Standard descriptive matrices in the identification of exophytophagous caterpillars


  • Aleksandra D. Trajković Faculty of Sciences and Mathematics, Department of Biology and Ecology, University of Niš, Višegradska 33, 18000, Serbia
  • Maja J. Lazarević Faculty of Sciences and Mathematics, Department of Biology and Ecology, University of Niš, Višegradska 33, 18000, Serbia
  • Saša S. Stanković Faculty of Sciences and Mathematics, Department of Biology and Ecology, University of Niš, Višegradska 33, 18000, Serbia
  • Miloš A. Popović Faculty of Sciences and Mathematics, Department of Biology and Ecology, University of Niš, Višegradska 33, 18000, Serbia
  • Marijana M. Ilić Milošević Faculty of Sciences and Mathematics, Department of Biology and Ecology, University of Niš, Višegradska 33, 18000, Serbia
  • Vladimir A. Žikić Faculty of Sciences and Mathematics, Department of Biology and Ecology, University of Niš, Višegradska 33, 18000, Serbia



autecology, morphology, Lepidoptera, polyphenism


Paper description:

  • Larvae of the order Lepidoptera display an abundance of macromorphological, autecological and behavioral profiles.
  • By defining and coding universal characters, an exhaustive description can be transformed into data matrices suitable for further analytical exploration.
  • The process of generating caterpillar identification tools takes phenotypic plasticity in the Lepidoptera order into account.
  • Incorporating the matrices into regular data processing, caterpillar specimens can provide the basis for inspecting character relationships and their states at intra- and interspecific levels.

Abstract: Identification of exophytophagous lepidopteran larvae is a necessity for researchers in biological disciplines ranging from biodiversity inventorying to research in parasitoid evolution and species monitoring. The lack of expertise in the field jeopardizes the outcomes of further investigations and recording of the multilevel plasticity of juvenile Lepidoptera. This paper offers an improvement to the existing haphazard approach by developing 41 simplified characters that include 150 morphological, behavioral and autecological states and their delineation, visual validation, and a descriptive matrix for 83 heterogeneous species. By combining the states into all possible identification scenarios, the matrix revealed 582 morphological, habitat and resource polyphenisms for the mentioned species. The categorical nature of the data implied the use of categorical principal component analysis to visualize the discriminative capacity without character relationship assumptions. The object-point biplot was used to derive the K value for K-mode clustering, while the cluster membership was introduced as a labeling variable to further inspect the grouping pattern. The results of this descriptive-analytic research indicate that descriptive matrices will allow continuous expansion and fine examination of many different species assemblages. From interactive identification keys to machine learning training, the presented framework can make data storage and interpretation significantly more attainable.


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Capinera, JL. Handbook of Vegetable Pests, 1st ed. San Diego, California: Academic Press; 2001. Order Lepidoptera: Caterpillars, Moths, and Butterflies; pp. 353-510.

Van Nieukerken EJ, Kaila L, Kitching IJ, Kristensen NP, Lees DC, Minet J, Mitter C, Mutanen M, Regier JC, Simonsen TJ, Wahlberg N. Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa. 2011;3148(1):212-21.

Clark BR, Godfray HC, Kitching IJ, Mayo S, Scoble MJ. Taxonomy as an eScience. Philos Trans Royal Soc. 2009;367(1890):953-66.

Dyar HG. The number of molts of lepidopterous larvae. Psyche. 1980;5(175-176):420-2.

Dyar HG. A Classification of lepidopterous larvae. Ann N Y Acad Sci. 1984;8(1):194-232.

Dyar HG. Descriptions of the larvae of some moths from Colorado. Proc U S Natl Mus. 1902;25 (1290):369-412.

Dyar HG. Life Histories of North American Geometridae - XLII. Psyche. 1903;10(325-326):190-200.

Frohawk, FW. Natural History of British Butterflies, Vol. 2, 1st ed. London: Hutchinson & Co.; 1924.

Hensley SD. Comparative Study of the Immature Stages of Three Species of the Diatraea Complex. [disseration]. [Oklahoma]: Oklahoma State University; 1960. 90 p.

Barbier R. Recherches sur la morphologie tegumentaire d'un insecte holométabole: Galleria mellionella L. (Lepidoptera, Pyralidae). [disseration]. [Rennes]: Université de Rennes; 1971. 205 p.

Fink LS. Color polymorphism in Sphingid caterpillars (Lepidoptera: Sphingidae). [disseration]. [Gainesville, Florida]: University of Florida; 1989. 273 p.

Capps, HW. Keys for the identification of some lepidopterous larvae frequently intercepted at quarantine. United States Department of Agriculture, Agricultural Research Service; 1956. 37 p.

Gerasimov, AM. Caterpillars. Fauna of the USSR. Lepidoptera Insects. Moskva, Leningrad: Zoological Institute, Academy of Sciences USSR, 1952. 338 p.

Hinton HE. The larvae of the Lepidoptera associated with stored products. Bull Entomol Res. 1943;34(3):163-212.

Aitken AD. A key to the larvae of some species of Phycitinae (Lepidoptera, Pyralidae) associated with stored products, and of some related species. Bull Entomol Res. 1963;54(2):175-88.

Carter DJ, Kristensen NP. Classification and keys to higher taxa. Vol. 1, Lepidoptera, Moths and Butterflies. Berlin: Walter de Gruyter; 1998. 358 p.

Gorham JR. Insect and mite pests in food: An illustrated key. US Department of Agriculture, Agricultural Research Service; 1991. 665 p.

Weisman DM. Keys for the identification of some frequently intercepted lepidopterous larvae. US Department of Agriculture, APHIS; 1986. 64 p.

Solis M. Key to selected Pyraloidea (Lepidoptera) larvae intercepted at US ports of entry: revision of Pyraloidea in "Keys to some frequently intercepted Lepidopterous larvae" by Weisman 1986. Washington, D.C.: USDA Systematic Entomology Laboratory; 2006. 58 p.

Peterson A. The Larvae of Insects, Part I, Lepidoptera and Hymenoptera. Ohio: Columbus; 1948. 315 p.

Hinton HE. On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Trans R Entomol Soc. 1946;97(1):1-37.

Godfrey GL, Jahn OL, Fleming, E. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera, Noctuidae) of America north of Mexico. Washington, D.C.: U.S. Dept. of Agriculture; 1972. 265 p.

Stehr FW. Immature Insects. Vol. 1. Iowa: Kendall/Hunt Publishing Company; 1987. 770 p.

Marquis RJ, Passoa CS, Lill TJ, Whitfield BJ, Le Corff J, Forkner ER, Passoa AV. Illustrated guide to the immature Lepidoptera on Oaks in Missouri. Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Assessment and Applied Sciences Team; 2019. 369 p.

Kitching IJ, Rawlins JE. The Noctuoidea. In: Kristensen NP, editor. Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol. 1: Evolution, Systematics and Biogeography. Berlin: W. de Gruyter; 1999. p. 355-402.

McGuffin WC. Guide to the Geometridae of Canada (Lepidoptera). III, IV and V. Subfamilies Archiearinae, Oenochrominae, and Geometrinae. Mem Ent Soc Can. 1998;120(S145):3-56.

Kristensen NP, editor. Lepidoptera, moths and butterflies. Vol. 1: Evolution, Systematics, and Biogeography. New York: Walter de Gruyter; 1999. 491 p. (Handbook of Zoology; Vol. 4; Part 35).

Beck H. Die Larven der europäischen Noctuidae - Revision der Systematik der Noctuidae (Lepid., Noctuidae) Vol. 1 and 2, Herbipoliana 5. Marktleuthen: Verlag Dr. Ulf Eitschberger; 1999. 859 p.

Beck H. Die Larven der europäischen Noctuidae - Revision der Systematik der Noctuidae (Lepid., Noctuidae). Vol. 3 and 4, Herbipoliana 5. Marktleuthen: Verlag Dr. Ulf Eitschberger; 2000. 525 p.

Ahola M, Silvonen K. Larvae of Northern European Noctuidae. Vol. 3. Finland: KuvaSeppala Yhtiot Oy; 2010. 672 p.

Wagner DL, Schweitzer DF, Sullivan JB, Reardon RC. Owlet Caterpillars of Eastern North America (Lepidoptera: Noctuidae). Princeton and Oxford: Princeton University Press;2011. 576 p.

Marquis RJ, Lill TJ, Forkner ER, Le Corf J, Landosky MJ, Whitfield BJ. Declines and Resilience of Communities of Leaf Chewing Insects on Missouri Oaks Following Spring Frost and Summer Drought. Front Eco Evol. 2019;7:396.

Ahola M, Silvonen K, Kronholm H, Lundsten KE, Sundell, PR. Description of the larva of Xestia atrata (Morrison, 1874) (Lepidoptera: Noctuidae) with notes on the biology of the species. Entomol Fenn. 2006;17(2):65-72.

Singh GP, Goel SC. Body chaetotaxy of three noctuid caterpillars. (Lepidoptera). Deut Entomol Z. 1987;34(4-5):373-81.

Duarte M, Robbins RK. Immature stages of Calycopis bellera (Hewitson) and C. janeirica (Felder) (Lepidoptera, Lycaenidae, Theclinae, Eumaeini): Taxonomic significance and new evidence for detritivory. Zootaxa. 2009;39-61.

Ahola M, Kuisma M, Leinonen R, Saarenmaa H, Silvonen K. Description of immature stages of Xestia brunneopicta (Matsumura, 1925), with a key to the mature larvae of the European species of Xestia (Pachnobia) (Lepidoptera, Noctuidae). Nota Lepidopterol. 2015;38(1):75-88.

Pires C, Marinoni L. DNA barcoding and traditional taxonomy unified through integrative Taxonomy: A view that challenges the debate questioning both methodologies. Biota Neotrop. 2010;10(2):339-46.

Sharkey M, Brown B, Baker A, Mutanen M. Response to Zamani et al. (2020): The omission of critical data in the pursuit of "revolutionary" methods to accelerate the description of species. Zookeys. 2021;1033:191-201.

Carter DJ. Pest Lepidoptera of Europe: with special reference to the British Isles. Vol. 31, Springer Science & Business Media; 1984. 438 p.

Eichlin TD, Cunningham HB. The Plusiinae (Lepidoptera: Noctuidae) of America north of Mexico, emphasizing genitalic and larval morphology: U.S. Department of Agriculture, Agricultural Research Service. 1978; 122 p.

Turčáni M, Patočka J, Kulfan J. How to identify larvae of the protected species: Dioszeghyana schmidtii (Diószeghy 1935) and survey its presence and abundance (Lepidoptera: Noctuidae; Hadeninae). J For Sci. 2010;56(3):121-9.

Gilligan TM, Passoa SC. LepIntercept - An identification resource for intercepted Lepidoptera larvae. Identification Technology Program (ITP). 2014- [cited 2023 Jan 14]. Available from:

Anderson S, Majka, G. Biodiversity and biosystematic research in a brave new 21st century information-technology world. Zookeys. 2009;22:1-4.

Rennwald E, Rodeland E. Lepiforums-Europaliste Lepiforum's Checklist of European Lepidoptera [Internet]. Version 6, January 1st, 2019- [cited 2023 Jan 15]. Available from: web application. [Internet]. [cited 2022 Jun 10]. Available from:

Popović M, Vasić N, Koren T, Burić I, Živanović N, Kulijer D, Golubović A. Biologer: An open platform for collecting biodiversity data. Biodivers Data J. 2020;8:e53014.

HabiProt Alciphron - database about the insects of Serbia. [Internet]. 2014- [cited 2021 Nov 1]. Available from:

Moyle D, Byrne C. The Caterpillar Key. [Internet]. 2020- [cited 2022 Jun 25]. Available from:

Huang Z. Clustering categorical data with k-Modes. In: Wang J, editor. Encyclopedia of Data Warehousing and Mining, 2nd ed. New York: IGI Global; 2009, p. 246-50.

Futahashi R, Banno Y, Fujiwara H. Caterpillar color patterns are determined by a two-phase melanin gene prepatterning process: new evidence from tan and laccase2. Evo Dev. 2010;12(2):157-67.




How to Cite

Trajković AD, Lazarević MJ, Stanković SS, Popović MA, Ilić Milošević MM, Žikić VA. Standard descriptive matrices in the identification of exophytophagous caterpillars. Arch Biol Sci [Internet]. 2023Apr.3 [cited 2024Apr.22];75(1):89-102. Available from:




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