Systematic Entomology (2012), 37, 223–228 DOI: 10.1111/j.1365-3113.2011.00602.

A new genus and species of scorpionfly (Mecoptera)

from Baltic amber, with an unusually developed
postnotal organ
1 2
W I E S Ł A W K R Z E M I Ń S K I and A G N I E S Z K A S O S Z Y Ń S K A - M A J
1
Institute of Systematic and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland and 2 Department of Invertebrate
Zoology and Hydrobiology, University of Łódź, Łódź, Poland

Abstract. A new genus and species of fossil scorpionflies (Mecoptera) Baltipanorpa

damzeni gen. et sp.n. is described from two well-preserved male specimens in Baltic
amber (middle Eocene: Lutetian). The most characteristic feature of the new taxon
is an unusually developed postnotal organ on abdominal tergum IV. This is the most
extremely developed example of this organ among Mecoptera and the only observation
of notal and postnotal organs among fossil scorpionflies. The following combination of
characters are provided to distinguish the new genus from other Panorpidae: Sc, short;
R1 and R2 two-branched; A1 joins posterior margin of wing only at same level as fork
of vein Rs; unusual shape of abdomen, abdominal segments I–IV strongly reduced,
abdominal segment V elongate and widened, segments VII and VIII strongly elongate;
notal and postnotal organs present, strongly developed process (postnotal organ) on
tergum IV, unknown in all described extant and fossil scorpionflies. Different types
of notal organs of Mecoptera are compared and their function and morphology are
discussed. Morphological analysis of notal and postnotal organs in extant species
permits us to conclude that B. damzeni sp.n. is characterized by the most developed
and complex notal organs in all Mecoptera.

Introduction element of these ecosystems, being at least three times more

Mecoptera, a small, holometabolous order of insects, com-
prises about 600 extant species grouped in nine families
(Cai et al., 2008; Bicha, 2010), which occupy all regions
of the world except Antarctica. The morphology and scav-
enging habits of both adults and larvae are primitive for
Holometabola (Grimaldi & Engel, 2005). The largest fam-
ily is Panorpidae with approximately 420 species, assigned
to four genera: Panorpa Linnaeus; Neopanorpa Weele; Lep-
topanorpa MacLachlan; and Sinopanorpa Cai & Hua (Cai
et al., 2008).
Mecoptera had a great share in faunas from the Late
Permian through the Cretaceous, and constituted a significant
Correspondence: Agnieszka Soszyńska-Maj, Department of Inver-
tebrate Zoology and Hydrobiology, University of Łódź, ul. Banacha
12/16, 90-237 Łódź, Poland. E-mail: agasosz@biol.uni.lodz.pl
widespread and diverse during the Mesozoic than at present
(Krzemiński & Krzemińska, 1996, 2003; Hoell et al., 1998;
Novokshonov, 2002). The level of extinction within this
group is unusual among holometabolous insects, although
it is somewhat mirrored among the Neuropterida (Willman,
1989; Grimaldi & Engel, 2005). Extant scorpionflies are
mostly carnivorous, but examination of fossil species indicates
that extinct Mecoptera were possibly also important plant
pollinators (Ren et al., 2009).
Baltic amber dates from the Lower Eocene–Ypresian
(50 Ma) to Lutetian (40 Ma). Until now representatives of
three families were found in Baltic amber (Weitschat &
Wichard, 2002): Bittacidae (five species; Krzemiński, 2007);
Panorpidae (two species); and Panorpodidae (two species;
Carpenter, 1954, 1955, 1976). The only known species of
Panorpidae in Baltic amber belong to the widely distributed
and abundant genus Panorpa: Panorpa obsoleta Carpenter,
1954 and Panorpa mortua Carpenter, 1954 (Carpenter, 1954).

© 2011 The Authors

Systematic Entomology © 2011 The Royal Entomological Society 223
224 W. Krzemiński and A. Soszyńska-Maj
Most Mecoptera (especially Panorpidae) are characterized
by interesting sexual and mating behaviour, including nuptial
feeding, pheromone emission or forced copulation (Thornhill,
1973, 1980, 1981, 1990; Thornhill & Sauer, 1991; Kock
et al., 2009). The structures important for the sexual behaviour
of some panorpids are processes occurring in the posterior
area of the third and fourth abdominal tergites of the males.
These used to be called, respectively, notal (= notorganus,
notorgan) and postnotal organs (Crampton, 1931; Mickoleit,
1971), ‘upper/lower part of the notal organ’ (Chu & Byers,
1978), wing-clasping organ (Penny, 1975) and clamp-like
structure (Byers & Thornhill, 1983), or just notal organ
(Thornhill, 1991). The basic role of the organ is to hold the
wings of the female, thereby prolonging or forcing copulation
(Byers & Thornhill, 1983; Thornhill, 1990; Thornhill & Sauer,
1991; Kock et al., 2009). However, Felt (1896) proposed that
the notal organ was a source of male pheromone licked up
by females (after Crampton, 1931). This was later rejected
by Mickoleit (1971) and Thornhill (1973), who found that
the pheromone gland is positioned in the genital bulb. The
process of the third tergum is sclerotized, projects backwards
and its length differs in individual species from half of
one tergite to even several tergites of the abdomen. The
second process located on the fourth tergum is always small,
more sclerotized than the former process, conical or hook
shaped (Byers & Thornhill, 1983; Thornhill, 1990). This
organ was found in four families of Mecoptera: Eomeropidae
(= Nothiothaumidae), Panorpidae, to a limited extent in the
Panorpodidae (strongly reduced) and in a few European species
of Boreidae (a rudimentary process, only on third tergite)
(Mickoleit, 1971; Penny, 1975). The greatest development of
these structures is observed in the ‘living fossil’ scorpionfly
Notiothauma reedi MacLachlan (Eomeropidae) and in the
family Panorpidae, with the exception of Sinopanorpa, where
this organ is not developed (Mickoleit, 1971; Cai et al.,
2008).
The first discovery of exceptionally well-developed notal
and postnotal structures in fossil Mecoptera is presented here,
along with the description of a new monotypic genus and
species of Panorpidae from Baltic amber.
Material and methods
The study was based on two inclusions in Baltic amber (dated
to the middle Eocene, ∼45 Ma, Lutetian) deposited in the
museum collection of the Institute of Systematic and Evolution
of Animals, Kraków, Poland (MP ISEA). The specimens were
studied with the use of a Leica MZFLII stereomicroscope,
under reflected and transmitted light. Photographs were made
with a Leica DFC295 camera attached to the microscope.
Drawings were made from the photographs. The terminology
of wing venation, the notal organ and genitalia follows that
of Chau & Byers (1978), Crampton (1931) and Webb et al.,
(1975), respectively.
Systematic Entomology © 2011 The Royal Entomological Society, Systematic Entomology, 37, 223–228
Systematic paleontology
Order Mecoptera Hyatt & Arms
Family Panorpidae Stephens
Genus Baltipanorpa gen.n.
Type species. Baltipanorpa damzeni gen. et sp.n.
Diagnosis. Both wings about as long as abdomen. Forewing
Sc short, not extending beyond one-half wing length; R1
forked before pterostigmal band to R1a and R1b ; R2 two-
branched; A1 joins posterior margin of wing only at same
level as fork of vein Rs; three crossveins between A1
and A2 present. Abdominal segments I–IV strongly reduced;
terga III and IV with long, narrow processes, extending to
two-thirds the length of segment VII, both set together and
forming a functional entity; process on tergum IV curved
upwards at apex; abdominal segment V elongate and widened;
segments VII and VIII strongly elongate, very thin at base,
in second half strongly widening and sinusoidally curved;
tergum VII in dorsal part produced in wide lobe; segment VIII
widened to a goblet shape, without additional lobe.
Etymology. Combination of Balti- for the occurrence of the
new genus in Baltic amber and Panorpa, type genus of the
family Panorpidae. The name is feminine.
Baltipanorpa damzeni gen. et sp.n.
(Figs 1, 2)
Material examined. Holotype: male, no. MP/3076; well
preserved and complete, except for the wings, which are
crumpled and the venation is partially poorly visible. Paratype:
male, no. MP/1/4/552/04; specimen without head, remainder
well preserved, wings complete with clearly visible venation,
except for the distal part. Both specimens in Baltic amber are
housed in the ISEA, Kraków, Poland.
Description. (Figs 1, 2). Male. Body length ∼16 mm.
Head: eyes round, separated by width of rostrum, bare,
facets of equal size, three ocelli present; antennae filiform
composed of 40 segments equal in length, covered by short,
thick setae; first flagellomere as long as next four combined,
last flagellomere rounded; scape wide and short, pedicel thicker
and rounded; rostrum long, narrow, typical for Panorpidae;
palpi not visible.
Wing venation: general model of wing venation similar to
venation of Panorpa; forewings (Fig. 3) ∼15 mm long, as long
as abdomen, membrane slightly infuscate, with clearly visible
darker apical and pterostigmal bands; Sc short, not extending
beyond one-half wing length; crossvein sc-r at four times its
length before tip of Sc; R1 forked before pterostigmal band into
R1a and R1b ; one crossvein between R1 and R2 and second
crossvein between R1 and R2 – 3 ; R2 two-branched; A1 joins
© 2011 The Authors

Fig. 1. Baltipanorpa damzeni gen. et sp.n. – holotype.
Fig. 2. Baltipanorpa damzeni gen. et sp.n. – abdomen of paratype.
posterior margin of wing only at same level as fork of vein
Rs; three crossveins between A1 and A2 (Fig. 4A, the complete
reconstruction of wing venation is impossible because only the
basal part of the forewings and cubital and anal parts of the
hindwings are well preserved; Fig. 4B).
Legs: long, tibia with two spurs, basitarsus a little shorter
than remaining tarsomeres, second tarsomere almost 1.5× as
© 2011 The Authors
Systematic Entomology © 2011 The Royal Entomological Society, Systematic Entomology, 37, 223–228
Baltipanorpa damzeni gen. et sp.n. 225
Fig. 3. Distal part of forewing of paratype of Baltipanorpa
damzeni gen. et sp.n.
Fig. 4. Anal parts of wings of holotype of Baltipanorpa
damzeni gen. et sp.n.: A, forewing; B, hindwing.
long as succeeding tarsomeres combined, a row of spines at
distal part of all tarsomeres; two terminal, pretarsal claws
visible.
Abdomen (Fig. 5): abdominal segments I–IV narrow; pro-
cess on segment III (notal organ) strongly widened at base,
projecting backwards, distal half with numerous short spines
on inner surface and four pairs of thin, long setae; first
seta longest, following pairs gradually shorter, last pair only
insignificantly longer than width of tergite III process; seg-
ment IV with long, thin and bare process (postnotal organ);
both processes almost equal in length, tergum IV process a lit-
tle longer and curved upwards at apex, abdominal segment V
elongate and widened, corresponding sternite almost one-third
shorter than tergite V; sternites and tergites of remaining seg-
ments fused; segment VI narrow in basal part, much widened
distally, dorsal part elongated in wide lobe; part of segment VI
inside segment V in holotype, clearly visible in paratype; seg-
ments VII and VIII strongly elongate, considerably thinner at
base, in distal half strongly widened and sinusoidally curved;
segment VII in dorsal part elongate in wide lobe, almost as
in previous segment; segment VIII widened in goblet shape,
without additional lobe; genital bulb elongate, very narrow at
base, which looks like pedicel; gonostylus in male genitalia
short and wide; inner structures of male genitalia not visible.
Diagnosis. The same as for the genus.
226 W. Krzemiński and A. Soszyńska-Maj
Fig. 5. Abdomen, lateral view, of Baltipanorpa damzeni gen. et sp.n.
Etymology. The specific name is derived from Jonas
Damzen, who found both inclusions and recognized their
importance.
Discussion
Baltipanorpa gen.n. differs significantly from all known extant
and fossil Mecoptera. The following combination of charac-
ters enable us to distinguish the genus from all other described
genera of Panorpidae: differences in wing venation in anal and
radial part (Figs 3, 4); distinctive shape of abdomen, abdominal
segments I–IV strongly reduced, abdominal segment V elon-
gate and widened, segments VII and VIII strongly elongate
(Fig. 5); strong development of postnotal organ on tergum IV
(which is even longer than notal organ on tergum III), unlike
anything observed in any previously described scorpionflies.
Characteristic differences in wing venation are present in
anal and distal parts. Short Sc in forewings, not extending
beyond one-half wing length, is similar to Panorpa alpina
(Rambur), and is in opposition to other Panorpa species and
other panorpid genera. Baltipanorpa gen.n. has two-branched
R1 , whereas in several previously described genera R1 is not
forked. R2 is two-branched, as in Panorpa, rather than three-
branched, as in Sinopanorpa (Cai et al., 2008; Tillier, 2008).
A1 joins posterior margin of wing only at same level as fork of
vein Rs, as in Neopanorpa, in a different way to Sinopanorpa
and Panorpa. There are three crossveins between A1 and A2
in Baltipanorpa gen.n., whereas in existing genera only one
(Neopanorpa) or two crossviens (Sinopanorpa and Panorpa)
are described (Cai et al., 2008).
In extant Mecoptera the posterior median process of the third
tergum is sclerotized, broad and flat in Panorpa, or elongate
and rod-shaped in Neopanorpa, and projects backwards (Issiki,
1933). Crampton (1931) proposed that this structure elongated
during the evolution of the Mecoptera. He suggested that
this is an orthogenetic tendency that probably started from
the most primitive and archaic scorpionfly, Notiothauma reedi
Systematic Entomology © 2011 The Royal Entomological Society, Systematic Entomology, 37, 223–228
MacLachlan, where the notal organ is a pincer-like structure. It
would reach its culmination in Panorpa takenouchii Miyake,
which has the longest notal organ among extant species.
Panorpid genera, as well as species, differ in the length of
the notal organ (Fig. S1). In European species of Panorpa the
notal organs are slightly produced and barely visible or even
completely reduced in Panorpa cognata Rambur (Engqvist &
Sauer, 2002). However, a few Panorpa species from eastern
Asia have very long processes, reaching almost the end
of abdomen Panorpa leucoptera (Uhler) and P. takenouchii
(Issiki). Leptopanorpa is characterized by a well-developed
but small, clamp-like structure (Lieftinck, 1936), and the
organ is not developed in Sinopanorpa (Cai et al., 2008). In
general, the longest notal organs in Mecoptera are found in
Neopanorpa spp., which was an important reason to separate
them from Panorpa (Weele, 1909), although the monophyly
of these groups relative to each other requires rigorous testing.
Neopanorpa species from the Oriental region are characterized
by short notal organs, reaching to only one half of the fourth
abdominal segment, e.g. in Neopanorpa byersi Webb & Penny
(Webb & Penny, 1979), whereas the longest reaching almost
to half of the seventh segment are observed in Neopanorpa
subreticulata Miyamoto & Makihara (Miyamoto & Makihara,
1979; Fig. S1). In the latter it is a bit shorter than in
B. damzeni sp.n.
In extant scorpionflies the second process (postnotal organ)
located on the fourth tergum is always very small, more
sclerotized than the notal organ, conical or hook shaped and
projects upwards. In species with very elongate processes of
the third tergum, the postnotal process is usually flat, barely
visible and reduced or only present as a small elevation with
a dense set of setae, as in P. leucoptera and P. takenouchii
(Issiki, 1933). No examples of any elongation of the process
on the fourth tergite appear in extant Mecoptera. The elongate
process on abdominal tergite IV, reaching two-thirds of
tergum VII, is only known from B. damzeni sp.n., and is an
unusually developed postnotal organ among Mecoptera.
As mentioned earlier, the function of notal and postnotal
organs is generally related to copulation and mating behaviour.
This was described in detail for Panorpa spp. (Mickoleit,
1971; Thornhill, 1973, 1980, 1981, 1990; Thornhill & Sauer,
1991). During mating in the European species Panorpa
communis Linnaeus, the male and female forms a V-shaped
position. The male first grasps the female’s leg or wing with
his genital claspers, and then repositions her to catch the
costa of the hindwing with his notal organ (Mickoleit, 1971;
Byers & Thornhill, 1983). It was proved that the function of
notal and postnotal organs is to enforce (without nuptial gift
or salivary mass) and prolong copulation (with nuptial gift or
salivary mass) to transfer more sperm, minimize the costs of
salivary mass production and avoid interruption of copulation
by an intruder. Between the third and fourth segments there
are muscles bringing the two processes together (Thornhill &
Sauer, 1991). Kock et al. (2009) proposed that the function
of the notal organ is to initiate and stabilize females during
mating in the V-shaped position. When the clamp-like structure
is reduced (as in P. cognata) the mating strategy is different.
© 2011 The Authors

The pre-mating period is prolonged to ensure that the female
is motivated to mate, the male looks for a sheltered location
and only one large salivary mass is produced to occupy the
female and prolong copulation (Engqvist & Sauer, 2002).
It is easy to imagine how males of species with small
notal organs, like P. communis or N. byersi catch the female’s
forewing costa using the notal organ as a pincer. It is more
difficult to understand how such elongated notal organs in
Mecoptera (as in Neopanorpa species or in P. takenouchii )
operate if the process on the fourth tergite is usually barely
developed. Issiki (1933) stated that the male can hold
the wings of females between the elongate notal process
and fourth tergite. The only description of other species
using the elongated organ comes from Thornhill (personal
communication), who observed flying males of species with
very elongated notal organs grabbing stationary females before
mating. In B. damzeni sp.n. the situation is more complicated
because not only is the process of the third tergite long,
but also the process on the fourth abdominal segment is
strongly elongated (even a bit longer than the process on
tergite III), and is extended to about two-thirds of abdominal
tergum VII. Males of B. damzeni sp.n. could hold the whole
surface of the female’s wing with four pairs of long seta on
the notal process, thereby immobilizing the female’s wing.
Taking into consideration Thornhill’s observations (personal
communication), we proposed another hypothesis. Males might
use the strongly elongated notal and postnotal organ to interrupt
a female mating with another male, or even to abduct her.
Some functional similarity of a plier-like organ of
B. damzeni sp.n. may be observed with structures on the tho-
rax in Boreidae. Their metamorphosed wings form four narrow,
hooked straps. Males of Boreus spp. hold the wingless female
legs on their backs during copulation (Byers & Thornhill,
1983). Even though these structures have evolved on differ-
ent body parts, their morphological similarity is considerable.
However, the absence of such developed notal structures in
any living scorpionfly makes any interpretation of their use
very speculative.
The shape of the abdomen of B. damzeni sp.n. differs
from other described Panorpidae genera. The abdominal
segment V of Baltipanorpa gen.n. is the longest among
Panorpidae. The abdominal segments VI–VIII are elongated,
similar to Panorpa species with a long notal organ, and
Neopanorpa species, however not so extremely long as in
Leptopanorpa spp. (Lieftinck, 1936). Additionally, VI–VIII
segments are markedly thinner at the base what is common
to species of Sinopanorpa spp. (Cai et al. 2008). According
to the unusual development of notal and postnotal organs,
a change in general plan of the abdomen was necessary.
The most elongated notal organs are positioned on a small
postnotal process, and lie on the abdomen in extant Mecoptera.
In the case of an elongation of the postnotal organ in
B. damzeni sp.n., the processes do not lie on the abdomen
but protrude from the body, forming an angle of about 45◦ .
The terminal segments of the abdomen are thinner at the
base and elongated, as in Sinopanorpa (Cai et al., 2008), but
are recurved upwards, as in all Panorpidae. However, the
© 2011 The Authors
Systematic Entomology © 2011 The Royal Entomological Society, Systematic Entomology, 37, 223–228
Baltipanorpa damzeni gen. et sp.n. 227
long and massive segment V is unique among Panorpidae.
It is interesting if the elongated postnotal organ induced the
peculiar shape of the abdomen. One hypothesis may be that
males could use the massive segment V to support the female
during copulation. On the other hand, in the extant Panorpidae
another side-by-side V-shaped mating position was observed
(Mickoleit, 1971; Byers & Thornhill, 1983). Nevertheless, such
support would be necessary if the females of Baltipanorpa
were wingless or if their wings were reduced.
Penny (1975) hypothesized that the notal organ was probably
acquired by a Mesozoic mecopteran ancestor some 200 million
years ago. Willmann (1987, 1989) suggested an earlier
acquirement of this structure and monophyly of Mecoptera.
However, Whiting et al. (1997) and Whiting (2002) proposed
the paraphyly of this order based on molecular studies. Most
recently, Grimaldi & Engel (2005) confirmed Mecoptera to
be paraphyletic. From the topology of the phylogenetic tree
they proposed and taking into consideration the presence of
a rudimentary notal organ in Boreidae, we can assume that
the notal organ was acquired by a mecopteroid ancestor in the
Early Triassic, more than 240 million years ago.
Supporting Information
Additional Supporting Information may be found in the online
version of this article under the DOI reference:
10.1111/j.1365-3113.2011.00602.x
Figure S1. The examples of notal and postnotal organs in
extant Mecoptera.
Please note: Neither the Editors nor Wiley-Blackwell
are responsible for the content or functionality of any
supporting materials supplied by the authors. Any queries
(other than missing material) should be directed to the
corresponding author for the article.
Acknowledgements
We would like to thank to Michał Grabowski (University of
Łódź) and Ewa Krzemińska (MP ISEA, Kraków) for their
suggestions and help in preparing the earlier versions of the
article. Special acknowledgments are due to Lars Vilhelmsen
for his very useful comments on the submitted article, and to
the anonymous reviewer for an extensive and valuable review.
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Accepted 1 August 2011
First published online 28 September 2011
© 2011 The Authors