Bulletin of the Czech Geological Survey, Vol. 77, No.

3, 167–182, 2002
© Czech Geological Survey, ISSN 1210-3527

Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

IVO CHLUPÁČ1 – VRATISLAV KORDULE2

1Institute of Geology and Palaeontology, Charles University, Albertov 6, 128 43 Praha 2, Czech Republic
2Dlouhá 104, 26 101 Příbram III, Czech Republic

A b s t r a c t . Rare non-trilobite arthropods, mostly with affinities with the Burgess Shale fauna, are described from the Middle Cambrian Jince For-
mation of the Barrandian area, Central Bohemia (Czech Republic): Tuzoia sp., another large Tuzoia-like arthropod (probably a new genus), Probosci-
caris hospes sp. n., Helmetia? fastigata sp. n., and Forfexicaris? sp. The status of Pilocystites primitius Barrande, 1887, formerly ranged with
echinoderms but here recognized as a Tuzoia fragment, is discussed. Most of described arthropods (except Forfexicaris? sp.) are thought to be nekto-
benthic animals of a shallow-water environment, possibly tolerant to some salinity fluctuations. Relationships, particularly to North American faunas,
indicate open migration possibilities between Laurentia and peri-Gondwana regions during the Middle Cambrian.

A b s t r a k t . Ze středního kambria Barrandienu (převážně jineckého souvrství) jsou popsány vzácné zbytky arthropodů se vztahy k burgesským
břidlicím západní Kanady. Jde o zástupce rodu Tuzoia Walcott (zde označené Tuzoia sp., sem patří i zbytek popsaný jako Pilocystites primitius Bar-
rande, 1887, dosud s rezervou řazený k ostnokožcům), dále zbytek jiného velkého členovce příbuzného rodu Tuzoia (patrně nový rod), prvý mimoa-
merický zástupce rodu Proboscicaris Rolfe (P. hospes sp. n.) a čeledi Helmetiidae – Helmetia? fastigata sp. n. K arthropodům nejistého
systematického postavení jsou řazeny kruhovité štítky prozatímně označené jako Forfexicaris? sp. Většina popsaných zbytků pochází ze svrchní části
jineckého souvrství z okolí Jinců (blízké podloží zóny Paradoxides gracilis až zóna Ellipsocephalus hoffi-Rejkocephalus), přibližný ekvivalent baltic-
ké trilobitové zóny Paradoxides paradoxissimus a báze zóny P. forchhammeri. Organismy jsou hodnoceny jako nektobentózní, snad tolerantní i k sa-
linitním výkyvům, Forfexicaris? sp. mohl patřit k pelagickým formám. Vztahy k severoamerickým faunám svědčí o otevřených možnostech migrací
mezi Laurentií a perigondwanskými oblastmi během středního kambria.

K e y w o r d s : Arthropoda, Phyllocarida and incertae sedis, Middle Cambrian, Barrandian area, Czech Republic

Introduction The laboratory work was partly assisted by a Ministry

of Education grant No. MSM-113100006.
The marine Middle Cambrian strata of the Jince For-
mation in the Barrandian area, Central Bohemia, are
world-famous particularly for well preserved trilobite and Systematic part
echinoderm faunas, accompanied by locally frequent ich-
nofossils, hyolithids, brachiopods, and acritarchs. Repre- Phylum Arthropoda
sentatives of other fossil groups such as ostracods, Class Malacostraca Latreille, 1806
gastropods, foraminifers, and algae as well as of some Subclass Phyllocarida Packard, 1879
problematical groups are rare. Remains of large non-trilo- Order and Family uncertain
bite arthropods are also very rare finds and mostly were
not correctly recognized in the past. Genus Tuzoia Walcott, 1912
Collecting activity during the last fifty years, particu-
larly in the Jince area, has revealed some remarkable finds T y p e s p e c i e s : Tuzoia retifera Walcott, 1912,
of non-trilobite arthropods with relationships to the fam- Burgess Shale, Middle Cambrian.
ous Middle Cambrian Burgess Shale fauna of North Amer- D i a g n o s i s (comp. Rolfe 1969): Carapace bivalve,
ica. These are the subjects of the present report. hinge line distinct. Outline of carapace valves subelliptical,
Regrettably, the new materials are incomplete in many with distinct anterior carapace horn. Sculpture reticulate,
respects, frequently allowing only a tentative determina- prominent lateral ridge, carinate, with smaller reticulae than
tion at the generic level. Consequently, open nomenclature on main areas of valves. Spines or spinules of different
is commonly used. length and number protrude from the margin of valves (ex-
The reference material is housed in the collections of cept the anterior margin, spines may occur also on the late-
the National Museum, Prague (inventory numbers pre- ral ridge). Doublure along the ventral margin is flat.
fixed by L), in the collections of the Czech Geological R e m a r k s . Tuzoia was introduced by Walcott (1912)
Survey, Prague (inventory numbers prefixed by VK = col- in describing the Burgess Shale fauna. Other species of this
lection of Vratislav Kordule, SZ = coll. M. Szabad). Three genus were described later from North America (particu-
specimens from the collection of the late Mr. J. Bouška are larly by Resser 1930, Robison and Richards 1981), China
deposited in the West Bohemian Museum in Plzeň (num- (Endo and Resser 1937, 1957) and South Australia (Glaess-
bers prefixed by MP-S). ner 1979). No representatives have been so far reported
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Ivo Chlupáč – Vratislav Kordule

Fig. 1. Tuzoia sp., incomplete right carapace valve MP-S01748 (length 155 mm). Jince Formation, Jince. A – entire fossil remain; B – dorsolateral part
of the same specimen with the dorsal margin and dorsolateral part of the lateral ridge; C – the same with the reticulate sculpture accentuated by
drawing. Scale 1:1.

from Europe, though Barrande (1887) described under the most appropriate approach (the abdomen and appendages
name Pilocystites primitius a fragment with characteristic are not known; see discussion in Glaessner 1979).
reticulate Tuzoia-like sculpture, which he regarded as an S p e c i e s : Tuzoia retifera Walcott, 1912 (synonyms:
echinoderm remnant (discussion in this paper). T. burgessensis Resser, 1930, T. canadensis Resser, 1930),
Tuzoia was classified by Raymond (1935) as a separate Middle Cambrian, Burgess Shale;
family Tuzoiidae, raised by Simonetta and Delle Cave T. australis Glaessner, 1979, Lower Cambr., South Aus-
(1975) to the rank of a separate order. This was followed tralia;
by Hou and Bergström (1997), who ranged this order with T. bispinosa Yuan et Zhao, 1999, Lower or Middle Cambr.,
arthropods of uncertain class within the Phyllum Schizo- China (Guizhou);
ramia. However, Rolfe (1969) classified Tuzoia as Phyllo- T. getzi Resser, 193O, Lower Cambr., Pennsylvania;
carida of uncertain order and family that seems to be the T. guntheri Robison et Richards, 1981, Middle Cambr., Utah;

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 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

T. magna Yuan et Zhao, 1999, Lower or Middle Cambr.,

China (Guizhou);
T. manchuriensis Resser et Endo in Resser, 1930, Middle
Cambr., China (Manchuria);
T. nodosa Resser, 1930, Lower Cambr., British Columbia;
T.? parva (Walcott, 1912), Middle Cambr., Burgess Shale;
T. polleni Resser, 1930, Lower Cambr., British Columbia;
T. praemorsa Resser, 1930, Middle Cambr., Burgess Shale;
T. sinensis P’an, 1957, Lower Cambr., South China (Yunnan). B
T. spinosa Resser, 1930, from the Lower Cambrian of
A
British Columbia is based on insufficient material. T. nitida
Resser et Howell, 1938, and T. vermontensis Resser et How-
ell, 1938, from the Lower Cambrian of eastern North Amer-
ica (Appalachians), are poorly known as commented by
Glaessner (1979). T.? parva can be regarded as a juvenile
ontogenic stage of Tuzoia as reported by Briggs (1977). C
Probable of separate generic position are Tuzoia? dun-
bari Resser, 1930, from the Lower Cambrian of Pennsyl- A
vania (marked by extremely developed spines, comp.
Resser 1930, Resser and Howell 1938) and Tuzoia? pe-
terseni Robison et Richards, 1981, from the Middle Cam-
brian of Utah, with a very long posterodorsal spine (lateral D
ridge and sculpture are unknown).
S t r a t i g r a p h i c r a n g e : late Lower up to late Mid-
Fig. 2. Tuzoia sp. from the Jince Formation, Jince area. A – incomplete
dle Cambrian. right carapace valve VK41, Vinice; B – fragmentary right carapace valve,
G e o g r a p h i c d i s t r i b u t i o n : North America L36483, counterpart, Vinice; C – fragment of the ventral doublure show-
(western and eastern parts), China, Australia, Central ing the external transition into the reticulate sculpture, SZ169; D – in-
Europe (Barrandian area, here reported). complete right carapace valve with partly preserved lateral ridge,
posteroventral spines and exposed doublure, VK40, Vinice. Scale A, B,
D × 0,5, C × 2. Comment to all drawings: dotted lines indicate second-
Tuzoia sp. ary damages of the specimens (broken limits etc.).
Pl. I, figs A-D; pl. II, figs A-H; text-figs 1–4

M a t e r i a l : 6 incomplete carapace valves showing diag- The other two more completely preserved specimens
nostic features of Tuzoia, other 8 carapace fragments with (VK40 – fig. 2D, pl. I, figs A-C, VK41 – fig. 2A, pl. II, figs
characteristic reticulate sculpture and few other characters. A-D) show at least partly preserved subelliptical outline of
D e s c r i p t i o n . The most completely preserved speci- carapace valves. A moderately developed anterior horn
men (MP-SO1748, fig. 1) represents the right carapace projection and the straight dorsal margin is shown in
valve with almost complete dorsal margin (length 120 VK41, the lateral ridge with sharp apex and markedly
mm), broken anterodorsal horn, well traceable lateral smaller and densely packed reticulae, and two spines (8.2
ridge and incomplete ventral part of the valve. The raised and 3.1 mm long) partly preserved on the posteroventral
longitudinal ridge (preserved length 139 mm) shows a margin can be observed in VK40. The typical reticulate
sharp crest-like apex for most part of its course. Pos- sculpture, consisting of reticulae of polygonal, markedly
terodorsal termination is protruded into a short and broad
horn-like projection, not prolonged into a spine. Anterior
and ventral margins are broken off; posterior margin forms
a gently posterior-bent arc. The characteristic sculpture
consists of marked reticulation of the whole surface. Indi-
vidual reticulae are of polygonal, most commonly pentago-
nal and hexagonal outline, obvious diameter 4–6 mm
(maximum 7 mm). Reticulae gradually diminish towards
the margins of the valve and a still more pronounced di-
minishing of reticulae is marked close to the lateral ridge
and on its surface (observable up to the distance of 4–5 mm
from the apex of the carina in dorsal and ventral direction).
Apart from wrinkles caused by compaction, unsharp, lon-
gitudinally projected depressions, interpreted as probable
traces of scavengers, radiate from the lateral ridge in dorsal, Fig. 3. Tuzoia sp., reconstruction of the left carapace valve from the Jince
posterodorsal and posteroventral directions. Formation, presumed length 110–180 mm.

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Ivo Chlupáč – Vratislav Kordule

Fig. 4. Tuzoia sp. incomplete right carapace valve MP-S01749, Jince Formation, Jince. Large specimen of preserved incomplete diameter 150 mm.
A – lightened from the dorsal side; B – the same with reticulate sculpture accentuated by drawing. Scale around 1:1.

variable outline (pentagonal and hexagonal shape obvious, dering the outer margin (the same situation is in the dou-
maximum diameter around 5 to 6 mm) is distinctive and blure fragment SZ169, fig. 2C, pl. ID).
well observable in both specimens. Remarkable is the Remarkable is the specimen MP-S01749 (fig. 4),
presence of doublure in the anterolateral border region of which is interpreted as an incomplete right carapace valve,
the valve of VK40 (fig. 2D, pl. I A, C), where it is exposed with partly preserved lateral ridge with sharp crest and
in the anterolateral border region 33 mm long. The dou- crowded smaller reticulae in its proximity. The broadly ar-
blure is smooth – except for two very narrow sharp lines cuate ventral margin lacks spines (they were evidently
(ridges) parallel to the outer margin of the valve. The con- more posterior).
nection of the doublure with the outer reticulate side of the All reported specimens exhibit the characteristic reti-
exoskeleton is indicated by imprints of small reticulae bor- culate sculpture consisting of polygonal reticulae, cover-

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 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

ing the entire surface, becomes markedly denser and com- All our specimens referred to Tuzoia sp. indicate large
posed of smaller reticulae near the margins and in proxim- dimensions: the extrapolated length of the carapace valves
ity of the lateral carina, where the ridges, forming the ranges between 110 and 160 mm, or even more (in
characteristic reticulae, are thickened. MP-S01749 is the preserved length of the incomplete cara-
The specimen L36481 (pl. IIH, preserved incomplete pace valve 145 mm, the total extrapolated length 170–180
length 71.5 mm, coll. Dr. M. Široký) represents fragments mm). These dimensions are larger than in specimens de-
of two carapace valves approached along the straight scribed from other areas and exceed even the measurements
hinge line. The very distinct polygonal reticulate sculpture of the type species T. retifera from the Burgess Shale.
diminishes and becomes denser close to the dorsal margin O c c u r r e n c e : Jince Formation, mostly the middle
and near the presumed lateral ridge (the ridge itself is bro- part. The lowermost find (VK41) derives from the upper
ken off). It may be recalled that this specimen was figured part of the Eccaparadoxides pusillus Zone, about 3 m be-
and misinterpreted as a fish fragment by Michálek (1978) low the Dawsonia bohemica “Horizon” at Vinice near
in a popular journal, which caused some erroneous stories Jince. Most specimens come from the interval between the
about fish remains in the Jince Cambrian. Eccaparadoxides pusillus and Paradoxides gracilis Zones,
All other specimens of Tuzoia sp. are fragmentary but namely from its upper part at Vinice and Vystrkov near
show a typical reticulate sculpture consisting of polygonal Jince (the single specimen from Velká Bába near Hos-
fields of different shape and density (most obvious pentago- tomice probably came from the same level). Later speci-
nal and hexagonal polygons, maximum diameter 6–7 mm). mens occur in the Paradoxides gracilis Zone (Vinice) and
Among these specimens, L36484 (pl. IIE, coll. Dr. R. the latest fragments come from the Ellipsocephalus hoffi-
J. Prokop) shows the straight dorsal margin preserved at a bearing layers at Koníček.
length of 49 mm long. The area of denser and smaller
reticulae, delimited by stronger ridges near the dorsal mar- Comments on Pilocystites primitius Barrande, 1887
gin, is locally up to 8 mm wide, but the increase of mar- Text-fig. 5
ginal density is typical and comparable e.g. with
MP-S01748 (fig. 1) or L36481 (pl. IIH). Barrande (1887) based the genus Pilocystites on a sin-
A remarkable remain is L36483 (text-fig. 2B, coll. Dr. gle fragmentary specimen named Pilocystites primitius
M. Široký). It is a negative counterpart of the dorsal and Barr. from the shales with “Paradoxides” belonging to his
posterodorsal parts of a large carapace valve (total pre- “faune primordiale” from Ginetz (recte Jince). He regard-
served length 111 mm), which shows damaged dorsal ed the fossil as an incomplete calyx of a cystoid and in its
margin terminated by a stout, horn-like spine (length 6 description he stressed the composition of irregular polyg-
mm). Configuration of this spine differs in form and sup- onal plates without distinct pores (Barrande 1887, p. 185).
pressed sculpture from the spine interpreted in VK41 as Barrande’s interpretation was understandable, as no simi-
the anterodorsal. Consequently, the spine in L36483 is in- lar arthropod remains with marked reticulate sculpture
terpreted as the posterodorsal, very similar to the termina- were known in his time from strata of comparable age (the
tion of the fragmentary remain described by Barrande as problematic Dictyocaris Salter, 1860 from much younger,
Pilocystites primitius (see discussion below). The polygo- Downtonian, strata, was no firm base for comparison).
nal reticulate sculpture shows only a slight tendency to di- Jaekel (1899) regarded Pilocystites as a doubtful root
minish near the margins (influenced by preservation). of a crinoid, Ubaghs in Beaver et al. (Treatise, 1967: S493,
Remarks. Although assigned to Tuzoia, the available fig. 323, 2) expressed doubts about the echinoderm nature
specimens do not show the entire outline of carapace of the fossil.
valves and the number and configuration of all marginal The original specimen of P. primitius is not deposited
spines. Consequently, a full diagnosis cannot be given and in the Barrande collection in the National Museum,
open nomenclature is used. Prague, and only the plaster cast – the plastotype L9049 –
Among the species described, T. retifera Walcott, 1912, exists (Barrande noted that the specimen is contained in a
T. guntheri Robison et Richards, 1981, and T. polleni Ress- block of shales with a large “Paradoxides” he found about
er, 1930, show analogies in reticulate sculpture, presence of forty years ago, i.e. in the 1840’s). Regrettably, all at-
a marked lateral ridge with diminished and denser reticulae tempts to find the original specimen were unsuccessful.
and configuration of the anterodorsal region. Although the In our interpretation, based on the plastotype of the
diminishing of the reticulation near and on the surface of the original specimen figured by Barrande (1887) in pl. 2, fig.
lateral ridge is more marked and occupies a broader band in 26 and refigured here in text-fig. 5, this is a fragment of a
our specimens, the individual variability of sculpture and its carapace valve of a tuzoiid arthropod, most likely conspe-
changes during ontogeny has not been studied in any larger cific with Tuzoia sp. as reported here.
set of individuals of Tuzoia, and, consequently, its systemat- The plastotype proper is not of very good quality but it
ic significance cannot be precisely evaluated. In our view, the shows a straight edge 51 mm long, representing in our
increase of reticulation density, especially near the lateral opinion the incomplete dorsal margin of a carapace valve.
ridge, can increase during the ontogeny, being more marked The horn-like process on the left edge of the cast can be in-
in large individuals. This is another reason for leaving our terpreted as the posterodorsal spine, comparable with that
specimens in open nomenclature. preserved in the specimen L36483 of Tuzoia sp. The

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Ivo Chlupáč – Vratislav Kordule

sculpture is developed even in

some other Cambrian arthro-
pods, e.g. Hurdia Walcott, 1912,
from the Burgess Shale, the re-
mains of a large tuzoiid arthro-
pod described in this paper, or
the curious Lower Cambrian
Retifacies Hou-Chen et LU,
A 1989, from the Chengjiang fau-
na (Hou and Bergström 1997). A
similar sculpture also occurs in
much later arthropods, e.g. the
enigmatic, late Silurian Dicty-
B ocaris and the Lower Carbonif-
erous phyllocarid Kulmocaris
Hahn et Brauckmann, 1977,
from the Viséan of Germany
(Hahn and Brauckmann 1977).
We cannot therefore see any
sufficient proof to replace, for
nomenclature reasons, the well
established and widely used Tu-
zoia Walcott, 1912, by the older
but ambiguous Pilocystites Bar-
rande, 1887. It is recommended
to use Pilocystites only for the
reported plastotype (eventually
C for the original specimen,
should that be found in future).
O c c u r r e n c e : After Bar-
rande (1887) Ginetz, i.e. Jince,
evidently the Jince Formation
(according to the large “Para-
doxides” reported by Barrande),
Fig. 5. Plaster cast (plastotype) of Pilocystites primitius Barrande, 1887, spec. Nr. L9049, length 51 mm,
Jince. A – original drawing figured by Barrande (1887, pl. 2, fig. 26); B – recent schematic drawing ac- exact locality unknown (for dis-
cording to the plastotype; C – recent photograph of the plastotype. cussion of Barrande’s localities
see Chlupáč 1999).

Large tuzoiid arthropod (n. gen.)

oblique margin on the left side is interpreted as a frag- Pl. III, figs A–C, text-fig. 6
mentary posterodorsal margin, all other edges of the rem-
nant being secondary. The most characteristic feature – the M a t e r i a l : The single specimen L36595 from Jince.
reticulate sculpture consisting of polygonal reticulae of D e s c r i p t i o n : The specimen is represented by two
unequal size (maximum diameter 5–6 mm) and shape, in separated, strongly sculptured parts interpreted as rem-
which pentagonal and hexagonal fields are obvious – ful- nants of two very large carapace valves in parallel posi-
ly agrees with Tuzoia. Also the diminishing reticulae near tion, without preserved margins.
the inferred dorsal and posterodorsal margins are analo- Each of the valve remnants – the left and the right parts
gous as observed e.g. in our specimen VK41 or in other of the fossil – shows an analogous and symmetrical arrange-
specimens of Tuzoia sp. ment of sculptural elements. The main median area of both
Although the conspecifity with Tuzoia sp. seems evi- “valves” that occupies most of the surface is flat and marked
dent, the plastotype cannot in our opinion affect the valid- by a typical sculpture consisting of polygonal reticulae anal-
ity of the generic name Tuzoia Walcott, 1912, even though ogous to those of Tuzoia: individual polygons of quadran-
Pilocystites Barrande, 1887, has priority. The reason is gular, pentagonal and hexagonal outline form a network of
that Pilocystites in its original concept (now based exclu- reticulae 4–7 mm (exceptionally up to 8 mm) in diameter,
sively on the plastotype) shows no characteristic features delimited by narrow ridges. Flat areas with this kind of
that unambiguously demonstrate the identity of both gen- reticulation are bordered on both sides (peripheral = outer,
era: the plastotype does not show the lateral ridge and mar- median = inner) by areas with clearly different sculpture,
ginal spines diagnostic of Tuzoia. Similar reticulate consisting of blunt and thickened ridges encircling distinct-

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 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

30 mm

Fig. 6. Schematic drawing of remains of the large tuzoiid arthropod Tuzoidae n. gen.? (L36595) from Vinice near Jince, Paradoxides gracilis Zone
(coll. M. Široký).

ly concave subcircular depressions terminated by median individual valves can be calculated as c. from 100 to
pits (pl. IIIC). The density of depressions is around 6–8 in 110 mm, length as 150 to 170 mm.
10 mm. The areas with large polygons and those with pit- 2. The second possibility differs in interpretation of the
ting are connected and distinguished by smaller reticulae main flat reticulate areas which would represent only
but the differences are very marked. the dorsolateral sectors of the valves, while the areas
Although the fossil is flattened, traces of the primary with dense sculpture on the outer margin of the fossil
morphology are partly preserved: the two median bands indicate merely bands along the lateral longitudinal
with dense sculpture are slightly raised, the main reticulate ridge (analogous to Tuzoia in which the ridge area is
areas are flat and the outer, marginal bands with dense distinguished by a denser reticulation). In this case, the
sculpture are moderately bent downwards on both sides of calculated dimensions would be substantially larger
the specimen. Remains of paradoxidid trilobites (librige- than in the first interpretation (the minimal dorsoven-
nae, thoracic segments, parts of cephala) are imprinted in- tral width would be estimated as c. 170–180 mm,
to the evidently thin reticulate areas of the shield. length as 220 to 250 mm).
M e a s u r e m e n t s : The total width of the fossil is As for sculpture, namely the broad bands with dense
227 mm; maximum length, measured parallel to the bands sculptural elements, they differ from all known represen-
of different sculpture, is 123 mm. The preserved transverse tatives of Tuzoia and point to a separate generic status.
width of bands with dense sculpture on the outer sides of However, the lack of primary margins prevents from mak-
both presumed valves is 26 mm and the inner bands with ing an unambiguous and sufficient diagnosis. Therefore,
dense sculpture attain a maximum width of 17 mm. the fossil, representing one of the most remarkable find-
R e m a r k s . As the original margins are not preserved, ings in the Middle Cambrian of Jince, is here left under in-
the interpretation of the fossil is ambiguous. Even in inter- formal designation.
pretation as two valves of a bivalve arthropod exist two O c c u r r e n c e : The single find, made by Dr. M.
possibilities: Široký at Vinice near Jince (lower part of the hillside), de-
1. The areas with dense sculpture situated in median part rives from the upper part of the Paradoxides gracilis Zone
of the fossil indicate areas in proximity and along the of the Jince Formation.
dorsal margins of valves, the large flat areas with large
reticulae represent most of the surface of both carapace Genus Proboscicaris Rolfe, 1962
valves, and the densely sculptured outer bands corres-
pond to marginal parts of valves near their ventral mar- Ty p e s p e c i e s : Proboscicaris agnosta Rolfe, 1962.
gin. In this case, the presumed dorsoventral width of D i a g n o s i s : Carapace valves subovate to subrectan-

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Ivo Chlupáč – Vratislav Kordule

10 mm

10 mm

Fig. 7. Proboscicaris hospes sp. n. – the holotype VK45a in-

C terpreted as the right carapace valve. A – the entire remain; B
– enlarged part of the left part; C – schematic drawing. Vinice
near Jince.

gular, with carapace horn anterior-produced into spatulate projection at about the mid-length of the carapace valve,
beak (Rolfe 1969). other irregularities could be due to pressure deformation.
R e m a r k s . With one exception, only isolated cara- The valve is markedly posterior-tapered. It reaches the
pace valves are preserved, reaching in the type species the maximum dorsoventral width of 23 mm slightly anterior
length of more than 100 mm and in the other species – P. of the mid-length of the valve and attains a width of only
ingens Rolfe, 1962 – up to 156 mm. The reticulate sculp- 9–10 mm near the posterior margin. The posterior margin
ture, resembling Tuzoia, is less marked and only locally exhibits two rounded projections the posteroventral of
preserved on the carapace valves of the type species as re- which is slightly longer and more markedly curved than
ported by Rolfe (1962). The genus has been so far known the posterodorsal one. The arcuate notch between both
exclusively from the Middle Cambrian of North America projections is moderately anterior-curved. The convexity
(Burgess Shale and Utah). Its occurrence in Bohemia is of the valve was modified by pressure but according to
the first of this kind in the Old World. mode of preservation it was moderate.
R e m a r k s . The assignment to Proboscicaris is based on
the presence of the anterodorsal “beak” and general shape of
Proboscicaris hospes sp. n. the carapace valve without any marked spines. The diagnos-
Text-fig. 7 A–C tic features of our species are the shape of the prolonged
valve, the broad anterior “beak” and two rounded posterior
H o l o t y p e : right carapace valve VK45a, b moderately projections separated by an arcuate notch. The last feature re-
deformed by pressure and preserved with its counterpart. sembles Pseudoarctolepis Brooks et Caster, 1956, in which
D e r i v a t i o n o f n a m e : from Lat. hospes = guest two posterior projections are also developed but the notch be-
(subst.). tween them is more marked and the posterodorsal projection
Ty p e l o c a l i t y : Vinice near Jince. is sharp and spin-formed (comp. Brooks and Caster 1956,
Ty p e s t r a t u m : Jince Formation (middle part), in- Robison and Richards 1981). The most characteristic feature
terval between the Eccaparadoxides pusillus and Paradox- of Pseudoarctolepis – the anteroventral-situated and pos-
ides gracilis Zones, Middle Cambrian. teroventral-directed hollow spine – is lacking in P. hospes.
M a t e r i a l : The holotype. Secondary pressure deformations are expressed in the
D e s c r i p t i o n . Carapace valve prolonged, semiellip- holotype as irregular wrinkles parallel with the ventral
tical in outline, tapering posteriorly. Dorsal margin in most margin and as irregular depressions and elevations on the
part of its course straight, without spines, anterodorsal part valve surface. No patterns of reticulate sculpture are de-
of the valve protruded into broad, anterior-rounded veloped and only very faint and small ridges near the pos-
“beak”. Anteroventral margin gently curved, ventral mar- terior margin (especially on the posteroventral projection)
gin gently curved with some irregularities: a slight ventral may be regarded as sculptural elements.

174
 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

A B

Fig. 8. Helmetia? fastigata sp. n. – the holotype VK42a (A) and its schematic drawing (B) supplemented by features preserved on the counterpart.
Rejkovice, Ellipsocephalus hoffi – Rejkocephalus Zone. Total width 37 mm.

O c c u r r e n c e : The only known specimen comes from terpreted as prolonged, bearing characteristic marginal
Vinice near Jince, middle part of the Jince Formation, the up- spines. The posterior, median spine is long and strong, mod-
permost part of the interval between the Eccaparadoxides erately tapering, pointed. The paired posterolateral margin-
pusillus and Paradoxides gracilis Zones (coll. V. Kordule). al spines are markedly smaller and shorter. The posterior
pair is approached to the median spine, the next, more ante-
Class Artiopoda Hou et Bergström, 1997 rior-placed pair is poorly preserved as a still smaller projec-
Subclass Conciliterga Hou et Bergström, 1997 tion. On one side of the shield, another longer and stouter,
Order Helmetiida Novozhilov, 1969 more anterior-placed spine is indicated and posterior-direct-
Family Helmetiidae Simonetta et Delle Cave, 1975 ed. However, this part of the shield is separated from the
Genus Helmetia Walcott, 1918 main part by a secondary fold, which makes the connection
unclear. The whole tail shield is flattened and crushed,
Ty p e s p e c i e s : Helmetia expansa Walcott, 1918, showing secondary folds and fissures, which also form its
Burgess Shale, Middle Cambrian, British Columbia. evidently secondary anterior delimitation.
D i a g n o s i s : see Stormer in Harrington et al. (1959). M e a s u r e m e n t s o f t h e h o l o t y p e : Sagittal
length of the preserved part of the shield: 60.5 mm, width
Helmetia? fastigata sp. n. c. 37 mm (dimensions affected by crushing), length of the
Text-fig. 8A, B median spine 13.5 mm, length of the posterolateral spines
2.5–3 mm.
H o l o t y p e : Crushed tail shield VK42a, b (with coun- R e m a r k s . Although the preservation of the holotype
terpart) figured in text-fig. 8A, B. is not sufficient for a full diagnosis, the configuration and
D e r i v a t i o n o f n a m e : From Latin fastigatus = position of marginal spines on the tail shield differs from
pointed (adj.). all previously described representatives of Helmetiidae
Ty p e l o c a l i t y : Rejkovice near Jince, railway cutting. and allows the establishment of a new species.
Ty p e s t r a t u m : Upper part of the Jince Formation, The secondary deformations strongly affect particular-
Zone with Ellipsocephalus hoffi, level with Lindinella ko- ly the anterior part of the holotype. They are manifested as
rdulei. asymmetrical curved folds and small ridges of two kinds
M a t e r i a l : The holotype and two fragmentary speci- and dimensions: the larger curved traces of more or less
mens possibly conspecific. longitudinal (exsagittal) direction strongly suggest the ich-
D e s c r i p t i o n . Only the tail shield is known. It is in- nofossil trails Helminthopsis, the markedly smaller ridges

175
Ivo Chlupáč – Vratislav Kordule

A B

C D

Fig. 9. Forfexicaris? sp., isolated carapace

valves. A, B – the largest specimen L36596 in
two different lightenings, Vinice near Jince
(length 23 mm), Paradoxides gracilis Zone; C,
D – the smallest specimen SZ168 with coun-
terpart (diameter 15.8 mm), Skryje Shale, Luh
near Skryje, Eccaparadoxides pusillus Zone.
In A, C, D the outline marked by a drawn line.

and irregularly curved traces are comparable with Gordia. H.? fastigata is the first representative of Helmetiidae
All these structures be of evidently organic origin represent outside North America and southwest China.
traces of worm-like animals, possibly scavengers concen- O c c u r r e n c e . Rejkovice near Jince, olive-green silty
trated on the surface of or near the shield at the time of its shales of the upper part of the Jince Formation, Zone with
burial (the traces are developed exclusively on the rem- Ellipsocephalus hoffi and Rejkocephalus, level with
nants of the shield and do not continue into the surround- Lindinella kordulei. In association with Ellipsocephalus
ing rock). Other deformations are caused by a clear hoffi (Schlotheim), Rejkocephalus rotundatus (Barrande),
imprint of a thorax of Ellipsocephalus in the lateral part of hyolithids, Lindinella kordulei Mergl et Šlehoferová, rare
the shield and by secondary crushing which caused a Konicekion tix Šnajdr and ichnofossils.
strong breakage of the anterior part of the fossil.
The family Helmetiidae includes in its present concept Arthropoda incertae sedis
three genera: Helmetia Walcott, 1918, Kuamaia Hou, 1987, Genus Forfexicaris Hou, 1999
and Rhombicalvaria Hou, 1987 (Hou and Bergström 1997).
As indicated by both authors, the separate position and T y p e s p e c i e s : Forfexicaris valida Hou, 1999,
generic status of Kuamaia and Rhombicalvaria are question- Lower Cambrian, Yunnan, China.
able, and this also concerns their relations to Helmetia,
which has nomenclature priority. Owing to this, our speci- Forfexicaris? sp.
men is tentatively ranged with Helmetia. Text-fig. 9A-D
The type species of Helmetia – H. expansa Walcott,
1918, as figured by Walcott (1931, cf. also figure in Con- M a t e r i a l : 8 subcircular plates interpreted as isolated
way Morris et al. 1982, Briggs et al. 1994), shows a clear- carapace valves flattened in silty shale.
ly greater distance between the posterior median spine and D e s c r i p t i o n : Valves subcircular, gently asymmetri-
the more anterior-placed lateral ones. The same difference cal in outline. Carapace very thin. Dorsal margin (at the
concerns both the so far described species of the Lower supposed hinge-line) slightly curved up to rectilinear for
Cambrian Kuamaia – K. lata Hou, 1987, and K. muricata some distance, thus differing from the broadly arcuate and
Hou et Bergström, 1997, which also show a shorter and markedly curved ventral margin. Anterodorsal corner
broader tail shield and a smaller median spine. The type rounded, somewhat narrower than the broadly arcuate pos-
and the only known species of Rhombicalvaria – R. acan- terodorsal one. Maximum length of valves in about 1/4 of
tha Hou, 1987 (comp. Hou 1987) exhibits – in contrast to the maximum width of valves measured perpendicularly
Kuamaia – considerably longer and slender marginal from the presumed dorsal margin. Surface of valves
spines, the distance between the median and the last pos- smooth, only slight indications of lines parallel with the
terolateral spine being substantially longer. margins (except the dorsal one).

176
 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

M e a s u r e m e n t s : The best-preserved specimen Biostratigraphy and accompanying fauna

(L36596) shows the dorsoventral width of the valve as
21.5 mm, its length being 23 mm. The maximum diame- The Jince Formation, introduced in the literature by
ters of other specimens range between 30 and 17 mm. Lipold and Krejčí (1860) for Barrande’s étage C of his
The smallest specimen SZ168 is 15.8 mm in diameter at “Silurian System” (1852) and later moderately redefined
the most. (Havlíček in Chlupáč, ed. 1968), corresponds too much of
R e m a r k s . As the extremities and the thorax remain the Middle Cambrian.
unknown, the interpretation and assignment are doubtful. According to recent state of knowledge, it can be sub-
The rounded outline and smooth surface of carapace divided into several biostratigraphic zones and subzones
valves are shared with several Cambrian arthropod taxa of (“horizons”); see Fatka and Kordule (1992), Kordule
diverse relationships. Most morphologically similar are (1996), Fatka (2000). The distribution and character of
carapace valves of the genera Forfexicaris Hou, 1999, Oc- faunas clearly reflect the cycle of marine ingression, start-
cacaris Hou, 1999, and Clypecaris Hou, 1999, all from the ing with low-diversity brachiopod and trilobite-dominated
Lower Cambrian Chengjiang fauna (Hou 1999). Among assemblages and continuing over two maxima of abun-
them, Forfexicaris seems to show greatest analogy in dance distinguished by the presence of higher diversity
available features, and, therefore, the remnants are tenta- faunas (even with miomerid = agnostid trilobites) and de-
tively assigned to it with reserve. clining into the regressive phase in the uppermost part of
Though the flattening cannot be excluded in all speci- the Jince Formation with dominating lingulids.
mens so far collected, the absence of any marked coarse The finds of described non-trilobite arthropods belong
and irregular wrinkles on the evidently very thin cuticle mostly to the upper parts of the Jince Formation, i.e. with-
points to the valves being originally flat in life. in the sequence with common trilobite and other faunas.
The described fossils were previously figured from The occurrence of Tuzoia sp., however, does not fully
the Jince Formation of the Jince area by Mergl and Šle- correspond with the abundance and diversity maxima of
hoferová (1991, pl. 11, figs 4, 5, MM284, YA1305), who other fossils. The oldest find derives from the upper part of
classified them with reserve as brachial valves of a pos- the Eccaparadoxides pusillus Zone. Most remains, how-
sible but doubtful inarticulate brachiopod Acrothele gi- ever, come from the interval between the E. pusillus and
gantea Želízko of uncertain systematic position (even Paradoxides gracilis Zones, particularly from its upper
with possibility of exclusion from Brachiopoda). part (designated by Kordule 1996 as the Subzone with
Mikuláš (2000) in his revision of Bohemian Middle Acanthocystites and Akadocrinus), where also Probosci-
Cambrian ichnofossils figured in pl. 13, figs 3, 4 and pl. caris hospes was found, but the total faunal diversity is
34, figs 6, 7 also subcircular very thin valves, which he low. The common accompanying trilobite is here Hydro-
also regarded with some reserve as shells of inarticulate cephalus minor (Boeck), frequently found even as com-
brachiopods with superficial ichnofossils designated as plete exoskeletons. It is joined by Rejkocephalus
fodinichnion igen. et isp. indet. rotundatus (Barrande), Peronopsis integra (Beyrich),
These subcircular thin shells are best interpreted as Acanthocystites briareus Barrande, Akadocrinus jani
arthropod carapace valves designated here as Forfexi- Prokop, Stromatocystites pentangularis Pompeckj, rare
caris? sp. Almost all show also analogous trace fossils on Konicekion radion Šnajdr, and very few other fossils. Ac-
their surface – possibly traces of scavengers (the rela- cording to Mikuláš (2000), Planolites and Teichichnus
tionships between arthropod remnants and associated dominate among ichnofossils.
ichnofossils will be discussed elsewhere). However, the Rare remains of Tuzoia sp., carapace valves of For-
rocks of the Jince Formation also contain thicker-shelled fexicaris? sp. and the unique find of the large tuzoiid
subcircular fossils with indicated apices whose brachio- arthropod (Tuzoidae n. gen.?), as described here, were
pod (inarticulate) nature is very probable – this concerns found within the upper part of the Jince Formation with
e.g. some specimens designated as Acrothele gigantea the second maximum fossil abundance, namely in the
Želízko deposited in collections of the National Museum, Paradoxides gracilis Zone. They occur together with nu-
Prague. merous trilobites with strongly dominant Paradoxides
The described thin-shelled subcircular fossils prob- gracilis (Boeck), frequent Conocoryphe sulzeri
ably do not occur so sporadically as the available mater- (Schlotheim), Ptychoparia striata (Emmrich), Hydro-
ial suggests. Their unattractive nature probably meant cephalus minor (Boeck), Peronopsis integra (Barrande),
that they were in most cases overlooked by fossil collec- echinoderms Lichenoides priscus Barrande, Stromato-
tors, even at classic localities. cystites sp., Etoctenocystis bohemica Fatka et Kordule,
O c c u r r e n c e : Jince Formation, Jince area: the slope rare ostracods Konicekion radion Šnajdr, hyolithids,
of Vinice, Paradoxides gracilis Zone, Rejkovice (railway Tommotium sp. etc.
cutting) and Koníček near Jince, the Ellipsocephalus hoffi In the present zonal subdivision of the Middle Cambri-
– Rejkocephalus Zone, Skryje-Týřovice area: Luh and an according to agnostid trilobites, the above mentioned
Plazy near Skryje, Eccaparadoxides pusillus Zone. beds with arthropod remains can be roughly correlated

177
Ivo Chlupáč – Vratislav Kordule

with the Ptychagnostus atavus and particularly with the P. ter life zone, dominated by polymerid trilobites.
punctuosus Zones (sensu Robison 1984), i.e. the higher The ichnofossils marked by common epi- and
part of the major Paradoxides paradoxissimus trilobite intrastratal traces indicate the widely distributed shallow-
biozone of the Baltic scheme. water environment of the Cruziana and/or Skolithos-
The latest described arthropods come from the Ellip- Cruziana Ichnofacies (comp. Mikuláš 2000).
socephalus hoffi – Rejkocephalus Zone, especially from The co-occurrence with complete exoskeletons of small
the locality at Rejkovice, where Helmetia? fastigata sp. n., and large trilobites and non- or incompletely disarticulated
Tuzoia sp., Forfexicaris? sp. and some other problematical echinoderms and the preservation of even thin shields ex-
remains were found in association with Ellipsocephalus clude any longer transport of the studied remains. In our
hoffi (Schlotheim), Rejkocephalus rotundatus (Barrande), view, the tuzoiid, helmetiid and probably also proboscicarid
frequent small hyolithids (often with opercula), bra- arthropods could be regarded as nektobenthic animals
chiopods Lindinella kordulei Mergl et Šlehoferová, rare whose habit might be compared with some trilobites: they
Konicekion tix Šnajdr, and few other fossils. Ichnofossils might be scavengers or filter-feeders rather slowly moving
(Mikuláš 2000) belong to Skolithos, Planolites, Teichich- above the sea floor (cf. consideration of Helmetia by Briggs
nus and Daedalus. The E. hoffi – Rejkocephalus Zone in Conway-Morris et al. 1982).
might correspond to the lower part of the Paradoxides A different habit can be presumed in Forfexicaris? sp.
forchhammeri Zone of the Baltic zonal scheme (the corre- with very thin cuticle. It might represent a pelagic element
lation, however, is still problematic). less dependent on bottom conditions and shallow-water
Rare remains tentatively reported as Forfexicaris? sp. from realms (this can be supported by its occurrence in the
the Skryje-Týřovice Cambrian in the NW flank of the Barran- Skryje Shale in association with more common agnostids).
dian, belong to the Eccaparadoxides pusillus Zone of the
Skryje Shale, corresponding roughly to the lower (but not the
lowest!) part of the Jince Formation of the Jince area. Conclusion
To summarize, the representatives of Tuzoia and allied
tuzoid arthropods occupy together with thin-shelled re- Rare remains of large non-trilobite arthropods in the
mains designated as Forfexicaris? sp. a rather long strati- Middle Cambrian of Bohemia are confined to the later
graphic interval within the Middle Cambrian, particularly phase of the marine transgression cycle.
in its higher part correlatable with the Paradoxides para- The described arthropods show clear resemblances to
doxissimus Zone. Rare finds of Helmetiids and other ex- other, particularly North American, Middle Cambrian fau-
ceptional arthropods occurring in later strata with the nas (especially the Burgess Shale of British Columbia and
index trilobite Ellipsocephalus hoffi can be correlated with coeval strata of Utah). Their rare occurrence and incomplete
the lower part of the Paradoxides forchhammeri Zone of preservation, however, prevent adequately based palaeogeo-
the Baltic scheme. graphical and biostratigraphical conclusions from being
drawn. The relationships indicate open migration possibili-
ties and faunal exchange over vast areas, probably even be-
Palaeoecological notes tween Laurentia and the presumed shelves of Gondwana, to
which also the Barrandian area belonged.
All the arthropods described here come from the Mid- All described arthropods lived in marine environment
dle Cambrian siliciclastic rocks, namely from siltstones with marked terrigenous influx. However, their remains
and silty-clayey shales. They occur in sequences marked are contained in finer-grained siltstone and combined silt-
by alternating influx of silt-clay-sand (greywacke) materi- stone-claystone layers pointing to temporary intervals of
al in irregular intervals of mm to tens of cm thickness relative stable and quieter sedimentation. These conditions
(lithologic characteristics of the Jince Formation: Kukal could be generally more suitable for preservation of thin
1971, Havlíček and Kukal in Chlupáč et al. 1998). In- carapaces of described arthropods. The occurrence of Hel-
terbeds of dark grey or grey-green greywackes are locally metia?, Tuzoia, and Forfexicaris? in siltstones of the El-
frequent but they are persistent on individual outcrops. De- lipsocephalus hoffi Zone with less diversified benthos may
scribed remains of arthropods derive exclusively from indicate a tolerance to conditions of decreased salinity.
shale or siltstone beds, they were newer found in A nektobenthic habit and a rather shallow-water envi-
greywacke layers. This suggests that the presence of such ronment are presumed for the discussed arthropods (ex-
arthropods corresponds to intervals of more stable condi- cept Forfexicaris? sp. as a possible pelagic element).
tions reflected in the fine-grained sedimentation. Biostratigraphic conclusions are premature, but the
Frequent remains of marine benthic animals, namely available finds come mostly from the upper part of the
large polymerid trilobites and less common but typical Jince Formation, where some decrease of diversity in ben-
echinoderms (carpoids, eocrinoids, edrioasteroids) and hy- thic fauna is already observed. The correlation with the
olithids, accompany the described arthropods. Agnostids middle and upper parts of the Paradoxides paradoxissimus
are also present, though they are not dominant. According Zone (most finds) and possibly with the lowest part of the
to the scheme established for the Jince Middle Cambrian Paradoxides forchhammeri Zone (the youngest finds) of
by Fatka (2000) our arthropods belong in the shallow-wa- the Baltic biostratigraphic scheme is most probable.

178
 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

A c k n o w l e d g e m e n t s . The authors are indebted to RNDr. M. Chengjiang Fauna, Yunnan, China. J. Paleont., 73, 1, 102–116. Tulsa.
Široký, Ing. L. Mandík and Mr. M. Szabad for offering the specimens Hou X. G., Bergström J. (1997): Arthropods of the Lower Cambrian
from their private collections for study. Dr. R. J. Prokop (National Chengjiang fauna, southwest China. Fossils Strata, 45, 116 pp. Oslo.
Museum, Prague) gave valuable information on echinoderms and evalu- Jaekel O. (1899): Stammesgeschichte der Pelmatozoen. 1. Thecoidea und
ation of Pilocystites primitius Barrande and supported, together with Dr. Carpoidea. 442 pp. Springer Berlin.
V. Turek, this study in the National Museum, Prague. Our particular Kordule V. (1996): Význam některých méně známých fosiliferních
thanks go to W. D. J. Rolfe (Edinburgh) for critical reading the manu- lokalit pro stratigrafii jineckého souvrství v českém středním kam-
script. Doc. Dr. O. Fatka (Institute of Geology and Palaeontology, briu. Věst. Čes. geol. Úst., 71, 1, 37–49.
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Handling editor: Petr Štorch

Explanation of plates posteroventral parts); E – fragmentary dorsal part of the right carapace valve
with posterior edge, L36484; F – fragmentary specimen SZp5964, a tooth-
like projection is probably a hyolithid shell attached to the Tuzoia fragment;
Plate I ⇒ G – fragment of a large specimen showing the typical reticulate sculpture,
Tuzoia sp., Jince Formation (upper part), Vinice near Jince (A-C), L36482; H – two fragmentary and approached carapace valves showing
Rejkovice (D). configuration of the dorsal margins and increasing density of reticulate
A, B, C – incomplete right carapace valve VK40 with partly preserved sculpture near the broken lateral ridges, L36481.
lateral ridge, posteroventral part and doublure; (B – enlarged figure with-
out the anterior part, C – enlarged anterior part with partly exposed dou- Plate III ⇒⇒⇒
blure); D – fragment of doublure SZ169. Large tuzoiid arthropod (n. gen.), Jince Formation, Vinice near Jince,
Paradoxides gracilis Zone. L36595 (coll. M. Široký), maximum diame-
Plate II ⇒⇒ ter 227 mm.
Tuzoia sp., Jince Formation near Jince (except F – Velká Bába near Hos- A, B – overall views in different light, with different sculpture and im-
tomice). printed remains of paradoxidid trilobites; C – strongly enlarged part of
A, B, C, D – incomplete right carapace valve VK41, Vinice; A – enlarged the left medium sector showing the marked difference between the “tu-
posterodorsal part, B – enlarged anterior part, C – enlarged dorsal part, D – zoiid” polygonal sculpture and the thicker and pitted parts.
the overall view showing the covered median part and broken antero- and All photos R. J. Duda

179
Ivo Chlupáč – Vratislav Kordule

10 mm

10 mm

10 mm

10 mm

Plate I

180
 Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic)

10 mm
A

10 mm

10 mm
E

10 mm

10 mm
F

10 mm

10 mm

10 mm

Plate II

181
Ivo Chlupáč – Vratislav Kordule

10 mm

10 mm

Plate III

182