near the depositional environment, the straighter abdomen and ovipositor suggest that the cricket was not ovipositing at its time of death.
Two new species of the family †Falsiformicidae Rasnitsyn, 1975, Siccibythus robustussp. nov. and Siccibythus aristovisp. nov., are described from the mid-Cretaceous Kachin amber based on three well-preserved specimens. The new species belongs to †Siccibythus Cockx & McKellar, 2016, based on the elongate pronotum and fore wing with cells 2+3rm and 3r not enclosed with tubular veins. We summarize the diversity and distribution for all species of †Falsiformicidae and provide a revised key to the family. These new findings enhance the diversity of †Falsiformicidae in Kachin amber.
This study investigates the coxosternal or ventral sternal region, in scorpions and spiders. We collected 10 specimens of Scorpions from provinces Baghdad in region Abo Kreab and Salahuddin provinces in Tikrit then 7 specimens of Spiders were collected from variant regions in provinces Baghdad and various genera and families were collected and processed for examination. The aim was to compare the morphology of the coxosternal between these two arachnid groups and analyze the underlying functional and evolutionary implications of any structural variations. So the result of this study on coxosternal region of scorpions in the Family: Buthidae like Androctonus crassicauda, is not wide the majority of members of this family have a triangular cut and the female genital cover is divided, coxosternal region in Orthochirus scrobiculosus do not extend forward and do not form lobes, edges of the scales grainy and it is finely grained the majority of individuals have a triangular cut, Mesobuthus eupeus the ventral side is shiny and wide with strong hairs and the sternum is triangular, coxosternal region in Family: Scorpionidae like Scorpio Maurus in the ventral side do not have lobes extending forward, the sternum is pentagonal. But in Family: Salticidae of spiders the coxosternal region in Hasarius adansoni, are Oval shape sternum or variable in Thyene imperialis, and Evarcha seyun, are circle shape sternum or variable, coxosternal region in Neoscona subfusca, family Araniedae are heart-shaped or triangular. The expected results include identifying distinct coxosternal shapes in scorpions compared to spiders. These variations might be linked to specific Roles like locomotion and internal organ protection. The determined geomorphology gets to render Understandings into the evolutionary relationships betwixt scorpions and spiders. This comparative analysis is expected to Add to a better understanding of how the coxosternal plays a role in the biology of scorpions and spiders. The known variations get bid important information along the development and practical adjustations inside these arachnids
Since the first description from China, the Mesozoic dictyopteran family Umenocoleidae was recorded from the Eurasian and South American continents with 18 genera and 31 species. Here, we describe one new genus and three species from the Jinju Formation, South Korea: Umenocoleus minimus sp. nov., Psedoblattapterix weoni gen. et sp. nov., and Petropterix koreaensis sp. nov.. We also discuss three new apomorphies of the Umenocoleidae and Alienopteridae, which are 1) M, Cu and PCu1 veins form an common vein near wing base, 2) M, Cu and PCu1 veins appress to R vein and 3) CuP+PCu1 vein are not as concave as in the other Holopandictyoptera (total group of extant Blattodea and Mantodea). On the basis of these new apomorphies, we support the inclusion of the family Alienopteridae into the superfamily Umenocoleoidea. We also support the exclusions of the genera Vitisma and Permoponopterix from the Umenocoleidae family and propose to transfer the genus Permoponopterix into the Protelytroptera.
Mantodea (praying mantises) is a group of exclusively predatory insects, which, together with nonraptorial blattodeans (cockroaches and termites) and groups exclusively found in the fossil record, form the group Dictyoptera. A central characteristic of Mantodea is the specialization of their first pair of legs as raptorial grasping appendages, but the evolution from walking to raptorial legs is not yet fully understood. Here, we trace the evolution of the raptorial appendages in Dictyoptera through time using a morphometric (morphospaces) approach. We also describe two new mantodean nymphs preserved in amber from the Cretaceous and Eocene, which expand the scarce mantodean fossil record. Blattodean and mantodean appendages appear distinct in morphospace, but several appendages of fossil non‐mantodeans can be considered raptorial, providing a potential transitional link between walking and raptorial morphotypes. Therefore, we discuss potential mantodean affinities for other predatory fossil dictyopterans. We examine changes across extant mantodeans, characterized by a straightening of the tibia especially associated with the rise of the diversification of the Mantidea and discuss whether a thickening of the femur could reflect an early adaptation to cursorial hunting.
The Crato Konservat-Lagerstätte is one of the main Mesozoic fossil sites from Gondwana, recording a wide diversity of terrestrial and non-marine aquatic fossils of great palaeobiological and evolutionary significance. This conservation deposit is recorded in a 9 m-thick interval of laminite, microbialite, and grainstone deposited in a lake system with variable water level, alternating moments of hypersaline and freshwater conditions. Despite numerous studies describing new species of plants, arthropods, fish, pterosaurs, birds, and many others, there remains a significant gap in our understanding of the most common and archetypal fossils, which are the rod-shaped macrofossils found on bedding surfaces in distinct stratigraphic intervals of the Crato Konservat-Lagerstätte. The rod-shaped macrofossils are up to 1.6 cm-long and 0.1 cm-wide, straight to curved compressions that preserve pyritized microfossils. Here we interpret the rod-shaped macrofossils as macroscopic organic aggregates that sank into the lakebed in a process called lake snow. During high organic productivity periods in the epilimnion, planktonic organisms thrived and produced exopolymers responsible for aggregation. Their concentrations in the limestone bedding planes reflect intensity of lake snow and environmental seasonality. Aggregates are prolate particles that are commonly oriented, suggesting their transport as bedload for short distances, which was facilitated by biostabilization by microbes and their exopolymers. Finally, pyritization was mediated by microbial communities living in the lakebed.
The fossil-rich Romualdo Formation (late Aptian/early Albian), Araripe Basin, northeastern Brazil, contains world-renowned Fossillagerstätten characterized by exceptionally preserved fossils. Macroinvertebrates in this formation are primarily represented by mollusks, echinoids, and decapod crustaceans. Mollusk shells are abundant in certain stratigraphic intervals, forming coquinas or shell pavements. Despite recent advances in our understanding of the taxonomy of certain groups, comprehensive taxonomic studies are lacking for almost the entire bivalve fauna. Therefore, a detailed taxonomic analysis is presented here. The described bivalves include four new genera (Araripenomia, Ciceromya, Inversatella, Australoeocallista), and six new species (Araripenomia infirma, Inversatella cearensis, Ciceromya edentulosa, Australoeocallista juazeiroi, Legumen kaririense, and Corbulomima delicata), in addition to Musculus maroimensis, Crassatella maroimensis, “Myrtea” sp. and “Tellina” sp. This bivalve fauna mainly consists of cosmopolitan and endemic brackish/marine genera, with Tethyan affinities. The fauna is not homogeneously distributed in the sedimentary succession of the Romualdo Formation, but is constrained to the third order highstand systems tract. Bivalves recorded from muddy facies are strongly dominated by infaunal and semi-infaunal suspension feeders. Assemblages of the sand-dominated facies, with dense shell accumulations of semi-infaunal to epifaunal byssate and infaunal suspension feeders, were formed under shallow, higher energy conditions. Despite the degree of generic endemicity, the mytilids, anomiids, crassateliids, astartids, tellinids, and corbulids are related to the bivalve fauna of the Early Cretaceous Riachuelo Formation of the Sergipe-Alagoas Basin, as previously demonstrated for the bakevelliids and echinoids. Indeed, the Romualdo bivalve fauna is, in part, a modified and impoverished brackish/marine fauna of the Riachuelo Formation.
The genus Vitisma Vršanský, 1999 is characterised by a shield-like, convex pronotum with transverse margins, scleroritized forewings with sometimes white macula, long clavus and rather simplified venation (simple Sc, straight and expanded M veins, CuA narrow and branched, A simple, R reaching apex). The hindwing venation is similar to the Blattulidae with a distinct pterostigma, but R1 is comb-like and CuA branched. The first record of Vitisma coriacea sp. n. in Burmese amber extends its palaeogeographic distribution to Gondwana, similar to other Umenocoleoidea. So far, the genus was only known from sediments in Spain, Russia, Mongolia, and possibly Japan. It also suggests that representatives of Vitisma inhabited warmer, tropical forest biomes.
A rich insect fauna is found in the Cretaceous (Aptian) Nova Olinda Member of the Crato Formation in the Araripe Basin, northeastern Brazil. The Crato Formation was deposited close to the centre of the supercontinent Gondwana, prior to significant rifting. In this paper, we analyzed remains of 1135 insect specimens, representing 55 families, within two limestone facies of the Nova Olinda Member: pale yellow and dark gray limestones. We collected taphonomical data and assessed the paleoecological significance of insect faunas in terms of their distribution and interaction with the paleoenvironment. The insect fauna is dominanted by fully terrestrial taxa despite preservation in an aquatic paleoenvironment. Considering both facies, the distribution of insect families is similar throughout the Nova Olinda Member, which excludes possible overlapping of one entomofauna over another. The pale yellow limestones near the top of the section are richer in fossil insect content and show a higher incidence of aquatic and semiaquatic insects preserved in dorsoventral view compared to the dark gray limestones. This suggests that the Crato insects preserved in the pale yellow limestones experienced short transport distances or prolonged decay or both. In contrast, insects preserved in the dark gray limestones appear proportionally with higher degrees of disarticulation than those preserved in the pale yellow limestones, indicating that these insects underwent a higher degree of postmortem transport in the biostratinomic stage. Compared to descriptions of other Early Cretaceous assemblages, the taxonomic literature of Crato Paleoentomofauna shows distinct differences. The Yixian and Zaza formations are described as being dominated by beetles and wasps. The Crato Formation, however, is relatively richer in Paleoptera species. Both the sedimentary facies studied and the paleoentomological content reveal a complex ecosystem inserted in a depositional setting, similar to modern long-standing wetlands, and where periodically flooded zones were surrounded by dry lands with xeromorphic vegetation
The known occurrences of Early Cretaceous freshwater bivalves in the Araripe Basin are confined to the 0.85- to 2-m-thick Caldas Bed of the Crato Formation. Herein, we record a new bivalve-dominated assemblage in a siltstone bed nearly 30 m below the upper boundary with the overlying Romualdo Formation. The assemblage is composed of ecologically incompatible bivalve mollusk species, being dominated by tiny individuals of Modiolus? sp., a brackish water form. These are directly associated with the freshwater bivalves Araripenaia elliptica and Monginellopsis bellaradiata, the latter two species commonly found in the Caldas Bed. Those bivalves are mixed with carbonized, partially comminuted plant remains. Specimens of Modiolus? sp. are usually articulated and are dispersed to loosely packed or forming cm-long clusters. A few shells of the freshwater forms are also articulated. The bivalves lived in a semi-confined embayment setting with fluctuating salinity. Background brackish water conditions must have existed long enough for the Modiolus? sp. larvae to settle and develop as juvenile individuals. Rapid salinity fluctuations associated with sea level and climate variations allowed A. elliptica and M. bellaradiata to thrive in the same area. Hence, the numerically dominant shells of the brackish water forms and the freshwater bivalves have been telescoped into the same bedding plane, generating an environmentally condensed, time-averaged benthic assemblage. Finally, the presence of these taxa considerably expands the vertical/temporal distribution of bivalve mollusks within the Crato Formation.
The Crato Formation is the main lacustrine unit of the Araripe Basin, known for its limestone layers since the nineteenth century, rich in well-preserved Early Cretaceous fossils. Since then, numerous biota has been studied, including abundant plant remains, thus granting the status of a Konservat Lagerstätte to the formation. Despite the paleontological interest, it is known that the host rocks of fossils are more than just limestone, yet include marls, shales, sandstones, and more. The study of these rocks complements environmental analyses of the Crato Formation, and they are essential in sedimentological and taphonomical research of lake deposits. In this subject, different authors carried out several studies, being the main approaches by means of analyses of fieldwork facies studies, petrography, mineralogy, microfossil studies, and organic and inorganic geochemistry. This effort involved data obtained from outcrops, quarries, and well logs. The current panorama deals with a very dynamic lacustrine system, sometimes fragmented in several lakes but eventually reassembled in a general tendency of enlargement. The main control was probably the rainfall regime under orbital forcing of cycles estimated to last 200–500 ka each. Alternating arid and slightly wet periods, six main carbonate units were recognized, each representing shallow to deep waterbodies fed by water, sediment, and nutrients through ephemeral deltas. Life thrived in four paleoenvironmental zones, from underwater saline lakes to the small hills around them. Once the lakes were very calm, the carcasses suffered little from transport; therefore, taphonomical studies have focused on chemical transformations inside the water column and sediment.
Dating back to the late Early Cretaceous, the macrofossil record of the iconic lotus family (Nelumbonaceae) is one of the oldest of flowering plants and suggests that their unmistakable leaves and nutlets embedded in large pitted receptacular fruits evolved relatively little in the 100 million years since their first known appearance. Here we describe a new fossil from the late Barremian/Aptian Crato Formation flora (NE Brazil) with both vegetative and reproductive structures, Notocyamus hydrophobus gen. nov. et sp. nov., which is now the oldest and most complete fossil record of Nelumbonaceae. In addition, it displays a unique mosaic of ancestral and derived macro- and micromorphological traits that has never been documented before in this family. This new Brazilian fossil-species also provides a rare illustration of the potential morphological and anatomical transitions experienced by Nelumbonaceae prior to a long period of relative stasis. Its potential plesiomorphic and apomorphic features shared with Proteaceae and Platanaceae not only fill a major morphological gap within Proteales but also provide new support for their unexpected relationships first suggested by molecular phylogenies.
Macrofossil evidence has demonstrated a first radiation of gnetophytes in the Early Cretaceous. However, the origin of the diversity of gnetophytes remains ambiguous because gnetalean macrofossils have rarely been reported from pre-Cretaceous strata. Here, we report a new putative gnetalean macrofossil reproductive shoot which possesses opposite phyllotaxy, long linear leaves more or less decurrent and having a prominent midvein and pedicled ovoid-ellipsoid and longitudinally striated chlamydosperms. Our new fossil is different from other known gnetalean macrofossils in the linear-lanceolate leaves with a midvein and pedicled chlamydosperms. As a result, we describe this new macrofossil reproductive shoot as new to science, i.e., Daohugoucladus sinensis gen. et sp. nov. Our new macrofossil displays additional morphological characters distinct from other known Mesozoic and modern gnetalean species and provides additional evidence of the origin and early evolution of female reproductive organs of gnetophytes.
Here a discussion is made regarding the validity and availability of the dinosaur name 'Ubirajara jubatus' or otherwise, according to the rules of the Internation Code of Zoological Nomenclature.
O Membro Crato e a Formação Codó são parte do registro Aptiano das bacias do Araripe e Parnaíba, respectivamente. A mineralogia das argilas e a litogeoquímica são ferramentas importantes para o estudo ambientes sedimentares, sendo poucos trabalhos realizados nessas unidades utilizando tais técnicas. Esse estudo tem como objetivo a caracterização mineralógica e litogeoquímica de lutitos provenientes dessas unidades visando reconstruções paleoambientais, paleoclimáticas e de proveniência sedimentar. Estudos de mineralogia, mineralogia das argilas e litogeoquímica foram realizados em amostras do Membro Crato e da Formação Codó. Montmorillonita, caulinita e illita foram identificadas como associação detrítica de argilominerais no Membro Crato, onde elevados teores de montmorillonita em relação a caulinita indicaram condições paleoclimáticas regionais áridas com baixas taxas de paleoprecipitação (< 250 mm/ano). Dados litogeoquímicos indicaram condições de paleosalinidade salobras, onde também foram observados aumento súbitos de paleosalinidade através de proxies de paleosalinidade (i.e., Rb/K), indicando entradas cíclicas de água marinha no ambiente. Duas sucessões de fácies foram construídas através de nove litofácies na pedreira Três Irmãos. A primeira se encontra na porção inferior e corresponde a barras de desembocadura deltaicas dominadas por maré, onde observou-se condições mais secas por conta de menores teores de caulinita em comparação à sucessão de fácies de laguna marginal localizada nas porções superiores. Na porção basal da Formação Codó foi identificado um paleolago anóxico hipersalino alcalino com base na ocorrência de 18 metros de folhelho composto por saponita autigênica, sendo tal argilomineral interpretado como originado da transformação in situ de outros argilominerais detríticos em condições hidroquímicas de alta salinidade e pH alcalino (~ 8 e 10). Processos hidrotermais foram responsáveis pelo enriquecimento do lago em magnésio com base em dados de litogeoquímica. Foi identificado um aumento súbito de salinidade através de dados litogeoquímicos na transição entre o paleolago hipersalino alcalino e uma sucessão de sabkha, sendo interpretado como o primeiro episódio relevante de entrada de água marinha no ambiente. Tal evento foi responsável por alterar as condições do lago de alcalinas para ácidas, inibindo a formação de esmectitas magnesianas. Os lutitos da sucessão de sabkha são compostos por uma associação detrítica de argilominerais constituída por montmorillonita, illita e caulinita. As associações detríticas de argilominerais observadas nas duas sucessões anteriores indicaram condições paleoclimáticas áridas com baixos valores de paleoprecipitação (< 250 mm). Contudo, acima da sucessão de sabkha descrita anteriormente e na base de uma sucessão sedimentar paleolagunar na porção superior da Formação Codó, um processo de humidificação paleoclimática regional foi identificado com base em dados de mineralogia das argilas, o que corroborou a hipótese de uma mudança paleoclimática regional associada com o estabelecimento precoce do cinturão equatorial úmido durante o Aptiano. Tal ambiente representa o intervalo marinho da Formação Codó e coincide com uma associação detrítica de argilominerais composta por caulinita, montmorillonita e illita. Uma área de proveniência composta de rochas pós-Arqueanas intermediárias foi interpretada para toda a sucessão com base em dados litogeoquímicos.
The Crato Formation is the main lacustrine unit of the Araripe Basin, known for its limestone layers since the nineteenth century, rich in well-preserved Early Cretaceous fossils. Since then, numerous biota has been studied, including abundant plant remains, thus granting the status of a Konservat Lagerstätte to the formation. Despite the paleontological interest, it is known that the host rocks of fossils are more than just limestone, yet include marls, shales, sandstones, and more. The study of these rocks complements environmental analyses of the Crato Formation, and they are essential in sedimentological and taphonomical research of lake deposits. In this subject, different authors carried out several studies, being the main approaches by means of analyses of fieldwork facies studies, petrography, mineralogy, microfossil studies, and organic and inorganic geochemistry. This effort involved data obtained from outcrops, quarries, and well logs. The current panorama deals with a very dynamic lacustrine system, sometimes fragmented in several lakes but eventually reassembled in a general tendency of enlargement. The main control was probably the rainfall regime under orbital forcing of cycles estimated to last 200–500 ka each. Alternating arid and slightly wet periods, six main carbonate units were recognized, each representing shallow to deep waterbodies fed by water, sediment, and nutrients through ephemeral deltas. Life thrived in four paleoenvironmental zones, from underwater saline lakes to the small hills around them. Once the lakes were very calm, the carcasses suffered little from transport; therefore, taphonomical studies have focused on chemical transformations inside the water column and sediment.
It is investigated the paleoecology of Araripemys barretoi Price, 1973 a pelomedusoid turtle from the Lower Cretaceous (Aptian) of Araripe and Parnaíba basins, Brazil. The analysis of forelimbs proportions allowed to interpret Araripemys barretoi as a turtle with a specialized morphology to live in large water bodies, as an agile swimmer. Several postcranial anatomical features suggest that Araripemys barretoi was able to live in aquatic environments of distinct magnitudes. This species was abundant throughout the Aptian in the Araripe and Parnaíba basins, where an epicontinental sea once existed, during the first marine ingressions related to the Equatorial Atlantic opening.
This chapter explores how the extant Odonata (dragonflies and damselflies) are only the small remaining part of the superorder Odonatoptera. The Odonatoptera are a clade with an impressive systematic diversity and morphological disparity since the Late Carboniferous. Their fossil record is rather well known because these insects inhabit humid environments, which are favorable to fossilization. Thus it is possible to follow the evolution of the morphology of the order, especially in the evolving complexity of their wing venation. It is this evolution through time that has rendered possible for this order to acquire highly specialized structures resulting in their impressive flying capacities.
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