Prosepididontus calopteryx HANDLIRSCH 1920, from the upper Lower Jurassic of Germ a ny, is transferred from Trichoptera to Grylloblattida: Geinitziidae. The anal loop like structure of Prosepididontus calopteryx is built of the concave CuP and two anal veins. Prosepididontidae HANDLIRSCH, 1920 is synonymised under Geinitziidae HANDLIRSCH, 1906.Prosepididontus calopteryx HANDLIRSCH 1920, from the upper Lower Jurassic of Germ a ny, is transferred from Trichoptera to Grylloblattida: Geinitziidae. The anal loop like structure of Prosepididontus calopteryx is built of the concave CuP and two anal veins. Prosepididontidae HANDLIRSCH, 1920 is synonymised under Geinitziidae HANDLIRSCH, 1906

Ansorge, J.

Rasnitsyn, A. P.

Revistes Catalanes amb Accés Obert

19
Identity of Prosepididontus calopteryxHANDLIRSCH 1920
(Insecta: Grylloblattida: Geinitziidae)
J. ANSORGE(1) andA.P. RASNITSYN (2)
(1) Institut für Geologische Wissenschaften, Ernst-Moritz-Arndt-Universität Greifswald, Friedrich-Ludwig-Jahn-Straße 17a, Greifswald, D - 17489.
E-mail: ansorge@mail.uni-greifswald.de
(2) Paleontological Institute, Russian Academy of Sciences, Russia 117647 Moscow. E-mail: rasna@online.ru
ACTA GEOLOGICA HISPANICA,v. 35 (2000),nº 1-2,p. 19-23
ABSTRACT
P rosepididontus calopteryx HANDLIRSCH 1920, from the upper Lower Jurassic of Germ a ny, is transferred from Trichoptera to
G ry l l o blattida: Geinitziidae. The anal loop like structure of P rosepididontus calopteryx is built of the concave CuP and two anal ve i n s .
Prosepididontidae HANDLIRSCH, 1920 is synonymised under Geinitziidae HANDLIRSCH, 1906.
Key wo rd s : Trichoptera. Prosepididontidae. Gry l l o blattida. Geinitziidae. Dobbertin. To a r c i a n .
I N T RO D U C T I O N
Geinitz (1880, 1884), Bode (1905) and first of all
Handlirsch (1906-08, 1920-21, 1939) described from the
Upper Liassic of Dobbertin and Braunschweig (Northern
Germany) about 38 species of Trichoptera which Han-
dlirsch assigned but one (Prosepididontus calopteryx) to
Necrotauliidae HANDLIRSCH, 1906. While revising
the type material one of us (J.A.) found amongst the
necrotauliids true Necrotaulius HANDLIRSCH 1906
( p r o b a bly stemgroup of Amphiesmenoptera = Tr i-
choptera + Lepidoptera), representatives of Trichoptera
(e.g. Liadotaulius HANDLIRSCH 1939) and most sur-
prising a number of Lepidoptera with well preserved
wing scales (e.g. Pseudorthophlebia platyptera HAN-
DLIRSCH 1906, ?Paratrichopteridium efossum HAN-
DLIRSCH 1939, ?Paratrichopteridium costale HAN-
DLIRSCH 1939). Finally Metatrichopteridium confusum
HANDLIRSCH 1939 was recognized as a representative
of Diptera: Hennigmatidae SHCHERBAKOV, 1995 (An-
sorge in prep.).
Since its first description from the upper Lower
Jurassic of Dobbertin (Mecklenburg/Germany) (Han-
dlirsch, 1920-21: 200: Fig. 170) P ro s e p i d i d o n t u s
calopteryx (Fig. 1) was always considered representing
its personal family Prosepididontidae HANDLIRSCH
1920 within the Trichoptera (Handlirsch, 1920-21, 1936,
1939; Laurentieux, 1953; Fischer, 1960, 1971; Martyno-
va, 1962; Müller, 1963, 1978, 1989; Malicky, 1973;
Sukatsheva, 1980, 1982, 1989; Carpenter, 1992; Novok-
shonov, 1997). 
S u k a t c h eva (1980, 1982, 1989) included Prosepidi-
dontidae within the Paleozoic trichopteran suborder Pro-
tomeropina. Novo k s h o n ov (1997: 85) rejected this view,
thinking Prosepididontidae to be a representative of any
of the living suborders.
After restudying the holotype of P ro s e p i d i d o n t u s
c a l o p t e r y x it became clear that this species can not be
a representative of the Trichoptera. The reasons for
excluding P. calopteryx from Trichoptera and the ar-
guments for the new placement are discussed in this
p a p e r.
R E D E S C R I P T I O N
O rd e r : G ry l l o blattida WALKER, 1914
Fa m i ly : Geinitziidae HANDLIRSCH, 1906
* 1906 Geinitziidae fam. nov. - Handlirsch: 426
1920 Prosepididontidae fam. nov. - Handlirsch: 200, syn. nov. 
GENUS P ro s e p i d i d o n t u sHANDLIRSCH 1920
P rosepididontus calopteryxHANDLIRSCH 1920
Figures 1-3
* 1920 P rosepididontus calopteryx gen. et sp. nov. - Handlirsch:
200, fig. 170
1936 P rosepididontus calopteryx HANDLIRSCH - Handlirsch:
1542: Fig. 1640
1939 P rosepididontus calopteryx gen. et sp. nov. - Handlirsch:
101, pl. 10, fig. 177
1953 P rosepididontus calopteryx - Laurentieux: 489
1960 P rosepididontus calopteryx HANDLIRSCH 1919 - Fi s-
cher: 9
1963 P rosepididontus calopteryx HANDLIRSCH - Müller: 223,
Fig. 306
1971 P rosepididontus calopteryx HANDLIRSCH 1919 - Fi s-
cher: 7
1978 P rosepididontus calopteryx HANDLIRSCH - Müller: 255,
Fig. 303
1982 P rosepididontus calopterys HANDL. - Sukatcheva: 6, 7,
Fig. 8
1989 unamed figure - Sukatcheva: 442, fig. 1
1989 P rosepididontus calopteryx HANDLIRSCH - Müller: 257:
Fig. 290
1992 P rosepididontus calopteryx HANDLIRSCH, 1920: - Car-
penter: 363, fig. 202,3
1997 P rosepididontus calopterys HANDL. - Novo k s h o n ov: 85
H o l o t y p e : Isolated wing (part and counterp a rt )
housed in Institute of Geological Sciences of Ern s t -
M o r i t z - A rn d t - U n iversity Greifswald (123/88) in a rather
poor state of preserva t i o n .
20
Figure 1. Prosepididontus calopteryx HANDLIRSCH 1920
from the Lower Toarcian of Dobbertin (Mecklenburg/Ger-
many) - original drawing of Handlirsch (1920-21, Fig. 170).
Figure 2. Prosepididontus calopteryx HANDLIRSCH 1920 from the Lower Toarcian of Dobbertin (Mecklenburg/Germany), holotype
housed in Institute of Geological Sciences University of Greifswald (123/88). Scale bar 1 mm.
Type locality: Fo rmer clay pit of Schwinz near Dob-
b e rtin (Mecklenbu rg / G e rm a ny ) .
Type horizon: Carbonate concretions within clay of
the „Green Series”, ammonite zone of H a r p o c e ras fal-
c i fe r u m ( L ower To a r c i a n ) .
D e s c r i p t i o n : Left fore wing, length 9,8 mm, width 3,5
mm, densely covered with microtrichiae. A number of
ve ry faint crossveins and spots is situated in the distal part
of the wing. At least two weak crossveins in the costal
area. Radius branches little before the middle of the wing,
the anterior branch of RP is forked distally, RA distally
not forked. Media branches at one third of the wing
length, distally forked dichotomous. The anterior deeply
f o r ked branch of CuA is fused with M5. CuA3 is wave d
s i g m o i d a l ly, connected with an oblique crossvein with
C u A2. The two convex anal veins form together with con-
c ave CuP an anal loop like stru c t u r e .
D i s c u s s i o n : The obvious reason of earlier authors to
consider P. calopteryx a trichopteran was the well deve-
loped anal loop like structure, that is, three forewing anal
veins meeting each other to form a single veinlet prior
reaching the hind wing margin. At the same time, some
other characters of P ro s e p i d i d o n t u s, part i c u l a r ly the nar-
r ow, pectinate, 3-branched RP, and especially the 3-
branched CuA, are unknown among caddisflies and
mecopteroid orders in general. This casts doubts on the
c u rrent assignments of P ro s e p i d i d o n t u s, the more so that
the anal loop is not unique in caddisflies and their sister
group of Lepidoptera (Willmann, 1989: 48). It has been
acquired independently by hemipterous and hy m e n o p t e-
rous insects, as well as by some booklice. These latter or-
ders have otherwise nothing to do with P ro s e p i d i d o n t u s,
so other groups should be considered in this connection.
Within all the variety of the pterygote orders known
to exist in the Mesozoic, not a single fits features of
Prosepididontus perfectly, even if to forget the anal loop
for the moment. Indeed, dragonflies, mayflies, or-
thopteroid and blattopteroid orders, webspinners, stone-
flies, earwings, beetles and hymenopterans all have a
r a d i c a l ly different wing structure and/or ve n a t i o n a l
scheme. Differences of the mecopteroid orders are men-
tioned above. Likewise, the neuropteroid wings, when
oligoneurous and not much modified, also have short or
no CuA fork and retain traces of compound origin of
their RP (as RP+MA), that is, the forked hind RP branch
and/or rudiment of MA base. Hemipterans and Mesozoic
booklice lack normally developed SC. Miomopterans al-
ways have each M and CuA two-branched. 
The general type of venation, esp. the 3-branched
CuA and the single concave CuP of P ro s e p i d i d o n t u s i n-
dicates a relationship only with gry l l o blattideans (Gry-
l l o blattida). Among the Mesozoic gry l l o blattideans only
Geinitziidae are similar to P ro s e p i d i d o n t u s in having M5,
that is, the convex, usually oblique hind branch of M that
l e aves M subbasally to connect it with CuA. In the Pa-
leozoic realm, more gry l l o blattideans retain M5, so as
some of the archaic Euryptilonidae MART Y N OV, 1940,
p a rt i c u l a r ly Blania falsa K U K A L OVÁ 1964, from the
L ower Pe rmian of Moravia (Czech Republik). Most of
other features are alike in P ro s e p i d i d o n t u s and Geinitzi-
idae as well, viz., long SC meeting C and bearing obl i q u e
veinlets (only two of them seen in P ro s e p i d i d o n t u s m i g h t
21
Figure 3. Prosepididontus calopteryx HANDLIRSCH 1920 from the Lower Toarcian of Dobbertin (Mecklenburg/Germany), drawing
of the holotype. Scale bar 1mm.
be due to its poor preservation state), few RP branches di-
rected obl i q u e ly upwa r d, twice dichotomous M, and
t h r e e - b r a n c h e d, pectinate backward CuA (the latter is pre-
sent in geinitziid genera other than G e i n i t z i a H A N-
DLIRSCH 1906). Unlike P ro s e p i d i d o n t u s, Geinitziidae
h ave not an anal loop like structure, although an initial
step in its direction is possible to see in the paratype of
S h u rabia fe rga n e n s i s RASNITSYN 1983, where the two
posterior anal veins join before reaching the hind wing
m a rgin (Rasnitsyn, 1983, fig. 4b). This makes it possibl e
to infer that P rosepididontus calopteryx represents not
more than an aberrant genus of Geinitziidae, the more so
that the geinitziid stratigraphic range (Upper and, possi-
bly, Middle Triassic of Central Asia, China, Australia and
South Africa, and Lower Jurassic of Germ a ny, England,
Central Asia, East Siberia, Mongolia, and China;
S t o r o z h e n ko, 1997) covers the upper Lower Jurassic, the
type stratum of P. calopteryx. Within Geinitziidae, P ro s e-
p i d i d o n t u s d i ffers from other genera, besides the anal
loop like structure, in that its SC is equidistant to C and R
at the level of the RS base, instead of being closer to the
l a t t e r. Otherwise it is similar to S h u rab i a M A RT Y N OV
because of CuA three-branched and M forking basally of
the RS base [and part i c u l a r ly to Sh. parvula R A S N I T-
SYN 1983 from the Asian Lower Jurassic (locality Ust
B a l ey) due to small size and relative ly close position of
the RS base and the first fork of M]. 
AC K N OW L E D G E M E N T S
A P R ’s visit to Germ a ny in September, 1998, was possibl e
due to ESF Grant and to the hospitality of Dr Wo l f gang Zessin,
Zoo Schwerin, Germ a ny. The research of J.A. is supported by a
postdoctoral grant of German Science Foundation DFG (An 311
1 - 1 ) .
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23


Identity of Prosepididontus calpterys Handlirsch 1920 (Insecta: Grylloblattida: Geinitziidae)

Prosepididontus calopteryx HANDLIRSCH 1920, from the upper Lower Jurassic of Germ a ny, is transferred from Trichoptera to Grylloblattida: Geinitziidae. The anal loop like structure of Prosepididontus calopteryx is built of the concave CuP and two anal veins. Prosepididontidae HANDLIRSCH, 1920 is synonymised under Geinitziidae HANDLIRSCH, 1906

Identity of Prosepididontus calpterys Handlirsch 1920 (Insecta: Grylloblattida: Geinitziidae)

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Identity of Prosepididontus calpterys Handlirsch 1920 (Insecta: Grylloblattida: Geinitziidae)

Abstract

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Full text

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Revistes CientÃfiques de la Universitat de Barcelona