A History of Entomological Classification: Michael S. Engel and Niels P. Kristensen

EN58CH29-Engel ARI 28 November 2012 20:12
A History of Entomological
Classification
Michael S. Engel1,∗ and Niels P. Kristensen2
by NORTH CAROLINA STATE UNIVERSITY on 01/15/13. For personal use only.
Annu. Rev. Entomol. 2013.58:585-607. Downloaded from www.annualreviews.org
1
Division of Entomology, Natural History Museum, and Department of Ecology &
Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045;
email: msengel@ku.edu
2
Natural History Museum of Denmark, DK-2100 Copenhagen Ø, Denmark;
email: npkristensen@snm.ku.dk
Annu. Rev. Entomol. 2013. 58:585–607 Keywords

The Annual Review of Entomology is online at Insecta, orders, phylogeny, taxonomy, entomology, systematics
ento.annualreviews.org
This article’s doi: Abstract

10.1146/annurev-ento-120811-153536
The classification of insects has attempted to most effectively communicate
Copyright c 2013 by Annual Reviews. information about this hyperdiverse lineage of life and, not surprisingly, has
All rights reserved
had a considerably rich historical development. This history can be coarsely
∗
Corresponding author segregated into four periods: the Pre-Linnean era, the first century span-
ning Linnaeus’s Systema Naturae to Darwin’s On the Origin of Species, the
Darwinian era up to the Cladistic Revolution, and the Hennigian era leading
to today. The major events of each of these episodes are briefly summarized
and some of the more notable researchers highlighted, along with their in-
fluence on our current understanding of insect relationships and how this is
reflected in the current classification of the Hexapoda.
585
INTRODUCTION
Classifications have existed as long as humanity. It can be argued that the first instinct is to clas-
sify. We classify those things that are important to us and for which we wish to find meaning and
value. Given the overarching importance of classification to every aspect of scientific entomology,
it is therefore worthwhile to consider the origins and evolution of our current system of insects.
Naturally, much has been and can be written about the history of entomology in general. An
excellent source book is Smith et al.’s (129) History of Entomology, published as a special issue of
the Annual Review of Entomology, and Wilson & Doner (142) provide a somewhat disjointed but
overall interesting account of the history of insect classification. Herein we restrict ourselves to
major works that focus on the higher-level classification of insects and make no attempt to be ex-
haustive. In selecting images we chose to highlight individuals or iconography not so prominently
displayed in other entomological works, for example, attempting to avoid repetition with those
in Willmann’s (141) and Grimaldi & Engel’s (45) historical accounts. In addition, we have given
cursory treatment to the current era of entomological classification, as a compromise had to be

made regarding what should be covered. We preferred to devote such pages to those eras which
form the foundation of present-day work rather than enumerate in detail the events surrounding
us. We therefore beg the reader’s forgiveness for glossing over many fascinating events and ideas,
regardless of how seemingly grand or quizzical, in the historical development of entomological
classification.
THE PRE-LINNEAN PERIOD

One of the earliest surviving classifications of the natural world is that of Aristotle (384–322 BC)
(6), disciple of Plato and mentor to the young Alexander the Great. Certainly there are earlier
accounts of insects, but Aristotle’s Historia Animalium (6) is the earliest surviving work in which
a comprehensive organization of insects is to be found, incorporating them among a larger ac-
count of all life (see sidebar Insects in Antiquity). It was this system, in one form or another, that
would persist for the next fifteen centuries. Naturally, Aristotle and other ancient authors—Greek,
Roman, or otherwise—proposed groups of lasting value. They identified and differentiated bees,
wasps, ants, butterflies, flies, beetles, and so forth. These systems of organization persisted for mil-
lennia and, although gradually augmented and adapted for different purposes, changed relatively
little. Thus, while Gaius Plinius Secundus’s (23–79 AD) Naturalis Historia was largely independent
in the biological accounts it provided, it preserved continuity in classification (129).
INSECTS IN ANTIQUITY
Parables from the Holy Bible, such as the Septuagint’s “Go to the bee, and learn how diligent she is, and what a noble
work she produces, whose labors kings and private men use for their use, she is desired and honored by all, and though
weak in strength she values wisdom and prevails” (Proverbs, 6:8), highlight the way in which people of antiquity
keenly observed insects. Insects also figure prominently in less flattering light, such as the plagues upon Egypt
(Exodus, 8:16–19; 10:4–6, 10:12–15). Although some of the notions regarding the origins of insect species were
remarkably fanciful, such as bees emanating from the carcasses of bulls, authors of antiquity simultaneously made
amazingly accurate observations. Historia Animalium rightly grouped whales among the mammals and provided
brief accounts of the dancing of honey bees, and it must be recalled that even before Aristotle, Anaximander had
proposed a mechanism of evolution, one that included humans!
586 Engel · Kristensen

Medieval Christian scholars did their best to copy what elements of knowledge they felt most
important, and perhaps one of the most enduring works of this time was Saint Isidore of Seville’s
(ca. 560–636) Etymologiae (7). This singular tome attempted to provide in one work an encyclopedic
treatment of human knowledge, including a classification and account of the natural world. The
Etymologiae included accounts of termites, silkworms, and cantharids, among others, classified
as De verminibus (vermin), while things such as bees, wasps, moths, butterflies, locusts, roaches,
cicadas, and true flies were classified as De minutis volatibus (tiny flying animals).
The eleventh century brought forth presumed lost works by authors of antiquity, as well as
improvements upon them or outright novel accounts by Arabic scholars. These discoveries opened
a floodgate of information, although it took centuries to spread across Europe. The Renaissance
saw little in terms of formal classification. Instead, a diversity of scholars sought to look at insects
anew, using the new rational tools of the era, as well as the latest in technological achievements
such as optics, and disseminated their findings using the newly developed printing press. This
was an exciting time for entomology, with magnificent works by Jan Swammerdam (1637–1680),
Marcello Malpighi (1628–1694), Antonie van Leeuwenhoek (1632–1723), and others investigat-
ing in greater and more accurate detail the anatomy of insects (129), but few considered major
classificatory arrangements across the Insecta. Of note was Ulisse Aldrovandi (1522–1605) of
Bologna, who produced the massive folio work De Animalibus Insectis Libri VII (3), a tome that
represented the first specialized text exclusively on entomology. Aside from classification, he also
gave a summary of insect morphology, metamorphosis, reproduction, behavior, and physiology,
among other biological annotations. Thomas Mouffet’s (1553–1604) Insectorum sive Minimorum
Animalium Theatrum (104), a work amalgamating the efforts of numerous naturalists, was a re-
lated attempt at classifying insects and their habits but lacked the finesse and detail of Aldrovandi’s
effort. However, many of the terms and names used by Mouffet were subsequently adopted by
Linnaeus as names for his genera.
Toward the beginning of the Enlightenment we find authors again more rigorously exploring
the taxonomy and classification of insects. John Ray (1627–1705) relied heavily on the anatom-
ical work of Swammerdam and attempted to develop a classification of insects based on their
morphology, biology, and ecology, summarized in his posthumously published Historia Insectorum
(114). Although his classification had overlapping categories, confusing to a modern reader, it was
immensely influential and many of the names he applied to groups were adopted by subsequent
authors, including Linnaeus. More remarkably, Ray brought forth a definition for species and
codified the notion that species be applied to all individuals from the same progenitor.
Three other individuals worth mentioning at this time are Maria Sibylla Merian (1647–1717),
René Antoine Ferchault de Réaumur (1683–1756), and August Johann Rösel von Rosenhof (1705–
1759). Merian published her findings in 1705 as the sumptuously illustrated Metamorphosis Insec-
torum Surinamensium (98). Réaumur was perhaps the most tireless observer of insect biology and
behavior of the time (115, 129). He openly disliked the efforts of developing classifications, instead
focusing his sharp intellect on the life histories of his subjects. Rösel von Rosenhof ’s engravings
and life-history accounts were subsequently bound together in four volumes as Insecten-Belustigung
(120, 129). All three would have a lasting impression on subsequent entomological authors, pro-
viding much original information about numerous insect species for the likes of Linnaeus and
Fabricius, who would never have the chance to observe those species directly or learn anything of
their habits.
THE FIRST CENTURY (1758–1859)

Around the middle of the eighteenth century, Carl Linnaeus (1707–1778) adopted uniformly his
system of binomial nomenclature and in the tenth, revised edition of his Systema Naturae put
www.annualreviews.org • A History of Entomological Classification 587

LINNEAN INSECTA
Linnaeus recognized all arthropods as his fifth class (in 1758), or class Insecta, segregating them into a series of
seven orders based on the nature (or outright absence) of their wings. Hence, based on the primacy Linnaeus gave
to wings over other characters of insect anatomy, we arrived at the familiar (in order of their appearance in his
system) Coleoptera, Hemiptera, Lepidoptera, Neuroptera, Hymenoptera, Diptera, and Aptera. Linnaeus did not
focus solely on single characters, however, and even in earlier editions highlighted the structure of the antennae
among other traits, but had to place emphasis somewhere. The orders were grouped by number of wings (0, 2, or 4),
then those in the alae 4 category were divided into superiores versus omnes, the former containing the Coleoptera (or
crustaceae totae, “fully hardened”) and Hemiptera (semicrustaceae, “partially hardened”) and the latter containing first
the Lepidoptera (imbricatae squamis, “overlapping scales”) and then those with membranous wings (membranaceae,
including the Hymenoptera and Neuroptera). Linnaeus then divided the orders into his numerous genera and
species.
forth a classification of insects (88). This system was in use in earlier editions and incorporated
much information from earlier writers such as Ray and Merian, among others, but retrospectively
is taken as the starting point for our modern system of zoological nomenclature. Insect orders
were consistently applied as early as the first edition (1735), recognizing four orders {Coleoptera,
Angioptera [later Gymnaptera in the second edition (1740)], Hemiptera, and Aptera}, but by the
time of the Fauna Svecica (1746) he had arrived at an arrangement of seven orders to which he
held for the remainder of this life (see sidebar Linnean Insecta). It is often overlooked how much
importance Linnaeus gave to the authors of antiquity, but as noted, the intellectual impact and
importance of ancient Greek and Roman texts and the improvements upon them made by the
early Arabic scholars were indeed still palpable during the Enlightenment.
Following upon the comments Linnaeus made in the second edition of Systema Naturae about
the essential nature of the structure of the mouthparts, his foremost entomology student Johan
Christian Fabricius (1745–1808), the first systematist considered to specialize on insects, published
in 1778 his Philosophia Entomologica (40), the first textbook of entomology. Herein he synthesized all
available information on insects, including a rational means for their description and an exposition
on the primacy of the structure of the mouthparts. Fabricius believed that mouthparts would serve
to establish a classification that is natural for genera and species (given that they are vital to the
nourishment of every insect and thereby affect their whole biology) but remain artificial at the
level of orders. He felt that a natural system for the orders was for later generations, noting
in 1778 that, “we are only novices in the science” (40, 129). Fabricius had already set forth his
system of insects in 1775 (Systema Entomologiae), his system of genera in 1776 (Genera Insectorum),
and then numerous works on species, culminating in individual accounts of the species of each
order. Given the primacy he placed on the mouthparts, he expanded the number of orders and
altered the names of those recognized by his mentor, ultimately segregating the world’s fauna
into Eleutherata (Coleoptera), Rhyngota (Hemiptera), Piezata (Hymenoptera), Antliata (Diptera),
Glossata (Lepidoptera), Ulonata (Orthoptera), Synistata (Neuroptera), and Odonata.
Baron Charles De Geer (1720–1778), whose works were modeled on those of Réaumur, was
the first to provide a meager attempt at uniting the systems of Linnaeus and Fabricius, noting
the artificiality of classifying together insects with haustellate and mandibulate mouthparts, de-
spite similarities in their wings. He therefore attempted a simple unification of the Linnean and
Fabrician systems, using both wings and mouthparts to arrive at a slightly expanded system of
orders, giving us the Dermaptera (although his use of this name differed greatly from our modern

concept) (33). This step was the impetus for subsequent authors to reconsider the composition of
the Linnean and Fabrician orders, eventually leading Guillaume Antoine Olivier (1756–1814) to
propose the Orthoptera (107) and much later to the disintegration of the Neuroptera. De Geer’s
commentator, Andres J. Retzius (1742–1821), simplified the arrangement of groups and character
suites (116) but erroneously ascribed suctorial forms to De Geer’s Dermaptera.
Linnaeus, Fabricius, and others such as Étienne-Louis Geoffroy (1727–1810) noted that while
their classifications at one level or another remained artificial, with each addition of information
to the hierarchy, they were getting closer to a natural system (43). Giovanni Antonio Scopoli
(1723–1788), however, was the first to note that the classification should be based on the whole
structure of the insect, and not just on the wings, the mouthparts, the antennae, or the habits
of the organisms in question (123). He was unable to develop a more synthetic and natural sys-
tem. A holistic approach only materialized with the French entomological luminary Pierre André
Latreille (1762–1833), whose 1796 Précis des Caractères Génériques des Insectes introduced the rank
of families and a synthetic approach to the classification, utilizing multiple traits, bringing to-
gether the works of Linnaeus, Fabricius, and others, and developing the first classification pro-
fessed to be truly natural (86). In 1796 Latreille recognized Coleoptera, Orthoptera, Hemiptera,
Neuroptera, Hymenoptera, Lepidoptera, Diptera, Suceurs, Thysanoures, and Parasites, among
other arthropods, all as belonging to Insecta. This system was regularly augmented, reflecting the
latest available information on insect diversity over the next 35 years, as well as discourse with
his colleagues (including Fabricius, who visited Paris many times over several years). Latreille
and his contemporaries in Paris, such as the famed Étienne Geoffroy Saint-Hilaire (1772–1844),
Jean-Baptiste Lamarck (1744–1829), and Georges Cuvier (1769–1832), were making profound
strides in detailed comparative anatomy, including the first codification of a set of criteria for
the recognition of homology, debating incipient advances toward evolutionary theory and the
classification of the natural world (4).
Lamarck (83, 84) expounded the philosophical view that animals tended toward a circular
process, one in which the arrangement of organisms diverged in opposite directions for some
characters, while meeting again in another character. Such concepts were carried forward by
Johann Gotthelf Fischer von Waldheim (1771–1853), who used Lamarck’s ideas in classifying
insects and other animals (41). In terms of entomological classifications, such circularity in natural
progression saw its greatest advocate in William Sharp MacLeay (1792–1865). MacLeay developed
and promoted the Quinarian system of classification, whereby all higher categories of the animal
kingdom were segregated into groups of five. These were each pictographically represented by
a series of five circles bordering each other and forming a complete ring (Figure 1a). It was his
notion that each of these most closely resembled the next bordering taxon in regard to a particular
character(s), but that this transition series eventually came back upon itself. He believed strongly
that no grouping could be considered natural if it did not tend to exhibit a circular series. He also
believed that the primary divisions of such series were in groups of 10, 5 of which were the primary
elements of the series, the others being minor (osculant) and intercalating or bridging between
those of the major series. Starting first with the classification of Scarabaeus, MacLeay eventually
developed this concept into a classification of all Animalia (91). MacLeay’s Horae Entomologicae; or,
Essays on the Annulose Animals was hugely influential, albeit quickly recognized to have fundamental
flaws as the numbers of species and the characters they embodied soon failed to truly fall into
nice categories of five. This did not deter others from ascertaining that the true flaw was the
reliance on five, rather than the overarching philosophy itself. Indeed, Edward Newman (1801–
1876) expounded a largely identical system but instead considered the fundamental elements to
organize themselves into groups of seven (105, 106). The Septenary system had its own followers
(like Swinton) but suffered the same fate as that advocated by the Quinarians.

a b
c d
Figure 1
(a) William S. MacLeay’s Quinarian arrangement of the insect orders from the Horae Entomologicae (91). (b) A phylogeny of insect
orders as presented by Hyatt & Arms (62) in 1890, with the disc representing the surface of the Earth and the lines of descent (as
interpreted from Scudder’s work) extending back into the geological past. (c) Carl J.B. Börner’s 1904 depiction of relationships among
the principal lineages of hexapods (12). (d ) Guy Chester Crampton’s 1938 remarkably modern-looking topology of insects (30).

At about the same time as MacLeay, William Kirby (1759–1850), who was principally interested
in the aculeate Hymenoptera and Coleoptera, was the first to recognize the order Strepsiptera
and later the Trichoptera. In collaboration with William Spence (1783–1860) he produced An
Introduction to Entomology, four volumes covering all aspects of insect science including a revised
system of the higher categories, incorporating many ideas from MacLeay, as they meshed nicely
with his idea of harmony in the orderly hand of God (69, 129).
In his doctoral dissertation of 1829, De Insectorum Systemate Naturali (18), Karl Hermann
Konrad Burmeister (1807–1892) set forth a brief explanation for a revised system of the insect
orders, and later expanded this into his textbook of general entomology, Handbuch der Entomologie
(published in five volumes between 1832 and 1855). Burmeister greatly disliked Latreille’s sys-
tem and belittled him for changing it so regularly. Instead, Burmeister emphasized the different
kinds of metamorphosis he observed and, estimating the world’s fauna to consist of about 80,000
species, segregated them into eight orders—Hemiptera, Dictyoptera, Neuroptera, Diptera, Lep-
idoptera, Hymenoptera, Orthoptera, and Coleoptera. Burmeister’s system still emphasized a very
limited set of characters, but did make rudimentary attempts to bring together diverse data such
as metamorphosis alongside wing and mouthpart structure (19).
Perhaps the last of the great entomological systematists to propose a classification of the orders
prior to the Darwinian era was John Obadiah Westwood (1805–1893), the first Hope Professor
at Oxford University. His two-volume An Introduction to the Modern Classification of Insects (1839–
1840) attempted to unify the systems of Latreille and MacLeay (137). Although not a Quinarian,
Westwood’s deference to MacLeay is apparent throughout his introductory remarks regarding
the “distribution of insects into orders.” Westwood employed the same idea of first recognizing
the “older orders” (those of the earlier authors), and then seeking to classify osculant groups that
bridged these grander groupings, but never restricting himself to any predetermined numerical ar-
rangement. Accordingly, Westwood indicated that the class of “Hexapod metamorphotic insects”
consisted of two descending series (subclasses): those with a “mouth of jaws” (Dacnostomata) ver-
sus “mouth with a sucker” (Antliostomata). The former included the series order Hymenoptera,
osculant order Strepsiptera, order Coleoptera, osculant order Euplexoptera (the earwigs), order
Orthoptera, order Neuroptera, and order Trichoptera, with Westwood hedging what might be
the osculant groups representing bridges between the last three. The Trichoptera then repre-
sented a “direct passage” to the arrangement of the other subclass, although this time presented
in ascending fashion as order Lepidoptera, order Homoptera, order Heteroptera, osculant order
Aphaniptera (fleas), osculant order Homaloptera (Hippoboscidae and Nycteribiidae), and finally
order Diptera. The Thysanoptera were perplexing to Westwood and he hesitatingly considered
them as osculant between Orthoptera and Neuroptera (enticingly reminiscent of the placement
of Paraneoptera as between a more basal orthopteroid stem and the holometabolans!). Westwood
never accepted evolution and was an ardent opponent of Darwinism. His system was clearly estab-
lished outside an evolutionary framework, but by describing and noting such consistent patterns of
characters, Westwood was strengthening the higher classification of hexapods while unwittingly
laying the early foundation toward a phylogenetic arrangement of the orders.
THE DARWINIAN ERA: HAECKEL TO HENNIG

Work on the higher classification of insects continued largely unchanged following the publication
of Charles R. Darwin’s (1809–1882) On the Origin of Species in 1859. To the extent authors
adopted evolutionary thinking, the innovation was evident in rhetoric rather than methodological
approach, with the emphasis continuing to be on the perceived significance of different characters.
Indeed, Darwin himself noted, “Systematists will be able to pursue their labours as at present” (32),

and the enduring patterns described by taxonomists for generations were now newly enlivened by
a powerful, synergistic explanation of descent. Refinements in methods of microscopical anatomy
from the late-nineteenth century onward led to numerous excellent studies on the structure and/or
developmental origin of selected organs or body regions of many kinds of insects, but the findings
went unnoticed by several systematists, who lived in different worlds, continuing to emphasize a
limited number of characters in the exoskeleton.
James Dwight Dana (1813–1895) emphasized the importance of cephalization in ordering the
Animal kingdom, and he emphasized organizing the insects on the basis of the degree to which
more complex biological functions were centralized to the anterior of the species (31). Dana was
not alone in this endeavor. Both Gustav Schoch (1833–1899) and František Klapálek (1863–1919)
attempted to organize the insects largely on the basis of the structure of the thorax, effectively
emphasizing their modes of locomotion, while John Bernhardt Smith (1858–1912) emphasized
an amalgamation of mouthpart and thoracic characters (142). Benjamin Cooke (1816–1883) and
Ferdinand Anton Franz Karsch (1853–1936), building on the earlier work of Newman, did the
same but organized around the nature of the pupa, and Carlo Emery (1848–1925) and Veit Graber
(1844–1892) more generally organized around the development and embryology of insects (142).
Such authors, then, placed emphasis on those traits they considered most critical in the evolutionary
history of the class, or outright established what they believed were plausible mechanisms for
driving macroevolutionary patterns.
As in many things, it was the controversial Ernst Heinrich Philipp August Haeckel (1834–1919)
who first depicted, in his 1866 Generelle Morphologie, an explicit phylogeny of the insect orders,
here embedded within a topology encompassing all Articulata (45, 141). From this time onward,
the classification of the orders was increasingly presented alongside a depiction or explanation
of their lines of descent, such as the 1876 trees of Paul Mayer (1848–1923), who also attempted
a reconstruction of the ancestral insect Protentomon (97). A somewhat later example was the
depiction of an explicit evolutionary tree (Figure 1b) by Alpheus Hyatt (1838–1902) and Jennie
M. Arms (1852–1937), who were also the first scientists to consider the Ephemeroptera as a distinct
order (62).
In terms of serious entomological inquiry, however, it was arguably Friedrich M. Brauer (1832–
1904) (16) who made the first truly conceptual leap toward classifying insects along the lines of
Darwinian evolution. After expounding upon the general principles of organic evolution and
the unifying aspect of Darwin’s theory, Brauer set forth his results of applying said principles
to the subject of insect diversity, effectively distinguishing between primarily wingless taxa, his
Apterygogenea (Collembola and Thysanura), and the secondarily wingless taxa that together with
the winged forms constituted his Pterygogenea, which were derived from the apterygogeneans
(16). He thereby emphasized suggestions made earlier by himself (15) and Lubbock (90), while
contradicting Mayer, who had considered the primarily wingless hexapods coordinate with the
winged insects rather than ancestral to them. Moreover, Brauer segregated the pterygotes into
two broad categories, Polynephria and Oligonephria along one axis and their development along
another axis (Ametabola + Hemimetabola versus Metabola). Brauer’s system had overlapping
means of segregating the groups. If followed completely, it left him with an arrangement of 16
orders that supported earlier attempts to divide the artificial “Hemiptera” (all sucking insects)
and “Neuroptera” of Linnaeus, and the orthopteroids, which were focused on first by De Geer,
Olivier, Latreille, Leach, and Burmeister. Difficulties remained, such as the Corrodentia of ear-
lier authors, which for Brauer consisted of termites, bark lice, and true lice. Brauer also seems
to have been the first to segregate what we today recognize as the Mecoptera into a distinct or-
der, the Panorpatae. This system was supported, although not necessarily agreed upon, by John
Henry Comstock (1849–1931) in his influential textbooks on entomology. Although bristling

at the aforementioned composition of Corrodentia, Comstock (26) accepted the arrangement

and even further united them in a retrograde system with mayflies, dragonflies and damselflies,
and stoneflies as the Pseudoneuroptera, an antique concept put forth by Erichson (39). Com-
Paraphyletic:
stock did, however, over the years recognize the artificiality of the grouping, eventually rec- describes a group of
ognizing the Isoptera, Ephemeroptera (as Ephemerida), Odonata, Plecoptera, Mallophaga, and organisms including its
Corrodentia (as effectively equivalent to Psocoptera) as distinct orders. Comstock also recognized members’ last
in time the validity of a separate distinction for the scorpionflies, recognizing an order Mecoptera common ancestor but
not all descendants of
(also spelled during the day as Mecaptera), as had Packard and Hyatt & Arms at about the same
the latter; the term was
time. Indeed, Alpheus Spring Packard (1839–1905) was extremely active in the years prior to not used in Hennig’s
and during Brauer’s era. Perhaps his most influential effort was his Guide to the Study of Insects early theoretical
(108) and its subsequent editions. Therein, as well as in various journal reports, he set forth a writings
hierarchical, evolutionary classification complete with superorders, orders, and suborders. He
rightly recognized Eurhynchota, which is equivalent to our modern Paraneoptera, and although
he retained them in a superordinal grouping named Phyloptera, Packard made prescient steps
toward the eventual demise of Linnaeus’s “Neuroptera,” recognizing Orthoptera, Dermaptera (as
Dermatoptera), Neuroptera (for Trichoptera and Planipennia), and Pseudoneuroptera (for
Odonata, Ephemeroptera, and Platyptera) (108–110), the last with suborders for the divisions
later elevated to orders in their own right. Each of the authors of this last quarter of the nine-
teenth century worked to unite diverse character systems, debated different means of distinguishing
homology from analogy, and considered how best to weight conflicting traits. Insect systematics
was becoming more holistic in its sources of evidence, a development that would be brought to
greater fruition in the early twentieth century.
The second half of the nineteenth century saw the rise in influence of paleontological evi-
dence on understanding insect interrelationships and on the classification of the higher categories.
Paleoentomology had been practiced for many years, but perhaps the most influential fossil seek-
ers were Oswald Heer (1809–1883) and Samuel H. Scudder (1837–1911), both of whom wrote
extensively on the subject, the latter producing one of the first world catalogs of fossil insects
(124). Neither established explicit phylogenies of the insects, but their more general works were
extensively imbued with fossil evidence and stimulated their contemporaries and followers (45,
129). Interestingly, despite his particular interest in Paleozoic insects and the obvious challenge of
placing them among the extant taxa, Scudder resisted establishing wholly new orders, preferring
instead to establish large, paraphyletic suborders or other categories, attempting to capture this di-
versity among the traditional system of his day. Instead, it was Friedrich Goldenberg (1798–1881)
and Charles Brongniart (1859–1899) who first broke that barrier, proposing groups such as the
now familiar Palaeodictyoptera, Megasecoptera, and Protodonata (23), although none contended
that there might have been lineages that did not leave any modern counterparts. Instead, these
groups were regarded as primordial progenitors of our living orders. The said authors influenced
their successors with their perception of the importance and the means of classifying fossil insects.
It was Anton Handlirsch (1865–1935), director of the Natural History Museum in Vienna, who
made the next major leap in the field, and he was certainly not shy about establishing extinct
orders or other categories of fossil lineages (23, 129). Handlirsch (47) brought the entire subject
into focus and made a critical leap toward fully integrating paleoentomology and neoentomology
in his three-volume monolith Die Fossilen Insekten. Handlirsch not only produced the then most
comprehensive account of the world’s diversity of fossil insects, but also founded a grand scheme
of insect evolutionary history in which paleontological evidence was paramount. Elements of
Handlirsch’s classification still linger with us today, although some of his ideas were certainly on
the fringes, including, for example, the amazing exhortation that Pterygota arose from a trilobite
ancestor (47).

This began a heyday for paleoentomological research, as well as its integration with the
study of living insect diversity. Handlirsch certainly kicked off this research program, but it was
the efforts of Andrey V. Martynov (1879–1938), Robin J. Tillyard (1881–1937), and Frank M.
Carpenter (1902–1994) that sustained it. All three were avid entomologists in their own right and
worked on their own favorite groups of living insects (interestingly, all three focused mostly on
the Mecoptera, Neuropterida, and Trichoptera). This keen interest in living insects made them
adept at interpreting insect fossils. While Martynov worked on the extensive Paleozoic and Meso-
zoic deposits of the Soviet Union, Tillyard developed his own ideas based on the abundance of
fossils from Australia and the United States, particularly the extensive Early Permian deposits
of Kansas (45). Indeed, it was Tillyard’s series of monographs on the insect fossils from Elmo,
Kansas, that helped further galvanize Carpenter’s already abiding passion for paleoentomology; in
fact, Carpenter would spend the majority of his 70-year career working on fossils from Kansas and
Oklahoma. Both Tillyard and Martynov proposed explicit classifications of the insects inspired by
their paleontological work, even recognizing additional extinct orders for taxa that did not easily
fit within the living lineages. Whereas some of Martynov’s (94) groupings have stood the test of
time, those of Tillyard were less fortunate. Indeed, Tillyard’s (131) ideas about the phylogeny
of insects and the affinities of particular fossils, such as Protohymenoptera (actually a group of
Megasecoptera), fueled considerable debate even in his own day (21). Unfortunately, both Tillyard
and Martynov died in their late fifties, curtailing their shining careers in entomology and their
championing of paleontological data.
While Carpenter did much to highlight the amazing diversity of insects in the geological past,
particularly those from the Paleozoic, he was remarkably reticent to create an explicit classifica-
tion based on such lines of evidence. Early in his career, he collaborated on the second edition of
Brues & Melander’s Classification of Insects, which was largely a key to families and less an exhorta-
tion of characters supporting a particular arrangement of orders or superordinal categories (17).
Carpenter (22) resurrected Brongniart’s discoveries of giant Paleozoic insects with his discovery
of even larger griffenflies (Protodonata) in North America, and highlighted the importance of
characters from such long-lost faunas. Nonetheless, he was a lumper, preferring to unite the or-
dinal groups of Handlirsch, Tillyard, Martynov, and other authors rather than use such evidence
to split the fossil lineages into their own categories. Carpenter preferred single, paraphyletic units
from which the still-extant orders derived and used fossil evidence to tweak individual elements of
the classification. Carpenter’s influence on insect classification was tremendous and wide ranging,
but he articulated only a single, all-encompassing, revised higher classification in his catalog of
fossil insects for the Treatise on Invertebrate Paleontology (23).
Although not working with insect fossils directly, Guy Chester Crampton (1881–1951) was
an eager advocate of incorporating a paleontological perspective into his intricate morphological
studies (28–30), which highlighted, for example, the phylogenetic importance of Grylloblatta, for
which he established the first ordinal name, Notoptera (27). Unfortunately, he was prone to
establishing numerous names for his hypothetical ancestors to particular lineages. Crampton’s
systems changed over the years, eventually appearing remarkably modern (Figure 1d ).
The influence of paleontology on insect phylogeny and classification was not uncontroversial.
For example, Carl J.B. Börner (1880–1953) did not hide his distrust of Handlirsch’s heavy re-
liance on fragmentary fossils. Börner was a careful anatomist, and it was he who recognized the
Zygentoma (silverfish) as distinct from the Archaeognatha (bristletails), noting (Figure 1c) that
the former were more closely allied to the Pterygota (12), a grouping Willi Hennig would later
dub the Dicondylia. Using similar (mouthpart) characters, he was the first to consider the Odonata
and Neoptera as composing a group, the Metapterygota (13). For Börner the fossil evidence was
simply too incomplete, relying heavily on wing venation and a meager record (at the time) of body

remains, as well as too few specimens for individual lineages, thereby hindering a comprehensive
perspective on a particular extinct group. He did not wholly abandon evidence from the geological
past, considering the Dictyoneurida of Handlirsch to be the progenitor of the Pterygota, but was
unwilling to give paleontology its due beyond this. At about the same time, Richard Heymons
(1867–1943) elucidated the finer aspects of insect development, discarding the coarse groupings
of Ametabola and Paurometabola and instead providing an arrangement of the orders that empha-
sized details of their ontogeny (55). His system split the insects into Epimorpha and Metamorpha,
with the former including Epimorpha typica (apterygotes, Orthoptera, Dermaptera, Isoptera,
Psocodea, Heteroptera), Hyperepimorpha (Homoptera in part), and Hemimetabola (Homoptera
in part, Odonata, Plecoptera), and the latter including Prometabola (Ephemeroptera) and the
Holometabola.
This early-twentieth century period in which the aforementioned authors worked was a re-
markable time. Amazingly, within a single decade, four extant higher-rank taxa were described
that were immediately or later recognized as having a pivotal significance in insect phylogeny:
Protura (127), Zoraptera (128), Grylloblattodea (134), and nannochoristid scorpionflies (130). It
was also during this same productive period that the fundamental structure of insect wings was
once again considered, and almost simultaneously three authors recognized the same division and
arrangement of orders. Auguste M. Lameere (1864–1942) (85), Crampton (28), and Martynov
(93) arrived at the same arrangement, although Martynov’s account is more detailed and the
names he accorded these divisions are the ones that gained almost universal acceptance—namely,
Paleoptera and Neoptera. This division, and the alternative arrangements of the basal winged
lineages (Ephemeroptera, Odonatoptera, Palaeodictyopterida, Neoptera), has remained one of
the lasting debates in insect phylogenetics, one in which Boris N. Schwanwitsch (1889–1957)
first emphasized the arrangement of the flight musculature and reversed Börner’s position, with
Ephemeroptera as closest to Neoptera (122). Despite the overwhelming support for Pterygota,
Lemche (87) argued for the diphyletic origin of wings, dividing the insects into the Plagioptera
(Palaeodictyopterida and Odonata) and Opisthoptera (Ephemeroptera and Neoptera) with sep-
arate origins. This position was resurrected and expounded by Matsuda (95) and La Greca (80)
but has not garnered further support. Although hypotheses indicating that the closest relatives
of hexapods were members of the myriapod assemblage had been predominant since the nine-
teenth century, suggestions of a closer relationship to crustaceans had been made intermittently,
and in 1926, neuroanatomical work led Hanström (48) to suggest that hexapods were subordinate
within the Crustacea, thereby foreshadowing much later developments. Although numerous other
contributions and authors debated the relationships of the orders and their implications for clas-
sification, it was not until the middle of the twentieth century that the next turning point would
be reached.
WILLI HENNIG, THE CLADISTIC REVOLUTION,

AND SUBSEQUENT DEVELOPMENTS
By the mid-twentieth century, the state-of-the-art accounts of higher insect classification were
arguably those presented in René G. Jeannel’s (1879–1965) (67) chapter on the subject in the
Traité de Zoologie and an article by Crampton (30) summarizing and revising his extensive previous
work. These works are largely congruent with each other (though names applied to supraordinal
groups differ), and the Jeannel chapter, largely reflecting Martynov’s views as summarized in a 1938
memoir (94) in Russian, was likely instrumental in making Martynov’s conclusions accessible to a
Western readership. It is noteworthy that in none of these works are the nonendopterygote winged
insects grouped together, and indeed Crampton (30) explicitly stated that “the usual division of

PHYLOGENETIC SYSTEMATICS AND CLADISTICS
The systematizing of organisms by synapomorphy alone, hence allowing only taxa that are monophyletic in the
strict sense (having “a history of their own”), was referred to by Hennig as phylogenetic systematics. Because such
systematizing is based exclusively on the branching pattern in the tree of life, it is also called cladistics (from Greek
klados, “branch”). Some contemporary systematists use the term cladistics in a restricted sense, e.g., for phylogenies
inferred from parsimony-based rather than so-called model-based methods (maximum likelihood, Bayesian analysis)
of character analysis. Arguably, cladistics remains an appropriate term for all approaches to systematizing organisms
according to the relative recency of common ancestry rather than to some kind of similarity assessment.
winged insects into Exopterygota and Endopterygota [concepts introduced by David Sharp (1840–
1922) before the turn of the century (126)], on the basis of the external or internal development
of the wings, is utterly meaningless from the standpoint of phylogeny” (text in brackets author’s
own). In contrast, this division continued to be emphasized in some academia, probably due
largely to its adoption in August D. Imms’s (1881–1949) globally influential General Textbook of
Entomology (including its 1957 edition revised by Richards & Davies, 63). More idiosyncratically,
Hermann Robert Weber (1899–1956) had continued Handlirsch’s advocacy of endopterygote
nonmonophyly, arguing that beetles, hymenopterans, and the remaining endopterygotes (Weber’s
Neuropteroidea) were independently derived from nonendopterygote ancestors (135, 136).
The core of what has been called the Hennigian, or cladistic, revolution in biological sys-
tematics is the restriction of the monophyly concept to taxa that include all descendants of their
members’ last common ancestor and the exclusion of all taxa that are nonmonophyletic in this
sense from the formal classification (see sidebar Phylogenetic Systematics and Cladistics). Al-
though Emil Hans Willi Hennig’s (1913–1976) book Grundzüge einer Theorie der phylogenetischen
Systematik is often believed to be the starting point of this revolution (49), his innovative views and
terminology were presented more clearly and more influentially in a major 1953 article (50) on in-
sect systematics, and his ideas continued to develop during the following years (118, 121). Formal
changes to insect classification were quite limited in Hennig’s overall cautious 1953 article, but
notable innovations were indeed made, including discarding such nonmonophyletic taxa as Aptery-
gota and Thysanura (in the sense of Archaeognatha + Zygentoma) and creating new names for
the putative monophyla Dicondylia (=Zygentoma + Pterygota) and Psocodea (=‘Psocoptera’ +
‘Mallophaga’ + Anoplura). Moreover, Hennig here tentatively preferred Börner’s (13) Metaptery-
gota (monophyletic Odonata + Neoptera) solution for the basal splitting event identifiable among
extant pterygotes, and he emphasized the spurious nature of the evidence invoked by Weber (135)
for endopterygote nonmonophyly. The monophyly of the Parametabola (=Paraneoptera), of the
Saltatoria (=Orthoptera), and of Mantodea + Blattodea + Isoptera was accepted, but the mono-
phyly of Paurometabola (=Polyneoptera) was not, and relationships within this assemblage were
left open. So too were the relationships between Coleoptera, Hymenoptera, Neuropteria, and
Mecopteria, while the Strepsiptera and Siphonaptera remained unassociated with any other en-
dopterygotes. A monophyletic ‘Eumetabola’ comprising parametabolans and endopterygotes was
tentatively accepted.
The impact of cladistic reasoning on insect phylogeny was modest in the years immediately fol-
lowing Hennig’s 1953 article (52). A contribution on insect phylogeny in Volume 3 of the Annual
Review of Entomology had been commissioned from the entomological polyhistorian Howard E.
Hinton (1913–1977), but his review (57) was devoted almost entirely to the Mecopterida (“panor-
poid orders”), on which his empirical work was at that time focused. Other “significant advances

made during the past two decades or so to our understanding of the phylogeny of the Insecta and
other classes of the subphylum Antennata” were summarized in a series of succinct statements:
Sister group: a taxon
(a) the polyphyletic nature of the old groups Myriapoda and Hexapoda; (b) the extreme isolation of A that shares with
the class Collembola and their lack of any close resemblance to the Insecta; (c) the close relationship taxon B a common
that exists between the Symphyla, Entotrophi (=Diplura), and Insecta . . . ; (d ) the Protura as a class ancestor not inferred
distinct from the Insecta; (e) the very great differences that exist between the Ephemeroptera and to be ancestral to other
known taxa
other pterygote insects, including the Odonata; ( f ) the isolation of the Dictyoptera (roaches, mantises)
from the old order Orthoptera and the relation of the Isoptera and Zoraptera with the Dictyoptera;
( g) the monophyletic origin of the Endopterygota; and (h) the close relation between the Megaloptera-
Neuroptera and the Coleoptera-Strepsiptera.
Several of these unreferenced statements were indeed in accordance with then widespread views,
but others (a, b, f: Zoraptera affinities, h) were not. While Hinton in principle endorsed Hennig’s
methodological approach, his reiterated advocacy for according micropterigid moths status as a
separate order (Zeugloptera) overlooked Hennig’s explicit statements about the proposal [made
first by Chapman in 1917 (25) and subsequently by Hinton in 1946 (56)] being incompatible
with phylogenetic-systematic reasoning. Hinton’s simultaneous elevation of boreid mecopterans
to full ordinal rank (Neomecoptera) also violated such reasoning, as it was based exclusively on
seeming autapomorphies of Boreidae and hence left out a nonmonophyletic Mecoptera. That
some subsequent findings (138) have supported the boreids’ closest relatives to be taxa outside the
Mecoptera sensu stricto is another matter.
Unsurprisingly, Hennigian cladistics first caught on in parts of continental Europe where
German is the native language or was (then) easily read by most scholars, and where the approach
was communicated to students in introductory university texts (51, 73). Tuxen (133) applied Hen-
nigian concepts to his 1959 advocacy of the monophyly of entognathan hexapods, but the center
for pioneering empirical contributions to insect classification was the University of Tübingen,
where Weber held the zoology chair from 1951 to his death in 1956. While personally and po-
litically Weber was controversial, it is unquestionable that he and his students from the 1930s
onwards had produced some of the finest-ever descriptive work in insect anatomy, but he had
reluctance, if not downright dislike, of drawing phylogenetic conclusions from the emerging find-
ings (9, 136). However, inspired by guest lectures at Tübingen from Hennig in the 1960s, one of
the last “Weber school” members, Gerhard Mickoleit, directed his own work, and subsequently
that of numerous research students, toward topics in insect anatomy that appeared promising for
clarifying phylogenetic issues. Early noteworthy examples are Mickoleit’s (99–101) own studies
on the pterothorax, supporting the monophyly of Mecopterida and Diptera + Mecoptera, and on
endopterygote ovipositors and their derivatives, in which for the first time substantial evidence was
provided in support of the overall generalized Neuropterida (=Raphidioptera + Megaloptera +
Neuroptera/Planipennia) as a monophylum, and also for the sister-group relationship between
Neuropterida and the Coleoptera. Moreover, there was exemplary Weber school work by Achtelig
on Neuropterida (1, see additional references in 45), in which initially putative evidence for Mega-
loptera paraphyly in terms of Raphidioptera was presented (a view Achtelig later questioned, 2),
and by Rähle (112), who provided unexpected evidence for a sister-group relationship between
the Embiodea and Phasmatodea, a finding later supported in some molecular analyses.
Hennig himself continuously worked on revising/expanding his account of insect phylogeny,
and in 1969 presented the work in book form: Die Stammesgeschichte der Insekten (53, 54; the
posthumously published English translation was updated and annotated by a group of special-
ists). Still with many reservations, he proposed a somewhat more resolved order-level phylogeny

in this book than in his 1953 article, including a monophyletic Ellipura, Palaeoptera, and Pau-
rometabola (=Polyneoptera excluding Plecoptera). A full resolution of the Paurometabola was also
suggested: (Embioptera + ((Notoptera + (Dermaptera + (Mantodea + (Blattariae + Isoptera)))) +
Clade: shorthand
term for groups that (Ensifera + (Caelifera + Phasmatodea)))). The suggested paraphyly of Orthoptera in terms of
are monophyletic in Phasmatodea had been suggested earlier by Sharov (125). The Zoraptera were suggested to be
the Hennigian sense the sister group to the Paraneoptera (=Psocodea + (Thysanoptera + Hemiptera)), while the
Holometabola/Endopterygota still had much the same modestly resolved arrangement as in 1953.
Hennig’s book prompted a review by Kristensen (74), in which a substantial amount of available
literature not considered by Hennig and some original observations were taken into account.
The result was somewhat retrograde in the sense that the evidence bearing on most of Hennig’s
newly recognized clades was shown to be more ambiguous/contradictory than evident from his
text. A monophyletic Palaeoptera was considered less probable than a monophyletic Odonata +
Neoptera, monophyly of Orthoptera was tentatively reinstated, but interrelationships within the
Plecoptera + ‘Paurometabola’ assemblage were otherwise considered unresolved [apart from the
long-accepted monophyly of Mantodea + (Blattodea + Isoptera)]. Kristensen (74, 75) found the
placement of the Strepsiptera to be a particular problem and even found the available evidence
for their assignment to Endopterygota questionable.
Soon after, the subject was treated in yet another book. H. Bruce Boudreaux’s (14) Arthropod
Phylogeny with Special Reference to Insects provided again a much (indeed fully!) resolved hexa-
pod tree. Details differed from those in Hennig’s proposals: There were suggested sister-group
relationships between Ephemeroptera and Neoptera, between Plecoptera and Embioptera, and
between Grylloblattodea and a clade comprising Zoraptera and (Isoptera + (Blattodea + Man-
todea)). Within Holometabola, the Neuropterida were the sister group to the Mecopterida rather
than to a Coleoptera + Strepsiptera clade. None of the evidence presented for these changes was
considered compelling in subsequent reviews by Kristensen (75–77).
New discoveries proved significant in the context of higher classification. The zygentoman
Tricholepidion described in 1961 from California was immediately recognized to have some re-
markably primitive characters (35, 143), but Boudreaux’s observation that it retained part of the
ligamentous head endoskeleton otherwise lost in Dicondylia proved particularly challenging. So
did the discovery of the aquatic larva of the small circum-Antarctic scorpionfly genus Nanno-
chorista, in conjunction with findings on their genital segments in both sexes (102, 103, 111).
These taxa were singled out in 1989 (76) as probably representing separate orders (a status ac-
tually earlier conferred upon Nannochorista by Hinton; see sidebar Nannomecoptera); however,
although considerable descriptive and analytical work has subsequently been devoted to them (61,
96, and references therein), the questions about the sister group of each (i.e., Tricholepidion as
related to other Zygentoma or all other Dicondylia; Nannochorista as related to which other sub-
group[s] in the Mecoptera + Diptera + Siphonaptera assemblage) arguably remain inconclusively
answered.
NANNOMECOPTERA
Howard Hinton had examined Nannochorista larvae some years before they were first illustrated (117) and described
(111), and in 1967 he was convinced (personal communication to N.P. Kristensen) that the taxon has “nothing to do
with Mecoptera,” a finding he intended to publish. However, the formal establishment of the order Nannomecoptera
for the genus only appeared in his posthumously published monograph on insect eggs (58), a most unlikely place
for a taxonomic action of that kind. Unsurprisingly, it was overlooked for many years.

Insect palaeontology was actively pursued in the second half of the twentieth century by sev-
eral workers, not least in Russia, where work by Aleksandr Grigorevich Sharov (1922–1973) and
Boris Borisovich Rohdendorf (1904–1977) was of importance for Hennig’s endeavor of establish-
ing times of origin of recognized high-rank clades (45). Jarmila Kukalová-Peck’s (78) extensive
paleoentomological studies led primarily to noteworthy (but controversial; 8) interpretations of
ancestral insect morphology (78). She integrated her paleontological work with observations on
extant taxa to advocate monophyly of Palaeoptera and eventually to establish a neopteran classifi-
cation with a monophyletic Orthoneoptera (=Plecoptera + Embioptera + Orthoptera) and Blat-
toneoptera (=Dermaptera + Grylloblattodea + Dictyoptera s.l.) as sister to the Paraneoptera +
Endopterygota (79). An upsurge in palaeontomology was prompted by discoveries of rich new
Lagerstätte (from the Cretaceous in particular) during the last decades of the twentieth century;
unfortunately, Carpenter & Burnham’s (24) review of the geological history of insects appeared as
this resurgence was still in its infancy and was understandably lacking. Spectacular new outcrops
provided an abundance of material, while the plethora of insect-bearing amber deposits, partic-
ularly those from the Cretaceous, that accumulated during the 1990s (and continue to accrue
to this day) provided volumes of taxa with enigmatic character combinations, with extralimi-
tal distributions, and with a fidelity of preservation unrivaled in the paleontological world (45).
Paleontological discoveries (e.g., 37, 59, 72), though at times controversially interpreted, increas-
ingly contributed to the total picture of insect origins and major evolutionary innovations. When
such taxa were cladistically analyzed alongside their modern counterparts, they achieved their
maximum potential for revising concepts of relationships (45). Also, some “new” concepts of or-
dinal monophyly, such as the paraphyly of Blattaria relative to termites (38), found support from
paleontological investigations.
EXPANDING CHARACTER SETS

The reexamination of the modest body of evidence drawn upon in attempts to classify high-rank
insect lineages in the 1950s and 1960s highlighted its conflicting nature, and hence the need for
procuring a much more extensive set of comparative data. This was particularly evident in the case
of the assemblage of lower Neoptera (i.e., Martynov’s Polyneoptera), which by that time appeared
(and still remains) the most difficult sector in the insect tree of life. Two attempts to systematize
polyneopterous groups using somewhat extensive sets of characters in integumental anatomy and
the then new analytical methods of numerical phenetics were made by Giles (44) and Kamp (68).
Whereas the results of such phenetic analyses had little relevance to phylogenetic systematists,
Blacklith & Blacklith’s (11) attempt to use a cladistic approach (the early Camin-Sokal method,
20) was more promising; its informativeness, however, was unfortunately seriously compromised
by the omission of Plecoptera and Embiodea from the analysis. The technical breakthrough in
electron microscopy (EM) (transmission EM in the mid-1950s, scanning EM a decade later)
added greatly to the precision and ease of morphological observation and documentation, but
few characters were thereby added that proved informative for higher-rank insect classification;
spermatozoa are the most notable exception (66).
Computer-aided approaches to phylogeny reconstruction, enabling analyses of sizable char-
acter sets, matured during the 1980s, and serious use of molecular characters in classification
of high-rank insect taxa developed toward the end of the decade, initially drawing almost en-
tirely on ribosomal (r)DNA. Molecular support for an unexpected Strepsiptera + Diptera (the
so-called Halteria) clade led to a much-publicized proposal that the former group evolved from
a dipteran-like ancestor through a homoeotic transformation interchanging the two pterotho-
racic segments (140), and the first “total evidence” study of endopterygote phylogeny (i.e., one

drawing on a somewhat sizable morphological data set plus rDNA data) was aimed primarily
at elucidating “the Strepsiptera problem” (139); here the Halteria again emerged. Although the
Halteria appeared novel, it was actually Lamarck (84) who first classified the Strepsiptera with the
Diptera, highlighting that there is truly nothing new under the sun. The Strepsiptera problem
remained for a while a battlefield between proponents of different approaches (parsimony versus
model-based) to the analysis of molecular data (60), but recent evidence from both morphology
and molecules has brought the Strepsiptera back full circle to being the sister group to Coleoptera
(42, 89).
A previously unimagined degree of arthropod polyphyly was advocated in Sidnie M. Manton’s
(1902–1979) extensive writings (92) focused on the functional anatomy of feeding and locomo-
tory structures. With respect to hexapods, a single origin of entognathy was disputed, and the
entognathan orders as well as the Thysanura and Pterygota were claimed to have been derived
independently from a multilegged ancestor whose limbs were nonarticulated (lobopodial), though
it was allowed that the last two could have shared “a very early common stage.” An earlier multi-
legged lobopodial ancestor was suggested to be shared (among extant taxa) only by the hexapod
and myriapod lineages and the Onychophora, and the three were collectively named the Uni-
ramia. Mantonian views had for some decades a considerable impact on parts of the international
(particularly Anglophone) zoological community, but they were never compatible with cladistic
reasoning, and their influence dwindled as they proved untenable in light of subsequent de-
velopments. Meanwhile, members of the outstanding Dijon school of morphology (the French
Weber school counterpart was founded by J. Robert Denis, 1893–1969) had procured an unsur-
passed body of factual information, particularly about the primarily apterous hexapods. Whereas
the Dijon workers, like their German colleagues, focused initially on problems in morphology
per se rather than on phylogeny, cladistic analyses of the basal hexapod lineages were eventu-
ally published by Bitsch & Bitsch (10). However, despite the substantial nature of the analyzed
morphological data sets, robust answers to the principal questions (i.e., on the monophyly of
Ellipura, on the closest relations of Diplura, on the position of Tricholepidion) did not emerge.
Only myriapod outgroups were considered in these analyses, but by the 1990s the hypothesis of
Crustacea + Hexapoda monophyly had become revived as a serious competitor to Hexapoda +
Myriapoda monophyly, primarily on the basis of molecular and neuroanatomical data (see
Reference 34 for a summary), and during the following decade it arguably outcompeted the
latter.
Developments since the turn of the millennium barely qualify as history (see sidebar Formal As-
pects of Naming). Suffice it to say that the continued and accelerated acquisition of large data sets,
particularly the phylogenomics revolution currently underway, is generating volumes of informa-
tion at a scale never before dreamt. Similarly, morphological studies are more imperative than ever
(45), and in some situations, such as with fossils or unique specimens, they can benefit markedly
from novel approaches such as synchrotron and X-ray computerized tomography (36, 81). A re-
cent review summarizes developments from both molecular and morphological approaches (132).
Equally significant, phylogenetically important taxa continue to be discovered, both as fossils and
as living species. The discovery of living and fossil mantophasmatodeans, likely the closest extant
relatives of the Grylloblattodea, highlights that taxa challenging even our higher-level concepts of
relationships may be found (5, 70). Few of the approximately 10,000 new insect species described
annually (65) will transform our notions of entire orders, but each species does inform us of its
unique combination of characters and biology, refining our knowledge of their close relatives
and thereby generating a cascade of refinements to monophyletic genera, families, superfamilies,
suborders, orders, and beyond. This century will be as dynamic in terms of higher entomological
classification as have been those preceding our current era.

FORMAL ASPECTS OF NAMING
The International Committee on Zoological Nomenclature (ICZN) does not regulate names above the level of
the family group (64). Johann Nepomuk Edler von Laicharting (1754–1797), however, typified the ordinal names
(82). Laicharting established a classification largely congruent with that of Fabricius but employed names based
on the included genera, nonetheless clearly proposing them as orders (“ordo”). This resulted in order names
such as Scaraboides (=Coleoptera L., Eleutherata Fabr.), Grylloides (=Ulonata Fabr.), Cimicoides (=Hemiptera
L., Rhyngota Fabr.), Papilionoides (=Lepidoptera L., Glossata Fabr.), Libelluloides (=Neuroptera L., Synistata
Fabr.), Vespoides (=Hymenoptera L., Piezata Fabr.), and Muscoides (=Diptera L., Antliata Fabr.). A Russian school,
which emphasizes the typification of higher names in the same manner as family-group names under the current
ICZN (113, 119), therefore uses as their starting point Laicharting’s monograph and even considers these names
as coordinate with existing familial names. Given the essentially universal usage of the ordinal names, particularly
those that have been in common parlance since the work of Linnaeus, the adoption of effectively unknown typified
names serves no good purpose. Arguably, the same applies to the more recently developed circumscription-based
nomenclature (71).
CONCLUSIONS
The history of systematics, as in all histories, is not just the story of grand discoveries or paradigm
shifts. It is the sum of all of its actors, toiling to document observations from the natural world,
steadily accruing information that is pieced together into enduring patterns, patterns which them-
selves lead us to explanations of greater or lesser degrees of generalization (46). Without the actions
of each and every systematist working passionately on their chosen lineage, the entire effort would
grind to a standstill. No contribution is too small, particularly in entomology where the numbers of
researchers remain meager. However, history also highlights to us the numerous false starts made
as part of the self-correcting body of knowledge we know as Science. These are natural and we are
all a part of this, with today’s hubris and certainty becoming tomorrow’s quaint anachronism or
outright failure. Although some of the assertions and concepts expounded by authors mentioned
herein seem quaint or bizarre (as may some of our own ideas some 100 years from now!), they
were all legitimate hypotheses and served to stimulate the standing intellectual dialogue. Science
needs its variation, even its future failures, just as much as it relies upon the solid descriptions that
underlie, sometimes secretly and unrecognizably, its most powerful theories. As we hope readers
will recognize, the history of entomological classification is a microcosm of such realities. From
the classification we can communicate about many descriptive patterns and explanatory processes
and build predictions, while it itself continues to evolve as new discoveries, new methods, and
new ideas (many certainly doomed to fall away in time) render our applied names all the more
meaningful.
SUMMARY POINTS
1. The history of science is not a steady progression. It has its transformative moments as
well as numerous false starts and reversals. Its entirety shapes the way we think today, and
the present is not an end product but instead merely a chapter in a continuing narrative.

2. Pre-Darwinian authors were astute and keen observers of the natural world, recognizing
enduring patterns of characters and, more often than not, correctly interpreting and
classifying the species before them. All this work was done with quaint tools and with
the tiniest fraction of material compared with what we have today.
3. The Darwinian paradigm shift transformed how authors interpreted their observed and
described patterns of characters, but it did not substantially augment how they actually
practiced systematics. Classifications were changing wildly, but this was not due to a
tectonic shift in the day-to-day practice of the science. Evolution was a conceptual and
explanatory change long before it became methodological.
4. With the rise of cladistic reasoning and numerical practices, systematics experienced
its greatest methodological shift. Emphasis on monophyly and more precise estimates
of genealogical relationship gave classifications far more predictive power than they
ever had before. Simultaneously, the diversity and magnitude of data sets have grown
exponentially, enlivening the field and strengthening the overarching classification of
insects.
DISCLOSURE STATEMENT
The authors are not aware of any affiliations, memberships, funding, or financial holdings that
might be perceived as affecting the objectivity of this review.
ACKNOWLEDGMENTS
We apologize to those entomological systematists whose efforts over the centuries have inspired
and guided us all, but for whom we were unable to devote space to the memory of their ac-
complishments. We are grateful to I.A. Hinojosa-Dı́az (University of Kansas) for assistance with
images and to Ted R. Schultz for inviting us to provide this overview, stimulating its production
and providing constructive criticism.
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Annu. Rev. Entomol. 2013. 58:585–607 Keywords

This article’s doi: Abstract

cursory treatment to the current era of entomological classiﬁcation, as a compromise had to be

THE PRE-LINNEAN PERIOD

586 Engel · Kristensen

THE FIRST CENTURY (1758–1859)

www.annualreviews.org • A History of Entomological Classification 587

588 Engel · Kristensen

www.annualreviews.org • A History of Entomological Classification 589

590 Engel · Kristensen

THE DARWINIAN ERA: HAECKEL TO HENNIG

www.annualreviews.org • A History of Entomological Classification 591

592 Engel · Kristensen

at the aforementioned composition of Corrodentia, Comstock (26) accepted the arrangement

www.annualreviews.org • A History of Entomological Classification 593

594 Engel · Kristensen

WILLI HENNIG, THE CLADISTIC REVOLUTION,

www.annualreviews.org • A History of Entomological Classification 595

PHYLOGENETIC SYSTEMATICS AND CLADISTICS

596 Engel · Kristensen

www.annualreviews.org • A History of Entomological Classification 597

598 Engel · Kristensen

EXPANDING CHARACTER SETS

www.annualreviews.org • A History of Entomological Classification 599

600 Engel · Kristensen

FORMAL ASPECTS OF NAMING

www.annualreviews.org • A History of Entomological Classification 601

602 Engel · Kristensen

www.annualreviews.org • A History of Entomological Classification 603

604 Engel · Kristensen

73. Kristensen NP. 1970. Systematisk Entomologi. Copenhagen: Munksgaard

www.annualreviews.org • A History of Entomological Classification 605

606 Engel · Kristensen

www.annualreviews.org • A History of Entomological Classification 607

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