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Basiala of Hennigmatinae: A, B, Trihennigma zavattierii gen. et sp. nov.: holotype MCNAM-PI 24607, part and inverted counterpart; C, Metatrichopteridium cladistorum (Shcherbakov, 1995): holotype PIN 3965/2789. Abbreviation: aialular incision.
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The first Triassic dipteran from South America is described based on an isolated wing from the lower Upper Triassic de-posits of Argentina (Mendoza Province, Potrerillos Formation, Quebrada del Durazno locality). Trihennigma zavattierii gen. et sp. nov. is a member of the Mesozoic family Hennigmatidae, previously recorded only from Eurasia. A key f...
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Context 1
... agree with Ansorge (2001) that the subcostal veinlets mentioned in the original description of Hennigma are artifacts of preservation and are absent in the holotype of M. cladistorum, as well as in other hennigmatids (Figs 1, 5, 6). However, his redrawing of the wing base is incorrect (Ansorge 2001: Fig. 3c): cu-a is certainly preserved as it was shown in Shcherbakov et al. (1995: Fig. 33) and can be seen in Trihennigma gen. nov.; moreover, in both genera, 2A is not marginal and a small alula is visible in M. cladistorum and is likely to be present in the new genus (Figs 5, ...
Context 2
... wings of both Metatrichopteridium and Trihennigma, the alula and alular incision are present, and the latter is situated just at the level of the humeral vein (Figs 5, 6; compare to Hennig 1968: Fig. 7). So, even according to Stary, it is a true alular incision. However, a true vein, pigmented, sclerotized, and without any break beyond anal loop-and thus undoubtedly not an "alular impression"-is clearly visible up to this incision. It is worth to mention that the shape of the considerably protruding anal lobe and the incision with callus in Mesozoic Metatrichopteridium are very similar to those in at least some Blephariceridae, which have no true alular incision according to Stary, because this angle at the base of the anal lobe is situated markedly beyond the humeral ...
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Citations
... It shows two r-m, the first connecting R 4+5 in the usual position and the second additional vein, connecting Rs and M 1+2 stems in the proximal part of the wing; the latter was considered a character of generic value (Szadziewski et al. 2018). In contrast to some other psychodomorph families, extant Tanyderidae and extinct Hennigmatidae, where it is a character of generic level (Lara & Lukashevich 2013), additional crossveins are extremely rare in Ptychopteridae. Although we have only a single wing on our disposal, we prefer to establish a new genus which is based not only on the additional crossvein but on atypical curved R 4 . ...
First Upper Triassic dipterans (Nematocera) are described based on isolated wings from Fuchsberg near Seinstedt (Sevatian, uppermost Norian of Lower Saxony, Germany): Crivoptychoptera nebrias gen. et sp. nov. (Psychodomorpha, Ptychopteridae), Ptychopteridae incertae sedis, Dohloboyia triassica gen. et sp. nov. (Bibionomorpha, Boholdoyidae) and Chironomidae incertae sedis (Culicomorpha). All taxa under discussion present the oldest occurrences of their families. Geological background and composition of the Upper Triassic insect fauna of Seinstedt are briefly discussed.
... Since 1990, the paleoentomology in Argentina has had a significant progress. Abundant insect fossils reported from different Triassic continental outcrops indicate high levels of richness, similar to those observed in only a few other basins in the world (e.g., Cairncross et al., 1995;Jell, 2004;Grimaldi & Engel, 2005;Zheng et al., 2018 Gallego, 1999Gallego, , 2006Gallego, , 2001Gallego, , 2009Martins-Neto et al., 2003, 2006a, 2006bGallego et al., 2011;Lara et al., 2012Lara et al., , 2015Lara et al., , 2019aLara et al., , 2019bLara et al., , 2021Lara et al., , 2023Lara & Lukashevich, 2013;Lara, 2016;Lara & Wang, 2016;Bustos-Escalona, 2020). About 90% of the Triassic fossil insect record in Argentina comes from the Cuyana and Ischigualasto-Villa Unión basins ( Fig. 2; Tab. 2). ...
... (Figs. 6, 7). (Lara & Lukashevich, 2013;Lara et al., , 2015Lara et al., , 2019aLara et al., , 2019bLara et al., , 2023Lara, 2016;Lara & Aristov, 2016;Lara & Wang, 2016;Lara & Bashkuev, 2020;Cariglino et al., 2021). Besides the numerous first taxonomic mentions of insect taxa at various levels (i.e., Order, Family, Genus and species) for South America, among some of the most remarkable findings are the largest collection of scytinopterid hemipterans (Lara, 2016;Wang et al., 2018;Lara et al., 2022Lara et al., , 2023; insect nymphs and larvae (Lara et al., 2019b); the earliest dipterans (Hennigmatidae, Lara & Lukashevich, 2013) and hymenopterans (Xyelidae, ; the first record of Eoscarterellidae (Hemiptera), which supports a Gondwanan origin for the family (Lara & Wang, 2016); the first record of adult grylloblattids (Geinitziidae, Lara & Aristov, 2017); and the earliest record of fossil insect oothecae (Dictyoptera) (Cariglino et al., 2020). ...
... (Lara & Lukashevich, 2013;Lara et al., , 2015Lara et al., , 2019aLara et al., , 2019bLara et al., , 2023Lara, 2016;Lara & Aristov, 2016;Lara & Wang, 2016;Lara & Bashkuev, 2020;Cariglino et al., 2021). Besides the numerous first taxonomic mentions of insect taxa at various levels (i.e., Order, Family, Genus and species) for South America, among some of the most remarkable findings are the largest collection of scytinopterid hemipterans (Lara, 2016;Wang et al., 2018;Lara et al., 2022Lara et al., , 2023; insect nymphs and larvae (Lara et al., 2019b); the earliest dipterans (Hennigmatidae, Lara & Lukashevich, 2013) and hymenopterans (Xyelidae, ; the first record of Eoscarterellidae (Hemiptera), which supports a Gondwanan origin for the family (Lara & Wang, 2016); the first record of adult grylloblattids (Geinitziidae, Lara & Aristov, 2017); and the earliest record of fossil insect oothecae (Dictyoptera) (Cariglino et al., 2020). ...
A state of the art of the insects collected and described from different Permian and Triassic units in South America is given, accompanied by their brief historical record (e.g., collectors, taxonomic revision) and new high-quality photos in color. Likewise, we succinctly provide data on the paleodiversity and paleoecology of the entomological assemblages from the different units. Finally, we discuss the South American paleoentomological records in relation to equivalent deposits worldwide. The thorough, updated review of fossil insect faunas in this paper will help settling the bases for future taxonomic, diversity and ecological studies during a time that comprised the evolutionary history of insects through two key episodes of the geological record: the end-Permian mass extinction (EPME, ⁓252 Ma) and the Carnian Pluvial Event (CPE, ⁓234–232 Ma).
... As mentioned in Lara et al. ( , 2021Lara et al. ( , 2022 and Lara & Bashkuev (2020), the frequency of these types of elements is probably associated to the robust and flexible exoskeleton (forewings), increasing their preservation potential in the fossil record. The remaining fossil insects, in abundance order, include membranous forewings of odonatans, mecopterans (Permochoristidae, Parachoristidae), orthopterans (Haglidae), dipterans (Hennigmatidae), grylloblattids (Geinitziidae), plecopterans, neuropterans, miomopterans, and elytra of beetles (Martins-Neto et al., 2008;Lara & Lukashevich, 2013;Lara et al., 2015Lara et al., , 2022Lara, 2016;Lara & Aristov, 2017;Lara & Wang, 2016;Lara & Bashkuev, 2020). ...
... A rich array of fossils has been studied from the Potrerillos and Cacheuta formations outcropping south of Cerro Cacheuta region, southernmost end of the Precordillera (Mendoza Province). These include well-known plant assemblages (Morel, 1994;Morel et al., 2010Morel et al., , 2011Bodnar et al., 2020;Cariglino et al., 2021), palynofloras Zavattieri & Prámparo, 2006), insect remains (Martins-Neto & Gallego, 1999;Martins-Neto et al., 2003, 2008Martins-Neto & Gallego, 2006;Martins-Neto et al., 2008;Gallego et al., 2011;Lara & Lukashevich, 2013;Lara et al., 2015;Lara, 2016;Lara & Wang, 2016;Lara & Aristov, 2017;Lara et al., , 2022Lara & Bashkuev, 2020), plant-insect interactions Cariglino et al., 2021, fish remains (cf. Neochallaia sp.), and spinicaudatan carapaces of Eustheria forbesi (Gallego, 1999;Tassi, 2016). ...
A new scytinopterid (Hemiptera: Cicadomorpha), Duraznoscytinum aristovi gen. et sp. nov., is described. The specimen was recovered from the upper section of the Potrerillos Formation (Upper Triassic) at the Quebrada del Durazno locality, south of the Cerro Cacheuta, Cuyana Basin, in Mendoza, Argentina. This represents the first record of a complete forewing (tegminous + clavus) of a scytinopterid for the Triassic of South America. Additionally, we briefly discuss the Triassic scytinopterid distribution worldwide, and provide preliminary morphological differences observed between the taxa recorded in the Southern Hemisphere. The new fossil described herein contributes to the knowledge on the hemipterofauna diversity present at the south of the Cerro Cacheuta area during Early Mesozoic times.
... The fossil record of Diptera in the Triassic is poor: only a little over a dozen sites from all continents except Antarctica have yielded dipterans [47]. Most such sites have yielded but a few dipteran wings. ...
Immature Diptera are more diverse and abundant in fresh water than any other insect order. The question arises whether the earliest dipterans, known from the Upper Buntsandstein of Europe (early Anisian, Middle Triassic), already developed in water and whether such mode of life was inherited from the ancestor of the order or if, alternatively, the dipteran larvae were ancestrally terrestrial. Nematoceran immatures have become common and diverse in the Mesozoic fossil record since the Middle Jurassic, but the vast majority of them represent only two culicomorphan families, Chaoboridae and Chironomidae. Earlier records and records of immatures of other families from the Mesozoic are extremely rare. A total of four larvae and about 30 pupae have been described from Anisian assemblages of France and Spain. Among these, one larva clearly belongs to Culicomorpha, the infraorder most closely associated with fresh waters, and one larva to Bibionomorpha, the infraorder most closely associated with terrestrial habitats, while the rest are hard to classify. Nevertheless, most of the pupae are morphologically similar to the semi-aquatic pupae of extant Limoniidae and Ptychopteridae from wet habitats. The oligopneustic respiratory systems of the Anisian larvae and their apparently allochthonous burial also suggest their semi-aquatic development, which appears to have been the ancestral mode of life of Diptera. The absence of dipteran immatures in lacustrine Konservat-Lagerstätten of the Madygen and the Cow Branch Formations (Ladinian–Carnian and Norian, respectively) is explained by the aquatic conditions being unfavorable for insect immatures and invertebrates in general.
... As such, of the four insect orders with known extant leaf miners, three are represented within this unit. With the exception of Lepidoptera, forewings attributed to the Diptera and Hymenoptera, as well as coleopteran elytra and abdomens were recovered from the site (Martins-Neto et al., 2008;Lara, 2016;Lara and Lukashevich, 2013;Lara et al., 2012Lara et al., , 2014. ...
... Dipterans from the Quebrada del Durazno are assigned to Trihennigma zavattierii Lara and Lukashevich (Fig. 4.1), and belong to the Hennigmatidae (Infraorder Psychodomorpha), an extinct lineage of which only the wings are known, precluding any inferences about their development or feeding strategies (Lara and Lukashevich, 2013). ...
We describe two ichnotaxa —Cuniculonomus parallelus Givulescu emend. Robledo et al. and Foliomacula diffussus ichnogen. et ichnosp. nov.— of insect leaf mines from the Late Triassic of Mendoza, Argentina. The specimens, from the Potrerillos Formation at the Quebrada del Durazno locality, consist of a linear mine on a Yabeiella leaf, and a blotch mine on a Heidiphyllum leaf. The earliest convincing records of leaf mining are all linear mines, dating back to the middle-late Triassic, with the first blotch mines appearing from the Cretaceous onwards. The discovery presented here therefore represents the earliest record of a blotch mine, and demonstrates that strategies of endophytic herbivory diversified considerably earlier than previously thought. The abundant associated paleoentomological fauna from the locality provides a rare opportunity to consider possible producers of these interactions. A definitive identification of the miners however, remains highly speculative, partly due to the paucity of direct analogues of insect lineages with leaf mining larvae this early in the fossil record. The exceptional abundance and quality of fossil preservation of both plants and associated insects at the Quebrada del Durazno locality also provides an opportunity to re-evaluate plant host diversity and possible adaptations to mitigate insect herbivory. The insect mine on Yabeiella is the first, and to date only, record of this type of interaction on a gnetalean host. We consider the presence of dense pubescence on Yabeiella and its affiliated Fraxinopsis seeds, as a potential passive defense mechanism against insect herbivory rather than as an adaptation to environmental conditions.
... In particular, there are rich yields of wings of Coleoptera (Taldycupedidae) and Blattodea (Mancusoblattidae, Mesoblattinidae) amongst the insect fauna of this deposit 32,34 . Notably, the Momonoki Formation yields true flies 36 , albeit undescribed, which have only been recorded from a few deposits from the Middle-Late Triassic [46][47][48][49][50][51] . Although more than 6000 specimens of fossil insects are recorded from this formation 30,36 , the taxonomic diversity is largely unexplored, and numerous terrestrial and aquatic insect specimens remain undescribed. ...
The Late Triassic saw a flourish of plant–arthropod interactions. By the Late Triassic, insects had developed all distinct strategies of herbivory, notably including some of the earliest occurrences of leaf-mining. Herein we describe exceptionally well-preserved leaf-mine trace fossils on a Cladophlebis Brongniart fern pinnule from the Momonoki Formation, Mine Group, Japan (Middle Carnian), representing the oldest unequivocal leaf-mines from East Asia. The mines all display a distinctive frass trail—a continuous meandering line, which later becomes a broad band containing spheroidal particles—demonstrating larval development. Although the shapes of the frass trails are generally comparable to those of Lepidoptera or Coleoptera, they cannot be unequivocally assigned to a specific extant leaf-mining taxon. Furthermore, elemental analyses by X-ray fluorescence (XRF) reveals that the frass trail comprises phosphate coprolites. The quantitative variations in P, S, and Si between coprolites and leaf veins may reflect physiological processes (e.g., consumption, absorption, and excretion) mediated by plant chemicals. Our findings reinforce the idea that leaf-mining had become a pervasive feeding strategy of herbivorous insects by the Late Triassic.
... In particular, there are rich yields of wings of Coleoptera (Taldycupedidae) and Blattodea (Mancusoblattidae, Mesoblattinidae) amongst the insect fauna of this deposit 31,34 . Notably, the Momonoki Formation yields true ies 36 , albeit undescribed, which have only been recorded from a few deposits from the Middle-Late Triassic [44][45][46][47][48][49] . ...
The Late Triassic saw a flourish of plant–arthropod interactions. By the Late Triassic, insects had developed all distinct strategies of herbivory, notably including some of the earliest occurrences of leaf-mining. Herein we describe exceptionally well-preserved leaf-mine trace fossils on a Cladophlebis fern pinnule from the Momonoki Formation, Mine Group, Japan (Middle Carnian), representing the oldest unequivocal leaf-mines from East Asia. The mines all display a distinctive frass trail – a continuous meandering line, which later becomes a broad blotch containing spheroidal particles – demonstrating larval development. The shapes of these mines generally resemble those of nepticuloid moths, although they are absent from extant fern-mining assemblages. Furthermore, elemental analyses by X-ray fluorescence (XRF) reveals that the frass trail comprises phosphate coprolites. The quantitative variations in P, S, and Si between coprolites and leaf veins may reflect physiological processes (consumption, absorption, and excretion) mediated by plant chemicals. Our findings reinforce the idea that leaf-mining had become a pervasive feeding strategy of herbivorous insects by the Late Triassic.
... The lower Upper Triassic Potrerillos Formation (Cuyana Basin, Mendoza Province) exhibits one of the best plant assemblages known in southwestern Gondwana, regarding both the quality and quantity of its fossil plants (Zamuner et al., 2001;Artabe et al., 2007 and references therein). Moreover, in recent years numerous exquisitely preserved insects have been recovered in different outcrops from this unit (Lara, 2016;Lara et al., 2012Lara et al., , 2014Lara et al., , 2015Lara and Lukashevich, 2013;Lara and Wang, 2016;Lara and Aristov, 2017;Lara and Baskhuev, 2020;Cariglino et al., 2020a), providing further insights about the biotic interactions between plants and arthropods . In decreasing order of commonness, the flora from the Potrerillos Formation comprises umkomasialean seed ferns, ginkgoaleans, gnetaleans, voltzialean conifers, cycadaleans, peltasperms, ferns and sphenophytes ( Fig. 5.1-5.12). ...
Plant-insect interactions are among the most common biotic relationships, both in diversity and abundance. The fossil record reflects these associations as damage traces on the leaves, stems or seeds, allowing researchers to identify the type of interaction and infer about its insect producer. Here, we present an update of the current knowledge of Permian and Triassic floras with evidence of plant-insect interactions from southwestern Gondwana and list other numerous units with high potential for plant-insect interaction analyses. When present, we also add information about the associated paleoentomofaunas, as they offer a unique opportunity for inferences about the producers of the damage. The Permian–Triassic time interval is of major interest due to salient biotic and abiotic events of regional and global significance. Therefore, we discuss the main abiotic factors that could have acted as key drivers controlling the interactions between plants and insects. With this contribution we hope to inspire other researchers to consider looking for evidence of arthropod damage in fossil floras and thus adding valuable knowledge for understanding the dynamics of ancient ecosystems.
... Numerous fossils insect reported in Triassic continental sediments places Argentina as the main paleoentomological region of South America and one of the most important from southwestern Gondwana (Lara, 2016;Lara et al., 2017). The significant paleoentomological richness of the Argentinian basins include thousands of specimens of insects classified into 12 orders and 91 species, comprising fragmentary and occasionally articulated specimens preserved as impressions/compressions of wings, and parts or complete bodies, attributable mainly to blattids, beetles, and hemipterans, and in less proportion to orthopterans, mecopterans, miomopterans, grylloblattids, plecopterans, dipterans, hymenopterans, odonatans and glosselytrodeans (Martins--Neto and Gallego, 1999Gallego, , 2001Gallego, , 2006Gallego, , 2009Martins-Neto et al., 2003, 2005, 2006a, 2006bGallego et al., 2011;Lara, 2016;Lara and Lukashevich, 2013;Lara and Wang, 2016;Lara and Aristov, 2016;Lara and Bashkuev, 2020;Lara et al., 2012Lara et al., , 2014Lara et al., , 2015Lara et al., , 2017Bustos-Escalona, 2020). All these insect remains have been collected in the Los Rastros Formation (Carnian), Ischigualasto Villa-Unión Basin (La Rioja Province), Potrerillos Formation (Carnian), Cuyana Basin, and Llantenes Formation (Norian), Malargüe Basin (Mendoza Province) (Lara, 2016;Bustos-Escalona, 2020). ...
Two new hemipteran taxa are described: Dysmorphoptiloides losrastrosensis sp. nov. and Gualoscarta obscura gen. et sp. nov. (Auchenorryncha: Dysmorphoptilidae). Likewise, we report the first specimen of a corixid (Heteroptera: Nepomorpha) from the Triassic of Gondwana. The fossil material studied in this paper comes from the Gualo locality, the Carnian Los Rastros Formation, Ischigualasto-Villa Unión Basin, La Rioja Province, Argentina. From new available material, we present a new forewing reconstruction of Saaloscytina carmonae Martins-Neto et al., 2006a and discuss its taxonomic position. As an additionally contribution, three additional genera from the Los Rastros Formation are revised: Gualoscytina Martins-Neto and Gallego, 2003, Argentinopheloscyta Martins-Neto and Gallego, 2003, and Australocicada Martins-Neto and Gallego, 2001 are synonymized to Apheloscyta Tillyard, 1922 (Apheloscyta mayae comb. nov.), Mesoscytina Tillyard, 1919 (Mesoscytina forsterae comb. nov.), and Fulgobole Shcherbakov, 2011 (Fulgobole arcucciae comb. nov.), respectively. Finally, the authors discuss the paleobiological and paleogeographical implications of this new material from the Triassic of Gondwana.
... 140 M. B. Lara and A. Bashkuev significant palaeoentomological potential of the Argentinean Triassic sequences is based on more than 500 specimens of insects classified into 12 orders and 87 species attributable mainly to dictyopterans, hemipterans and coleopterans, and in less proportion to orthopterans, mecopterans, miomopterans, grylloblattids, plecopterans, dipterans, hymenopterans, odonatans and glosselytrodeans (Lara 2016). The insects recorded as fragmentary and occasionally articulated specimens with different preservation modes as impressions/molds of wings, part of bodies and complete bodies, come from Ischichuca (~Anisian-Ladinian) (Martins-Neto & Gallego 2009) and Los Rastros (Carnian) (Martins-Neto et al. 2005;Martins-Neto et al. 2006a;Martins-Neto et al. 2006b) formations, Bermejo Basin, San Juan and La Rioja provinces; Cerro de Las Cabras (~Anisian-Ladinian) and Potrerillos (Carnian) formations, Cuyana Basin, Mendoza Province (Martins-Neto et al. 2007;Martins-Neto et al. 2008;Gallego et al. 2011;Lara et al. 2012;Lara & Lukashevich 2013;Lara et al. 2014;Lara et al. 2015;Lara & Aristov 2016;Lara & Wang 2016;Lara et al. 2017); Llantenes Formation (Norian), Tronquimalal Group, Llantenes Depocenter, Malargüe Basin, Mendoza Province (Brauckmann et al. 2010) (Gallego 1997;Martins-Neto & Gallego 2001;Martins-Neto et al. 2003). ...
... In recent explorations carried out between 2015 -2017 in to the outcrops of the Carnian Potrerillos Formation at Puesto Miguez (= Minas de Petróleo) and Quebrada del Durazno localities (southern flank of the Cerro Cacheuta or Cacheuta peak), Cuyana Basin, Mendoza Province, several assemblages of fossil insects were collected. This insect fauna is one the most important within the Basin due to its diversity and exquisite preservation which include hemipterans, as the most abundant order, and in less proportion plecopterans, odonatans, orthopterans, grylloblattids, mecopterans, coleopterans, miomopterans, dipterans and hymenopterans (Lara & Lukashevich 2013;Lara et al. 2015;Lara 2016;Lara & Aristov 2016;Lara & Wang 2016;Lara et al. 2017). The insects collected belong to second stage (Anisian-Carnian) sensu Shcherbakov (2008), which marks the time of the rise of the typical Triassic entomofauna and appearance of many Meso-Cenozoic groups. ...
... The insects collected belong to second stage (Anisian-Carnian) sensu Shcherbakov (2008), which marks the time of the rise of the typical Triassic entomofauna and appearance of many Meso-Cenozoic groups. Besides, in the Argentinean Triassic localities have been also re-corded one of the earliest representatives of Diptera (Trihennigma zavattierii Lara & Lukashevich, 2013) and Hymenoptera (Potrerilloxyela menendezi Lara et al., 2014), supporting that the Triassic Period is marked by the beginning of the modern ecosystems. Therefore, the Potrerillos Formation offers an opportunity to study the diversity global recovery of insect taxa after of the end-Permian extinction. ...
New insect taxa, Tipuloidea cuyana sp. nov., Cuyanoscytinum arcucciae gen. et sp. nov. (Hemiptera: Scytinopteridae), Duraznochorista zavattierii gen. et sp. nov. (Mecoptera: Permochoristidae), and Potrerillopanorpa macloughlini gen. et sp. nov. (Mecoptera: Parachoristidae), are described from the Upper Triassic Potrerillos Formation, at the Quebrada del Durazno locality, in the Cuyana Basin, Mendoza Province, Argentina. The genera Tipuloidea Wieland, 1925 and Argentinocicada Martins-Neto & Gallego, 1999 (Chiliocyclidae) are transferred back from Chiliocyclidae to Scytinopteridae Handlirsch, 1906. The diagnosis and description of Tipuloidea rhaetica Wieland, 1925 are revised and emended. The family Parachoristidae is reported for the first time in the Triassic of South America. Argentinopanorpa miguezi Lara et al., 2015 is tentatively transferred to Parachoristidae based on new data. Distribution data of respective families in Gondwana is updated.
