Criscione and Grimaldi—Earliest Belostomatid fossils from Triassic North Carolina


Table 1. List of fossil belostomatid species. Species


Age


Odrowazicoris polonicus Tarsabedus menkei


Lethonectes naucoroides


Aenictobelostoma primitivum Mesobelostomum deperditum Mesonepa primordialis Stygeonepa foersteri


Nettelstedtia breitkreutzi


Araripebelostomum martinsnetoi Neponymphes godoii Lethocerus vetus


Lethopterus multinervosus Sinobelostoma liui Iberonepa romerali Unnamed Specimen Unnamed Specimen Unnamed Specimen


Lethocerus sulcifemoralis Unnamed Specimen


Propoissonia beskonakensis Lethocerus turgaicus Manocerus stagnans


Early Jurassic Early Jurassic Early Jurassic Middle Jurassic Late Jurassic Late Jurassic Late Jurassic Late Jurassic


Early Cretaceous Early Cretaceous Early Cretaceous Early Cretaceous Early Cretaceous Early Cretaceous Paleocene


early Eocene early Eocene Oligocene


late Oligocene


Diplonychus microcephalum= Sphaerodema microcephalum Lethocerus americanus Belostoma bakeri


middle Miocene middle Miocene


1167


Formation Zagaje


Solnhofen Solnhofen Solnhofen Bed 103


Crato Crato Crato


Dzun-Bain


Huachi-Huanhe La Huérguina Paskapoo Fur Fur


Oligocene–Miocene — Miocene


Late Pleistocene Late Pleistocene


Abrosimovka Shanwang Shanwang


“Lower Lias” “Lower Lias” Todilto


Locality Poland


Dorset, England Dorset, England


New Mexico, USA Germany Germany Germany


Schwarze Quarry, Nettelstedt, Germany


Brazil Brazil Brazil


Gansu Province, China Las Hoyas, Spain Alberta, Canada Denmark Denmark


Mongolia


Bechlejovicer Diatomite Czech Republic Enspel


Germany Turkey


W. Siberia, Russia


Rancho La Brea Asphalt California, USA McKittrick Asphalt


California, USA


group to Belostomatidae. In addition, all species of Lethocerinae are emergent-brooders, meaning that their eggs are deposited on emergent vegetation and attended to by the males. In contrast, the more derived members of Belostomatinae are back-brooders, in which eggs are deposited on the backs of their male mates. Horvathiniinae is placed between Lethocerinae and Belostomati- nae because of itsmore intermediatemorphological characteristics. Brooding behavior has not been observed in this group and its phylogenetic position is less certain (Lauck andMenke, 1961). The relationships within subfamily Belostomatinae are


mostly resolved based on morphology, with the exception of the genus Limnogeton. Its position has been questioned due to its lack of raptorial forelegs and natatorial mid and hind legs. Lauck and Menke (1961) noted the possibility that these characters may indicate a more basal position within Belostomatidae. However, it has been placed within subfamily Belostomatinae because it exhibits back-brooding behavior (Voelker, 1968). This position was supported in a recent phylogenetic analysis of Nepomorpha (Brożek, 2014). This paper describes the earliest known species of family


Belostomatidae, Triassonepa solensis n. gen. n. sp., from the Late Triassic Cow Branch Formation of southern Virginia and northern North Carolina. Based on both the species assemblage within the formation and the known habitats of modern belos- tomatids, probable chemical and environmental conditions of the deposit are discussed.


Materials and methods


Locality and material.—The fossils described in this study are from the Cow Branch Formation of southern Virginia and northern North Carolina, a Late Triassic (Carnian/early Norian, 230–220 Ma) deposit outcropping in the former Solite Quarry where >30 Van Houten cycles are preserved. Three cycles have yielded insect fossils, but one cycle has produced the majority of


Popov, Dolling, and Whalley, 1994 Popov, Dolling, and Whalley, 1994 Polhemus, 2000


Author(s); Reference Popov, 1996


Germar, 1839; Lakshminarayana, 1984 Germar, 1839; Lakshminarayana, 1984 Popov, 1971


Popov, Rust, and Brauckmann, 2000 Nel and Paicheler, 1992 Nel and Waller, 2006 Chou and Hong, 1989


Zamboni, 2001 Popov, 1989


Shandong Province, China Zhang, 1989


Nel and Paicheler, 1992 Popov, 1971


Rust and Ansorge, 1996 Říha and Kukalová, 1967 Wedmann, 2000


Martínez-Delclòs, Nel, and Popov, 1995 Mitchell and Wighton, 1979 Larsson, 1975


Shandong Province, China Zhang and Zhang, 1994 Leidy, 1847; Miller, 1983


Montandon, 1913; Miller, 1983


the insects in this formation (Olsen et al., 1978; Fraser et al., 1996). The Cow Branch Formation is significant because it preserves the oldest fauna of freshwater insects, which are preserved as thin, two-dimensional, silvery films in a matrix of very fine-grained, black shale. Preservation is often excellent, with many specimens fully articulated and microscopic details visible. In addition to 11 orders of insects (Fraser and Grimaldi, 2003; Grimaldi et al., 2005), the Solite Quarry has produced numerous amphibious reptiles (Tanytrachelos ahynis Olsen, 1979), a gliding reptile (Mecistotrachelos apeoros Fraser et al., 2007), fish fossils, dinosaur footprints, and many plant species (Olsen et al., 1978).


Methods.—Specimens were viewed using a Nikon SMZ1500 microscope, fitted with a fiber optic ring light. This non-direc- tional, diffuse light source was necessary to illuminate the silvery, carbonized film by which the insects are preserved. In order to examine the minute details, specimens were wetted with 70% ethanol to increase contrast between the fossil and the black shale matrix. Photographs were taken using two separate setups. Specimens viewed at the American Museum of Natural History (AMNH) were photographed with a Nikon 16MP camera and Nikon Elements NIS software on a Nikon SMZ1500 stereomicroscope. Specimens housed at the Virginia Museumof Natural History (VMNH) were photographed with a Canon 6D DSLR camera using Canon Utility 2 software. All specimen measurements were taken using ImageJ software. Total lengths of adults were measured from the anterior margin of head to the distal margin of the 7th abdominal segment, in order to exclude the 8th abdominal segment, which protrudes to varying degrees in different specimens.


Repositories and institutional abbreviations.—Most of the material examined in this study is housed at the Virginia Museum of Natural History (VMNH), Martinsville, Virginia,


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