Journal of Paleontology, 91(6), 2017, p. 1166–1177 Copyright © 2017, The Paleontological Society 0022-3360/17/0088-0906 doi: 10.1017/jpa.2017.48
The oldest predaceous water bugs (Insecta, Heteroptera, Belostomatidae), with implications for paleolimnology of the Triassic Cow Branch Formation
Julia Criscione1 and David Grimaldi2
1Department of Earth and Planetary Sciences, Rutgers University, 610 Taylor Road, Piscataway, NJ, 08854, USA ⟨
juliecriscione@gmail.com⟩ 2Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th St, New York, NY, 10024, USA ⟨
grimaldi@amnh.org⟩
Abstract.—A new genus and species of predaceous water bugs, Triassonepa solensis n. gen. n. sp., is described from the Triassic Cow Branch Formation of Virginia and North Carolina (USA) based on ~36 adult specimens and 51 nymphs. This species is the oldest known member of the extant family Belostomatidae. It is placed in a new genus based on the unique structure of the raptorial foreleg, in which the tarsus is elongate and opposed to the tibia + femur. The fossil record of this family is reviewed and the paleoenvironmental implications of the species assemblage preserved in the Cow Branch Formation are discussed.
Introduction
Jurassic, when the family first diversified. The oldest of these is Odrowazicoris polonicus Popov, 1996, an isolated wing in Hettangian-age beds of the Holy Cross Mountains of Poland. A number of other Jurassic species have been described from Europe and the United States, including: Tarsabedus menkei Popov, Dolling, and Whalley, 1994; Aenictobelostoma primitivum Polhemus, 2000; Stygeonepa foersteri Popov, 1971; and Nettelstedtia breitkreutzi Popov, Rust, and Brauckmann, 2000. The Early Cretaceous also harbored diverse species and genera, including three taxa from the Crato Formation Platten- kalk of Brazil, and a species possessing unique paddle-shaped metathoracic legs, Iberonepa romerali Martínez-Delclòs, Nel, and Popov, 1995, from Las Hoyas, Spain.
The Heteroptera, or sucking bugs, have a long fossil record, potentially spanning back to the Permian. The first putative heteropteran from this period is Paraknightia magnifica Evans, 1943 from New South Wales (Evans, 1950). However, the first definitive heteropteran, Arlecoris louisi Shcherbakov, 2010, was recently described from the earliest Middle Triassic (early Anisian) of the northern Vosges Mountains of France. This species is also the earliest member of the infraorder Nepomorpha Popov, 1968, a group containing the majority of truly aquatic heteropterans (Belostomatidae Leach, 1815;Nepidae Latreille, 1802;Naucoridae Leach, 1815; Corixidae Leach, 1815; and Notonectidae Leach, 1815). The Nepomorpha have the best fossil record of all Hetero- ptera (Grimaldi andEngel, 2005), no doubt because of their aquatic habits, but the fossil record of belostomatids is not yetwell studied. Modern Belostomatidae are medium- (9mm) to very large-sized (110mm adult body length) swimming bugs that are efficient predators, grabbing prey with raptorial forelegs, injecting potent salivary secretions, and siphoning out the liquefied internal tissues of their prey with a sharp beak. The earliest described belostomatid fossils are from the
Though belostomatids are widespread in Cenozoic sedi-
ments, many are still undescribed, including a Paleocene specimen from Alberta, Canada (Mitchell and Wighton, 1979), two unnamed early Eocene specimens from Denmark (Larsson, 1975; Rust and Ansorge, 1996), and a late Oligocene specimen from Germany (Wedmann, 2000). The extant genus Lethocerus Mayr, 1853 became fairly diverse during the Miocene, with two species (L. sulcifemoralis Říha and Kukalová, 1967, and L. turgaicus Popov, 1971) found in the Oligocene–Miocene of the Czech Republic and the Miocene of Russia, respectively. Additionally, two modern species, Lethocerus americanus Leidy, 1847 and Belostoma bakeri Montandon, 1913, are reported to occur in Late Pleistocene asphalt deposits of California (Miller, 1983). See Table 1 for a complete list of fossil belostomatid species. Today, belostomatids have a worldwide distribution,
although the majority of species are found in the tropics. They are represented by three subfamilies consisting of nine genera and ~146 species (Schuh and Slater, 1995). The subfamily Belostomatinae Lauck and Menke, 1961, is by far the largest group within Belostomatidae and contains six genera: Abedus Stål, 1862; Appasus Amyot and Serville, 1843 (Polhemus, 1995); Belostoma Latreille, 1807; Diplonychus Laporte, 1833; Hydrocyrius Spinola, 1850; and Limnogeton Mayr, 1853. The subfamily Horvathiniinae Lauck and Menke, 1961, is mono- generic, consisting of nine species in the genus Horvathinia Montadon, 1911. The third subfamily, Lethocerinae Lauck and Menke, 1961, consists of three genera: Lethocerus Mayr, 1853; Benacus Stål, 1861 (Goodwyn, 2006); and Kirkaldyia Montandon, 1909 (Goodwyn, 2006). The current belostomatid phylogeny is based on morphology
and reproductive behaviors: (Lethocerinae, (Horvathiniinae (Belostomatinae))) (Lauck and Menke, 1961; Mahner, 1993; Smith, 1997). Lethocerinae is the most basal taxon because it retains many characters of Nepidae (‘water scorpions’), the sister
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