Introduction to Lepidoptera

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Introduction to Lepidoptera Taxonomic Workshop for Early Detection of Important Tortricidae and Other Lepidopteran Agricultural and Silvicultural Pests UMass Amherst 15-17 July 2013 Todd M. Gilligan, Ph.D. Colorado State University Department of Bioagricultural Sciences and Pest Management 1177 Campus Delivery Fort Collins, Colorado 80523 USA tgilliga@gmail.com / tgilliga@rams.colostate.edu

Introduction to Lepidoptera Short review Overview of Lepidoptera Placement in Arthropoda Recent morphological and molecular phylogenies Characters to define Lepidoptera Major pest families

First Some review Domain Kingdom Phylum Class Order Family Genus species Specific epithet (specific name) Treated as Latin (italicized) Species = binomial (Genus species Author) Allocapnia sequatchie Kondratieff & Kirchner Allocapnia vivipara (Claassen)

Review terminology Synapomorphy = shared derived character state Character = particular attribute of an organism (usually physical) Monophyly = a group of species descended from a single ancestor and which includes all species descended from that ancestor Systematics / Phylogenetics = reconstructing the evolutionary history of organisms Molecular phylogenetics = reconstructing the evolutionary history of organisms using DNA data

Order Lepidoptera 157,424 described species (as of 2011) Largest lineage of plant-feeding organisms Sister group to the Trichoptera (caddisflies) Characterized by many (24) synapomorphies Large radiation approx. 100 MYA at the same time as the flowering plants One of the most studied and monographed insect orders Large number of pest species

Phylum Arthropoda Regier et al. 2010

Class Insecta Wheeler et al. 2001

Superorder Amphiesmenoptera Trichoptera + Lepidoptera Heterogametic females WZ female/zz male in Ditrysia Pair of glands on sternite V Dense, long setae on wings Forewing with anal veins forming a double Y Larva with a fused hypopharynx and prelabium with silk glands Fusion of 9th tergite and sternite into a ring in male genitalia (= tegumen and vinculum)

Order Lepidoptera Kristensen 1999

Grimaldi & Engel 2005

Order Lepidoptera Traditional groupings Microlepidoptera Primitive moths up to or including the pyraloids Larvae internal feeders Majority of families, minority of species Macrolepidoptera Advanced moths from pyraloids to noctuoids Larvae external feeders Majority of species, minority of families

Order Lepidoptera Synapomorphies (24 listed in Kristensen et al. 2007): loss of median ocellus intercalary sclerite on the antenna maxillary palp with points of flexion between segments 1 and 2, 3 and 4 terminal segment of labial palp with vom Rath s organ (sensillary depression) protibia with epiphysis and only a single spur wings with covering of dense scales metathoracic spiracle with a single, anterior, external lip first abdominal tergite is extensively desclerotized with lateral lobes articulating with second sternite male valvae are primarily undivided anal cerci are absent

Order Lepidoptera Scales Modified setae (specialized cells arising from a socket) Occur in several orders; wings covered in scales unique to the Lepidoptera

Order Lepidoptera Grimaldi & Engel 2005

Order Lepidoptera Epiphysis Located on protibia Used to clean the antennae Lost in many butterflies (except Papilionidae)

Order Lepidoptera vom Rath s organ Sensory structure located in a depression at the terminal end of the labial palp Also termed labial palp pit Detects atmospheric carbon dixoide

Order Lepidoptera Two recent molecular phylogenies of higher Lepidoptera (Ditrysia) Regier, J. C., et al. 2009. Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary Biology 9: 280. doi:10.1186/1471-2148-9-280. Mutanen, M., N. Wahlberg & L. Kaila. 2010. Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proceedings of the Royal Society B. doi:10.1098/rspb.2010.0392.

Order Lepidoptera Remember the morphological phylogeny by Kristensen (1999)?

Regier et al. 2009

Mutanen et al. 2010

Order Lepidoptera Molecular phylogeny results Ditrysia (higher Lepidoptera) monophyletic Most ditrysian groups not supported Butterflies placed in the microlepidoptera(!) Short branches support hypothesis of rapid radiation during diversification of angiosperms

Pest Lepidoptera Three superfamilies contain most pest species: Noctuoidea Approx. 1,000-1,500 pest species Pyraloidea Approx. 750 pest species Tortricoidea Approx. 680 pest species Numbers from Zhang (1994), Index of economically important Lepidoptera

Noctuoidea Most diverse superfamily More than 40,000 described species Family/subfamily taxonomy has changed several times in the past decade All have a thoracic tympanum Recent molecular work recognizes six families More later in family summaries

Noctuoidea Superfamily Noctuoidea (6 families) Family Oenosandridae (4 genera, 8 species) Australia only Family Notodontidae (704 genera, 3,800 species) Family Erebidae (1,760 genera, 24,569 species) Includes Lymantriinae and Arctiinae Family Euteliidae (29 genera, 520 species) Family Nolidae (186 genera, 1,738 species) Family Noctuidae (1,089 genera, 11,772 species) Includes many (but not all) pest species (Numbers from van Nieukerken et al. 2011)

Pyraloidea Snout moths More than 15,000 described species Treated as microlepidoptera (adults) or macrolepidoptera (larvae) All have an abdominal tympanum Crambidae formally treated as subfamily More later in family summaries

Pyraloidea Superfamily Pyraloidea (2 families) Pyralidae (1,055 genera, 5,921 species) Crambidae (1,020 genera, 9,655 species) (Numbers from van Nieukerken et al. 2011)

Tortricidae Bell moths or tortrix moths Possibly highest percentage of pests(?) More than 10,000 described species Female with flat ovipositor lobes Three subfamilies MUCH more later

Tortricidae Superfamily Tortricoidea (1 family) Tortricidae (1,077 genera, 10,361species) (Numbers from www.tortricid.net)

Selected references Grimaldi, D. & M. S. Engel. 2005. Evolution of the insects. Cambridge University Press. 755 pp. Kristensen, N. P. (ed.) 1999. Handbook of Zoology: Vol. 4. Arthropoda: Insecta. Part 35, Lepidoptera, moths and butterflies. Vol. 1. Evolution, systematics, and biogeography. W. de Gruyter, Berlin. 491 pp. Kristensen, N. P. (ed.) 2003. Handbook of Zoology: Vol. 4. Arthropoda: Insecta. Part 36, Lepidoptera, moths and butterflies. Vol. 2. Morphology, physiology, and development. W. de Gruyter, Berlin. 564 pp. Mutanen, M., N. Wahlberg & L. Kaila. 2010. Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proceedings of the Royal Society B. doi:10.1098/rspb.2010.0392. Regier, J. C., et al. 2009. Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary Biology 9: 280. doi:10.1186/1471-2148-9-280. Scoble, M. J. 1992. The Lepidoptera: form function and diversity. Oxford University Press. 404 pp. van Nieukerken, E., et al. 2011. Order Lepidoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 212 221. Zhang, B.-C. 1994. Index of economically important Lepidoptera. CAB International, Wallingford, UK. 599 pp.