Mantophasmatodea now in the Jurassic

Abstract

The Mantophasmatodea is the most recently discovered insect order. The fossil records of all other 'polyneopteran' orders extend far in the past, but the current absence of pre-Cenozoic fossils of the Mantophasmatodea contradicts a long evolutionary history, which has to be assumed from the morphological distinctness of the group. In this paper, we report the first Mesozoic evidence of a mantophasmatodean from the Middle Jurassic of Daohugou, Inner Mongolia, China. Furthermore, the new fossil shares apomorphic characters with Cenozoic and recent Mantophasmatodea, suggesting a longer evolutionary history of this order.

Full text

ORIGINAL PAPER
Mantophasmatodea now in the Jurassic
Di-ying Huang & André Nel & Oliver Zompro &
Alain Waller
Received: 26 March 2008 / Revised: 15 May 2008 / Accepted: 19 May 2008 / Published online: 11 June 2008
#
Springer-Verlag 2008
Abstract The Mantophasmatodea i s the most recently
discovered insect order. The fossil records of all other
‘polyneopteran’ orders extend far in the past, but the
current absence of pre-Cenoz oic fossils of the Mantophas-
matodea contradicts a long evolutionary history, which has
to be assumed from the morphological distinctness of the
group. In this paper, we report the first Mesozoic evidence
of a mantophasmatodean from the Middle Jurassic of
Daohugou, Inner Mongolia, China. Furthermore, the new
fossil shares apomorphic characters with Cenozoic and
recent Mantophasmatodea, suggesting a longer evolutionary
history of this order.
Keywords Insecta
.
Mantophasmatodea
.
Middle Jurassic
.
Evolution
.
Polyneoptera
.
Phylogeny
Introduction
The Mantophasmatodea, also named African rock crawlers,
heelwalkers, or gladiators, is the most recently discovered
order of insects (Klass et al. 2002). Extant Mantophasmatodea
are known from Namibia, Tanzania, and South Africa
(Picker et al. 2002;Zomproetal.2003). While nearly all
the ‘polyneopteran’ orders are known to be very ancient, with
oldest records ranging at least between the Early Carbonifer-
ous to the Triassic, fossil rock crawlers are only described
from the Eocene Baltic amber (Zompro 2001, 2005;Klasset
al. 2002;ArilloandEngel2006). No pre-Cenozoic insects
can be attributed to this newly discovered order. The
affinities of these insects were long considered controversial
due to the few supporting morphological apomorphies and
contradictions in molecular analyses (Klass et al. 2002; Terry
and Whiting 2005; Cameron et al. 2006;Klass2007).
Mantophasmatodea have to be considered the sister group of
the modern Dictyoptera (Klass et al. 2003), Mantodea
(Dallai et al. 2003), Orthoptera (Tilgner 2002;Zompro
2005), Grylloblattodea (Terry and Whiting 2005;Kjeretal.
2006; Arillo and Engel 2006), or Phasmatodea (Cameron
et al. 2006). Hamilton (2003) even synonymized the taxon
with the Carboniferous Cnemidolestodea but Zompro (2005)
convincingly rejected this opinion. Gorochov (2007)sug-
gested, without argument, that the Mantophasmatodea could
be a subgroup of the Permo-Triassic order Titanoptera.
However, on the basis of the wing venation, Béthoux (2007)
argued that Titanoptera belongs to the orthopteroid lineage.
In conclusion, we agree with Damgaard et al. (2008,pp.
459
–460) that the ‘sister-group relationship of Mantophas-
matodea to Grylloblattodea or some other neopteran lineage
is far from settled.’
Naturwissenschaften (2008) 95:947–952
DOI 10.1007/s00114-008-0412-x
Communicated by G. Mayr
D.-y. Huang (*)
Nanjing Institute of Geology and Palaeontology,
State Key Laboratory of Palaeobiology and Stratigraphy,
Chinese Academia of Sciences,
Nanjing 210008, People’s Republic of China
e-mail: huangdiying@sina.com
A. Nel (*)
:
A. Waller
CNRS UMR 5202, CP 50,
Entomologie, Muséum National d’Histoire Naturelle,
45 Rue Buffon,
75005 Paris, France
e-mail: anel@mnhn.fr
O. Zompro
Sungaya-Verlag,
Helsinkistraße 52,
24109 Kiel, Germany
e-mail: o.zompro@t-online.de

The Palaeozoic and Mesozoic fossils that are currently
included into the Grylloblattodea share no apomorphies with
their modern representatives. Grylloblattodea were considered
to be paraphyletic by Storozhenko (2002) who studied the
fossil ‘grylloblattids’ and are not characterized by any
apomorphy. The Mesozoic and Palaeozoic ‘grylloblattodea’
is a ‘waste basket’ that replaces the former polyphyletic group
‘Protorthoptera,’ and should be completely reevaluated. Until
now, there was still no evidence supporting the existence of
Mesozoic Grylloblattodea sensu stricto, with the same
applying to the Palaeozoic and Mesozoic fossils currently
attributed to the Phasmatodea, and Tilgner (2001) considered
the oldest unambiguous Phasmatodea to be Cenozoic. Pre-
Cenozoic fossils that have been assigned to Phasmatodea
have problematic relationships (Nel et al. 2004).
The fossil reported in this paper shows several apomorphic
features of the Mantophasmatodea. It comes from the Middle
Jurassic (165 Mya) of Daohugou, Inner Mongolia, China,
which has also yielded plants, insects, co nchostracans,
anostracans, spiders, salamanders, pterosaurs, and mammals
(Huang et al. 2006).
Materials and methods
The female holotype (NIGP 142171) is deposited in the
Nanjing Institute of Geology and Palaeontology, Chinese
Academy of Sciences. It is exquisitely fossilized as part and
counterpart on a slab of volcanic tuff. The exam was
realized without alcohol and under alcohol.
Systematic paleontology
Order Mantophasmatodea (Klass et al. 2002)
Family Mantophasmatidae (Klass et al. 2002)
Subfamily Raptophasmatinae (Zompro 2005)
Emended diagnosis This subfamily is supposed to be
uniquely Eocene (Baltic amber). The present discovery
implies that its geological range is wider. The head of the
Eocene Raptophasmatinae is triangular in anterior view, which
is not the case in Juramantophasma; thus, this character has
to be excluded from the diagnosis of the subfamily.
Genus Juramantophasma gen. n.
Type species Juramantophasma sinica sp. n.
Diagnosis The insect was d iagnosed as Raptophasmatinae,
with head round in an anterior aspect; mandibles armed
with strong teet h, especially the right one ; sclerotized
elongate dorsal process of third tarsomere spine-like; both
female cerci and paraprocts hook-like; pronotum more or
less rounded and mesonotum shorter than wide; metanotum
with postero-lateral angles; hind legs distinctly not saltatorial;
arolium large, projecting beyond claws; pro- and mesofemora
lacking definite, prominent spines; profemora with small
granulae; and ventro-apical spines absent on tibiae.
Juramantophasma differs f rom Raptophasma in i ts
profemora with small granulae, instead of being with only
small bristles, and in the round head, which is triangular in
Raptophasma. Praedatophasma (Zompro et al. 2002)has
also a round head but differs from Juramantophasma in the
presence of spines on the body and of ventral spines on the
prothoracic tibiae (Zompro et al. 2002).
Etymology Genus named after Jura and
Mantophasma.
Juramantophasma sinica sp. n.
Material Only the holotype was included (NIGP 142171a, b).
This specimen is a nearly complete adult female in dorsal–
ventral compression that split in the median section of the insect.
Thus, we designate 142171a as the part preserved with head
structures and postabdominal appendages and 142171b as the
counterpart with thorax and most parts of the legs (Fig. 1a–l).
Locality and age Middle Jurassic Jiulongshan Formation at
Daohugou, Ningcheng County, Inner Mongolia, Northeast
China.
Etymology The species was named after the Latin name for
China.
Diagnosis The insect was diagnosed as for genus.
Description The female holotype was 34 mm long. The
hypognathous head covered with fine fur-like setae, 5.1 mm
long and 4.4 mm wide; large eyes, 1.7 mm long and
Fig. 1 Juramantophasma sinica gen. and sp. n., holotype Nigpas
142171a-b. a General habitus of part . b General habitus of
counterpart. c Details of head under alcohol (142171a) showing
antenna, eyes, mandibles, and maxillary hooks. d Female postabdo-
minal appendages of 142171a, showing ovipositor, cerci, and para-
procts. e Enlargement of right middle leg of 142171a, showing setae
arrangements of femora and tibia. f Enlargement of thorax (142171b),
showing pronotum, mesonotum, and metanotum. g 142171a, showing
arrangements of eggs. h Details of left hind tarsi of 142171b, showing
euplantulae of tarsomeres, dorsal process of third tarsomere and
pretarsal arolium with dorsal setae rows. i Details of right hind tarsi of
142171b, showing dorsal process of third tarsomere and pretarsal
arolium. j Details of right middle tarsi of 142171b, showing dorsal
process of third tarsomere and pretarsal arolia. k Enlargement of g,
showing details of eggs. l Enlargement of g, showing small points
within circular ridge of eggs. an antenna, ce cercus, ch chorion, cr
circular ridge, dp dorsal process of third tarsomere, e eye, fe femur, ma
mandible, mh maxillary hook, go gonapophyse 8 of ovipositor, pa
pretarsal arolium, ti tibia. Scale bars 10 mm in a and b, 2 mm in e and
j, 1 mm in c, d, f, i, k; 500 μminl
b
948 Naturwissenschaften (2008) 95:947–952

1.3 mm wide, but no visible ocelli (Fig. 1a–b); antennae
filiform, with basal five antennomeres preserved; large
scape, 0.8 mm long; pedicel, 0.25 mm long; first flagellomere,
0.95 mm long; second flagellomere, 0.8 mm long; both
mandibles with three strong teeth but not symmetrical; left
mandible stronger, broader, and with larger teeth than the
right one, with apical tooth and first marginal tooth
relatively sharp, second marginal tooth very blunt; right
mandible armed with a sharper apical tooth, first marginal
tooth similar to left one, second marginal tooth smaller than
Naturwissenschaften (2008) 95:947–952 949

left one, with a nearly orthogonal tip; all teeth bearing clear
cutting margins, a topmost ridge visible in all teeth of the
right mandible; maxillary hooks (laciniae) with two apical
teeth; and an api cal maxillary hook distinctly larger than the
second (Fig. 1c).
The thorax (Fig. 1f) is covered with fine short setae
elongate 11.0 mm long; pronotum, 3.5 mm long and
4.5 mm wide, more or less rounded rather than square,
with slightly protruding anterior and posterior margins and
arched lateral margins; mesonotum trapezoidal, distinctly
shorter than wide, 3.2 mm long, 4.8 mm wide; metanotum
rectangular, distinctly shorter than wide, 3.3 mm long and
4.0 mm wide, with small postero-lateral angles pointed
backward.
The legs are covered with fine setae (Fig. 1e); all coxae
protruding laterally due to taphonomic compression, mid
coxae larger than hind ones; anterior margins in wave shape
with a dense row of setae; hind coxae with rather straight
anterior margins and a row of dense setae; trochanters
smaller than coxae, fore- and mid-trochanters more or less
triangular, fore trochanters smallest, hind trochanters more
rounded, setae on troc hanters arranged in bands; fore
femora, 5.4 mm long and 2.2 mm wide; fore tibia,
5.3 mm long; mid femora, 4.5 mm long and 1.7 mm wide;
tibia, 3.9 mm long; hind femora, 6.0 mm long and 1.5 mm
wide; tibia, 7.0 mm long; tibiae with inner side lacking hair
but limited by two rows of long setae, and outer side
covered with small setae; femora with inner side without
hair and limited by two rows of small setae, median part
without hairs, and outer side setose, basal part without
setae; sclerotized elongate dorsal process of third tarsomere
spine-like, slightly curved, but not pointed apically; four
basal tarsomeres bearing very large euplantulae; arolia of
all legs fan-like, very large, and broad (Fig. 1h –j), those of
the forelegs the smallest, mid-legs the largest; ten dorsal
processes, each process armed with a long seta; claws
hook-shaped and sharp near apex in all legs, very large in
fore legs; all tarsi rather short, in hind legs first tarsomere
broadest and longest, first to fourth tarsomeres gradually
shorter and narrower, fifth tarsomere elongate bu t shorter
than the first one, with increased width.
Abdomen is 21 mm long and 5 mm wide, filled by 28
visible eggs (approximately 3.3 mm long and 1.6 mm
wide), anterior most part of abdomen with only two rows
of eggs, middle part with four rows of eggs, posterior
part with three rows of eggs (Fig. 1k–l); apex of
abdomen very well preserved, with all structures clearly
visible; last abdominal tergum lob e-like; seco nd last
tergum narrower but longer than third last, wi th distinct
postero-lateral a ngles; third last tergum very short, short-
est in its median section; a thin but sclerotized gonapo-
physe 8 of ovipositor almost covered by last abdominal
tergum, with tapering shape and a cut tip, arm ed with
marginal hair pointed inner and backward; cerci one-
segmented, hooked, w ith a strong base, posterior section
abruptly thinner and more or less incurved; two hooked,
sharp, incurved at apex, and strong structures are visible
between the cerci, but they are s horter than them. They
probably correspond to the gonoplacts IX ‘gl9’ sensu
Klass et al. (2003;Fig.1d).
Discussion
The general body shape and leg structures of this insect
correspond to those of a ‘Polyneoptera.’ Because it has
unsegmented cerci, the fossil could not be considered to
belong to any extant group of Grylloblattodea, which have
segmented cerci. Even the few fossil taxa currently
attributed to the Grylloblattodea, with preserved body
structures, have multi-segmented cerci (i.e., Blattogryllidae
Rasnitsyn 1976). Among Hexapoda with unsegmented
cerci, the presence of large mandibles excludes the Diplura.
The plesiomorphic condition of five-segmented tarsi of
Juramantophasma is present not only in the Mantophas-
matodea but also in the Mesozoic Dermaptera, whereas all
recent representatives of the other ‘polyneopteran’ orders
have unsegmented cerci.
Juramantophasma exhibits several characters currently
considered as apomorphies of the Mantophasmatodea, i.e., a
third tarsomere with a sclerotized elongated dorsal process;
enlarged and fan-like pretarsal arolia with a clearly visible row
of dorsal setae, identical to those of the modern Mantophasma
zephyra (Zompro et al. 2002
, as figured in Klass et al. 2002
and Beutel and Gorb 2006, Fig. 3b); last tarsomere making a
right angle with the others, keeping it up in the air; female
gonoplacs (valves 3) short and claw-shaped; and egg with a
circular ridge. All of these characters are completely different
from those of the modern Phasmatodea, Grylloblattodea (see
Zompro 2001; Beutel and Gorb 2001 and 2006, Fig. 4), and
Dermaptera (see Haas and Gorb 2004), which exclude
affinities with these insects.
The following characters of our fossil are further shared
with the Mantophasmatodea: hypognathous head, wings
absent, antennae probably long, and filiform after the shape
of the preserved part, ocelli not visible, probably absent,
pronotum longer than meso- and metanotum; hind legs very
thin but longer than the middle and forelegs; fore femora
are the widest; four basal tarsomeres very short (long in
Phasmatodea, except for ground dwelling forms, e.g.,
Agathemera, some Aschiphasmatidae, Heteropterygidae,
and Eurycanthinae), and bearing very large euplantulae
(very small in modern Grylloblattodea); abdominal seg-
ments transverse; ovipositor elonga te, nearly reaching the
apex of abdomen; egg large, elongate, and a chorion with a
pattern of small spots and a central gibbosity.
950 Naturwissenschaften (2008) 95:947–952

The phylogenetic relationships of Juramantophasma
within the Mantophasmatodea are difficult to establish
because of the lack of a phylogenetic analysis of this order
based on morphological characters. The available analyses
are based on molecular data only (Klass et al. 2003;
Damgaard et al. 2008). We can only base our analysis on
typologica l c ompa ri son with other Mantophasm at odea .
Juramantopha sma shares with the Mantophasmatidae
(Klass et al. 2002) hind legs distinctly not saltatorial,
arolium large, projecting beyond claws. The absence of
ventro-apical spines on the tibiae is a difference to the
Mantophasmatinae, as Zompro (2005) characterized this
group on the basis of the presence of two ventro-apical
spines on tibiae. Juramantophasma shares with the Baltic
amber Raptophasmatinae (Zompro 2005), its prothoracic
tibiae lacking definite ventral spines.
The mandibles of Juramantophasma seems to be less
specialized than those of the recent Mantophasmatodea,
especially the left one with three strong teeth, instead of one
apical tooth well separated from two distinctly smaller teeth
(Baum et al. 2007). Nevertheless, the difference is rather
weak, especially with the mandibles of the recent Man-
tophasma. Unfortunately, little is known on the mandibles
of the Baltic amber Mantophasmatodea. The mandibles of
this Jurassic taxon look similar to those of the recent
Ensifera. Juramantophasma was probably a predator (e.g.,
mandible with strong teeth; broad fore legs with strong
hook-like claws) but less specialized than the recent rock
crawlers, supporting Gorochov’s(2006) hypothesis on the
Jurassic diversification of predatory ‘polyneopterous’
groups that ‘tried to occupy’ the niches that remained
empty between the extinction of the Titanoptera and the
diversification of the Mantodea.
At this stage, it is not possible to infer anything on the
relationships between Mant ophasmatodea and the other
‘polyneopterous’ orders from the present discovery because
Juramantophasma belongs to the Mantophasmatodea sensu
stricto and also because of the great confusion in the
systematics and phylogenetic affinities of the fossil taxa
currently included in the Grylloblattodea.
Nevertheless, the present discovery of a middle Jurassic
Mantophasmatodea does not fit well the assumption of Arillo
and Engel (2006, p. 8) that the ‘Mantophasmatodea and
perhaps also their sister Grylloblattodea as they differentiated
from stem-group families such as Blattogryllidae in the
Jurassic or Early Cretaceous.’ Mantophasmatodea are clearly
much older than the middle–late Jurassic Blattogryllidae.
Juramantophasma documents that the Mantophasmatodea
were present by 165 Ma ago, 120 Ma o lder than the
previous record, and before the modern lineages of the
Orthoptera are known to appear (Béthoux and Nel 2002),
which contradict s the hypothesis of Tilgner (2002).
This suggests that these insects could be much older
(Palaeozoic). The difficulties in resolving the phylogenetic
relationships of the Mantophasmatod ea within the ‘ Poly-
neoptera
’ are probably due to the great antiquity and rapid
diversification of the latter (Whitfield and Kjer 2008).
Despite this new Mesozoic record, the Mantophasmato-
dea apparently remained a cryptic order in recent and past
worlds, compared to the predaceous mantises that began to
diversify in the Early Cretaceous (Grimaldi and Engel
2005).
Acknowledgment We thank Dr. P. Grandcolas and Prof. N. P.
Kristensen for their helpful criticisms and comments. H.D.Y. is
pleased to acknowledge this project supported by the NSFC (grants
no. 40672013 and 40632010), the Major Basic Research Projects of
MST of China (2006CB806400), the State Key Laboratory of
Palaeobiology and Stratigraphy (Nigpas, no. 073101), and the
MNHN, Paris, for a grant as invited Maître de Conférence. We
sincerely thank Dr. Gerald Mayr, Dr. Karl Kjer, and two anonymous
referees for the useful comments on the first version of this paper.
References
Arillo A, Engel MS (2006) Rock crawlers in Baltic amber (Notoptera:
Mantophasmatodea). Am Mus Novit 3539:1– 10
Baum E, Dressler C, Beutel RG (2007) Head structures of
Karoophasma sp. (Hexapoda, Mantophasmatodea) with phylo-
genetic implications. J Zool Syst Evol Res 45:104–119
Béthoux O (2007) Cladotypic taxonomy applied: titanopterans are
orthopterans. Arthropod Systematics & Phylogeny 65:135–156
Béthoux O, Nel A (2002) Venation pattern and revision of Orthoptera
sensu nov. and sister groups. Phylogeny of Palaeozoic and
Mesozoic Orthoptera sensu nov. Zootaxa 96:1–88
Beutel RG, Gorb SN (2001) Ultrastructure of attachment special-
izations of hexapods (Arthropoda): evolutionary patterns inferred
from revised ordinal phylogeny. Z Zool Syst Evol 39:177–207
Beutel RG, Gorb SN (2006) A revised interpretation of the evolution of
attachment structures in Hexapoda with special emphasis on
Mantophasmatodea. Arthropod Systematics & Phylogeny 64:3–25
Cameron SL, Barker SC, Whiting MF (2006) Mitochondrial genomics
and the new insect order Mantophasmatodea. Mol Phyl Evol
38:274–279
Dallai R, Frati F, Lupetti P, Adis J (2003) Sperm ultrastructure of
Mantophasma zephyra (Insecta: Mantophasmatodea). Zoomor-
phology 122:67–76
Damgaard J, Klass K-D, Picker MD, Buder G (2008) Phylogeny of
the heelwalkers (Insecta: Mantophasmatodea) based on mtDNA
sequences, with evidence for additional taxa in South Africa. Mol
Phyl Evol 47:443–462
Gorochov AV (2006) New and little known orthopteroid insects
(Polyneoptera) from Fossil resins: communication 1. Paleontol J
40:646–654
Gorochov AV (2007) The first representative of the suborder
Mesotitanina from the Paleozoic and notes on the system and
evolution of the order Titanoptera (Insecta: Polyneoptera).
Paleontol J 41:621–625
Grimaldi D, Engel MS (2005) Evolution of the insects. Cambridge
University Press, Cambridge, p 755
Haas F, Gorb SN (2004) Evolution of locomotory attachment pads in the
Dermaptera (Insecta). Arthropod Structure & Development 33:45–66
Hamilton KGA (2003) Cnemidolestodea: an ancient insect order
resurrected. Biodiversity 4:22–24
Naturwissenschaften (2008) 95:947–952 951

Huang D-Y, Nel A, Shen Y-B, Selden P, Lin Q-B (2006) Discussions
on the age of the Daohugou fauna —evidence from invertebrates.
Prog Nat Sci 16(Special Issue):308–312
Kjer KM, Carle FL, Litman J, Ware J (2006) A molecular phylogeny
of Hexapoda. Arthropod Systematics & Phylogeny 64:35–44
Klass K-D (2007) Di e Stammesgeschichte der Hexapoden: eine
kritische Diskussion neuerer Daten und Hypothesen. Denisia
20:413–450
Klass K-D, Zompro O, Kristensen NP, Adis J (2002) Mantophasmatodea:
a new insect order with extant members in the Afrotropics. Science
296:1456–1459
Klass K-D, Picker MD, Damgaard J, Van Noort S, Tojo K (2003) The
taxonomy, genitalic morphology, and phylogenetic relationships
of South African Mantophasmatodea (Insecta). Entomol Abh
61:3–67
Nel A, Marchal-Papier F, Béthoux O, Gall J-C (2004) A “stick insect-
like” from the Triassic of the Vosges (France) (Insecta: “pre-
Tertiary Phasmatodea”). Ann Soc Entomol Fr 40:31–36
Picker MD, Colville JF, Van Noort S (2002) Mantophasmatodea now
in South Africa. Science 297:1475
Storozhenko SY (2002) Chapter 2.2 .2.2.1. Order Grylloblattida
Walker, 1914 (= Notoptera Crampton, 1915, = Grylloblatto dea
Brues et Melander, 1932, + Protorthoptera Handlirsch, 1906, =
Paraplecoptera Martynov, 1925, + Protoperlaria Tillyard, 1928).
In: Rasnitsyn AP, Quicke DLJ (eds) History of insects. Kluwer,
Dordrecht, pp 278–284
Terry MD, Whiting MF (2005) Mantophasmatodea and phylogeny of
the lower neopterous insects. Cladistics 21:240–257
Tilgner E (2001) The fossil record of Phasmida (Insecta: Neoptera).
Insect Syst Evol 31:473–480
Tilgner E (2002) Mantophasmatodea: a new insect order? Science
297:731
Whitfield JB, Kjer KM (2008) Ancient rapid radiations of insects:
challenges for phylogenetic analysis. Annu Rev Entomol
53:449–472
Zompro O (2001 ) The Phasmatodea and Raptophasma n. gen.,
Orthoptera incertae sedis, in Baltic amber (Insecta: Orthoptera).
Mitt Geol-Pal Inst Univ Hamburg 85:229–261
Zompro O (2005) Inter- and intra-ordinal relationships of the
Mantophasmatodea, with comments on the phylogeny of poly-
neopteran orders (Insecta: Polyneoptera). Mitt Geol-Pal Inst Univ
Hamburg 89:85–116
Zompro O, Adis J, Weitschat W (2002) A review of the order
Mantophasmatodea (Insecta). Zool Anz 241:269–279
Zompro O, Adis J, Bragg PE, Naskrecki P, Meakin K, Wittneben M,
Saxe V (2003) A new genus and species of Mantophasmatidae
(Insecta: Mantophasmatodea) from the Brandberg Massif, Namibia,
with notes on behaviour. Cimbebasia 19:13–24
952 Naturwissenschaften (2008) 95:947–952