Arthropod Systematics & Phylogeny
75 – 8169 (2)

75© Museum für Tierkunde Dresden, eISSN 1864-8312, 21.07.2011

Notes on mating and oviposition of a primitive representativeof the higher Forficulina, Apachyus chartaceus (de Haan)(Insecta: Dermaptera: Apachyidae)
Shota Shimizu * & Ryuichiro Machida
Sugadaira Montane Research Center, University of Tsukuba, Sugadaira Kogen, Ueda,
Nagano 386-2204, Japan[ooarikui@sugadaira.tsukuba.ac.jp][machida@sugadaira.tsukuba.ac.jp]* Corresponding author
Received 22.iii.2011, accepted 08.iv.2011.
Published online at www.arthropod-systematics.de on 21.vii.2011.

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Abstract

Mating, oviposition, and selected details of the egg surface in the basalmost clade of the higher Forficulina, Apachyidae,
were described, using Apachyus chartaceus (de Haan, 1842) as a representative. The mating of A. chartaceus is of the endto-end type with the partners being dorsoventrally reversed. Throughout copulation, the male tightly holds the female’s
postabdomen using his forceps. This manner of mating is unique in Dermaptera, likely autapomorphic to Apachyidae, andperhaps correlated with life under bark. The eggs of A. chartaceus have an adhesive substance, by which they attach to thesubstratum; this is also found in basal Forficulina but is uniquely plesiomorphic for higher Forficulina. A. chartaceus does
not show intensive maternal care; this is overall a secondary condition or may be accidentally caused under rearing, while
the absence of some aspects of brood care (especially transport of eggs) is more likely plesiomorphic. Apachyus thus showsa mixture of unique apomorphic features and features that are uniquely plesiomorphic for higher Forficulina.

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Key words

Apachyus chartaceus, Apachyidae, Dermaptera, higher Forficulina, mating, oviposition, rearing, egg structures.

1. 		 Introduction
Dermaptera comprise the 11 families Karschiellidae,
Diplatyidae, Pygidicranidae, Apachyidae, Anisolabididae, Labiduridae, Forficulidae, Spongiphoridae, Chelisochidae, Arixeniidae, and Hemimeridae (Haas &Klass 2003), and about 2,000 species have been described (Sakai 1996). Hemimeridae and Arixeniidae
have curious features due to specialisation to their epizoic ways of life, and have often been dealt with as
independent suborders, i.e., Arixeniina and Hemimerina, respectively (Nakata & Maa 1974). The greaterpart of Dermaptera is composed of the nine remaining
families, to which the typical earwigs belong, being
named the Forficulina (Jarvis et al. 2005). Among theForficulina, the Karschiellidae, Diplatyidae and Pygidicranidae, all of which retain many ancestral char-acteristics, are called the “basal Dermaptera”, which
are surely paraphyletic, and the six remaining, morederived families are grouped into the “higher Forficulina”. The latter group is most likely a monophylum(Haas & Klass 2003; Klass 2003), though possibly
only with inclusion of Hemimeridae and Arixeniidae,
which might be in different subordinate positions
within the “higher Forficulina”. Apachyidae has been
regarded as the basalmost clade of the higher Forficulina, possessing primitive features concerning thewing structure and reproductive system (Haas 1995).
This agrees with the results of Jarvis et al. (2005),
who suggested Apachyidae to be the sister group of
the non-pygidicranid Forficulina including Hemimerina from their molecular-based phylogenetic analysis.

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Shimizu & Machida: Mating and oviposition of apachyid Dermaptera

Based on characteristics of the female postabdomen,
Klass (2001: pp. 278, 304) very tentatively proposed
Apachyidae to be the sister group of the epizoic Hemi­
meridae. In contrast, the preliminary results of Col­gan et al.’s (2003) molecular-based phylogenetic
study of Dermaptera, which included Apachyus peter­soni Borelli, 1925, suggest that Apachyidae is close to
Pygidicranidae, although the support for this relationship is weak. Therefore, Apachyidae is a very significant group for reconstructing the origin and ground­plan of the higher Forficulina or Dermaptera and perhaps also the origin of the epizoic lifestyle.
Apachyidae includes only two genera, Apachyus
and Dendroiketes, with altogether 15 species limited
to the Oriental, Australian, and Ethiopian zoogeographical regions (Steinmann 1989). The knowledge
on the biology of the group has remained sketchy andfragmentary. On this background, we tried to clarify
the behavior in mating and oviposition of Apachyidae, using Apachyus chartaceus (de Haan, 1842) asan exemplar. Knowledge of reproductive biology will
also be helpful in establishing a rearing technique forapachyid species, which in turn would ease further
studies on the life history of these earwigs.
Apachyus chartaceus is distributed on the Peninsula of Malaysia, Borneo and Sumatra (Kirby 1904;
Burr 1911). It has an extremely dorsoventrally flattened habitus, and the adults are about 3 cm in bodylength (Figs. 1 – 3). Both the forewings (sclerotized
tegmina) and the hindwings are well developed, andthey can fly (our pers. observ.). The forceps are large,
crescent-shaped without medial teeth, and similar inshape between males and females (Figs. 1 – 3). Between the bases of the paired forceps is a large, semicircular process, the “anal process” (Burr 1910),
which according to Klass (2001: p. 278) consists ofthe posterior part of tergite X, tergite XI, and perhaps
a dorsal telson sclerite. The combination of large crescent-shaped forceps and anal process reminds us of acap-opener (Figs. 1 – 3, 6).

meter) with wet soil (Fig. 4), in which a cylinder madeof non-woven pulp-polyethylene fabric was put. Different from the adults, the juveniles could not walkon the vertical plastic surface, and settled on the fabric cylinder. A. chartaceus was fed on several killed
Drosophila flies every several days. For mating, a paircollected from the field or a pair raised to adulthoodin the laboratory was kept in a single rearing vessel
for several days. The A. chartaceus individuals reared
stayed in the space between the vessel wall and fabriccylinder, and their behavior could be easily observed
through the transparent vessel wall.
After being cleaned in physiological saline,E­phrussi-Beadle’s solution (0.75% NaCl + 0.035%
KCl + 0.021% CaCl2), the eggs were pricked with afine needle and fixed in FAA (ethyl alcohol : formaline : acetic acid = 15 : 5 : 1) for 12 h.
For scanning electron microscopic observation,
the eggs fixed with FAA were dehydrated in a graded
ethyl alcohol series, transferred to and dipped in acetone for a time, in order to remove oil from the inside,which often emerges onto the egg surface and makes
it dirty. The eggs processed in acetone were hydrated
in a graded ethyl alcohol series and then postfixedwith 1% OsO4 for 1 h. The fixed eggs were again dehydrated in a graded ethyl alcohol series, dried witha critical point dryer TOUSIMIS Samdri®-PVT-3D,
coated with gold, and observed under a scanning electron microscope TOPCON SM-300.

2. 		 Material and methods
The Apachyus chartaceus specimens were collectedin tropical rain forests in Ulu Gombak, the State of
Selangor, Malaysia. We collected 10 juveniles, threemales and seven females in February 2009 and fourjuveniles, one male and two females in April 2010. All
individuals were hiding between the trunk and bark ofliving or dead trees.
The Apachyus chartaceus specimens were rearedin the laboratory at 25°C. They were kept separately
in a plastic vessel (10 cm in height and 8 cm in dia­3. 		 Results
The Apachyus chartaceus specimens stayed in the
narrow space between the fabric cylinder and vessel
wall (Fig. 4). Without the cylinder in the vessel, they
did not mate or lay eggs. A pair mated while placedon the vertical surface of the fabric cylinder (Fig. 5).
Mating was of the end-to-end type with the male andthe female being dorsoventrally reversed (Fig. 5A,B).
Throughout the copulation, the male tightly held his
partner’s postabdomen with his forceps in a symmetrical manner (Fig. 5C,C’). The duration of mating
was relatively long, and in one case lasted at least five
hours. The pair repeatedly mated during several days.
Even when disturbed, the pair ran around but maintained the copulation.
Among 14 females – including five that emerged
from among the 14 juveniles – three deposited eggs:
the numbers of eggs deposited were 7, 10 and 25. The
eggs were laid one by one and with some interspace
onto the wall of the rearing vessel and fabric cylinder, and they were tightly adhered to the substratum.
Once we observed a female to bring a mass of sev-

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Arthropod Systematics & Phylogeny 69 (2)

1

2

FC

3

4

Figs. 1 – 4. Apachyus chartaceus (de Haan, 1842) and a vessel for rearing it. 1: Male. Scale bar = 1 cm. 2: Female. Scale bar = 1 cm.
3: Ultimate instar juvenile. Scale bar = 1 cm. 4: Rearing vessel. Scale bar = 5 cm. FC = fabric cylinder.

eral eggs on her anal process (Fig. 6); the next day
the egg mass was found to fall on the ground, where
the eggs remained as a cluster. It is known that females of the higher Forficulina take great care of theireggs and hatched juveniles (overview in Matzke &Klass 2005). However, we did not once observe any
evidence of active egg-caring behavior of Apachyus
chartaceus females.

The eggs of Apachyus chartaceus were ellipsoidin shape, with the anterior half slightly swollen, anda little less than 1 mm in length (Fig. 7). They werewhite since the whitish yolk was visible through thetransparent chorion (Fig. 7). The eggs were covered
with a secretion, which was abundant in the posterior
half. This secretion or “adhesive substance” acted asa “glue” adhering the egg to the substratum. When

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Shimizu & Machida: Mating and oviposition of apachyid Dermaptera

O
P

O

P

O

5A

P

5B

5C
O

P

6

5C’
F
E

AP

Ab10

6’

7A

7B

AS

Figs. 5 – 7. Mating and eggs of Apachyus chartaceus (de Haan, 1842). 5: Mating. 5A: Male (left, ventral view) and female (right,
dorsal view). 5B: Male (left, dorsal view) and female (right, ventral view). 5C: Lateral view. 5C’: Enlargement of the connection.
Scale bars = 1 cm. 6: Female with an egg mass on her anal process. Scale bar = 1cm. 6’: Enlargement of the postabdomen. Scalebar = 1 mm. AP = anal process; Ab10 = 10th abdominal segment; E = eggs; F = forceps. 7: Egg. 7A: Moistened egg with swollen
adhesive substance (AS). An arrowhead shows the approximate position of the circular spot seen light microscopically (Fig. 8A)or the chorionic elevation seen scanning electron microscopically (Fig. 8B) revealed on the anterior pole of the egg. 7B: Dried egg
with desiccated adhesive substance. Scale bar = 500 μm.

79Arthropod Systematics & Phylogeny 69 (2)

8A

8C

8B

8D

Fig. 8. Structures around the anterior pole of the egg of Apachyus chartaceus (de Haan, 1842). A: Light micrograph showing a circular spot (arrow). Scale bar = 100 μm. B: Scanning electron micrograph showing a low chorionic elevation, which is light microscopically detected as a whitish circular spot as shown in A. Scale bar = 20 μm. Arrow = central depression with porous structure;arrowheads = small pores circularly arranged; star = low chorionic elevation. C: Enlargement showing the central depression of
porous structure. Scale bar = 5 μm. D: Enlargement showing two of the small circularly arranged pores. Scale bar = 2 μm.

the egg was artificially moistened, the largely swollen
adhesive substance was soft and sticky (Fig. 7A), but
once dry, it shrunk and became hard (Fig. 7B), yielding adhesive power. Light microscopic observation
revealed a circular spot about 85 μm in diameter at
the anterior pole of the egg (Fig. 8A). Under a scanning microscope, this structure proved to be a lowele­vation of the chorion (Fig. 8B). In the center ofthis chorionic elevation, there was a depression about
10 μm in diameter, in whose center a porous structure
was visible (Fig. 8B,C). Around the chorionic elevation, small pores about 1–2 μm in diameter were circularly arranged (Fig. 8B,D).

4. 		 Discussion
The copulation of Apachyidae was observed for the
first time: it is 1) of the end-to-end type, 2) with themale tightly and symmetrically grasping the female’s
postabdomen using his forceps throughout mating,and 3) with the pair keeping a posture in which they
are dorsoventrally reversed.

The posture in Apachyus chartaceus is peculiar
among Dermaptera and may be regarded as an autapomorphy of Apachyidae. In other Dermaptera, the pairs
do not dorsoventrally reverse their postures, but usually the males twist their postabdomen during mating(Günther & Herter 1974; Briceño & Eberhard
1995; Kamimura 2006). The functional advantage
might be correlated with the life of A. chartaceus under bark, where abdominal twisting would be difficult,as it needs additional vertical space. And thanks to theabnormal mating posture the animals do not need toleave their sheltered micro-habitat for mating and thus
avoid exposure to predators.
It is also unusual that the males hold their partners
with their forceps. Apart from Apachyus chartaceus
we only know the example of Pseudomarava prom­
inens Steinmann, 1990 (Spongiphoridae) in Briceño& Eberhard (1995): the male of P. prominens, which
possesses crescent-shaped forceps suitable for holding
like A. chartaceus, grasps the female prior to copulating and inserts his penis without twisting his abdomen. However, once the end-to-end copulation without dorsoventral reverse has succeeded, the male releases the female, different from A. chartaceus, where
the grasping continues throughout mating.

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Shimizu & Machida: Mating and oviposition of apachyid Dermaptera

The pairs of Apachyus chartaceus did not ceasemating if disturbed, and only ran around to escape,maintaining the connection. This may be unusual in
Dermaptera, in which disturbed pairs usually stop
copulating, and we have few examples for keeping
the copulation under disturbance: Tagalina papua(de Bormans, 1903) (Pygidicranidae) in Matzke &Klass (2005) and Euborellia cincticollis (Gerstaecker, 1883) (Anisolabididae) in Knabke & Grigarick(1971).
The eggs of Apachyus chartaceus have an adhesive substance in their posterior half, and are deposited and stuck to the substratum. Among Dermaptera,an adhesive substance which attaches the egg to thesubstratum has been reported only in two families ofthe basal Dermaptera, i.e., the Diplatyidae and Pygidicranidae: for Diplatyidae, Diplatys greeni Burr,
1904 (Green 1898), a Papua New Guinean gen. sp.
(Matzke & Klass 2005), and Diplatys flavicollis(Shiraki, 1907) (Shimizu & Machida 2009, 2011);
for Pygidicranidae, Tagalina papua (de Bormans,
1903), Tagalina burri Hincks, 1955, and Paracra­
nopygia siamensis (Dohrn, 1863) (Matzke & Klass
2005: figs. 2 – 10). In both families, being more posteriorly restricted than in A. chartaceus, the adhesive
substance is located at the posterior pole of the egg.
In the Papua New Guinean diplatyid (Matzke &Klass 2005) and D. flavicollis (Shimizu & Machida2009, 2011), the adhesive substance forms a stalk, andShimizu & Machida (2009) named it the adhesive
stalk. The present finding of the adhesive substancein A. chartaceus is the first report for the higher Forficulina. We encountered a case where the eggs werecarried as a cluster on the female’s anal process. Thismight have been an accident, but it could alternatively
be a step preceding deposition on the substratum.
Diplatyidae, Pygidicranidae and Karschiellidae
have complicated external female genitalia, which includes the presence of an ovipositor (2 pairs of gon­apophyses and 1 pair of gonoplacs, with much variation) and usually IXth-segmental accessory glands– all corresponding with the ground plan of InsectaDicondylia (Klass 2003). The accessory glands ofabdominal segment IX are in many other Insecta thesource for substances added to the egg surface either
for adhesion or other purposes. Matzke & Klass(2005) proposed that in the basal Dermaptera the accessory glands likewise yield the secretions for egg
attachment and the gonapophyses provide the ability
for targeted deposition of the eggs (which due to their
attachment cannot be translocated later, as it is done inhigher Forficulina). Female genitalia are strongly simplified, the ovipositor is reduced, and the accessory
glands are absent in all higher Forficulina studied sofar (Giles 1961; Bhatnagar 1964; for Eudermaptera
K.-D. Klass & K. Schneider pers. comm.). Unfortu-

nately, female genitalia have not yet been studied in
an apachyid, and it is thus unknown for the groupwhether accessory glands are present and what is thedegree of differentiation of the ovipositor (which, if
present, must be short, since no valves are seen to surpass the subgenital plate). Both the presence and theabsence of accessory glands in Apachyus would beof special interest – the former in a phylogenetic context (perhaps a unique plesiomorphy of Apachyidae
among higher Forficulina), the latter in a functional
context (where do then the secretions for egg attachment come from?).
At the anterior pole of the eggs of Apachyus
chartaceus, small pores are circularly arranged, andat the center of the pore group a single pore with porous inner structures is present. Our knowledge on
egg structures of Dermaptera is very scanty, and wehave only a study on those of Forficula auricularia L.,
1758 by Chauvin et al. (1991). The pore structures
in A. chartaceus eggs closely resemble those found in
F. auricularia. Chauvin et al. (1991) identified thecircularly arranged pores and the single central porewith inner porous structures as aeropyles and a micropyle, respectively, although they did not show any evidence.
For very few species of Diplatyidae and Pygidicranidae a few elements of maternal brood care havebeen reported (Matzke & Klass 2005: table 3): (1)Mothers remain associated with eggs and (2) withyoung first instar nymphs, and both (3) eggs and (4)nymphs are defended. These are likely groundplan
elements of brood care in Dermaptera (but there are
no data on Karschiellidae). In higher Forficulina (5)transport of eggs, (6) cleaning of eggs with mouthparts, (7) assistance to nymphs in hatching from theeggs, and (8) grasping of nymphs with mouthparts(to collocate them) have additionally been reported.
These elements have most likely evolved at the baseof or within the higher Forficulina. In contrast, in thisstudy, we did not observe any behavior suggestive ofmaternal care in Apachyus chartaceus. The absenceof the behavioural elements (1) – (4) may be interpreted as secondary. The elements (6), (7), and (8) ofbrood care could be either primarily absent (not yet
evolved in this basal lineage of higher Forficulina) or
secondarily absent (included in a general reductionof brood care as evidenced by absence of features(1)–(4)). However, it is possible that the lack of caring elements is an effect of rearing in unusual andstressful conditions. The absence of (5), egg transport, may plausibly be considered primary, since egg
transport may be unlikely to co-occur with eggs fixed
to the substrate. On the other hand, diplatyids havebeen observed to translocate freshly laid eggs prior tothe hardening of the adhesive substance (Shimizu &Machida 2011).

Arthropod Systematics & Phylogeny 69 (2)5. 		 Conclusions
The present study revealed that Apachyus chartaceus
has an apomorphic copulation posture including dorsoventrally reversed partners and a continued graspby male cerci, both unique among Dermaptera; its
eggs are attached to the substratum by an adhesive
substance, which is uniquely plesiomorphic among
higher Forficulina but usual in basal Dermaptera; maternal brood care has not been observed at all, whichis unique among forficuline earwigs and at least partly
apomorphic. The mixture of unique apomorphic features and features that are uniquely plesiomorphic forhigher Forficulina makes Apachyidae a highly interesting study taxon.

6. 		 Acknowledgments
We are grateful to Dr. M. Nishikawa of Ehime University, Japan, for his help in the identification of specimens, to Dr. Y.
Kamimura of the Keio University, Japan, for his valuable comments on the mating behavior of Dermaptera, and to Dr. K.-D.
Klass of the Senckenberg Natural History Collections Dresden and an anonymous reviewer for valuable comments to themanu­script. We also thank Messrs. Kim Hong Yap, Mohamad
Fadil and Shahrul Nazly B. Mahmud for their kind assis­
tance in collecting materials in Malaysia. This study was supported by a Grant-in-Aid from the JSPS (Scientific Research
C: 21570089) to RM. This is contribution No. 220 from theSugadaira Montane Research Center, University of Tsukuba.

7. 		 References
Bhatnagar R.D.S. 1964. The morphology of the male andfemale terminalia and external genitalia of the earwig
Labidura riparia (Pallas) (Dermaptera, Labiduridae). –The Entomologist 97: 106 – 112.
Briceño R.D., Eberhard W.G. 1995. The functional morpho­
logy of male cerci and associated characters in 13 speciesof tropical earwigs (Dermaptera: Forficulidae, Labiidae,
Carcinophoridae, Pygidicranidae). – Smithonian Contributions to Zoology 555: i – iii + 1 – 63.
Burr M. 1910. Dermaptera (earwigs). In: Shipley A.E. (ed.),The Fauna of British India Including Ceylon and Burma. –Taylor and Francis, London. xviii + 217 pp. + 10 plates.
Burr M. 1911. Dermaptera. – Genera Insectorum 122: 1 – 112+ plates 1 – 9.
Chauvin G., Hamon C., Vancassel M., Vannier G. 1991.
The eggs of Forficula auricularia L. (Dermaptera, Forficu­
lidae): Ultrastructure and resistance to low and high temperatures. – Canadian Journal of Zoology 69: 2873 – 2878.
Colgan D.J., Cassis G., Beacham E. 2003. Setting the molecular phylogenetic framework for the Dermaptera. – Insect Systematics & Evolution 34: 65 – 79.

81
Giles E.T. 1961. The female reproductive organs and genital
segments of Anisolabis littorea (White) (Dermaptera, Labiduridae). – Transactions of the Royal Society of New Zealand (Zoology) 1: 293 – 302.
Green E.E. 1898. Further notes on Dyscritina, Westw. – Transactions of the Royal Entomological Society of London 5:
381 – 390 + plates 18 – 19.
Günther K., Herter K. 1974. 11. Ordnung Dermaptera (Ohr­würmer). Pp. 1 – 158 in: Helmcke J.-G., Starck D., Wer­
muth H. (eds.), Handbuch der Zoologie IV (2) 2/11. – Walter de Gruyter, Berlin.
Haas F. 1995. The phylogeny of the Forficulina, a suborder ofthe Dermaptera. – Systematic Entomology 20: 85–98.
Haas F., Klass K.-D. 2003. The basal phylogenetic relationships in the Dermaptera. In: Klass K.-D. (ed.), Proceedings of the 1st Dresden meeting on insect phylogeny:“Phylogenetic Relationships within the Insect Orders”(Dresden, September 19 – 21, 2003). – Entomologische Ab­
hand­lungen 61: 132 – 133.
Jarvis K.J., Haas F., Whiting M.F. 2005. Phylogeny of earwigs (Insecta: Dermaptera) based on molecular & morphological evidence: Reconsidering the classification of Dermaptera. – Systematic Entomology 30: 442 – 453.
Kamimura Y. 2006. Right-handed penises of the earwig Labi­
dura riparia (Insecta, Dermaptera, Labiduridae): Evolutionary relationships between structural and behavioral
asymmetries. – Journal of Morphology 267: 1381 – 1389.
Kirby W.F. 1904. Vol. I. Orthoptera, Euplexoptera, Cursoria,
et Gressoria (Forficulidae, Hemimeridae, Blattidae, Mantidae, Phasmidae). A Synonymic Catalogue of Orthoptera. –Bri­tish Museum, London. 501 pp.
Klass K.-D. 2001. The female abdomen of the viviparous
earwig Hemimerus vosseleri (Insecta: Dermaptera: Hemi­
meridae), with a discussion of the postgenital abdomen of
Insecta. – Zoological Journal of the Linnean Society 131:
251 – 307.
Klass K.-D. 2003. The female genitalic region in basal earwigs (Insecta: Dermaptera: Pygidicranidae s.l.). – Entomo­
logische Abhandlungen 61: 173 – 225.
Knabke J.J., Grigarick A.A. 1971. Biology of the African
earwig, Euborellia cincticollis (Gerstaecker) in California
and comparative notes on Euborellia annulipes (Lucas). –Hilgardia 41: 157 – 194.
Matzke D., Klass K.-D. 2005. Reproductive biology andnym­phal development in the basal earwig Tagalina papua(Insecta: Dermaptera: Pygidicranidae), with a comparisonof brood care in Dermaptera and Embioptera. – Entomologische Abhandlungen 62: 99 – 116.
Nakata S., Maa T.C. 1974. A review of the parasitic earwigs(Dermaptera: Arixeniina; Hemimerina). – Pacific Insects
16: 307 – 374.
Shimizu S., Machida R. 2009. Eggs and their deposition andcare of a primitive earwig Diplatys flavicollis (Shiraki) (Dermaptera: Diplatyidae). – Proceedings of the Arthropodan
Embryological Society of Japan 44: 29 – 32.
Shimizu S., Machida R. 2011. Reproductive biology and post­embryonic development in the basal earwig Diplatys flavi­
collis (Shiraki) (Insecta: Dermaptera: Diplatyidae). – Arthro­
pod Systematics & Phylogeny 69: 83 – 97.
Steinmann H. 1989. World Catalogue of Dermaptera. – Aka­
démiai Kiadó, Budapest. 934 pp.

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Zeitschrift/Journal: Arthropod Systematics and Phylogeny
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Band/Volume: 69
Autor(en)/Author(s): Shimizu Shota, Machida Ryuichiro
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