Extinct Family Research Articles

New genus of falsiformicid wasps (Hymenoptera, Vespoidea: Falsiformicidae) with brachypterous male from middle Cretaceous Kachin amber in Myanmar

Falsiformicidae is a poorly known extinct family of aculeate wasps. It is known only as inclusions in various Cretaceous ambers. Brachypterous male Falsiformix pedestris Zhang and Rasnitsyn gen. et sp. nov. is described in the family Falsiformicidae based on a unique fossil from the middle Cretaceous Kachin amber. Occurrence and meaning of the male flightlessness in Hymenoptera is discussed. A combination of the flightless males with flying females occurs when mating takes place shortly after the eclosion, before adults disperse from confined space where they developed. An equally rare even if less exotic strategy with males and females both being flightless depends on the female pedestrian habits rather than on the pre-dispersal mating. Wide spatial (Angara Land, Laurussia Land and Gondwana Land) and narrow temporal distribution (Cenomanian and Turonian ages only) indicate Falsiformicidae as a possibly important stratigraphical indicator.

Revision and phylogenetic placement of one of the earliest freshwater gobies from the Lower Oligocene of Central Europe

ABSTRACT Gobioid fishes (Teleostei) are renowned for their remarkable species diversity in marine and freshwater environments, making their evolutionary history a fascinating study subject. Among their oldest known fossils is the small-sized freshwater goby ‘Gobius’ gracilis from the Lower Oligocene of Central Europe, which has not undergone a detailed investigation since its first description. Here, we present a comprehensive re-analysis of ‘G.’ gracilis using type specimens and new material from museum collections. We show that this species possesses a unique combination of traits, leading to the establishment of a new genus, †Laubeichthys gen. nov. Phylogenetic analyses using Bayesian Inference (BI) recover †Laubeichthys gen. nov. as a stem member of the extinct family †Pirskeniidae and suggest a sister relationship between the family Thalasseleotrididae (ocean sleepers) and †Laubeichthys gen. nov. + †Pirskeniidae. Maximum parsimony analyses do not contradict such a relationship, but the trees are less resolved. Today, the Thalasseleotrididae includes four marine species that are restricted to the Southwest Pacific. In contrast, †Pirskeniidae and †Laubeichthys gen. nov. inhabited freshwater environments in Central Europe. Our findings provide new insights into the evolutionary history and historical biogeography of gobioid fishes and highlight the dynamic nature of their past diversity and distribution.

Morphometrics of the mandible of Metoposaurus krasiejowensis Sulej, 2002 and its ecological implications

Amphibians, due to their ecological plasticity, are some of the best environmental indicators among vertebrates nowadays and in the fossil record. One such example is the extinct family Metoposauridae Watson, 1919. Metoposaurids were abundant amphibians in Late Triassic Pangea. The remains of the genus Metoposaurus Lydekker, 1890 have been found in Germany, Poland and Portugal with three species, respectively Metoposaurus diagnosticus (Meyer, 1842), Metoposaurus krasiejowensis Sulej, 2002 and Metoposaurus algarvensis Brusatte, Butler, Mateus and Steyer, 2015. Since the majority of studies concern the skull and the pectoral girdle, in this work M. krasiejowensis has been analysed through a morphometric study of the mandible. This was made possible by the high abundance of fossils found in Krasiejów (SW Poland) in the last 20 years. The characteristics considered are the morphology of the mandible corpus and its most relevant bones, the adaptation to stress during biting and the dermal ornamentation. The results reveal that not only do these characters have great intraspecific variability, but that at least two groups of a single population of M. krasiejowensis probably had different lifestyles, one more aquatic and the other more terrestrial.

A New Intriguing Teleost from the Albian Muhi Quarry, Central Mexico, and Early Euteleostean Diversification

The Albian Muhi Quarry of Central Mexico has produced a variety of fishes, among which the teleosts are the most diversified, including representatives of stem groups together with a variety of crown groups. A new genus and species, †Xeneichthys yanesi, is described based on a unique combination of characters, such as a dorsoventral elongation and narrowing of infraorbitals 2 and 3, preopercle, opercle, and cleithrum; absence of some bones, such as the infraorbital 5, interopercle, and pelvic plate and fin; presence of an elongated urostyle and a membranous outgrowth or stegural on the first uroneural; and cycloid and ctenoid scales on the flanks. Due to its combination of characters, †Xeneichthys yanesi is interpreted as belonging to a new extinct family, †Xeneichthyidae, which is considered as an Euteleostei or Euteleosteomorpha incertae sedis. The fish faunas of another Albian quarry, Tlayúa of Tepexi de Rodriguez, Puebla, and Muhi Quarry are compared, and although both contain taxa that can be assigned to stem teleosts (e.g., †ichthyodectiforms and †crossognathiforms), both also contain crown teleosts (e.g., elopomorphs, clupeomorphs, and euteleosts)—all of which are endemic to their respective quarries and make them important centers of diversification of fish faunas during the Early Cretaceous.

Filling the gap in the evolution of the genus Echinochara Peck (Clavatoraceae, Charophyta)

Echinochara is a poorly known charophyte genus of the extinct family Clavatoraceae that thrived in the Middle Jurassic–Early Cretaceous of North America, Europe, and North Africa. It represents the most ancient and plesiomorphic genus of this family. However, the fossil record of Echinochara is discontinuous, showing several gaps, one of them especially significant in evolutionary terms between the Kimmeridgian and the early Barremian. The new species Echinochara pontis sp. nov., described here from the late Berriasian of the Maestrat Basin (NE Spain), bridges, in part, this gap, shedding light on the evolution of the genus during the earliest Cretaceous. The fructification (utricle) has bilateral symmetry, formed by two superimposed series of bract cells that abaxially cover the gyrogonite. The inner series has three long cells that are born by a short basal cell and reach the oospore apex. The outer series shows a central fan of bract cells, composed of a small basal cell, triangular in shape, that bears three long cells growing upwards, which are in turn flanked by two long bract cells on each side. The central basal cell and these two lateral cells are directly attached to the base of the utricle. The thallus is corticated with cells twisting at an angle of 30°. The utricle of Echinochara pontis displays a combination of features present in Echinochara peckii (Bathonian?–Kimmeridgian) and Echinochara lazarii (Barremian–early Aptian). For instance, the bilateral symmetry of the utricle and the morphology of the inner bract-cell series allow comparison with E. lazarii, whilst the morphology of the outer bract-cell series is closer to E. peckii. Moreover, the thallus of the new species shows cortical cells with a twisting angle intermediate between that of Echinochara peckii (40°–45°) and Echinochara lazarii (not twisted). The unique combination of morphological characters in the utricle of Echinochara pontis fits well with its intermediate chronostratigraphic position between the two former species. Echinochara pontis formed monospecific meadows in brackish settings. A preference for brackish settings is also observed for Echinochara peckii and in the older populations of Echinochara lazarii, suggesting that this habitat is probably a basal character in the evolution of the genus Echinochara.

First report of adult †Burmecaelidae with description of a new species from mid-Cretaceous Burmese amber (Orthoptera: Caelifera)

An adult male of †Burmecaelinus armis Uchida, Husemann and Kotthoff, 2023, and a newly identified genus and species †Moban zhengzheminigen. et sp. nov., are reported based on two well preserved adult specimens from mid-Cretaceous Burmese amber. †M. zhengzheminigen. et sp. nov. can be distinguished by following characters: body with irregular small dark spots; pronotum is in uniform width, with dispersed shallow dents and two distinct transverse sulci, posterior angle is protruding forward; posterior thoracic process is shark-tooth shaped; pronotal disc without four longitudinal carinae; tip of apical spurs on metatibia not curved. The morphological diagnosis of the family †Burmecaelidae was emended, and the posterior thoracic process of adult †Burmecaelidae species is possibly specialized scutellum. We proposed a hypothesis that this extinct family was a sister group of Tridactyloidea and species of †Burmecaelidae lived in humid environment near water and had affinity for wet and semi-aquatic habitats.

Flat‐footed Females and Missing Males: A New Genus and Species of Pleciofungivoridae (Diptera: Bibionomorpha) Represents the First Record of the Family from Mid‐Cretaceous Kachin Amber

AbstractA new genus and species, Cretopleciofungivora simpsoni gen. et sp. nov., from the extinct family Pleciofungivoridae (Diptera: Bibionomorpha), is discovered in mid‐Cretaceous Kachin amber. Previously, this family was known only from imprints in sedimentary rocks of the Jurassic and the Lower Cretaceous. Discovery of a representative of Pleciofungivoridae in Kachin amber confirms the presence of the family in the Upper Cretaceous. The new species has a unique structure of fore tarsus, with lobed and extended tarsal segments II to IV, a feature hitherto known only in a few species of extant Sciaroidea. Although not particularly rare, the new species is currently known only from female specimens. Possible reasons for this phenomenon, very unusual in Sciaroidea, are briefly discussed, including parthenogenesis as a potentially plausible hypothesis.

Stable isotope analysis evidences dietary changes in toxodontids (Mammalia, Notoungulata) across the Neogene-Quaternary from Argentina

Toxodontidae was an extinct family of South American notoungulates, widely studied systematically, but not so much from an ecological perspective. This work aims to improve the knowledge about variations in diet and habitats of different representatives of this group in the context of biotic and abiotic events throughout the last 20 My. The tooth enamel carbon isotope composition (δ13C) of 7 genera recorded in 11 localities of Central, West and Northeastern Argentina, from the Early Miocene to the Late Pleistocene, was analyzed. The diet of the toxodontids studied in this work range from mainly C3 resources during the Miocene, mixed C3-C4 in the Pliocene, to a significant consumption of C4 plants during the Pleistocene. Differences in diet during the Late Pleistocene were also recorded, associated with geographic (i.e., Northeastern vs. Central) and temporal (i.e., glacial vs. interglacial periods) variations, evidencing the adaptive capacity of the group under ecological and environmental pressures.

Nuttalliellidae in Burmese amber: implications for tick evolution.

Ticks are composed of 3 extant families (Argasidae, Ixodidae and Nuttalliellidae) and 2 extinct families (Deinocrotonidae and Khimairidae). The Nuttalliellidae possess one extant species (Nuttalliella namaqua) limited to the Afrotropic region. A basal relationship to the hard and soft tick families and its limited distribution suggested an origin for ticks in the Afrotropics. The Deinocrotonidae has been found in Burmese amber from Myanmar and Iberian amber from Spain, suggesting a wider distribution of the lineage composed of Deinocrotonidae and Nuttalliellidae. The current study describes 8 fossils from mid-Cretaceous (ca. 100 Ma) Burmese amber: 2 Deinocroton species (Deinocroton bicornis sp. nov.; Deinocroton lacrimus sp. nov.), 5 Nuttalliella species (Nuttalliella gratae sp. nov., Nuttalliella tuberculata sp. nov., Nuttalliella placaventrala sp. nov., Nuttalliella odyssea sp. nov., Nuttalliella tropicasylvae sp. nov.) and a new genus and species (Legionaris nov. gen., Legionaris robustus sp. nov.). The argument is advanced that Deinocroton do not warrant its own family, but forms part of the Nuttalliellidae comprising 3 genera, Deinocroton, Legionaris nov. gen. and Nuttalliella). Affinities of Burmese tick fossils to the Australasian region, specifically related to rifting of the Burma terrane from northern Australia ~150 million years ago, suggest that Nuttalliella had a much wider distribution than its current limited distribution. The distribution of Nuttalliella likely stretched from Africa over Antarctica and much of Australia, suggesting that extant members of this family may still be found in Australia. Considerations for the geographic origins of ticks conclude that an Afrotropic origin can as yet not be discarded.

Primitive new termites (Blattodea, Termitoidae) in Cretaceous amber from Myanmar.

Mastotermitidae, the first-diverging extant family of termites, has only one relic extant species; however, this family had greater richness during the Mesozoic and Cenozoic eras. Fossil termites from the Cretaceous provide information on the early evolution of termites and the transition between extinct families. Herein, two new Mastotermitidae species found in upper Cretaceous (Cenomanian) Kachin amber are reported. One is a female imago described as Angustitermesreflexusgen. et sp. nov. and assigned to the subfamily Mastotermitinae. The other is Mastotermesreticulatussp. nov., which is described from an isolated forewing. With the comparison especially of the antenna and venation, these new mastotermitids further increase our knowledge of the diversity and morphology of Mastotermitidae during the Mesozoic.

Unravelling the mishmash: A new phylogeny for the family Empheriidae (Psocodea, Trogiomorpha) with a new genus and species from Cretaceous Charentese amber

The order Psocodea, including barklice, booklice, and parasitic lice, is diverse and widely distributed since the Cretaceous. That is particularly the case for the speciose extinct family Empheriidae (Psocodea, Trogiomorpha, Atropetae), recently fused with the ‘Archaeatropidae’. Understanding the evolution of barklice is dependent in part on studying this family, as its representatives have been found from the Early Cretaceous to the Eocene, surviving the K/Pg extinction event. The phylogenetic relationships of Empheriidae in relation to other families, such as Lepidopsocidae or Psoquillidae, have been extensively debated. However, distinguishing diagnostic characters for the Empheriidae has proven challenging. In this study, we describe the new empheriid Santonipsocus mimeticusgen. et sp. nov. from Cretaceous Charentese amber (France). It is the third empheriid species found in this locality. The new genus is compared with the other genera in the family, and Proprionoglaris guyoti and Proprionoglaris axioperierga are revised based on the type material and new specimens. We explore the phylogeny of Empheriidae, both the relationships with other families and the inner relationships between the genera, through maximum parsimony analysis and Bayesian inference analysis. Our results suggest that Empheriidae may represent a paraphyletic evolutionary grade to the rest of Atropetae. The phylogenetic relationships between genera align with the biogeography of the family and support previous hypotheses. In addition, we discuss the possible biology of the members of the family, shedding light on the evolutionary history of Empheriidae.

New Sinoalidae (Hemiptera, Cercopoidea) in Cenomanian Kachin amber, with notes on its paleobiogeographic implications

The froghopper extinct family Sinoalidae shows high paleo-diversity in mid-Cretaceous Kachin amber biota. The sinoalid genus Cretadorus Chen, 2020 is revised in this study. Araeoanasillus Poinar et Brown, 2023, tentatively placed in Sinoalidae with subfamily and tribe incertae sedis in original paper, is proposed herein to be synonymized under Cretadorus, leading to Cretadorus leptosomus (Poinar et Brown, 2023) comb. nov. A new species of this genus, Cretadorus qingyui sp. nov., is described and illustrated on the basis of an almost complete fossil trapped in Kachin amber. The Gondwanan origin of Burma Terrane suggests that Sinoalids in Kachin amber are likely one of residual groups of Gondwanan biotae, and so this family had wide paleobiogeographic distribution on the earth in the Jurassic. In the Early Cretaceous, the distribution range of Sinoalidae has reduced: it at least probably disappeared in northeastern Eurasia. This hemipteran lineage was extinct in the late Mesozoic owing to food crisis after the angiosperm floristic revolution.

An assessment of the anseriform affinities of “Telmabates” howardae

"Telmabates" howardae, identified from a tibiotarsus discovered in the early Eocene Las Flores Formation in Chubut Province, Argentina, was initially classified under the extinct family Presbyornithidae. However, this classification was later dismissed, and it was deemed incertae sedis. In this study, we suggest a new genus for "Telmabates" howardae and evaluate its relationships within Anseriformes. Its familial relationships remain unclear due to the location of the distal opening of the canalis extensorius, the shape of the sulcus m. fibularis (smooth and laterally bounded by a ridge, similar to that of Saintandrea or Dendrocygna), and the medial position of the condyles. However, unlike Anatidae and Romainvillidae, the condylus medialis is smaller and blunt, whereas in these taxa, it is large and narrow. This suggests that this species was more terrestrial and less adapted to swimming than Anatidae.

First record of fossil psocodeans in copula from mid-Cretaceous Burmese amber.

We report the first discovery of barklice preserved in copula from amber dating back to the mid-Cretaceous, thus documenting the oldest preserved reproductive behaviour of Psocodea. The new finding provides new insight into the evolution of copulatory behaviour in Trogiomorpha. Moreover, we describe and illustrate the new fossil material of Burmempheria densuschaetae (= Latempheria kachinensis) in the extinct family Empheriidae and Longiantennum fashengi in the extinct family Archaeatropidae, two diverse and widely distributed groups during the Cretaceous. No significant sexual dimorphism is recorded, apart from variation in size where males are larger. The taxonomy of Cretaceous empheriids from Burmese amber is revised and discussed.

A new species of Protopsyllidiidae (Hemiptera, Sternorrhyncha) from the Middle Jurassic of northeastern China.

Based on two compression fossils from the Middle Jurassic Jiulongshan Formation of Daohugou Village, Ningcheng County, Inner Mongolia, China, a new fossil species, Poljanka curticapillata L, Luo & Yao, sp. nov., of the extinct family Protopsyllidiidae is described. It is characterized by the flagellomeres 16 slender and 6 small subsections at apex, the presence of genae, the basal tarsomere almost equal to the apical tarsomere of fore and middle legs, and basal tarsomere about 2.11 times as long as apical tarsomere of hind leg, and the forewing veins R longer than M + Cu1, Cu1 with curved Cu1a and Cu1b, and Cu1a/Cu1b ratio 3.463.63. These body features and combination of forewing venation characteristics differ the new taxon from all other known species of Poljanka Klimaszewski, 1995. We also provide a key to the species of Poljanka and revise the diagnosis of the genus.

Fossil Trichoptera embedded in mid-Cretaceous Burmese amber

The paper gives an overview of Trichoptera found as adults in mid-Cretaceous Burmese amber from about 100 million years ago. Fifty-eight extinct species are listed, three of which are still described here: Paduniella cretaceasp. nov., Palerasnitsynus vilarinoisp. nov., Palleptocerus kuranishiisp. nov. The extinct subfamily Palerasnitsyninaestat. nov. of the family Xiphocentronidae is established and the extinct Bipectinata orientaliscomb. nov. is transferred from the family Calamoceratidae to the family Odontoceridae. The extinct family LepidochlamidaeWang et al., 2022, stat. nov. is transferred to the superfamily Leptoceroidea. The fifty-eight caddisflies of Burmese amber are distributed among twenty-one genera and fourteen families, of which fifteen genera and four families are also extinct. The large time distance between extinct and extant organisms makes the assignment to the extant genera and families difficult, because the higher taxa are defined according to the species living today and often do not or hardly correspond to the earlier species and their adaptations. Furthermore, in line with the hypothesis of a Gondwanan origin of Burmese amber, some embedded Trichopterans are discussed as relict descendants of Gondwanan Trichoptera, e.g. the family Palleptoceridae and the Xiphocentronid subfamily Palerasnitsyninae.

Taxonomic revision and type specimens’ location of jianghanichthyids (Ostariophysii: Cypriniformes) from Buxin Formation, Sanshui Basin, China

ABSTRACT The Paleogene nominal species Osteochilus sanshuiensis, Osteochilus longipinnatus, and Osteochilus laticorpus from Sanshui Basin, China, were originally assigned to the extant genus Osteochilus, in the family Cyprinidae. They were lately considered to belong to Catostomidae or the extinct family Jianghanichthyidae. However, the type specimens of the above nominal species have been deemed to be lost, which hindered the corroborated taxonomic assignment of the species. After multiple attempts and inter-institutional efforts, a series of original type specimens were recovered from a warehouse. We herein report the resurfacing and relocation of the specimens, as well as clarify the taxonomic status of those nominal species. Based on the type specimens and our recently collected specimens in the same locality, we propose to revise the three nominal species as one species †Jianghanichthys sanshuiensis comb. nov. One of the best-preserved original type specimens, SYSU R. 741177 is designated to be the lectotype.

A new Liassophlebiidae (Odonata: Heterophlebioidea) from strata close to the Triassic-Jurassic boundary in Somerset, UK

ABSTRACT Liassophlebiidae is an extinct family of damsel-dragonflies found in Upper Triassic and Lower Jurassic strata of Europe, Asia and Antarctica. Whilst Liassophlebiidae is well represented by Lower Jurassic fossils, their lowest occurrence in the Upper Triassic has hitherto only been suggested by three fragmentary specimens. These were originally ascribed to two species: Liassophlebia withersi and Liassophlebia batheri, but the latter is now considered nomen dubium. Here we describe a fourth, better preserved specimen that is likely to be Rhaetian (Late Triassic) in age. The specimen, BRSMG Cg3101 a+b, was collected from Bowdens Quarry, Somerset, UK, from the lower part of the White Lias Formation. The specimen comprises an incomplete forewing attributed to Liassophlebia due to: the small number of antenodals, antesubnodals and crossveins between RP and MA based RP3/4 in the base of RP2 opposite the subnodus; a straight and elongate secondary longitudinal vein in the postdiscoidal area; numerous cells and secondary veins in radial and median areas. The specimen is likely to represent a new species and provides stronger evidence than the previous three specimens of the presence of Liassophlebiidae during the late Rhaetian. Its stratigraphical position suggests that Liassophlebiidae arose in the immediate aftermath of the Triassic-Jurassic mass extinction.

New genus of Empheriidae (Psocodea; Trogiomorpha) from mid-Cretaceous Burmese amber

We characterise, describe and illustrate a new fossil genus and species, Paralellopsocus elongatus gen. et sp. nov., a small barklouse belonging to the extinct family Empheriidae (Trogiomorpha: Atropetae), based on specimens preserved in Burmese amber. These findings shed more light on Empheriidae and its possible senior synonymy with Archaeatropidae, as we discuss their classification and the value of their diagnostic characters. This new discovery further increases the number of described taxa in Empheriidae as the family was probably diversified and widely distributed during the Cretaceous.

EOCENE RHAMPHOSIDAE (TELEOSTEI: SYNGNATHIFORMES) FROM THE BOLCA LAGERSTÄTTE, ITALY

The Rhamphosidae is an extinct family of syngnathiform fishes from the lower Eocene deposits of Europe, primarily known from specimens derived from the Ypresian Konservat-Lagerstätte of Bolca (Verona province, Italy). A descriptive analysis of 28 specimens of Rhamphosus from Bolca revealed the existence of six species, showing a greater taxonomic diversity compared to the previous scenario of only two species (the type species Rhamphosus rastrum and Rhamphosus biserratus). Four new species are established herein: Rhamphosus bloti n. sp., characterized by a peculiar and unique rostrum with a discoid shape; Rhamphosus brevirostris n. sp., which exhibits a moderately large size associated with a relatively short rostrum; Rhamphosus longispinatus n. sp., characterized by having the longest dorsal-fin spine and rostrum of all the Rhamphosus species; Rhamphosus tubulirostris n. sp., which shows a peculiar slim rostrum, long-based dorsal and anal fins, and a unique squamation characterized by the presence of numerous bucklers. The species from Bolca are also compared to the only other known Rhamphosus species, Rhamphosus rosenkrantzi, from the Fur Formation, Denmark. A number of morphological features support the assignment of the Rhamphosidae to the syngnathiform clade Dactylopteroidei, together with the extant families Dactylopteridae and Pegasidae, representing the sister group to the Pegasidae.