Ever since the description of Zorotypus guineensis Silvestri, 1913 and Grylloblatta campodeiformis Walker, 1914, which served for the erection of the new orders Zoraptera Silvestri, 1913 and Grylloblattodea Brues & Melander, 1932 (= Notoptera Crampton, 1915) all newly described insect species could be assigned to known orders. The insect order Mantophasmatodea, described by K.-D. Klass, O. Zompro, N.P. Kristensen, and J. Adis in 2002 (Science 296: 1456–1459).The initial description of Mantophasmatodea in KLASS et al. (2002) was based on two museum specimens: an ethanol preserved female collected in Namibia in 1909, and a dried male collected in Tanzania in 1950. These have been described as two different species, Mantophasma zephyra Zompro, Klass, Kristensen & Adis, 2002, and Mantophasma subsolana Zompro, Klass, Kristensen & Adis, 2002.Extant Mantophasmatodea are only known from Africa south of the equator, these approximately 45 million year old fossils show that in the early Tertiary mantophasmatodeans also inhabited Europe (ZOMPRO et al. 2002).The first living specimens of Mantophasmatodea were found in February and March 2002, when an international expedition of entomologists, directed by E. Marais (National Museum in Windhoek, Namibia) and O. Zompro, searched the Brandberg (Namibia, Erongo Province).No doubt Mantophasmatodeans are beautiful and very interesting creatures on Earth which have been recently discovered. Their life style and unique behaviours are amazing. Their evolution or affinities have different point of view among the scientists. But majority believes in their relation with Grylloblattodea. Recently Grylloblattodea and Mantophasmatodea have been put into a new clade called, Xenonomia.But the conflict regarding their ordinal status is still there.

1. Mantophasmatodea- Discovery, Description, Life history,
Behaviour, Affinities and, Ordinal status
SOUREN MONDAL
M.Sc. ENTOMOLOGY
INDIAN AGRICULTURAL RESEARCH INSTITUTE
msouren46@gmail.com

2. HISTORY OF DESCRIPTION
• Ever since the description of Zorotypus guineensis Silvestri, 1913 and Grylloblatta campodeiformis Walker, 1914, which served for the
erection of the new orders Zoraptera Silvestri, 1913 and Grylloblattodea Brues & Melander, 1932 (= Notoptera Crampton, 1915) all newly
described insect species could be assigned to known orders.
• The insect order Mantophasmatodea, described by K.-D. Klass, O. Zompro, N.P. Kristensen, and J. Adis in 2002 (Science 296: 1456–1459).
• The initial description of Mantophasmatodea in KLASS et al. (2002) was based on two museum specimens: an ethanol preserved female
collected in Namibia in 1909, and a dried male collected in Tanzania in 1950. These have been described as two different species,
Mantophasma zephyra Zompro, Klass, Kristensen & Adis, 2002, and Mantophasma subsolana Zompro, Klass, Kristensen & Adis, 2002.
• The Tanzanian specimen had been stored at the Zoological Museum of the University of Lund (Sweden) until 1986, when R. Danielsson,
entomology curator in Lund, sent it to The Natural History Museum in London in order to have it identified.
• Over the years, J. Marshal, a curator in London, showed it to many insect systematists, but only O. Zompro (Max-Planck-Institute for
Limnology, Plön, Germany), a specialist of Phasmatodea who was visiting The Natural History Museum in summer 2001, realized that this
could be a member of an unknown insect order.
• A little later he found the female specimen from Namibia in the Museum für Naturkunde of the Humboldt-University, Berlin, where it had
been stored among the unidentified Phasmatodea.
• For confirmation of the importance of this discovery, Zompro sent these two specimens to Klass, then working at the Zoological Museum
of the University of Copenhagen, who then conducted the morphological studies and character discussion that eventually showed that
the status of a new order was actually appropriate.

3. • Extant Mantophasmatodea are only known from Africa south of the equator, these approximately 45 million year old fossils show that in
the early Tertiary mantophasmatodeans also inhabited Europe (ZOMPRO et al. 2002).
• The first living specimens of Mantophasmatodea were found in February and March 2002, when an international expedition of
entomologists, directed by E. Marais (National Museum in Windhoek, Namibia) and O. Zompro, searched the Brandberg (Namibia,
Erongo Province).
• Mantophasmatodea showed up in several places in Namibia. This dramatic expansion of the known distribution area and number of
putative species of Mantophasmatodea led Klass and Picker to compose an international research team for the study of
Mantophasmatodea. After extensive cooperative collecting and field work in July–September 2003, the members of the team (VAN
NOORT 2003) have started to study the taxonomy of Mantophasmatodea, their skeleto-muscular system, circulatory system, and
ovaries, the structure of their eggs and egg pods, their embryonic and nymphal development, their chromosome sets, their
phylogenetic relationships (those within Mantophasmatodea and those to other insects, based on morphological and molecular data),
their behaviour, life history and ecology, and their biogeographic history at the species and population levels.

4. DISTRIBUTION OF MANTOPHASMATODEA
SPECIES IN AFRICA.
Klass et al. 2003

5. • The holotype of Praedatophasma maraisi Zompro & Adis, was
collected in a dry, stony, mountainous area in southwestern Africa.
• The fossil Raptophasma lived in a much different area and environment
(Pre-European mainland, Eocene woodlands some 45 Million years
ago). Generally, the Baltic amber fauna and flora has been studied quite
extensively during the last few decades, with several reconstructions of
the palaeoenvironment attempted (LARSSON 1978; WICHARD &
WEITSCHAT 1996; WEITSCHAT & WICHARD 1998). Palaeoclimatic data
show that the Eocene (Early Tertiary), when the Baltic amber was
formed, was characterized by very warm, tropical to subtropical
conditions extending all the way into high palaeolatitudes. This is
verified by numerous geological and palaeoclimatological indicators.
• Localities harboring Mantophasma-todea show a wide range of
vegetation density. Some species frequently are found in tufts of grass
(Poaceae) or Cape reed (Restionaceae, which superficially resemble
grasses); they occur either at the bottom between the culms or on top
of the culms, where they are well camouflaged by their mottled and
striped coloration.
HABITAT
Source: Roth et al. 2014

6. Colouration
The basic color of the body is brown, gray, green, or yellow; different tints of these colors, a whitish component of varied
extent, and— in some species—black dots form a pattern of mottles and/or longitudinal stripes. Coloration varies between
and also within species. Nymphs resemble the adults in appearance.
• Mottled-type colouration: Characterized by a complicated patchwork of colouration components
that are more or less strongly contrasted against a bright whitish or greyish and partly reddish
background colour. The elements of this patchwork show various tints of black, brown, dark grey,
red, and dark violet, cover a wide range from fairly bright to very dark.
• Uniform type colouration: In the respective specimens colouration is much simpler,
because most of the dark elements present in the mottled-type specimens are either
entirely absent or absent in some body regions.
Longitudinal ribbons: The epidermal pigmentation together with or without patches of darkened cuticle
can form dorsal, pleural, and ventral ribbons that run along the entire body but are occasionally broken
or vestigial.
Two basic types of colouration are distinguishable: mottled and (essentially) uniform.
Body Size
DESCRIPTION OF THE TAXON
Image source:
Klass et al. 2003
The consistently wingless Mantophasmatodea have a uniform body shape, which superficially resembles certain grasshoppers
or stick insects. The body length (without the antennae) ranges from 0.35 to 0.94 in (9–24 mm), males usually being somewhat
smaller than females.

7. According to KLASS et al. (2002), they are hemimetabolous insects
with slight sexual dimorphism.
• Orthognathous with generalized mouthparts.
• The mandibles bear three small teeth in the molar area.
• The maxillar palpus is five, the labial palpus three segmented.
• The tentorium lacks a perforation; its anterior pit is far above
the anterior mandibular articulation.
• An epistomal sulcus is lacking.
• The subgenal sulcus has an unusual course, extending from
the posterior mandibular articulation directly to the anterior
tentorial pit and thence downwards/backwards to anterior
mandibular articulation.
• The head capsule is posteroventrally closed by a weak
submentum, a gula is not produced.
The head
Metamorphosis
Image source: Wipfler et al. 2018

8. • Long and filiform.
• The antennae show a unique structure: they have 14 basiflagellomeres
and 7 distiflagellomeres, which differ in their width, shape, and
equipment with sensilla .
• The first and sixth distiflagellomeres each bear a tiny distal gland.
The antennae
The compound eyes are well developed, albeit of varied size, but ocelli are
lacking.
Compound eyes and ocelli
• Each tergum overlaps narrowly the following, the terga are decreasing in
length.
• The prothoracic pleuron is large and fully exposed. The metathorax lacks a
spinal apodeme.
• The coxae are narrowed and slightly elongated. The pro- and mesofemora are
broadened with their ventral edges being bristled or spinose.
the thorax
Zompro et al. 2003
Zompro et al. 2002
Image source: Klass et al. 2003

9. • The femora are distinctly thickened in the forelegs and somewhat thickened in the
midlegs, but they are very slender in the hind legs.
• The tibiae of the forelegs and midlegs bear on their inner surfaces (opposing the
femora) two rows of short thorns, which render the legs suitable for grasping other
insects.
• The tarsi comprise five tarsomeres, but the three basal ones are fused (with grooves
indicating the borders).
• A large adhesive lobe (arolium) originates from between the claws.
• The hindlegs are elongated, and used for small jumps. The tarsi consist of five
segments, a large arolium is present.
• The four basal tarsomeres bear euplantulae. The third tarsomere bears a small,
triangular process dorsoapically.
Leg
All known species are apterous, lacking any rudiments of wings.
Wing
• The abdomen consists of 10 well-developed segments and a reduced eleventh segment.
Abdominal segments are transverse.
• The cerci are one-segmented.
• In the male, the subgenital plate features a median projection.
• The small spiracles I–VIII are located in the pleural membrane, with intrinsic occlusor
and extrinsic (coxosternal) dilator muscles.
Abdomen Ramel 2021
Zompro et al. 2003
Zompro et al. 2003

10. Male Genitalia
In the male, the coxosternum IX is not subdivided, and forms a
subgenital lobe with median spatulate process. Styli are absent. The
phallic region features membranous lobes around the gonopore and the
transverse, medially asymmetrically produced sclerite articulates with
the anterolateral corners of tergum X. The cerci are prominent, clasping,
and not forming a differentiated articulation with tergum X.
In the female the ovipositor projects markedly beyond the short
subgenital lobe formed by the coxosternum VIII. The gonoplacs are
short and strongly sclerotized. The gonapophyses VIII are markedly
blunt distally, the gonapophyses IX are largely fused with the gonoplacs;
its composite formation with ventral keel interlocking with dorsal
groove on gonapophyses VIII. The gonangulum bears the usual 3
articulations.
Female genitalia
Image source: KLASS et al. 2002

11. • The abdominal ganglion VII is free from the terminal ganglion.
Digestive system
• The foregut has a large proventricle which is armed with weak, papillose
sclerites that terminate in three successive whorls of weakly sclerotized
lobes.
Circulatory system
• The heart lacks lateral arteries in the mid abdomen. The abdomen has a
ventral diaphragm.
Nervous system
Source: Roth et al. 2014

12. LIFE HISTORY
• Mantophasmatodea are annual and univoltine species.
Females oviposit in pods of foam that form a cocoon,
which hardens due to incorporated sand and contains
eggs from both ovaries.
• These insects may occasionally oviposit eggs from a
single ovary.
• These pods are laid superficially (0.5 - 2 cm deep) in
sandy soil.
• females do not hesitate to lay eggs at approximately the
same spot.
• In the pods, the eggs exhibit a regular vertical
arrangement similar to that of bottles in a beer crate.
• Pregnant Mantophasmatodea females laid eggs almost
immediately if sufficient soil was provided.
• Individual females laid up to four egg pods over a short
period, i.e., a few days, and 50 - 100 eggs total during
their adulthood.
EGG
Image source: Roth et al.
2014

13. In nature, the egg stages survive at least eight months of a dry
summer or winter within the resistant egg pods. Other than the direct
influence of rainfall on egg development, there is no information on
which additional factors induce the end of dormancy in eggs after the
dry period.
The first instar nymphs hatch the following year during the
rainy season (i.e., in mid-summer for Namibia/East Africa and
autumn/winter (May or later) for South Africa) .
They reach adulthood after 3.5-4 months ; the adults can
survive at least two months in the laboratory.
DIAPAUSE
NYMPH
ADULT
Image source: http://www.biodiversityexplorer.info/mantophasmatodea/recognition/index.htm
1st instar
4th or 5th instar
Adult female

14. Unique behaviours of The Gladiators
(Roth et al. 2014)
❑ Flexibility: Both sexes are secondarily wingless and highly flexible along the longitudinal
axis. This flexibility enables the species to clean their external genitalia with their
mouthparts. but it is also used to attack large prey.
❑ Camouflage: The known species’ ground color is brownish, grey, or greenish, which
indicates camouflage.
❑ Reluctancy to escape behaviour: Heelwalkers rarely show fast escape behaviors.
However, in certain cases, a strange “curling” behavior can be seen when the
insects are knocked out of a bush. This behavior is particularly conspicuous
because, after dropping, Mantophasmatodea typically do not immediately move
away. Heelwalkers sometimes jump for small distances, such as a few centimeters.
Occasionally, Mantophasmatodea actively drops in the vegetation. In most cases,
they land upright on their legs ("cat-like") on the ground or in the correct direction
in small branches of shrubs and grasses.
Source: https://www.google.com/url?sa=i&url=https%3A%2F%2Fgenent.cals.ncsu.edu%2Finsect-
identification%2Forder-
mantophasmatodea%2F&psig=AOvVaw1dU3I7nwOw2QBSyG1sDEDY&ust=1685473127598000&s
ource=images&cd=vfe&ved=0CBEQjRxqFwoTCOiG9Omam_8CFQAAAAAdAAAAABAE
Source: Roth et al. 2014

15. Jumping style
Source: Roth et al. 2014
Cat-like landing

16. ❑ Multifunctional arolium:
• Both sexes possess very large arolia on the pretarsi of all legs, which are, however, mostly held in an upright
position; this highly characteristic position ("heelwalker") is quite unique among insects.
• When cleaning body parts or eating large prey, the animals sometimes hung upside-down and often used only
a single hindleg arolium to adhere to smooth plastic surfaces.
• Effective adhesion is supported by secretions produced in large glands in the arolia. Secretion production is
typically energy-consuming, which may be one reason to maintain the large arolia in an upward position when
not in use.
• They frequently clean their arolia but also antennae and genital segments using their mouthparts.
Source: Eberhard et al. 2009
Source: Roth et al. 2014

17. ❑ Waving of entenna: While walking or hiding, these insects
continuously wave their antennae in a manner similar to that of
cockroaches.
❑ Occasionally, males walk with their abdomens upwards, similarly to
scorpions (e.g., when approaching a sexual partner).
❑ Diurnal habit has been observed in Mantophasma specimens.
Occasionally, these insects are seen sitting in the upper parts of grasses
and outer shrub branches during the hottest portion of the day (11
a.m. - 5 p.m.);For all species, mating and drumming behaviors are
predominantly observed at night, including dusk and dawn.
Male, bending their abdomen upward like a scorpion
Image Source: Roth et al. 2014

18. PREY AND FEEDING BEHAVIOR
• Mantophasmatodea are carnivorous insects.
Similarly to many other predatory insects, they may
be generalists regarding their prey because they
consume different types of arthropods, mainly
spiders and insects.
• A hide-and-wait strategy (in combination with slow
stalking) is very likely and is concordant with the
nocturnal activity of most species.
• Apart from hunting, when Mantophasmatodea
stalk or walk quickly over very short distances to
catch their prey, they predominantly sit and hide,
and they rarely walk in vegetation. Most species
hide in the centers of bushes or grass tussocks
during the day.
Image Source: Roth et al. 2014

19. DRUMMING BEHAVIOUR
Both sexes use single vibrational calls, which
are produced by tapping the abdomen onto the
substrate for mate location and recognition.
The male subgenital plate has a semicircular
lip-like process of taxon-specific extension,
which temporarily contacts the surface during
drumming (=‘drumstick’).
The Females do not develop this abdominal
process and tap with the middle part of their
ventral abdomen. Female calls consist of single
repeated pulses, whereas male calls are more
complex pulse trains that likely contain coding
for species recognition.
Image source : https://zoologie.uni-
greifswald.de/storages/uni-
greifswald/fakultaet/mnf/biologie/zoologie/allg_und_syst_
zool/images_other/Images_Research/Mantophasmatodea/
K_biedouwense_female_1_hp.jpg
Male subgenital process
Image Source: Roth et al. 2014

20. COURTSHIP AND MATING BEHAVIOUR
• Mating generally begins in the evening or at night. In
contrast, mating of several Mantophasma specimens
was observed at any time of day.
• Vibrational communication(drumming) is part of a
heelwalker's sexual behavior.
• As they near each other, rapid antennation (i.e.,
flickering of their antennae) occurs, which indicates
odour-mediated sexual communication. This approach
is often interrupted by phases without movement.
Image Source: Roth et al. 2014
Typical mating position of Mantophasmatodea

21. • Arolia are used during copulation, this behavior is used during
mounting to stabilize their position.
• The male cerci are used as claspers to bring the genitalia of sexes
close together.
• The male genitalia evaginate and insert into the vagina.
• During prolonged mating, males remain mounted regardless of
the female’s behavior. During copulation, females are also can be
observed walking, stalking, catching prey, and excreting. Even if a
female dropped down from the grass, the male remained sitting
on the female's abdomen afterwards.
• The males generally appear to be at risk of attack by the female,
as indicated by their rapid mounting. Female defend herself
against a mounted male by both leg kicking and biting.
Image Source: Roth et al. 2014

22. SEXUAL DIMORPHISM
• The one-segmented cerci are short in the female but fairly long and curved in the male, which uses them as accessory
copulatory structures.
• The tenth abdominal tergum, which bears the muscles moving the cerci, is much wider in the male than in the female.
While the middle part of the abdomen is widest in females (because it harbors the ovaries), in males the hindmost part
usually is widest due to the expanded tenth tergum.
Image source :
Klass et al. 2003
Widest middle part
Widest hindmost part
Adult female Adult male
Short cerci
Long, curved cerci

23. AFFINITIES
• Mantophasmatodeans exhibit several characters which are similar to members of other polyneopteran insect orders; for instance, they
have an enlarged arolium (a cushion-like pad between the tarsal claws) which is superficially similar to Timema, a basal genus of stick
insects.
• Tilgner (2002) suggested that mantophasmatodeans may simply be ‘‘aberrant members of the order Orthoptera’’ and argued that the
characters used to exclude them from this group were misinterpreted, however, he also failed to provide any formal phylogenetic
analysis supporting this conclusion. The difficulty in placing the Mantophasmatodea among insect orders highlights the confusion
regarding the phylogenetic relationships among polyneopteran insects in general.
Tilgner, 2002; Whiting et al., 2003
• A recent investigation detailing the sperm structure of Mantophasmatodea zephyra suggested similarities with Mantodea.
• They also possess a ‘‘configuration of sclerites and lobes’’ in the proventriculus (a small region of the digestive tract) that is similar to
Grylloblattodea, but also paralleled in other insects.
Klass et al., 2002
Dallai et al., 2003

24. Terry et al., 2005
These data demonstrate that Mantophasmatodea is neither the sister group of Mantodea nor Phasmatodea, and strongly support the
sister taxon relationship of Grylloblattodea and Mantophasmatodea.
Grylloblattodea is a small, cryophilic order of insect (25 extant species, five genera) confined to north-western North American and north-
eastern Asia. Extant representatives are wingless scavengers and ⁄or predators (Storozhenko, 1979) adapted to cool and cold
environments. The most diverse genus (Grylloblatta) is distributed throughout the mountains of Western Canada and the North-western
USA and is adapted to the extreme temperatures associated with glaciers and ice caves.
On the other hand, extant Mantophasmatodea are confined to the Karoo-Namib region of southern Africa (Picker et al., 2002), and are
adapted to the hot, arid conditions prevalent in this area. Although the paleontological evidence indicates a much wider historic
distribution (Zompro et al., 2002).
This disjunction of distributions and the small size of both groups suggest an ancient divergence with subsequent extinctions of
intermediate lineages. This may be a common theme throughout Polyneoptera and could explain many of the difficulties regarding the
phylogenetic reconstruction of relationships.
The results of these phylogenetic analyses, coupled with the ecological disparity between these two groups and the formal recognition of
Grylloblattodea for nearly a century, strongly supports the validity of the ‘‘gladiator’’ as a new insect order. We have named the clade
comprising Grylloblattodea + Mantophasmatodea as ‘‘Xenonomia’’, derived from the Greek ‘‘xenos’’ (stranger or outsider) and
‘‘onoma’’ (name). This is in reference to the fact that the ordinal names for both orders include portions of the names for orders to which
they are not closely related.

27. Bläser, M., & Predel, R. (2020)

28. The insect order Mantophasmatodea was described by K.-D. Klass, O. Zompro, N.P. Kristensen, and J. Adis in 2002 (Science 296: 1456–
1459).
The new insect order Mantophasmatodea by KLASS et al. (2002) was based on two lines of morphological evidence: first, that these
insects lack the sets of apomorphies that characterize the other insect orders; second, evidence on the sistergroup relationship of
Mantophasmatodea was conflicting – though Notoptera (= Grylloblattodea) and Phasmatodea appeared to be the most likely candidates.
According to KRISTENSEN (1991, 1995), the Neoptera comprises 10 major lineages, whose relationships have remained unresolved: the
Plecoptera, Orthoptera, Phasmatodea, Dictyoptera, Dermaptera, Embioptera, Notoptera, Zoraptera, Acercaria (= hemipteroid orders),
and Endopterygota (= Holometabola). Although all presently known Mantophasmatodea are entirely wingless (KLASS et al. 2002;
ZOMPRO et al. 2002), their assignment to the Pterygota-Neoptera was shown to be supported by several apomorphies (KLASS et al.
2002). Consequently, the Mantophasmatodea now form an 11th clade in this basal neopteran polytomy.
JUSTIFICATION OF ITS ORDINAL STATUS
Eberhard, 2010
Since Mantophasmatodea (heelwalkers) were described in May 2002 (Klass et al., 2002), most research on the new insect order has been
on their phylogenetic placement within the Hexapoda and on their taxonomy (Picker et al., 2002, Dallai et al., 2003, Klass et al., 2003,
Klass, 2004, Machida et al., 2004, Predel et al., 2004, Terry and Whiting, 2005, Zompro, 2005, Cameron et al., 2006, Beutel and Gorb,
2006, Baum et al., 2007, Klass and Eulitz, 2007, Klug and Klass, 2007, Damgaard et al., 2008). Although the ordinal status of
Mantophasmatodea was disputed (Arillo and Engel, 2006, Tilgner, 2002), They agreed with the arguments of Klass et al., 2003, Klass
et al., 2002 and treat Mantophasmatodea as an insect order.

29. CONCLUSION
No doubt Mantophasmatodeans are beautiful and very interesting creatures on Earth which have
been recently discovered. Their life style and unique behaviours are amazing. Their evolution or
affinities have different point of view among the scientists. But majority believes in their relation
with Grylloblattodea. Recently Grylloblattodea and Mantophasmatodea have been put into a new
clade called, Xenonomia.But the conflict regarding their ordinal status is still there.

30. REFERENCES
1. Klass, K. D., Picker, M. D., Damgaard, J., van Noort, S. I. M. O. N., & Tojo, K. (2003). The taxonomy, genitalic morphology, and phylogenetic
relationships of Southern African Mantophasmatodea (Insecta). Entomologische Abhandlungen, 61(1), 3-67.
2. Zompro, O., Adis, J., & Weitschat, W. (2002). A review of the order Mantophasmatodea (Insecta). Zoologischer Anzeiger-A Journal of Comparative
Zoology, 241(3), 269-279.
3. Klass, K. D., & Grossmann, C. (2009). Mantophasmatodea. In Encyclopedia of Insects (pp. 599-600). Academic Press.
4. Cranston, P. S., & Gullan, P. J. (2009). Phylogeny of insects. In Encyclopedia of insects (pp. 780-793). Academic Press.
5. Klass, K. D., Picker, M. D., Damgaard, J., van Noort, S. I. M. O. N., & Tojo, K. (2003). The taxonomy, genitalic morphology, and phylogenetic
relationships of Southern African Mantophasmatodea (Insecta). Entomologische Abhandlungen, 61(1), 3-67.
6. Terry, M. D., & Whiting, M. F. (2005). Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics, 21(3), 240-257.
7. Klass, K. D., Zompro, O., Kristensen, N. P., & Adis, J. (2002). Mantophasmatodea: a new insect order with extant members in the
Afrotropics. Science, 296(5572), 1456-1459.
8. Eberhard, M. J., Lang, D., Metscher, B., Pass, G., Picker, M. D., & Wolf, H. (2010). Structure and sensory physiology of the leg scolopidial organs in
Mantophasmatodea and their role in vibrational communication. Arthropod Structure & Development, 39(4), 230-241.
9. Roth, S., Molina, J., & Predel, R. (2014). Biodiversity, ecology, and behavior of the recently discovered insect order Mantophasmatodea. Frontiers
in zoology, 11(1), 1-20.
10. Bläser, M., & Predel, R. (2020). Evolution of neuropeptide precursors in Polyneoptera (Insecta). Frontiers in Endocrinology, 11, 197.