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POLYNEOPTERA 217
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7.44. The caloneurodeans, such as Paleuthygramma tenuicornis (Paleuthygrammatidae) from the Permian of
Russia, were enigmatic relatives of early Orthoptera and Phasmatodea that became extinct probably at the end
of the Permian. Length 24 mm; redrawn from Carpenter (1992).

and unique folding mechanism; a prognathous head (lacking a

CALONEURODEA: THE CALONEURODEANS
Little is known of this extinct Paleozoic order of poly-
neopteran insects (Figure 7.44). The order is noteworthy for
the secondary loss of the anal area in the hind wing, the
unbranched and nearly parallel (or fused in a few groups)
veins CuA and CuP in both the fore- and hind wings, and,
where known, unsegmented cerci. Other features of the order
include the strongly convex and concave wing veins; five-
segmented tarsi; long, multisegmented antennae; and fore-
and hind wings having a similar shape, venation, and texture.
Presently there are nine families, many with a single genus,
and the systematics of the group requires considerable revi-
sion. Nothing is known of caloneurodean biology aside from
the fact that they were terrestrial, apparently primitively
resembling cursorial orthopterids. The caloneurodeans are
known only from the Late Carboniferous and Permian.
Copyright 2005. Cambridge University Press.
gula); absence of ocelli; a subgenital plate formed by an
enlarged seventh sternum in females; and a vestigial ovipositor.
The uniquely expanded anal fan of the hind wing may eventu-
ally prove to be independently derived from that of other poly-
neopterans, in which case Dermaptera would be classified
elsewhere. Earwigs are overflowing with unique features and
the chronology of these is even apparent in the fossil record.
Dermaptera are distributed globally except Antarctica and
the extreme Arctic, but most of the nearly 1,900 described
species occur in tropical to warm-temperate habitats (e.g.,
Steinmann, 1986, 1989a,b,c, 1993) (Figure 7.46). Earwigs typ-
ically live in riparian habitats, in crevices, in leaf litter, or
under the bark of trees. Most species are nocturnal and
omnivorous; only a very few species are strictly herbivorous
or carnivorous (Chopard, 1938). The cercal forceps are used
to capture prey and are employed in mating and in folding
the hind wings under the tegmina (Kleinow, 1966). Female

earwigs exhibit extended maternal care over the eggs and

DERMAPTERA: THE EARWIGS
Almost anyone can immediately recognize an adult earwig
with its stout, terminal forceps (Figure 7.45) and the short,
tegminous forewings. Less conspicuous are the three-
segmented tarsi; very distinctive, greatly expanded hind wing
vannus (Figure 7.50), with a dramatically reduced remigium
early instar nymphs, carefully cleaning them to protect from
invasive fungi (Herter, 1943; Rentz and Kevan, 1991). After
two molts, however, nymphs must fend for themselves, or
they will be eaten by the mother.
Earwigs have a long geological history. The oldest fossils to
date are tegmina from the Late Triassic–Early Jurassic of
England and Australia (e.g., Jarzembowski, 1999). As already

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218
7.45. The modification of cerci into forceps is the most recognizable
trait of the earwigs (order Dermaptera). The forceps are used to
defend, to capture prey, and to assist in folding the fanlike wings under
the short tegmina. Not to the same scale.
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EVOLUTION OF THE INSECTS
discussed and what should be of little surprise, some traits
believed by neontologists to be diagnostic for an earwig do
not apply to the earliest earwigs. Basal, extinct members of
the order all had five-segmented tarsi, well-developed
ovipositors, veined tegmina, and long, multisegmented cerci.
Traditionally, primitive earwigs from the Late Jurassic and
Early Cretaceous were classified in the suborder Archi-
dermaptera (e.g., Bei-Bienko, 1936; Vishniakova, 1980;
Carpenter, 1992), a paraphyletic stem group to modern
Dermaptera. Archidermaptera in a restricted sense contains
the Jurassic–Cretaceous families Protodiplatyidae, Turanovi-
idae, and Dermapteridae, which comprise the basalmost lin-
eage of the order (Willmann, 1990a; Haas and Kukalová-Peck,
2001; Engel, 2003c) (Figure 7.47). Archidermapterans are a
sister group to the Pandermaptera, which comprise two fur-
ther suborders: Eodermaptera, for the Jurassic-Cretaceous
families Semenoviolidae and Turanodermatidae, and the
Neodermaptera, which contains all the modern lineages.
Eodermapterans share with Neodermaptera the derived
development of unsegmented, forcipate cerci but primitively
retain venation in their tegmina, presence of ocelli, and pen-
tamerous tarsi. The Neodermaptera have three-segmented
tarsi, no ocelli, and lost venation in their tegmina. They first
appear in the Early Cretaceous (e.g., Popham, 1990; Engel
et al., 2002) but may have originated in the latest Jurassic
since there is a putative, undescribed pygidicranoid from the
Jurassic of central Asia (Rasnitsyn and Quicke, 2002). Cer-
tainly, definitive neodermapterans (Figure 7.48) and recog-
nizable pygidicranids are known by the mid-Cretaceous
(Grimaldi et al., 2002; Engel and Grimaldi, 2004c) (Figure
7.49). Tertiary earwigs, mostly of the Forficulidae, are pre-
served in several deposits, and a relatively unexplored diver-
sity of species is known in Baltic (Burr, 1911), French (Nel
et al., 2002c); and Dominican ambers (Figure 7.51).
The internal phylogeny and classification of Neo-
dermaptera has been in constant flux, with dramatically
different arrangements of families and superfamilies by con-
temporaneous authors. Recent phylogenetic work has begun
to shed light on relationships within the suborder (e.g., Haas,
1995; Haas and Kukalová-Peck, 2001; Colgan et al., 2003)
(Figure 7.51). Within Neodermaptera, the infraorder Proto-
dermaptera, including the superfamilies Pygidicranoidea
and Karschielloidea, is basal but perhaps paraphyletic (e.g.,
Haas, 1995; Haas and Kukalová-Peck, 2001). Unlike all other
earwigs, the protodermapterans have ventral cervical scle-
rites of equal size, carinae on the femora, and a segmented
pygidium. Almost all Cretaceous records of Dermaptera are
protodermapterans. The infraorder Epidermaptera, includ-
ing all other Neodermaptera, is characterized by the enlarge-
ment of the posterior ventral cervical sclerite, rounded
femora, and fusion of the three pygidial sclerites (Popham,
1985).
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POLYNEOPTERA 219

7.46. Representative Recent earwigs. Assembled from Genera Insectorum.

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220 EVOLUTION OF THE INSECTS

Although two additional suborders (or infraorders) have

been traditionally recognized for the parasitic earwigs, both
are actually derived Epidermaptera. The families Hemimeri-
dae and Arixeniidae are ectoparasites with a suite of paedo-
morphic characters (i.e., the retention of nymphal traits in
the adult). The Hemimeridae is perhaps closely related to
Apachyidae (Klass, 2001b), while Arixeniidae is related to, if
not derived from within, Spongiphoridae (Popham, 1985).
Hemimerids live on murid rodents in Africa. Two genera are
recognized: Hemimerus, which consists of nine species living
on Cricetomys rats, and Areomerus, which consists of two
species living on Beamys rats (Rehn and Rehn, 1936; Nakata
and Maa, 1974). The five species of arixeniids live in the
roosts of Cheiromeles bats (Molossidae) in southeast Asia,
where they feed on secretions from the skin or on other
insects invading the roost (Medway, 1958; Nakata and Maa,
1974). Hemimerids were at one time considered a distinct
order, called Diploglossata, Dermodermaptera, or Hemime-
rina (e.g., Verhoeff, 1902; Brues and Melander, 1915; Popham,
1961). As with many ectoparasitic insects, fossil Hemimeri-
dae and Arixeniidae have yet to be discovered, and the fami-
lies are probably no older than the mid-Tertiary.
Dermaptera are believed by some to stem from the Per-
mian order Protelytroptera owing to tegminous forewings
and the large, uniquely formed anal fan (Kukalová-Peck,
1991; Haas and Kukalová-Peck, 2001). Some families have
erroneously been placed in Protelytroptera, specifically the
Cretaceous Umenocoleidae, which is actually a highly spe-
cialized lineage of roaches (Figures 7.70, 7.71). The possibility
exists that Permian Protelytroptera were earwig progenitors,
but definitive evidence is still required to establish this.

7.47. An archidermapteran, Microdiplatys campodeiformis (Pro-

todiplatyidae), from the Late Jurassic of Karatau in Kazakhstan. Archi-
dermaptera lacked the cercal forceps and primitively retained some
veins in the forewings, though the tegminous forewings were short as
in modern earwigs. PIN 2904/441; length, excluding cerci, 10 mm.

7.48. (right) A neodermapteran earwig from the Early Cretaceous of

Brazil. AMNH; length 5 mm.

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POLYNEOPTERA 221

7.50. Phylogenetic relationships among major lineages of earwigs.

Significant characters indicated in Table 7.2. Relationships based on
Haas and Kukalová-Peck (2001). Thick dark lines are the known extent
of fossils, lighter thick lines indicate fossils possibly belonging to those
groups.

TABLE 7.2. Significant Characters in Dermaptera

Phylogenya

1. Cerci unsegmented, forcipate

2. Trimerous tarsi
3. Ocelli lost
4. Tegminal veins lost
5. Ovipositor reduced
6. Posterior, ventral cervical sclerite enlarged
7. Three pygidial subsegments fused
8. Reduction to single penal lobe and single virga
9. Expanded regions of anal and intercalary veins distinctly
separated
a
Numbers correspond to those on phylogeny, Figure 7.50.

7.49. The earliest pygidicranoid earwig, Burmapygia resinata (Pygidi-

cranidae), in mid-Cretaceous amber from Myanmar. Pygidicranids are
among the most primitive of living earwigs. AMNH Bu274; length 6
mm; from Engel and Grimaldi (2004c).

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222 EVOLUTION OF THE INSECTS

All rights reserved. May not be reproduced in any form without permission from the publisher, except fair uses permitted under U.S. or applicable copyright law.

7.51. A forficulid earwig in Miocene amber from the Dominican Republic. The hind wings of earwigs are
remarkably distinctive, with most of the veins fused at the base and a large anal fan comprising most of the
wing. Morone Collection, M3400; body length 6 mm. Photo: R. Larimer.

deviate little from freezing (Atchison, 1979). These insects are

GRYLLOBLATTODEA: THE ICE CRAWLERS
Grylloblattodea (or Notoptera) represent an interesting, relict
lineage today confined entirely to the Northern Hemisphere.
In a class renowned for its overwhelming diversity, the Gryl-
loblattodea, along with Zoraptera and Mantophasmatodea,
hold the distinction of being the least diverse of insect orders.
Today there are 26 species classified into five genera within a
single family – Grylloblatta from the northwestern United
States and southwestern Canada; Grylloblattella and Gryl-
loblattina from the Russian Far East; Galloisiana from
Japan, Korea, China, and Russia; and Namkungia from
Korea but which probably makes Galloisiana paraphyletic
(Storozhenko, 1997, 1998; Storozhenko and Park, 2002).
These soft-bodied, wingless insects are typically found in leaf
litter or under stones in cold temperate forests, often at
higher elevations, although some blind Asian species have
been discovered in caves (Namkung, 1982; Nagashima, 1990).
Copyright 2005. Cambridge University Press.
omnivorous and typically scavenge dead arthropods but rely
on plant material when frozen carcasses become scarce
(Pritchard and Scholefield, 1978; Nagashima et al., 1982).
Modern Grylloblattodea have numerous defining fea-
tures, such as a median, eversible sac on the first abdominal
sternum; the loss of ocelli (also in Mantophasmatodea and
perhaps shared through common ancestry in these two line-
ages); and the asymmetrical male genitalia. Unique among
all hexapods is the presence of a spina on the metathoracic
sternum. Although difficult to place in the greater scheme of
insect phylogeny, ice crawlers are probably basal members of
Orthopterida and were also thought to be “living fossils” even
by their discoverer (Walker, 1914, 1937). Wings are absent in
modern grylloblattids and other characteristics are generally
primitive: the five-segmented tarsi; long, multisegmented
cerci; and an ovipositor composed of three stout pairs of
“valvulae” intermediate to the orthopterid ovipositor. The

Species are active at cold temperatures, and several studies
have indicated optimal temperatures around 1–4°C (Mills
and Pepper, 1937; Henson, 1957) (Figure 7.52). Although ice
crawlers prefer low temperatures, they are not impervious to
freezing. Individuals of Grylloblatta can be killed by ice for-
mation within the body at around8°C (Morrissey and
Edwards, 1979). During winter months when night tempera-
tures drop below freezing, ice crawlers likely survive under
the snow-pack and near the soil where temperatures may
third pair of valvulae are, in fact, the gonoplacs. In Grylloblat-
todea the gonoplac is greatly developed and incorporated
into the ovipositor, representing an intermediate stage
among orthopterids where the function of the reduced, sec-
ond gonapophyses is assumed by the gonoplac. Gonoplac
development in ovipositor construction is a typical
orthopterid feature.
The presence of enlarged coxae, which is typical of basal
Dictyoptera, has at times been used to support a relationship

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