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Milbert's Tortoiseshell, Aglais Milberti (Lepidoptera: Nymphalidae): a Facultative Trogloxene in Alpine Caves

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Milbert's Tortoiseshell, Aglais Milberti (Lepidoptera: Nymphalidae): a Facultative Trogloxene in Alpine Caves Taylor et al. Milbert's Tortoiseshell, Aglais milberti (Lepidoptera: Nymphalidae): A facultative trogloxene in alpine caves Steven J. Taylor1, Jean K. Krejca2, Michael E. Slay3, and Terry L. Harrison4 1Illinois Natural History Survey, University of Illinois, 1816 S Oak St., Champaign, IL 61820 USA. Email: [email protected] 2Zara Environmental LLC, 118 W. Goforth Rd., Buda, TX 78610 USA. Email: [email protected] 3The Nature Conservancy, 601 North University Avenue, Little Rock, AR 72705 USA. Email: [email protected] 4Department of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801 USA. Email: [email protected] Key Words: Arthropoda, Insecta, Nymphalinae, Mountain View Cave, Broken Cave, Great Basin National Park, Snake Range, White Pine County, Nevada, USA, trogloxene Use of caves as a habitat by moths has been reported for the herald moth, Scoliopteryx libatrix (Linnaeus) (Noctuidae)1, grannyʼs cloak moth, Speiredonia spp. (Noctuidae)2,3, the hop vine moth, Hypena humuli Harris (Noctuidae)4, guano moths (Tineidae)5,6, and Geometridae (e.g., Triphosa haesitata [Guenee])1,7–9. However, most other Lepidoptera – the butterflies, moths and skippers – are almost never thought of as trogloxenes or troglophiles. Milbertʼs Tortoiseshell, Aglais milberti (Godart) (Lepidoptera: Nymphalidae: Nymphalinae) is a boreal North American brush-footed butterfly, most closely related to the Palearctic butterfly Aglais urticae (Linnaeus)10. It occurs from Alaska to Newfoundland, extending to the south – in the western United States west of the Great Plains – to southern California and New Mexico. In the eastern United States it is less frequently recorded further south than Wisconsin, Minnesota, Michigan, Ohio and west to Pennsylvania and New York11. This butterfly is reported as being a riparian species12,13, and occurs in meadows, roadsides, and clearings14,15. Adults are strong fliers12, and migrate to alpine habitats to forage in the summer16, where they can be common17–19. Nectar sources include phlox and bee plant20, and, in the eastern United States, Burdock, Joe-Pye Weed, and Shrubby Cinquefoil21. Adults return to lower altitudes to overwinter, with eggs being laid at lower altitudes in the spring16. Larvae feed on herbaceous vegetation, especially Urtica sp.14,15,22,23. We have recently completed a bioinventory of caves of Great Basin National Park (White Pine County, Nevada) in which we observed and collected butterflies in two caves located above timberline. In Mountain View Cave (length 16.2 meters, elevation 3413 meters), two specimens of A. milberti were collected on 18 July 2007 from dry bedrock ceiling in the twilight zone, where the 2009 Speleobiology Notes 1: 20-23 20 Taylor et al. temperature was between 2.07 and 15.57 ○C, the relative humidity was between 34.6 and 80.9%, and light was less than 4 lux (the collection was between two stations where the parameters were measured, thus a range is given). Surface conditions were 20.39 ○C, 25.9% relative humidity, and light 94600 lux. In Broken Cave (length 32.9 meters, elevation 3407 meters), one specimen of Milbertʼs Tortoiseshell was collected on 10 July on a dry rock wall and another on 16 July 2007 on a dry bedrock ceiling (Figure 1), with others observed on 16 July 2007. Here the collections were in the twilight zone where on 16 July 2007, air temperature was 8.27 ○C, relative humidity was 48.2%, and light level was 1 lux. Surface conditions on 16 July 2007 were air temperature 18.28 ○C, relative humidity 38.0%, and light 9860 lux. All specimens at both sites were observed to be roosting with wings folded over their dorsum as in Figure 1. Figure 1. Milbertʼs Tortoiseshell, Aglais milberti (Nymphalidae) roosting on the ceiling of Broken Cave (16 July 2007). Our study of caves in Nevadaʼs Snake range included multiple visits to 26 caves ranging in elevation from 1724 to 3413 meters, yet we observed A. milberti in only two of these caves, both of which were above timberline. These 2009 Speleobiology Notes 1: 20-23 21 Taylor et al. butterflies were resting on dry bedrock ceilings and walls in the twilight or dark zones of the caves, functioning as facultative trogloxenes. Careful examination of western North American caves above timberline during the period when A. milberti has migrated to the alpine zone to forage should yield additional observations of facultative cave use by this species. Acknowledgements: We thank Gretchen M. Baker, Ben M. Roberts, and Meg A. Horner, all of the National Park Service, for guidance and assistance in all stages of our research in Great Basin National Park. Field assistance at Broken and Mountain View caves was also provided by Christy A. M. Slay, Patrick M O'Brien, and Shawn C. Thomas. Literature Cited: 1. McKillop, W. B. Scoliopteryx libatrix (Noctuidae) and Triphosa haesitata (Geometridae) in caves in Manitoba, Canada. Journal of the Lepidopterists Society. 47,106–113 (1993). 2. Pavey, C. R. & Burwell, C. J. Cohabitation and predation by insectivorous bats on eared moths in subterranean roosts. Journal of Zoology, London 265, 141–146 (2005). 3. Weinstein, P. & Slaney, D. Invertebrate faunal survey of Rope Ladder Cave, Northern Queensland: a comparative study of sampling methods. Journal of the Australian Entomological Society 34, 233–236 (1995). 4. Mingari, J. & Rudy, C. Hop vine moths, Hypena humuli, in a Wisconsin cave. Occasional Papers of the Moth Photographersʼ Group, No.1. http://mothphotographersgroup.msstate.edu/Files/Live/CR/CaveMoths.sht ml (accessed 15 February 2009) (2006). 5. Bradley, J. D. Tineid moths from the Bat Caves, Selangor. Journal of Natural History 7, 675–682 (1973). 6. Robinson, G. S. Cave dwelling tineid moths: a taxonomic review of the world species (Lepidoptera: Tineidae). Transactions of the British Cave Research Association 7, 83–120 (1980). 7. Graham, R. E. The moth Traephosa haesitata of California and Nevada Caves. Cave Notes 4, 42 (1962). 8. Graham, R. E. Spatial biometrics of subterranean demes of Triphosa haesitata (Lepidoptera: Geometridae). Caves and Karst 10, 21–29 (1968). 9. Graham, R. E. The twilight moth, Triphosa haesitata (Lepidoptera: Geometridae) from California and Nevada caves. Caves and Karst 10, 41– 47 (1968). 10. Nylin, S., Nyblom, K., Ronquist, F., Janz, N., Belicek, J. & Källersjö, M. Phylogeny of Polygonia, Nymphalis and related butterflies (Lepidoptera: Nymphalidae): a total evidence analysis. Zoological Journal of the Linnaean Society 132, 441–468 (2001). 11. Opler, P. A., Pavulaan, H., Stanford, R. E. & Pogue, M., coordinators. Butterflies and Moths of North America. Bozeman, Montana: Big Sky 2009 Speleobiology Notes 1: 20-23 22 Taylor et al. Institute. http://www.butterfliesandmoths.org/ (accessed 15 February 2009) (2006). 12. Fleishman, E. & Murphy, D. D. Patterns and processes of nestedness in a Great Basin butterfly community. Oecologia 119, 133–139 (1999). 13. Fleishman, E., Murphy, D. D. & Brussard, P. F. A new method for selection of umbrella species for conservation planning. Ecological Applications 10, 569–579 (2000). 14. Emmel, T. C. The ecology and distribution of butterflies in a montane community near Florissant, Colorado. American Midland Naturalist 72, 358–373 (1964). 15. LaFontaine, J. D. & D. M. Wood. Butterflies and Moths (Lepidoptera) of the Yukon. pp. 723-785 in H.V. Danks and J.A. Downes (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp. (1997). 16. Scott, J. A. & Epstein, M. E. Factors affecting phenology in a temperate insect community. American Midland Naturalist 117, 103–118 (1987). 17. Goehring, D. M., Daily, G. C., Dasgupta, S. & Ehrlich, P. R. Range occupancy and endangerment: a test with a butterfly community. American Midland Naturalist 157,106–120 (2007). 18. Klots, A. B. New butterfly subspecies from Wyoming (Nymphalidae, Pieridae). American Museum Novitiate 1054, 1–6 (1940). 19. Spalding, J. B. The aeolian ecology of White Mountain Peak, California: windblown insect fauna. Arctic Alpine Research 11, 83–94 (1979). 20. Johnson, K. & Nixon, E. S. The Rhopalocera of northwestern Nebraska. American Midland Naturalist 78, 508–528 (1967). 21. Voss, E. G. The Butterflies of Emmet and Cheboygan counties, Michigan with other notes on northern Michigan butterflies. American Midland Naturalist 51, 87–104 (1954). 22. Grimble, D. G. & Beckwith, R. C. A survey of the Lepidoptera fauna from the Blue Mountains of eastern Oregon. Journal of Research on Lepidoptera 21, 83-102 (1992). 23. Slansky, F., Jr. Relationship of larval food-plants and voltinism patterns in temperate butterflies. Psyche 81, 243–253 (1974). 2009 Speleobiology Notes 1: 20-23 23 .
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