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<I>Anthocharis Sara</I> University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Center for Systematic Entomology, Gainesville, Insecta Mundi Florida 2018 A review of three species-level taxa of the Anthocharis sara complex (Lepidoptera: Pieridae: Pierinae: Anthocharidini) Todd L. Stout Florida Museum of Natural History, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/insectamundi Part of the Ecology and Evolutionary Biology Commons, and the Entomology Commons Stout, Todd L., "A review of three species-level taxa of the Anthocharis sara complex (Lepidoptera: Pieridae: Pierinae: Anthocharidini)" (2018). Insecta Mundi. 1133. https://digitalcommons.unl.edu/insectamundi/1133 This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. March 30 2018 INSECTA 0615 1–38 urn:lsid:zoobank.org:pub:322DABD4-EDB4-43D1-9140- A Journal of World Insect Systematics 4238E3C6C22F MUNDI 0615 A review of three species-level taxa of the Anthocharis sara complex (Lepidoptera: Pieridae: Pierinae: Anthocharidini) Todd L. Stout The McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History Gainesville, FL 32611-2710 Date of issue: March 30, 2018 CENTER FOR SYSTEMATIC ENTOMOLOGY, INC., Gainesville, FL Todd L. Stout A review of three species-level taxa of the Anthocharis sara complex (Lepidoptera: Pieridae: Pierinae: Anthocharidini) Insecta Mundi 0615: 1–38 ZooBank Registered: urn:lsid:zoobank.org:pub:322DABD4-EDB4-43D1-9140-4238E3C6C22F Published in 2018 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any non-marine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. Insecta Mundi will not consider works in the applied sciences (i.e. medical entomology, pest control research, etc.), and no longer publishes book reviews or editorials. Insecta Mundi publishes original research or discoveries in an inexpensive and timely manner, distributing them free via open access on the internet on the date of publication. Insecta Mundi is referenced or abstracted by several sources, including the Zoological Record and CAB Abstracts. Insecta Mundi is published irregularly throughout the year, with completed manuscripts assigned an individual number. Manuscripts must be peer reviewed prior to submission, after which they are reviewed by the editorial board to ensure quality. One author of each submitted manuscript must be a current member of the Center for Systematic Entomology. Guidelines and requirements for the preparation of manuscripts are available on the Insecta Mundi website at http://centerforsystematicentomology.org/insectamundi/ Chief Editor: David Plotkin, [email protected] Assistant Editor: Paul E. Skelley, [email protected] Head Layout Editor: Robert G. Forsyth Editorial Board: J. H. Frank, M. J. Paulsen, Michael C. Thomas Review Editors: Listed on the Insecta Mundi webpage Printed copies (ISSN 0749-6737) annually deposited in libraries CSIRO, Canberra, ACT, Australia Museu de Zoologia, São Paulo, Brazil Agriculture and Agrifood Canada, Ottawa, ON, Canada The Natural History Museum, London, UK Muzeum i Instytut Zoologii PAN, Warsaw, Poland National Taiwan University, Taipei, Taiwan California Academy of Sciences, San Francisco, CA, USA Florida Department of Agriculture and Consumer Services, Gainesville, FL, USA Field Museum of Natural History, Chicago, IL, USA National Museum of Natural History, Smithsonian Institution, Washington, DC, USA Zoological Institute of Russian Academy of Sciences, Saint-Petersburg, Russia Electronic copies (Online ISSN 1942-1354, CDROM ISSN 1942-1362) in PDF format Printed CD or DVD mailed to all members at end of year. Archived digitally by Portico. Florida Virtual Campus: http://purl.fcla.edu/fcla/insectamundi University of Nebraska-Lincoln, Digital Commons: http://digitalcommons.unl.edu/insectamundi/ Goethe-Universität, Frankfurt am Main: http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:hebis:30:3-135240 Copyright held by the author(s). This is an open access article distributed under the terms of the Creative Commons, Attribution Non-Commercial License, which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited. http://creativecommons.org/licenses/by-nc/3.0/ Layout Editor for this article: Robert G. Forsyth 0615: 1–38 2018 A review of three species-level taxa of the Anthocharis sara complex (Lepidoptera: Pieridae: Pierinae: Anthocharidini) Todd L. Stout The McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History Gainesville, FL 32611-2710 [email protected] Abstract. A combination of five characters supports the arrangement of three species in the Anthocharis sara complex (Lepidoptera: Pieridae: Pierinae: Anthocharidini) consisting of Anthocharis sara Lucas, 1852 (Sara oran- getip), Anthocharis thoosa Scudder, 1878 (southwestern orangetip), and Anthocharis julia W.H. Edwards, 1872 (Julia orangetip). These five characters include adult phenotypes, fifth instar larval coloration, pupal cone curva- ture and color, number of overwintering cycles, and interspecific contact zone behavior. This taxonomic treatment is generated from extensive life history studies from 12 western U.S. states through most of the sara complex distribution. Key words. Sara orangetip, Anthocharis thoosa, Anthocharis julia, caterpillar, pupae. Introduction Over the last half century, there have been several interpretations regarding the number of species in the Anthocharis sara complex ranging from one to four. These interpretations have primarily depended on adult wing characters. This paper will review five separate characters of adult phenotypes, fifth instar larval coloration, pupal cone curvature and color, number of overwintering cycles, and interspecific contact zone behavior to support the arrangement of three species-level taxa- Anthocharis sara Lucas, 1852, Anthocharis thoosa Scudder, 1878, and Anthocharis julia W. H. Edwards, 1872. Historical review. The initial impetus for the reexamination of the sara complex was provided by Geiger and Shapiro (1986) wherein the taxa Anthocharis sara and Anthocharis stella W. H. Edwards, 1879 (Stella orangetip), were shown to represent distinct species and it was suggested that A. julia was also a distinct species. Subsequently, Opler (1999) recognized four species within the sara complex (A. stella, A. sara, A. thoosa, and A. julia). Opler’s treatment is an expansion on Geiger and Shapiro (1986). The Pelham catalogue (2008, 2017) currently places all named subspecies of the complex under A. sara pending further research which is the main purpose of this paper. Materials and Methods Extensive rearing studies of 96 populations of all subspecies of the sara complex were performed between 1995 and 2016 to photograph fifth instar caterpillars, pupae, and adults. Photographic equip- ment included a Canon Rebel XTi and Canon EOS 60D with a 100-mm fixed macro lens. Eggs were mostly obtained from field-collected gravid females. Eggs and larvae were also located in the field on mustards (Brassicaceae) including Descurainia pinnata, Walter, 1894, Guillenia lasiophylla Rydberg, 1923, Isatis tinctoria Linnaeus, 1753, Streptantella longirostris Watson, 1906, Cardamine californica Nuttall, 1891, Brassica nigra Linnaeus, 1833, Arabidopsis thaliana Linnaeus, 1842, Boechera hirsuta Linnaeus, 1772, Boechera holboellii Hornem, 1827, Boechera sparsifloraNuttall, 1838, Boechera peren- nans Watson, 1887, Boechera retrofracta Graham, 1827, Boechera microphylla Nuttall, 1838, Turritis glabra Linnaeus, 1753, Boechera stricta Graham, 1829, and other unidentified species of Boechera (formerly Arabis). Rearing methods. Species identification of pierid ova was facilitated in the field by three factors: Oviposition location, color of the freshly laid ova, and numbers of ova laid in close proximity. These 2 • INSECTA MUNDI 0615, March 2018 STOUT factors were helpful because Pontia sisymbrii Boisduval, 1852, Pontia protodice Boisduval and Le Conte, [1836], Euchloe ausonides Lucas, 1852, Euchloe olympia W. H. Edwards, 1852, Euchloe hyantis W. H. Edwards, 1871, Anthocharis cethura Felder and Felder, 1865, and Anthocharis lanceolata Lucas, 1852, share the same host plants with the sara complex at varying locations. Females of A. sara, A. thoosa, and A. julia frequently lay eggs singly on the pedicels of rock cresses whereas females of E. hyantis, E. olympia, and E. ausonides usually oviposit on unopened flower buds. Contrastingly, females of P. sisymbrii prefer to oviposit one or multiple eggs on the lower leaves and stems. Similar to females of the sara complex, females of P. beckerii W. H. Edwards, 1871, P. protodice, and P. occidentalis Reakirt, 1866, also prefer to oviposit one or several eggs on host pedicels whereas sara complex females usually oviposit a single egg on a pedicel. Freshly laid fertile ova of North American Pontia spp., Euchloe spp., and Anthocharis spp. change color from a lighter whitish, yellowish, or greenish
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