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Fossil Spiders (Araneae) from the Eocene Kishenehn Formation of Montana, USA

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Fossil Spiders (Araneae) from the Eocene Kishenehn Formation of Montana, USA Palaeontologia Electronica palaeo-electronica.org Fossil spiders (Araneae) from the Eocene Kishenehn Formation of Montana, USA Matthew R. Downen and Paul A. Selden ABTRACT The Kishenehn Formation contains a diverse assemblage of terrestrial arthropod fossils and represents a tropical ecosystem in North America during the Eocene (46 Ma). Most of the fossils are small insects, but spiders have also been recovered and recently made available for study. The fossil spiders are small and preserved as com- pressions in a thinly laminated oil shale, but abundant setae and other morphological features are preserved in relatively high detail. Here, the fossil spiders are described for the first time and include new species of orbweaving spiders from the family Aranei- dae and a ground-dwelling spider from the family Gnaphosidae. Most of the spiders in the assemblage resemble extant orbweaving spiders like Neoscona: Greenwalta- rachne pamelae gen. & sp. nov. and Consteniusi leonae gen. et sp. nov.. A single gna- phosid, distinguished by widely separated cylindrical spinnerets, is likely a juvenile. A single male spider belonging to Araneomorphae is too poorly preserved to discern at family level. The fossils described here are the oldest fossil spiders recovered from Montana thus far. Matthew R. Downen. Department of Geology, University of Kansas, Ritchie Hall Earth, Energy & Environment Center 1414 Naismith Drive, Room 254 Lawrence, KS 66045, USA. [email protected] Paul A. Selden. Department of Geology, University of Kansas, Ritchie Hall Earth, Energy & Environment Center 1414 Naismith Drive, Room 254 Lawrence, KS 66045, USA and Natural History Museum, Cromwell Road, London SW7 5BD, UK. [email protected] Keywords: Fossil-Lagerstätte; lacustrine; new genus; new species; taxonomy Submission: 24 October 2020. Acceptance: 18 November 2020. http://zoobank.org/5A215F0E-AE94-4250-9D22-E38DE45250E7 Downen, Matthew R. and Selden, Paul A. 2020. Fossil spiders (Araneae) from the Eocene Kishenehn Formation of Montana, USA. Palaeontologia Electronica, 23(3):a56. https://doi.org/10.26879/1135 palaeo-electronica.org/content/2020/3237-kishenehn-fossil-spiders Copyright: December 2020 Paleontological Society. This is an open access article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ DOWNEN & SELDEN: KISHENEHN FOSSIL SPIDERS INTRODUCTION and spiders, currently including 14 species of ara- neids in two genera: Araneus Clerck, 1757, and The Kishenehn Formation of Montana Tethneus Scudder, 1885, although many of these (Eocene: 46 Ma) is one of several Cenozoic lacus- fossil spiders from the Florissant Formation are in trine deposits of North America that are considered need of revision (World Spider Catalog, 2020). Fossil-Lagerstätten. A diverse assemblage of fos- Gnaphosids are ground-dwelling spiders that sils is represented and includes terrestrial arthro- do not weave webs to capture prey; instead, they pods, plants, aquatic molluscs, and mammals actively hunt on the ground (Jocqué and Dippe- (Pierce and Constenius, 2014). Small insects naar-Schoeman, 2007). Gnaphosids and their rela- including flies (Diptera), true bugs (Hemiptera), and tives are distinguished from the aerial web- extremely minute fairy wasps (Hymenoptera: spinning spiders by possessing only two tarsal Mymaridae) are the most abundant fossils, while claws, instead of three, and differing leg lengths. vertebrates are rare (Greenwalt and Labandeira, Gnaphosids themselves are distinguished easily 2013; Greenwalt and Engel, 2014; Greenwalt et from other similar spiders by their widely separated al., 2014; Lapolla and Greenwalt, 2015). The cylindrical spinnerets and claw tufts at the end of exceptional preservation has yielded even a blood- each leg (Platnick, 1990). There are currently 2522 engorged mosquito with preserved biomolecules species within Gnaphosidae, which also makes derived from hemoglobin (Greenwalt et al., 2013). them one of the most diverse spider families. The Relatively few fossil spiders have been recovered fossil record of Gnaphosidae comes mostly from this deposit and, hitherto, none have been Baltic amber with 13 species (Menge, 1854; Wun- described formally. This paper is a taxonomic sur- derlich, 2011). Five species are from the Florissant vey of the fossil spider assemblage from the Kish- Formation (34 Ma), but many of these need revi- enehn Formation. sion (Petrunkevitch, 1922). The assemblage consists of new species of orbweaving spiders (Araneidae), a ground dwelling GEOLOGICAL SETTING AND FOSSIL spider from the family Gnaphosidae, and an inde- PRESERVATION terminate araneomorph. Most of the spiders in the assemblage are conspecific and similar to extant The Kishenehn Formation is located in north- spiders in the araneid spider genus Neoscona western Montana and is composed of two mem- Simon, 1864. A single very small spider is likely a bers: the Coal Creek Member and the overlying juvenile gnaphosid, based on cylindrical spinner- Pinchot Conglomerate Member. The Coal Creek ets. A single male spider is too poorly preserved to Member is represented by an 1150 m thick succes- discern at family level and is left as an indetermi- sion of sandstone, siltstone, and oil shale (Conste- nate araneomorph. Whereas none of the spiders nius et al., 1989). Fossil insects and spiders are described here extend the age range of any fami- found in the middle of the Coal Creek Member in lies, they do contribute to comparing spider assem- the oil shale. The fossils are preserved as com- blages preserved in Eocene lacustrine deposits pressions in extremely thin and delicate lamina- across North America during the Eocene. tions representing varves. A thin layer of surface The fossil record of Araneidae extends back silicates obscures many of the fossils (Greenwalt to the Cretaceous with the earliest araneid, Meso- et al., 2014). zygiella dunlopi Ortuño, 2006, preserved in amber The depositional setting and paleoenviron- from Álava, Spain (Penney and Ortuño, 2006). Ara- ment is interpreted as a freshwater lacustrine sys- neids are one of the most diverse groups of spiders tem in a subtropical to tropical environment. today (3100 species; World Spider Catalog, 2020), Fishes, including bowfins (Amiidae) and suckers with many fossils described from amber (91 spe- (Catostomidae), are indicative of freshwater habi- cies). The Cenozoic record of Araneidae in North tats (Wilson, 1988). Sycamores are the most abun- America currently includes araneids from the dant plant fossils and represent humid and warm Dominican and Chiapas ambers and one lacustrine conditions, whereas other plants like cattails and deposit, the Florissant Formation of Colorado waterferns are suggestive of paludal environments (Scudder, 1890; Petrunkevitch, 1922, 1971; Wun- (Constenius et al., 1989). Relatively few mammal derlich, 1982, 1986, 1988). The Florissant Forma- fossils have been recovered but include flying tion (Eocene: 34 Ma) is a well-known lacustrine lemurs, which also support a tropical paleoenviron- deposit with many exceptionally preserved insects ment (McKenna, 1990). 2 PALAEO-ELECTRONICA.ORG MATERIALS AND METHODS sils to fluoresce, and thus, increasing the contrast between matrix and cuticle/fossil material. Materials Abbreviations: AME anterior median eye(s), car The specimens consist of a part only and are carapace, ch chelicera, cx coxa, fe femur, L length, preserved in oil shale from the Coal Creek Member lb labium, LE lateral eye(s), mt metatarsus, mx of the Kishenehn Shale in northwestern Montana. maxilla, op opisthosoma, pa patella, Pd pedipalp, Greenwalt et al. (2013) lists specific sites of collec- PME posterior median eye(s), sp spinnerets, st tion. Specimens (PAL 583.1, PAL 20412.1, PAL sternum, ta tarsus, ti tibia, W width. 32280, PAL 20480) are deposited in the Depart- ment of Paleobiology, National Museum of Natural SYSTEMATIC PALEONTOLOGY History, Washington, D.C., with specimen labels Order ARANEAE Clerck, 1757 beginning with PAL, and one specimen (CM Suborder OPISTHOTHELAE Pocock, 1892 56099) is deposited in the Carnegie Museum of Infraorder ARANEOMORPHAE Smith, 1902 Natural History, Pittsburg, PA. Araneomorphae incertae sedis Methods (Figure 1) Due to the coating of silicates on the surface Remarks. Parts of this fossil spider are preserved of the fossils, specimens were submerged in 95% with high fidelity (the legs), whereas other portions ethanol. The specimens were studied using a are poor in detail (the palps and carapace). The Leica M205C stereomicroscope, photographed palps, carapace, and some of the leg joints have with a Canon EOS 5D MkII digital camera attached been replaced with pyrite which contributes to a to the microscope and captured with DSLR Assis- loss of detail. Mygalomorph spiders typically have tant software (www.kaasoft.com) on an Apple Mac- large porrect chelicerae and robust legs. The fossil Book Pro computer. Drawings were made using a here has relatively small chelicerae that are not drawing tube attached to the microscope. Photo- porrect, and legs that are relatively slender with graphs were manipulated using Adobe Photoshop heavy spination. The pedipalps are modified, indi- software. Fossils were also imaged using an Olym- cating
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