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Home , Allocapnia vivipara, Amphiesmenoptera, Ditrysia, Lepidoptera, Microlepidoptera, Oenosandridae

Introduction to

Taxonomic Workshop for Early Detection of Important 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 Management 1177 Campus Delivery Fort Collins, Colorado 80523 USA [email protected] / [email protected]

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   Specific epithet (specific name)  Treated as (italicized)  “Species” = binomial (Genus species Author) ◦ sequatchie Kondratieff & Kirchner ◦ Allocapnia vivipara (Claassen) Review terminology

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

 157,424 described species (as of 2011)  Largest of plant-feeding organisms  to the Trichoptera ()  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 orders  Large number of pest species Phylum Arthropoda

Regier et al. 2010 Class Insecta

Wheeler et al. 2001 Superorder  Trichoptera + Lepidoptera ◦ Heterogametic females  WZ female/ZZ male in ◦ Pair of glands on sternite V ◦ Dense, long setae on wings ◦ Forewing with anal veins forming a double “Y” ◦ with a fused hypopharynx and prelabium with 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 ◦  “Primitive” up to or including the pyraloids  Larvae internal feeders  Majority of families, minority of species ◦  “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 ◦ 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 (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: ◦  Approx. 1,000-1,500 pest species ◦  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 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 (4 genera, 8 species) ◦ only  Family (704 genera, 3,800 species)  Family (1,760 genera, 24,569 species) ◦ Includes and  Family (29 genera, 520 species)  Family (186 genera, 1,738 species)  Family (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  formally treated as subfamily  More later in family summaries… Pyraloidea Superfamily Pyraloidea (2 families)  (1,055 genera, 5,921 species)  Crambidae (1,020 genera, 9,655 species)

(Numbers from van Nieukerken et al. 2011) Tortricidae

 “Bell moths” or “ moths”  Possibly highest percentage of pests(?)  More than 10,000 described species  Female with flat 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 . 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.) : 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.