Basic adult and larval morphology

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 Pest Management 1177 Campus Delivery Fort Collins, Colorado 80523 USA [email protected] / [email protected]

Lepidoptera morphology

 Adult morphology ◦ Head: compound eyes, ocelli, proboscis, palpi, scaling (vestiture) ◦ Thorax: tegulae, wings (venation, coupling method, pattern), tympanal organs, scaling and tufts, male secondary structures ◦ Abdomen: specialized scaling, tympanal organs, genitalia  Larval morphology ◦ Head: stemmata, labial palpi, mandibles, antennae, chaetotaxy ◦ Thorax: prothoracic shield, thoracic prolegs, spiracles, primary and secondary setae, chaetotaxy ◦ Abdomen: prolegs, spiracles, primary and secondary setae, chaetotaxy Adult morphology Adult morphology

 Diagnostic structures of the three body parts ◦ Head: compound eyes, ocelli, proboscis, palpi, scaling (vestiture) ◦ Thorax: tegulae, wings (venation, coupling method, pattern), tympanal organs, scaling and tufts, male secondary structures ◦ Abdomen: specialized scaling, tympanal organs, genitalia Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture)

Grimaldi & Engel 2005 Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Head

 Proboscis ◦ Basal are mandibulate ◦ = families with a proboscis (and larval spinneret) ◦ Proboscis developed with maxillary galeae becoming elongate, concave, and held together with processes called ligulae ◦ This occurred in conjunction with the evolution of angiosperms to allow feeding on nectar and pollen

Grimaldi & Engel 2005 Grimaldi & Engel 2005 Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Head

 Antennae (antenna)  Compound eye  Ocelli (ocellus)  Proboscis  Labial palpi (palpus)  Maxillary palpi  Scaling (vestiture) Thorax

 Tegulae and scaling (crests)  Wings  Legs

Scoble 1992 Thorax

 Tegulae and scaling tegula (crests)  Wings  Legs

posterior crest Thorax

 Tegulae and scaling (crests)  Wings ◦ Venation  Legs

Grimaldi & Engel 2005 Thorax Jugal coupling

 Wing coupling ◦ Jugal coupling (basal families)  Jugum (FW) ◦ Amplexiform coupling (most and )

Amplexiform coupling Thorax

 Wing coupling male female ◦ Frenulo-retinacular coupling  Frenulum (HW) and retinaculum (FW) retinaculum ◦ Usually male = one bristle, female = two or more bristles (this varies among families)

frenulum Grimaldi & Engel 2005 Thorax

 Tegulae and scaling (crests)  Wings  Legs

Scoble 1992 Tympanal organs

 On thorax or abdomen, depending on family  Metathoracic ◦ and other families  Abdominal ◦ and Geometroidea

Tympanal organs

 Evolved 8 separate times? (or only a few times – see placement of Pyraloidea in Regier et al. 2009 phylogeny)  , Hedylidae, , few , , , Geometroidea, Pyraloidea, Noctuoidea  Appears to have evolved around the same times as diversification of bats (approx. 60-70 MYA) Abdomen

 Female genitalia  Male genitalia

 We will discuss genital terminology during the tortricid lecture and for specific families  It is important to note that some terms are not homologous across families Abdomen

 Female genitalia ◦  Common opening for copulation and oviposition ◦ “”  Separate copulatory opening; sperm migrates through an external fold to the ovipore ◦  Separate copulation and oviposition openings, connected by ductus seminalis  98% of all Lepidoptera

Grimaldi & Engel 2005 Grimaldi & Engel 2005 Abdomen

 Female genitalia  Male genitalia Abdomen

 Female genitalia  Male genitalia Sex scaling

 Wings, legs, abdomen, thorax  Usually found in males Larval morphology

photo by Lukjonis on Flickr Larval morphology

 Diagnostic structures of the three body parts ◦ Head: stemmata, labial palpi, mandibles, antennae, chaetotaxy ◦ Thorax: prothoracic shield, thoracic prolegs, spiracles, primary and secondary setae, chaetotaxy ◦ Abdomen: prolegs, spiracles, primary and secondary setae, chaetotaxy Larval morphology Head

 Epicranial notch  Stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

Stehr 1987 Head

 Epicranial notch  Stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

Stehr 1987 Head

 Epicranial notch  Stemmata ◦ Sometimes incorrectly termed “ocelli” ◦ Typically 6 stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

Stehr 1987 Head

 Epicranial notch  Stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

Stehr 1987

D. W. McCoy, USDA/APHIS Head

 Epicranial notch  Stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

D. W. McCoy, USDA/APHIS Head

 Epicranial notch  Stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

Stehr 1987

D. W. McCoy, USDA/APHIS Head

 Epicranial notch  Stemmata  Antennae  Mandibles  Spinneret  Chaetotaxy

Stehr 1987 Thorax

 Prothoracic shield  Thoracic legs  Spiracle  Primary and secondary setae  Chaetotaxy

Stehr 1987 Thorax

 Prothoracic shield  Thoracic legs  Spiracle  Primary and secondary setae  Chaetotaxy Thorax

 Prothoracic shield  Thoracic legs  Spiracle  Primary and secondary setae  Chaetotaxy Thorax

 Prothoracic shield  Thoracic legs  Spiracle ◦ On the prothorax (T1)  Primary and secondary setae  Chaetotaxy Thorax

 Prothoracic shield  Thoracic legs  Spiracle  Primary and secondary setae  Chaetotaxy Thorax

 Prothoracic shield  Thoracic legs  Spiracle Names and positions of primary setae:

 Primary and D = dorsal SD = subdorsal secondary setae L = lateral SV = subventral  Chaetotaxy V = ventral Thorax

 Prothoracic shield  Thoracic legs  Spiracle  Primary and Prespiracular pinaculum (L-group) secondary setae on prothorax (T1)  Chaetotaxy Abdomen

 Spiracles  Abdominal prolegs  Crochets  Primary and secondary setae  Anal shield  Chaetotaxy

Stehr 1987 Abdomen

 Spiracles ◦ On segments A1-8  Abdominal prolegs  Crochets  Primary and secondary setae  Anal shield  Chaetotaxy Abdomen

 Spiracles  Abdominal prolegs ◦ Segments A3-6, A10 ◦ (Plusiinae) A5-6, A10 ◦ Geometridae A6, A10  Crochets  Primary and secondary setae  Anal shield  Chaetotaxy Abdomen

 Spiracles  Abdominal prolegs ◦ Segments A3-6, A10 ◦ Noctuidae (Plusiinae) A5-6, A10 ◦ Geometridae A6, A10  Crochets Plusiinae prolegs A5-6, A10  Primary and secondary setae  Anal shield  Chaetotaxy

Geometridae prolegs A6, A10 Abdomen

 Spiracles  Abdominal prolegs  Crochets ◦ Arrangement  Circle  Ellipse  Mesoseries ◦ Length  Uniordinal  Biordinal  Triordinal  Primary and secondary setae  Anal shield  Chaetotaxy

Stehr 1987 Abdomen Primary setae Secondary setae

 Spiracles  Abdominal prolegs  Crochets  Primary and secondary setae  Anal shield  Chaetotaxy

Geometridae Arctiinae Tortricidae Lymantriinae Noctuidae Abdomen

 Spiracles  Abdominal prolegs  Crochets  Primary and secondary setae  Anal shield  Chaetotaxy Abdomen

 Spiracles  Abdominal prolegs  Crochets Names and positions of primary setae:  Primary and secondary D = dorsal setae SD = subdorsal  Anal shield L = lateral SV = subventral  Chaetotaxy V = ventral Selected references

 Common, I. F. B. 1990. of Australia. Melbourne University Publishing. 535 pp.  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.  Peterson, A. 1948. Larvae of insects. Part 1: Lepidoptera and Hymenoptera. Columbus, Ohio. 315 pp.  Scoble, M. J. 1992. The Lepidoptera: form function and diversity. Oxford University Press. 404 pp.  Stehr, F. W. 1987. Immature Insects, Volume 1. Kendall/Hunt, Dubuque, Iowa. 754 pp.  Wagner, D. L. 2005. Caterpillars of eastern North America: A guide to identification and natural history. Princeton University Press. 512 pp.