Sam W. Heads's research while affiliated with University of Illinois, Urbana-Champaign and other places

Publications (84)

Article
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A fossil coral of the genus Solenastrea Milne Edwards & Haime (Anthozoa: Scleractinia: Faviidae) is reported from Pleistocene deposits exposed near the town of Terra Ceia in Beaufort County, North Carolina. The specimen is similar in gross and corallite morphology to Solenastrea hyades (Dana) which is common in Neogene and Quaternary sediments of N...
Article
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Two new species of the genus Archelcana Sharov (Orthoptera: Elcanidae: Archelcaninae) are described from the Toarcian (Lower Jurassic) deposits of Uerschterhaff, near Sanem, southern Luxembourg. Archelcana numbergerae sp. nov. and Archelcana tina sp. nov. are the first fossil insects to be formally described from the Uerschterhaff locality and the...
Article
In our recent paper discussing the impacts of dragonfly behaviour on their taphonomy from the Lower Cretaceous Crato Formation of Brazil (Barling et al., 2021), we figured eight specimens (Fig. 3, p. 145) illustrating dragonflies in various states of preservation. In the caption to this figure we listed the temporary laboratory numbers assigned to...
Article
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During the Mid-Miocene Climatic Optimum [MMCO, ~14 to 17 million years (Ma) ago], global temperatures were similar to predicted temperatures for the coming century. Limited megathermal paleoclimatic and fossil data are known from this period, despite its potential as an analog for future climate conditions. Here, we report a rich middle Miocene rai...
Article
The relative completeness of Odonata (dragonflies and damselflies) fossils from the Crato Formation is investigated and compared to other fossil insect groups from the same formation. Tagma completeness is measured as present, partial, or absent, with some additional subdivision of body components (head, thorax, limbs, individual wings, anterior an...
Article
We describe a new Eocene fossil assassin bug, Aphelicophontes danjuddi gen. et sp. nov. (Reduviidae: Harpactorinae), from the Green River Formation of Colorado. The specimens informing this description are extraordinarily preserved, particularly in the external and internal structures of the adult male genitalia. We discuss implications of the new...
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The fossil heteropteran Miocoris fagi Cockerell, 1927 stat. rev. is redescribed and transferred from Reduviidae to the nominate tribe in Nabidae. The fossil Nabidae are surveyed and enumerated. One taxon, Sinanabis brevipes Zhang, 1986 stat. rev., is removed from Nabidae and is considered incertae familiae.
Article
The Crato Formation Lagerstätte of NE Brazil is well known for the exceptional preservation of its fossil assemblage. A wide variety of plants and animals are preserved with a high degree of fidelity, revealing structures of labile tissues at the micron scale, or even smaller. Chemical analyses and descriptions of replacement fabrics observed in in...
Article
Xu, C.P., Fang, Y., Heads, S., Zhang, Q., Jarzembowski, E.A. & Wang, B. XX. 2019. The first Orthoptera (Insecta) from the Triassic of China. Alcheringa XXX, X–X. ISSN 0311-5518. The extinct family Locustavidae (Insecta: Orthoptera) is rare in the fossil record, and to date no representative has been reported from China. In this paper, Hejiafanga to...
Article
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A new genus and species of pygmy grasshopper (Orthoptera: Tetrigidae) is described from Eocene Baltic amber. Danatettix hoffeinsorum gen. et sp. nov. is assigned to the subfamily Batrachideinae based on antennae with more than 19 antennomeres, sulcate mesofemora, and rectangular paranota. This species is readily distinguished from other batrachidei...
Article
A new genus and species of Elcanidae (Insecta: Orthoptera) is described from mid-Cretaceous Burmese amber. Elcanonympha diana gen. et sp. nov. is known from a single, complete nymph and is distinguished from other elcanids by the pronotum cut by a single transverse sulcus, the dorsal margin of the metafemoral genicula with a row of prominent apical...
Article
Mud crickets (family Ripipterygidae) comprise a small family of orthopterans distributed throughout much of the Neotropics, but knowledge of this groups' biology, ecology and distribution remains poor in comparison to most orthopteran taxa. Here we review the state of knowledge of nymphs in the genus Mirhipipteryx Günther 1969 and report the genus...
Article
The fossil cricket Araripegryllus romualdoi was described by Freitas et al. (2016) based on a single, very poorly preserved specimen from the Romualdo Member of the Lower Cretaceous Santana Formation; a deposit famed for the exceptional preservation of vertebrates in carbonate concretions (Martill, 1996, 2007). While abundant and diverse in the und...
Article
A new genus and species of the Elcanidae (Orthoptera, Elcanoidea), Cascadelcana virginiana n. gen. n. sp., is described based on a forewing specimen from the Upper Triassic (Norian) Cow Branch Formation in the Solite Quarry Lagerstätte near the North Carolina-Virginia boundary, USA. It is distinguished from other elcanid species by its RP+MA1 with...
Article
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A new genus and species of fossil caddisfly (Insecta: Trichoptera) from the Lower Eocene (Ypresian) Green River Formation of Colorado is described. Litholimnephilops yinani gen. et sp. nov. is the first adult caddisfly to be described from the Green River Formation, and is characterized by large adult body size, presence of ocelli, dark leg spines,...
Article
Two new species of the extinct family Araripeneuridae are described from the Lower Cretaceous Crato Formation of Brazil: Caririneura macrothoracica sp. nov. and Cratoneura minor sp. nov. Caririneura damianii Martins-Neto, 1992a and C. crassatella Martins-Neto and Vulcano, 1997 are transferred to the genus Araripeneura Martins-Neto and Vulcano, 1989...
Article
Two new species of Dentridactylus Günther (Orthoptera: Tridactylidae) are described and illustrated: Dentridactylus quadratus sp. nov. and Dentridactylus truncatus sp. nov. The new species represent the first record of the subfamily Dentridactylinae from Colombia and the second record for the Americas. Both come from the south of the country and ar...
Article
A new genus and species of Elcanidae (Orthoptera: Elcanoidea) is described from the Yixian Formation of northeastern China. Jeholelcana yanensis gen. et sp. nov., represents the first Archelcaninae to be formally described from the Mesozoic of China. The new genus with both forewing and hind tibiae well preserved differs from all other Archelcanina...
Article
The name Gondwanagaricites magnificus, incorrectly published in a previous paper, is made valid and new MB numbers assigned.
Article
The Cretaceous lacewing family Babinskaiidae comprises nine species in seven genera, of which two new genera and one new species are described herein: Parababinskaia elegans gen. et sp. nov. from the late Aptian Crato Formation of Brazil, and Pseudobabinskaia gen. nov. (with the only species Pseudobabinskaia martinsnetoi Lu et al., 2017, comb. nov....
Article
A new fossil mushroom is described and illustrated from the Lower Cretaceous Crato Formation of northeast Brazil. Gondwanagaricites magnificus gen. et sp. nov. is remarkable for its exceptional preservation as a mineralized replacement in laminated limestone, as all other fossil mushrooms are known from amber inclusions. Gondwanagaricites represent...
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Six heteropteran species are reported for the first time from Illinois based on recently caught specimens from various bioinventories: the flat bug Neuroctenus pseudonymus Bergroth (Aradidae: Mezirinae), the stilt bug Metacanthus multispinus (Ashmead) (Berytidae: Metacanthinae), the leaf-footed bugs Anasa repetita Heidemann and Chelinidea vittiger...
Article
Tridactylus galla was described by Henri de Saussure (1895) on the basis of a single adult female collected during Vittorio Bottego’s first expedition to the Horn of Africa in 1892 and 1893. The species appears in lists compiled by Fenizia (1896), Lucas (1898) and Kirby (1906), but aside from a brief mention by Gunther (1995), is entirely overlooke...
Poster
Mud crickets and pygmy mole crickets are small insects that belong to the superfamily Tridactyloidea (Orthoptera: Caelifera). These crickets are mostly associated with freshwater ecosystems; they display a great potential for research in climatic change because of their association with these aquatic environments that are currently threat. However,...
Article
Two new genera and species of Megaloptera are described from the Lower Cretaceous Crato Formation of Brazil. Cratocorydalopsis brasiliensis gen. et sp. nov. and Lithocorydalus fuscata gen. et sp. nov. are both placed within the family Corydalidae. The specimens represent the first Cretaceous examples of adult megalopteran body fossils not preserved...
Article
Henri de Saussure described 82 species currently placed in the superfamilies Rhaphidophoroidea, Stenopelmatoidea and Tettigonoidea. The names are listed alphabetically, and the location of the type material (if known) and the current nomenclatural combination are given. When there is primary type material in the Muséum d’histoire naturelle de Genèv...
Article
Fossil insects from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil are preserved as goethite replacements in laminated limestones of lacustro-lagoonal origin. They display remarkable degrees of morphological detail down to the macromolecular level in some examples. We document the fidelity of preservation and reveal an astonishi...
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Three new species of Ripipteryx Newman (Orthoptera: Tridactyloidea: Ripipterygidae) are described from Colombia; namely R. diegoi sp. n. (Forceps Group) and R. guacharoensis sp. n. (Marginipennis Group) from Parque Nacional Natural Cueva de los Guacharos in Huila, and R. gorgonaensis sp. n. (Crassicornis Group) from Parque Nacional Natural Gorgona...
Article
Orthoptera is the most diverse order among the polyneopteran groups and includes familiar insects, such as grasshoppers, crickets, katydids, and their kin. Due to a long history of conflicting classification schemes based on different interpretations of morphological characters, the phylogenetic relationships within Orthoptera are poorly understood...
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A new genus and species of pygmy grasshopper (orthoptera: Tetrigidae) is described from Early Miocene (Burdigalian) Dominican amber. Electrotettix attenboroughi Heads & Thomas, gen. et sp. n. is assigned to the subfamily Cladonotinae based on the deeply forked frontal costa, but is remarkable for the presence of tegmina and hind wings, hitherto un...
Article
The Muséum d'histoire naturelle de Genève (MHNG) collection, which includes that of Henri de Saussure, contains many type specimens of Orthoptera from the Americas. Most of these were deposited by the North American entomologists Lawrence Bruner and Morgan Hebard, who received loans from or worked directly on the collection, even though their main...
Article
Henri de saussure described 403 species or subspecies in the superfamily Grylloidea. The names are listed alphabetically, and the location of the type material (if known) and the current nomenclatural combination are given. When there is primary type material in the Muséum d’histoire naturelle de Genève (MHNg), the sex, verbatim label data and cond...
Article
The first fossil representative of Melanoplus differentialis (Orthoptera: Acrididae: Melanoplinae) is documented from Late Pleistocene asphalt deposits of the McKittrick tar pits in southern California, USA. The specimen, an incomplete but well preserved adult male, is identified as M. differentialis based on the structure of the terminalia and in...
Article
: Three new genera and four new species of the extinct mantidfly subfamily Mesomantispinae (Insecta: Neuroptera: Mantispidae) are described from the Lower Cretaceous Yixian Formation of Liaoning and the Middle Jurassic Jiulongshan Formation of Inner Mongolia: Archaeodrepanicus nuddsi gen. et sp. nov., A. acutus gen. et sp. nov., Sinomesomantispa mi...
Article
AbstractA new species of the cricket genus Proanaxipha Vickery & Poinar (Orthoptera: Gryllidae: Pentacentrinae) from Early Miocene Dominican amber is described and illustrated. Proanaxipha madgesuttonae sp. n. is distinguished from congeners by: (1) head capsule bearing a distinctive posteriorly bilobed colour spot on the vertex; (2) presence of cr...
Article
The first record of a protogrylline orthopteran (Ensifera: Grylloidea: Protogryllidae) from the Middle Jurassic of India is provided. Protogryllus (Protogryllus) lakshmi, new species, was recovered from the Kota Formation in Andhra Pradesh and is compared with other species in the genus. General comments are provided on Protogryllidae and the diver...
Article
An annotated list of the Orthoptera (Insecta) species described by Henri de Saussure, with an account of the primary type specimens housed in the Museum d'histoire naturelle de Geneve, Part 1: The Tridactyloidea (Caelifera). - Henri de Saussure described thirty species or subspecies in the superfamily Tridactyloidea (Orthoptera: Caelifera). The nam...
Article
Two new mole crickets of the genus Scapteriscus Scudder (Orthoptera: Grylloidea: Gryllotalpidae) are described from Colombia: S. cerberus, sp. nov. and S. zeuneri, sp. nov. The first species can be recognized by its robust appearance, large pronotum and small separated ocelli. In contrast, the second species is characterized by its large ocelli whi...
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A new species of the genus Ripipteryx (Orthoptera: Tridactyloidea: Ripipterygidae) from the Toledo District of southern Belize is described and illustrated. Ripipteryx mopana sp. n. is placed in the Scrofulosa Group based on its elaborately ornamented frons and is readily distinguished from its congeners by the fusion of the superior and inferior f...
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The fossil orthopteran Brauckmannia groeningae Martins-Neto (Orthoptera, Ensifera) from the Early Cretaceous Crato Formation of Brazil, currently misplaced at both the genus and family level, is transferred to the family Schizodactylidae and assigned to the extant genus Schizodactylus Brullé; ergo, Brauckmannia enters synonymy under Schizodactylus...
Article
Two new Tridactyloidea of the families Ripipterygidae and Tridactylidae are described from the Early Miocene (Burdigalian) amber of the Dominican Republic, namely: Archaeoellipes engeli n. gen., n. sp. (Tridactylidae: Tridactylinae) and Mirhipipteryx antillarum n. sp. (Ripipterygidae). These new taxa represent the fi rst record of Tridactyloidea in...
Article
A new species of Ripipteryx Newman (Orthoptera: Tridactyloidea: Ripipterygidae) is described from a fragment of primary Andean forest on the slopes of the extinct Pasochoa volcano in Pichincha Province, northern Ecuador. Ripipteryx pasochoensis sp. nov. is distinguished from congeners by the morphology of the male terminalia, particularly the struc...
Article
A new species of Ripipteryx Newman (Orthoptera: Tridactyloidea: Ripipterygidae) is described from a fragment of primary Andean forest on the slopes of the extinct Pasochoa volcano in Pichincha Province, northern Ecuador. Ripipteryx pasochoensis sp. nov. is distinguished from congeners by the morphology of the male terminalia, particularly the struc...
Article
A new spider cricket (Orthoptera: Gryllidae: Phalangopsinae) is described from an adult female preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Araneagryllus dylani gen. et sp. nov. represents the first fossil record of Phalangopsinae, and is assigned to the tribe Luzarini, subtribe Amphiacustina stat. nov. A cladistic an...
Article
Gastrimargus Saussure, 1884 includes 23 species of economically important locustine grasshoppers (Acrididae: Oedipodinae: Locustini) distributed throughout Africa, Australia and Southeast Asia (Ritchie, 1982). Whilst reviewing oedipodine genus-group names in the Orthoptera Species File Online (Eades & Otte, 2009) we encountered an unused senior hom...
Article
A new genus and species of Tridactylidae (Orthoptera: Caelifera: Tridactyloidea) is described from mid-Cretaceous Burmese amber. Burmadactylus grimaldii gen. et sp.nov. is the first tridactylid to be formally described from a Cretaceous amber and is assigned to the extant subfamily Dentridactylinae. The new genus is distinguished from all other Den...
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The morphology of the enigmatic, Mesozoic, aquatic insect family Chresmodidae is redescribed and its phylogenetic affinities among the polyneopterous orders discussed. Study of the complete venation of both fore-and hind wings observed in some specimens from the Spanish Barremian, permit us to postulate the hypothesis that the family belongs to the...
Article
Eoproscopia martilli gen. et sp. nov. is described from the Early Cretaceous (Aptian) Crato Formation Lagersttte of Ceará State, north-east Brazil. The new taxon is assigned to the extant family Proscopiidae and represents the first occurrence of the group in the fossil record. Eoproscopia is similar to crown group proscopiids in its stick-like hab...
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The morphology of the enigmatic, Mesozoic, aquatic insect family Chresmodidae is redescribed and its phylogenetic affinities among the polyneopterous orders discussed. Study of the complete venation of both fore- and hind wings observed in some specimens from the Spanish Barremian, permit us to postulate the hypothesis that the family belongs to th...
Article
Recent records of the ground-hopper Tetrix subulata (L.) from Yorkshire are summarised. Two colour forms were present. Many of the populations were on acid soils suggesting the species has a broad pH tolerance and may have been overlooked in habitats previously considered unsuitable. The possibility of range extension northwards in this species and...
Article
A new species of Yuripopovia Jarzembowski (Coleorrhyncha: Progonocimicidae) is described from the Lower Cretaceous Vectis Formation (Wealden Group) of the Isle of Wight, southern England. Yuripopovia vectense sp. nov. differs from Yuripopovia woottoni Jarzembowski in the following characters: Sc undivided, branching from R basally; R1 ,R 2 and RS e...
Article
A new species of katydid is described from the Lesser Antillean island of St Maarten, Dutch West Indies. Phoebolampta caeruleotergum sp. n. (Tettigoniidae: Phaneropteri-nae) differs from congeners in having a deeply incised humeral sinus, ventral valvulae of the ovi-positor broad, at least half the width of the gonoplac, and male cerci shorter than...
Chapter
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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 chapter...
Article
HEADS, S. W. 2005. A new caddisfly larval case (Insecta, Trichoptera) from the Lower Cretaceous Vectis Formation (Wealden Group) of the Isle of Wight, southern England. Proceedings of the Geologists' Association, 117, 307-310. A new species of trichopteran larval case is described and named Piscindusia complexa isp. nov. The holotype specimen occur...
Article
Two new genera and four new species of Palaeontinidae (Cicadomorpha), Baeocossus finchae gen. et sp. nov., Colossocossus loveridgei gen. et sp. nov., C. rugosa gen. et sp. nov. and Parawonnacottella penneyi sp. nov. are described from the Nova Olinda Member of the Upper Aptian (Lower Cretaceous) Crato Formation of Ceará, northeast Brazil. The diagn...
Article
  A new species of palaeoleontid neuropteran Baisopardus cryptohymen sp. nov. is described. The holotype is remarkable for the preservation of fine body structures and colour patterning on all of its four wings. The specimen is nearly complete, lacking only the abdomen and the right mesothoracic leg, and is preserved as a black-brown compression in...
Article
Two new Palaeontinidae (Insecta: Cicadomorpha), Baeocossus fortunatus n. sp. and Colossocossus bechlyi n. sp. are described from the Lower Cretaceous (Aptian/Albian) Nova Olinda Member of the Crato Formation, Northeast Brazil. The diagnoses of the genera Baeocossus MENON, HEADS & MARTILL, and Colossocossus MENON, HEADS & MARTILL are revised and the...

Citations

... In Asia, Mesozoic (late Cretaceous) ants are reported from Kazakhstan and Myanmar (Lapolla et al. 2013;Barden 2017) while Cenozoic ants are reported from India (early Eocene), Iran (Ataabadi et al. 2019) and China (early Eocene, early Miocene, middle Miocene) (Zhang 1989;Lapolla et al. 2013;Wang et al. 2021). Fossil Dolichoderinae are known from Myanmar (Perrichot 2019), and they are also distributed in Thailand nowadays (Jaitrong and Nabhitabhata 2005). ...
... 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(Barling et al., , 2020(Barling et al., , 2021Os es et al., 2016;Bezerra et al., 2018Bezerra et al., , 2020Bezerra et al., , 2021Dias 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. ...
... In addition to the assassin bug, this fossil also has a rove beetle larva next to it, which belongs to Coleoptera, and although it is documented that the assassin bugs prey on Coleoptera as well (Youssef and Abd-Elgayed, 2015;Loko et al., 2022), there is no direct evidence that this assassin bug was preying on the rove beetle larva in this fossil, but the leg of the assassin bug was in contact with the carapace of the larva. Coincidentally, the assassin bug fossil documented by Swanson also shows Coleoptera (Swanson et al., 2021), and the leg of the assassin bug is also in contact with the body of the beetle. ...
... Fossil insects and spiders from the Lower Cretaceous Crato Formation of Brazil are preserved as 3D goethite replacements with internal voids filled with calcite and can be relatively high relief (Selden et al., 2006;Downen & Selden, 2021). The preservation of arthropods in the Crato Formation was likely facilitated by microbial mats (Varejão et al., 2019;Barling et al., 2020;Iniesto et al., 2021;Dias & Carvalho, 2022). However, the internal anatomy of the cephalothorax of the Crato spiders are preserved in contrast to the Aix-en-Provence spider molds. ...
... Fossil Dolichoderinae are known from Myanmar (Perrichot 2019), and they are also distributed in Thailand nowadays (Jaitrong and Nabhitabhata 2005). In Asia, fossil Orthoptera are reported from Iran (Oligocene) (Ataabadi et al. 2019), Kazakhstan (middle-late Jurassic), Japan (early Cretaceous) (Gu et al. 2010), China (Triassic, mid-late Jurassic) (Gorochov 2003;Gu et al. 2010;Xu et al. 2020) and South Korea (early Cretaceous) (Kim et al. 2021). In Southeast Asia, fossil Grylloidea are reported from Myanmar (middle-late Cretaceous) (Poinar et al. 2020;Yuan et al. 2022). ...
... The genus is readily distinguished from Mirhipipteryx, the only other genus in the family, by its comparatively larger size (body 5.5-14.0 mm long), interocular distance at least half the width of the compound eyes, metatarsus approximately equal in length to the metatibial spurs, and the distinctly sclerotized lateral valvulae of the phallus (Günther 1969;Heads 2010). Species of the genus are usually black or very dark brown, often with contrasting white, yellow and occasionally red markings (Heads 2010). ...
... Enquanto na Formação Crato os insetos correspondem a uma fundamental parcela da fauna registrada, na Formação Romualdo estes animais não são frequentes, provavelmente pela unidade se tratar de sucessões sedimentares formadas em ambiente marinho, o que diminui o potencial de preservação dos insetos. Até o momento tem-se apenas o registro de um táxon de Insecta, correspondendo a um Grylloidea reconhecido em uma concreção carbonática no nível dos folhelhos negros (Freitas et al., 2016;Heads et al., 2018). O baixo grau de preservação do espécime, provavelmente por ter sido transportado por longas distâncias do habitat terrestre até o ambiente marinho, impede seu enquadramento em níveis taxonômicos menores. ...
... Ripipterygidae are characterized by unsegmented cerci (the cerci are two-segmented in Tridactylidae), uninflated mesotibiae (the mesotibiae are distinctly inflated in Tridactylidae) and modified paraproctal lobes (in Tridactylidae, the paraproctal lobes are unmodified and cylindrical in shape) [2][3][4][5][6][7]. Ripipterygids mostly existnear freshwater habitats, such as streams, lakes, ponds, sandy banks, and floodplain areas, where they live mostly in sandy areas [5,8,9]. ...
... Within the Order Orthoptera, the phylogenetic position of the extinct superfamily Elcanoidea remains unsettled; they are considered to belong to the Suborder Ensifera (Zessin, 1987;Kukalov a-Peck, 1991;Carpenter, 1992;Gorochov, 1995;Gorochov et al., 2006), or treated as a sister group to the Suborder Caelifera (B ethoux and Nel, 2002), while Gorochov and Rasnitsyn (2002) even proposed a possibility of being a sister group to all other orthopterans. The Elcanoidea is divided into two families, Elcanidae and Permelcanidae (Grimaldi and Engel, 2005;Gorochov et al., 2006;Fang et al., 2018b). The Permelcanidae includes six species of three genera, known from the Lower Permian to the Lower Triassic (Tillyard, 1937;Sharov, 1968), while the Elcanidae contains more than 50 species belonging to fifteen genera usually from the Mesozoic (Gorochov et al., 2006;Cigliano et al., 2018). ...
... Elcanidae is a diverse group of Mesozoic orthopterans that can be easily distinguished from other Orthoptera by the tegminal venation and the special spurs on the metatibia. The fossil records of the Elcanidae mainly extended from Triassic to Cretaceous and distributed in Europe, Asia, North and South America (Handlirsch, 1906;Sharov, 1968;Martins-Neto, 1991;Grimaldi & Engel, 2005;Gorochov et al., 2006;Poinar et al., 2007;Peñalver & Grimaldi, 2010;Fang et al., 2015;Fang et al., 2018a, b;Heads et al., 2018;Tian et al., 2019a, b;Gu et al., 2020;Kim et al., 2021). Recently, a new record from France has been assigned to the Elcanidae, suggesting that elcanid-form insects may exist to the Paleogene (Schubnel et al., 2020). ...