Professional Documents
Culture Documents
AZRAreviewarticle
AZRAreviewarticle
net/publication/342591184
CITATION READS
1 8,102
5 authors, including:
Some of the authors of this publication are also working on these related projects:
All content following this page was uploaded by Shashank P. R. on 03 December 2020.
1929). Insects were often perceived to arise spontaneously (a conception, often dubbed as “the
theory of spontaneous generation”, which was later disproved by various workers).
Table-1: Important books on insect classification in past and their authors (based on ENGEL
and KRISTENSEN, 2013)
Author Book Title
Aristotle (384-322 BC) Historia Animalium
St. Isidore of Seville (ca. 560-636 BC) Etymologiae
UlisswAldrovandi of Bologna (1522-1603) De Animalibus Insectin Libri VII
Thomas Mouffet (1553-1604) Insectorumsive Minimorum Animalium
Theatrum
John Ray (1627-1705) Historia Insectorum
Maria SibyllaMerian (1647-1717) Metamorphosis Insectorum Surinamensium
August Johann Rösel von Rosenhof Insecten-Belustigung
(1705-1759)
Carl Linnaeus (1707-1778) Systema Naturae, Fauna Svecica
Johan Christian Fabricius (1745-1808) Philosophia Entomologica, Systema
Entomologiae, Genera Insectorum
PierreAndrè Latreille (1762-1833) Précis des Caractères Génériques des
Insectes
William Sharp MacLeay (1792-1865) Horae Entomologicae (Essays on the
Annulose Animals)
William Kirby (1759-1850) and An Introduction to Entomology
William Spence (1783-1850)
Karl Hermann Konrad Burmeister De Insectorum Systemate Naturali,
(1807–1892) Handbuch der Entomologie
John Obadiah Westwood (1805–1893) An Introduction to the Modern
Classification of Insects
Ernst Heinrich PhilippAugust Haeckel Generelle Morphologie
(1834–1919)
Alpheus Spring Packard (1839–1905) Guide to the Study of Insects
Anton Handlirsch (1865–1935) Die Fossilen Insekten
Frank M. Carpenter (1902–1994) Brues & Melander’s Classification of Insects
(2nd ed.), Treatise on Invertebrate
Paleontology
August D. Imms (1881–1949) General Textbook of Entomology
Emil Hans Willi Hennig (1913–1976) Grundzügeeiner Theorie der
phylogenetischen Systematik,
Die Stammesgeschichte der Insekten
Saint Isidore of Seville (ca. 560-636 BC), in his book Etymologie, recognized two
separate groups, De verminibus and De minutisvolatibus (though both these groups were of
insects, these were not included formally under a common insect group) (Barney et al., 2006)
(Fig. 1).
JOURNAL OF APPLIED ZOOLOGICAL RESEARCHES 03
Fig. 1: Two distinct and mutually unrelated groups of “Insects” proposed by Saint Isidore of Seville
Invention of microscope and printing press during 15 century (GUENTHER, 2002) paved
th
way for remarkable revisions of the insect classification. Marcello Malphighi (1628-1694)
and Antoine van Leeuwenhoek (1632-1723) studied anatomy of insects using microscope.
Inspired by Malphighi study on silkworm, Jan Swammerdam (1637–1680) became pioneer
in microscopic study of insect anatomy (SMITH, MITTLER and SMITH, 1973) and has put
forward a revolutionary classification of insects in 1669 based on their modes of
development, which is in use still today. Ulisse Aldrovandi (1522-1605) of Bologna is
credited to write the first specialized text exclusively on the study of insects (OGILVIE,
2008). Aldrovandi in 1602 distinguished insects based on their habitat mainly land and
water (terrestrial and aquatic) and attempted classification of insects based on wing and leg
morphology. Thomas Mouffet (1553-1604) attempted to classify insects based on their
habits in his book Insectorum sive Minimorum Animalium Thetrum (KRISTENSEN, 1999).
An influential work of this era on classification of insects based on their morphology,
biology, ecology and anatomy was produced by John Ray (1627-1705) in his book Historia
Insectorum (RAVEN, 1942). Maria Sibylla Merian (1647-1717), Furchault de Reaumur
(1683-1756) and Rösel von Rosenhof (1705-1759) extensively studied the life histories of
insects and along with John Ray, they influenced the research work of Linnaeus (ENGEL and
KRISTENSEN, 2013).
Gustav Schich (1833-1899) and Franstiśek Klapá lek (1863-1919) emphasized on the
structure of the thorax and mode of locomotion, John Bernhardt Smith (1858-1912) focused
on mouth parts and thoracic characters, Benjamin Cooke (1816-1833) emphasized on nature
of the pupa, Carlo Emery and Vein Graber (1844-1892) considered development and
embryology more important for classifying the insects (WILSON and DONER, 1937).
Ernst Heinrich Philipp August Haeckel (1834-1919) for the first time proposed an
explicit phylogeny of the insect orders (termed articulata) in his book Generelle
Morphologie (WILLMANN, 2003). Paul Mayer (1848-1923) attempted to reconstruct an
ancestral insect Protentomon and suggested parallel evolution of wingless hexapods and
winged hexapods. Alpheus Hyatt (1838-1902) and Jennie M. Arms (1852-1937) are credited
for constructing an explicit evolutionary tree of insects and they considered Ephemeroptera
as a distinct order. Friedrich, M. Brauer (1832-1904) was the first to classify insects based on
Darwinism (BRAUER, 1869; BRAUER, 1885) (Fig. 7).
In his classification, Brauer put all primarily wingless taxa under Apterygogenea
and all secondarily wingless taxa and winged forms under Pterygogenea. He along with
Labbock suggested independent evolution of Apterygogenea and Pterygogenea. He
recognised 16 orders. Earlier he put termites, bark lice and true lice under Corrodentia and
also accepted Erichson’s grouping of mayflies, dragonflies, damselflies, stoneflies and
Comstock’s grouping of corrodentia under Pseudoneuroptera. But later he recognised the
artificiality of these groups and classified them as distinct orders, Isoptera, Ephemeroptera
(then called as Ephemerida), Odonata, Plecoptera, Mallaphaga and Corrodentia (equivalent
to Psocoptera). He also suggested a new order Panorpatae (now known as Mecoptera or
Mecaptera) for scorpionflies same time with Packard, and Hyatt and Arms. Alpheus Spring
Packard (1839-1905) proposed an evolutionary classification in which hierarchy was
generated by creating superorders, orders and suborders. He recognized Eurynchota
(modern Paraneoptera) under superorder Phyloptera. He also recognized Orthoptera,
Dermaptera, Neuroptera (for Trichoptera and Planipennia) and Pseudoneuroptera (for
Odonata, Ephemeroptera and Platyptera) as orders (SMITH et al., 1973; ENGEL and
KRISTENSEN, 2013).
The classification system of insects in this era showed a progressive shift in pattern
over previous systems as it involved evolution of systems based on diverse characters,
analogy-homology distinction and dealing with problems concerned with assigning
weightage to conflicting characters, etc. This era witnessed the rise of the field of
Palaeoentomology. Scientists started exploring palaeontological evidences to understand
JOURNAL OF APPLIED ZOOLOGICAL RESEARCHES 07
Anton Handirsch (1865-1935) recognized that it is not necessary that all fossil
lineages must leave modern counterparts rather some become extinct without leaving any
successor. He proposed few extinct orders too. Handlirsch wrote one of the most
comprehensive accounts of fossil insects in the form of three volume monolith named Die
FossilenInsekten integrating Palaeontology and Neoentomology. He proposed evolutionary
history of insects based on palaeontological evidence. His accounts also suffered few
drawbacks like he wrote that pterygotes arose from a trilobite ancestor which was found to be
incorrect (GRIMALDI, 2001).
Four new extant higher-rank taxa were described in the beginning of 20th century
that played a pivotal role in studying insect phylogeny. These are Protura, Zoraptera,
Grylloblattodea and nammochoristid scorpion flies. Based on the fundamental structure of
insect wings, Martynov proposed division of winged insects (Pterygota) into Palaeoptera
and Neoptera (ENGEL and KRISTENSEN, 2013). Most entomologists supported common
origin of pterygotes but Lemche suggested a diphyletic origin of winged insects dividing
insects into two broad groups, Plagioptera and Opisthoptera, and their further sub-divisions
as shown in Fig. 9 (MATSUDA, 1981).
As the name of this era suggests, this period marks the occurrence of Hennigian
Revolution (or Cladistic Revolution) that led to the development of the field of Phylogenetic
Systematics. Phylogenetic Systematics deals with organising (more precisely
systematising) organisms solely on the basis of synapomorphy into taxa where each taxon
includes organisms that are strctly monophyletic in nature. This very concept of
Phylogenetic Systematics was materialised by Emil Hans Willi Hennig’s (1913–1976) in his
book Grundzügeeiner Theorie der phylogenetischen Systematik (ENGEL and
KRISTENSEN, 2013). He discarded taxa Apterygota except Thysanura, where the latter
included the non-monophyletic taxa Archaeognatha and Zygentoma. He proposed names for
putative monophyletic taxa that included Zygentoma and Pterygota as “Dicondyla” and that
included Psocoptera, Mallophaga and Anoplura as “Psocodea”. He included monophyletic
Odonata along with Neoptera within Boner’s Metapterygota. He accepted monophyly of
Parametabola (Paraneoptera), Saltatoria (Orthoptera) and the clade containing Mantodea
along with Blattodea and Isoptera.
Stammesgeschichte der Insekten”. In this book he proposed more resolved and natural order-
level phylogeny of insects. He recognized monophyletic nature of Ellipura, Palaeoptera, and
Paurometabola (this includes Polyneoptera minus Plecoptera). He proposed relationship
among different taxa within monophyletic Paurometabola as represented in Fig. 10
(HENNIG, 1969; 1981).
Fig. 10: Tree showing relationship among different Taxa under Monophyletic Clade
Paurometabola as proposed by Hennig (branch length not up to scale)
insufficient number of evidences or literature cited in favour of many of his findings. In this
regard, Hennig ambiguously placed Strepsiptera under Endopterygota that was questioned
by Kristensen. Also the unresolved status of assemblage/clade containing Plecoptera plus
Paurometabola was brought into picture (KRISTENSEN, 1975).
Later, H. Bruce Boudreux provided a much resolved hexapod tree consisting of both
insects and non-insect hexapods. He suggested sister group relationships between
Ephemeroptera and Neoptera, between Plecoptera and Embioptera, between
Gryllablattodea and clade comprising of Zoraptera and assemblage of Isoptera and
Blattodea-Mantodea pair within. Among Holometabolans, he suggested sister group
relationship between Neuropterida and Mecopteroida (BOUDREUX, 1979).
In the same era, insect Palaeontology achieved new heights. Works of Aleksander
Grigorevich Sharav (1922-1973) and Boris Borisovich Rohdendorf (1904-1977) helped
Hennig to establish time of origin of the recognized high-rank clades (GRIMALDI and
ENGEL, 2005). KUKALOVÁ-PECK (2008) worked extensively on morphology of fossil
insects and studied origin of extant insect taxa based on paleontological evidences. She
established monophyly of Palaeoptera, included monophyletic clade Orthoneoptera
(consisting of Plecoptera, Embioptera, and Orthoptera) within neoptera, established
monophyly of Blatoneoptera (consisting of Dermaptera, Gryllablattodea and Dictyoptera),
and recognized sister group relationship among Orthoneoptera, Blatoneoptera and clade
consisting of Paraneoptera plus Endopterygota (KUKALOVA-PECK and
BRAUCKMANN, 1992 and KUKALOVA-PECK, 2008).
Table-2: List of some important fossil insect taxa (based on GRIMALDI, 2001)
S. Order Fossil Taxa Site of Discovery Period
No.
1. COLLEMBOLA Rhyniellapraecursor Scotland Devonian
2. COLLEMBOLA Permobrya mirabilis South Africa Permian
3. ARCHAEOGNATHA Monura (Dasyleptus) Europe Carboniferous
4. ARCHAEOGNATHA Triassomachilis Europe Triassic
5. Lepidothrichidae* Lepidotrix Baltic Eocene amber
6. EPHEMEROPTERA Lithoneura Illinois Carboniferous
7. EPHEMEROPTERA Protereisma Kansas, Oklahoma Permian
8. Palaeodictyopteroidea** Thuringopteryx Germany Triassic
9. MANTODEA Jersimantis New Jersey Cretaceous amber
10. ISOPTERA Meiatermes Spain Cretaceous
11. ISOPTERA Valditermes England Cretaceous
12. ISOPTERA Cretatermes carpenteri Canada Cretaceous
13. ISOPTERA Carinatermes New Jersey Cretaceous amber
14. EMBIOPTERA Burmitembia Burma Cretaceous amber
*Extinct family. **Extinct super order.
Table-2 shows list of some of the important insect fossils described from these fossil deposits
(GRIMALDI, 2001).
To gain wider and more accurate insights, modern entomologists use sequence data
from both Genomic DNA as well as Mitochondrial DNA. Using Mitochondrial DNA
sequence has some specific advantages as it is maternal in origin, simple in structural design
as it doesn't contain introns and have lost recombination ability, making it a reliable tool to
trace point of divergence and extant of accumulation of genetic changes in an organism over
time (DESALLE, 2017). For instance, mitochondrial Cyochrome Oxidase-I gene is being
used universally to generate DNA Barcode of different species among invertebrates
(FOLMER et al., 1994). DNA Barcodes are molecular signatures based on specific DNA
sequences that are unique for each individual species. Also, the availability of in silico tools
and publically accessible data bases containing DNA sequences over internet has
revolutionized the area of molecular analyses to establish more reliable phylogenetic
relationships among taxa (as well as individual organisms) (WILSON et al., 2017).
This classification has also been accepted and followed by Royal Entomological
Society. Many modern zoologists and entomologists like RUPPERT, FOX and BARNES
(2004); BRUSCA and BRUSCA (2003); ANDERSON (2001) follow or have proposed
somewhat similar classification of insects with minor modifications.This classification
takes into account the monophyly of each taxa to avoid coming together of non-
monophyletic taxa and to allow closely related taxa to be grouped together. In this regard,
monophyletic order Dictyoptera includes three suborders Blattodea, Mantodea and Isoptera
that were demoted from the rank of independent orders, the reason that led these suborders to
get their names mentioned in Table-3 as well. Name of many orders have kept as it is to avoid
confusion and to allow easy comparison with respect to older classifications. For many such
reasons, present classification of insects also represents phylogeny of insect taxa. This
classification is simple, easy-to-follow yet accurate in comparison to earlier classifications
of insects.
Table-3: Modern Classification of Class Insecta (BARNARD, 2011)
Class INSECTA
SubclassApterygota
OrderArchaeognatha or Microcoryphia (Bristletails)
Order Zygentoma (Silverfish and firebrats)
Subclass Palaeoptera
Order Ephemeroptera (Mayflies or upwing flies)
Order Odonata (Dragonflies and Damselflies)
Subclass Polyneoptera
Order Dermaptera (Earwigs)
Order Dictyoptera (Cockroaches, termites and mantids)
Suborder Blattodea, Blattaria, or Blattoptera (Cockroaches)
Suborder Mantodea (Mantids)
Suborder Isoptera (Termites)
Order Embioptera (Webspinners)
Order Grylloblattaria (Rock crawlers)
Order Mantophasmatodea (Heelwalkers)
Order Orthoptera (Grasshoppers, crickets and bush-crickets)
Order Phasmida (Stick insects)
Order Plecoptera (Stoneflies)
Order Zoraptera (Zorapterans)
Subclass Paraneoptera
Order Hemiptera (True bugs)
Order Phthiraptera (Sucking and biting lice)
Order Psocoptera (Booklice and barklice)
Order Thysanoptera (Thrips)
Subclass Endopterygota
Order Coleoptera (Beetles)
Order Diptera (True flies)
Order Hymenoptera (Ants, bees and wasps)
Order Lepidoptera (Butterflies and moths)
Order Mecoptera (Scorpionflies)
Order Megaloptera (Alderflies)
Order Neuroptera (Lacewings)
Order Raphidioptera (Snakeflies)
Order Siphonaptera (Fleas)
Order Strepsiptera (Stylops)
Order Trichoptera (Caddisflies or sedge flies)
JOURNAL OF APPLIED ZOOLOGICAL RESEARCHES 13
REFERENCES
ANDERSON, D. T. 2001. Invertebrate Zoology, 2nd edn. Oxford University Press, Victoria, pp 1-
476.
APPEL, T. A. 1987. The Cuvier-Geoffroy Debate: French Biology in the Decades Before Darwin.
Oxford Univ. Press, Oxford, U.K., pp 1-334.
BARNARD, P. C. 2011. The Royal Entomological Society Book of British Insects. Wiley-Blackwell.
pp 1-396.
BARNEY, S.A., LEWIS, W. J., BEACH, J.A. and BERGHOF, O. 2006. The Etymologies of Isidore of
Seville. Cambridge Univ. Press, Cambridge, U.K., pp 1-488.
BOUDREAUX, H. B. 1979. Arthropod Phylogeny with Special Reference to Insects. Wiley, New
York. pp 1-320.
BRAUER, F. 1869. Betrachtungenuber die Verwandlung der Insektenim Sinne der Descendenz-
Theorie. Verhandl. k.-k. Zool. Bot. Ges., 19: 299–318.
BRAUER, F. 1885. Systematische-ZoologischeStudien. Sitzungsber. kais. Akad. Wissen., Math.-
Naturwissen. Klass., Abt. 1, Mineral. Bot. Zool. Anat. Geol. Palaontol., 91: 237–414.
BRUSCA, R. C. and BRUSCA, G. J. 2003. Invertebrates. Sinauer Associates, Sunderland,
Massachusetts. pp 1-922.
CARPENTER, F. M. 1992. Superclass Hexapoda. In Treatise on Invertebrate Paleontology, Part R,
Arthropoda 4, Vols. 3–4, ed. Kaesler, R.L. Geol. Soc. Am. Univ. Kans, Lawrence, pp 1-655.
CLARK, J. F. M. 2009. Bugs and the Victorians. Yale University Press. pp 1-322.
CRAMPTON, G. C. 1938. The Interrelationships and Lines of Descent of Living Insects. Psyche, 45:
165–181.
DANA, J. D. 1864. The classification of animals based on the principle of cephalization. No. II.
Classification of Insects. Am. J. Sci. Ser. 2. 37: 10–33.
DESALLE, R., SCHIERWATER, B. and HADRYS, H. 2017. MtDNA: The Small Workhorse of
Evolutionary Studies. Frontiers in Bioscience, Landmark. 22: 873-887.
ENGEL, M. S. and KRISTENSEN, N. P. 2013. A history of entomological classification. Annual
Review of Entomology, 58: 585-607.
FOLMER, O., BLACK, M., HOEH, W., LUTZ, R., and VRIJENHOEK, R. 1994. DNA Primers for
Amplification of Mitochondrial Cytochrome C Oxidase Subunit I from Diverse Metazoan
Invertebrates. Molecular Marine Biol. & Biotech. 3(5): 294-299.
14 JOURNAL OF APPLIED ZOOLOGICAL RESEARCHES
FREEMAN, R. B. 1977. The Works of Charles Darwin: An Annotated Bibliographical Handlist. 2nd
edn. Folkstone, Dawson, pp 1-235.
GRIMALDI, D. 2001. Insect Evolutionary History from Handlirsch to Hennig, and Beyond. J.
Paleont., 75(6): 1152–1160.
GRIMALDI, D. and ENGEL, M. S. 2005. Evolution of the Insects. Cambridge Univ. Press,
Cambridge, U.K., pp 1-760.
GUENTHER,A. H. 2002. International Trends inApplied Optics. SPIE Press, Bellingham, WA., pp 1-
697
HENNIG, W. 1969. Die Stammesgeschichte der Insekten. Kramer, Frankfurt, pp 1-436.
HENNIG, W. 1981. Insect Phylogeny. Wiley, New York, pp 1-514.
HEYMONS, R. 1909. Die verschiedenen Formen der Insecten-metamorphose und ihreBedeutungim
Vergleichzur Metamorphose anderer Arthropoden. Ergeb. Fortschr. Zool., 1: 137–188.
KRISTENSEN, N. P. 1975. The Phylogeny of Hexapod “Orders”. A Critical Review of Recent
Accounts. Z. Zool. Syst. Evol. 13: 1–44.
KRISTENSEN, N. P. 1999. Lepidoptera, Moths and Butterflies: Evolution, Systematics and
Biogeography. Handbuch der Zoologie: Eine Naturgeschichte der Stämme des Tierreiches.
Arthropoda: Insecta. Walter de Gruyter, Berlin, 4(35): 1-5
KUKALOVA-PECK, J. 2008. Phylogeny of higher taxa in Insecta: finding synapomorphies in the
extant fauna and separating them from homoplasies, Evol. Biol. 35: 4–51.
KUKALOVA-PECK, J. and BRAUCKMANN, C. 1992. Most Paleozoic Protorthoptera are
Ancestral Hemipteroids: Major Wing Braces as Clues to a New Phylogeny of Neoptera
(Insecta). Canadian J. Zool., 70: 2452–2473.
LINNAEUS, C. 1758. Systema Naturae per Regna TriaNatura, Secundum Classes, Ordines, Genera,
Species, cum Characteribus, Differentiis Synonymis, Locis. Laurentii Salvii, Stockholm. 10th
ed., Rev. pp 1-1384.
MATSUDA, R. 1981. The origin of insect wings (Arthropoda :Insecta). International Journal of
Insect Morphology and Embryology. 10(5, 6): 387-398.
MILLER, W., MAKOVA, K. D., NEKRUTENKO, A. and HARDISON, R. C. 2004. Comparative
Genomics. Annual Review of Genomics and Human Genetics. 5: 15-56.
OGILVIE, B. W. 2008. Nature's Bible: Insects in Seventeenth-Century European Art and Science,
Tidsskrift for kulturforskning, 7(3): 5-21.
PARSONS, G. L. 2015. Recent Classification of Orders of Insects. Presented at seminar to
th
Entomology Department on 12 October 2015 at Michigan State University.
PARSONS, G. L., CASSIS, G., MOLDENKE, A. R., LATTIN, J. D., ANDERSON, N. H., MILLER,
J. C., HAMMOND, P., and SCHOWALTER, T. D. 1991. Invertebrates of the H. J. Andrews
Experimental Forest, Western Cascade Range, Oregon. V: An annotated list of insects and
other arthropods, U.S. Department of Agriculture, Forest Service, Pacific Northwest Research
Station. pp 1-168.
RACINE, V. 2013. Essay: The Cuvier-Geoffroy Debate. Embryo Project Encyclopedia.
http://embryo.asu.edu/pages/essay-cuvier-geoffroy-debate.
RAVEN, C. E. 1942. New Light on John Ray. Proc. Linnean Society London. 154 (1): 3-10
RICHARDS, O. W and DAVIES, R.G. 1957. Imms' A General Textbook of Entomology: Including the
th
Anatomy, Physiology, Development and Classification of Insects (9 edn.), Methuen, London.
pp 1-886.
RICHARDS, O. W and DAVIES, R.G. 1977. Imms' General Textbook of Entomology: vol. 2:
Classification and Biology (10th edn.). Chapman & Hall, London. pp 421-1354.
RUPPERT, E. E., FOX, R. S., and BARNES, R. D. 2004. Invertebrate zoology: A functional
evolutionary approach. Thomson-Brooks/Cole: Belmont, C.A., pp 1-963.
SCHLICK-STEINER, B. C., STEINER, F.M., SEIFERT, B., STAUFFER, C., CHRISTIAN, E. and
JOURNAL OF APPLIED ZOOLOGICAL RESEARCHES 15