Annals of the Entomological Society of America, XX(X), 2020, 1–20

doi: 10.1093/aesa/saaa042
Review Review

Are Insects Heading Toward Their First Mass Extinction?

Distinguishing Turnover From Crises in Their Fossil Record

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Sandra R. Schachat1,5, and Conrad C. Labandeira2,3,4
Department of Geological Sciences, Stanford University, Stanford, CA 94305, 2Department of Paleobiology, Smithsonian Institution,
1

Washington, DC 20013, 3Department of Entomology, University of Maryland, College Park, MD 20742, 4College of Life Sciences,
Capital Normal University, Beijing 100048, People’s Republic of China, and 5Corresponding author, email: schachat@stanford.edu

Subject Editor: Lee Dyer

Received 15 July 2020; Editorial decision 2 October 2020

Abstract
Time and again, over hundreds of millions of years, environmental disturbances have caused mass extinctions
of animals ranging from reptiles to corals. The anthropogenic loss of species diversity happening now is often
discussed as the ‘sixth mass extinction’ in light of the ‘Big Five’ mass extinctions in the fossil record. But in-
sects, whose taxonomic diversity now appears to be threatened by human activity, have a unique extinction
history. Prehistoric losses of insect diversity at the levels of order and family appear to have been driven by
competition among insect lineages, with biotic replacement ensuring minimal net losses in taxonomic diver-
sity. The end-Permian extinction, the ‘mother of mass extinctions’ in the seas, was more of a faunal turnover
than a mass extinction for insects. Insects’ current biotic crisis has been measured in terms of the loss of abun-
dance and biomass (rather than the loss of species, genera, or families) and these are essentially impossible
to measure in the fossil record. However, should the ongoing loss of insect abundance and biomass cause the
demise of many insect families, the current extinction event may well be the first sudden loss of higher-level in-
sect diversity in our planet’s history. This is not insects’ sixth mass extinction—in fact, it may become their first.

Key words:  biodiversity, Cretaceous, paleontology, paleoentomology, Permian

Insects have been largely neglected in studies of the current extinc-
tion crisis. Studies of extinction risk are overwhelmingly biased
in favor of vertebrates, particularly birds and mammals (Ceballos
et al. 2017). The current extinction crisis is often referred to as ‘the
sixth extinction’ (Barnosky et al. 2011), a reference to the ‘Big Five’
mass extinctions that have been identified in animals’ fossil record:
the end-Ordovician event at 444 million years ago (Ma), the Late
Devonian event around 372 Ma, the end-Permian event at 252 Ma,
the end-Triassic event at 201 Ma, and the end-Cretaceous event at
66 Ma (Jablonski 1994, Raup 1994). Because the fossil record of
animals is biased in favor of marine invertebrates with hard body
parts, such as brachiopods, bivalves, and gastropods (Kidwell and
Flessa 1996), studies that compare the current extinction crisis with
the Big Five mass extinctions tend to focus on shelly marine inver-
tebrates due to their extensive fossil record and on vertebrates due
to the wealth of available data on extant lineages (Barnosky et al.
2011). Insects, which do not live on the ocean floor, do not have
mineralized shells, and are not ‘charismatic megafauna’, do not
lend themselves particularly well to comparisons between the cur-
rent extinction crisis and the Big Five mass extinctions. However,
insects’ tremendous ecological, economic, and aesthetic importance
(Scudder 2017) means that their plight cannot be ignored.
As of July 2020, the Red List of Threatened Species main-
tained by the International Union for Conservation of Nature
and Natural Resources contains evaluations for 91% of described
mammal species, 100% of described bird species, and fewer than
1% of described insect species (IUCN 2020). While various studies
have focused on genera (Salcido et al. 2020), families (Houghton
and Holzenthal 2010), guilds (Lister and Garcia 2018), and com-
munities (van Klink et  al. 2020), studies and media coverage of
insect decline often focus on individual species that are not repre-
sentative of larger clades. The monarch butterfly Danaus plexippus
(Linnaeus, 1758) (Lepidoptera: Nymphalidae) is well studied
(Gustafsson et  al. 2015, Thogmartin et  al. 2017) but, as one of
a handful of butterfly species that migrate across long distances
(Gilbert and Singer 1975), offers few clues to the potential extinc-
tion risks that other butterflies face. The honey bee Apis mellifera
Linnaeus, 1758 (Hymenoptera: Apidae) has also received tremen-
dous attention (Suryanarayanan and Kleinman 2013, Staveley
et  al. 2014) but, as a domesticated eusocial species that is often
transported by humans on a semiannual basis and has been intro-
duced in much of its current range, does not necessarily represent
other bee species. The cave beetle Anophthalmus hitleri Scheibel,
1937 (Coleoptera: Carabidae), named in honor of Adolf Hitler and

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2 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
currently endangered because neo-Nazis pay large sums of money
for poached specimens (Jóźwiak et al. 2015), provides no clues to
the plight of other ground beetle species.
Because extinction history and extinction risk remain unknown
for the vast majority of insect species (Dunn 2005, Sánchez-Bayo
and Wyckhuys 2019, Didham et  al. 2020), changes in abundance
and biomass—rather than taxonomic diversity—are the most access-
ible clues to global patterns of insect decline (Shortall et  al. 2009,
Schuch et al. 2012, van Klink et al. 2020). Recent studies, primarily
spanning temperate latitudes, have found precipitous declines in in-
sect abundance and biomass, suggesting that insects are in the midst
of an anthropogenic extinction crisis (Hallmann et al. 2017, Lister
and Garcia 2018). Because a small fraction of species can dominate
the abundance and biomass in their community, changes in the se-
verity of extinction risk at the species level cannot be extrapolated
from abundance and biomass data.
The sudden increase in concern about insect extinction raises
the question of whether the current crisis can be compared to
events in the fossil record, as has been done for many vertebrates
and for marine invertebrates. Indeed, the fossil record of insects
contains a cataclysmic extinction (of the paleopterous superorder
Palaeodictyopteroidea) during one of the Big Five—a loss reminis-
cent of the extinctions of trilobites, ammonites, and nonavian dino-
saurs (Labandeira 2005). At least one insect guild also experienced
turnover during the most recent of the Big Five mass extinctions,
the end-Cretaceous event (Donovan et  al. 2014, Labandeira et  al.
2016b).
However, three of the five largest extinctions identified in the
fossil record of insects—at the Late Pennsylvanian (approximately
304 Ma), Late Jurassic (approximately 155 Ma), and later Early
Cretaceous (approximately 113 Ma)—are uncorrelated with mass
extinctions in other taxa (Labandeira 2005). The worst extinction
events for clams, sharks, and tetrapods like us are not necessarily
the worst events for insects, and vice-versa. Comparisons of the ex-
tant entomofauna with extinct entomofaunas should be made with
caution, limited to inferences permitted by the data available from
the fossil record.
Strengths of the Insect Fossil Record
The fossil record of insects provides considerably more information
about insects’ evolutionary history than could be gleaned from in-
ferences limited to extant taxa and their relationships. The earliest
verified occurrences of relevant fossil insect taxa are used to time-
calibrate divergences among extant lineages (Kohli et  al. 2016,
Evangelista et al. 2017). When incorporated as terminals, fossils can
resolve conflicts among topologies generated from extant taxa alone
(Engel 2001).
The fossil record sheds light on the timing of behavioral ad-
vances in addition to divergence times. For example, extant plant–
insect interactions often occur between plant and insect lineages
of very different ages, and insects can switch hosts over time (Vea
and Grimaldi 2016). These phenomena raise the question of when
those interactions originated—a question that only fossils can an-
swer (Ren et al. 2009, Winkler et al. 2010, Schachat and Labandeira
2015, Peris et al. 2017). Eusociality provides another example of this
type of insight: termites and ants are by far the two most speciose in-
sect groups that are strictly eusocial, and they diversified around the
same time despite having originated 30–40 Myr apart and belonging
to very distantly related higher taxa (Engel et al. 2016, Barden 2017,
Barden and Engel 2019). Other insect fossils demonstrate the ancient
origins of behaviors ranging from camouflage (Pérez-de la Fuente
et al. 2012; Wang et al. 2012, 2016a) to parental care (Laza 2006,
Cai et al. 2014, Wang et al. 2015a; Gao et al. 2019).
Fossils also shed light on bizarre and fascinating episodes of in-
sect history that have left no trace in the extant fauna. Over 100
million years ago, a number of insect groups—first Paleoptera, then
neopterous Hemimetabola, then Holometabola—reached body sizes
that exceed those of any extant insect (Grimaldi and Engel 2005).
Early representatives of many insect groups, from ants (Barden and
Grimaldi 2012) to Pterygota as a whole (Pecharová et al. 2015), did
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not resemble their extant relatives. Some extinct fossil groups also
exhibit remarkable convergence with very distantly-related extant
taxa (Labandeira et al. 2016c), raising the sorts of questions about
contingency and inevitability that have long fascinated paleontolo-
gists (Sepkoski 2017).
Limitations of the Insect Fossil Record
The fossil record of insects contains fewer genera and species than
one might expect given their abundance and the hundreds of mil-
lions of years that have passed since they originated. Mineralized
body parts, like shells and teeth, are most easily preserved as fossils.
A typical insect exoskeleton is not nearly as thick or as durable as
a clam’s shell or as a vertebrate’s tooth. Although insects have a far
richer fossil record than, for example, earthworms—which lack min-
eralized body parts and do not live in microhabitats that facilitate
fossilization—the fossil record of insects contains larger temporal
and taxonomic gaps than the fossil records of vertebrates and of
shelled, aquatic invertebrates (Benton and Simms 1995, Kidwell and
Flessa 1996).
Barely 46,000 insect specimens have been described from the
entire fossil record (Clapham et  al. 2016), amounting to only 124
individuals per million years since the earliest evidence of winged
insects (Pterygota) (Prokop et  al. 2005, Haug and Haug 2017,
Schachat et al. 2018). For a modern perspective, the 124 described
fossil insects per million years from across the globe falls within the
range of insect species diversity in a typical home in Raleigh, North
Carolina (Bertone et al. 2016). Put another way, the less than 50,000
described insect specimens in the entire fossil record from the whole
planet approximately equals the number of specimens collected by
interception traps in a single study of modern arthropod diversity
(Basset and Arthington 1992).
Due to the incompleteness of the insect fossil record, the lines of
inquiry pursued with fossil marine invertebrates are not always feas-
ible with fossil insects. The current decline of animal groups such as
bivalves and vertebrates is typically compared to the Big Five mass
extinctions with one of two goals. The first of these goals is to de-
termine whether the current crisis could reach the severity of any of
the Big Five mass extinctions (Barnosky et al. 2011). The second goal
is to compare the patterns and causes inferred from the Big Five to
the patterns and causes observed in the current crisis (Harnik et al.
2012, Payne and Clapham 2012, McCallum 2015). Neither of these
goals can be achieved with the insect fossil record.
In addition to being sparse, the insect fossil record is fragmentary
across space and in time. Most described insect fossils come from
‘konzentrat Lagerstätten’, or fossil assemblages where the depos-
itional conditions permitted vast numbers of insects to be preserved.
Here, we arbitrarily define konzentrat Lagerstätten as assemblages
with 100 or more described insect specimens. Fossil insects have
been described from approximately 2,500 assemblages and only 90
of these assemblages are konzentrat Lagerstätten (Fig. 1). However,
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 3

66% of all described insect occurrences in the fossil record come
from these 90 konzentrat Lagerstätten.
The 90 konzentrat Lagerstätten in the insect fossil record cannot
provide complete coverage of insect taxa for the past 320 million
years. Konzentrat Lagerstätten are unevenly distributed in time: the
longest gap between successive konzentrat Lagerstätten is 22 mil-
lion years, between the 218-million-year-old Ipswich Lagerstätte and
the 196-million-year-old Issyk-Kul Lagerstätte. In contrast, there
are eight konzentrat Lagerstätten from the last 126,000 yr alone
[Titusville (the United States), Omolon (Russia), Upton Warren
Because fossil insects are relatively rare and occur so unevenly
in time and space, the severity of insect extinctions in deep time is
remarkably difficult to measure. This renders the timing of insect ex-
tinctions similarly difficult to constrain. These limitations preclude
studies of whether the current insect decline is comparable in magni-
tude to any of the Big Five and prevent precise determinations of the
causes of insect extinctions in the fossil record.
Abundance and biomass data are not available for fossil insects
(Fig.  2). For marine animals that live in a circumscribed depositional
setting, it is possible to measure changes in animal abundance through

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(the United Kingdom), Malaya Konkovaya River (Russia), Thrupp
House Farm (the United Kingdom), Magnolia Bluff (the United
States), Ziegler Reservoir (the United States), Kalaloch (the United
States)], encompassing the latest Pleistocene and Holocene epochs.
These Lagerstätten occur within ‘unlithified’ sediments that have not
hardened into rocks and that therefore have a low probability of
persisting in the rock record millions into the future (Clapham et al.
2016). The Priabonian Stage of the Eocene Epoch, which lasted for
four million years (37.8–33.9 Ma), also contains eight konzentrat
Lagerstätten, due primarily to the preservation of amber near the
Baltic Sea [Bembridge Marls (United Kingdom), Bitterfeld amber
(Germany), Rovno amber (two Lagerstätten in Ukraine), Baltic
amber (one Lagerstätte each in Russia, Poland, Lithuania, and
Denmark)].
The distribution of konzentrat Lagerstätten is even more in-
consistent across space than time. The vast majority of konzentrat
Lagerstätten—90%—occur at paleolatitudes over 20° from the
paleoequator, and over half of konzentrat Lagerstätten—52%—
occur at paleolatitudes over 43° from the paleoequator (Fig. 1). Not
a single konzentrat Lagerstätte occurs at paleolatitudes between 10°
and 20° from the paleoequator. This overwhelmingly temperate dis-
tribution of fossil insects is comparable with that of fossil plants
and contrasts with the primarily equatorial distribution of nonavian
dinosaurs (Rees et al. 2004). The dearth of tropical and subtropical
konzentrat Lagerstätten is troubling because the richness of extant
insect species is highest, and disproportionately undescribed, in the
tropics (Stork 2018).
time. If animals become scarce or absent in such deposits, it can be in-
ferred that these animals really were scarce or absent in the particular en-
vironment in which those deposits formed (Payne et al. 2006). The link
between decreased abundance and increased extinction risk, central to
many studies of the extinction crisis in living insects, has also been exam-
ined for marine animals in the fossil record (Payne et al. 2006, Simpson
and Harnik 2009). However, the fossil record of insects does not permit
such inferences. This is not only because the fossil record of insects is
relatively depauperate and uneven compared to that of many marine
taxa, but also because insects overwhelmingly tend not to live in the sorts
of settings, such as lake basins, where fossils most easily form. If a lake
deposit containing insect fossils were studied in sufficient detail to deter-
mine changes in insect abundance through time, the apparent changes
could reflect true changes in insect abundance near the lake but also
could be confounded by abundance in nearby habitats. For example, if
insects are typically swept into a lake by flash floods (Smith 2012), a de-
crease in flash flooding over time would cause a decrease in the number
of insects brought into this depositional setting.
Biases Inherent to the Insect Fossil Record
The fossil record is biased toward certain orders and families, and
this is no less true for insects as for other taxa (Smith 2012). The
biases discussed in this section are inherent to the fossil record: there
are biases as to which taxa arrive in depositional environments and
become preserved there. The biases discussed in the next section are
due to human activity: there are biases as to which taxa are priori-
tized for curation in museum collections and which taxa are formally

90.0
Amber Kap København
Adpression Taimyr Alberta,
Ust-Baley (Siberia) Manitoba
66.3 ●
Issyk-Kul ● (Canada)
Álava Baltic
Isady Sai-Sagul (Spain) ●
Kuznetsk ● Fushun ●● ●
Madygen ●
Soyana Posidonia Karatau New Menat Oise
Chekarda Jersey (France)
23.5
Paleolatitude

Tikhie ●●
●●
Obora Gory Myanmar Dominican
Wettin Lebanon ●
Elmo ●
0.0 Commentry Tanzania ●
Mazon Crato
Creek
-23.5

Ipswich
-66.3 Belmont

-90.0
Paleozoic Mesozoic Cenozoic
Pennsylv. Permian Triassic Jurassic Cretaceous Paleogene Neogene

Age
323 299 252 201 145 66 23 0
(Ma) Late End- End- Late Later Early End-
Pennsylvanian Permian Triassic Jurassic Cretaceous Cretaceous
insect turnover event event insect turnover insect turnover e v en t

Fig. 1. The ages and paleolatitudes of the 90 konzentrat Lagerstätten that contain at least 100 described insect occurrences. The points representing each
Lagerstätte are semitransparent, such that darker points represent multiple Lagerstätten with nearly identical ages and paleolatitudes.
4 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX

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Fig. 2.  Data from the fossil record can only be compared at the coarser taxonomic scales of order and family, whereas modern ecological data allow comparisons
at the genus, species, and especially the population scale. All icons are from PhyloPic.org. Mantoida nitida Newman, 1838 (Mantodea: Mantoididae), license at
http://creativecommons.org/publicdomain/mark/1.0/. Trachypachus gibbsii Leconte, 1861 (Coleoptera: Trachypachidae) by T. Michael Keesey and Yves Bousquet,
and Somatochlora tenebrosa (Say, 1839) (Odonata: Corduliidae) and Eristalis tenax (Linnaeus, 1758) (Diptera: Syrphidae) by Gareth Monger, license at http://
creativecommons.org/licenses/by/3.0/. Hyalophora cecropia (Linnaeus, 1758) (Lepidoptera: Saturniidae) by Mali’o Kodis and Jim Vargo, license at http://
creativecommons.org/licenses/by-nc-sa/3.0/.

described, plus biases as to which taxonomic experts work on the
material and which taxa journals are interested in publishing. One
overarching question that arises from this series of biases is whether
they cancel each other out, as was found to be the case in a canon-
ical study of trilobites (Adrain and Westrop 2000), or reinforce each
other. The answer to this question may vary with the taxonomic and
temporal scale of the data considered.
The need for an insect to live near a body of water—or, alterna-
tively, in a specific microhabitat such as a tree surface that exudes
certain resins—causes a bias in the insect fossil record: habitat.
Although a number of tree species exude resin that has been pre-
served as amber (Labandeira 2014a), many insect species do not
live near these trees. The major amber deposits are formed from the
resin of vastly different trees, ranging from gymnosperms to angio-
sperms, introducing another bias (Penney 2010). Because strata typ-
ically form in basins such as bodies of water, higher-altitude habitats
and environments lacking internally drained ponds and lakes pro-
vide vanishingly few opportunities for fossilization. Paleobotanists
have extensively discussed the role of upland and dryland habitats in
plant evolution (Chaloner 1958, Falcon-Lang and Bashforth 2004,
Thomas and Cleal 2017, Cleal and Cascales-Miñana 2019), but this
topic is considered for insects only rarely (Shcherbakov 2008).
Within a given habitat, there are also biases in the body parts most
likely to be preserved. The thorax, abdomen, antennae, legs, palps, and
other structures consist of multiple segments that can easily become dis-
articulated in transit to a depositional setting that would permit fossil-
ization (Smith 2000). Wings are the insect body part that fossilize most
easily. Indeed, the two earliest definitive fossils of winged insects are
isolated wings (Brauckmann and Schneider 1996, Prokop et al. 2005), as
is common for many insect taxa throughout earth history.
Certain morphological features cause taxonomic biases. Lepidoptera
(moths and butterflies) have a notoriously incomplete fossil record
(Sohn et al. 2015), due in large part to their characteristic thickly scaled
wings and delicate bodies. The scales cause wings to float, preventing
them from sinking into sediments at the bottom of bodies of water
where they can be buried and preserved. Lepidoptera are one of the five
overwhelmingly speciose insect orders in the modern fauna (Grimaldi
and Engel 2005), but their fossil record contains only 2.6% as many
described specimens as that of Coleoptera (beetles), 4.5% as many
described specimens as that of Diptera (true flies), 6.3% as many de-
scribed specimens as that of Hymenoptera (sawflies, wasps, ants, and
bees), and 6.8% as many described specimens as that of Hemiptera
(aphids, whiteflies, scale insects, and bugs).
Because the smallest insect bodies and body parts are those that
can most easily fit into a drop of tree resin or become completely
buried in a body of water, the insect fossil record is overwhelmingly
biased against large taxa (Labandeira 2014a). Gigantic dragonfly re-
latives such as the Meganisoptera and Palaeodictyopteroidea, present
from the Middle Pennsylvanian to the Early Permian (315–272 Ma),
are among the most spectacular of fossil insects but are extremely
rare. Indeed, although paleopterous, orthopteroid, and holomet-
abolous groups independently evolved to be larger than any insect
alive today, there are only 54 wings known in the entire fossil record
that are longer than those of any extant insect species (Clapham and
Karr 2012). Furthermore, four described fossilized wings measure at
least 30 cm but there are no described wings between 26 and 30 cm
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 5

in length, although insects with 30-cm wings presumably had ances-
tors with wing lengths in this range.
Publication Biases
The smallest insects, which fossilize most easily (Smith 2012), are
the most difficult to study, with minor exceptions of certain amber
deposits (Labandeira 2014a) and compression–impression deposits
occurring in unusual environments (Greenwalt et  al. 2014). Their
body size makes them difficult to notice, whether at a mountainside
outcrop or in an amber specimen, and renders them difficult to view
this Lagerstätte (Rasnitsyn and Zhang 2004a,b; Ansorge et al. 2006;
Rasnitsyn et  al. 2006; Rasnitsyn and Zhang 2007, 2010; Zhang
and Rasnitsyn 2008; Li et al. 2013, 2015; Zhang et al. 2014; Wang
et  al. 2014, 2015b, 2016b, 2017; Kopylov et  al. 2020; Rasnitsyn
and Öhm-Kühnle 2020). The influence of investigator preference
may depend on the amount of time that a Lagerstätte has been
known, which can vary greatly. In Europe alone, Baltic amber has
been known for centuries (Weitschat and Wichard 2010) whereas
the amber deposits of Álava in Spain and Oise in France have been
known for only a few decades (Nel et al. 1999, Alonso et al. 2000).

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and image in a laboratory setting. The small size of a typical fossil-
ized insect also reduces the number and quality of characters avail-
able for analysis in a taxonomic study. This ultimately leads to a bias Comparisons with Other Animals
toward medium-sized insects, which are small enough to fossilize
Comparisons with Marine Invertebrates
relatively intact but large enough to be noticed by researchers and
In general, shelled marine invertebrates have a far more complete
display an informative suite of apomorphies.
fossil record than insects do (Foote and Sepkoski 1999). Marine
There also is a bias toward description of specimens that rep-
groups such as bivalves, gastropods, and brachiopods have hard
resent the first and last occurrences of higher-level taxa (Schachat
shells that facilitate fossilization, and these animals tend to spend
et  al. 2019). First occurrences are of unparalleled importance for
their lives atop or within the layer of sediment where fossilization
time-calibration of divergences in phylogenies (Kohli et  al. 2016,
can occur (Kidwell and Flessa 1996).
Evangelista et al. 2017) and last occurrences inform debates about
The oxygen content of the oceans has decreased steadily for dec-
when extinct taxa disappeared (Béthoux et  al. 2010, Shcherbakov
ades (Helm et al. 2011), increasing extinction risk for many marine
2011). Because of this publication bias favoring first and last occur-
animals (Keeling et  al. 2010). From first principles one would ex-
rences, the described fossil history of many groups contains large
pect insects to fare better than marine invertebrates during a biotic
gaps between very early and very late occurrences. For example, the
crisis caused by low oxygen concentrations. Seawater contains far
extinct family Lophioneuridae, related to Thysanoptera, has 32 de-
less oxygen than the atmosphere (Vaquer-Sunyer and Duarte 2008).
scribed fossil specimens from the first 32 million years of its known
Furthermore, oxygen levels can fall in the ocean due to factors such
history and 11 described fossil specimens from the last 77 million
as decreased solubility and increased stratification that are irrele-
years of its known history, but not a single described fossil specimen
vant in the atmosphere (Breitburg et  al. 2018). Lastly, small mol-
from the intervening 91 million years.
ecules such as oxygen travel 10,000 times faster through air than
Because of the paleoentomological focus on systematics, few publi-
water (Vogel 1994). For these reasons, breathing air is far easier than
cations about fossil insects focus on faunas (Solórzano Kraemer 2007,
breathing water.
Penney 2010, Guénard et al. 2015) rather than taxa. Publications typ-
These are major reasons why deoxygenation of the ocean is a
ically describe new genera and species (Nel and Jarzembowski 2019,
major concern in the context of anthropogenic global warming
Zhao et al. 2019), or, less commonly, describe first occurrences of higher-
(Keeling et  al. 2010) whereas deoxygenation of the atmosphere is
level taxa (Haug et al. 2015, Zheng et al. 2018, Montagna et al. 2019).
not. Furthermore, the atmosphere is sufficiently well mixed that at-
Only a handful of paleoentomological studies have quantified clades’
mospheric oxygen levels do not vary geographically in the absence
relative prevalence within a deposit (Solórzano Kraemer 2007, Grimaldi
of a large change in altitude. In contrast, the ocean is poorly mixed
et al. 2010, Perrichot et al. 2010) as is often done for marine inverte-
(Paulmier and Ruiz-Pino 2009). The ocean appears to have con-
brates (Hsieh et al. 2019). The commercial operations that mine amber
tained anoxic regions throughout earth history (Meyer and Kump
from major deposits (Penney 2010) also obstruct the methods for con-
2008) and has occasionally contained massive anoxic zones during
ducting a paleontological census. As a result, only a small fraction of
intervals that may (Lau et al. 2016) or may not (Jenkyns 1980) be
specimens from each deposit are mentioned in the published literature.
associated with the Big Five mass extinctions.
Barely 5 % of the fossil insects from the Late Jurassic Karatau deposit,
Other consequences of increased pCO2 and the resulting global
for instance, have been formally described (Grimaldi and Engel 2005).
warming can cause extinction in the oceans but not on land. Ocean
Approximately 1,000 fossils insects have been excavated from each
acidification makes it more difficult for animals such as corals to
of the Obeshchayushchiy, Kzyl-Zhar, and Orapa deposits (Zherikhin
produce calcium carbonate for their skeletons (Andersson and
2002), which date to the notoriously fossil-poor Late Cretaceous (100–
Gledhill 2013). Warming also changes the distribution of salinity in
66 Ma), but only 75, 57, and 38 specimens have been formally described
the ocean (Durack and Wijffels 2010). Deoxygenation, acidification,
from each of these deposits, respectively.
and changes in salinity caused by increased pCO2 are not new prob-
Investigator preference and expertise can also impact the emer-
lems: all of these have been implicated in at least one of the Big Five
gent faunistic view of a Lagerstätte. For example, an examination
(Kiessling and Simpson 2011, van Soelen et al. 2018, Zhang et al.
of the tens of thousands of fossil insects collected from Daohugou
2018).
(Jurassic) has shown that the three most abundant orders in this
To put it simply, environmental changes can cause more physio-
Lagerstätte are Orthoptera, Neuroptera, and Diptera (Huang
logical stress for animals in the ocean than on land. An increase in
et  al. 2006). However, Hymenoptera dominate the small fraction
pCO2 large enough to prevent clams from breathing and constructing
of Daohugou insects that have been published. This anomaly may
their shells might not pose any challenge to insect physiology.
be attributable in large part to Alexandr Rasnitsyn, who became
established as a preeminent expert in fossil Hymenoptera (e.g.,
Rasnitsyn 1967, 1988) long before the first description of fossils Comparisons with Tetrapods
from Daohugou (Wang et al. 2000, Yuan 2000) and who has con- On land, tetrapods are one of the few groups that predominate in
tributed to many publications describing fossil Hymenoptera from terms of biomass, although their biomass pales in comparison to
6 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
that of insects (Bar-On et  al. 2018). Because tetrapods have a far
more complete fossil record than insects, it is tempting to look to
the tetrapod fossil record for clues as to how insects may have fared
during mass extinctions.
Several differences limit the relevance of the tetrapod fossil record
to that of insects. Perhaps, the most obvious is the minimum overlap
in body size between insects and tetrapods. Currently, most insects
are smaller than the smallest living tetrapod (Finlay et  al. 2006,
Rittmeyer et al. 2012). Insects’ smaller body size makes it easier for
them to hide and to survive on limited quantities of food. Insects’
small size also allows them to occur at higher abundances within a
community (Finlay et al. 2006) and to reach reproductive maturity
quicker, which leaves insects with shorter generation times. During
brief environmental disturbances such as drought, extreme heat, or
the aftermath of an asteroid impact, insects’ small body size may
allow them to take refuge in microhabitats that are not available to
tetrapods. Arthropod faunas in modern forests can contain just as
many species and genera in leaf litter as on vegetation (Brühl et al.
1998), but this is not the case for tetrapods (Gardner et al. 2007).
Within a given timespan in a particular habitat, the average in-
sect species can adapt to changing environmental conditions faster
than the average tetrapod species. Insects occur at higher densities,
providing more variation on which natural selection can act, and in-
sects’ greater number of generations per unit of time allows evolution
to happen faster. The frequency with which insects enter diapause
also has been invoked as a mechanism through which they manage
to survive environmental crises, particularly those of limited dur-
ations (Whalley 1987, Tuljapurkar and Istock 1993). Consequently,
an environmental crisis that causes a mass extinction of tetrapods
need not cause a mass extinction of insects.
Data, Methods, and Terminology
Extinctions in deep time will be discussed here in terms of the geo-
logical periods of the Phanerozoic Eon (541 Ma to the present),
Fig. 3.  A schemtic showing ‘stem’ and ‘crown’ representatives of nested clades. A crown group is the monophyletic clade that includes all extant descendants of
a given ancestor. The stem group is the most comprehensive paraphyletic grade that includes extinct lineages that are more closely related to the crown group
at hand than to any other crown group.
which are from oldest to youngest: the Cambrian (541–485 Ma),
Ordovician (485–444 Ma), Silurian (444–419 Ma), Devonian (419–
359 Ma), Mississippian (359–323 Ma), Pennsylvanian (323–299
Ma), Permian (299–252 Ma), Triassic (252–201 Ma), Jurassic (201–
145 Ma), Cretaceous (145–66 Ma), Paleogene (66–23 Ma), Neogene
(23–2.6 Ma), and Quaternary (2.6 Ma to the present) Periods. Of
note, some authors denote the interval from 359 to 299 Ma as the
Carboniferous Period and many others divide the Carboniferous
into two Periods, the Mississippian and the Pennsylvanian. Here,
the Mississippian and the Pennsylvanian are treated as separate
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periods rather than subperiods in accordance with the latest geo-
chronology (Walker et al. 2013) and because the Pennsylvanian con-
tains definitive insect body fossils whereas the only fossil insects that
might be Mississippian in age occur in very close proximity to the
Mississippian/Pennsylvanian boundary (Brauckmann and Schneider
1996, Prokop et al. 2005).
Subordinate intervals are discussed here as belonging to the early,
middle, or late portions of their respective periods. For example, all
fossils and events from the Changhsingian Age of the Lopingian
Epoch will be discussed here as a part of the Late Permian. The early,
middle, and late intervals within each geologic period were desig-
nated long before the advent of reliable absolute dating methods,
such as uranium-lead dating, and are not necessarily of equal length.
For example, the Early Permian lasts for more than half of the
Permian, and the Middle Triassic begins and ends within the earliest
third of the Triassic. The Cretaceous consists of an early and a late
interval with no formal Middle. The early, middle, and late intervals
within each period are demarcated just below the period designa-
tions in Fig. 1.
The concept of a ‘stem’ and ‘crown’ group within each clade is
illustrated in Fig.  3. Crown representatives of many extant orders
appeared during the Triassic, but stem groups typically extend back
to the Paleozoic (Grimaldi and Engel 2005, Labandeira 2006).
Some of the figures presented here include insect occurence data from
the fossil record. These figures were generated with data downloaded
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 7

from the Paleobiology Database on 22 March 2020, compiled pri-
marily by Clapham et al. (2016). To visualize family-level relationships
among insect communities before and after mass extinction bound-
aries, we conducted nonmetric multidimensional scaling (NMDS) of in-
sect Lagerstätten using vegan (Oksanen et al. 2019) version 2.5–6 in R
(R Development Core Team 2020) version 3.6.2. Because Lagerstätten
contain varying numbers of described occurrences, we subsampled each
Lagerstätte down to 100 occurrences and iterated this procedure 100
times. We generated plots of our NMDS results in ggplot2 version 3.2.1
(Wickham 2009) illustrating each subsampled insect community and
pattern also seen in the plant and tetrapod records from this interval
(DiMichele and Aronson 1992, Huttenlocker et  al. 2018). The in-
sect extinction event of the Late Jurassic is overshadowed by many
contemporaneous diversification events, such as the mid-Mesozoic
family-level radiation of parasitoid wasps (Labandeira 2018,
Schachat et al. 2019). The insect extinction event of the later Early
Cretaceous occurred during one of the most spectacular diversifica-
tion events in insect history (Labandeira 2014b) when total insect
diversity reached one of its highest peaks (Schachat et al. 2019).

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an 84% confidence ellipse for each Lagerstätte, following the methods
outlined by Schachat et al. (2020). The ‘Big Five’ Mass Extinctions in Deep Time
The End-Ordovician Event
The first of the Big Five mass extinctions, the end-Ordovician event,
The Meaning of a Mass Extinction
occurred 444 million years ago (Sheehan 2001). This event is known
The insect fossil record contains major insect extinctions that do entirely from the marine realm and predates all putative fossil insect
not overlap with the Big Five. Labandeira (2005) identified insect occurrences by tens of millions of years. In addition to the lack of
extinctions at the Late Pennsylvanian, Late Jurassic, and later Early fossil evidence of any insects before this extinction, there are also
Cretaceous. The Late Pennsylvanian insect extinction (Labandeira no fossils of vascular plants or tetrapods. Bryophytes are inferred
and Phillips 2002) occurred as the world became warmer and dryer to have diversified before this extinction but their depauperate fossil
(Joeckel 1999) and coal swamps shrank (DiMichele et  al. 2006), record does not contain any macroscopic remains (Taylor et  al.
causing rapid turnover (DiMichele et  al. 2001) in which approxi- 2009). The end-Ordovician event is of no relevance to insects.
mately two-thirds of all coal-swamp plant species disappeared
(DiMichele and Phillips 1996). The Late Jurassic and later Early The Late Devonian Event
Cretaceous extinctions are the two main pulses in the long de-
Controversy surrounds the timing, magnitude, and duration of the
cline of several major insect lineages, particularly Neuroptera and
Late Devonian event (McGhee Jr 2013). Extinctions have been noted
Mecoptera, and other lineages such as certain odonate families and
near the beginning of the Givetian Stage at 388 Ma (Bosetti et  al.
bizarre, flea-like insects (Labandeira and Sepkoski 1993, Bechly et al.
2011); later in the Givetian, approximately 384 Ma (Aboussalam
2001, Ren et al. 2009, Gao et al. 2013, Lin et al. 2019). Although
and Becker 2011, Marshall et al. 2011); at the Frasnian/Famennian
Neuroptera and Mecoptera have not gone extinct, their current di-
boundary, 372 Ma (McGhee 1996); and at the end of the Devonian,
versity is considerably less than what it was during the Mesozoic
359 Ma (Kaiser et al. 2016). This loss of biodiversity may in fact be
(Labandeira 2010), and entire suborders or superfamilies have gone
extinct within both orders (Yang et al. 2014, Lin et al. 2019).
The difference between a mass extinction and turnover is the key
to understanding these three events. Changes in taxonomic diversity
in deep time are measured by changes in extinction rates, origination
rates, and total taxonomic diversity (Alroy and Hunt 2010). The
extinction rate is the proportion of taxa within an interval that do
not survive into the next interval. The origination rate is the propor-
tion of taxa within an interval that were not present in the previous
interval.
One of the two things can happen when extinction rates increase.
If origination rates do not rise along with extinction rates, total taxo-
nomic diversity will drop. Under this scenario, many taxa have gone
extinct with insufficient new taxa to replace them. If extinction rates
are high enough and total taxonomic diversity decreases sharply
enough, a mass extinction occurs.
Alternatively, origination rates can increase alongside extinction
rates. Under this scenario, many taxa have gone extinct but many
new taxa have originated at approximate replacement levels, causing
total taxonomic diversity to remain relatively stable. Such an extinc-
tion event occurs within the context of biotic replacement and is not
a mass extinction. The term ‘turnover’ conveys an increase in origin- Fig. 4. Variations in origination and extinction rates, and the resulting
ation that compensates for the increase in extinction. changes in total taxonomic diversity. In (a), extinction and origination both
The distinction between turnover and a mass extinction increased: the extinction that occurred is not immediately obvious in the
shows why the Late Pennsylvanian, Late Jurassic, and later Early diversity curve because the extinction event happened within the context
of evolutionary turnover. In (b), extinction increased while origination
Cretaceous insect extinction events identified by Labandeira (2005)
decreased, causing a net loss of taxonomic diversity as seen in the diversity
are not analogous to the current insect extinction crisis. The Late
curve: a true biotic crisis, or mass extinction. In (c), total taxonomic diversity
Pennsylvanian extinction of insects that lived in coal swamps and has decreased, but not due to an increase in extinction: extinction decreased,
adjacent humid habitats was part of a transition in which other in- rather than increased, but origination decreased even more than extinction,
sect groups diversified in more xeric habitats (Labandeira 2006), a causing a ‘mass depletion’ sensu Bambach et al. (2004).
8 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
more of a ‘depletion’, caused in large part by low origination rates
(Fig. 4), than a true mass extinction (Bambach et al. 2004).
During the approximately 75 million years that passed between
the end-Ordovician event and the Late Devonian event, terrestrial eco-
systems changed dramatically: in many habitats, microbial soil crusts
seemingly devoid of macroscopic animal life were replaced by for-
ests with 30-m-tall trees (McGhee Jr 2013). Early tetrapods ambled
through shallow bodies of water, perhaps occasionally venturing onto
land (Ahlberg and Clack 2006). Though vertebrates may not have been
terrestrial by this time, Middle-to-Late Devonian landscapes were un-
questionably home to a wide range of terrestrial arthropod groups
including early centipedes, scorpions, and springtails, extinct groups
such as trigonotarbids and arthropleurans—and most likely to primi-
tively wingless insects as well (Shear et al. 1984).
The fossil record of insects before the Late Devonian event is min-
imally informative: the one and only insect fossil from this interval
whose affinity and age have not been questioned is a small frag-
ment assigned to Archaeognatha (Shear et  al. 1984), a primitively
wingless order that is widely agreed to represent the most early-
diverging known branch of Insecta (Misof et al. 2014). The presence
of Archaeognatha in Middle Devonian landscapes is unsurprising,
given the presence of a collembolan in the Early Devonian (Whalley
and Jarzembowski 1981). In addition to this probable Devonian
archaeognathan fragment, there is another possible Devonian
archaeognathan fragment (Labandeira et al. 1988) whose antiquity
was questioned by authors who did not examine the original ma-
terial (Jeram et al. 1990) and a headless fragment of an animal that
appears to belong to either Archaeognatha or Zygentoma (Fayers
and Trewin 2005). Regardless of whether these additional fragments
represent Devonian insects, they do not change the emergent view of
Devonian insect life.
Until recently, the Late Devonian event was considered by some
authors to have occurred within a biotic crisis that suppressed in-
sect abundance for tens of millions of years. A 407-million-year-
old mandible was interpreted, controversially, as belonging to a
winged insect (Engel and Grimaldi 2004). The next known fossils
of winged insects are approximately 80 million years younger than
this mandible (Brauckmann and Schneider 1996, Prokop et  al.
2005), raising the question of why Pterygota are absent from the
fossil record for several tens of millions of years of their supposed
history. Shortly after the 407-million-year-old mandible was at-
tributed to a winged insect, estimates of atmospheric oxygen in
Earth’s deep past showed that the gap in the insect fossil record
coincides with reduced pO2 (Berner 2006). These findings suggest
that the Late Devonian event occurred near the midpoint of a long
interval in which reduced oxygen availability suppressed insect
abundance (Ward et al. 2006).
More recently, the 407-million-year-old mandible was redescribed
as belonging to a centipede (Haug and Haug 2017). In a relevant
development, the same model that reconstructed low oxygen levels
near the Late Devonian event was re-run with updated input of iso-
topic data and estimates of low oxygen were not recovered at or near
the Late Devonian (Schachat et  al. 2018). Estimates of unremark-
able oxygen levels are consistent with the various described occur-
rences of arachnids throughout this interval (Shear 2000, Gess 2013)
which suggest that winged insects could have become abundant on
the landscape and could have been preserved in the Devonian rock
record if they had already originated (Schachat et al. 2018). An en-
vironmental crisis no longer explains why Pterygota are not known
within 45 million years of the Late Devonian event, but an explan-
ation is no longer needed because there is no evidence that Pterygota
existed at that time.
Like Collembola, Archaeognatha appear to have originated be-
fore the Late Devonian event. Accordingly, Archaeognatha could not
have gone extinct during the Late Devonian, because they still exist
today. That Archaeognatha survived is all that can be confidently
said about how insects fared during the Late Devonian event.
The End-Permian Event
The end-Permian event at 252 million years ago was the most severe
extinction crisis in the history of life (Raup and Sepkoski 1982) and
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the most ecologically impactful (McGhee et al. 2004, 2012; Wagner
et al. 2006). Over 80% of all marine genera are estimated to have
perished (Stanley 2016). The end-Permian was also the only time
when a supraordinal clade of insects went extinct (Labandeira and
Sepkoski 1993). This has led certain paleontologists to communicate
the unique severity of the end-Permian event by describing it as the
only extinction event that was severe enough to extinguish insects
(Bowring et  al. 1998). Indeed, all methods used to estimate insect
diversity in deep time show a major loss of family-level diversity at
the end-Permian (Labandeira and Sepkoski 1993, Jarzembowski and
Ross 1996, Shcherbakov 2008, Clapham et al. 2016, Dmitriev et al.
2018, Schachat et al. 2019). However, the attribution of this loss of
insect diversity to environmental causes, rather than evolutionary
dynamics unique to the Insecta, is questionable.
Two paleopterous groups appear to have gone extinct near the
end-Permian. These contain the majority of the paleopterous or-
ders that ever existed; their disappearance constitutes a major loss
of morphological diversity and evolutionary history. The order
Meganisoptera, also known as Protodonata, is widely agreed to
form the stem-group (Fig.  3) to crown Odonata (Nel et  al. 2001,
Grimaldi and Engel 2005). The superorder Palaeodictyopteroidea
contains four extinct orders—Palaeodictyoptera, Permothemistida,
Megasecoptera, and Diaphanopterodea—and its relationships with
other lineages of Paleoptera are still in dispute (Béthoux 2008).
Meganisoptera and Palaeodictyopteroidea are the two lineages
whose representatives reached the largest body sizes in insect his-
tory: Meganisoptera reached wing lengths of at least 33  cm and
Palaeodictyopteroidea reached wing lengths of at least 22.6  cm,
whereas the longest insect wing length known after the Permian/
Triassic boundary is merely 18 cm. But not all Palaeodictyopteroidea
were gigantic, and certain lineages evolved progressively smaller
body sizes throughout the Permian with some reaching the size of a
mosquito (Wootton 1981, Van Dijk and Geertsema 1999).
Meganisoptera and Palaeodictyopteroidea are also known for
their feeding strategies. Meganisoptera were specialized hawker
predators whose visual acuity allowed them to patrol ancient for-
ests (Nel et al. 2018a) and Palaeodictyopteroidea had piercing-and-
sucking mouthparts that are structurally reminiscent of Hemiptera
but evolved independently (Labandeira 1997, 2019; Pecharová et al.
2015). During the Pennsylvanian Period (323–299 Ma), the first geo-
logic period with definitive evidence of insects, Palaeodictyopteroidea
were abundant and diverse, comprising appropriately half of the de-
scribed insect species from this time (Carpenter 1971).
The end-Permian marks the disappearance of insect groups that
had previously been abundant and diverse. However, Meganisoptera
and Palaeodictyopteroidea were in decline throughout the Permian
(Fig. 5). In some regions, Paleoptera became rare before the Permian
began (Rasnitsyn et  al. 2004, Schneider et  al. 2004, Shcherbakov
2008). The Palaeodictyopteroidea are inferred to have been outcom-
peted by the Hemiptera due to the similarity of their mouthparts
and feeding strategies (Labandeira 1997, Grimaldi and Engel 2005,
Schachat and Labandeira 2015). The extinction of the Meganisoptera
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
Fig. 5.  The proportion of described insect occurrences from Pennsylvanian,
Permian, and Triassic konzentrat Lagerstätten that belong to the
Palaeodictyopteroidea and to other paleopteran lineages, i.e., stem and
crown Ephemeroptera and Odonata.
is unremarkable in that many major lineages of insects have stem
representatives (Fig. 3) that went extinct near the Permian/Triassic
boundary (Labandeira and Sepkoski 1993); not all of these stem
groups are treated as separate, highly ranked taxonomic groups
such as orders. The end-Permian extinction of Meganisoptera and
Palaeodictyopteroidea resembles the end-Permian extinction of tri-
lobites in that the taxonomic diversity of all three of these once-
abundant groups was already in decline before the Permian began
(Lerosey-Aubril and Feist 2012).
The slow pace of faunal turnover near the end-Permian be-
lies the unparalleled significance of this transition. Only two
evolutionary entomofaunas have been identified in deep time:
the Paleozoic Insect Fauna and the Modern Insect Fauna
(Jarzembowski and Ross 1996, Zherikhin 2002, Labandeira
2005). These faunas do not form monophyletic clades, or even
paraphyletic grades; they are the polyphyletic assemblages that
dominated insect communities of their times, akin to the three
marine faunas identified by Sepkoski (1981). Stem representa-
tives of Paleoptera, Polyneoptera, and Paraneoptera prevailed
in the Paleozoic Insect Fauna. The Modern Insect Fauna, with
us to this day, is epitomized by crown Odonata, Polyneoptera,
Paraneoptera, and especially Holometabola (Labandeira 2005).
One of the most notable transitions between the Paleozoic
and Modern Insect Fauna is the ecological replacement of the
Palaeodictyotperoidea by the Hemipteroidea; both major clades pos-
sessed stylate mouthparts and piercing-and-sucking feeding habits
(Labandeira 2019). Another ecological replacement was that of
various basal and stem-group orthopteroids of the Archaeorthoptera,
basal Orthoptera, many ‘roachoids’, and perhaps Reculida of the
Paleozoic Insect Fauna by Orthoptera, Phasmatodea, and possibly
Titanoptera of the Modern Insect Fauna.
The Paleozoic Fauna dominated insect communities until the
end-Permian and the modern fauna dominated insect communities
thereafter, but the turnover between these two faunas was gradual.
Consequently, the end-Permian event is best conceptualized in an
entomological context as the approximate midpoint of the most im-
portant global shift in taxonomic composition that occurred in in-
sect history.
9
The current consensus is that the end-Permian event was caused
by the Siberian Traps, a massive ‘large igneous province’ whose
magma burned through overlying coal deposits, releasing CO2 into
the atmosphere from both lava and coal (Ganino and Arndt 2009,
Svensen et al. 2009, Ogden and Sleep 2012). The geographic ranges
of many groups, including insects, shifted poleward in an apparent
response to global warming (Shcherbakov 2008). Changing envir-
onmental conditions would have posed many challenges to animal
physiology at this time. And yet, controversy continues to surround
the chemical composition of the atmosphere and the timing of the
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environmental perturbations (Cui and Kump 2015, Shen et al. 2019).
The fossil records of plants and tetrapods, which are more com-
plete than that of insects, show evidence of widespread ‘disaster forms’
or ‘disaster taxa’ during the Early Triassic. Disaster taxa include
the small pole-like tree Pleuromeia Corda, (1852) (Pleuromeiales:
Pleuromeiaceae), a lycopsid whose closest living relative is the diminu-
tive quillwort Isoetes Linnaeus, 1753 (Isoetales: Isoetaceae), and the
tetrapod Lystrosaurus Cope, 1870 (Anomodontia: Dicynodontia:
Lystrosauridae), a therapsid whose closest living relatives are mam-
mals. These became widespread after the extinction event (Benton
1983, Grauvogel-Stamm and Ash 2005, Looy et al. 2005). Disaster
taxa are emblematic of the low alpha, beta, and gamma diversity
that characterized plant and tetrapod communities for several mil-
lion years after the end-Permian event (Benton and Newell 2014).
A debate persists, however, as to whether plants experienced a mass
extinction at the end-Permian (Cascales-Miñana and Cleal 2014,
Nowak et al. 2019) as tetrapods did (Maxwell 1992), with doubts
surrounding the geographic scale of the purported plant extinction
(Rees 2002). Nevertheless, the end-Permian event depressed plant
and tetrapod diversity for an extended interval and fundamentally
different communities emerged in its wake.
The relevance of plants and especially tetrapods to insects at
the end-Permian event is tenuous. The number of described insect
species that exist today vastly exceeds the number of described and
estimated total species of plants and tetrapods (Mora et al. 2011),
and there is no reason to doubt that insects were already far more
diverse than plants and tetrapods at the end of the Permian. The
number of described insect species from each of the nine Permian
konzentrat Lagerstätten exceeds the global estimated tetrapod diver-
sity for the entire Permian (Benson and Upchurch 2013). The highest
number of estimated plant species from all Permian stages is ap-
proximately equal to the number of insect species described thus far
from the Elmo Lagerstätte of the Early Permian of Kansas (Nowak
et  al. 2019). Because insects already were relatively diverse at the
end-Permian, an insect community would have had to lose far more
species, genera, and families than a plant or tetrapod community in
order to become dominated by a single ‘disaster taxon’.
The Lagerstätte-driven nature of the insect fossil record pre-
cludes definitive statements about the dominance of ‘weedy’ disaster
taxa after the end-Permian event. Disaster taxa are defined by their
occurrence across multiple assemblages and in multiple environ-
ments. The Madygen assemblage from southwestern Kyrgyzstan
is the oldest insect konzentrat Lagerstätte that follows the end-
Permian event and was deposited approximately 20 million years
after the extinction boundary (Fig. 1), when terrestrial ecosystems
are believed to have recovered (Chen and Benton 2012). Along with
the Ipswich assemblage from northeastern Australia, Madygen is
one of only two insect konzentrat Lagerstätten from the entirety
of the Triassic. The occurrence of the same species at both Triassic
Lagerstätten would be unlikely due to the 14.25 million years and
over 100° of paleolatitude that separate these assemblages. In the
event that co-occurrence of insect taxa were elevated between these
10
Lagerstätten, the two assemblages are insufficient to establish a
global pattern. Finally, insects’ poleward range shifts occurred on
a timescale that, in geochronological terms, was essentially in-
stantaneous (Shcherbakov 2008). Consequently, differences among
Lagerstätten probably reflect shorter-term climatic fluctuations as
much as they reflect biogeography and long-term changes in gamma
diversity.
Like the Triassic, the Late Permian contains two insect konzentrat
Lagerstätten. These intervals contain one Lagerstätte each from the
Northern and Southern Hemispheres, from relatively similar paleo-
latitudes: 40.05° to 40.26° north of the equator and 60.76° to
78.65° south of the equator (Fig. 1).
The NMDS plot comparing described insect communities of Late
Permian and Triassic Lagerstätten does not suggest a major family-
level extinction at the Permian/Triassic boundary: the Lagerstätten
are not arranged chronologically, nor are the Permian Lagerstätten
clearly separated from the Triassic Lagerstätten (Fig. 6). The Permian
Isady Lagerstätte is closer to the three other Lagerstätten than any of
these are to each other.
When the prevalence of different insect orders is visualized for the
Late Permian and the Triassic, no clear differences emerge (Fig. 6).
The clearest differences correspond not with time but with geog-
raphy. Various polyneopteran orders—the extinct order Reculida,
and the extant orders Plecoptera, Blattodea, Grylloblattodea, and
Phasmatodea—are better-represented among described occurrences
from the Northern Hemisphere Lagerstätten of Isady and Madgyen
than would be expected from extant diversity. In contrast, Belmont
and Ipswich, the Lagerstätten from the Southern Hemisphere, con-
tain far more described occurrences of Hemiptera.
Any pattern gleaned from a comparison of these Lagerstätten
should be treated with great caution. The four available Lagerstätten
represent a small fraction of insect diversity—especially for nearly
40 million years of evolutionary history. The limitations of the
fossil record for quantifying the impact of the end-Permian event
on insect diversity are exacerbated by the finding that insects from
this interval appear to have been most diverse, and most modern,
at tropical latitudes where they had the lowest probability of pres-
ervation (Shcherbakov 2008). Debate continues as to whether ob-
served changes in Permian insect assemblages are better ascribed to
Fig. 6.  Left: Nonmetric mulitdimensional scaling showing family-level differences in insect communities from Late Permian and Triassic konzentrat Lagerstätten,
each labeled with its age in Ma. Points and ellipses are color-coded by period, with the Permian in light red and the Triassic in purple. Each ellipse corresponds to
subsampled occurrences from the same locality following Schachat et al. (2020). Right: The proportions of described insect occurrences belonging to the orders
present in these Lagerstätten. The Lagerstätten are ordered from oldest (left) to youngest (right).
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
a decline in diversity or a change in habitat types where preservation
occurred (Shcherbakov 2008).
Plant–insect interactions provide a record of insect ecological di-
versity from habitats and intervals in which insect body parts are
not preserved (Labandeira 2013). Insect herbivory on fossilized
plants from the Late Permian through Middle Triassic has been
documented only rarely (Scott et  al. 2004, Krassilov and Karasev
2008, Prevec et al. 2009, McLoughlin 2011, Kustatscher et al. 2014,
Wappler et al. 2015, Bernardi et al. 2017, Cariglino 2020), and re-
markably few localities contain plants from both the Permian and
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Triassic. A recent study conducted with plants from the same geo-
graphic region found no decrease in Middle Triassic herbivory com-
pared to the Late Permian (Labandeira et al. 2016a). However, the
absence of Early Triassic (252–247 Ma) fossils from this region pre-
cludes strong conclusions about how its herbivores fared during the
end-Permian event.
Given the dearth of Late Permian Lagerstätten and the ab-
sence of any Lagerstätten from the Early–Middle Triassic, insect
diversity in the aftermath of the world’s greatest extinction event
remains poorly known. Nevertheless, the similarities between Late
Permian and Late Triassic insect communities—and the absence of
Palaeodictyopteroidea and Meganisoptera from both intervals—
suggests that the loss of insect diversity by the end of the Permian
was due to earlier biotic replacement.
For insects, the end-Permian event is best conceptualized as a
turnover event rather than a mass extinction (Fig. 7), analogous to
the late Trempealeauan eclipse at the end of the Cambrian Period
(Jun-yuan and Teichert 1983). The fossil record indicates that many
animal phyla originated in rapid succession around 540 million
years ago during the ‘Cambrian Explosion’ (Marshall 2006). For
decades, it was believed that early representatives of many phyla
were lost in a mass extinction at the end-Cambrian (Newell 1967).
However, as additional research was conducted with more precise
stratigraphic correlations (Palmer 1965) and abandonment of the
layer-cake stratigraphic concept (Labandeira 1986), it was realized
that these taxa went extinct slowly over many millions of years,
not suddenly in a single cataclysmic event. The concept of a mass
extinction therefore holds limited relevance to species loss toward
the end of the Cambrian because these losses occurred as a series
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
Paleozoic Insect Fauna Modern Insect Fauna
Origin of
Pterygota
End-Permian
eclipse
Fig. 7.  Turnover from the Paleozoic Insect Fauna (red) to the Modern Insect Fauna (blue) before and after end-Permian, in phylogenetic context. All icons are from
PhyloPic.org. Mantoida nitida Newman, 1838 (Mantodea: Mantoididae), license at http://creativecommons.org/publicdomain/mark/1.0/. Petrobius brevistylis
Carpenter, 1913 (Archaeognatha: Machilidae) by Birgit Lang, Somatochlora tenebrosa (Say, 1839) (Odonata: Corduliidae) by Gareth Monger, Palomena prasina
(Linnaeus, 1761) (Hemiptera: Pentatomidae) by Dave Angelini, and Trachypachus gibbsii Leconte, 1861 (Coleoptera: Trachypachidae) by T. Michael Keesey and
Yves Bousquet, license at http://creativecommons.org/licenses/by/3.0/.
of smaller extinctions (Palmer 1984, Westrop 1988, Rieboldt 2005,
Taylor 2006). Many other taxa that appeared to have gone extinct at
the end of the Cambrian were later found to have persisted into the
Ordovician for tens of millions of years after the presumed extinc-
tion event (Van Roy et al. 2010). This slow loss of ancient lineages,
barely 50 million years after their phyla originated, is now known as
the ‘late Trempealeauan eclipse’ (Fang et al. 2019).
Insect diversity near the end-Permian recapitulates the sequence
of events documented for the late Cambrian and can therefore be
described as a turnover event. Just as many animal phyla originated
in a geologic instant during the Cambrian, many orders of winged
insects appeared in quick succession around 320 million years ago
(Brauckmann and Schneider 1996, Prokop et al. 2005, Nel et al. 2013,
Schachat et  al. 2018, Haug et  al. 2015). Less than 75 million years
after these first appearances, entire orders, some comprising stem
representatives of higher taxa—from Paleoptera to Orthopteroidea
to Hemipteroidea to some Holometabola (Labandeira and Sepkoski
1993, Labandeira 2011, Prokop et al. 2017)—went extinct while the
Modern Insect Fauna came to overshadow the Paleozoic Insect Fauna.
As additional stratigraphic and paleontological data become avail-
able, it becomes even more clear that these extinctions of the Paleozoic
Insect Fauna occurred as the culmination of long declines, with some
taxa restricted to marginal habitats before going extinct (Shcherbakov
2008). These extinctions were neither simultaneous nor sudden.
The word ‘turnover’ conveys the slow twilight of the Paleozoic
Insect Fauna, which began long before the environmental crisis at
the end-Permian and continued long after ecosystems recovered.
This term differentiates the extinction dynamics of insects from the
rapid loss of marine biodiversity associated with the canonical end-
Permian extinction.
The End-Triassic Event
The end-Triassic event at 201 million years ago is known primarily
from marine fossils (Raup 1992, Tanner et al. 2004). Extinctions are
attributed to volcanism in the Central Atlantic Magmatic Province
(CAMP), which changed the chemical composition of the oceans
and atmosphere (Dal Corso et  al. 2014). The Triassic was a time
11
Holometabola
Hymenoptera,
Coleoptera,
Lepidoptera,
Diptera, etc.
Paraneoptera
Thysanoptera,
Hemiptera,
Psocodea
Polyneoptera
Orthoptera,
Downloaded from https://academic.oup.com/aesa/advance-article/doi/10.1093/aesa/saaa042/6056214 by guest on 11 January 2021
Dictyoptera, etc.
Paleoptera
Ephemeroptera,
Odonata
Apterygota
Present
of climatic extremes long before the onset of CAMP volcanism
(Whiteside et al. 2015). Near the temporal midpoint of the Triassic,
tens of millions of years before the end-Triassic event, the Carnian
Pluvial Event (approximately 235 Ma) evidently caused mega-
monsoons and seven degrees of global warming (Dal Corso et  al.
2015).
On land, tetrapods are the group whose fate during the end-
Triassic event has been most intensely studied. Controversy remains
as to whether dinosaurs gradually outcompeted non-ornithodiran
archosaurs throughout the Triassic or diversified during the Jurassic
after archosaurs were disproportionately extirpated at the end-
Triassic (Brusatte et al. 2010, Irmis 2010). A clearer consensus has
emerged for plants.
It appears that plants did not experience a mass extinction at
the end-Triassic. Plant communities changed drastically shortly be-
fore the end of the Triassic (Lindström 2016)—most likely due to
the environmental perturbations caused by volcanism—but these
changes appear transient, with no corresponding mass extinction at
higher taxonomic levels (Kürschner and Herngreen 2010, Bonis and
Kürschner 2012, Lucas and Tanner 2015). The severity of genus-
and species-level plant extinction at the end-Triassic varies greatly
with latitude (Lindström 2016). Acute, global ‘fern spikes’ indicate
severe disruption of plant communities at the end-Permian and end-
Cretaceous. In contrast, the ‘fern spikes’ near the end-Triassic are
patchy, and variable in duration (Lindström 2016). Plant communi-
ties changed near the end-Triassic, but not nearly as severely as they
did at the end-Permian or end-Cretaceous.
Migrating end-Triassic plant communities may have little in
common with the extinction that ravaged marine ecosystems at that
time. End-Triassic plant migrations are perhaps better understood
as analogous to the range shifts that occurred within Miocene and
Pliocene plant communities as C4 grasslands expanded and sub-
sequently migrated in synchrony with glacial–interglacial cycles
(Edwards et al. 2010). Climate change caused certain biomes to ex-
pand and others to contract but caused no increase in the family-level
extinction rate (McElwain and Punyasena 2007, Cascales-Miñana
and Cleal 2014).
12 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX

These Triassic shifts in plant communities caused by climate
change may be responsible for turnover among herbivorous tetra-
pods (Bond and Grasby 2017). Insects, however, appear to have been
less impacted by these shifts: previous paleoentomological studies
have not noted any mass extinction of insects at or near the end-
Triassic (Zherikhin 2002, Nel et al. 2018b).
When family-level similarities between described insect communities
of Triassic and Early Jurassic konzentrat Lagerstätten are visualized in
an NMDS plot, the Triassic Lagerstätten are easily distinguishable from
the Jurassic Lagerstätten (Fig. 8). However, both Triassic Lagerstätten
who survived the event may have had to seek shelter from environ-
mental hazards such as fire (Wolbach et al. 1988), acid rain (Ohno
et al. 2014), and extreme temperatures (Schulte et al. 2010). If pri-
mary productivity collapsed during an ‘impact winter’ scenario
immediately after the asteroid impact (Vellekoop et al. 2016), det-
ritivorous insects would have been the most reliable food source for
tetrapods (Sheehan and Fastovsky 1992), and insectivorous tetra-
pods tend to have smaller body sizes than co-occurring herbivorous,
omnivorous, and carnivorous tetrapods (Pineda-Munoz et al. 2016).
This trend of increased extinction risk for larger taxa at the end-

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are more similar to the Jurassic Sai-Sagul Lagerstätte than they are to
each other. As was the case for the end-Permian event (Fig. 6), the se-
verity of the end-Triassic event is obscured by the dearth of Triassic
Lagerstätten. Nevertheless, the NMDS plot suggests that the differences
that accumulated in insect communities before and after the extinction
event are no greater than would be expected given the tens of millions
of years that separate them.
At the level of the insect order, one of the most noticeable differ-
ences between the Triassic and Early Jurassic is that Diptera are more
prevalent among the described occurrences from all six Early Jurassic
konzentrat Lagerstätten than from either Triassic Lagerstätte (Fig. 8).
However, the Ipswich Lagerstätte of the Triassic is the only assem-
blage with a described beetle fauna whose prevalence approximates
that seen today. The Jurassic Lagerstätten contain fewer described
beetle occurrences than Ipswich and the younger faunas. The extinct
order Reculida is better-documented in the Posidonia Lagerstätte
from the Jurassic than in either of the Triassic Lagerstätten. It is not
immediately clear whether Jurassic entomofaunas are taxonomically
more modern than those of the Triassic.
The End-Cretaceous Event
The end-Cretaceous event, occurring at 66 Ma, perhaps is best known
for the extinction of nonavian dinosaurs. The event was triggered by
an asteroid impact whose environmental fallout would have made life
difficult for insects and plants as well as dinosaurs (Alvarez et al. 1980,
Schulte et al. 2010, Hull et al. 2020). However, insects seem not to have
struggled as mightily as dinosaurs and other tetrapods.
Tetrapods with larger body sizes had the greatest risk of extinc-
tion during the end-Cretaceous event (Archibald 1996). Individuals
Cretaceous also holds for relatively small-bodied tetrapod groups
such as snakes and lizards. The largest body size for both of these
groups shifted across the boundary from multiple kilograms to ap-
proximately 500 g (Longrich et al. 2012). However, the largest in-
sect that ever lived had an estimated body mass more than ten times
smaller than this, below 50 g (Dorrington 2016).
The small body size of insects as compared to tetrapods leaves no
reason to expect that larger-than-average insects would have faced
the elevated extinction risk that larger-than-average snakes and liz-
ards did. Any habitat that would provide sufficient shelter to smaller
snakes and lizards should also provide sufficient shelter to insects of
all sizes. And because detritivorous insects vary greatly in body size
(Ruesink and Srivastava 2001), a collapse of primary productivity
that favors smaller tetrapods for their insectivorous diet need not
favor smaller insects.
Heterotrophs of all sizes, from gigantic to unicellular, suffered
from elevated extinction rates at the end-Cretaceous (Hull et  al.
2011). Ecosystems recovered from the end-Cretaceous event much
faster on land than in the oceans (Beerling et al. 2001). Terrestrial
ecosystems were, nevertheless, impacted by the end-Cretaceous
event. In addition to the loss of large vertebrates, a ‘fern spike’ epit-
omizes an abrupt and dramatic shift in dominance of plant commu-
nities from angiosperms to ferns as terrestrial ecosystems collapsed
(Beerling et  al. 2001). Nevertheless, the end-Cretaceous does not
mark a regime shift in plant communities as it does for tetrapods
and marine organisms. Although plant communities changed at the
end-Cretaceous (Fastovsky and Bercovici 2016), such as the replace-
ment of mesic communities by mire communities on well-drained
substrates in North America (Tschudy et al. 1984), losses of higher

Fig. 8.  Left: Nonmetric mulitdimensional scaling showing family-level differences in insect communities from Triassic and Early Jurassic konzentrat Lagerstätten,
each labeled with its age in Ma. Points and ellipses are color-coded by period, with the Triassic in purple and the Jurassic in blue. Each ellipse corresponds to
subsampled occurrences from the same locality following Schachat et al. (2020). Right: The proportions of described insect occurrences belonging to the orders
present in these Lagerstätten. The Lagerstätten are ordered from oldest (left) to youngest (right).
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
taxa were strictly local and not a single plant family went extinct
(Cascales-Miñana and Cleal 2014).
Insects likely experienced the end-Cretaceous event similarly to
plants, with a momentary drop in alpha and beta diversity that did
not impact higher-level taxonomic diversity (Labandeira et al. 2002).
Unfortunately, the insect fossil record preserves minimal evidence of
insect diversity shortly before, during, and after the end-Cretaceous.
Canadian amber deposits from the provinces of Alberta and Manitoba,
the last insect konzentrat Lagerstätten that predate the end-Cretaceous
event, formed approximately ten million years before the Cretaceous
ended, and Menat, the first insect konzentrat Lagerstätte that postdates
the end-Cretaceous event, formed over five million years after the
Cretaceous ended (Fig. 1). By this time, plant communities and many
plant–insect interactions had unquestionably recovered (McElwain and
Punyasena 2007, Wappler et al. 2009).
Because the temporal distribution of konzentrat Lagerstätten
precludes estimates of insect diversity for the time when plants and
tetrapods reeled from the end-Cretaceous event, the next best exer-
cise would be to compare insect communities from Lagerstätten on
either side of the end-Cretaceous. Due to the nature of the insect
fossil record, this is not advisable either. The end-Cretaceous occurs
near the midpoint of a 16-million-year gap in the insect fossil record.
Another approach would be to calculate the average rate
of insect community turnover per 16 million years for the Late
Cretaceous and then compare this to the rate of turnover between
the last Cretaceous Lagerstätte and the first Paleogene Lagerstätte.
However, here the preservational modes of Lagerstätten would cause
a significant problem. All five of the Late Cretaceous Lagerstätten
are amber deposits whereas Menat is an adpression deposit con-
sisting of rocks formed at the bottom of a body of water (Fig. 1).
Insects preserved in adpression deposits such as Menat tend to be
less complete than those preserved in amber; fewer visible body
parts results in fewer taxonomic characters, which results in fewer
opportunities to identify a specimen’s affinities (Karr and Clapham
2015). This taphonomic bias can cause insect communities preserved
in adpression deposits to appear less diverse than they really were
(Karr and Clapham 2015). Furthermore, amber deposits come from
tree resin and adpression deposits come from aquatic settings, such
that these two types of deposits represent fundamentally different
habitats (Martínez-Delclòs et  al. 2004, Labandeira 2014a). Lastly,
amber deposits tend to preserve smaller insects than adpression de-
posits (Martínez-Delclòs et al. 2004, Labandeira 2014a).
In lieu of insect body fossils, the fossil record of plants is the only
direct source of information about how insects fared before, during,
and after the end-Cretaceous event. Analyses of insect herbivory
on leaves from various Cretaceous–Paleogene assemblages have
shown that specialized modes of herbivory, especially piercing and
sucking, mining, and galling, led to an increased risk of extirpation
(Labandeira et  al. 2016b). Plant–herbivore interactions recovered
on different timescales in different regions (Labandeira et al. 2002,
Wilf et  al. 2006, Wappler et  al. 2009, Donovan et  al. 2014), with
insect herbivores at localities more distant from the Chicxulub im-
pact crater faring better (Wappler et al. 2009, Donovan et al. 2017).
Where specialized herbivores appear to have been extirpated at the
end-Cretaceous, new herbivores with convergent feeding habits col-
onized localities within the same region on varying timescales (Wilf
et al. 2006, Donovan et al. 2014).
To summarize, not a single insect Lagerstätte from before the
end-Cretaceous can provide a meaningful comparison to Menat,
the only available insect Lagerstätte deposited within ten million
years of the end-Cretaceous event. Because entomofaunas changed
13
substantially during the Late Cretaceous, as insects adapted to radi-
ating angiosperm taxa that took over plant communities (Zherikhin
2002), the only insect Lagerstätte deposited within ten million years
of the end-Cretaceous event cannot be meaningfully compared with
any Cretaceous adpression Lagerstätten, which all predate the eco-
logical expansion of angiosperms (Fig. 1).
The End-Cretaceous Event as a Template for a
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Future Insect Extinction
Various methods for estimating insect diversity in deep time have
reconstructed a massive loss of family-level diversity near the end-
Cretaceous (Clapham et  al. 2016, Schachat et  al. 2019). However,
this apparent mass extinction is better attributed to the complete
absence of adpression konzentrat Lagerstätten from the entire Late
Cretaceous (100–66 Ma) fossil record of insects. The insect families
that had their last known appearance during the Early Cretaceous
(145–100 Ma) and those that had their first known appearance
during the Paleogene may well have coexisted. But if these families
have a low potential of being preserved in amber due to their habitat
preferences or a large body size, the fossil record will obscure their
coexistence—with an apparent extinction resulting as an artifact of
this gap in our knowledge.
An NMDS comparison of described insect communities of
Lagerstätten from the Late Cretaceous through early-Middle
Paleogene (Fig. 9) supports the consensus in the paleoentomological
literature that the end-Cretaceous was hardly catastrophic for in-
sects (Whalley 1987, Labandeira and Sepkoski 1993, Jarzembowski
and Ross 1996, Zherikhin 2002, Labandeira 2005). The clusters
of Late Cretaceous and Paleogene Lagerstätten (Fig. 9) do not sug-
gest a fundamental change in insect community composition at the
end-Cretaceous. For example, the Fushun amber Lagerstätte from
the Paleogene of eastern China is nearly indistinguishable from
the Oise amber Lagerstätte from France in terms of both age and
paleolatitude (Fig. 1), but the insect community described thus far at
Fushun is noticeably more similar to that of the Cretaceous amber
Lagerstätte from Manitoba, Canada than to that of Oise (Fig. 9).
The documented prevalence of particular insect orders within
konzentrat Lagerstätten does not change meaningfully from the
Late Cretaceous to early-Middle Paleogene (Fig.  9). The insect or-
ders Hemiptera, Hymenoptera, Coleoptera, and Diptera consistently
dominate these faunas. Along with Lepidoptera, which have an in-
famously depauperate fossil record (Sohn et al. 2015), these four or-
ders are by far the most diverse in modern ecosystems (Grimaldi and
Engel 2005). The clearest shift in the ordinal-level composition of
these Lagerstätten is the decline of Neuroptera at approximately 90
million years before the present. This is a well-documented phenom-
enon that has not been linked to an environmental crisis or to species
loss in any other taxon (Labandeira and Sepkoski 1993, Grimaldi
and Engel 2005).
There is little a priori reason to suspect that insects would suffer
a mass extinction during an event such as the end-Cretaceous, in
which not a single plant family went extinct (Cascales-Miñana and
Cleal 2014). Herbivorous insects that lived in distant habitats from
the impact crater, in places such as Patagonia, Argentina and Menat,
France, appear to have re-populated the communities that suffered
during the end-Cretaceous. Although Pangaea began to split apart
long before this time, the continents were not as widely dispersed
across the planet as they are now (Barron 1987) and latitudinal tem-
perature gradients were much gentler than they are today (Sewall
14 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX
Fig. 9.  Left: Nonmetric mulitdimensional scaling showing family-level differences in insect communities from Late Cretaceous through early-Middle Paleogene
konzentrat Lagerstätten, each labeled with its age in Ma. Points and ellipses are color-coded by period, with the Cretaceous in green and the Paleogene in
orange. Each ellipse corresponds to subsampled occurrences from the same locality following Schachat et al. (2020). Right: The proportions of described insect
occurrences belonging to the orders present in these Lagerstätten. The Lagerstätten are ordered from oldest (left) to youngest (right).
and Sloan 2001). These factors considerably reduced impediments
to insect dispersal.
Conclusions
1. Changes in the fossil record of insect diversity are currently
understood only at the levels of family and order. This lack of
genus- and species-level resolution can be attributed to several
factors. First is insufficient support for taxonomic studies of
fossil insects. Second is the difficulty of tracking a single genus
through time, caused by the infrequency with which insects are
preserved as fossils. Third is the rarity and inaccessibility of in-
sect morphological characters, such as genitalia, used to distin-
guish genera and species.
2. It is safe to assume that numerous families of insects would have to
go extinct for the anthropogenic biodiversity crisis to leave its mark
in the fossil record. The current decline of insects is primarily meas-
ured by the loss of abundance and biomass. Because the insect fossil
record is unsuited to estimates of these two metrics, it is essentially
impossible to compare the current plight of insects with mass insect
extinctions in deep time.
3. The five major extinction events identified in the fossil record
of insects—the Late Pennsylvanian, end-Permian, Late Jurassic,
later Early Cretaceous, and end-Cretaceous—are best under-
stood as examples of turnover, rather than as mass extinction
events. Major groups of insects went extinct during each of these
events, but these events were not sudden, and replacement of
the extinct insect groups occurred quickly. Two major insect
losses overlap with Big Five mass extinctions, the end-Permian
and end-Cretaceous, but both commenced well before the en-
vironmental crises associated with these extinctions. The other
Big Five mass extinction that occurred after the origin of winged
insects, the end-Triassic, is long recognized to have no associa-
tion with any large-scale change in the entomofauna.
4. Insects fare better than tetrapods and marine invertebrates during
mass extinctions. The small body size of insects, their unparal-
leled abundance, and their ability to enter diapause may give
them an advantage over tetrapods during environmental crises.
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The relatively mild effects of increased pCO2 and decreased pO2
in the atmosphere, compared to the oceans, shield insects from
many of the physiological challenges that marine invertebrates
face during catastrophic climate change. The extinction history
of insects is most comparable to that of plants. The end-Permian
is the only one of the Big Five mass extinctions that coincides
with global losses at or above the level of family; all other ex-
tinction events appear to have been local or restricted to lower
taxonomic levels.
5. Insects do not appear to have ever suffered a sudden mass ex-
tinction. If the current biodiversity crisis extinguishes diverse
higher-level clades, then perhaps humans will have caused
insects’ first true mass extinction. However, at present, there are
no indications that we are anywhere near a crisis of this severity.
Acknowledgments
C. Kevin Boyce, William A.  DiMichele, S.  Augusta Maccracken, Pedro
M.  Monarrez, Jonathan L.  Payne, Pulkit Singh, and three anonymous re-
viewers provided valuable feedback on this manuscript. Matthew E. Clapham
compiled the fossil insect dataset downloaded from the Paleobiology Database.
SRS received funding from the Coleman F.  Fung Interdisciplinary Graduate
Fellowship, Vice Provost of Graduate Education, Stanford University; from
the Harriet Benson Fellowship Award, Department of Geological Sciences,
Stanford University; from the McGee/Levorsen Research Grant, Department
of Geological Sciences, Stanford University; and from the Winifred Goldring
Award, which is a joint initiative of the Paleontological Society and the
Association for Women Geoscientists. This is contribution 375 of the
Evolution of Terrestrial Ecosystems consortium at the National Museum of
Natural History in Washington, D.C.
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