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Two trilobite faunas of late Ordovician (Sandbian and Katian) age are described from the siliciclastic Karagach Formation, western Tarbagatai Range, eastern Kazakhstan. They comprise 15 families and 24 genera and include the new taxa Agerina acutilimbata sp. nov., Birmanites akchiensis sp. nov., Dulanaspis karagachensis sp. nov. and Kimakaspis kovalevskyi gen. et sp. nov. Most of the Karagach Formation yields graptolites characteristic of the Diplograptus foliaceus [multidens] Biozone, which are associated with the older trilobite fauna; the uppermost part, which is the source of the younger trilobite fauna, contains Orthograptus quadrimucronatus and Dicranograptus hians which suggest a younger age, equating with the lowermost Ensigraptus caudatus Biozone, and the base of the Katian Stage. Most of the trilobite genera in both faunas have a wide geographical distribution in the late Ordovician, although Dulanaspis and Sinocybele are characteristic of low latitude eastern peri-Gondwanan faunas.
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Late Ordovician trilobites from the Karagach
Formation of the western Tarbagatai Range,
Mansoureh Ghobadi Pour
, Lucy M. E. McCobb
, Robert M. Owens
Leonid E. Popov
Department of Geology, Faculty of Sciences, Golestan University, Gorgan, Iran
Department of Geology, National Museum of Wales, Cathays Park, Cardi, CF10 3NP Wales, UK
ABSTRACT: Two trilobite faunas of late Ordovician (Sandbian and Katian) age are described
from the siliciclastic Karagach Formation, western Tarbagatai Range, eastern Kazakhstan. They
comprise 15 families and 24 genera and include the new taxa Agerina acutilimbata sp. nov.,
Birmanites akchiensis sp. nov., Dulanaspis karagachensis sp. nov. and Kimakaspis kovalevskyi gen. et
sp. nov. Most of the Karagach Formation yields graptolites characteristic of the Diplograptus
foliaceus [multidens] Biozone, which are associated with the older trilobite fauna; the uppermost part,
which is the source of the younger trilobite fauna, contains Orthograptus quadrimucronatus and
Dicranograptus hians which suggest a younger age, equating with the lowermost Ensigraptus caudatus
Biozone, and the base of the Katian Stage. Most of the trilobite genera in both faunas have a wide
geographical distribution in the late Ordovician, although Dulanaspis and Sinocybele are character-
istic of low latitude eastern peri-Gondwanan faunas.
KEY WORDS: Katian, nileid biofacies, peri-Gondwana, Sandbian, taxonomy
The Tarbagatai Range lies on the border between eastern
Kazakhstan and Xinjiang Province, north-western China, and
is an area from which little has been published on Ordovician
trilobites. The rst were discovered by Vasilievsky in 1912 and
were subsequently described as Remopleurides sibiricus by
Vasilievsky in Weber (1948); otherwise, only three Upper
Ordovician taxa, Dindymene brevicaudata Kolobova, 1972
from the Karagach Formation (upper Sandbian), Dionide
tarbagataica Koroleva, 1979 (precise horizon and locality
unknown) and Pliomerina rigida Kolobova, 1972 from the
upper Kulunbulak Formation (upper Katian), have been
described, whilst others remain known only from lists of
preliminary identications (e.g. Nikitin 1972), which are not
always reliable. Fortey & Cocks (2003) recently reported a
deep-water Caradoc trilobite fauna from the vicinity of
Karagach village, which was examined by Fortey amongst
extensive collections made by the late M. K. Apollonov. Part
of this collection, comprising six samples derived from three
levels in the siliciclastic Karagach Formation (Sandbian to
lower Katian; Caradoc), is the subject of this present paper. In
spite of a relatively small number of specimens, the fauna is
taxonomically diverse, comprising 24 genera. The Karagach
Formation has yielded graptolite faunas indicative of the
foliaceus and clingani biozones (Tsai 1976), discussed below.
1. Stratigraphy and age
The section of the Karagach Formation sampled by the late
Mikhail Apollonov is situated about 35 km east of the town of
Ayaguz, on the east side of the Ayaguz River, about 7 km
north of Akchii village (Figs 1, 2). According to Nikitin (1972,
p. 206, text-g. 69), the Karagach Formation overlies a discon-
tinuous sequence of ne siliciclastic rocks belonging to the
Middle Ordovician Naiman Formation, and underlies the
Upper Ordovician Zhartas Formation, which comprises
andesitic volcanic rocks and tus with units of volcaniclastic
sandstones and argillites. Tsai (1976) estimated the thickness of
the Karagach Formation to be over 1200 m, and subdivided it
into four informal units (see Figs 2–3) which in ascending
order are: (1) limestones about 10 m thick; (2) dark grey
siltstones with a few beds of ne-grained sandstones about
200 m thick; (3) alternating sandstones and siltstones about
130 m thick; (4) siltstones about 675 m thick with a few beds of
sandstones and calcareous siltstones in the uppermost 30 m of
the unit.
Graptolites occur sporadically in the sequence, and were
described by Tsai (1976). They provide relatively good age
constraints for the trilobite faunas, and have been recovered
from all three fossiliferous horizons in the Karagach Forma-
tion. Those from the two lower ones (samples 201, 202,
205) are considered to be indicative of the Diplograptus
foliaceus [multidens] Biozone. Sample 201 (unit 2) has yielded
Dicranograptus ramosus (Hall, 1847), D. nicholsoni diapason
Gurley, 1896, Pseudoclimacograptus scharenbergi (Lapworth,
1876), Hustedograptus teretiusculus (Hisinger, 1840) and Diplo-
graptus diminutus (Ruedemann, 1947); samples 202 and 205
(unit 3) have yielded Pseudoclimacograptus scharenbergi and
Dicranograptus sp. From the top of unit 4 of the Karagach
Formation, samples 203, 203a, 203b have yielded Orthograptus
quadrimucronatus (Hall, 1865) and Dicranograptus hians (Hall,
1905). These two species occur together at the base of the
Ensigraptus caudatus Biozone at the Black Knob Ridge sec-
tion, Oklahoma, the stratotype section for the base of the
Katian Stage (Goldman et al. 2007).
Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 101, 161–187, 2011 (for 2010)
2011 The Royal Society of Edinburgh. doi:10.1017/S1755691010010078
The two trilobite assemblages in the Karagach Formation
occur at three levels; both are considered to be characteristic of
outer shelf environments. The lower one, from units 2 (sample
201) and 3 (samples 202 and 205) includes the pelagic cyclo-
pygids Cyclopyge, Microparia (Heterocyclopyge) and Symphys-
ops, which co-occur with Dindymene, Dionide?, Dulanaspis,
Raphioampyx?, Shumardia and a trinucleid (Kimakaspis gen.
nov.). This association indicates a somewhat deeper environ-
ment from that of unit 4. However, neither association
includes three-segmented raphiophorids, which are other-
wise widespread in deep water environments across eastern
Kazakhstanian terranes.
The upper part of unit 4 (samples 203, 203a, 203b) contains
a medium diversity trilobite assemblage which includes the
genera Agerina, Amphilichas, Ceraurinella?, Dulanaspis, Illae-
nus, Nileus, Remopleurides, Stenopareia, Sphaerocoryphe and
Sinocybele, which may represent a nileid biofacies. This fauna
occurs together with a brachiopod assemblage dominated by
Christiania, in association with Acculina, Glyptambonites and
Leptaena (Ygdrasilomena) among others.
2. Significance of the fauna
In their brief review of the trilobite assemblage from the
Karagach Formation, Fortey & Cocks (2003) pointed out that
it includes geographically widespread taxa (e.g. Dionide,
?Raphioampyx, Shumardia, Eodindymene, Microparia and
Cyclopyge), and this is true mainly for the lower assemblage
from the Diplograptus foliaceus [multidens] Biozone. The
assemblage from the uppermost Karagach Formation con-
tains, among other taxa, Dulanaspis and Sinocybele which,
together with the occurrence of Pliomerina reported by
Kolobova (1972) from the somewhat younger Kulunbulak
Formation, is clear evidence of anity with the mid to
late Ordovician ‘eastern’ peri-Gondwanan Eokosovopeltis
Pliomerina fauna, distributed through central Asia, the
Taurides in Turkey, Tarim, North and South China, Sibumasu
and eastern Australia (Zhou & Zhou 2006, p. 387; Edgecombe
& Webby 2006, p. 414). The latter authors (2006, p. 414)
noted that Sinocybele should be regarded as an additional
biogeographically sensitive indicator of this fauna.
Most of the genera recorded from the Karagach Formation
also occur in other parts of Kazakhstan; in particular, they are
known from the Anderken and Dulankara formations of
Chu-Ili Range (Chugaeva 1958) and from the approximately
contemporaneous Besharyk (=Ichkebash) Formation of the
Dzhebagly Mountains (Kolova 1936). However, biofacies dif-
ferentiation of late Ordovician trilobite faunas of Kazakhstan
remains poorly understood, which makes further comparison
dicult. There is little doubt, however, that all these faunas
belong to separate early Palaeozoic terranes (Fortey & Cocks
2003, Popov et al. 2009) and typical Kazakhstanian genera
such as Birmanites and Dulanaspis are represented in the
assemblage from the Karagach Formation by local endemic
A small trilobite fauna described by Kolobova (1983)
from the Anderken Regional Stage of Koryk and Malkeldy
in the Chingiz Range contains one species (Encrinuroides
septemcostatus Kolobova) which is probably common to the
Karagach Formation, but the species of Nileus from that
assemblage is dierent, and there are no other genera in
Figure 1 Part of eastern Kazakhstan, showing position of the Karagach section in the western Tarbagatai
3. Systematic palaeontology
Most of the specimens are deposited in the Department of
Geology, National Museum of Wales, Cardi (NMW, Acces-
sion number 2005.32G). Some illustrated and referred type
specimens are deposited in the F. N. Chernyshev Central
Geological Scientic Research and Exploration Museum
(CNIGR), St Petersburg. The terminology and systematic
classication follows Whittington & Kelly (in Kaesler 1997).
Measurements are in millimetres. Abbreviations: L=length
of exoskeleton, Cl, Cw=cranidial length and width; CPl,
CPw=cephalic length and width; Gl, Gw=glabellar length and
maximum width; GPw=posterior glabellar width; ORl=sagit-
tal length of occipital ring; Pl, Pw=pygidial length and width;
PAl, PAw=pygidial axis length and width.
Family Shumardiidae Lake, 1907
Genus Shumardia Billings, 1862
Type species. Shumardia granulosa Billings, 1862, Middle
Ordovician, Darriwilian Stage, Shumardia Limestone, Que´bec,
Remarks. Following Waisfeld et al. (2001), Shumardia and
Conophrys are considered to be separate genera, and Shumar-
doella Pek & Vaneˇk, 1989 is regarded as the junior subjective
synonym of Shumardia.
Shumardia a. extensa Weir, 1959
(Fig. 4a–h)
a. 1997 Shumardia (Shumardoella) extensa Weir; Hammann
& Leone, p. 44, pl. 4, gs 1–8.
Material. NMW 2005.32G.259, exoskeleton, internal
mould (L, 5·0, Cl, 1·8, Pl, 1·9, PAl, 1·25); NMW 2005.32G.261,
NMW 2005.32G.262 (Cl, 1·6, Cw, 2·6, Gw, 1·5, GPw, 1·0,
ORl, 0·4), cranidia, internal moulds; NMW 2005.32G.260,
pygidium with thorax, internal mould; NMW 2005.32G.264,
pygidium, internal mould (Pl, 1·6, Pw, 2·4, PAl, 1·1, PAw, 1·0);
all from sample 201. NMW 2005.32G.147, cephalon with
thorax, internal mould (Cl, 1·9, Cw, 2·9, Gw, 1·7); NMW
2005.32G.148, pygidium, internal mould; both from sample
Description. Cranidium semioval, about 65% as long as
wide. Glabella about 60% as wide as cranidium, with maxi-
mum width at about one quarter glabellar length from the
anterior margin, comprising not more than 150% of posterior
glabellar width. Axial furrows deep, shallowing near anterior
margin of glabella. Preglabellar furrow weakly dened. Ante-
rolateral glabellar lobes separated from rest of glabella by very
shallow, anteromedially directed glabellar furrows fading
anteriorly, their posterior margins situated at about mid-length
Figure 2 Schematic geological map of the area east of Ayaguz River showing distribution of the Middle to
Upper Ordovician rocks and the positions of fossil localities (modied from Nikitin, 1972).
of the cranidium and their width (trans.) almost equal to
occipital ring length (sag.). S1 short (trans.), curved, dening
small bacculae. Occipital ring occupying about 20–25% of
cranidial length, with a straight, transverse posterior margin,
separated from the glabella by a narrow, straight occipital
furrow. Preglabellar eld very short (sag.). Fixigenae sub-
triangular with posterior border slightly widening (exsag.)
abaxially and a straight posterior margin slightly inclined
posteriorly abaxially. Posterior border furrow transverse,
narrow. Librigenae and hypostoma unknown.
Thorax of six segments with wide axis, about half as wide
(tr.) as thoracic segments and lacking a macropleural segment.
Pygidium parabolic, about 65% as long as wide, with axis
occupying up to 45% of maximum pygidial width and up to
70% of pygidial length. Axis subconical, with four axial rings
separated by deep, transverse ring furrows, deepening abaxi-
ally, and a terminal piece. Axial furrows deep, converging
posteriorly. Pleural elds gently convex with four pairs of
pleural ribs separated by weakly dened pleural furrows,
fading abaxially. Doublure wide (sag.), occupying almost one
third of pygidial length.
Remarks. The morphology of Shumardia a. extensa,
including the presence of distinct bacculae, the absence of a
macropleuric thoracic segment, an elongate pygidium with
an axis not extending to the margin, and a curved convex
posterior pygidial margin, are diagnostic for Shumardia (for
details see Waisfeld et al. 2001). In their cranidial and pygidial
morphology, they are almost identical to those described from
Sardinia by Hammann & Leone (1997) from the Cyclopygid
Bed of the Punta S’Argiola Member of the Domusnovas
Formation (Katian) as Shumardia (Shumardoella) extensa
Weir, 1959, and they may well be conspecic. However, the
attribution of the specimens from Kazakhstan and Sardinia to
this species is questionable, because the original description of
S. extensa is based on a single cranidium.
Shumardia a. extensa diers from the type species, S.
granulosa Billings, 1862 revised by Whittington (1965), in
having a more elongate cranidium, a larger glabella with
evenly rounded anterior margin, and smaller anterolateral
lobes not strongly projecting laterally, a preglabellar eld
lacking a median depression, and a parabolic pygidial outline
with only four axial rings. Shumardia a. extensa is similar to
Figure 3 Stratigraphical section through the Upper Ordovician deposits on the eastern side of Ayaguz River,
showing the horizons of the fossil localities (numbers as in Fig. 2) and stratigraphical ranges of trilobite and
graptolite species.
S. lacrima Koroleva, 1964 from the late Sandbian/early Katian
Mailisor Formation of northern Kazakhstan, but has a more
elongate cranidium, a shorter (sag.) occipital ring, a distinctly
wider pygidium with a shorter pygidial axis bearing only three
axial rings (instead of six), and a smaller number of pygidial
pleural ribs (three pairs instead of six). This species diers
from S. analoga Koroleva, 1964, also from the Mailisor
Formation, in having an evenly rounded anterior cranidial
margin, a wider glabella with anterolateral glabellar lobes
which terminate posteriorly at cranidial mid-length, a very
short preglabellar eld, xigenae with a subtriangular (not
subrectangular) outline, and a shorter pygidial axis with only
three axial rings.
Shumardia a. extensa diers from S. tarimuensis Zhang,
1981 from the Saergan Formation (upper Darriwilian to
lower Sandbian) of Tarim and the Chedao Formation (lower
Sandbian) of north-western China (Zhou & Dean 1986) in
having a more elongate cranidium with a very narrow pre-
glabellar eld and evenly rounded anterior glabellar furrow, a
signicantly wider glabella with smaller anterolateral glabellar
lobes, xigenae with a subtriangular outline, and a pygidium
with a suboval, not subtriangular outline.
Family Leiostegiidae Bradley, 1925
Genus Agerina Tjernvik, 1956
Type species. Agerina erratica Tjernvik, 1956; Lower
Ordovician, Floian Stage, Billingen Regional Stage, Latorp
Formation, Na¨rke, Sweden.
Agerina acutilimbata sp. nov.
(Fig. 5a–c)
Derivation of name. From the pointed anterior margin of
Holotype. NMW 2005.32G.135, articulated exoskeleton,
external mould (L, 6·4, W, 3·9); sample 203a.
Paratypes. NMW 2005.32G.137, partly disarticulated
exoskeleton, external mould (CPl, 2·8, CPw, 3·5, Gl, 2·1, Gw,
1·4); NMW 2005.32G.138, cranidium, internal mould (Cl, 2·4,
Figure 4 Shumardia a. extensa Weir, 1959: (a–b) NMW 2005.32G.259, incomplete exoskeleton, internal
mould, dorsal and lateral views, 15, 16, sample 201; (c) NMW 2005.32G.260, pygidium with attached
thoracic segments, internal mould, 9, sample 201; (d) NMW 2005.32G.147, cranidium with attached thorax,
internal mould, 13, sample 205; (e–f) NMW 2005.32G.261, cranidium, internal mould, dorsal and lateral
views, 22, sample 201; (g) NMW 2005.32G.148, pygidium, internal mould, 16, sample 205; (h) NMW
2005.32G.262, cephalon with two thoracic segments, internal mould, 15, sample 201.
Cw, 2·8, Gl, 2·1, Gw, 1·4); NMW 2005.32G.139, pygidium,
external mould (Pl, 1·4, Pw, 2·0, PAl, 1·5, PAw, 0·5); sample
Diagnosis. Cephalon semioval, about 80% as long as wide;
elongate, parallel axial furrows with evenly rounded anterior
margin conned to the anterior border furrow; three pairs of
very weakly dened glabellar furrows; occipital ring transverse
with a small median tubercle; convex cephalic border of equal
width with short median spine; medially (exsag.) situated
palpebral lobes about 25% of cephalic length; eight thoracic
segments, and pygidium with conical axis about 35% of
maximum pygidial width, four axial rings and very short
terminal piece; pleural elds with three to four pairs of ribs;
border narrow, fading posteriorly abaxially.
Description. Exoskeleton elongate suboval, about 165% as
long as wide with maximum width at the posterior margin of
the cephalon, and lateral margins converge gently posteriorly.
Cephalon semioval, about 80% as long as wide. Cranidium
about 85% as long as wide with a strongly convex, elongate
subrectangular glabella. Glabellar furrows short (tr.), very
weakly dened; S1 oblique, and dene very small subtriangular
L1; S2 situated at about glabellar mid-length; S3 opposite
anterior termination of palpebral lobes. Frontal margin of
glabella gently rounded, almost conned to anterior border
furrow. Occipital ring strongly convex, transverse, with a small
median tubercle; occipital furrow narrow, deep and transverse.
Cephalic border convex, of constant width, bearing a short,
subtriangular median spine, dened by a deep and narrow
border furrow. Postocular xigena short (exsag.), narrows
abaxially with bluntly rounded lateral termination. Palpebral
lobes semioval, maximum length (exsag.) about 25% that of
cephalon (sag.), with posterior termination at about 30%
cephalic length from the posterior margin. Preocular facial
sutures slightly divergent anteriorly. Librigenae wide (tr.) and
convex, with a short (tr.) posterior border. Genal spines
present but poorly preserved.
Thorax with eight segments. Axis about 35% of thoracic
width, pleurae long (exsag.) with a wide and deep pleural
furrow, tapering and terminating at inner margin of facet.
Outer portion of pleura bends posteroventrally, and terminates
with a rounded distal extremity; prominent facet present on the
anteriolateral part of pleura, about 50% of its width (tr.).
Pygidium small, transverse, semioval, about 60% as long as
wide with a conical axis occupying about 35% (tr.) of the
anterior margin and about 75% (sag.) of pygidial length. Axis
with four rings and a small terminal piece, dened by deep
axial furrows, slightly converging posteriorly. Posteriormost
axial ring very short (sag.) and separated from a short terminal
piece of about equal length by a very faint axial furrow. Pleural
eld with three to four pairs of pleural ribs, slightly curved
backwards and bisected by faint interpleural furrows. Pleural
furrows deep and narrow. Border narrow, fading antero-
medially, separated from the pleural eld by a shallow, weakly
dened border furrow.
Remarks. In their revision of Agerina, Adrain & Fortey
(1997) suggested that this generic name should be applied to
taxa which have a parallel-sided glabella with eaced glabellar
furrows, and a pygidial border ornamented by subparallel,
raised terrace ridges, similar to the type species and to Agerina
pamphylica Dean, 1973 from the Middle Ordovician Sobova
Formation of the Taurus Mountains, southern Turkey. In its
cephalic morphology, A. acutilimbata sp. nov. is similar to this
group of species, but the short median spine on the anterior
margin of the anterior cranidial border is unique for Agerina.
A. acutilimbata diers from A. pamphylica in having very
poorly dened glabellar furrows and a granulose, not pitted
ornament on the cranidium and librigenae.
Family Asaphidae Burmeister, 1843
Subfamily Asaphinae Burmeister, 1843
Genus Birmanites Sheng, 1934
Type species. Ogygites birmanicus Reed, 1915; Upper
Ordovician, Sandbian Stage, Hwe Mawng Beds, northern
Shan States, Burma.
Species assigned. In addition to the type species, the
genus includes: (1) Birmanites hupeiensis Yi, 1957, Middle
Ordovician Miapo and Shihtzupu formations, south China; (2)
Niobe lata Angelin, 1851, Katian Stage, Jonstorp Formation,
Va¨stergo¨tland, Sweden (Kielan 1960) and Sqort Tepe Forma-
tion, south-east Turkey (Dean & Zhou 1988); (3) Ogygites
pamiricus Balashova, 1966, upper Darriwilian Stage,
Kozynsdy Formation, eastern Pamir, Tajikistan; (4) Opsima-
saphus asiaticus Petrunina in Yaskovich & Repina, 1975,
Upper Ordovician olistolith, Madygen, Turkestan Range,
Kyrgyzstan; (5) Ogygites kolovae Chugaeva, 1958, lower
Katian Stage, Dulankara Formation, Chu-Ili Range,
Kazakhstan; (6) Ogygites almatiensis Chugaeva, 1958,
Sandbian Stage, Anderken Formation, Chu-Ili Range,
Kazakhstan; (7) Ogygites sp., Abdullaev in Abdullaev &
Khaletskaya, 1970, Upper Ordovician, Beshtor Formation,
Pskem Range, Uzbekistan.
Birmanites pamiricus Balashova, 1966, from the Tremado-
cian Stage of eastern Pamir, Tajikistan is based on a single
poorly preserved external mould of a cephalon, and its precise
generic attribution cannot be made (Balashova 1966, p. 208,
pl. 3, g. 9). Therefore, this taxon is considered here as a nomen
dubium. However, the binomen Birmanites pamiricus
Balashova, 1966, is a senior homonym of Ogygites pamiricus
(see Balashova 1966, p. 215) because the latter is a valid species
of Birmanites.
Birmanites akchiensis sp. nov.
(Fig. 7a–e)
Derivation of name. From Akchii village south of the type
Holotype. NMW 2005.32G.181, external mould of cranid-
ium (Cl, 26·6, Cw, <30·3, Gl, 14·6, Gw, 16·6); sample 203b.
Paratypes. NMW 2005.32G.182, librigena, external
mould, sample 203b; NMW 2005.32G.183, hypostome, inter-
nal mould, sample 203a; NMW 2005.32G.184, pygidium,
external mould (Pl, 39·0, PAl, 26·7, PAw, 26·70), sample 203b;
NMW 2005.32G.185, pygidium, internal mould (Pl, 40·3,
PAl, 26·5, PAw, 10·8), sample 203b; NMW 2005.32G.200,
pygidium, internal mould, sample 203a.
Diagnosis. Cranidium of about equal length and width,
with broadly and evenly rounded anterior margin; preglabellar
eld smooth, with very weakly expressed longitudinal median
ridge; pygidium about 70% as long as wide, with axis occupy-
ing 20% of pygidial width and 75% of maximum pygidial
length, and terminating with a short postaxial ridge; axial
furrows are shallow and wide, almost straight; 12 well-dened
axial rings; pleural eld attened, with six pairs of pleural ribs
which do not reach border.
Description. Cranidium as long as wide with maximum
width at posterior margin, only slightly exceeding maximum
width of preglabellar eld. Anterior margin evenly curved.
Surface covered by a Bertillon pattern of terrace ridges.
Glabella gently convex, about 60% cranidial length (sag.) and
35% its width (tr.), with maximum width at occipital ring.
Axial furrows shallow and narrow, shallowing in front of
palpebral lobes. S1 transverse, deep and wide, situated oppo-
site mid-length of palpebral lobe, distally directed backwards,
but not reaching occipital furrow, and partially dening sub-
triangular L1. S2 shallow and wide, situated opposite anterior
end of palpebral lobe. Preoccipital tubercle present between
rst lateral glabellar lobes (L1). Occipital furrow narrow,
deepens abaxially. Occipital ring slightly wider than preoccipi-
tal glabella, and about 15% glabellar length (sag.). Palpebral
lobes semicircular, very large, about 20% cranidial length
(exsag.) and situated towards posterior end of cranidium.
Postocular xigena wide (tr.) and very short (exsag.), with a
shallow and narrow posterior furrow and relatively broad
(exsag.) posterior border, widening abaxially. Preocular xi-
genae very long (sag.) and wide (tr.), about 35% of cranidial
length and about 90% of maximum cranidial width (tr.).
Palpebral area of xigena reaches same height as glabella;
palpebral furrow shallow and wide, strongly curved (exsag.),
separating a prominent palpebral rim from the rest of the
palpebral area. Preocular facial sutures at rst only slightly
divergent, then strongly diverging before nally curving
Figure 5 (a–c) Agerina acutilimbata sp. nov.: (a, c) NMW 2005.32G.135, holotype, articulated exoskeleton, latex
cast of external mould, dorsal and oblique views, 19, 15, sample 203a; (b) NMW 2005.32G.137, paratype,
partly disarticulated exoskeleton, latex cast of external mould, 15, sample 203. (d) Dindymene brevicaudata
Kolobova: NMW 2005.32G.177, articulated exoskeleton, internal mould, dorsal view, 9, sample 201.
anteromedially. Postocular facial sutures strongly divergent to
almost transverse.
Librigenae wide with broadly convex lateral margin and a
broad, at subtriangular genal spine directed backwards. Eye
socle low, ridge-like, separated from rest of librigena by a very
shallow furrow. Surface of librigenae covered with ne con-
centric terrace ridges. Doublure very wide, with subparallel
terrace ridges.
Hypostome elongate, about 80% as wide as long, with a
gently convex anterior margin. Anterior wings very small,
curved dorsally. Anterior lobe of middle body subquadrate,
slightly convex. Well-dened maculae at beginning of middle
furrows, at posterolateral angles of the middle body. Middle
furrows deep and wide, merge posteriorly to create transverse
furrow. Posterior lobe of the middle body transverse, gently
convex with an eaced posterior border furrow. Posterior
border deeply cut by a large U-shaped notch; posterior fork
broad-based, pointed posteriorly. Posterolateral regions elon-
gate subtriangular, with lateral margins almost posteriorly
directed. Lateral borders attened, widening posteriorly.
Border furrows deep and wide. Surface covered by ne
transverse terrace ridges. Thorax unknown.
Pygidium semielliptical, about 70% as long (sag.) as wide
(tr.). Axis narrow, conical, with 12 axial rings and a very small
terminal piece. Axis about 20% of maximum pygidial width
and about 75% of pygidial length. Axial furrows shallow and
wide, almost straight, converging posteriorly. A short post-
axial ridge is recognisable. Pleural eld attened with six pairs
of at pleural ribs, widening abaxially and shallowing and
dying out towards border. Anteriormost three pairs curved
gently backwards; three posterior pairs almost straight and run
posterolaterally. Pleural furrows shallow and wide. Doublure
very wide, extending to axial termination posteriorly and very
close to axial furrows laterally. Surface of pygidium and
doublure is covered by coarse terrace ridges, which are
subparallel laterally and convex posteriorly.
Remarks. This species is most similar to B. kolovae
(Chugaeva, 1958, p. 39, pl. 3, gs 9–16; text-g. 6), but is easily
distinguished by having an evenly rounded rather than pointed
anterior cranidial margin, and a less transverse pygidium with
a longer axis bearing 12 axial rings. Birmanites akchiensis
diers from B. almatyensis (Chugaeva, 1958, p. 42, pl. 4,
gs 1–6; text-g. 8) in possessing a very weak median ridge
bisecting the preglabellar eld, in having a rounded anterior
cranidial margin and a less transverse pygidium bearing a short
postaxial ridge. It diers from B. hupeiensis Yi, 1957 in the
absence of a prominent medial longitudinal ridge on the pre-
glabellar eld, in having a longer glabella, a pygidium with
almost straight axial furrows and 12 well dened axial rings.
Incomplete preservation of the cranidium in B. asiaticus
(Petrunina in Yaskovich & Repina, 1975) makes it dicult to
compare it with B. akchiensis in cephalic morphology, but the
new species clearly diers from the former in having a pygid-
ium with a narrow axis ending with a short post-axial ridge
and bearing 12 axial rings, and in possessing a attened pleural
eld with only six (instead of eight) pairs of pleural ribs. The
pygidium of B. akchiensis shows similarities in the number of
axial rings and pleural ribs to the specimens from Pamir
described by Balashova (1966) as Ogygites pamiricus. How-
ever, it diers from the latter in having a more elongate
pygidium and pleural ribs that terminate some distance from
the border. More detailed comparison with O. pamiricus is
impossible due to signicant deformation of the specimens,
poor illustrations and schematic description. B. pamiricus
described by Balashova in the same paper does not appear to
be conspecic with Ogygites pamiricus (see above).
Family Cyclopygidae Raymond, 1925
Subfamily Cyclopyginae Raymond, 1925
Genus Cyclopyge Hawle & Corda, 1847
Type species. Egle rediviva Barrande, 1846; Upper
Ordovician, Sandbian Stage, Beroun Series, Vinice Formation,
Beroun, Czech Republic.
Cyclopyge cf. tereki Koroleva, 1967
(Fig. 7j–k, m)
?1982 Cyclopyge quadrangularis Kielan; Koroleva, p. 117,
pl. 24, gs 1–5 [non Kielan 1960, p. 83, pl. 9, gs 1–9]
Material. NMW 2005.32G.186, cephalon with attached
thorax, internal mould (CPl, 6·5, CPw, 3·7); NMW
2005.32G.203–206, cranidia, internal moulds; NMW
2005.32G.187, pygidium with attached thorax, internal mould
(Pl, 1·9, Pw, 5·3, PAl, 0·8, PAw, 5·6); NMW 2005.32G.188,
pygidium with attached thorax, external mould (Pl, 1·8, Pw,
4·1, PAl, 1·0, PAw, 1·1); NMW 2005.32G.207–208, pygidia
with attached thorax, external moulds; all sample 205. NMW
2005.32G.201, 202, pygidia with attached thoracic segments,
internal moulds; NMW 2005.32G.209, axial shield, internal
mould; all sample 201.
Description. Cranidium with subcircular outline, glabella
95% as long (sag.) as wide (trans). Shallow S1 inclined gently
backwards, and lies about one third of the way forwards from
posterior margin of glabella. Thorax with six segments, axis
tapering gently backwards. Pygidium transverse, 40–45% as
long (sag.) as wide (trans.), although these values are approxi-
mate due to some distortion. Axis with blunt termination, with
one ill-dened ring. Weak rst pleural furrow, otherwise
pleural areas smooth. Ill-dened, broad, weakly convex
border. No postaxial ridge.
Remarks. This species is similar to Cyclopyge tereki from
the Sandbian Stage of north-east Kazakhstan (Koroleva 1967,
p. 89, pl. 10, gs 15, 16; 1982, p. 115, pl. 23, gs 8–11) in having
one ill-dened ring on the pygidial axis, but diers in being
more transverse in outline, having a less well-dened pygidial
border, and lacking a postaxial ridge. Cyclopyge umbonata
bohemica Marek (1961, p. 22, pl. 1, gs 6–9) from the
Darriwilian Dobrotiva´ Formation, Prague district, Czech
Republic, also has only one, very ill-dened pygidial axial ring,
and has a shallow border furrow, but has a proportionately
longer (sag.) postaxial area. The specimens described here
resemble those from the Tarbagatai Mountains described by
Koroleva (1982, p. 117, pl. 24, gs 1–5) as Cyclopyge quadran-
gularis Kielan. The gures are dark and dicult to interpret,
but the pygidium in her g. 1 has a similar outline to that of
the present specimens, the axis lacks well-dened rings, and a
trace of a weak pygidial border can be seen on the right hand
side. These Tarbagatai Mountains specimens are possibly
conspecic with C cf. tereki. They are not conspecic with C.
quardangularis (Kielan 1960, p. 83, pl. 9, gs 1–9) from the
Staurocephalus clavifrons Zone (Katian Stage) of the Holy
Cross Mountains, Poland, which diers in that the glabella is
more elongated, with S1 closer together and closer to the
posterior (Kielan 1960, pl. 9, gs 6, 7) and the pygidium has a
more tapered axis with one ring clearly dened (Kielan 1960,
pl. 9, g. 7).
Genus Microparia Hawle & Corda, 1847
Type species. Microparia speciosa Hawle & Corda,
1847 from Upper Ordovician, Katian Stage, Kra´lu˚v Dvu˚r
Formation, Kra´lu˚v Dvu˚r, Czech Republic.
Subgenus Heterocyclopyge Marek, 1961
Type species. Cyclopyge pachycephala Hawle & Corda,
1847 from Upper Ordovician, Sandbian Stage, Vinice
Formation, Trubı´n near Beroun, Czech Republic.
Microparia (Heterocyclopyge) sp.
(Fig. 7l)
Material. NMW 2005.32G.189, pygidium, internal mould
(Pl, 4·6, Pw, 6·5, PAl, 2·6, PAw, 2·6), sample 205.
Remarks. The distinctive triangular pygidial axis with
three well-dened rings and a triangular terminal piece nds
parallels in such M. (Heterocyclopyge) species as M. (H.)
abunda Zhou, McNamara, Yuan & Zhang, 1994 (gs 5C, D,
F–I, 6A–H, 7A, C, E, H, M), from the early Sandbian
Charchaq Group, Tarim and M. (H.) shelvensis Whittard, 1961
(pl. 24, gs 3–4; Owens 2002, pl. 3, g. 4) from the Llanvirn
Series (Darriwilian) of the Welsh Borderland and central
Wales. It diers from the rst in the subquadrate pygidial
outline (see, e.g. Zhou et al. 1994, gs 5F, I; 6A, C), and from
the second in having deeper, broader pleural furrows. A very
similar pygidial axis also occurs in Cyclopyge kossleri Kloucˇek,
1916 from the Darriwilian S{a´rka Formation of the Prague
district (Marek 1961, pl. 1, gs 15–17) and the Llanvirn
(Darriwilian) Llanfallteg Formation of Pembrokeshire, Wales
(Fortey & Owens 1987, g. 37 a–b), but the glabella of this
species has deep S1, typical of Cyclopyge. The triangular
pygidial axis in C. kossleri is not shared by other Cyclopyge
species, so it should perhaps be transferred to Microparia
(Heterocyclopyge) – the lack of deep S1 in species currently
attributed to the latter may simply be due to eacement. The
presence in C. kossleri of both deep S1 and a triangular
pygidial axis suggests derivation of M. (Heterocycolpyge) from
Subfamily Pricyclopyginae Fortey & Owens, 1987
Genus Symphysops Raymond 1925
Type species. Aeglina armata Barrande, 1872, from Upper
Ordovician, Katian Stage, Kra´lu˚v Dvu˚r Formation, Kra´lu˚v
Dvu˚r, Czech Republic.
Symphysops cf. djebaglensis (Kolova, 1936)
(Fig. 7i)
cf. 1936 Cyclopyge djebaglensis Kolova, p. 37, pl. 2, gs 1–3.
Material. NMW 2005.32G.171, pygidium, incomplete
internal mould; NMW 2005.32G.190, pygidium, external
mould (Pl, 9·8, Pw, 18·5, PAl, 5·8, PAw, 5·9); sample 205.
Description. Pygidium almost semicircular in outline,
about half as long as wide. Axis convex, subconical, about 60%
as long as the pygidium, about as long as wide, with a
well-dened rst axial ring, very poorly-dened second and
third rings, and a terminal piece. Pleural elds gently convex,
eaced with two very weakly dened pairs of pleural ribs.
Border wide, gently convex; border furrow wide and shallow.
Remarks. These pygidia show close anity to that from
the Besharyk Formation of the Dzhebagly Mountains, south-
ern Kazakhstan described and gured by Kolova (1936, p. 37,
pl. 11, g. 3) as Cyclopyge djebaglensis. They dier from this
specimen in apparently having much more weakly dened
pleural ribs, although preservation is poor. Pygidia gured as
Cyclopyge (?) by Weber (1932, p. 9, pl. 4, gs 52–54), also
from the Dzhebagly Mountains, may be conspecic, but the
material is too incomplete and the gures too small for detailed
Family Nileidae Angelin, 1854
Genus Nileus Dalman, 1827
Type species. Asaphus (Nileus) armadillo Dalman, 1827,
Middle Ordovician, Darriwilian Stage, Holen Limestone
Formation, Asaphus expansus Biozone, O}stergo¨tland, Sweden.
Nileus sp.
(Fig. 11f, h)
Material. NMW 2005.32G.193, cephalon with attached
thoracic segments, external mould (CEl, 5·7, CEw, 7·0, Cw
6·6), sample 203; NMW 2005.32G.194, pygidium, internal
mould (Pl, 5·9, Pw, 10·5, PAw, 5·0), sample 203a.
Description. Cephalon semioval, about 80% as long as
wide, strongly convex (sag., tr.). Cranidium slightly elongate,
subrectangular, strongly eaced with glabella not clearly
separated from xigenae. Palpebral lobes large, almost semi-
circular, occupying almost 60% of cephalic length, with
anterior terminations situated just 15% from the anterior
cranidial margin. Preocular facial suture very short, weakly
divergent, curved adaxially at anterior cephalic margin. Post-
ocular facial sutures divergent at an angle of 50( to axis.
Postocular xigenae small, subtriangular, poorly dened.
Librigenae with genal angles narrow (tr.) and steeply inclined
laterally. Visual surface of eyes vertical and smooth.
Thorax with a wide axis, about 60% of thoracic width, and
very weakly dened axial furrows.
Pygidium transversely semioval, about 55% as long as wide,
with a short axis, which is weakly dened anteriorly and
occupies almost half of anterior pygidial width. Pygidial
border wide, gently concave in cross section. Pygidial doublure
wide, concave and ornamented by strong terrace lines, with
U-shaped anterior margin mainly conned to axis.
Remarks. This species has a strongly transverse, eaced
pygidium with a broad, concave border which resembles
that of Nileus transversis Koroleva, 1982, which is also
known from the Sandbian, Anderken Regional Stage of the
south-western Chingiz Range in Kazakhstan. However, the
cranidial characters of the latter are unknown, so further
comparison is not possible. Another species from the Upper
Ordovician of the south-western Chingiz Range is N. kasach-
stanicus Koroleva, 1982. Nileus sp. can be readily distinguished
from it in having almost completely eaced cranidial axial
furrows, large palpebral lobes that occupy more than half
of the cephalic length, and facial sutures with very short
preocular branches.
Family Remopleurididae Hawle & Corda, 1847
Genus Remopleurides Portlock, 1843
Type species. Remopleurides colbii Portlock, 1843, Upper
Ordovician Katian Stage, (Ashgill Series), Killey Bridge
Formation, Desertcreat, Co. Tyrone, Northern Ireland (see
Whittington, 1950, for revision of species).
Remopleurides cf. sibiricus Vasilievsky in Weber, 1948
(Fig. 7f–h)
?1948 Remopleurides sibiricus Vasilievsky (ms.) in Weber; p. 20,
pl. 3, g. 7.
1982 Remopleurides cf. sibiricus Weber, Koroleva, p. 72, pl. 10,
g. 4.
Material. NMW 2005.32G.179, external mould of cranid-
ium; NMW 2005.32G.136, 178, 198–199, pygidia with
attached thoracic segments, internal and external moulds;
samples 203, 203a.
Remarks. The cranidium has a gently convex, strongly
eaced glabella with a relatively short, declined anterior
tongue and ornamented by very ne, transverse terrace ridges;
a short (sag.) transverse occipital ring tapering distally; and a
palpebral rim maintaining almost constant width along its
entire length (exsag.). Pygidia attached to up to nine thoracic
segments, mostly exfoliated, and the position of a thoracic
median spine cannot be determined. The cranidial features
suggest attribution to Remopleurides sibiricus. The description
of this species given in Weber (1948) is based on a manuscript
by Vasilievsky, who collected the specimens from an un-
identied, presumably Upper Ordovician unit in the Ayaguz
river basin in 1912. According to Weber (1948, p. 100), this
material had been lost by 1940, and a holotype was not
assigned in the original description of the species. The locality
data given by Weber (1948) are very general, but specimens
described in the present paper may be topotypic, or may have
originated in close proximity to the type locality. The speci-
mens described by Koroleva (1982) are probably conspecic
with those from the Karagach Formation, and are therefore
included in the synonymy list. Remopleurides sibiricus requires
revision, but the available material is inadequate to discuss its
Family Illaenidae Hawle and Corda, 1847
Genus Illaenus Dalman, 1827
Type species. Entomostracites crassicauda Wahlenberg,
1818; Middle Ordovician, Darriwilian Stage, Dalby Limestone
Formation (basal part – see Jaanusson 1963, pp. 35–36),
Fja¨cka, Dalarna, Sweden.
Figure 6 (a–b, d) Illaenus sp.: (a) NMW 2005.32G.142, pygidium, internal mould, dorsal view, 1·8, sample
203a; (b) NMW 2005.32G.141, pygidium, internal mould, dorsal view, 1·8, sample 203a; (d) NMW
2005.32G.140, cranidium, external mould, dorsal view, 5·5, sample 203. (c) Dionide sp. NMW 2005.32G.265,
disarranged articulated specimen, latex cast of external mould, 5, sample 201. (e) Raphioampyx?sp.:
NMW 2005.32G.162, disarticulated exoskeleton, dorsal view, 6, sample 201. (f) Aethedionide sp., NMW
2005.32G.191, pygidium, internal mould, dorsal view, 7·5, sample 205. (g–h) Stenopareia sp.: (g) NMW
2005.32G.145, cranidium, internal mould, dorsal view, 1; (h) NMW 2005.32G.144, pygidium, internal mould,
dorsal view, 3·5; both sample 203a. (i) Amphilichas a. nasutus Webby, 1974: NMW 2005.32G.192, incomplete
cranidium, internal mould, dorsal view, 3, sample 203b.
Illaenus sp.
(Fig. 6a, b, d)
Material. NMW 2005.32G.140, external mould of cranid-
ium, sample 203; NMW 2005.32G.141–143, pygidia, internal
moulds, sample 203a.
Remarks. A number of species attributed to Illaenus and
often described in open nomenclature can be found in publi-
cations by Weber (1948), Lisogor (1965) and Koroleva (1965).
Due to poor illustrations and only brief descriptions, their
precise taxonomic discrimination is dicult, and they require
revision. The unnamed species from the Karagach Formation
is distinctive among them in having a strongly eaced cranid-
ium with posteriorly placed palpebral lobes, and a strongly
convex pygidium with a relatively long, conical axis occupying
about one third of its length. Among the late Ordovician
species of the genus from Kazakhstan, the cranidium of
Illaenus sp. is most similar to the specimen described by Weber
(1948) as Illaenus sp. 17 from the Upper? Ordovician of
Karatas, east of Astana (53(30#N, 72(30#E), in possessing a
subquadrate outline, a strongly curved lateral prole and
indistinguishable axial furrows. The pygidia of Illaenus sp.
show similarity in outline and in their relatively long subconi-
cal axis to I. renhuaensis Yin in Yin & Lee, 1978 (revised by
Zhou et al. 1984) from the Shihtzupu Formation of Guizhou
Province, South China.
Genus Stenopareia Holm in Schmidt, 1886
Type species. Illaenus linnarssoni Holm, 1882, Upper
Ordovician, Katian Stage, Boda Limestone Formation,
Dalarna, Sweden.
Stenopareia sp.
(Fig. 6g, h)
Material. NMW 2005.32G.145, cranidium, internal mould
(Pl, 47·6, Pw, 56); NMW 2005.32G.146, cranidium, internal
mould (Cl, 10·6, Cw, 13·3); NMW 2005.32G.144, pygidium,
internal mould (Pl, 20·5, Pw, 21·1); all sample 203a.
Description. Cranidium moderately convex (sag., tr.),
about 80% as long as wide with evenly rounded anterior
margin. Glabella gently convex (tr.) with shallow, subparallel
axial furrows that become obsolete at cranidial mid-length.
Palpebral lobe long (sag.), gently convex abaxially with
anterior margin situated slightly posterior to cranidial mid-
length. Prominent lunette present adjacent to axial furrow.
Posterior section of xigena very small, subtriangular. Other
cranidial features are not suciently preserved for description.
Pygidium moderately convex, semielliptical, about 85–95%
as long as wide. Axis exceeding half of anterior pygidial width,
well dened by shallow, broad axial furrows in the anterior
third of pygidial length in the smaller specimen, but becoming
almost eaced in the larger specimen (Fig. 6h). Pygidial
doublure dorsally concave, narrow, maintaining equal width
posteriorly and laterally, ornamented by ne, subparallel
terrace ridges.
Remarks. In its pygidial morphology, this unnamed
species resembles, and may be conspecic with, the specimens
described by Apollonov (1974) as Stenopareia sp. 2 from the
Zharky Beds (Katian) of north-eastern central Kazakhstan. In
common with them, it has a relatively elongate pygidium with
length exceeding 80% of pygidial width, and a narrow pygidial
doublure maintaining constant width along the entire pygidial
margin. It diers from S. pulchrum Apollonov, 1974, which
occurs at the same locality, in possessing relatively large,
posteriorly placed palpebral lobes, and a pygidial axis that is
well dened by axial furrows on the anterior third of the
pygidial length in juvenile specimens. As Apollonov (1974)
described neither the pygidial doublure morphology, nor illus-
trated the pygidium of S. pulchrum, further comparison with
this species is not possible.
Stenopareia sp. diers from the specimens described as S.
a. bowmanni (Salter, 1848) by Zhou & Dean (1986) from the
Chedao Formation (Sandbian) of eastern Gansu Province of
northwest China, in having signicantly larger palpebral lobes,
and a more elongated pygidium with narrow doublure that
maintains constant width posteriorly and laterally.
Family Styginidae Vogdes, 1890
Genus Dulanaspis Chugaeva, 1956
Type species. Dulanaspis levis Chugaeva, 1956; Upper
Ordovician, Katian Stage, Dulankara Formation, Chu-Ili
Range, Kazakhstan.
Dulanaspis karagachensis sp. nov.
(Fig. 8a–d)
Derivation of name. From the Karagach Formation.
Holotype. NMW 2005.32G.150, incomplete cephalon, in-
ternal mould (Cl, 10·3, Cw, 11·0, Gl, 7·1, Gw, 6·4); sample 205.
Paratypes. NMW 2005.32G.153, pygidium with attached
thoracic segments, internal mould, sample 203a; NMW
2005.32G.152, pygidium, internal mould (Pl, 9·3, Pw, 14·5,
PAl, 2·4; PAw, 3·2), sample 203a; NMW 2005.32G.151, pygid-
ium, internal mould, sample 203b; NMW 2005.32G.154,
pygidium, internal mould, sample 203a; NMW 2005.32G.155,
pygidium, internal mould, sample 203b.
Diagnosis. Glabella slightly elongate, with maximum
width at anterior margin; cranidial axial furrows strongly
divergent anteriorly; very narrow cephalic border; large semi-
circular eye; very weakly dened posterior cephalic border;
pygidium transverse, semioval, with axis completely lacking
segmentation; long, well-dened postaxial ridge.
Description. Cephalon strongly transverse, semioval.
Cranidium about equal length and width, with maximum
width at mid-length of palpebral lobe. Glabella pestle-shaped,
gently convex, about 110% as long (sag.) as wide (tr.) with
maximum width at evenly curved anterior margin, extends as
far as weak anterior border furrow. A very small, weakly
dened L1 is present at the base of the preoccipital glabella.
Axial furrows deep and narrow, almost parallel as far as point
opposite mid-length (exsag.) of palpebral lobe, then diverging
anteriorly. Occipital ring narrow, transverse, separated from
the glabella by a shallow, weakly dened occipital furrow.
Cephalic border very narrow (exsag.) with a weakly dened,
shallow border furrow. Palpebral lobes semicircular, posteri-
orly placed with anterior margins situated slightly posterior of
cephalic mid-length. Posterior border almost eaced. Post-
ocular xigenae narrow (tr.) and short (exsag.). Preocular
facial sutures diverge immediately in front of palpebral lobes,
and run more or less parallel to axial furrow. Postocular facial
sutures very short, divergent posteriorly. Librigenae gently
convex, subtriangular, with posterior margins strongly curved
posteriorly abaxially and with acute genal angles.
Thorax with narrow, convex axis dened by deep and
narrow axial furrows. Parts of eight segments are preserved
(Fig. 8d); pleurae curve gently backwards, and terminate in
blunt points.
Pygidium moderately and evenly convex, semioval, about
65% as long as wide. Axis subconical, short, moderately
convex, eaced, about 30% of pygidial length (sag.) and about
25–30% of maximum pygidial width. Axial furrows wide and
shallow, strongly converging posteriorly. A ne, narrow post-
axial ridge is present. Pleural elds evenly convex, smooth.
Border wide, gently concave, separated from the pleural elds
by a very shallow, weakly dened border furrow. Doublure
gently concave dorsally, with ne, parallel terrace ridges. A
pair of large, oblique, elongate suboval muscle scars is present,
situated posterolaterally of posterior termination of pygidial
Remarks. This species diers from others (e.g. D. anderk-
ensis Chugaeva, 1958; D. costata Chugaeva, 1958 and D. levis
Chugaeva, 1956) from Kazakhstan in having a narrow (tr.)
postocular xigena, a very narrow cephalic border, and a more
transverse pygidium with a semioval, not parabolic outline. It
also diers from D. costata in the completely eaced pygidial
axis and pleural elds without any traces of segmentation, and
unlike D. anderkensis and D. levis, it has a long, well dened
postaxial ridge.
Styginidae gen. et sp. indet.
(Fig. 8g)
Material. NMW 2005.32G.156, pygidium with parts of
four thoracic segments and an associated librigena, internal
mould (Pl, 13·5, Pw, 21·1, PAl, 8·0, PAw, 4·0); sample 205.
Description. Pygidium gently convex, slightly transverse,
suboval in outline and about 65% as long as wide. Axis
narrow, about 60% length (sag.) of pygidium and occupying
about 10% of the anterior pygidial width, with six well-dened
rings and a seventh, poorly-dened, posterior ring. Terminal
piece occupying posterior third of axis, with bluntly rounded
posterior margin. Ring furrows narrow (exsag.), well dened
on anks of axis and almost obsolete sagittally. Axial furrows
deep and narrow, very slightly converging posteriorly. Pleural
elds weakly convex with six pairs of at axial ribs, which
widen distally and are separated by narrow, deep pleural
furrows which extend to margin. Border very weakly dened,
relatively narrow, weakly concave. Pygidial surface nely
Remarks. This specimen is clearly a moult, but without an
associated cephalon, its generic aliation is uncertain. The
pygidium resembles those from the Anderken Formation
(Sandbian) of the Chu-Ili Range assigned to Bronteopsis (?)
extraordinaria by Chugaeva (1958, p. 107, pl. 11, gs 12–14) in
having a narrow, elongate axis occupying about two thirds of
pygidial length, six pairs of weakly convex, almost at pleural
ribs widening distally, and a nely granulose ornament. It
diers in having ring furrows that are almost obsolete sagit-
tally, and a narrower axis occupying only 10% of the maxi-
mum width, and with axial furrows only slightly convergent
In describing Bronteopsis (?) extraordinaria, Chugaeva
(1958, p. 107, pl. 11, gs 10, 11) associated a cephalon of a
blind form with the pygidium discussed above, and later Prˇibyl
& Vaneˇk (1971) made this the type species of their genus
Chugaevia. The xigena of the Karagach specimen has a wide
doublure, the base of a genal spine and a poorly preserved
eye surface, and assuming that the cephalon and pygidium
were correctly associated by Chugaeva (1958) it cannot be
attributed to Chugaevia.
Kolova (1936, p. 34, pl. 1, g. 14) described and gured a
pygidium broadly similar to ours from the Ordovician of
Kara Sai, Dzhebagly Mountains, as Bronteus a. andersoni
Nicholson & Etheridge, 1879. It is immediately distinguished
in having a much shorter (sag.), more tapering axis with more
(seven) pairs of ribs which appear to be more convex in cross
Family Trinucleidae Hawle & Corda, 1847
Subfamily Trinucleinae Hawle & Corda, 1847
Genus Kimakaspis gen. nov.
Derivation of name. From Kimak, the ancient Turkish
tribe who inhabited the area between Irtysh River and
Tarbagatai Range.
Type and only species. Kimakaspis kovalevskyi gen. et sp.
nov. Upper Ordovician, Sandbian Stage, Karagach Forma-
tion, sample 201, east side of Ayaguz river west of Karagach
village, Tarbagatai Range, Kazakhstan.
Diagnosis. Fringe with pits on upper lamella dominantly
radially arranged in relatively narrow, deep sulci, containing
pits E
and I
. I pits interior of I
in radial alignment with
sulci frontally, randomly distributed and out of radial align-
ment laterally and posterolaterally. Fringe with E
developed abaxially very close to E
, diverging from it
posterolaterally until maximum curve of fringe, then converg-
ing to it posteriorly. I
and I
complete; I
very close to I
frontally, with median two I
pit pairs merging; I
in distance from I
laterally, but cannot be traced anterolater-
ally due to irregular arrangement of I pits between I
and I
Apart from I
and I
, I pits irregularly arranged. Around four
irregular arcs between I
and I
at anterolateral corners, up to
eight posterolaterally. Lateral eye ridges and tubercles absent.
Pygidium with ve axial rings plus terminal piece.
Remarks. In possessing two arcs of E pits, which lie very
close together, Kimakaspis is similar to Bergamia Whittard,
1955 and Tretaspis M‘Coy, 1849. On the lower lamella of
Bergamia, the I pits are arranged in a strong, regular, radial
pattern, which only becomes slightly irregular in the genal
angle of the fringe. In contrast, the lower lamella of
Kimakaspis has regular, radially arranged I pits only frontally;
abaxially, the irregular distribution of I pits makes it imposs-
ible to trace radial rows of pits in a straight line from I
to I
The pygidia of the two taxa are similarly short (Bergamia has
up to six axial rings) and wide, but the axis of Bergamia
Figure 7 (a–e) Birmanites akchiensis sp. nov.: (a) NMW 2005.32G.182, paratype, left librigena, latex cast of
external mould, dorsal view, 1·5, sample 203b; (b) NMW 2005.32G.181, holotype cranidium, latex cast
of external mould, dorsal view, 2·5, sample 203b; (c) NMW 2005.32G.183, paratype, hypostome, latex cast of
external mould, ventral view, 3·5, sample 203a; (d) NMW 2005.32G.184, paratype, incomplete pygidium, latex
cast of external mould, dorsal view,1·75, sample 203b; (e) NMW 2005.32G.185, paratype, incomplete
pygidium, internal mould, dorsal view, 1·75, sample 203b. (f–h) Remopleurides cf. sibiricus Vasilievsky in
Weber, 1948: (f) NMW 2005.32G.178, pygidium with attached thoracic segments, internal mould, dorsal view,
4·75, sample 203; (g) NMW 2005.32G.136, pygidium with attached thoracic segments, latex cast of external
mould, dorsal view, 5·75, sample 203; (h) NMW 2005.32G.179, incomplete cranidium, latex cast of external
mould, dorsal view, 5, sample 203. (i) Symphysops cf. djebaglensis (Kolova, 1936): NMW 2005.32G.190,
pygidium, latex cast of external mould, dorsal view, 3·5, sample 205. (j–k, m) Cyclopyge cf. tereki Koroleva,
1967: (j) NMW 2005.32G.186, cranidium and incomplete thorax, internal mould, 6; (k) NMW 2005.32G.187,
pygidium and thorax, internal mould, dorsal view, 5·5; (m) NMW 2005.32G.188, pygidium and thorax, latex
cast of external mould, 9. All sample 205. (l) Microparia (Heterocyclopyge) sp., NMW 2005.32G.189,
pygidium, internal mould, dorsal view, 4, sample 205.
continues onto the posterior border, while in Kimakaspis the
axis ends at the posterior margin.
Tretaspis has pits E
and I
in radial sulci on the upper
lamella, with pits I
outside the sulci but in alignment with
them (Hughes et al. 1975), while Kimakaspis has pits E
in radial sulci, and the I pits inside this are only aligned
with the sulci anteriorly, becoming more irregularly distrib-
uted laterally. Tretaspis also possesses lateral eye ridges and
tubercles, both of which are absent from Kimakaspis.
Although most species of the Katian/Ashgill genus Nanki-
nolithus Lu, 1954, have just one arc of E pits, some have at
least a partial E
arc. N. yanhaoi Zhou & Hughes, 1989, has an
arc of small E
pits, although this is present only anteriorly
(11–12 pits per half fringe), whereas Kimakapsis has only E
anterior of the glabella, with E
appearing close to E
abaxial of the axial furrows, and pits in both arcs are of equal
size. N. granulatus (Wahlenberg, 1818) also diers in having E
anteriorly, although in this species E
continues posteriorly
Figure 8 (a–d) Dulanaspis karagachensis sp. nov.: (a) NMW 2005.32G.150, holotype, incomplete cephalon,
internal mould, dorsal view, 3, sample 205; (b) NMW 2005.32G.152, paratype, pygidium, internal mould,
dorsal view, 3, sample 203a; (c) NMW 2005.32G.151, paratype, pygidium, internal mould, dorsal view, 3·25,
sample 203b; (d) NMW 2005.32G.153, paratype, pygidium with attached thoracic segments, internal mould,
dorsal view, 4, sample 203a. (e) Ceraurinella?a. kasachstasnica (Chugaeva, 1958), NMW 2005.32G.157,
cranidium, internal mould, dorsal view, 2·75, sample 203a. (f, i) Sphaerocoryphe sp.: (f) NMW 2005.32G.160,
cranidium, external mould, dorsal view, 5, sample 203a; (i) NMW 2005.32G.161, pygidium, internal mould,
dorsal view, 6, sample 203b. (g) Styginidae gen. et sp. indet.: NMW 2005.32G.156, pygidium with parts of four
thoracic segments, and associated librigena, latex cast of external mould, 5, sample 205. (h) Encrinuroides cf.
septemcostatus Kolobova, 1983: NMW 2005.32G.163, cranidium, latex cast of external mould, dorsal view, 5,
sample 203b. (j–n) Sinocybele weberi (Kolova, 1936): (j) NMW 2005.32G.165, cranidium, internal mould, dorsal
view, 5·25, sample 203a; (k) NMW 2005.32G.166, cranidium, latex cast of external mould, dorsal view, 5·5,
sample 203; (l) NMW 2005.32G.168, pygidium, latex cast of external mould, dorsal view, 5, sample 203b; (m)
NMW 2005.32G.169, cranidium, internal mould, dorsal view, 2·75, sample 203a; (n) NMW 2005.32G.167,
cranidium, latex cast of external mould, dorsal view, 5·5, sample 203.
Figure 9 Kimakaspis kovalevskyi gen. et sp. nov.: (a–b) NMW 2005.32G.241, paratype, articulated exoskeleton,
latex cast, dorsal and lateral views, 4·5; (c–d) NMW 2005.32G.240, holotype, articulated exoskeleton, internal
mould, lateral and dorsal views, 4; (e) latex cast of previous, ventral view, 3; all from sample 201; (f) NMW
2005.32G.242, paratype, incomplete enrolled exoskeleton, internal mould, (3·75) sample 205.
rather than being restricted to the anterior of the fringe. N.
granulatus also diers from Kimakaspis in having a wider
fringe with more I arcs anteriorly (at least four in the former,
only three in the latter) and along the posterior margin (up to
11, rather than 8). Zhou & Hughes (1989) note that the few
Nankinolithus species with partial E
arcs are middle Ashgill
(Katian) in age, whereas older members of the genus (early
Ashgill) have just E
Although Kimakaspis is older than known species of
Nankinolithus that possess E
, it is more similar to this genus
than to Bergamia and Tretaspis, in the irregularity of some I
arcs laterally (i.e. not all pits are aligned into concentric arcs)
and because the I arcs are only distributed in a regular radial
pattern anteriorly. However, where E
is developed in
Nankinolithus, its distribution is dierent to that of
Kimakaspis; the E
pits are developed anteriorly and are
generally smaller than those of E
, whereas E
abaxially in Kimakaspis, from pits of similar size to E
that are
initially twinned with them. It may be that the development of
in both taxa is entirely homologous, although the strong
similarities in the pattern of the I arcs suggests a genuine
relationship between them, and Kimakaspis may represent the
ancestral stock from which Nankinolithus evolved.
A similar irregular arrangement of pits in the I arcs is seen
in Declivolithus Prˇibyl & Vaneˇk, 1967, from the Bohdalec
Formation (Katian) of the Czech Republic and similar hori-
zons in Morocco, although this genus has a much broader
‘harpiform’ (Hughes et al. 1975) fringe and has only one E arc.
The type material of Jianxilithus latimarginis Zhang & Zhou
in Lu et al., 1976, the single species assigned to this late
Sandbian–early Katian genus from China is incompletely
preserved, but Zhou & Hughes (1989) note that the ‘‘fringe is
reminiscent of Declivolithus’’, in having only E
and randomly
distributed inner I arcs. However, Jianxilithus lacks a well-
dened pseudofrontal glabellar lobe, so additional material is
required to clarify its relationship to Declivolithus and other
genera. In addition to lacking E
, Jianxilithus diers from
Kimakaspis in the shape of the glabella and in possessing eye
Kimakaspis kovalevskyi gen. et sp. nov.
(Figs 9, 10, 11e, g, 13i)
Derivation of name. After the late O. P. Kovalevsky, in
appreciation of his contribution to the study of the Ordovician
geology and palaeontology of the Tarbagatai Range.
Holotype. NMW 2005.32G.240, articulated exoskeleton,
internal mould (L, 6·4, W, 3·9), sample 201.
Paratypes. NMW 2005.32G.216, 241, 252, disarticulated
exoskeletons internal and external moulds; NMW
2005.32G.254, incomplete exoskeleton; NMW 2005.32G.216,
cephalon, external mould; NMW 2005.32G.272, external
mould of damaged articulated exoskeleton; NMW
2005.32G.248–251, 253 incomplete cephala, internal moulds;
NMW 2005.32G.247, pygidium with attached thoracic seg-
ments, internal mould; all sample 201. NMW 2005.32G.242,
incomplete enrolled exoskeleton; NMW 2005.32G.243–245,
fragments of fringe; sample 205.
Diagnosis. As for genus.
Description. Cephalon sub-semicircular in outline, al-
though one large specimen (Fig. 9a) has angular anterolateral
corners. Fine reticulate sculpture covering glabella and genal
lobes. Glabella has maximum width (tr.) at around midlength
(sag.) of pseudofrontal lobe, equal to approximately 60% of its
length (sag.). Axial furrows tapering gently anteriorly; rela-
tively deep and narrow (tr.) anteriorly, rapidly widening pos-
terior of 3p. Pseudofrontal lobe of glabella prominent, inated,
suboval, overhangs fringe slightly; separated from posterior of
glabella by wide (sag.), relatively shallow, transverse furrow 3p
faintly impressed on sides of pseudofrontal lobe, just posterior
of mid-lobe length (sag.). 2p furrows are deep, elongate, oval
pits at posterior of pseudofrontal lobe; 1p furrow pits and
occipital pits dene corners of gently convex occiput, which is
separated from occipital ring by shallow furrow. Occipital ring
narrow (sag.), convex, tapers gently abaxially. No occipital
spine. Area adaxial of 1p, 2p and 3p furrows swollen to form
a composite lateral glabellar lobe, delimited adaxially by axial
furrow. Posterior cephalic border narrow (exsag.), convex,
separated from genal lobes by relatively deep, narrow furrow.
Eye tubercles and ridges absent.
Fringe with E
, and I
, with numerous additional I pits
laterally. No F pits present. Of E pits, only E
is present
medially, comprising around seven pits in front of glabella.
Just abaxial of axial furrows, E
developed very close to E
diverging from E
posterolaterally until maximum antero-
lateral curve of fringe; then E
and E
converge posteriorly,
merging so that just a single pit is present at the posterior limit
of E arcs. Girder strong and clearly expressed, of consistent
width along the length of the fringe. I
and I
complete; I
close to I
frontally, with median two I
pit pairs merging; I
diverging from I
laterally, but cannot be condently traced
anterolaterally due to irregular arrangement of I pits between
and I
. Abaxially, I
splits to form two concentric arcs of
pits, running very close to each other, but gradually diverging
posteriorly; the I arc outside I
does not cut it o. Another
concentric arc of pits runs outside this arc, but it stops and
does not continue along the front of the fringe. Inside I
laterally, there is a roughly concentric arc of pits that may
represent a continuation of I
, but some of the pits are out of
alignment so that it is not a perfectly regular arc. At the
anterolateral corners of the fringe, there are four arcs between
and I
, but although roughly concentric, these are not
arranged in perfectly regular curves and increasing numbers of
additional pits become introduced to the middle part of the
fringe posteriorly, producing up to eight arcs of I pits postero-
laterally. Pits on upper lamella dominantly radially arranged in
relatively narrow, deep sulci (27–30 per half fringe), containing
pits E
and I
. I pits interior of I
in radial alignment with
sulci frontally, but randomly distributed and out of alignment
laterally and posterolaterally. In one specimen, two pairs of
anterolateral sulci are interspersed with deep single pits next to
the border, producing Y-shaped inter-radial ridges. Genal
spines long, extending beyond pygidium.
Thorax of six segments; distal-most ends of pleurae
deected downwards; deep apodemal pits, continuous with
shallow pleural furrows directed obliquely backwards; deep
articulating pits in axial furrows; prominent articulating half
ring on segments 2–6 (Fig. 9d, e), but specimens do not show
whether this is developed on anterior-most segment.
Pygidium subtriangular, transverse, length (sag.) one-
quarter to one-third maximum width (tr.). Axis gently convex;
maximum width one-quarter maximum pygidium width;
tapers gently to posterior margin, but does not continue onto
vertical, relatively wide posterior margin. Axis dened by wide,
shallow axial furrows. Axis with ve rings plus terminal piece,
and up to four interpleural furrows.
Remarks. The presence of two Y-shaped inter-radial
ridges on the fringe of one specimen of K. kovalevskyi is
reminiscent of Stapeleyella Whittard, 1955. However, the latter
has this distinctive ridge pattern consistently developed all
around the fringe, and the isolated occurrence of these ridges
in a single specimen of K. kovalevskyi is thought to reect a
deformity in fringe growth, an occurrence that Hughes (1971)
recognised as not uncommon in Trinucleus fimbriatus.
Family Dionididae Gu¨rich, 1907
Genus Dionide Barrande, 1847
Type species. Dione formosa Barrande, 1846, Upper
Ordovician, Sandbian Stage, Vinice Formation, Trubı´n, Czech
Dionide sp.
(Fig. 6c)
Material. NMW 2005.32G.271, cephalon with attached
thoracic segments, internal mould; NMW 2005.32G.265,
articulated specimen, comprising incomplete cranidium,
thorax and pygidium, external mould (L, 10·4, Cl, 4·2, Cw, 6·9,
Pl, 3·8, Pw, 6·6), sample 201.
Remarks. The only late Ordovician species of the genus
described from the Tarbagatai Range is Dionide tarbagataica
Koroleva, 1979, from an unidentied locality and unit. It is
based on a single cephalon, and the pygidium is unknown.
The articulated specimen has a slightly transverse pygidium
with ten axial rings plus terminal piece. This is narrower (tr.)
and has a narrower axis than other known species from
Kazakhstan (e.g. D. ammophila Koroleva, 1979; D. altaica
Koroleva, 1979, D. plana Koroleva, 1979 and D. kasachstanica
Chugaeva, 1958). The Karagach cranidium diers from most
of these, and also from D. tarbagataica, in having much
narrower (tr.), longer (sag.) xigenae with more evenly curved
antero-lateral margins that reach a much more anterior pos-
ition relative to the glabella, forming a virtually continuous arc
with its anterior margin. D. altaica has similar xigenae, but its
glabella is subrounded and of roughly equal length and width,
whilst that of Dionide sp. is suboval, with its width (tr.) around
four-fths of its length (sag.).
Genus Aethedionide Zhou & Ju in Qiu et al., 1983
Type species. Aethedionide fusiformis Zhou & Ju in
Qiu et al., 1983; Upper Ordovician, late Katian Stage,
Huangnehkang Formation, western Zhejiang Province, South
Aethedionide sp.
(Fig. 6f)
Material. NMW 2005.32G.191, pygidium, internal mould
(Pl, 4·7, Pw, 4·8, PAw 1·4); sample 205.
Remarks. This pygidium is subtriangular and of almost
equal length and width, with an axis bearing 18 axial rings plus
terminal piece, and a pleural eld with 16 pairs of ribs. It is
similar to that of Aethedionide fusiformis Zhou & Ju in Qiu
et al., 1983 in its narrow axis and large number of axial rings
and pleural ribs, but diers in being more elongate, and in
lacking distinct median tubercles on the axis.
Family Raphiophoridae Angelin, 1854
Genus Raphioampyx Baldis & Baldis, 1995
Type species. Raphioampyx argentinus Baldis & Baldis,
1995, from Sandbian Stage, Las Aguaditas Formation, San
Juan, Argentina.
Raphioampyx? sp.
(Fig. 6e)
Material. NMW 2005.32G.162, disarticulated exo-
skeleton, comprising cranidium, four thoracic segments and
pygidium; sample 201.
Remarks. The triangular cranidium with short (tr.) xi-
genae, short anterior spine and smooth, gently convex, rhom-
boidal glabella, is consistent with assignment to Raphioampyx,
as are the deep, concave axial furrows. However, the single
specimen at hand is assigned only tentatively to Raphioampyx
because the posterior border and occipital ring are not pre-
served, so it is impossible to determine whether these are fused
to form a continuous structure, a character that is considered
diagnostic of the genus (Baldis & Baldis 1995; Turvey 2007).
Four thoracic segments are preserved, and because the speci-
men is disarticulated behind the cranidium, it is possible that
the full complement is ve, which is typical for Raphioampyx.
Turvey (2007) described R. sinankylosus from the lower
Darriwilian of northern Hunan, south China, and suggested
that Lonchodomas jiantsaokouensis Lu, 1975, from the
Caradoc (Sandbian–Katian) of south China and Thailand
(Fortey 1997), L. nanus Zhou, in Zhou et al., 1982 (see Zhou &
Dean 1986), from the Sandbian of Gansu and Ningxia,
northwest China and Ampyx reedi Yin, 1937, from the
Darriwilian of western Yunnan, south China, also belong to
Raphioampyx. The Kazakhstanian specimen is too incomplete
for detailed comparison, although L. nanus clearly diers from
it in having much wider (tr.) xigenae.
Family Cheiruridae Hawle and Corda, 1847
Subfamily Cheirurinae Hawle and Corda, 1847
Genus Ceraurinella Cooper, 1953
Type species. Ceraurinella typa Cooper, 1953, Upper
Ordovician, Sandbian Stage, Edinburg Formation, Virginia,
Ceraurinella?a. kasachstanica (Chugaeva, 1958)
(Fig. 8e)
Material. NMW 2005.32G.157, cranidium, internal mould
(Gl, 12·4. Gw, 11·9); NMW 2005.32G.158, 159, incomplete
internal moulds of cranidia; sample 203a.
Description. Glabella subrectangular, about 113% as long
as wide (excluding occipital ring), with maximum width across
mid part of the anterior lobe; gently convex in sagittal prole
and moderately convex across the anterior lobe. Anterior
margin of glabella with a small median indentation. Three
pairs of deep, narrow glabellar furrows; S1 directed slightly
backward as far as one third of glabellar width then curved
strongly backwards to merge with the occipital furrow. S2 and
Figure 10 Kimakaspis kovalevskyi gen. et sp. nov. Schematic drawing
showing arrangement of pits on the fringe.
S3 of equal length (tr.), almost parallel, inclined slightly
backwards. L2 slightly shorter (tr.) than L1 and L3. Axial
furrows deep and narrow, slightly concave opposite L2 and
L3, bearing deep anterior pits opposite to S3. Occipital ring
wide medially, tapered abaxially with a small median tubercle
and almost transverse posterior margin. Occipital furrow deep,
convex anteriorly and widening medially. Anterior cranidial
border narrow, convex, almost transverse medially. Fixigenae
incompletely preserved. Glabella tuberculate, xigenae with
coarse pits.
Remarks. These specimens are assigned provisionally to
Ceraurinella mainly because of similarity in cranidial morpho-
logy, including a subparallel-sided glabella, S1 strongly curved
backwards and merged with the occipital furrow, subparallel
S2 and S3 of about equal size. They dier, however, from the
type species, C. typa, in having longer (tr.), slightly posteriorly
inclined S2 and S3, the outline of the anterior glabellar margin,
and the almost transverse median part of the anterior cranidial
margin. They broadly resemble the cranidium described by
Chugaeva, 1958 (p. 83, pl. 9, g. 8) as Cheirurus [possibly=
Ceraurinella?] kasachstanicus from the Duklankara Formation
(lower Katian) of the Chu-Ili Range, but are distinguished
from it in having S2 and S3 slightly posteriorly inclined
adaxially and a narrower (tr.) frontal lobe of the glabella with
a far more prominent median indentation on its anterior
margin. Another similar species is Ceraurus exornatus
Lisogor (1965, p. 176, pl. 2, gs 1, 2) from the Angrensor
Formation (Katian) of north-eastern central Kazakhstan,
but unlike Ceraurinella? kasachstanica, it has a wider (tr.)
frontal lobe with an evenly rounded, convex anterior mar-
gin, S3 slightly inclined anteriorly, S1 strongly shallowing
Subfamily Deiphoninae Raymond, 1913
Genus Sphaerocoryphe Angelin, 1854
Type species. Sphaerocoryphe dentata Angelin, 1854,
Upper Ordovician, Katian Stage, Upper Jonstorp Formation,
Va¨stergo¨tland, Sweden.
Sphaerocoryphe sp.
(Fig. 8f, i)
Material. NMW 2005.32G.160, cranidium, external
mould (Cl, 3·5, Cw, 6·9, Gl, 2·4, Gw, 3·1); sample 203a; NMW
2005.32G.161, pygidium, internal mould (Pl, 3·3, Pw, 5·2,
PAw, 2·8); sample 203b.
Description. Cranidium transverse, about half as long
(sag.) as wide (tr.), with glabella dominated by a large, almost
spherical anterior lobe, which overhangs anterior cranidial
margin and occupies about 70% of cranidial length (sag.). L1
very small, tubercle-like, S1 merging with occipital furrow
adaxially of L1 to form a wide (sag., exsag.), depressed strip,
which separates a convex occipital ring from frontal lobe.
Axial furrows deep and narrow, strongly divergent anteriorly.
Fixigenae strongly convex, palpebral lobes small, situated
slightly anterior to base of frontal lobe. Posterior border
transverse, ridge-like, separated from xigenae by a deep and
wide border furrow. Genal spines relatively short and slender,
bending strongly posteriorly.
Librigenae, hypostome and thorax unknown.
Pygidium transverse, subtrapezoidal in outline, about 65%
as long as wide (excluding spines). Sagittal length around 40%
of exsagittal length. Axis subconical, strongly convex with two
narrow (sag.), convex axial rings and with bluntly rounded
terminal piece occupying posterior half of the axis. Pleural eld
weakly convex with two pairs of broad, indistinct pleural ribs
terminated with pygidial spines. Anterior pair of spines almost
transverse, exceeding in length half of pygidial width; posterior
pair of spines broad and robust, about twice as long as
pygidium, diverging posterolaterally for anterior half of their
length, then curving evenly backwards. Span (tr.) of anterior
pair of spines slightly longer than that of posterior pair of
spines. Short, straight lateral margin between anterior and
posterior spines.
Remarks. Although there have been previous reports of
Sphaerocoryphe in Kazakhstan (e.g. Nikitin 1972), the speci-
mens from the Karagach Formation are the rst representa-
tives of the genus formally described. Tripp et al. (1997)
summarised the characters of the thirty or so known species of
Sphaerocoryphe, noting that establishing relationships within
the genus is dicult, due to dierent combinations of charac-
ters occurring in dierent species. Unfortunately, one of the
most useful discriminating characters that they identied, the
presence or absence of one or two pairs of proxigenal spines,
cannot be applied to the Karagach species because they are not
preserved on the single available cranidium. Another useful
character is the length (sag.) of the anterior lobe of the
glabella, expressed as a percentage of the sagittal length of the
cranidium (Tripp et al. 1997). The value for Sphaerocoryphe
sp. is 70%, and it falls into a group of species for which the
range is 65–75%. Another distinctive character that it possesses
is the presence of robust, almost transverse anterior pygidial
spines. Four other species of Sphaerocoryphe share this com-
bination of glabellar and pygidial morphology: S dentata
Angelin, 1854, S. pemphis Lane, 1971, S. punctata (Angelin,
1854) and S. murphyi Owen et al., 1986.
Sphaerocoryphe sp. most closely resembles S. thomsoni
Reed, 1906, from the South Threave Formation, Drummuck
Subgroup (Ashgill) at Thraive Glen, Girvan, Ayrshire, which
Kielan-Jaworowska et al. (1991) and Tripp et al. (1997) regard
as a junior subjective synonym of the type species, S. dentata
Angelin, 1854, and this synonymy is accepted in the present
paper. The cranidia of S. dentata and Sphaerocoryphe sp. share
convex cheeks, a narrow, transverse, ridge-like posterior bor-
der, a wide, deep posterior border furrow and a wide (sag.),
shallow strip separating L1 and the occipital ring from the
inated frontal lobe of the glabella. Although the genal spines
in S. dentata curve strongly posteriorly, those of Sphaero-
coryphe sp. curve back at a sharper, more acute angle. The
posterior border furrow in S. dentata fades out abaxially,
resulting in a ‘bridge’ in the genal angle between the border
and xigenal eld (Tripp et al. 1997); in contrast, the posterior
border furrow continues without interruption around the genal
angle into the lateral border furrow in Sphaerocoryphe sp. The
pygidium of S. dentata diers in having four axial rings plus a
small terminal piece as opposed to only three axial segments in
total in Sphaerocoryphe sp. The pygidial spines are of similar
size and arrangement in both species, although the anterior
pair in S. dentata is relatively straight for half of their length
(tran.) and then curve back posteriorly, while those of Sphae-
rocoryphe sp. are apparently straight and almost transverse for
their entire length.
The cephalon of S. pemphis Lane, 1971, from the Balclatchie
Formation (Sandbian) at Dow Hill, near Girvan, diers from
that of Sphaerocoryphe sp. in having a posterior border that
widens (sag.) towards the genal angle, as opposed to being
transverse and uniformly narrow. The pygidium of S. pemphis
is also dierent, having four axial rings plus a terminal piece,
instead of three axial segments in Sphaerocoryphe sp. The
former also has a deep anterior interpleural furrow behind the
rst axial ring, and there is no gap between the broad bases of
the anterior and posterior pairs of pleural spines, whereas the
interpleural furrows in Sphaerocoryphe sp. are very weak, and
there are short, straight lateral margins separating the two
pairs of spines on each side of the pygidium.
Sphaerocoryphe punctata (Angelin, 1854), from the Katian
of Sweden, diers from Sphaerocoryphe sp. in having a nar-
row, shallow posterior border furrow, which fades out abaxi-
ally, while the latter has a broad, prominent posterior border
furrow that is continuous with the lateral furrows. S. punctata
also has a strong axial furrow alongside the occipital ring,
while that of Sphaerocoryphe sp. is relatively weak.
Sphaerocoryphe murphyi Owen et al., 1986, from the Raheen
Formation (late Caradoc) of County Waterford, Republic of
Ireland, has xigenae that are more weakly swollen than those
of Sphaerocoryphe sp. as well as having a narrower posterior
border furrow, and genal spines that are straight and diver-
gent, as opposed to curving strongly posteriorly in the latter.
The palpebral lobes are also positioned farther forward in S.
murphyi. The pygidium of S. murphyi has posterior spines that
are more robust and more strongly divergent than those of the
Karagach species, resulting in a longer span (tran.) between the
tips of the posterior spines than between the anterior, almost
horizontally directed, pair. In contrast, the anterior pair of
pygidial spines in Sphaerocoryphe sp. extends more trans-
versely and has a span slightly greater than that of the
posterior pair of spines. Further specimens of Sphaerocoryphe
sp. are needed in which the proxigenal spines are preserved in
order to establish in more detail its relationship to other
known species.
Subfamily Acanthoparyphinae Whittington & Evitt, 1954
Acanthoparyphinae gen. indet.
(Fig. 12e)
Material. NMW 2005.32G.149, incomplete cranidium,
external mould; sample 203b.
Remarks. The posterior part of this cranidium is missing.
Part of the left hand L1 is preserved, and is dened by a deep
S1, of which the adaxial end is deected backwards. Weak,
subparallel S2 and S3 are directed backwards at a similar angle
to S1, and dene elongate (trans.) L2 and L3. These characters
are consistent with those of Acanthoparyphinae, but generic
assignment is not possible on the basis of this specimen.
Family Encrinuridae Angelin, 1854
Subfamily Encrinurinae Angelin, 1854
Genus Encrinuroides Reed, 1931
Type species. Cybele sexcostata Salter, 1848; Upper
Ordovician, Ashgill Series (Katian Stage), Sholeshook Lime-
stone Formation, Pembrokeshire, Wales. Neotype selected and
redescribed by Whittington (1950).
Encrinuroides cf. septemcostatus Kolobova, 1983
(Fig. 8h)
Material. NMW 2005.32G.163, incomplete cranidium,
external mould (Cl, 6·8, Gl, 5·1, Gw, 4·9); NMW
2005.32G.164, incomplete cranidium, external mould; sample
Description. Glabella moderately convex with maximum
width slightly anterior of S3. Anterior margin almost semi-
circular. Glabellar base narrow, slightly exceeding 60% of
maximum glabellar width. Three pairs of transverse glabellar
furrows, deep abaxially, rapidly shallowing adaxially. L1 nar-
row, ridge-like, less than half as long as L2 (exsag.); L3 and L2
about equal length (exsag.). Frontal lobe almost semicircular,
with a narrow, shallow longitudinal median furrow on its
anterior part. Small pit possibly present in preglabellar furrow.
Axial furrows deep and narrow, strongly divergent anteri-
orly. Glabella ornamented with numerous ne, irregularly
distributed tubercles, with a single transverse row at L1 and a
single median tubercle opposite S1. Tubercles numerous on L2
and L3, arranged in two irregular rows on each. Occipital
furrow deep and wide (sag.), slightly convex anteriorly. Occipi-
tal ring strongly convex (sag., exsag.), smooth, slightly wider
than base of preoccipital glabella. Fixigenae wide (tr.), sub-
triangular, moderately convex, ornamented with irregular,
coarse tubercles and pits. Anterior cranidial border narrow,
with four pairs of strong tubercles and two pairs of ne
tubercles between them. Palpebral lobes high, raised, smooth,
with their anterior ends situated opposite the middle part of L2
and the posterior ends opposite S1. Posterior cranidial border
narrow (exsag.), ridge-like, turning gently backwards abaxi-
ally, separated from xigenae by deep and narrow (exsag.)
border furrow.
Remarks. These cranidia closely resemble those of
Encrinuroides septemcostatus Kolobova (1983, p. 255, pl. 1,
gs 9, 12) from the Anderken Regional Stage (Sandbian) of the
Chingiz Range in the shape of the glabella, characteristics of
the glabellar furrows and the ornament of numerous ne
tubercles covering the glabella and xigenae, and in having the
anterior cephalic border visible in dorsal view. Without an
associated pygidium, they cannot condently be assigned to
E. septemcostatus. Both the present cranidia, and those of E.
septemcostatus are immediately distinguished from that of
E. sexcostatus (Salter) (see Whittington 1950, p. 535, pl. 68,
gs 7–10) in having a less expanded frontal glabellar lobe
which does not overhang the anterior cephalic margin.
Subfamily Cybelinae Holliday, 1942
Genus Sinocybele Sheng, 1974
Type species. Sinocybele baoshanensis Sheng, 1974, Middle
Ordovician, Darriwilian, Pupiao Formation, western Yunnan,
Remarks. Chen & Zhou (2002, p. 223), Zhou & Zhou
(2006, p. 393) and Edgecombe & Webby (2006, p. 414) have
shown that Sinocybele is a senior subjective synonym of
Koksorenus Koroleva (type species K. kazakhstansensis
Koroleva, 1992). The present authors accept the arguments of
these authors and use Sinocybele herein.
Sinocybele weberi (Kolova, 1936)
(Figs 8j–n, 11a, d)
1936 Cybele weberi Kolova, p. 35, pl. 1, gs 15, 16.
1948 Cybele weberi Kolova; Weber, p. 64, pl. 10, gs 23, 24.
Lectotype (here selected). CNIGR 60/4263 (Fig. 11d) dor-
sal exoskeleton, internal mould; paralectotype: CNIGR 61/
4263, incomplete pygidium, external mould. Both from lower
Katian Stage, Besharyk Formation, Dzhebagly Mountains.
Material. NMW 2005.32G.165, cranidium, internal
mould; sample 203a; NMW 2005.32G.166, cranidium, external
mould (Cl, >4·5, Cw, 9·1, Gl, 2,5, Gw, 2·5), sample 203; NMW
2005.32G.167, cranidium, external mould (Cl, >3·6, Cw, 6·4,
Gl, 2,9, Gw, 3·2), sample 203; NMW 2005.32G.169, cranid-
ium, internal mould, sample 203a; NMW 2005.32G.168,
pygidium, external mould (Pl, >6·7, Pw, >8·8, PAl, 4·5, PAw,
2,3); sample 203b; NMW 2005.32G.196, pygidium, external
mould; sample 203; NMW 2005.32G.197, pygidium, internal
mould; sample 203; NMW 2005.32G.171, pygidium, external
mould, locality as for lectotype.
Description. Cranidium almost semicircular, about twice
as wide as long. Glabella strongly forwardly expanded, with
inated frontal lobe, slightly overhanging anterior margin of
cranidium. S1 strongly curved backwards, separating small,
elongate suboval L1; S2 and S3 small, pit-like. S4 visible on
some internal moulds (Fig. 8m). Glabellar base about 40% of
maximum glabellar width (tr). Axial furrows deep and wide,
subparallel between S3 and occipital furrow, then straight,
broadly divergent anteriorly. Glabellar surface ornamented
with irregular tubercles, with sparse larger ones interspersed
with numerous small ones. Occipital furrow narrow (sag.),
convex anteriorly, pit-like abaxially. Occipital ring strongly
convex (sag., tr.), strongly tapered abaxially with a posteriorly
placed median tubercle. Fixigenae wide (tr.), subtriangular,
convex, with narrow genal spines directed posterolaterally;
ornamented by deep, polygonal pits and sparse coarse tuber-
cles. Eye stalks anteriorly directed, with bases situated on
anterior part of xigena, located approximately opposite the
widest part of glabella. Border furrow deep and narrow (sag.).
Posterior border narrow (sag.), transverse proximally, curving
posteriorly opposite the base of palpebral lobes and then
widening abaxially. Librigenae unknown.
Thorax with 12 segments, sixth segment macropleural,
bearing long spines directed backwards far beyond posterior
end of pygidium.
Pygidium subtriangular in outline with a narrow, strongly
convex axis occupying about 25% of maximum pygidial width
and bearing 14 axial rings and a short terminal piece. Anterior
Figure 11 (a, d) Sinocybele weberi Kolova, 1936: (a) CNIGR 61/4263, paralectotype, incomplete pygidium, latex
cast of external mould, 9·5; (d) CNIGR 60/4263, lectotype, incomplete dorsal exoskeleton, latex cast of
external mould, 6·75. Both from Besharyk Formation (lower Katian), Dzhebagly Mountains. (b–c) Diacan-
thaspis? sp.: NMW 2005.32G.267a–b, dorsal exoskeleton, latex cast of external mould, and internal mould,15,
sample 201. (e, g) Kimakaspis kovalevskyi gen. et sp. nov. NMW 2005.32G.216, paratype, disarticulated
exoskeleton: (e) dorsal view, 5; (g) pygidium and thoracic segments, detail, 8·5, sample 201. (f, h) Nileus sp.:
(f) NMW 2005.32G.193, cephalon with attached thoracic segments, latex cast of external mould, dorsal view,
6·5, sample 203; (h) NMW 2005.32G.194, pygidium, internal mould, dorsal view, 4, sample 203a.
axial rings complete; ring furrows on posterior part of axis
almost obsolete medially. Axial furrows narrow, well dened.
A narrow postaxial ridge is present between terminal piece and
posterior pygidial margin. Pleural elds with three pairs of
ribs, separated by wide, shallow pleural and interpleural
furrows, in which shallow pits are present (Fig. 11a, d).
Anterior pleural bands narrow, depressed; posterior pleural
bands and far more prominent, convex, upstanding and extend
distally into what appear to be short spines. Pleural ribs
become progressively less backwardly curved so that third pair
runs more or less exsagittally.
Remarks. Kolova’s (1936, pl. 1, gs 15, 16) and Weber’s
(1948) photographs of the exoskeleton and a pygidium of
Cybele weberi were strongly retouched, and details of the
outline and characteristics of the pleural ribs of the pygidium
can hardly be seen. Restudy of the type specimens, re-
illustrated here as Figure 11a and d, shows that they have the
morphology described above. Because there is no signicant
dierence in cranidial and pygidial morphology (genal spines
are not preserved in the types) between specimens from the
Dzhebagly Mountains and those from the Tarbagatai Range,
they are considered conspecic. Between them they show
morphological features characteristic of Sinocybele, including
the macropleural sixth thoracic segment and three well devel-
oped pairs of pygidial pleural ribs (e.g. Edgecombe & Webby
2006, p. 414).
Koksorenus kazakhstanensis Koroleva (1992, g.1a,б, г, в)
from the Karamolinsk Formation (Sandbian), Stepnyak dis-
trict, north central Kazakhstan, is similar to S. weberi, but the
illustrations are unsatisfactory. Detailed comparison is not
possible without reference to the original specimens, and an
attempt by one of us (LEP) to examine them (registered under
the accession number IGNA 1101) during a visit to the
Geological Museum of the Institute of Geological Sciences,
Almaty in 2006 was unsuccessful, because they were not
deposited in the museum, and their present whereabouts is
unknown. The holotype (Koroleva 1992, g.1a,б) has the
glabella much more forwardly expanded than that of S. weberi,
as does the crandium from Inner Mongolia attributed to
Sinocybele kazakhstanensis by Zhou & Zhou (2006, g. 4N),
and this suggests that dierent species are represented. Sino-
cybele gaoluoensis (Zhou in Zhou et al., 1977, p. 260, pl. 79,
gs 1a–b, 2) from the Katian Linxiang Formation of western
Hubei, China is similar to S. weberi, but has a more inated
anterior glabellar lobe, longer, narrower posterior part of the
glabella, longer genal spines and apparently longer, broader
pygidial spines.
Cybele psemica (Abdullaev in Abdullaev & Khaletskaya,
1970) from the Beshtor Formation (late Sandbian/early Ka-
tian) of the Pskem Range, Uzbekistan was attributed to
Sinocybele by Zhou & Zhou (2006, p. 386) and by Edgecombe
& Webby (2006, p. 414). It has a similar glabella to that of S.
kazakhstanensis but the holotype cranidium (Abdulleav in
Abdullaev & Khaletskaya 1970, pl. 4, gs 7, 8 – incorrectly
numbered in plate caption: cf. text, p. 45) has a long, back-
wardly oblique ocular ridge, and the pygidium has deep,
narrow pleural and interpleural furrows. These morphological
features indicate that this species should not be assigned to
Subfamily Dindymeninae Henningsmoen in Moore, 1959
Genus Dindymene Hawle & Corda, 1847
Type species. Dindymene friderciaugusti Hawle &
Corda, 1847; Upper Ordovician, Katian Stage, Kra´lu˚v Dvu˚r
Formation, Czech Republic.
Dindymene brevicaudata Kolobova, 1972
(Figs 5d, 12a–d, f–g)
1972 Dindymene brevicaudata Kolobova; p. 244, pl. 55, gs 13,
Holotype. CNIGR Museum 9/9559, Sandbian Stage,
Karagach Formation, east side of Ayaguz River about 7 km
north of Akchii village, Tarbagatai Range, Kazakhstan.
Material. NMW 2005.32G.176 (L, 7·2, W 7·3, Cl, 2·6,
Cw, 7·3, Gw, 2·1) NMW 2005.32G.177, NMW 2005.32G.210
(L, 7·2, W 7·3, Cl, 2·6, Cw, 7·3, Gw, 2·1), and NMW
2005.32G.211, articulated exoskeletons, internal moulds;
NMW 2005.32G.217, cephalon with attached thorax, external
mould; NMW 2005.32G.215, pygidium, external mould;
NMW 2005.32G.245, pygidium, internal mould; all sample
201. NMW 2005.32G.173, pygidium, external mould (Pl, 3·8,
Pw, 4·2, PAl, 2·7, PAw, 1·9); sample 205.
Description. Exoskeleton slightly elongate suboval, with
maximum width at posterior cephalic margin. Cephalon
almost semicircular, about 50–55% as long as wide. Preoccipi-
tal glabella strongly convex (sag., tr.), expanding anteriorly,
with strongly convex anterior margin, bordered laterally by
narrow and deep axial furrows converging towards occipital
furrow. Base of a median glabellar spine situated at about 25%
of preoccipital glabellar length from the deep, transverse
occipital furrow. Occipital ring narrow with a convex posterior
margin. Fixigenae strongly convex, declined outwards. Pos-
terior border furrow deep, transverse; posterior border narrow
(sag.), convex. Genal spines small, directed posterolaterally.
Cephalon ornamented by coarse, widely spaced tubercles, also
visible on internal moulds.
Thorax of ten segments; deep and wide axial furrows
converging posteriorly, and narrow, convex axis occupying
about 20% of thoracic width. Inner portion of pleurae trans-
verse, subparallel, subdivided into at anterior bands and
more prominent posterior bands. Outer portion of pleurae
directed posteroventrally at fulcrum, produced into wide and
at pleural spines and terminating with rounded distal
extremities. Anterior segments are less curved and directed
posterolaterally; posterior segments curved more strongly
backwards, to lie parallel to pygidial pleurae.
Pygidium slightly transverse, about 90% as long as wide
(including spines). A subconical axis occupies about 45% of
maximum pygidial width, and has six or seven axial rings. First
three axial rings are separated by deep and wide (sag.) ring
furrows, connected to posteriorly directed pleural ribs.
Remaining posterior axial rings fused together to form a
subtriangular structure with very ne ring furrows, which do
not merge with axial furrows. Terminal piece very small,
subtriangular, pointed posteriorly. Axial furrows almost
straight, converging posteriorly. Pleural eld with three pairs
of strong pleural ribs, terminating in three pairs of short, blunt
pygidial spines and separated by wide, deep interpleural fur-
rows. Two anterior pairs of pleural ribs posterolaterlly directed
distally, and then bent strongly posteriorly abaxially. Third
pair of pleural ribs posteriorly directed, with inner margins
conned to axial furrows.
Remarks. Our specimens are topotypes, and are identical
to the type specimens of D. brevicaudata in almost all details,
but Kolobova’s (1972, pp. 244–245) description includes two
apparent points of dierence. Firstly, she noted the presence of
short, posterolaterally directed librigenal spines, which are not
present on our specimens. We are unable to conrm their
presence by examination of her gures (1972, pl. 55, gs 12,
13), and it is possible that the material was misinterpreted.
Secondly, our specimens have six or seven pygidial axial rings,
but Kolobova (1972, p. 245) stated that three or four are
present in D. brevicaudata. The holotype (Kolobova 1972,
pl. 55, g. 13) has a well-preserved thorax comprising ten
segments, behind which the pygidium is slightly displaced. On
the latter, although the illustration is small, at least ve
pygidial axial rings can be discriminated, and it seems
that these were miscounted in the original description. We
have, therefore, no reason to exclude our material from
D. brevicaudata.
Dindymene(?) araneosa Lisogor (1965, p. 181, pl. 2, g. 7)
from the Akzhar Formation (Sandbian, foliaceus Biozone)
of south eastern central Kazakhstan, known from a single
pygidium does not belong to Dindymene. It is a cybeline, and
probably belongs to Lyrapyge (Fortey 1980, p. 100).
Figure 12 (a–d, f–g) Dindymene brevicaudata Kolobova, 1972: (a) NMW 2005.32G.210, articulated exoskeleton,
internal mould, dorsal view 8·5, sample 201; (b) NMW 2005.32G.211, articulated exoskeleton, internal mould,
dorsal view 8·5, sample 201; (c–d) NMW 2005.32G.217, cephalon with attached thoracic segments, internal
mould, dorsal and lateral views, 9·5 and 14, sample 201; (f) NMW 2005.32G.215, pygidium, latex cast of
external mould (probably belongs to the same individual as c–d), dorsal view, 12, sample 201; (g) NMW
2005.32G.173, pygidium, latex cast of external mould, dorsal view, 11, sample 205. (e) Acanthoparyphinae
gen. indet.: NMW 2005.32G.149, incomplete cranidium, latex cast of external mould, dorsal view, 4, sample
Figure 13 (a–h) Dindymene a. longicaudata Kielan, 1960: (a, d, f) NMW 2005.32G.213, articulated exoskel-
eton, latex cast of external mould, (a) dorsal view, 10; (d) lateral view, 11·5, (f) lateral oblique view, 11·5,
sample 201; (b) NMW 2005.32G.212, thorax and pygidium, internal mould, 9, sample 201; (c) NMW
2005.32G.214, thorax and pygidium, internal mould, dorsal view,5·5, sample 201; (e, g) NMW 2005.32G.180,
(e) exoskeleton with pygidium disarticulated, internal mould, dorsal view, 6·5; (g) xigena of same specimen:
detail of coarsely tuberculate ornament and transverse genal spine, 12, sample 201; (h) NMW 2005.32G.255,
pygidium with attached thoracic segments, internal mould, 8·5, sample 201. (i) Kimakaspis kovalevskyi gen. et
sp. nov.; NMW 2005.32G.272, paratype, latex cast of external mould of damaged articulated specimen, dorsal
view, 5·5, sample 201.
Dindymene a. longicaudata Kielan, 1960
(Fig. 13a–h)
Material. NMW 2005.32G.180 (Cl, 5·0, Cw, 10·7, Gw, 4·2,
Pl, 3·6, Pw, 4·8, Aw, 1·6), dorsal exoskeleton, internal mould;
NMW 2005.32G.213, dorsal exoskeleton, external mould (L,
7·8, W 5·5, Cl, 2·7, Cw, 6·0, Gw, 2·6, Pl, 2·0, Pw, 2·2, Aw, 1·0);
NMW 2005.32G.218–224, dorsal exoskeleton, external
moulds; NMW 2005.32G.212, 214, 256, pygidium and
attached thoracic segments, external and internal moulds.;
NMW 2005.32G.255, pygidium with attached thoracic seg-
ments, internal mould; all from sample 201. NMW
2005.32G.172, pygidium, external mould; sample 205.
Remarks. In addition to Dindymene brevicaudata
Kolobova, sample 201 contains articulated exoskeletons of
Dindymene with almost identical cephalic characters, but very
dierent pygidial morphology. This is characterised by a long,
narrow axis with 9 to 12 rings. Only two anterior axial rings
are complete and separated by deep, transverse furrows,
whereas succeeding axial rings merge axially and are separated
by transverse furrows that almost fade medially, adaxially, and
deepen towards the axial furrows. The pleural eld bears three
pairs of pleural ribs, but only the two anterior pairs are
connected to pleural ribs, with the second pair of pleural ribs
running almost parallel to the axial furrows. The third pair of
pleural ribs is very short and not connected to any axial ring,
and is terminated by a pair of short spines.
This distinct pygidial morphology matches in many details
that of Dindymene longicaudata Kielan, 1960 (e.g. pl. 30, g. 2)
from the upper Katian (Ashgill) Staurocephalus clavifrons
Biozone of the Holy Cross Mountains, Poland. The only
dierence is in the clear separation of the third pair of the
pleural ribs from the axis. The cephalic morphology also
resembles that of D. longicaudata, although the tubercles on
the glabella appear to be a little larger and sparser (but note
that we have only internal moulds available), and the short
genal spines are slightly larger and broader based.
There are a number of external moulds of isolated cranidia
of Dindymene in samples 201 (NMW 2005.32G.225–239, 257,
258), and 205 (NMW 2005.32G.174, 175), but because of the
almost identical cranidial morphology of D. brevicaudata and
D. a. longicaudata, both of which occur in these samples, we
are unable decide to which of these they belong.
Family Lichidae Hawle & Corda, 1847
Genus Amphilichas Raymond, 1905
Type species. Platymetopus lineatus Angelin, 1854, Upper
Ordovician, Katian Stage, Boda Limestone Formation,
Dalarna, Sweden.
Amphilichas a. nasutus Webby, 1974
(Fig. 6i)
Material. NMW 2005.32G.192, incomplete cranidium,
internal mould; sample 203b.
Remarks. This cranidium has an anterior median glabellar
lobe which expands strongly forwards, bounded by deep
longitudinal furrows, which converge gently anteriorly until
cranidial mid-length, then curve strongly and diverge anteri-
orly adaxially. The surface is ornamented with sparse coarse
and numerous ne tubercles, strongly resembling that of
Amphilichas nasutus Webby, 1974 (pl. 34, gs 3–8) from the
Upper Ordovician, lower Eastonian of New South Wales, but
the latter has a greater preponderance of large tubercles. The
cranidium from the Karagach Formation also resembles those
described by Chugaeva (1958) as Amphilichas karakanensis
var. disjunctus from the Upper Ordovician, Sandbian,
Anderken Formation of the Chu-Ili Range, but diers in
having a median glabellar lobe which is strongly expanded
anteriorly, an unevenly curved anterior margin, and coarse,
more strongly dierentiated, tuberculate ornament.
Family Odontopleuridae Burmeister, 1843
Subfamily Odontopleurinae Burmeister, 1843
Genus Diacanthaspis Whittington, 1941
Type species. Diacanthaspis cooperi Whittington, 1941,
Upper Ordovician, Martinsburg Formation, Virginia, USA.
Diacanthaspis? sp.
(Fig. 11b, c)
Material. NMW 2005.32G.267a–b, 268, 269, 270. Three
articulated specimens (counterpart internal and external
moulds, one internal mould, one external mould), one incom-
plete cranidium, all sample 201.
Description. Glabella tapers forwards, median lobe
parallel-sided, rounded anteriorly, delineated by wide and
shallow axial furrows. Glabella lateral lobes poorly dened but
two can be discriminated; occipital ring transverse, wider (tr.)
than median glabellar lobe. Occipital furrow narrow, well-
dened. Librigena with narrow border dened by shallow
border furrow; up to ten stout lateral marginal spines, decreas-
ing progressively in length anteriorly. Genal spine slender,
tapering gently, directed posterolaterally. Cranidium appar-
ently granulose, but too poorly preserved to distinguish size
range of granules. Palpebral lobe and eye not preserved.
Thorax with nine segments, posterior band of each pleura
extends into a long spine. Spines become progressively longer
and more strongly curved backwards towards posterior. Pair
of prominent tubercles on each axial ring and two to three on
each posterior band of pleurae.
Pygidium of two(?) segments, the rst with a well-dened
axial ring bearing two stout tubercles. First pair of pleurae
with a stout tubercle, and extend laterally into major spines,
which are directed posteriorly and are slightly shorter than
about half the width (tr.) of posteriormost thoracic spines.
Terminal pygidial segment less well-dened, tuberculate and
bearing one to two pairs of short, slender, posteriorly-directed
Remarks. These specimens are very small holaspides (sag-
ittal length 2·8–3·9 mm). Condent generic assignment is not
possible because of the poor preservation of the cranidia and
pygidia, but such morphological features as can be seen
suggest that they belong probably to Diacanthaspis.
4. Acknowledgements
MGP acknowledges support from the Golestan University,
and from the National Museum of Wales, Cardi. LMcC, RO
and LP acknowledge support from the National Museum of
Wales. We are grateful to the reviewers, Jonathan Adrain and
Greg Edgecombe, for their constructive comments, which
much improved the manuscript.
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