242 Artem Kouchinsky et al.

Laurentia 249
Baltica 249
Biology and affinities of Anabaritids 249
Systematic descriptions 251
Genus Anabarites Missarzhevsky, in Voronova & Missarzhevsky, 1969 253
Anabarites trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969 255
Anabarites ex gr. trisulcatus Missarzhevsky, in Voronova & Missarzhevsky,
1969, form 1 258
Anabarites ex gr. trisulcatus Missarzhevsky, in Voronova & Missarzhevsky,
1969, form 2 259
Anabarites ex gr. natellus (Val’kov & Sysoev, 1970) 261
Anabarites cf. signatus Mambetov, in Missarzhevsky & Mambetov, 1981 263
Anabarites latus (Val’kov & Sysoev, 1970) 265
Anabarites cf. latus (Val’kov & Sysoev, 1970) 267
Anabarites modestus Bokova, 1985 268
Anabarites tripartitus Missarzhevsky, in Rozanov et al., 1969 269
Anabarites ex gr. tripartitus Missarzhevsky, in Rozanov et al., 1969 270
Anabarites missarzhevskyi (Vasil’eva, 1986) 270
Anabarites ternarius Missarzhevsky, in Rozanov et al., 1969 271
Anabarites ex gr. ternarius Missarzhevsky, in Rozanov et al., 1969 272
Anabarites tristichus Missarzhevsky, in Rozanov et al., 1969 273
Anabarites compositus Missarzhevsky, in Rozanov et al., 1969 274
Anabarites valkovi (Bokova, in Bokova & Vasil’eva, 1990) 276
Anabarites hexasulcatus (Missarzhevsky, 1974) 278
Anabarites hariolus (Val’kov, 1987) 280
Anabarites korobovi (Missarzhevsky, in Rozanov & Missarzhevsky, 1966) 280
Anabarites rectus Vasil’eva, in Rudavskaya & Vasil’eva 1984 284
Genus CAMBROTUBULUS Missarzhevsky, in Rozanov et al., 1969 285
Cambrotubulus ex gr. decurvatus Missarzhevsky, in Rozanov et al., 1969 286
Genus SELINDEOCHREA Val’kov, 1982 286
Selindeochrea tecta Val’kov, 1982 288
Selindeochrea tricarinata (Missarzhevsky, in Voronova & Missarzhevsky,
1969) 288
Genus ACULEOCHREA Val’kov & Sysoev, 1970 289
Aculeochrea ornata Val’kov & Sysoev, 1970 290
Aculeochrea rugosa (Val’kov & Sysoev, 1970) 290
Genus MARIOCHREA Val’kov, 1982 291
Mariochrea sinuosa Val’kov, 1982 292
Acknowledgements 293
References 294

Introduction
Anabaritids, or angustiochreids, are a group of early skeletal
fossils known from the Precambrian–Cambrian transitional
and Lower Cambrian beds worldwide (Fig. 1). These fossils
occur as tubular skeletons (named tubes or thecae herein)
or internal moulds thereof. Their main feature is a dis-
tinct triradial symmetry expressed by lobes and grooves
and by other longitudinal morphological elements (Fig. 2).
Some forms with bilaterally symmetrical and circular cross-
sections, however, are included in the group, because of traces
of triradial symmetry or similarities in the wall microstruc-
ture and initial parts to those of triradially symmetrical ana-
baritids. Anabaritid tubes often change their transverse pro-
file during growth (e.g. Fig. 2G–H, J–K, P–Q, U–V), but their
initial part has a circular transverse profile, is goblet-like and
open at the apical end (see Figs 2X, 8A–B, L–M; 15C–E, F,
L–O; 38E–G, L–N, Q–R; 42A–D, F).
Rosén’s (1919) illustration of triradially symmetrical
sections in limestones of the Swedish Caledonides is the
first record of anabaritids, but neither formal description nor
name was given. The first illustration of anabaritids from
Siberia was published by Sysoev (1965; fig. 2) under the
name Hyolithellus sp. (recognised herein as Anabarites tri-
stichus Missarzhevsky, in Rozanov et al., 1969). The order
Angustiochreida was erected by A. K. Val’kov and V. A.
Sysoev in 1970 (the name first appeared in a conference ab-
stract by Val’kov & Sysoev 1969). However, the informal
name anabaritids, which is currently much more widely
 Systematics of Lower Cambrian fossil Anabaritids 243

Figure 1 Palaeogeographical map for the early Early Cambrian (after Steiner et al. 2007) showing crustal blocks, from which anabaritids are
reported.

used, originated from the family name Anabaritidae Mis-
sarzhevsky, 1974. The erection of the Angustiochreidae by
Val’kov & Sysoev (1970) with Anabarites as the type genus is
invalid according to International Commission on Zoological
Nomenclature (ICZN) rules (cf. Glaessner 1979; Bengtson
et al. 1990). V. V. Missarzhevsky initially described anabar-
itids in his Doctoral Candidate thesis in 1967 as a group of
serpulid worms, but his first publications on anabaritids did
not appear until 1969 (Rozanov et al. 1969; Voronova & Mis-
sarzhevsky 1969). Anabarites trisulcatus Missarzhevsky, in
Voronova & Missarzhevsky, 1969, was the first valid genus
and species assigned to this group. To date, 72 species (in-
cluding 10 nomina nuda) and 19 genera (including 2 nomina
nuda) of anabaritids have been formally described (see Ap-
pendix 1 in the Supplementary Material). Among them, 53
species (including 10 nomina nuda) and 18 genera (with one
nomen nudum) were originally reported from Siberia.
Appendix 1 lists all of the species and genera assigned
to anabaritids in the present study, with the originally given
name and occurrence, as well as available information on
other forms of anabaritids reported to co-occur within the
same samples. Appendix 2 contains a list of Siberian samples
examined by A. Kouchinsky and referred to in this
work, together with their localities and originally spe-
cified stratigraphical positions. The appendices are pub-
lished electronically as “Supplementary data” available on
Cambridge Journals Online: http://www.journals.cup.org/
abstract_S1477201909002715.
Anabaritids and the Lower
Cambrian stratigraphy of the
Siberian platform
Anabaritids are known from areas of shallow-water carbon-
ate sedimentation worldwide. However, ca. 70% of the di-
versity has been described from the Siberian Platform (see
Appendix 1 in Supplementary Material). Anabaritids are the
earliest metazoans with mineralised skeletons recognised on
the Siberian Platform, where they range into the Tommotian
Stage of the Lower Cambrian (Tables 1 & 2).
The Tommotian Stage biostratigraphy on the Siberian
Platform is based mainly on archaeocyaths (Rozanov &
Zhuravlev 1992; Rozanov et al. 1992). Ever since the
Tommotian Stage was first defined in the Lena–Aldan region

Table 1 Selected forms of anabaritids from the lower and upper parts of the Manykayan Stage of the Siberian Platform discussed in the
Systematic Descriptions.

Beds Forms
Upper Anabarites compositus Missarzhevsky, in Rozanov et al., 1969; A. hariolus (Val’kov, 1987); A. hexasulcatus (Missarzhevsky,
1974); A. kelleri Missarzhevsky, 1989; A. cf. korobovi (Missarzhevsky, in Rozanov & Missarzhevsky, 1966); A. cf. latus
(Val’kov & Sysoev, 1970); A.? licis (Missarzhevsky, in Rozanov et al., 1969); A. missarzhevskyi (Vasil’eva, 1986); A. modestus
Bokova, 1985; A. ternarius Missarzhevsky, in Rozanov et al., 1969; A. ex gr. ternarius; A. tripartitus Missarzhevsky, in Rozanov
et al., 1969; A. ex gr. tripartitus; A. tristichus Missarzhevsky, in Rozanov et al., 1969; A. valkovi (Bokova, 1990); A.? volutus
(Missarzhevsky, in Rozanov et al., 1969); Cambrotubulus decurvatus Missarzhevsky, in Rozanov et al., 1969; Cambrotubulus
ex gr. decurvatus; Mariochrea sinuosa Val’kov, 1982; Selindeochrea tecta Val’kov, 1982; S. tricarinata (Missarzhevsky, in
Rozanov et al., 1969).
Lower Aculeochrea ornata Val’kov & Sysoev, 1970; A. rugosa (Val’kov & Sysoev, 1970); Anabarites convexus (Val’kov & Sysoev, 1970);
A. latus (Val’kov & Sysoev, 1970); A. natellus (Val’kov & Sysoev, 1970); A. ex gr. natellus (Val’kov & Sysoev, 1970); A.
trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969; A. ex gr. trisulcatus; Cambrotubulus decurvatus
Missarzhevsky, in Rozanov et al., 1969; Cambrotubulus ex gr. decurvatus.
244 Artem Kouchinsky et al.

Figure 2 Cross-sections of internal moulds of some anabaritids that are included in the Systematic Descriptions, together with the key
morphological features of the thecae and internal moulds: A, Anabarited trisulcatus Missarzhevsky, in Rozanov., 1969; B–C, Anabarites latus
(Angustiochrea lata Val’kov & Sysoev, 1970), variation in transverse profiles; D, Anabarites ex gr. natellus (Lobiochrea natella Val’kov & Sysoev,
1970); E, Anabarites tripartitus Missarzhevsky, in Rozanov et al., 1969; F, Anabarites ex gr. tripartitus Missarzhevsky, in Rozanov et al., 1969;
G–H, Anabarites missarzhevskyi (Udzhaites missarzhevskyi Vasil’eva, 1986), at the apex (G) and at the aperture (H); I, Anabarites ternarius
Missarzhevsky, in Rozanov et al., 1969; J–K, Anabarites ex gr. ternarius Missarzhevsky, in Rozanov et al., 1969, at the apex (J) and aperture (K);
L–M, Anabarites valkovi (Jakutiochrea valkovi Bokova, in Bokova & Vasil’eva, 1990), at the apex (L) and aperture (M); N, Anabarites
hexasulcatus (Anabaritellus hexasulcatus Missarzhevsky, 1974), with first- and second-order grooves shown; O, Anabarites isisticus
Missarzhevsky, 1974; P–Q, Anabarites hariolus (Anabaritellus hariolus Val’kov, 1987), at the apex (P) and aperture (Q); R, Anabarits biplicatus
(Kotuites biplicatus Missarzhevsky, 1989); S, Anabarites? licis (Tiksitheca licis Missarzhevsky, in Rozanov et al., 1969); T, Selindeochrea tecta
Val’kov, 1982; U–V, Selindeochrea cf. tecta Val’kov, 1982, at the apex (U) and aperture (V); W, Selindeochrea tricarinata (Anabarites tricarinatus
Missarzhevsky, in Rozanov et al., 1969). X, Anabarites tristichus Missarzhevsky, in Rozanov et al., 1969, internal mould with notches and partly
preserved wall with flanges (septum, or tooth-like protrusion of the inner wall is shown at the aperture; smooth surface of the initial part is
shown at the apex); Y, Anabarites tricarinatus Missarzhevsky, in Rozanov et al., 1969, the upper part of a theca with prominent longitudinal
keels; Z, Mariochrea sinuosa Val’kov, 1982, apertural view of the internal mould showing a concentric invagination of the aperture. Scale bar:
A–H, J–W, Y, Z = 500 μm; I & X = 1 mm.

of the Siberian Platform (Rozanov et al. 1969), there has been siliferous beds. These sub-Tommotian beds are often re-
considerable discussion on the correlation of the lowermost ferred to as the Manykayan, or Nemakit–Daldynian Stage,
Tommotian Nochoroicyathus sunnaginicus Biozone across with the lower boundary at the base of the Anabarites tri-
the Platform and on the age of the sub-Tommotian fos- sulcatus Biozone and the upper boundary at the base of

Table 2 Forms of anabaritids reported from the Tommotian Stage of the Lena–Aldan region.

Biozone Forms
D. lenaicus Anabarites isiticus Misarzhevsky, 1974.
Dokidocyathus regularis Anabarites rectus Vasil’eva, in Rudavskaya & Vasil’eva, 1984; A. tristichus Missarzhevsky, in
Rozanov et al., 1969; Cambrotubulus decurvatus Missarzhevsky, in Rozanov., 1969; Tiksitheca
korobovi (Missarzhevsky, in Rozanov & Missarzhevsky, 1966); T. licis Missarzhevsky, 1969.
Nochoroicyathus sunnaginicus Anabarites tristichus; Cambrotubulus decurvatus; Tiksitheca korobovi; T. licis.

Source: after Sokolov & Zhuravleva 1983; Rozanov et al. 1992.

 Systematics of Lower Cambrian fossil Anabaritids 245

Figure 3 Map of the localities on the Siberian Platform referred to in the text. Samples derive from the localities indicated by stars with
numbers (see also Appendix 2 of Supplementary Material): 1, 1571/2; 2, M419/12, 92–1b/18, 1282, M423/13, 92–2/21, 92–2/25 and 92–2/26; 3,
M321/26, M321/31, M321/34, M314/5–10, M314/7 and M314/12; 4, 1404, 96-4/4, 96-4/4(2), 96–5a/34.5 and 96-5a/34.75; 5, 1326, B-483,
B-484, B-489; 6, 806/3-4; 7, 769; 8, B-335, B-335b, B-337, B-339, 907, M71-2/62, M71-2/66 and M71-2/69, 882 and 1487; 9, 11/16.05 and
11/16.1; 10, 106 and 106b; 11, M302/1-2; 12, A-380k; 13, 1733; 14, 355a and 369; 15, 316/4.

the Tommotian Stage (Missarzhevsky 1989; Rozanov et al.
1992, 2008; Khomentovsky & Karlova 1993, 2002; Kho-
mentovsky et al. 1998; Kaufman et al. 1996; Figs 3 & 4).
Despite priority issues favouring the Nemakit–Daldynian
Stage (Khomentovsky & Karlova 2002), the term Manykayan
Stage is also widely used in English. Since neither is formally
accepted, we prefer to use the term Manykayan Stage herein.
The latter appears to be roughly equivalent to the recently
defined Fortunian Stage (Landing et al. 2007), provided both
of them have currently provisional upper boundaries.
Biostratigraphical correlation of these Precambrian–
Cambrian transitional strata across the Platform is contro-
versial for lack of archaeocyaths and is based on the earliest
skeletal fossils, such as anabaritids. The abundant anabaritids
from these beds are especially useful for an analysis of their
taxonomic diversity and morphological variability. Taking
into account the existing taxonomic and biostratigraphical
ambiguities, we avoided using information on the distribu-
tion of anabaritids from published lists of fossils without
illustrations from the Siberian Platform outside the Lena–
Aldan region. Some of the samples with anabaritids lis-
ted in Appendix 2 (Supplementary Material) are shown in
Fig. 4, their stratigraphical ages constrained by carbon iso-
tope chemostratigraphy.
The first skeletal tubular remains, referred to as Cam-
brotubulus sp. and Anabarites sp., have been found in the
upper Turkut Formation, 30m below the boundary with the
Kessyusa Formation, at the Khorbusuonka River, northern
Siberian Platform (Karlova & Vodanyuk 1985; Karlova 1987;
Khomentovsky & Karlova 1993; Khomentovsky et al. 1998).
The local Precambrian–Cambrian transition is approximated
by the sequence boundary between the Turkut and Kessy-
usa formations. The negative carbon isotopic excursion N,
which spans the Precambrian–Cambrian boundary around
the world (Brasier et al. 2000; Amthor et al. 2003) with a
negative δ 13 C spike lower than −4‰ (Brasier et al. 1994b;
Knoll et al. 1995; Kaufman et al. 1996), is reported from
the uppermost unit of the Turkut Formation, although the
minimum value is only −2.6 ‰ (Knoll et al. 1995).
The upper boundary of the Anabarites trisulcatus
Biozone is not distinct in practice, because it is often situ-
ated in dolomitic and terrigenous rocks unsuitable for the
preservation and extraction of skeletal fossils. The base of
the overlying Purella Biozone (Fig. 4) is marked by the
246 Artem Kouchinsky et al.

Figure 4 δ 13 C profiles through the Manykayan (Nemakit–Daldynian)–Tommotian beds, with an indication of the stratigraphical position of
some of the samples discussed in the text. Reference curve, lower part, from the Aldan River (Lena–Aldan region), adapted from Brasier et al.
1994b. Selinde section (after Kouchinsky et al. 2005), with biostratigraphical scale through the lower Pestrotsvet Formation after Val’kov, 1982.
Bol’shaya Kuonamka section (Kouchinsky et al. 2001), lower part, with biostratigraphical scale after Val’kov, 1975. Kotuj section is adapted from
Kaufman et al. 1996. Profiles are not to the same vertical scale, shown for correlation only.

appearance of scaly shells of Purella cristata Missarzhevsky,
1974, a possible senior synonym of P. antiqua (Abaimova,
1976), and by an increasing diversity of other bilaterally
symmetrical skeletal fossils (Missarzhevsky 1989; Kho-
mentovsky & Karlova 1993, 2002; Khomentovsky et al.
1998). The upper limit of the Purella Biozone is often re-
ferred to as the base of the Tommotian Stage in the Lena–
Aldan region (Khomentovsky & Karlova 1993, 2002; Kho-
mentovsky et al. 1998; Brasier et al. 1994a, b; Zhuravlev
1998). Therefore, where the fossil record permits, it is pos-
sible to distinguish lower and upper Precambrian–Cambrian
transitional beds of pre-Tommotian age on the Siberian Plat-
form (Fig. 4).
On the basis of biostratigraphy (e.g. Val’kov 1982,
1987; Missarzhevsky 1983, 1989) and chemostratigraphy
(Brasier et al. 1994a, b; Knoll et al. 1995; Kaufman et
al. 1996; Kouchinsky et al. 2001, 2005) several biozones
have been suggested (Fig. 4), unrecognised in the Lena–
Aldan region, between the Purella and Nochoroicyathus
sunnaginicus Biozones. Since biostratigraphical correlation
at this level across the Siberian Platform is controversial (e.g.
Khomentovsky & Karlova 1993, 2002; Rozanov et al. 1969,
1992, 2008), the assignement of this interval can vary from
the upper Purella Biozone of the Manykayan Stage to the
Dokidocyathus regularis Biozone of the Tommotian Stage.
Detailed carbon isotopic profiling from these strata,
where available, show, however, isotopic signatures dif-
ferent from those of the Nochoroicyathus sunnaginicus to
D. regularis biozones in the Lena–Aldan region (Brasier
et al. 1994a, b). The lower boundary of the Tommotian Stage
coincides either with the base of the Pestrotsvet Formation
(Khomentovsky & Karlova 1993, 2002) or – in the stra-
totype region – is situated in the uppermost Ust’–Yudoma
Formation (Rozanov et al. 1992; Zhuravlev 1998). There is
an interval with several positive peaks, preceding a sharp
drop at the lower boundary of the Tommotian Stage in the
Medvezhin (Knoll et al. 1995; Kaufman et al. 1996) and
Emyaksin formations (Kouchinsky et al. 2001) of the west-
ern and eastern flanks, respectively, of the Anabar uplift
on the northern Siberian Platform, as well as in the lower
Pestrotsvet Formation in the Selinde section (Kouchinsky
et al. 2005) of the southeastern Siberian Platform, and within
the upper Sukharikha Formation of the Sukharikha River
section (Kouchinsky et al. 2007). Chemostratigraphical and
biostratigraphical correlation suggests that beds with this
succession of positive peaks were deposited earlier than
the basal Pestrotsvet Formation in the Lena–Aldan region
(Fig. 4).
Selected forms of anabaritids from beds attributed
herein to the Manykayan Stage of the Siberian Platform and
discussed in the Systematic Descriptions section, below, are
listed in Table 1. Few species are known from the Tommo-
tian Stage in the Lena–Aldan region, where the age is directly
supported by archaeocyaths. These are shown in Table 2 (data
 Systematics of Lower Cambrian fossil Anabaritids 247

Table 3 Forms of anabaritids reported from outside the Siberian Platform (see palaeogeography in Fig. 1).

China ∗ Anabaritellus cylindriculus Qian & Ding, 1988 in Ding & Qian, 1988; Anabaritellus cf. hexasulcatus Missarzhevsky; ∗ A.
xishuiensis Qian, 1977; ∗ Anabarites acuminatus Chen & Xiong, in Xing et al., 1984; ∗ A. gracilis Chen, 1984; ∗ A.
longissimoides Chen & Xiong, in Xing et al., 1984; ∗ A. obliquasulcatus Qian, 1978; ∗ A. primitivus Qian & Jiang, 1980 in
Luo et al., 1982; ∗ A. rotundus Qian, 1977; ∗ A. sulcoconvex Qian, 1978; ∗ A. tenuistriatus Qian, 1989; A. ternarius
Missarzhevsky, in Rozanov et al., 1969; A. tristichus Missarzhevsky, in Rozanov et al., 1969; A. trisulcatus
Missarzhevsky, in Voronova & Missarzhevsky, 1969; ∗ A. undulatus Qian, 1978; Cambrotubulus decurvatus
Missarzhevsky, in Rozanov et al., 1969; ∗ Tiksitheca fangxianensis Duan, 1983; ∗ T. huangshandongensis Qian
et al., 1979; ∗ T. irregularis Qian & Ding, in Ding & Qian, 1988; T. korobovi (Missarzhevsky, in Rozanov &
Missarzhevsky, 1966); ∗ T. minuta Yue, in Xing et al., 1984. (∗ , reported from China only)
Western Gondwana Anabarites tristichus; A. trisulcatus; Cambrotubulus corniformis Elicki, 1994; C. decurvatus; Tiksitheca licis
Missarzhevsky, in Rozanov et al., 1969.
Laurentia Anabarites trisulcatus.
Kazakhstan Anabarites signatus Mambetov, in Missarzhevsky & Mambetov, 1981; Tiksitheca licis.
Western Mongolia Anabaritellus hexasulcatus Missarzhevsky, 1974; Angustiochrea magna Bokova, 1992 [nomen nudum]; Anabarites
tribaculatus Bokova, 1992 [nomen nudum]; A. tripartitus Missarzhevsky, in Rozanov et al., 1969; A. trisulcatus;
Jakutiochrea portentosa Bokova & Val’kov, in Val’kov, 1987; Cambrotubulus decurvatus; Tiksitheca licis; Udzhaites
cf. missarzhevskyi Vasil’eva, 1986.
Eastern Gondwana Anabarites sexalox Conway Morris & Bengtson, in Bengtson et al., 1990; A. trymatus Conway Morris & Bengtson, in
(Australia) Bengtson et al., 1990.
Baltica Anabarites cf. sexalox.
Avalonia ?Anabarites tricarinatus; A. tripartitus; Anabarites trisulcatus; Tiksitheca korobovi; T. licis.

from Sokolov & Zhuravleva 1983 and Rozanov et al. 1992).
No fossils assigned to anabaritids have been reported from
deposits younger than the Tommotian Stage on the Siberian
Platform. The age of Anabarites tristichus (Hyolithellus sp.
in Sysoev (1965; fig. 2)) from ‘the Atdabanian Horizon’ in
a locality on the left bank of the Kotuj River, 6 km down-
stream of the mouth of the Kotujkan River, western flanks
of the Anabar Uplift is questionable, for it is not clear from
which level of the outcrop the specimen derives.
archaeocyaths therein, the entire formation may be regarded
as pre-Tommotian (cf. Brasier et al. 1996) and corresponding
to the Fortunian Stage (Landing et al. 2007). However, some
workers propose a Tommotian–Atdabanian age, based on the
controversial biostratigraphical correlation with the Siberian
Platform (Esakova & Zhegallo 1996; Khomentovsky &
Gibsher 1996). Cambrotubulus decurvatus and Anabarites
trisulcatus have also been reported from the Khairkhan Form-
ation, where archaeocyaths probably indicate a Botomian age
(Esakova & Zhegallo 1996).

Anabaritids from outside the Kazakhstan

Siberian platform
Outside the Siberian Platform (Fig. 1) anabaritids are re-
ported to range into the Atdabanian–Botoman equivalent
strata. It is not clear whether the lack of records from post-
Tommotian deposits on the Siberian Platform is due to pre-
servation, ecology, or palaeobiogeography.
Anabarites signatus Mambetov, in Missarzhevsky & Mam-
betov, 1981, and Tiksitheca licis are known from the lower
Chulaktau Formation of the Lesser Karatau Range in Kaza-
khstan (Missarzhevsky & Mambetov 1981; Table 3). Their
age is indicated as Tommotian (Missarzhevsky & Mambetov
1981; Missarzhevsky 1989) or Atdabanian (Landing 1988,
p. 690).

Western Mongolia
The earliest anabaritids, A. trisulcatus and C. decurvatus,
occur in the upper member of the Tsagaan Oloom Forma-
tion (Brasier et al. 1996; Esakova & Zhegallo 1996; Kho-
mentovsky & Gibsher 1996), whereas Anabarites tripartitus
has been reported from the uppermost part of this member
(Khomentovsky & Gibsher 1996). This upper member of
the Tsagaan Oloom Formation is correlated with the Ana-
barites trisulcatus Biozone (Brasier et al. 1996; Esakova &
Zhegallo 1996; Khomentovsky & Gibsher 1996). These and
other forms of anabaritids are found in the overlying Bayan
Gol Formation (Table 3; Brasier et al. 1996; Esakova &
Zhegallo 1996; Khomentovsky & Gibsher 1996). According
to the position of the high positive carbon isotope values in
the upper part of the Bayan Gol Formation and the absence of
China
Most species of Anabarites described from China (Table 3)
have been regarded as synonyms of A. trisulcatus (Conway
Morris & Chen 1989; Qian & Bengtson 1989: 124; Steiner
et al. 2004). These forms occur mostly in the Lower Meishu-
cunian Stage (Anabarites trisulcatus–Protohertzina anabar-
ica Assemblage Zone or equivalents) of the Yangtze Plat-
form. The Lower–Middle Meishucunian Stage is broadly
correlated with the Manykayan Stage of Siberia (Brasier
1989; Qian 1989, 1999; Qian & Bengtson 1989; Bengtson
& Yue, 1992; Peng & Babcock 2001; Zhu et al. 2001; Qian
et al. 2001, 2002; Steiner et al. 2003, 2004, 2007; Chen
& Peng 2005) and Fortunian Stage (Landing et al. 2007).
Anabarites cf. trisulcatus are also indicated from the Middle
248 Artem Kouchinsky et al.
and Upper Meishucunian Stages of China (Li et al. 2007).
Anabarites cf. trisulcatus extend into younger strata in the
Jiangshan region of the Zhejiang Province and Aksu–Wushi
region in the Tarim area of the Xinjiang Province correlated
with the Qiongzhusian Stage (Atdabanian equivlent) (Xiao
1989; Xiao & Duan 1992; Qian et al. 2001), and in the
Chaohu region of the Anhui Province correlated with the
Qiongzhusian and Canglangpuan Stages (Atdabanian and
Botomian equivalents respectively) (Feng 2005).
Forms probably synonymous to A. trisulcatus described
from the Lower Meishucunian strata include: A. obliquasul-
catus Qian, 1978, A. rotundum Qian, 1977, and A. sul-
coconvex Qian, 1978 from the Kuanchuangpu Formation of
the Shaanxi Province; A. undulatus Qian, 1978 from the
Zhongyicun Member, Zhujiaqing Formation of Yunnan; A.
acuminatus Chen & Xiong, 1984 (in Xing et al., 1984) and
A. longissimoides Chen & Xiong, 1984 (in Xing et al., 1984)
from the Tianzhushan Member, Dengying Formation of the
Hubei Province, and A. tenuistriatus Qian, 1989 from the
Huangshadong Member, Tongying Formation of Hubei (see
Appendix 1 of Supplementary Material).
Anabarites tristichus Missarzhevsky occurs in the
Lower Meishucunian Stage and has been found in the basal
Gezhongwu Formation of the Guizhou Province (Peng &
Babcock 2001). Anabarites gracilis Chen, 1984 of probably
the same age is described from West Hubei and synonymised
herein with A. tristichus.
Rare fragments of A. tripartitus Missarzhevsky or A.
trymatus Conway Morris & Bengtson, in Bengtson et al.,
1990 with depressions on the three lobes were identified
(but not illustrated) by Steiner et al. (2004) from the Lower
Meishucunian Stage lower–middle Kuanchuanpu Formation
in the Zhangjiagou section, Hexi area of the south Shaanxi
Province. These forms can be also compared with A. valkovi
(Bokova, in Bokova & Vasil’eva, 1990: see Systematic De-
scriptions, below).
Anabarites cf. hexasulcatus (Missarzhevsky) is reported
from the Lower Meishucunian Stage of the Yangtze Platform
(Brasier 1989; Qian 1999), as well as the morphologically
similar Anabaritellus xishuiensis Qian, 1977 described from
the Yuhucun Formation of the Guizhou Province and Ana-
baritellus cylindriculus Qian & Ding, in Ding & Qian, 1988
from the lower Yangjiaping Formation of the Hunan Province
(Appendix 1).
Lower Meishucunian Anabarites ternarius Missar-
zhevsky illustrated by Xing et al. 1984 (pl. 14, fig.16) occurs
in Bed 34 of the Maidiping section, Emei, Sichuan Province
(fig. 7–3 therein) and is also indicated by Steiner et al. 2007
(supplement) from the lower–middle Kuanchuanpu Forma-
tion of Southern Shaanxi.
Cambrotubulus ex gr. decurvatus and Tiksitheca spp. are
reported from the Meishucunian and Qiongzhusian Stages
of the Yangtze Platform (Brasier 1989; Qian 1999; Li
et al. 2007). By comparison with Anabarites, identification
of these fossils is less clear and comparison with the Siberian
type material is more problematic, because of a defficiency
of distict features in them and unsatisfactory preservation.
Along with them, tubular shells attributed to Conotheca
Missarzhevsky are commonly reported from China and else-
where by various authors (e.g. Qian & Bengtson 1989). Their
affinity with anabaritids is usually implied. Indeed, Cambro-
tubulus and Tiksitheca are often difficult to distinguish from
Conotheca, when their apical parts are not preserved (e.g.
Qian & Bengtson 1989: fig. 85; Steiner et al. 2004: fig. 3–
17) (see Remarks in Systematic Descriptions, below, for the
genus Cambrotubulus). It may, therefore, be that Cambrotu-
bulus and Tiksitheca are more common in China and else-
where, but this question needs investigation beyond the scope
of this paper.
Several forms have been described as Tiksitheca Missar-
zhevsky from the Yangtze Platform (Appendix 1). These are
Lower Meishucunian T. irregularis Qian & Ding, in Ding
& Qian, 1988, from the lower Yangjiaping Formation of
Hunan; T. huangshandongensis Qian et al. 1979, from the
Huangshandong Formation of Hubei; and T. minuta Yue, in
Xing et al., 1984, from the upper Dengying Formation of
Shaanxi. Tiksitheca fangxianensis Duan, 1983 from the Xi-
haoping Member, uppermost Dengying Formation of Hubei
is attributed to the Qiongzhusian Stage (Steiner et al. 2003).
Eastern Gondwana
Anabarites trymatus Conway Morris & Bengtson, in Bengt-
son et al., 1990, and Anabarites sexalox Conway Morris
& Bengtson, in Bengtson et al., 1990, occur in the South
Australian Stansbury Basin, Yorke Peninsula (upper Kulpara
Limestone and Parara Limestone formations) and in the Ar-
rowie Basin, Flinders Ranges (Ajax Limestone Formation) in
beds correlated with the upper Atdabanian–Botomian Stages
(Bengtson et al. 1990; Gravestock et al. 2001; Table 3).
Western Gondwana
During the Cambrian Period the western part of the Gond-
wana supercontinent consisted of a complex arrangement of
crustal blocks (Fig. 1). Anabaritids have been reported from
a number of areas in Europe and the Middle East (Table 3).
Anabaritids (although non-identified) are shown ranging
from the middle into the upper part of the Rio Huso Group
in the Valdelacasa Anticline of Central Spain correlated with
the pre-Atdabanian beds of Siberia (Vidal et al. 1999: figs 1
& 2). Cambrotubulus cf. decurvatus has been reported from
the Heraultia Limestone of the Atdabanian–Botomian equi-
valent of the Massif Centrale in France (Kerber 1988: 136,
pl. 1, fig. 26).
Cambrotubulus cf. decurvatus and Tiksitheca licis are
reported from the Upper Torgau Member of the Zwethau
Formation of the Torgau–Doberlung Syncline in Germany
(Elicki 1994) and correlated with the lower Atdabanian
(middle Issendalenian Stage: Elicki 2005). Cambrotubulus
decurvatus, C. corniformis Elicki, 1994 and Tiksitheca licis
are reported from the Upper Ludwigsdorf Member of the
Charlottenhof Formation of the Gorlitz Syncline in Germany
(Elicki 1994) and assigned to the upper Atdabanian (higher
Banian Stage: Elicki 2005).
From the Lesser Himalaya of India, Anabarites tri-
sulcatus and Tiksitheca licis have been reported from the
basal unit of the lower Tal Formation (Kumar et al. 1987)
given a Precambrian–Cambrian boundary age (Brasier &
Singh 1987; Hughes et al. 2005). In the Elburz Mountains
of Iran Anabarites trisulcatus, Cambrotubulus decurvatus
and Tiksitheca licis are found in the Middle Dolomite Mem-
ber of the Soltanieh Formation (Brasier 1989; Hamdi 1995)
correlated with the Manykayan Stage (Hamdi 1995). These
forms co-occur with A. cf. tristichus in the lower part of
the overlying Upper Shale Member (Brasier 1989; Hamdi
 Systematics of Lower Cambrian fossil Anabaritids
et al. 1989) correlated by Hamdi (1995) with the Tommotian
Stage.
Avalonia
Southeastern Newfoundland, southern New Brunswick,
Nova Scotia, eastern Massachusetts and Rhode Island in
North America, as well as England and Wales belonged to
an independent small continent, Avalonia, during the Cam-
brian (Fig. 1). Forms assigned to Tiksitheca korobovi, T. li-
cis, Anabarites trisulcatus and A. tripartitus were found in
southeastern Newfoundland and synonymised by Landing
(1988) with Tiksitheca korobovi (Table 3). They occur from
the Watsonella crossbyi Biozone to the middle part of the
Camenella baltica Biozone (Landing et al. 1989), with the
latter probably ranging into the lower Atdabanian (Brasier
et al. 1992; Geyer & Shergold 2000). The entire interval is
contained within the Terreneuvian Series beginning with the
Fortunian Stage (Landing et al. 2007). Anabarites cf. tri-
partitus (Tiksitheca korobovi sensu Landing 1988) occurs in
the Sunnaginia imbricata Biozone of the Weymouth Forma-
tion of eastern Massachusetts (Landing 1988; Landing et al.
1989). Anabaritellus tricarinatus (synonymised by Landing
& Murphy 1991 with Anabaritellus hexasulcatus, Anabar-
ites isiticus, Anabarites ternarius, Coleoloides trigeminatus
and Selindeochrea tecta) is reported from mainland Nova
Scotia and southeastern Newfoundland (Landing & Murphy
1991), but the affinity of this fossil with anabaritids is ques-
tionable (see Remarks in Systematic Descriptions, below, for
the genus Selindeochrea).
Problematic fossils described by Hinz (1987) from the
Strenuella and Protolenus Limestone (upper Lower Cam-
brian Protolenid–Strenuellid Biozone) of Comley, England
as Anabarites compositus Missarzhevsky bear little resemb-
lance to this species, but rather to Selindeochrea tricarinata
(Missarzhevsky). The material includes 33 specimens (ac-
cording to Hinz 1987: 72), but their affinity with anabaritids
is questionable.
Laurentia
Laurentian localities of anabaritids are limited to the Wer-
necke Mountains of the Canadian Northwestern Territory,
where Anabarites trisulcatus occurs in the Ingta Forma-
tion (former basal Vampire Formation in Nowlan et al.
1985; Table 3) containing fossils of the lowermost Cam-
brian Anabarites-Protohertzina Biozone (Pyle et al. 2006)
that can be attributed to the lower Fortunian Stage (Landing
et al. 2007).
Baltica
Anabaritids are known in thin sections from the cent-
ral Scandinavian Caledonides. They derive from two or
three small localities in the upper part of the Gärdsjön
Formation, attributed to the Holmia kjerulfi-group Zone,
a possible equivalent of the upper Atdabanian Stage
(Bergström & Ahlberg 1981; Ahlberg 1984). Recent in-
vestigation of these beds has not changed their age de-
termination, nor did they provide more variety, than those
originally figured (Rosén 1919). Judging from the cross-
sections, these forms are similar to Anabarites sexalox
(Fig. 5; Anabarites cf. sexalox in Table 3).
249
Biology and affinities of
Anabaritids
The biological affinity and phylogeny of anabaritids is un-
certain. The uncertainty is mainly due to lack of data on
anabaritid soft parts, paucity and variability of morpholo-
gical characters, as well as a lack of reliable evidence on the
ancestors and descendents of anabaritids in the palaeontolo-
gical record and modern biota.
Their possible affinity with tubicolous polychaetes was
initially discussed by Missarzhevsky (Voronova & Missar-
zhevsky 1969; Rozanov et al. 1969) on the basis of the su-
perficial morphological similarity to sedentary polychaetes
of the family Serpulidae. Although some serpulids carry three
longitudinal ribs on the outer surface of their tubes, the in-
ner wall in serpulids is cylindrical and smooth, reflecting
the ability of the animals to move freely in their tubes. This
is in contrast to most anabaritids, in which the inner sur-
face of the wall is convoluted by lobes and grooves (Fig. 2),
usually more prominent than corresponding morphological
elements of the outer wall. Furthermore, some forms, such as
Anabarites tristichus (see Figs 26–28), have serially repeated
internal tooth-like projections that would have prevented free
movement of the body along the tube.
The thecae of anabaritids grew by periodical accre-
tion of mineralised material at their aperture and demon-
strate a characteristic microstructure of the fibrous wall
(Kouchinsky & Bengtson 2002). Cambrotubulus decurvatus,
Tiksitheca licis and Anabarites tristichus differ from each
other only in the prevailing angles of inclination of the mi-
crostructural fibres. In Cambrotubulus (see Fig. 43) the wall
is built of fibre bundles and has a smooth outer surface.
In Tiksitheca (see Fig. 39E) and Anabarites tristichus (see
Fig. 26A) it produces flanges. The microstructures of
Selindeochrea tecta (see Figs 46G & 47A–E) and S. tri-
carinata (see Fig. 48A–E) are also very similar.
A chevron-like pattern of the growth lamellae has
been observed in longitudinal sections through the walls
of Tiksitheca licis and Anabarites tristichus (Kouchinsky &
Bengtson 2002). This pattern resembles that which was ex-
clusively known from serpulids, the earliest indubitable re-
cord of which is Mesozoic (Weedon 1994; Vinn 2008). The
chevron-like pattern and evidence for accretionary growth
(transverse flanges, growth lamellae) suggests that the wall
was formed by building on the leading edge by a glandu-
lar epithelium localised at the tube aperture (Kouchinsky &
Bengtson 2002). This mechanism was suggested previously
on the basis of the presence of external longitudinal keels
in S. tricarinata (Bengtson et al. 1990). Nevertheless, the
presence of internal tooth-like structures in anabaritid tubes
suggests a different relationship between tube and soft body
from that in the Serpulida.
According to Missarzhevsky (1974), the promin-
ent lobes, internal tooth-like structures and longitudinal
twisting of some anabaritid tubes suggest a fixed position
of the organism inside the tube, or at least that the soft parts
adjacent to the interior wall were firmly attached. Hence, he
concluded that the tube reflected a tri-fold radial symmetry of
an organism with an uncertain affinity to cnidarians. Twisting
was interpreted as a growth reaction of the sedentary organ-
isms to periodical (seasonal?) changes in the environment
(Missarzhevsky 1974).
250 Artem Kouchinsky et al.

Figure 5 Anabarites cf. sexalox Conway Morris & Bengtson, in Bengtson et al., 1990. A–D, cross-sections of hexasulcate tubes from the
Swedish Caledonides (collected by Dr Christian Skovsted). Scale bars = 200 μm.

The cnidarian affinity has been accepted by most sub-
sequent authors. Exceptions are Glaessner (1976), who main-
tained the originally published idea of a polychate relation-
ship, and Abaimova (1978), who questioned the cnidarian
model on the grounds of the mode of growth of the tubes. Al-
though she found that anabaritids are morphologically most
similar to Palaeozoic conulariids, which are usually inter-
preted as cnidarians (a recent analysis suggests that they are
the sister group of extant coronate scyphozoans: Van Iten
et al. (2006)), she noted the absence of attachment disks and
septa in the initial part of anabaritids, unlike conulariids, as
well as the difference in the anabaritid three-fold symmetry
from the four-fold symmetry common in the conulariids. No
opercula that could protect the animal inside the anabaritid
tube were found, nor any attachment structures, although the
initial parts may occasionally show a tapered open apex (see
Figs 2X & 15).
Important to Abaimova’s (1978) discussion of the affin-
ity is her reconstruction of the anabaritid theca as an internal,
supporting skeleton, especially in forms with prominent
longitudinal keels, such as Selindeochrea tricarinata. She
argued (Abaimova 1978: 82) that these could only have been
formed from the outside, although accretion on the aper-
tural edge of the wall would be sufficient to produce such
structures (Kouchinsky & Bengtson 2002). Those transversal
flanges and longitudinal keels may be interpreted as an ad-
aptation to a sedentary mode of life, with the tubes em-
bedded in soft sediment (Qian & Bengtson 1989) or in a
microbial mat (Chen & Peng 2005). Keels would have, in
addition, prevented the tube from rolling (Qian & Bengtson
 Systematics of Lower Cambrian fossil Anabaritids
1989). Although the three- or six-fold symmetrical longit-
udinal structures of the inner wall most probably reflect
symmetrical folding of the secretory epithelium, radial sym-
metry in other tissues remains speculative (Bengtson et al.
1990).
Val’kov (1982) interpreted the anabaritids as closely
related to scyphozoans. The main basis for this interpreta-
tion was the morphological similarities to polyps of mod-
ern scyphozoans and to the tests of conulariids, as well
as the serial constrictions in some anabaritid tubes (cf.
Figs 14J–L, 50 & 51 herein), which were taken as evidence
for strobilation.
Indirect support for a cnidarian affinity was provided by
the report of globular phosphatised embryos with tetrameral
symmetry co-occurring with a diverse complex of anabarit-
ids in the lower Manykayan Stage of Siberia (Kouchinsky
et al. 1999). The morphology of these globules is consistent
with that of a cnidarian actinula larva with eight incipient
tentacles. Kouchinsky et al. (1999) speculated that the ana-
baritids might represent a triradial cnidarian with tetraradial
symmetry in the embryo, but it cannot be excluded that the
embryos belong to a different adult organism not fossilised
at the site.
More recently, Chen & Peng (2005) have argued that
the aragonitic skeleton and tendency towards hexaradial sym-
metry in some anabaritids, i.e. the subdivided main lobes in
forms such as A. hexasulcatus (see Fig. 33), A. isiticus (see
Fig. 35), A. hariolus (see Fig. 36) and A. sexalox (see Fig. 34)
suggest that they represent a stem lineage of hexacorals. An
aragonitic composition, however, is the rule among skeletal
fossils appearing in the latest Neoproterozoic and earliest
Cambrian and probably reflects the sea chemistry at the time
of origin of the skeleton (Bengtson 2005; Porter 2007). The
origin of hexacorals remains an unresolved question (Stan-
ley 2003), but in the absence of more compelling synapo-
morphies, the occasional presence of six lobes in triradially
symmetrical anabaritid tubes is not likely to be homologous
with the hexaradial/biradial symmetry of a hexacoral basal
skeleton.
Fedonkin (1985, 1986) drew attention to the presence of
Vendian disk-like fossils (e.g. Tribrachidium and Albumares)
with triradial symmetry, implying their possible phylogenetic
relationship with the anabaritids. Again, lacking other con-
vincing synapomorphies, the number of radials alone is quite
likely to be an effect of convergence in radially symmetrical
organisms.
Thus, whereas the affinity of anabaritids has been
looked for mostly within annelids (notably serpulids) or cnid-
arians (possibly close to the Scyphozoa), the available evid-
ence is currently inconclusive. A further search for soft tis-
sues in suitable lithologies (anabaritids commonly co-occur
with phosphatised metazoan embryos) probably provides the
best hope of resolving this question.
Systematic descriptions
The scarcity of morphological criteria and the uncertain af-
finity of the group make the taxonomy of anabaritids rather
unstable. We evaluate here taxonomically important features
in order to find a balance between the taxonomical oversplit-
251
ting of anabaritids, which has hitherto resulted in 72 species
and 19 genus names, and unsubstantiated lumping.
Missarzhevsky (Missarzhevsky 1967; Rozanov et al.
1969; Voronova & Missarzhevsky 1969) stressed the import-
ance of the triradial symmetry of Anabarites tubes, as well
as the transformation of their cross-section during ontogeny.
Val’kov & Sysoev (1969, 1970) distinguished the order An-
gustiochreida Val’kov & Sysoev, 1970 with three families
(Table 4). They drew attention to the unusual type of radial
symmetry and the variety of cross-sections and sculptures.
Eventually, Missarzhevsky (1974) recognised a single family,
Anabaritidae Missarzhevsky, 1974 (Table 4), of problematic
affinity within the Cnidaria.
Val’kov (1982) attributed the order Angustiochreida
Val’kov & Sysoev, 1970, to a new subclass Angustiomedu-
sae within the class Scyphozoa (Phylum Cnidaria), and re-
cognised four families (Table 4). Later, however, Val’kov
(1987) placed the order within the class Trilobozoa (Phylum
Cnidaria?). This later placement was inspired by Fedonkin’s
(1985, 1986) establishment of the Trilobozoa to encompass
tri-fold symmetrical Vendian non-skeletal organisms and,
possibly, anabaritids.
Conway Morris & Chen (1989) published their obser-
vations on the variability of anabaritids from South China
and allied them to a single genus Anabarites Missarzhevsky,
in Rozanov et al., 1969, of the family Anabaritidae Missar-
zhevsky, 1974. This view was maintained by Qian & Bengt-
son (1989) and Bengtson et al. (1990), who admitted a high
level of intraspecific variability in anabaritids.
Missarzhevsky (1989) postulated a trend of increasing
hardness in the evolution of anabaritid tubes by means of
more intense folding of the walls, and subdivided the family
Anabaritidae into four subfamiles (Table 4).
Esakova (1989), in her Doctoral Candidate thesis ab-
stract, considered only triradially symmetrical forms as ana-
baritids and described five families (Table 4). With modi-
fication, these results were formally published by Esakova
& Zhegallo (1996), with the main family-level characters
represented by cross-sectional shapes and external morpho-
logical elements of the tubes (table 8 therein).
Bokova (1990, 1992) classified anabaritids on the basis
of a hierarchical system of morphological characters and
described two suborders and five families within the order
Angustiochreida (Table 4). Suborders were distinguished by
the type of radial symmetry (infinite or three-fold). Famil-
ies were based on the general morphology and sculpture of
the tubes. Genera were resolved by the outline of the cross-
sections, elements of the sculpture and their combinations,
expansion rate and wall thickness. Curvature and size, de-
tails of the sculptural elements and their location, and details
of the cross-section were regarded as species-level features.
The Bokova 1990 paper is the published abstract of Bokova’s
Doctoral Candidate thesis. It contains new names, but no de-
scriptions. Bokova, 1992 is her actual Doctoral Candidate
thesis. It contains descriptions, but is formally unpublished.
Therefore, new taxa described in these two works are un-
available under the ICZN rules. Nevertheless, some of these
names without adequate descriptions (nomina nuda), such
as Anabarites tribaculatus and Angustiochrea magna, were
later employed by Khomentovsky & Gibsher (1996: fig. 13)
and Brasier et al. (1996: fig. 6a).
Hence, there is considerable disagreement surround-
ing anabaritid taxonomy, because of different assessments
252 Artem Kouchinsky et al.

Table 4 Classification of anabaritids by different authors.

Taxa
Val’kov & Sysoev, 1970 Order Angustiochreida Val’kov & Sysoev, 1970.
1. Family Lobiochreidae Val’kov & Sysoev, 1970. Genera: Longiochrea Val’kov & Sysoev, 1970 and Lobiochrea
Val’kov & Sysoev, 1970.
2. Family Aculeochreidae Val’kov & Sysoev, 1970. Genera: Jakutiochrea Val’kov & Sysoev, 1970 and
Aculeochrea Val’kov & Sysoev, 1970.
3. Family Angustiochreidae Val’kov & Sysoev, 1970 (invalid, because the designated type genus is Anabarites
Missarzhevsky, in Voronova & Missarzhevsky, 1969; cf. Glaessner, 1979; Bengtson et al. 1990).
Genera: Anabarites Missarzhevsky, 1969 and Angustiochrea Val’kov & Sysoev, 1970.

Missarzhevsky, 1974 Family Anabaritidae Missarzhevsky, 1974. Genera: Anabarites Missarzhevsky, in Voronova & Missarzhevsky,1969;
Tiksitheca (pars) Missarzhevsky, in Rozanov et al., 1969; Cambrotubulus (?) Missarzhevsky, in Rozanov et al.,
1969; Aculeochrea Val’kov & Sysoev, 1970; Anabaritellus Missarzhevsky, 1974.

Val’kov, 1982 1. Family Aculeochreidae Val’kov & Sysoev, 1970. Genera: Aculeochrea Val’kov & Sysoev, 1970 and Jakutiochrea
Val’kov & Sysoev, 1970.
2. Family Anabaritidae Missarzhevsky, 1974. Genera: Cambrotubulus Missarzhevsky, in Rozanov et al., 1969;
Tiksitheca Missarzhevsky, in Rozanov et al., 1969; Anabarites Missarzhevsky, in Voronova & Missarzhevsky,
1969; Anabaritellus Missarzhevsky, 1974.
3. Family Angustiochreidae Val’kov & Sysoev, 1970. Genera: Angustiochrea Val’kov & Sysoev, 1970;
Selindeochrea Val’kov, 1982; Mariochrea Val’kov, 1982; Gastreochrea Val’kov, 1982.
4. Family Lobiochreidae Val’kov & Sysoev, 1970. Genera: Lobiochrea Val’kov & Sysoev, 1970 and Longiochrea
Val’kov & Sysoev, 1970.

Missarzhevsky, 1989 Family Anabaritidae Missarzhevsky, 1974.

1. Subfamily Cambrotubulinae Missarzhevsky, 1989. Genus Cambrotubulus Missarzhevsky, in Rozanov et al., 1969.
2. Subfamily Tiksithecinae Missarzhevsky, 1989. Genera: Tiksitheca Missarzhevsky, in Rozanov et al., 1969;
Mariochrea Val’kov, 1982; Longiochrea Val’kov & Sysoev, 1970; Kugdatheca Missarzhevsky, in Rozanov et al.,
1969.
3. Subfamily Anabaritinae Missarzhevsky, 1989. Genera: Anabarites Missarzhevsky, in Voronova & Missarzhevsky,
1969; Anabaritellus Missarzhevsky, 1974; Selindeochrea Val’kov, 1982.
4. Subfamily Lobiochreinae Missarzhevsky, 1989. Genera: Lobiochrea Val’kov & Sysoev, 1970; Jakutiochrea
Val’kov & Sysoev, 1970; Aculeochrea Val’kov & Sysoev, 1970.

Esakova, 1989; Esakova 1. Family Lobiochreidae Val’kov & Sysoev, 1970. Genera: Lobiochrea Val’kov & Sysoev, 1970; Longiochrea
& Zhegallo, 1996 Val’kov & Sysoev, 1970; Kugdatheca Missarzhevsky, in Rozanov et al., 1969; Tiksitheca Missarzhevsky, in
Rozanov et al., 1969; possibly, Cambrotubulus Missarzhevsky, in Rozanov et al., 1969.
2. Family Mariochreidae Esakova, 1996. Genera Mariochrea Val’kov, 1982 and Gastreochrea Val’kov, 1982.
3. Family Anabaritidae Missarzhevsky, 1974. Genera: Anabarites Missarzhevsky, in Voronova & Misssarzhevsky,
1969; Jakutiochrea Val’kov & Sysoev, 1970; Selindeochrea Val’kov, 1982; Sexangulatus Fedorov,
in Pel’man et al., 1990; Udzhaites Vasil’eva, 1986.
4. Family Aculeochreidae. Genus Aculeochrea Val’kov & Sysoev, 1970.
5. Family Anabaritellidae Esakova, in Esakova & Zhegallo, 1996. Genus Anabaritellus Missarzhevsky, 1974.

Bokova, 1990, 1992 Order Angustiochreida Val’kov & Sysoev, 1970.

Suborder Cambrotubulina Bokova, 1990 [nomen nudum].
Family Cambrotubulidae Bokova, 1990 [nomen nudum]. Genus Cambrotubulus Missarzhevsky, in Rozanov et al.,
1969.
Suborder Angustiochreina Bokova, 1990 [nomen nudum].
1. Family Anabaritidae Missarzhevsky, 1974. Genera: Anabarites Missarzhevsky, in Voronova & Missarzhevsky,
1969; Angustiochrea Val’kov & Sysoev, 1970; Angustites Bokova, 1990 [nomen nudum]; Anabaritellus
Missarzhevsky, 1974; Udzhaites Vasil’eva, 1986; Tiksitheca Missarzhevsky, in Rozanov et al., 1969;
Lobiochrea Val’kov & Sysoev, 1970.
2. Selindeochreidae Bokova, 1990 [nomen nudum]. Genus Selindeochrea Val’kov, 1982.
3. Aculeochreidae Val’kov & Sysoev, 1970. Genera: Aculeochrea Val’kov & Sysoev, 1970 and Jakutiochrea Val’kov &
Sysoev, 1970.
4. Mariochreidae Esakova, in Esakova & Zhegallo, 1996 (published as nomen nudum in Esakova, 1989).
Genera: Mariochrea Val’kov, 1982; Gastreochrea Val’kov, 1982; Longiochrea Val’kov & Sysoev, 1970.
 Systematics of Lower Cambrian fossil Anabaritids
of variability (Table 4). Given also that the affinity of ana-
baritids and their phylogenetic history are still obscure, the
suprageneric taxonomy is very unstable. Instead of using
formal criteria (even if hierarchically organised), analysis of
ontogenetic and preservational variability in natural fossil
associations of anabaritids is attempted herein in order to
define species and outline genera.
A consistent shape of the cross-section and regularities
in the morphology of thecae (or their internal moulds) during
ontogeny are considered herein the most significant features
for recognition of anabaritid species. The internal moulds of-
ten carry more pronounced grooves than the outer wall, and
the relief may become more pronounced towards the aper-
ture in ontogeny (Fig. 2B–C, G–H, J–M, P–Q, U–V). The
wall is usually ornamented with longitudinal or transverse
sculptural elements (see Figs 26A, 29D, 34, 37L–P, 39E,
48C, 51C–E). The shape of the cross-section in ontogeny is a
function of size and growth mode. Therefore, cross-sections
close in size but significantly different in shape may be at-
tributed to different species (e.g. compare Anabarites hexa-
sulcatus (Missarzhevsky, 1974) and A. hariolus (Val’kov,
1987) below). Lack of transitional morphologies within the
same size class within a given association indicates different
species rather than intraspecific morphological variation.
This case is considered below for the association of A. hexa-
sulcatus, A. ternarius Missarzhevsky, in Rozanov et al., 1969,
A. tripartitus Missarzhevsky, in Rozanov et al., 1969 and
Selindeochrea tecta Val’kov, 1982, as represented by sev-
eral specimens of internal moulds with similar diameters but
different transverse profiles from samples 355 and 369.
Parameters of the thecae, such as their curvature, ex-
pansion rate and size, are often variable within populations
and seem, therefore, to be less important for distinguish-
ing species. For instance, variation in these parameters is
common in fossil associations of Cambrotubulus ex gr. de-
curvatus Missarzhevsky, in Rozanov et al., 1969, but the
presence of several species of Cambrotubulus is neverthe-
less possible (see below). In some forms, however, these
features are rather consistent and characteristic, and so have
been considered to be useful for systematics in these cases.
For example, Anabarites biplicatus (Missarzhevsky, 1989)
can be distinguished as a separate species because it oc-
curs in large quantities as bilaterally symmetrical, curved,
small-sized tubes without larger individuals interpreted as
its further growth stages in the same fossil association
(see below).
The diagnostic characteristics of genera and most of the
species described herein are shown in Table 5. The Material
sections in the Systematic Descriptions, below, list only those
specimens figured herein. Institutional abbreviations for the
material studied are: GIN, Geological Institute of the Russian
Academy of Sciences; SMNH, Swedish Museum of Natural
History. Occurrences are given in the stratigraphical context
discussed above. Assessments of synonymy are based on
A.K.’s personal observations and available illustrations of
sufficient quality. The notation “ex gr.” [ex gregarii] is used
herein to denote that a form belongs to an unresolved species
complex. Appendix 1 of the Supplementary Material gives
information on type species and holotypes. We are inclined
to regard anabaritids as diploblastic-grade metazoans similar
to, or located within, the Cnidaria. There is, however, no firm
evidence for that, and we therefore leave them as incertae
sedis.
253
Genus ANABARITES Missarzhevsky, in Voronova &
Missarzhevsky, 1969
1967 Anabarites [nomen nudum] Missarzhevsky: 20.
1969 Anabarites Missarzhevsky; Voronova & Missar-
zhevsky: 209.
1969 Anabarites Missarzhevsky; Rozanov et al.: 155.
?1969 Kugdatheca Missarzhevsky; Rozanov et al.: 111.
?1969 Tiksitheca Missarzhevsky; Rozanov et al.: 114.
1970 Anabarites Missarzhevsky; Val’kov & Sysoev: 97.
1970 Angustiochrea Val’kov & Sysoev: 97.
1970 Jakutiochrea Val’kov & Sysoev: 98.
?1970 Lobiochrea Val’kov & Sysoev: 96.
1974 Anabaritellus Missarzhevsky: 187.
1977 Anabarites Missarzhevsky; Qian: 259.
1977 Anabaritellus Missarzhevsky; Qian: 260.
1978 Anabarites Missarzhevsky; Qian: 15.
?1978 Tiksitheca Missarzhevsky; Qian: 17.
1981 Anabarites Missarzhevsky; Missarzhevsky & Mam-
betov: 73.
1982 Angustiochrea Val’kov & Sysoev; Val’kov: 67.
1982 Anabarites Missarzhevsky; Val’kov: 73.
1982 Anabaritellus Missarzhevsky; Val’kov: 76–77.
1982 Jakutiochrea Val’kov & Sysoev; Val’kov: 78.
?1982 Lobiochrea Val’kov & Sysoev; Val’kov: 65–66.
?1982 Tiksitheca Missarzhevsky; Val’kov: 73.
1986 Udzhaites Vasil’eva: 103.
1987 Jakutiochrea Val’kov & Sysoev; Val’kov: 111.
1988 Tiksitheca Missarzhevsky; Landing: 690 (pars.).
1989 Jakutiochrea Val’kov & Sysoev; Missarzhevsky: 193.
1989 Anabarites Missarzhevsky: 191.
1989 Anabaritellus Missarzhevsky: 192.
1989 Jakutiochrea Val’kov & Sysoev; Missarzhevsky: 193.
1989 Kotuites Missarzhevsky: 194.
1989 Anabarites Missarzhevsky; Conway Morris & Chen:
619–620.
1989 Anabarites Missarzhevsky; Qian & Bengtson: 125
(pars).
1989 Anabarites Missarzhevsky; Landing et al.: 757 (pars).
?1989 Tiksitheca Missarzhevsky: 190.
?1989 Kugdatheca Missarzhevsky: 190–191.
1990 Anabarites Missarzhevsky; Bengtson et al.: 192–193
(pars).
?1990 Sexangulatus Fedorov; Pel’man et al.: 27.
1991 Anabaritellus Missarzhevsky; Landing & Murphy:
392 (pars).
TYPE SPECIES. Anabarites trisulcatus Missarzhevsky, in
Voronova & Missarzhevsky, 1969.
DIAGNOSIS. Calcareous tubes (thecae) or internal moulds
with triradially symmetrical distribution of longitudinal ele-
ments of the sculpture. The triradial symmetry develops dur-
ing ontogeny and can be combined with bilateral symmetry.
Outer wall without prominent external longitudinal keels,
transversal folds, or concentric invagination of the aperture.
OCCURRENCE. Manykayan and Tommotian Stages of the
Siberian Platform; Meishucunian–Canglangpuan Stages of
China; lower part of the Lower Cambrian of Laurentia; Lower
Cambrian of Gondwana, Avalonia, Western Mongolia; Tom-
motian and ?Atdabanian equivalents of Kazakhstan; upper
Atdabanian equivalent of Baltica.
254 Artem Kouchinsky et al.

Table 5 Diagnostic characteristics of the genera and species of anabaritids described herein.

ANABARITES Missarzhevsky, in Rozanov et al., 1969

Internal moulds and thecae with three-fold symmetry that develops during ontogeny and can be combined with a bilateral symmetry. The
outer wall without prominent external longitudinal keels, transversal folds, or concentric invagination of the aperture.
A. trisulcatus Missarzhevsky, in Rozanov Slowly expanding tubes and internal moulds with three rounded lobes separated by shallow
et al., 1969 narrow grooves. Growth lines often curve in the grooves towards the aperture.
A. latus (Val’kov & Sysoev, 1970) Rapidly expanding internal moulds with three rounded lobes separated by deep grooves.
A. tripartitus Missarzhevsky, in Rozanov Slowly expanding internal moulds with three rounded lobes separated by deep grooves.
et al., 1969
A. missarzhevskyi (Vasil’eva, 1986) Internal moulds with a triangular transverse cross-section and deep narrow grooves; rhomboidal
transverse profile of the lobes becomes rounded at the apex of the mould.
A. ternarius Missarzhevsky, in Rozanov Internal moulds with narrow extended lobes and wide V-shaped grooves in between.
et al., 1969
A. tristichus Missarzhevsky, in Rozanov Internal moulds with three lobes separated by three chains of notches situated in grooves.
et al., 1969
A. compositus Missarzhevsky, in Rozanov Internal moulds with three longitudinal bands of chevron-like structures attached.
et al., 1969
A. valkovi (Bokova, in Bokova & Vasil’eva, Internal moulds with three chains of notches in the middle part of three distinct lobes separated
1990) by wide grooves.
A. hexasulcatus (Missarzhevsky, 1974) Internal moulds with six grooves separating rounded lobes. Secondary grooves appear early on
the lobes delimited by the first-order grooves.
Anabarites hariolus (Val’kov, 1987) Internal moulds with deep grooves and three T- or Y-shaped lobes carrying a secondary groove
along their distal portions. The secondary grooves are shallow and disappear towards the apex.
A. korobovi (Missarzhevsky, in Rozanov & Internal moulds and thecae curved in one plane, with a rounded-triangular cross-section
Missarzhevsky, 1966) extended in the direction of the convex side.
A. rectus Vasil’eva, in Rudavskaya & Internal moulds with almost circular transverse profile and three consistently preserved
Vasil’eva, 1984 longitudinal bands of diagenetic phosphatic material attached.
SELINDEOCHREA Val’kov, 1982
Internal molds and thecae with three prominent lobes. Prominent longitudinal keels produced by the walls extend from distal portions
of the lobes.
S. tecta Val’kov, 1982 Internal moulds twisted through several revolutions and carrying wide V-shaped grooves
between lobes narrowing distally.
S. tricarinata (Missarzhevsky, in Rozanov Internal moulds almost straight or slightly twisted, with rounded lobes and flattened depressions
et al., 1969) in between.
ACULEOCHREA Val’kov & Sysoev, 1970
Internal moulds with rounded triangular cross-section, composed of segments produced by concentric folds. One or several tubercles occur
along lower margins of the folds. Each segment is subdivided by wide and shallow longitudinal grooves into three lobes.
A. ornata Val’kov & Sysoev, 1970 A single tubercle situated in the lower middle of every fold on each lobe.
A. rugosa (Val’kov & Sysoev, 1970) Multiple tubercles situated along lower margin of every fold.

MARIOCHREA Val’kov, 1982

Internal moulds subdivided into stacked segments with rounded triangular transverse profile and edges perpendicular to the growth axis.
The upper face of each segment in bigger specimens can show a cast of a concentric invaginations of the wall with a central
rounded-triangular or clover-leaf-shaped aperture.

CAMBROTUBULUS Missarzhevsky, in Rozanov et al., 1969

Internal moulds and thecae with circular transverse profile at any stage of growth and straight growth lines.

REMARKS. Anabarites Missarzhevsky, 1969, was described
as a slowly expanding tube divided into three convex lobes
by narrow grooves (Voronova & Missarzhevsky 1969).
Angustiochrea Val’kov & Sysoev, 1970, has a much
wider angle of expansion, but in itself this feature is not con-
sidered sufficient to distinguish the genus. The width of the
theca may represent ecological variation related to a sedent-
ary mode of life and thus remains unimportant taxonom-
ically. In Angustiochrea lata Val’kov & Sysoev, 1970 (see
Figs 15–17) there is a considerable variation in expansion
rate, regarded herein as intraspecific (see below).
Jakutiochrea Val’kov & Sysoev, 1970, which carries
three longitudinal rows of tooth-like protrusions on the in-
ner tube wall, shows features otherwise typical of Anabarites
Missarzhevsky. The tooth-like protrusions can hardly be used
as generic characters, because they occur in forms as dif-
ferent morphologically as A. compositus Missarzhevsky, in
Rozanov et al., 1969 (see Figs 29–31), Aculeochrea rugosa
(Val’kov & Sysoev, 1970) (see Fig. 50A–B, D–E), Ana-
barites valkovi (Bokova, in Bokova & Vasil’eva, 1990; see
Fig. 32) and Anabarites sexalox Conway Morris & Bengtson,
in Bengtson et al., 1990.
 Systematics of Lower Cambrian fossil Anabaritids
Udzhaites Vasil’eva, 1986, fits the diagnosis of Ana-
barites used herein, and in its early growth stage the trans-
verse profile of U. missarzhevskyi Vasil’eva, 1986, is similar
to that of Anabarites tripartitus Missarzhevsky, 1969 (see
Fig. 22G–H).
Anabaritellus Missarzhevsky, 1974, was originally
characterised as a six-foliate form. Internal moulds of Ana-
baritellus hexasulcatus Missarzhevsky, 1974, are subdivided
by first- and second-order grooves into six lobes (see
Fig. 33). This feature can potentially be taxonomically im-
portant, because it may reflect the symmetry of the soft body.
There are other forms, however, that suggest a less funda-
mental importance of this feature. Anabarites sexalox Con-
way Morris & Bengtson, in Bengtson et al., 1990, shows a
series of concave impressions in the middle of each of the
three lobes in the earlier stages of growth, and secondary
grooves and lobes at later stages (Bengtson et al. 1990: figs
128 & 129). This is similar to Anabarites valkovi (Bokova,
1990), although the latter does not show secondary lobes at
later stages of growth (see Fig. 32). It seems plausible, there-
fore, that the sixfoliate walls evolved convergently (Bengtson
et al. 1990). Consequently, the recognition of a genus Ana-
baritellus Missarzhevsky, 1974, based only on this feature,
is not tenable.
Kotuites Missarzhevsky, 1989, is a possible junior syn-
onym of Anabarites Missarzhevsky, in Rozanov et al., 1969,
because its bilateral symmetry appears to be a variable fea-
ture (see Fig. 38). Anabarites signatus Mambetov, in Missar-
zhevsky & Mambetov, 1981 and Tiksitheca korobovi (Missar-
zhevsky, in Rozanov & Missarzhevsky, 1966) are similarly
curved in one plane, although, unlike the condition in Kotu-
ites, none of the lobes of Anabarites signatus is extended.
Tiksitheca korobovi is very similar to K. biplicatus, but the
latter apparently has narrower thecae with relatively better-
defined grooves (see Figs 37 & 38). At an early stage of
ontogeny, Kotuites has a regular tri-fold symmetry typical of
Anabarites.
Tiksitheca Missarzhevsky, in Rozanov et al., 1969 and
Kugdatheca Missarzhevsky, in Rozanov et al., 1969, were
initially regarded as allied to hyoliths (Rozanov & Missar-
zhevsky, 1966; Rozanov et al. 1969), but later were reinter-
preted as anabaritids (Missarzhevsky, 1982), albeit without
distinct grooves but having rounded triangular cross-sections.
Apparently the only difference between these two genera is
that Kugdatheca is twisted (see Fig. 39A & D). The only diff-
erence with Anabarites is, then, a lack of longitudinal groo-
ves. It is not clear, therefore, whether this feature is sufficient
to distinguish Tiksitheca and Kugdatheca from Anabarites.
Sexangulatus Fedorov, in Pel’man et al., 1990, is rep-
resented by poorly preserved internal moulds (a paratype is
shown in Fig. 12E) having a rounded hexagonal transverse
profile, which is the main feature distinguishing it from Ana-
barites (Pel’man et al. 1990). An incipient hexasulcate condi-
tion is known from internal moulds of Anabarites rotundum,
A. ex gr. trisulcatus, form 2 (see Fig. 10), A. compositus
Missarzhevsky, in Rozanov et al., 1969 (see Fig. 30F) and
A. tristichus Missarzhevsky, in Rozanov et al., 1969 (see
Fig. 26H), while a hexagonal transverse profile occurs in A.
ex gr. natellus (Val’kov & Sysoev, 1970; see Fig. 13C, E–F).
According to the expanded diagnosis of the genus Anabarites
accepted herein, such a transverse profile is not diagnostic
for the proposed monospecific genus Sexangulatus, which
can therefore be synonymised under the question mark with
Anabarites and represents a problematic erection.
255
Lobiochrea Val’kov & Sysoev, 1970, was originally
described from internal moulds with a rounded triangular
transverse profile. This diagnosis of Lobiochrea Val’kov &
Sysoev, 1970 was changed by Missarzhevsky (1989: 193),
who included in the genus only those forms with a rounded
hexagonal cross-section. The holotype of L. natella Val’kov
& Sysoev, 1970 has, however, a transverse profile interme-
diate between triangular and hexagonal (see Fig. 14N). In
our material there are forms with rounded hexagonal cross-
sections described by Bokova (1990, 1992) as Lobiochrea
formosa [nomen nudum] and herein as Anabarites ex gr. na-
tellus (Val’kov & Sysoev, 1970: see Fig. 13). Lobiochrea
known as internal moulds of variable transverse profile can
also be tentatively synonymised with Anabarites.
Anabarites trisulcatus Missarzhevsky, in Voronova &
Missarzhevsky, 1969 (Figs 6, 7A–E, 11A–L, O–V, 12D)
1967 Anabarites trisulcatus Missarzhevsky: 20 [nomen
nudum].
1969 Anabarites trisulcatus Missarzhevsky; Voronova &
Missarzhevsky: 209, pl. 1(8–9).
1969 Anabarites trisulcatus Missarzhevsky; Rozanov
et al.: 156, pl. 8(10).
?1970 Anabarites trisulcatus Missarzhevsky; Val’kov &
Sysoev: 97, pl. 1(3–5).
1975 Anabarites trisulcatus Missarzhevsky; Matthews &
Missarzhevsky: pl. 2(4, 16).
?1975 Anabarites trisulcatus Missarzhevsky; Val’kov:
pl. 13(3–5).
?1977 Anabarites rotundum Qian; 260, pl. 1(11–12).
1977 Anabarites trisulcatus Qian; 259, pl. 1(9–10, 18–19).
1978 Anabarites trisulcatus Missarzhevsky; Qian: 15,
pls 3(2–3, 12–13), 4(1–2).
1978 Anabarites obliquasulcatus Qian: 16, pl. 3(6–8).
1978 Anabarites sulcoconvex Qian: 16, pl. 3(9–10).
?1978 Anabarites undulatus Qian: 16–17, pl. 3(11).
?1981 Anabarites signatus Missarzhevsky & Mambetov:
73, pl. 3(11, 17, 18).
1982 Anabarites trisulcatus Missarzhevsky; Val’kov: 74,
pl. 11(15–17).
1982 Anabarites trisulcatus Missarzhevsky; Luo et al.:
171, pl. 14(7, 9).
1982 Anabarites primitivus Qian & Jiang, 1980; Luo et al.:
172, pl. 14(10).
?1982 Anabarites grandis Val’kov: 74–75, pl. 11(18).
?1984 Anabarites trisulcatus Chen: 54, pl. 1(18).
?1984 Anabarites cf. trisulcatus Chen: 54–55, pl. 1(19–20).
1985 Anabarites trisulcatus Missarzhevsky; Nowlan et al.:
242, fig. 6.
1989 Anabarites trisulcatus Missarzhevsky; Qian &
Bengtson: 125–127, fig. 84.
1989 Anabarites trisulcatus Missarzhevsky; Conway
Morris & Chen: 628–629, fig. 12c-k.
1989 Anabarites trisulcatus Missarzhevsky: pls 13(19) and
14(1, 3–4).
1989 Anabarites sulcoconvex Qian, 1978; Qian: 147, pl.
23(3–9).
1989 Anabarites tenuistriatus Qian, 1989: 145, pl. 23(1–2).
?1989 Anabarites sulcatus (Bokova, 1985); Qian: 146, pl.
23(10–15).
?1989 Anabarites trisulcatus Missarzhevsky; Qian: 147, pl.
23(16–19) & 24(1–4).
256 Artem Kouchinsky et al.
Figure 6 Anabarites trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969. SMNH X3579–X3588, respectively. A, internal mould
(note growth lines curved upward, towards the aperture in the groove); sample M419/12. B, cross-section of the theca; sample 92-1b/18. C–J,
internal moulds from sample M419/12. Scale bars: A, C–J = 500 μm; B = 200 μm.
?1989 Anabarites rotundus Qian; Conway Morris & Chen:
620–628, figs 6–9, 12a, b.
?1989 Anabarites trisulcatus Missarzhevsky; Hamdi et al.:
fig. 3h.
1990 Anabarites valkovi Fedorov; Pel’man et al.: 26, not
pl. 2(5).
1991 Anabarites trisulcatus Missarzhevsky; Kho-
mentovsky & Karlova, pl. 1, fig. 2.
1991 Anabarites valkovi Fedorov; Khomentovsky & Kar-
lova, pl. 1, fig. 1.
2002 Anabarites trisulcatus Missarzhevsky; Qian et al.:
text-fig. 4(15–16).
2004 Anabarites trisulcatus; Steiner et al.: fig. 3(14).
2004 Anabarites trisulcatus forme sulcoconvex; Steiner
et al.: fig. 3(15).
2004 Anabarites trisulcatus forme obliquasulcatus;
Steiner et al.: fig. 3(16).
2005 Anabarites; Chen & Peng: figs 3, 4.
2005 Anabarites rotundus Qian, 1977; Feng: fig. 2A, B.
2005 Anabarites trisulcatus Missarzhevsky, 1969; Feng:
fig. 2C, D.
?2005 Anabarites sp.; Feng: fig. 2E–H.
2006 Anabarites trisulcatus Missarzhevsky, 1969; Pyle
et al.: 815, fig. 6(1–4).
2007 Anabarites trisulcatus; Steiner et al.: fig. 2D,
E, F, I.
MATERIAL. Represented by 9 specimens from the type local-
ity, sample M419/12, and one transverse section from sample
92-1b/18 (Fig. 6B), SMNH X3579-X3588.
OCCURRENCE. Manykayan (Nemakit–Daldynian)–?Tom-
motian Stages of the Siberian Platform; Meishucunian–
Canglangpuan Stages of China, lower part of the Lower Cam-
brian of Laurentia, Western Gondwana, Avalonia; Lower
Cambrian of Western Mongolia; possibly, Tommotian equi-
valent in Kazakhstan.
DIAGNOSIS. Anabarites species having slowly expanding
tubes, with internal moulds expressing three rounded lobes
separated by shallow grooves (Figs 2A & 6B). Growth lines
in the grooves often curve towards the aperture.
DESCRIPTION. Irregularly curved, slowly expanding internal
moulds with tri-fold transverse profiles. Three rounded lobes
are separated by narrow and shallow, but distinct, grooves. In
some specimens, growth lines in the grooves curve towards
the aperture (Fig. 6A & C). Some possibly preserve remnants
of phosphatised walls (Fig. 6C, H & J).
REMARKS. The type material of Anabarites trisulcatus
(sample M419/12), falls within the range of variation of A.
rotundus Qian, 1977, of Conway Morris & Chen (1989).
Additional material of A. cf. trisulcatus Missarzhevsky, in
Rozanov et al., 1969 from samples 1326 (Fig. 7A–E) and
B-489 (Fig. 8K, N, P–R) shows considerably more variation,
close to the range of variation described by Conway Morris
& Chen (1989) in A. rotundus. In contrast to A. rotundus,
there are only rather irregular and less pronounced swellings
(Figs 7E & 8N). Forms of A. rotundus with regular swellings
and with the incipient hexasulcate condition are, however,
rare in the material described by Conway Morris & Chen
 Systematics of Lower Cambrian fossil Anabaritids 257

Figure 7 A–E, Anabarites trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969. A–B, internal mould, lateral views. C, internal
mould, apertural view. D–E, internal mould, lateral views. F–G, Anabarites cf. signatus Mambetov, in Missarzhevsky & Mambetov, 1981, internal
moulds partly covered with phosphatic crust (arrowed). Sample 1326. SMNH X3603 (A–B), SMNH X3604 (C–E), SMNH X3605 (F), X3606 (G).
Scale bar = 500 μm.

(1989). More specimens are needed from the type locality to
produce a result statistically comparable to A. rotundus, with
159 specimens examined (Conway Morris & Chen 1989).
An incipient hexasulcate condition occurs in A. ex gr. tri-
sulcatus, form 2 (see Fig. 10), named by Bokova (1990) as
A. trapezialis [nomen nudum]. Although the regularity of
swellings in A. rotundus cannot be readily recognised in A.
trisulcatus, this does not seem sufficient to distinguish these
two species. Taking into account the synonymy list of Con-
way Morris & Chen (1989), we tentatively synonymise A.
trisulcatus with A. rotundus and other such forms from the
Yangtze Platform, namely A. obliquasulcatus Qian, 1978 (see
Fig. 11O–T) and A. sulcoconvex Qian, 1978 (see Fig. 11E–L).
Steiner et al. (2004) regarded A. obliquasulcatus and A. sul-
coconvex as intraspecific varieties of A. trisulcatus, because
they occur together and with transitional forms.
The picture labelled as showing Anabarites valkovi
Fedorov, in Pel’man et al., 1990, in pl. 2, fig. 5 of Pel’man
et al. (1990), in fact mistakenly shows an A. bisulcatus Fe-
dorov, in Pel’man et al., 1990 (Fedorov, pers. comm., 2004).
According to Pel’man et al. (1990: 26), the only difference
between A. valkovi (see Fig. 12D) and A. trisulcatus is that in
the longitudinal grooves of the former there are well-defined
growth lines curved towards the aperture. Such a pattern of
growth lines, however, was originally described from A. tri-
sulcatus by Voronova & Missarzhevsky (1969: 210), also
confirmed by Rozanov et al. (1969: 156) and herein. These
two forms, therefore, can be synonymised.
Anabarites grandis Val’kov, 1982, differs from A. trisul-
catus in being several times larger and having a straight aper-
ture (Val’kov 1982: 74–75, pl. 11(18)). On average, tubes of
A. trisulcatus are 2–3 mm long and 0.4–0.6 mm wide (Voro-
nova & Missarzhevsky 1969). Tubes of A. grandis reach a
length of 14 mm and a diameter of 3.5 mm (Val’kov 1982).
Fragments of a similar size belonging to A. trisulcatus oc-
cur, however, in sample M419/12 (A.K., pers. obs.). The
situation with the growth lines is less clear, because they
can also be straight in A. cf. trisulcatus (from samples 1326
258 Artem Kouchinsky et al.

Figure 8 A–J, L–M, O, R, Anabarites cf. signatus Mambetov, in Missarzhevsky & Mambetov, 1981. K, N, P–Q, Anabarites cf. trisulcatus
Missarzhevsky, in Voronova & Missarzhevsky, 1969. SMNH X3589 (A–B), SMNH X3590 (C), SMNH X3591 (D–E), SMNH X3592 (F–G), SMNH
X3593 (H–I), SMNH X3594-X3602 (J–R, respectively). A–I, sample B-484. A–B, internal mould with partly preserved outer phosphatic crust
(arrowed). A, close-up of B showing the initial part. C, internal mould with phosphatic crust and growth lines curved towards the aperture in
grooves (arrow). D–I, different views of three internal moulds from the same sample. J–Q, internal moulds; sample B-489; initial part is
preserved in L & M (close-up of L). R, cross-section of an internal mould. Scale bar: A = 150 μm; B–L, N–R = 500 μm; M = 300 μm.

(Fig. 7A–C) and B-489 (Fig. 8D–E, K & Q)). It seems, there- MATERIAL AND OCCURRENCE. Two specimens from sample
fore, that there is not enough evidence to define A. grandis 1326 and 5 from B-489. Lower Manykayan Stage, SMNH
as a separate species. X3607-X3614.

DESCRIPTION. Slowly expanding, almost straight or gently

Anabarites ex gr. trisulcatus Missarzhevsky, in
Voronova & Missarzhevsky, 1969, form 1 (Fig. 9)
1990 Angustites orbicularis Bokova [nomen nudum].
curved internal moulds. In the early stages of growth, the
internal mould does not carry grooves and the transverse
profile is circular (Fig. 9A–E, I–N). At a later stage three
sharp narrow furrows appear. In one of the specimens they
are interrupted and above the interruption they are slightly
 Systematics of Lower Cambrian fossil Anabaritids 259

Figure 9 Anabarites ex gr. trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969, form 1 (Angustites orbicularis Bokova, 1992
[nomen nudum]), internal moulds. SMNH X3607 (A–B), SMNH X3608 (C–E), SMNH X3609 (F–G), SMNH X3610-X3612 (H–J, respectively), SMNH
X3613 (K–L), SMNH X3614 (M–N). A–E, sample 1326. F–N, sample B-489. N, close-up of M. Scale bar: A–M = 500 μm; N = 200 μm.

displaced clockwise (Fig. 9G, arrowed). Growth lines are
thin and straight (Fig. 9B, M–N).
REMARKS. This form is different from other A. trisulcatus-
like forms in having, at a later growth stage, a consistent
pattern of thin and sharp furrows on the internal mould.
Co-occurring with A. cf. trisulcatus, A. ex gr. trisulcatus
(form 2) and A. cf. signatus, this form may represent a sep-
arate species.
Anabarites ex gr. trisulcatus Missarzhevsky, in
Voronova & Missarzhevsky, 1969, form 2 (Fig. 10)
1990 Angustites trapezialis Bokova [nomen nudum].
MATERIAL AND OCCURRENCE. Four specimens from sample
B-489 and five from sample 1326; SMNH X3615-X3623.
Lower Manykayan Stage.
DESCRIPTION. Internal moulds are slightly irregularly
curved and twisted. In one specimen the early part is cir-
cular in cross-section (Fig. 10N–P), and at a later stage three
lobes with flattened distal parts appear. Along each outer
portion of the lobe there may be a shallow depression (Fig.
10D–I, J (black arrow), L, Q–W). The resultant transverse
profile is variable, from almost rounded triangular (Fig. 10I)
to tri-lobate with incipient secondary grooves in the middle
of the lobes (Fig. 10C–D, L, P, S–T). The growth lines are
angled towards the aperture, in the grooves and often on the
flattened sides of the lobes (Fig. 10G–H, J–K, U–X).
REMARKS. Anabarites ex gr. trisulcatus Missarzhevsky
(form 2) is different from Anabarites ex gr. natellus
(Val’kov & Sysoev, 1970) in having more prominent lobes
and sometimes an incipient hexasulcate transverse pro-
file, as well as in lacking longitudinal low ridges. These
two forms co-occur in samples 1326 and B-489. Some
260 Artem Kouchinsky et al.

Figure 10 Anabarites ex gr. trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969, form 2 (Angustites trapezialis Bokova, 1992
[nomen nudum]). SMNH X3615 (A–C), SMNH X3616 (D–F), SMNH X3617 (G–I), SMNH X3618 (J–L), SMNH X3619 (M), SMNH X3620 (N–P), SMNH
X3621 (Q–S), SMNH X3622 (T–W), SMNH X3623 (X). A–I, sample 1326. A–B, E–F, G–H, internal moulds, lateral views. C–D, I, internal moulds,
apertural views. J–P, sample B-489. K, internal mould, view of the first-order groove (arrowed). J, internal mould, view of the second-order
groove (arrowed); growth lines curved towards the aperture in the grooves. L, view of the apertural end. M, internal mould. N–O, views of the
internal mould showing the early part without grooves. P, view of the apertural end. Q–W, sample 1326. Q, internal mould, view of the lobe. R,
internal mould, view of the groove. S, view of the apertural end of the same specimen. U, internal mould, view of the groove. V, internal mould,
lateral view. W, close-up of V showing growth lines curved towards the aperture in grooves (right arrow) and on lobes (left arrow). T, view of the
apertural end of the same specimen. X, fragment of the phosphatised outer mould of Anabarites ex gr. trisulcatus Missarzhevsky, in Voronova &
Missarzhevsky, 1969, form 2; sample A-380k (misprinted as A-830k, as in Val’kov 1987). Scale bar: A–V, Y–Z = 500 μm; W = 200 μm; X = 1 mm.

internal moulds of A. ex gr. trisulcatus Missarzhevsky,
in Rozanov et al., 1969 (form 2), from samples B-
489 and 1326 show cross-sections similar to A. hexasul-
catus (see below), but with less distinct secondary lobes
(Fig. 10J–L, Q–S). Internal moulds with the well-defined
six lobes and sharp secondary grooves typical of A. hexasul-
catus have not been found in samples B-489 and 1326. Other
specimens of A. ex gr. trisulcatus (form 2) have rounded
triangular to hexagonal profiles. Anabaritellus hexasulcatus
Missarzhevsky, 1974, reported from the earliest complex of
anabaritids on the Siberian Platform, may well represent
Anabarites ex gr. trisulcatus (form 2). Anabarites ex gr.
 Systematics of Lower Cambrian fossil Anabaritids 261

Figure 11 A–B, Anabarites undulatus Qian, 1978 (considered herein A. trisulcatus Missarzhevsky, in Voronova & Missarzhevsky, 1969);
holotype (A, apertural view). C–D, Anabarites rotundum Qian, 1977 (considered herein A. trisulcatus Missarzhevsky); holotype (C, apertural
view). E–L, Anabarites sulcoconvex Qian, 1978 (considered herein A. trisulcatus Missarzhevsky). E–F, paratype specimen 33697. G, holotype. H,
specimen 89425. I–J, specimen 89428. K–L, specimen 89427. M–N, Anabaritellus xishuiensis Qian, 1977 (considered herein ?Anabarites
hexasulcatus (Missarzhevsky, 1974)), holotype. O–T, Anabarites obliquasulcatus Qian, 1978 (considered herein A. trisulcatus Missarzhevsky).
O–Q, holotype. R–S, paratype specimen 33694. T, paratype specimen 33693. U–V, Anabarites tenuistriatus Qian, 1989 (considered herein A.
trisulcatus Missarzhevsky), holotype. W–Z, Tiksitheca huanshadongensis Qian, Chen & Chen, in Qian et al., 1979 (considered herein as repre-
senting problematic erection owing to poor preservation). W–X, holotype. Y–Z, paratype specimen 51729. Scale bar: A–B = 1 mm; C–Z = 500 μm.

trisulcatus (form 2) co-occurs with A. ex gr. trisulcatus
(form 1), and they may represent separate species.
Anabarites ex gr. natellus (Val’kov & Sysoev, 1970)
(Fig. 13)
1970 Lobiochrea natella Val’kov & Sysoev: 96, pl. 1(2).
1975 Lobiochrea natella Val’kov & Sysoev; Val’kov: pl.
13(2).
1982 Lobiochrea natella Val’kov & Sysoev; Val’kov: 66, pl.
8(1–5).
1989 Lobiochrea trialata Val’kov [nomen nudum].
1990 Lobiochrea trialata Val’kov [nomen nudum]; Bokova.
1990 Lobiochrea formosa Bokova [nomen nudum].
MATERIAL. One specimen from sample B-483 (indicated as
holotype of Lobiochrea formosa [nomen nudum] by Bokova,
1990) and one specimen from sample B-489.
OCCURRENCE. Forms consistent with the description of Lo-
biochrea natella Val’kov & Sysoev, 1970, are known from
262 Artem Kouchinsky et al.

Figure 12 A, Tiksitheca curvata Fedorov, in Pel’man et al., 1990 (considered herein Anabarites? licis (Missarzhevsky, in Rozanov
et al., 1969)); holotype. B, Cambrotubulus crassus Fedorov, in Pel’man et al., 1990 (considered herein C. ex gr. decurvatus Missarzhevsky, in
Rozanov et al., 1969); holotype. C, Anabarites bisulcatus Fedorov, in Pel’man et al., 1990 (considered herein A. biplicatus (Missarzhevsky,
1989)); paratype specimen figured in Pelman et al. 1990 (fig. 2(9)). F, Anabarites bisulcatus Fedorov, in Pel’man et al., 1990 (considered herein A.
biplicatus (Missarzhevsky, 1989)); holotype. D, Anabarites valkovi Fedorov, in Pel’man et al., 1990 (considered herein A. trisulcatus
Missarzhevsky, in Voronova & Missarzhevsky 1969); holotype. E, Sexangulatus denudatus Fedorov, in Pel’man et al., 1990 (considered herein
?Anabarites sp. and representing problematic erection owing to poor preservation); paratype. Scale bar: A–B = 1 mm; C–F = 500 μm.
 Systematics of Lower Cambrian fossil Anabaritids 263

Figure 13 Anabarites ex gr. natellus (Val’kov & Sysoev, 1970), internal moulds. A–C, holotype of Lobiochrea formosa Bokova, 1990 [nomen
nudum], sample B-483. A–B, lateral views with a longitudinal ridge indicated by the arrows. C, view of the apertural end with tubes of smaller
diameter stacked inside. D–F, internal mould from sample B-489; SMNH X3625. D, lateral view. E–F, apertural views at the upper and lower
ends of D. Scale bar = 500 μm.

the Siberian Platform, lower Manykayan Stage. Lobiochrea
trialata Val’kov [nomen nudum] has been reported from the
Allatheca cana Biozone of the Selinde section on the south-
eastern Siberian Platform (Val’kov 1989) (Figs 3 [locality 14]
& 4), attributed herein to the upper part of the Manykayan
Stage.
DESCRIPTION. Straight or curved internal moulds with
rounded-hexagonal (Fig. 13C, E–F) transverse profiles. The
internal moulds may carry on the flattened sides three longit-
udinal narrow low ridges, each with a narrow central groove
(arrowed in Fig. 13A–B).
REMARKS. Lobiochrea natella Val’kov & Sysoev 1970, was
described as rounded triangular in cross-section (Val’kov
& Sysoev 1970; Val’kov 1982). There is apparently vari-
ation in the shape of the cross-section of the internal
moulds from rounded triangular to rounded hexagonal,
possibly depending on the growth stage and preserva-
tion. The holotype is intermediate in cross-sectional shape
(Fig. 14N).
Anabarites cf. signatus Mambetov, in Missarzhevsky
& Mambetov, 1981 (Figs 7F–G & 8A–J, L–M, O, R)
MATERIAL AND OCCURRENCE. Five specimens from sample
B-484 (SMNH X3589-X3693); 5 from sample B-489
(SMNH X3594, X3596, X3597, X3599 & X3602); 2
from sample 1326 (SMNH X3605-X3606). Manykayan
Stage.
DESCRIPTION. Internal moulds subdivided into three roun-
ded lobes by three longitudinal distinct grooves. The moulds
264 Artem Kouchinsky et al.

Figure 14 Holotype specimens. A–D, Jakutiochrea convexa Val’kov & Sysoev, 1970 (considered herein Anabarites convexus): A, apertural view;
B–D, lateral views of the mould. E–G, Angustiochrea lata Val’kov & Sysoev, 1970 (considered herein Anabarites latus): E, apertural view; F–G,
lateral views of the mould. H, N Lobiochrea natella Val’kov & Sysoev, 1970 (considered herein Anabarites natellus). H, lateral view, N, apertural
view. I–L, Aculeochrea ornata Val’kov & Sysoev, 1970: I, apertural view; J–L, lateral views. M, Cambrotubulus sibiricus (Val’kov, 1968)
(considered herein Cambrotubulus ex gr. decurvatus Missarzhevsky, in Rozanov et al., 1969). Scale bar = 500 μm.
 Systematics of Lower Cambrian fossil Anabaritids 265

Figure 15 Anabarites latus (Angustiochrea lata Val’kov & Sysoev, 1970). SMNH X3630 (A–E), SMNH X3631 (F), SMNH X3632 (G–K), SMNH
X3633 (L), SMNH X3634 (M–O). A–B, internal mould, lateral views. C, view of the apertural end of the same specimen. E, juvenile specimen of
possibly the same species enlarged from C with the outer surface of the wall replicated by phosphate (arrow). D, initial part of E, enlarged; the
wall is dissolved and situated in the space between the internal mould and the outer phosphatic crust (the latter is arrowed). F, internal mould
of a juvenile specimen with the initial part from sample 1326. H–I, internal mould of a juvenile specimen with grooves. G, view on the apertural
end of the same specimen. J–K, close-up of the apertural region of I; sample B-489. L, internal mould of a specimen at the same developmental
stage as E; sample 96-4/4. M, view of the apical end of a juvenile specimen of possibly the same species covered with a phosphatic crust. N,
close-up of M. O, close-up of N; sample 96-4/4. Scale bar: A, B, F–I, L = 500 μm; C = 300 μm; D = 50 μm; E = 150 μm; J–K = 30 μm;
M = 300 μm; N = 50 μm; O = 10 μm.

are curved in a single plane. Some specimens show a lon-
gitudinally orientated fibrous texture and remains of a phos-
phatic crust on the outer surface of the walls (Figs 7F &
8A–B).
REMARKS. Anabarites signatus Mambetov, in Missar-
zhevsky & Mambetov, 1981, is curved in a single plane,
and this represents the major difference with tubes of A. tri-
sulcatus, which are often curved irregularly (Missarzhevsky
& Mambetov 1981). In samples B-484 and B-489 (Fig. 8A–J,
L–M), forms curved in one plane co-occur with less regularly
curved ones (Fig. 8K, N, P–Q), but no specimen curved in
one plane has been found within the type material of A. trisul-
catus. Neither have regularly curved moulds been reported
from A. rotundus.
Anabarites latus (Val’kov & Sysoev, 1970)
(Figs 14E–G, 15–17)
1970 Angustiochrea lata Val’kov & Sysoev: 98, pl.
1(6–7).
1975 Angustiochrea lata Val’kov & Sysoev; Val’kov: pl.
13(6–7).
1982 Angustiochrea lata Val’kov & Sysoev; Val’kov: 67, pl.
8(6–17).
1989 Anabarites latus (Val’kov & Sysoev); Missarzhevsky:
pl. 13(4, 6–7).
MATERIAL. Six specimens from sample 1326 (SMNH
X3630, X3631, X3635-X3638), 4 from sample B-489
(SMNH X3632, X3639-X3641), 3 from sample B-484
266 Artem Kouchinsky et al.

Figure 16 Anabarites latus (Angustiochrea lata Val’kov & Sysoev, 1970); internal moulds; sample 1326. SMNH X3635 (A–B), SMNH X3636
(C–D), SMNH X3637 (E–G), SMNH X3638 (H–J). A, apertural view. B, lateral view. C–D, lateral views. E, apertural view. F–G, lateral views. H,
apertural view. I–J, lateral views. Scale bar = 500 μm.

(SMNH X3642 & X3643), 2 from sample 96-4/4 (SMNH
X3633 & X3634), and holotype from sample 1326.
OCCURRENCE. Lower Manykayan Stage of the Siberian
Platform.
DIAGNOSIS. Anabarites species having rapidly expanding
internal moulds with three rounded lobes separated by deep
grooves (Fig. 2B–C).
DESCRIPTION. Internal moulds, rapidly expanding and ir-
regularly curved. Deep grooves separate three longitud-
inal lobes. Variability: the grooves may be narrower (see
Figs 15A–C; 16H–J & 17A–C, G–L) or wider (see Figs 16A–
G & 17D–F), depending on the expansion and extension rates
of the lobes. Initial portions of the moulds have a circular
transverse profile, which changes rapidly during ontogeny
(Fig. 15). Growth lines produce obtuse angles angled in the
grooves towards the aperture (Fig. 16B).
REMARKS. The variability (in samples B-484, B-489 and
1326) of both the expansion rate and the width of the grooves
in the internal moulds is considerable (see Figs 16 & 17).
The “classical” form with a very broad theca (Fig. 14E–G)
is known only from the Malaya and Bol’shaya Kuonamka
rivers (see Fig. 3, localities 4 and 5). Initially, this species
had been assigned to a new genus, Angustiochrea Val’kov
& Sysoev, 1970, which differed from Anabarites Missar-
zhevsky, in Voronova & Missarzhevsky, 1969, by a much
 Systematics of Lower Cambrian fossil Anabaritids 267

Figure 17 Anabarites latus (Angustiochrea lata Val’kov & Sysoev, 1970); internal moulds. SMNH X3639 (A–C), SMNH X3640 (D–F), SMNH
X3641 (G–I), SMNH X3642 (J–L), SMNH X3643 (M). A–I, sample B-489. B–C, E–F, G–H, lateral views. A, D, I, apertural views. J–M, sample B–484.
J–K, M, lateral views. L, part of the apertural end. Scale bar: A–K, M = 500 μm; L = 300 μm.

wider angle of expansion (Val’kov & Sysoev 1970). The rate
of expansion is quite variable in these forms, however, and
was probably dependent on environmental conditions during
growth. No other discernible difference exists between An-
gustiochrea and Anabarites, and it is therefore reasonable to
synonymise the two genera.
Anabarites cf. latus (Val’kov & Sysoev, 1970) (Fig. 18)
1989 Anabarites latus (Val’kov & Sysoev, 1970); Missar-
zhevsky: pl. 12(3).
1990 Angustiochrea magna Bokova [nomen nudum].
1996 Angustiochrea magna Bokova; Khomentovsky & Gib-
sher [nomen nudum].
MATERIAL. Three specimens from samples B-339, B-335,
M71-2/69 (SMNH X3644-X3646, respectively), and two
from M71-2/62 (SMNH X3647 & X3648).
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform, and possibly Lower Cambrian of Western
Mongolia.
DESCRIPTION. Almost straight or slightly curved, gently
twisted internal moulds with tri-lobate transverse profile
(Fig. 18B). Grooves are distinct, narrow and shallow; and
are absent in the earlier stages of growth (Fig. 18E–J), where
the internal moulds have circular cross-sections (Fig. 18I).
The growth lines in the grooves tend to curve towards the
apex (Fig. 18D).
REMARKS. This form differs from other species in having
the growth lines in the grooves curved towards the apex.
The rate of expansion is within the variation of A. latus, but
the grooves in the internal moulds are consistently shallower
and narrow. Anabarites cf. latus is common in samples B-
335, M71-2/62, and M71-2/69. It does not co-occur with A.
latus, from which it differs in having shallower grooves and
268 Artem Kouchinsky et al.

Figure 18 Anabarites cf. latus (Val’kov & Sysoev, 1970) (Angustiochrea magna Bokova, 1992 [nomen nudum]), internal moulds. SMNH X3644
(B–D), SMNH X3645 (E–F), SMNH X3646 (G), SMNH X3647 (H–J). A, holotype of Angustiochrea magna Bokova, 1992 [nomen nudum], lateral
view; sample B-339. B, apertural view; sample B-335. C–D, lateral views of the same specimen, with growth lines. E–F, lateral views; sample
M71-2/69. G, lateral view; sample M71-2/62. H, J, lateral views with preserved aperture. I, apical view; sample M71-2/62. Scale bar = 500 μm.

less prominent lobes; it may, therefore, represent a different
species.
Anabarites modestus Bokova, 1985 (Fig. 19)
1985 Anabarites modestus Bokova: 15–16, pl. 1(1).
2002 Anabarites modestus Bokova; Kouchinsky & Bengt-
son: fig. 9H-J.
MATERIAL AND OCCURRENCE. Six specimens from
sample 1282 (SMNH X3649-X3654 and holotype); upper
Manykayan Stage.
DESCRIPTION. Almost straight or irregularly curved internal
moulds with partly phosphatised remains of walls (Fig. 19A-
E). Cross-section is rounded triangular (Fig. 19A, G, I, L), but
at the initial part it is circular. The grooves along each of the
three flattened sides form very shallow and wide depressions.
Growth lines are straight (Fig. 19B, D [arrowed], E).
REMARKS. The stratigraphical position of sample 1282 (col-
lected by A.K. Val’kov in 1966; results published in Val’kov
1975, but previously discussed by Meshkova 1974) in the
upper part of Member 9 (Rozanov et al. 1969, English edi-
tion: 44) is in agreement with Bokova (1985) and Shishkin
(1974), who both collected from the same level of this sec-
tion, as well as with A.K., who collected from section M419
and the vicinity in 1992. Nevertheless, the diversity of fossils
from sample 1282 reported by Bokova (1985) has not been re-
peated in A.K.’s samples, although many moulds very similar
to A. modestus have been found. The age of sample 1282 is
considered pre-Tommotian. Carbon isotopic profiling shows
that this level in the Manykay Formation is well below the
positive excursion I’ in the overlying Medvezhin Forma-
tion (Kaufman et al. 1996), and it can be assigned to the
Purella Biozone, again in accordance with biostratigraphy
(Khomentovsky & Karlova 1993, 2002: fig. 4).
Anabarites modestus Bokova, 1985, differs from A. tri-
sulcatus and the other species described above in having less
distinct longitudinal grooves, in the form of wide shallow
longitudinal depressions (Bokova, 1985: 16), and straight
growth lines. In most cases it is difficult to distinguish
A. modestus from weathered internal moulds of Tiksitheca
Missarzhevsky, in Rozanov et al., 1969 or Anabarites Mis-
sarzhevsky of similar size, and there is little support for re-
cognition of this species beyond material of the type sample,
1282.
 Systematics of Lower Cambrian fossil Anabaritids
Figure 19 Anabarites modestus Bokova, 1985; sample 1282. SMNH
X3649 (D–E), SMNH X3650 (F–G), SMNH X3651 (H–I), SMNH X3652 (J),
SMNH X3653 (K), SMNH X3654 (L–M). A, view of the apertural end of
B, which is the holotype specimen representing internal mould with
phosphatised theca, lateral view. C, close-up from the area arrowed in
B, showing fibrous composition of the wall. E, internal mould with
remains of phosphatised theca. D, close-up of E with fibrous phos-
phatised wall and growth lines (arrow). F, internal mould, lateral view.
G, apertural view. H, internal mould with remains of phosphatised
wall. I, apertural view. J–K, M, internal moulds in lateral view. L,
apertural view. Scale bar: A–B, E–M = 500 μm; C = 100 μm;
D = 200 μm.
269
From the same locality and level Fedorov (1984) de-
scribed a species named Angustiochrea rara Fedorov, 1984.
It was compared to Anabarites latus (Val’kov & Sysoev,
1970) and is different from the latter in having a lower angle
of expansion (10–15◦ ), shallower grooves and straight growth
lines (Fedorov 1984: 8). The described form co-occurs with,
and is similar to, Anabarites modestus, but differs from it in
showing more distinct grooves with occasional depressions,
between the growth lines. We refer A. rara to the genus Ana-
barites Missarzhevsky, in Voronova & Missarzhevsky, 1969,
according to its synonymy accepted herein. Since we have
had no access to the type material of Anabarites rarus for
comparison, we cannot conclude whether A. rarus and A.
modestus are synonymous. If so, A. rarus (Fedorov, 1984)
would be a senior synonym of A. modestus Bokova, 1985.
Weakly developed grooves are known from Ana-
barites kelleri Missarzhevsky, 1989. This form often oc-
curs on the northern Siberian Platform within the upper
Manykayan beds, particularly in the lower Medvezhin and
lower Emyaksin formations. Anabarites kelleri is represen-
ted by 1–2 mm straight internal moulds with tri-lobate or
rounded triangular cross-section, shallow and often indis-
tinct longitudinal grooves, and thin, almost straight, growth
lines. Missarzhevsky (1989: 191–192) noted its very stable
morphology (variability of length and apertural width not
exceeding 15% in the type material) and mass occurrence.
Hence, it is likely that A. kelleri Missarzhevsky, 1989 repres-
ents a separate species.
Anabarites tripartitus Missarzhevsky, in Rozanov
et al., 1969 (Fig. 20)
1967 Anabarites tripartitus Missarzhevsky: 20 [nomen
nudum].
1969 Anabarites tripartitus Missarzhevsky: Rozanov et al.:
158, pl. 8(11, 16, 20).
1975 Anabarites tripartitus Missarzhevsky; Matthews &
Missarzhevsky: pl. 2(3).
1982 Anabarites tripartitus Missarzhevsky; Val’kov: 75,
pl. 12(1–6).
1988 Tiksitheca korobovi (Missarzhevsky); Landing: 690,
fig. 9(13).
1989 Anabarites korobovi (Missarzhevsky); Landing et al.:
757, fig. 5(13–14).
1989 Anabarites tripartitus Missarzhevsky: pl. 13(8).
1989 Anabarites tripartitus Missarzhevsky; Khoment-
ovsky & Karlova: 57, pl. 4(6).
1990 Anabarites tripartitus Missarzhevsky; Khoment-
ovsky et al.: pl. 11(4).
1990 Anabarites tripartitus Missarzhevsky; Pel’man et al.:
pl. 3(8).
?1996 Anabarites tripartitus Missarzhevsky; Esakova &
Zhegallo: 92–93, pl. 3(5–8).
MATERIAL. Four specimens from sample 1733; SMNH
X3656-X3659.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform; lower part of the Lower Cambrian of Avalonia
and, probably, in Western Mongolia.
DIAGNOSIS. Anabarites species having slowly expanding in-
ternal mould with three rounded lobes separated by deep
grooves (see Fig. 2E).
270 Artem Kouchinsky et al.
Figure 20 Anabarites tripartitus Missarzhevsky, in Rozanov et al., 1969; internal moulds; sample 1733. SMNH X3656 (A–C), SMNH X3657
(D–E), SMNH X3658 (F–H), SMNH X3659 (I–K). A–B, D–E, G–H, I–K, lateral views; note a faint ridge along the lobe in B (arrowed). C, F, J,
apertural views. Scale bar = 500 μm.
DESCRIPTION. Internal moulds slightly curved, with three
rounded lobes and deep grooves between them (Fig. 20).
The lobes become more prominent towards the apertural end.
Along the middle part of each lobe there is a narrow, rounded,
low ridge (Fig. 20B, arrowed). Within the grooves, growth
lines are gently curved towards the aperture.
REMARKS. Anabarites tripartitus is a rather distinct form in
all samples (355, 369, 1733) where it has been encountered.
Some moulds of A. tripartitus from the first two samples have
wider lobes than the figured specimens (Fig. 20), but they are
narrower than in Anabarites ex gr. tripartitus Missarzhevsky
(Fig. 21A). Angustiochrea aspera Vasil’eva, in Bokova &
Vasil’eva, 1990 (Fig. 21E–F) has more pronounced lobes
than A. tripartitus of the same size, but its lobes are wider and
shorter than those of A. ternarius Missarzhevsky, in Rozanov
et al., 1969 (see below).
Anabarites ex gr. tripartitus Missarzhevsky, in
Rozanov et al., 1969 (Fig. 21)
1990 Anabarites tribaculatus Bokova [nomen nudum].
1996 Anabarites tribaculatus Bokova [nomen nudum]; Kho-
mentovsky & Gibsher; Brasier et al.
MATERIAL. Three specimens from sample B-355b; SMNH
X3660-X3662.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform and the lower Lower Cambrian of Western
Mongolia.
DESCRIPTION. Internal moulds twisted clockwise
(Fig. 21C–D), with three rounded lobes having swollen
profile and deep intervening grooves (Fig. 21A). The moulds
expand very slowly.
REMARKS. Differs from Anabarites tripartitus in having in-
ternal moulds with lobes widened distally (see Fig. 2F).
Anabarites missarzhevskyi (Vasil’eva, 1986) (Fig. 22)
1986 Udzhaites missarzhevskyi Vasil’eva: 103–104,
pl. 1(a–d).
MATERIAL. Three specimens from sample 96-5a/34.75;
SMNH X3663-X3665.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
DIAGNOSIS. Anabarites species having internal moulds with
triangular transverse cross-section and deep narrow grooves;
the rhomboidal transverse profile of the lobes becomes roun-
ded at the apical part of the mould (see Fig. 2G–H).
DESCRIPTION. Slightly curved internal moulds with three
deep narrow grooves. Lobes have a rhomboidal or triangular
transverse profile (Fig. 22D, F). The grooves become shal-
lower towards the apex (Fig. 22A–C, E). In this direction the
lobes also acquire a rounded shape (Fig. 22E, G–H).
REMARKS. From sample 96-5a/34.75 A. missarzhevskyi is
available as phosphatised internal moulds, clearly morpho-
logically different from Selindeochrea cf. tecta (see Fig. 47),
which in the same sample is represented by mostly glauc-
onitic internal moulds. The basal part of A. missarzhevskyi
has a transverse profile similar to A. tripartitus (Fig. 20) and
there is no evidence of longitudinal keels attached distally
 Systematics of Lower Cambrian fossil Anabaritids
Figure 21 Anabarites cf. tripartitus Missarzhevsky, in Rozanov
et al., 1969 (Anabarites tribaculatus Bokova, 1992 [nomen nudum]),
internal moulds; sample B-335b. A, cross-section; SMNH X3660. B–D,
lateral views, SMNH X3661 (B) & SMNH X3662 (C–D). E–F,
Angustiochrea aspera Vasil’eva, in Bokova & Vasil’eva, 1990; internal
moulds. E, specimen from Bokova & Vasil’eva (1990: pl. 15(3)). F,
specimen from Bokova & Vasil’eva (1990: pl. 15(4)). Scale bar:
A = 300 μm; B–F = 500 μm.
to the lobes, otherwise typical of Selindeochrea Val’kov,
1982.
Anabarites ternarius Missarzhevsky, in Rozanov
et al., 1969 (Figs 23 & 24)
1967 Anabarites ternarius Missarzhevsky: 20 [nomen
nudum].
1969 Anabarites ternarius Missarzhevsky: Rozanov et al.:
159, pl. 8(15, 18).
1982 Anabarites ex gr. ternarius Missarzhevsky; Val’kov:
75–76, pl. 12(12–21).
?1984 Anabarites ternarius Missarzhevsky; Xing et al.: pl.
14(16).
1989 Selindeochrea ternaria (Missarzhevsky); Missar-
zhevsky: pls 13(5), 14(7).
271
Figure 22 Anabarites missarzhevskyi (Udzhaites missarzhevskyi
Vasil’eva, 1986); internal moulds; sample 96–5a/34.75. SMNH X3663
(A–E), SMNH X3664 (F), SMNH X3665 (G–H). A–C, different views of
the same internal mould. D, apertural view. E, apical view. F,
cross-section. H, lateral view at early stage of development. G,
apertural view. Scale bar: A–C, H = 500 μm; D–G = 300 μm.
MATERIAL. Three specimens from sample 355; SMNH
X3666-X3668.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform and probably Lower Meishucunian Stage of the
South China Platform.
DIAGNOSIS. Anabarites species having internal moulds with
narrow extended lobes, straight in cross-section and wide V-
shaped grooves in between (see Fig. 2I).
DESCRIPTION. Fragments of internal moulds twisted anti-
clockwise, with very narrow extended lobes, straight in cross-
section, and very wide grooves in between. The lobes do not
significantly change their width distally (see Fig. 24). There
is undulation on the lobes with low folds inclined at ca. 45◦
towards the aperture. Growth lines are apparently pointed
towards the aperture in the grooves (Fig. 23A).
272 Artem Kouchinsky et al.

Figure 24 Anabarites ternarius Missarzhevsky, in Rozanov et al.,

1969; sample 355a. A, transversal profile of internal mould, SMNH
X3668. B, transversal profile of the apical end of the specimen in Fig.
23B. Scale bar = 500 μm.

Figure 23 Anabarites ternarius Missarzhevsky, in Rozanov et al.,

1969; sample 355a. A–B, two fragments of internal moulds; SMNH
X3666 & X3667, respectively. Scale bar = 500 μm.

REMARKS. This form is distinguished from other species

of Anabarites by its very extended lobes. In samples 355
and 369 such forms as A. tripartitus, A. hexasulcatus and
Selindeochrea tecta co-occur with A. ternarius and are rep-
resented by several specimens of internal moulds having sim-
ilar diameters but different transverse profiles. Thus, they
represent different species rather than intraspecific morpho-
logical variation.
Figure 25 Anabarites ex gr. ternarius Missarzhevsky, in Rozanov
Anabarites ex gr. ternarius Missarzhevsky, in et al., 1969; internal moulds; sample 1571/2. SMNH
Rozanov et al., 1969 (Fig. 25) X3669 (A–B) and SMNH X3670 (C–D). A, apertural view; B, C,
MATERIAL AND OCCURRENCE. Two specimens from sample lateral views; D, apical view. Scale bar: A, D = 300 μm; B,
1571/2 (SMNH X3669 & X3670); ?upper Manykayan Stage. C = 500 μm.
 Systematics of Lower Cambrian fossil Anabaritids 273

DESCRIPTION. Internal moulds twisted clockwise, with nar-

row and very extended lobes. Each lobe is additionally curved
and its distal portion is turned clockwise in the apertural view
(Fig. 25A). The early part has a transverse profile with smal-
ler and slightly twisted rounded lobes (Fig. 25D).
REMARKS. The form is different from A. ternarius in being
twisted in the opposite sense and in having lobes additionally
turned in distal portions.

Anabarites tristichus Missarzhevsky, in Rozanov

et al., 1969 (Figs 26–28)
1965 Hyolithellus sp. Sysoev: 13, Fig. 2.
1967 Anabarites tristichus Missarzhevsky: 20 [nomen
nudum].
1969 Anabarites tristichus Missarzhevsky: Rozanov et al.:
156–157, pl. 8(1, 14 & 19).
1975 Jakutiochrea tristicha (Missarzhevsky); Val’kov: pl.
13(9).
1975 Anabarites tristichus Missarzhevsky; Matthews &
Missarzhevsky: pl. 2(8).
1982 Jakutiochrea sp. Val’kov: 78–79, pl. 13(20).
1982 Jakutiochrea tristicha (Missarzhevsky); Val’kov: pl.
13(17–19).
1983 Anabarites tristichus Missarzhevsky; Sokolov &
Zhuravleva: 160, pl. 51(2).
1984 Anabarites gracilis Chen: 62, pl. 1(9).
1987 Jakutiochrea solita Val’kov: 111–112, pl. 14(1–5).
1987 Jakutiochrea lenta Val’kov: 114, pl. 14(7–8).
?1987 Jakutiochrea portentosa Val’kov: 113, pl. 14(6).
1989 Anabarites trisulcatus Missarzhevsky; Brasier: pl.
7.4(9).
1989 Anabarites tristichus Missarzhevsky; Khomentovsky
& Karlova: 56, pl. 6(4).
1989 Jacutiochrea tristicha (Missarzhevsky); Missar-
zhevsky: pl. 13(3, 16–17).
2002 Jacutiochrea tristicha (Missarzhevsky); Kouchinsky
& Bengtson: figs 2–5.
MATERIAL. Two specimens from sample M321/31 (SMNH
X3673 & X3676), 2 from sample 96-5a/34.75 (SMNH
X3674-X3675), 2 from sample 92-2/25 (SMNH X3671-
X3672), and 1 specimen from sample 92-2/26 (SMNH
X3409).
OCCURRENCE. Upper Manykayan Stage–D. regularis
Figure 26 Anabarites tristichus Missarzhevsky, in Rozanov
Biozone of the Tommotian Stage of the Siberian Platform;
et al., 1969. A, theca preserved in celestite–barite; specimen SMNH
Lower Meishucunian Stage of the South China Platform.
X3409 (see Kouchinsky & Bengtson, 2002). B, close-up of A (arrowed)
DIAGNOSIS. Anabarites species having internal moulds with showing septum-like protrusion of the wall; sample 92-2/26. C–H,
three lobes separated by three grooves carrying longitudinal SMNH X3671–X3676, respectively. C–G, internal moulds, lateral
rows of notches. views. C–D, sample 92–2/25. E, sample M321/31. F–G, sample
96–5a/34.75. H, internal mould, transverse profile; sample M321/31.
DESCRIPTION. Almost straight, slightly twisted internal Scale bar: A, C–H = 500 μm; B = 20 μm.
moulds, with three flattened lobes separated by shallow and
wide depressions (grooves) (Fig. 26H). The middle of each internal moulds are produced by tooth-like internal protru-
depression has longitudinally elongated notches from the sions of the wall beneath the flanges (Fig. 26B).
apex to the aperture (Fig. 26C–G). One specimen is repres-
ented by a straight fragment of theca with walls preserved in REMARKS. In forms from the type locality of Anabarites tri-
celestite and barite, but without apex and aperture (Fig. 26A– stichus (sample M321/31), variation in both the roundness of
B; Kouchinsky & Bengtson 2002). The wall has annular the lobes and the concavity of depressions between them sug-
flanges, 20–30 μm high and 100–250 μm apart, except in the gests that the other two described forms, Jakutiochrea solita
narrower parts of the tube, where the flanges are absent. More Val’kov, 1987 (Fig. 27) and J. lenta Bokova & Val’kov, 1987,
subdued annular rugae are situated between the flanges. The in Val’kov, 1987 (Fig. 28), actually represent morphological
thickness of the wall is 10–15 μm. The notches observed on and preservational forms of A. tristichus. The latter (samples
274 Artem Kouchinsky et al.
Figure 27 Jakutiochrea solita Val’kov, 1987 (considered herein
Anabarites tristichus Missarzhevsky, in Rozanov et al., 1969). A–B,
holotype specimen; sample 769. Scale bar = 500 μm.
M321/31 and 92-2/26) has more flattened lobes than J. sol-
ita (samples 769 and B-335), but this feature varies between
internal moulds of the same size. Examination of type collec-
tions did not reveal the presence of A. tristichus-like forms
in either sample 1326 or other samples that yield the earliest
complex of anabaritids from the lower Manykayan Stage.
However, eroded internal moulds of Aculeochrea rugosa
(Val’kov & Sysoev, 1970) from sample 1326 do show pits
similar to those seen in moulds of Anabarites tristichus (see
Fig. 50). From this complex, Val’kov & Sysoev (1970) de-
scribed Jakutiochrea convexa (see Fig. 14A–D), with a roun-
ded triangular transverse profile and notches located in the
middle of the convex lobes.
Anabarites compositus Missarzhevsky, in Rozanov
et al., 1969 (Figs 29–31)
1967 Anabarites compositus Missarzhevsky: 20
[nomen nudum].
1969 Anabarites compositus Missarzhevsky: Rozanov
et al.: 158, pl. 8(5, 9).
1975 Anabarites compositus Missarzhevsky; Matthews
& Missarzhevsky: pls 2(18–19), 4(10–11).
non 1987 Anabarites compositus Missarzhevsky; Hinz: 72,
pl. 14(37–38), text-fig. 3I.
1989 Aculeochrea composita (Missarzhevsky); Missar-
zhevsky: pl. 14(11–12).
MATERIAL. Eight specimens from sample 96-5a/34.5.
SMNH X3677-X3684.
Figure 28 A–B, lateral views of the holotype specimen of
Jakutiochrea lenta Bokova & Val’kov, in Val’kov, 1987; sample 907.
C–D, lateral views of the holotype specimen of Jakutiochrea
portentosa Bokova & Val’kov, in Val’kov, 1987. E, enlarged area with
parallel lines of impressions of septum-like protrusions, arrowed;
sample 769. Both forms are synonymised herein with Anabarites
tristichus Missarzhevsky, in Rozanov et al., 1969. Scale bar:
A–D = 500 μm; E = 200 μm.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
DIAGNOSIS. Anabarites species characterised by internal
moulds with three longitudinal bands of chevron-like struc-
tures.
DESCRIPTION. Straight or slightly curved internal moulds,
with remnants of phosphatised walls. Cross-section of the
theca is rounded hexagonal; internal moulds show three
flattened lobes (Fig. 30F). The surface of the internal mould is
usually ornamented with three longitudinal bands of chevron-
like structures usually pointing towards the apex and situated
in the central region of the flattened lobes (Figs 29 & 31).
Growth lines, produced by flanges of the wall (preserved
as phosphatised remains on the internal moulds), are almost
straight and superimposed on the chevrons (Figs 29 & 31).
Where the tube wall is partly phosphatised, the growth lines
display small flanges crossing the surface of the theca and
overlapping the chevron-like structures (Figs 29D–K & 31).
One mould is preserved with extended chevrons that pro-
duce folds running around the circumference (Fig. 29F–G).
Longer fragments of the moulds show that the chevron-like
elements may be transformed into cordate structures, but ori-
entated in the opposite direction, i.e. with the lobes towards
the apex (Figs 29C, J & 31A). Other internal moulds may
show notches extending along the lobes and partly covered
by the chevrons (Figs 29I & 31). These notches represent
imprints of the tooth-like protrusions that can be seen in
positive relief in phosphatised walls (Fig. 30). There is one
such protrusion beneath each chevron-like element (Fig. 31).
 Systematics of Lower Cambrian fossil Anabaritids 275

Figure 29 Anabarites compositus Missarzhevsky, in Rozanov et al., 1969; sample 96-5a/34.5. SMNH X3677 (A–C), SMNH X3678 (D–E), SMNH
X3679 (F–G), SMNH X3680 (H, K–M), SMNH X3681 (I–J). C, internal phosphatic crust with chevron-like structures. A, close-up of C, lateral view of
the chevrons. B, close-up of C, plane view of the chevrons. D, phosphatised theca preserved with growth lines (flanges). E, close-up of D. F,
internal mould with ring-like folds. G, close-up of F. H, internal mould with remains of phosphatised theca (arrow). K, close-up of H,
phosphatised wall with flanges overlaps chevron-like structures (arrow), lateral view. L, close-up of H, phosphatised theca with flanges overlaps
chevron-like structures (arrow), plane view. M, close-up of L (chevron indicated by arrow). J, internal mould with chevron-like structures and
notches below them; the outer phosphatic crust is arrowed. I, close-up of the lower part of the mould in J showing notches. Scale bar: A, B, E, G,
I, K, M = 100 μm; L = 200 μm; C, D, F, H, J = 500 μm.
276 Artem Kouchinsky et al.

Figure 30 Anabarites compositus Missarzhevsky, in Rozanov et al., 1969; sample 96-5a/34.5. SMNH X3682 (A–E) and SMNH X3683 (F–H). A,
inner side of a fragment of the outer phosphatic crust with septum-like outgrowths. B, close-up of A. C, bilayered structure of the inner
phosphatic crust; boundary between the two layers is indicated by an arrow. D, close-up of C. E, close-up of A, lateral view. H, internal
phosphatic crust with remains (arrows) of the theca (aperture to the bottom). F, view of the internal cavity of the theca. G (represents a fragment
of F tilted), showing septum-like outgrowths (arrowed). Scale bar: A, H = 200 μm; B, E = 50 μm; C = 7 μm; D = 2 μm; F, G = 100 μm.

A thin crust, reflecting the internal surface of the original
tube, shows septum-like protrusions directed inwards and
obliquely inclined towards the aperture (Fig. 30A–B, F–G).
REMARKS. The tooth-like structures resemble the protru-
sions in Anabarites tristichus (see Fig. 26). Anabarites com-
positus does not seem to be a preservational form of A. tri-
stichus, because the chevron-like structures are consistent,
and in our sample A. compositus does not co-occur with A.
tristichus. The affinity of problematic fossils from the upper
Lower Cambrian of England misidentified as A. compositus
Missarzhevsky (Hinz, 1987: pl. 14: 37–38) is questionable,
but they resemble Selindeochrea tricarinata (Missarzhevsky)
(see also Brasier 1989: 135). The age of the holotype
of Anabarites compositus, from the Medvezhin Formation
(sample M314/12; see Appendix 2 of Supplementary Ma-
terial), was indicated as “Dokidocyathus regularis Biozone
(Lapworthella tortuosa Subzone)” by Rozanov et al. (1969).
There are, however, no chemostratigraphical data from sec-
tion M314 that directly show its age to be either Tommotian
or pre-Tommotian, nor can the biostratigraphy provide a re-
liable conclusion in this respect (Missarzhevsky 1989).
Anabarites valkovi (Bokova, in Bokova & Vasil’eva,
1990) (Fig. 32)
1987 Jakutiochrea cf. tristicha Missarzhevsky; Val’kov:
114–115, pl. 14(9–10).
1990 Jakutiochrea valkovi Bokova; Bokova & Vasil’eva:
29–30, pl. 15(5).
 Systematics of Lower Cambrian fossil Anabaritids
Figure 31 Anabarites compositus Missarzhevsky, in Rozanov et al.,
1969; sample 96-5a/34.5, SMNH X3684. A, internal phosphatic crust
with chevron-like structures on the outer surface and septum-like
outgrowths (arrow) on the inner side. B, close-up of A, with a partly
phosphatised flange overlapping a chevron-like structure (aperture to
the right). C–D, close-ups of B. Scale bar: A = 250 μm; B = 100 μm;
C–D = 50 μm.
MATERIAL AND OCCURRENCE. Two specimens from sample
907 (SMNH X3687 & X3688); 2 specimens from samples
B-337 and B-339 (SMNH X3686 & X3685, respectively);
upper Manykayan Stage.
DIAGNOSIS. Anabarites species characterised by internal
moulds carrying a longitudinal series of notches in the middle
part of three distinct lobes separated by wide grooves (see
Fig. 2L–M).
DESCRIPTION. Internal moulds slightly curved. Three dis-
tinct lobes with flattened middle portions separated by wide
grooves. Each lobe carries an aligned succession of small and
longitudinally elongated (Fig. 32F) notches in the middle
277
Figure 32 Anabarites valkovi (Jakutiochrea valkovi Bokova, in
Bokova & Vasil’eva, 1990); internal moulds. SMNH X3685 (A–B),
SMNH X3686 (C–F), SMNH X3687 (G–H), SMNH X3688 (I–J). A, lateral
view. B, enlarged apical end of A; sample B-339. D–E, lateral views. C,
apertural view. F, close-up of E with a chain of impressions from
tooth-like protrusions; sample B-337. G–J, sample 907. G–H, lateral
views. J, fragment of an internal mould. I, view on the upper end of the
mould. Scale bar: A, C–E, G–J = 500 μm; B = 300 μm; F = 100 μm.
(Fig. 32E–F, H, J). The moulds have thin, straight growth
lines (Fig. 32A–B).
REMARKS. Anabarites valkovi (from samples 907, B-
337, and B-339) differs from other forms, because
it carries notches in the middle part of the lobes.
278 Artem Kouchinsky et al.

Figure 33 Anabarites hexasulcatus (Anabaritellus hexasulcatus Missarzhevsky, 1974). C, view of the major groove of the internal mould. B,
view of the secondary groove of the internal mould. A, view of the apertural end. D, view of the apical end; sample 355a; SMNH X3689. F,
holotype specimen, view of the major groove of the internal mould. G, view of the secondary groove of the internal mould. E, view of the
apertural end; sample M71–2/69. I, view of the major groove of the internal mould. J, view of the secondary groove of the internal mould. H,
view of the apertural end. K, view of the apical end; sample 879; SMNH X3690. N, view of the major groove of the internal mould. M, view of the
secondary groove of the internal mould. L, view of the apertural end; sample M71–2/62; SMNH X3691. O, view of the major groove of the
internal mould. P, view of the upper end of the mould; sample 907; SMNH X3692. Q, view of the secondary groove of an internal mould; sample
M71-2/69; SMNH X3693. Scale bar = 500 μm.

Jakutiochrea convexa Val’kov & Sysoev, 1970 (see Fig. 14B– Anabarites hexasulcatus (Missarzhevsky, 1974)
D) also shows distinct pitted lines on the convex sides of the (Fig. 33)
mould, but A. valkovi has consistently more prominent and
distinct lobes. This form is included within the genus Ana- 1974 Anabaritellus hexasulcatus Missarzhevsky: 187, pl.
barites, because the only significant difference from other 23(6–7).
species of the genus are the notches on the internal moulds, 1975 Anabarites hexasulcatus (Missarzhevsky); Matthews
reflecting tooth-like protrusions of the inner wall. & Missarzhevsky: pl. 2(11).
 Systematics of Lower Cambrian fossil Anabaritids
Figure 34 Anabarites sexalox Conway Morris & Bengtson, in
Bengtson et al., 1990; sample UNEL1873. A, overall view of the theca
replaced by phosphate. B, apertural view of another specimen (note
arrowed internal protrusions of the apertural margin). Scale bar:
A = 500 μm; B = 300 μm.
?1977 Anabaritellus xishuiensis Qian: 260, pl. 1(8).
1982 Anabaritellus hexasulcatus Missarzhevsky; Val’kov:
77, pl. 13(1–16).
1987 Anabaritellus hexasulcatus Missarzhevsky; Val’kov:
109, pl. 13(17–18).
?1988 Anabaritellus cylindriculus Qian & Ding; Ding &
Qian: 44–45, pl. 3(7–8).
1989 Anabaritellus hexasulcatus Missarzhevsky: pl. 14(9).
1989 Anabaritellus hexasulcatus Missarzhevsky; Kho-
mentovsky & Karlova: 57, pl. 6(8).
1996 Anabaritellus hexasulcatus Missarzhevsky; Esakova
& Zhegallo: 96, pl. 3(17).
MATERIAL. One specimen from sample M71-2/69 (SMNH
X3693); 4 specimens from samples 355, 879, M71-2/62 and
907 (SMNH X3689-X3692, respectively); holotype from
sample M71-2/69.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform; reported from the lower part of the Lower Cam-
brian of Western Mongolia and Lower Meishucunian Stage
of the South China Platform.
DESCRIPTION. Irregularly curved internal moulds having six
grooves, first- and second-order. The latter are typically shal-
lower than the first-order ones. In the grooves casts of growth
279
Figure 35 Anabarites cf. isiticus Missarzhevsky, 1974; internal
moulds; sample 1571/2. SMNH X3781 (A–C), SMNH X3782 (D–F),
SMNH X3783 (G–I). B, view of the major groove. C, view of the
secondary groove. A, view of the apertural end. F, view of the major
groove. E, view of the secondary groove. D, view of the apertural end.
I, view of the major groove. G, view of the secondary groove. H, view
of the upper end. Scale bar: A, D, H = 300 μm; B, C, E–G = 500 μm.
lines are curved towards the aperture (Fig. 33O). As represen-
ted by the internal moulds, there is a range of variation from
the different samples. The type specimen shows second-order
grooves that are significantly less developed than the first-
order ones (Fig. 33E–G). Forms from other samples have
more prominent lobes (Fig. 33A) and some of them have
deeper and broader second-order grooves (Fig. 33H).
DIAGNOSIS. Anabarites species characterised by internal
moulds with six grooves separating rounded lobes (see
Fig. 2N). Second-order grooves appear early on the lobes
delimited by the first-order grooves.
REMARKS. Unlike A. sexalox Conway Morris & Bengtson,
in Bengtson et al., 1990, from Australia (Fig. 34), the internal
moulds of A. hexasulcatus from sample M71-2/69 show no
notches at any stage of growth. In this sample there is also
no transition between A. tristichus, having notches between
the lobes, and A. hexasulcatus of the same size. Anabar-
ites valkovi from sample 907 co-occurs with A. hexasulcatus
of similar size. They are morphologically different in hav-
ing three lobes with notches and six lobes without notches,
respectively. Internal moulds of A. isiticus Missarzhevsky,
1974 have prominent lobes and approach the tranverse pro-
file of A. sexalox (Fig. 34), although the notches are miss-
ing. Missarzhevsky (1974: 187) reported a maximum dia-
meter of 0.5–0.6 mm in his form, that is ca. half as big as
illustrated originally (pl. 23(11–12) therein), as well as in
the fragments reported herein (Fig. 35). Anabarites isiticus
280 Artem Kouchinsky et al.

appears on the Siberian Platform later than A. hexasulcatus,

in the Tommotian Stage, but earlier than the Australian A.
sexalox. Anabaritellus xishuiensis Qian, 1977 is a twisted
internal mould with a longitudinal depression along each
of the three lobes (Fig. 11M–N). The secondary lobes are
not distinct. There is no specimen of A. hexasulcatus in our
material combining indistinct, but strongly twisted, second-
ary lobes. The form appears morphologically different from
Anabaritellus cylindriculus Qian & Ding, in Ding & Qian,
1988 (pl. 3(7–8)). The latter has more prominent longitudinal
ridges, while having 2–3 times smaller diameter. Question-
ably synonymised herein with Anabarites hexasulcatus, this
fossil is similar to Coleoloides trigeminatus Missarzhevsky,
in Rozanov et al., 1969 (see ‘Remarks’ in the description of
Selindeochrea below).

Anabarites hariolus (Val’kov, 1987) (Fig. 36)

1987 Anabaritellus hariolus Val’kov: 110, pl. 13(14–16).

MATERIAL. Three specimens from sample 806/3-4 (SMNH
X3694-X3696); 1 specimen from sample 96-5a/34.5 (SMNH
X3697).
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
DIAGNOSIS. Anabarites species with deep grooves and three
T- or Y-shaped lobes, each carrying a secondary groove along
the distal portion. The secondary grooves are shallow and
disappear towards the apical part.
DESCRIPTION. Internal moulds are almost straight and ex-
pand rapidly. Three very deep grooves separate T- or Y-
shaped lobes that carry a secondary groove along their distal
portions (Fig. 36). Secondary grooves are not developed at
the apical part preserved in one of the specimen (Figs 2P, Q
& 36N).
REMARKS. Anabarites hariolus does not co-occur with A.
hexasulcatus, and comparison of these two forms from dif-
ferent samples shows that specimens of similar size have
different cross-sections. While small A. hariolus already
have their characteristic cross-section, A. hexasulcatus of the
same size display only weakly developed grooves (compare Figure 36 Anabarites hariolus (Anabaritellus hariolus Val’kov, 1987).
Figs 33 & 36). One specimen of A. hariolus (Fig. 36K–N) A–J, sample 806/3–4. SMNH X3694 (A–C), SMNH X3695 (D–F), SMNH
shows changes of cross-section during growth that are differ- X3696 (G–J). A, view of the major groove of a theca replicated by a
ent from those in A. hexasulcatus. In the latter, the secondary phosphatic crust. B, view of the secondary groove of the theca
groove appears in the initial part, where it is relatively narrow replicated by a phosphatic crust. C, view of the apertural end. D, view
and not as broad as in A. hariolus. In A. hariolus the trans- of the major groove of a theca replicated by a phosphatic crust. E,
verse profile at the early stage is reminiscent of A. tripartitus view of the secondary groove of the theca replicated by a phosphatic
(Fig. 36N). During growth the lobes flatten, on account of crust. F, view of the apertural end. G, view of the major groove of theca
the median secondary wide depression (groove). The depres- replicated by a phosphatic crust. H, view of the secondary groove of
sion quickly becomes deeper, while lobes continue to grow theca replicated by a phosphatic crust. I, view of the apertural end. J,
outwards (Fig. 36M–N). view of the apical end. K–M, SMNH X3697, sample 96–5a/34.5. K,
view of the major groove of a theca replicated by a phosphatic crust. L,
Anabarites korobovi (Missarzhevsky, in Rozanov & view of the secondary groove of the theca replicated by a phosphatic
Missarzhevsky, 1966) (Fig. 37) crust. M, view of the apertural end. N, view of the apical end. Scale
bar: A, B, D, E, G, H, K, L = 500 μm; C, F, I, J, M, N = 300 μm.
1966 Semielliptotheca korobovi Missarzhevsky; Rozanov &
Missarzhevsky: 109–110, pl. 12(10), fig. 66.
1969 Tiksitheca korobovi (Missarzhevsky); Missarzhevsky, 1983 Tiksitheca korobovi (Missarzhevsky); Sokolov &
in Rozanov et al., 114–115. Zhuravleva: 161, pl. 51(8).
1974 Tiksitheca korobovi (Missarzhevsky); Meshkova: 60, 1989 Tiksitheca korobove (Missarzhevsky); Missarzhevsky:
pl. 9(3). pl. 12(8).
 Systematics of Lower Cambrian fossil Anabaritids 281

Figure 37 Anabarites korobovi (Semielliptotheca korobovi Missarzhevsky, in Rozanov & Missarzhevsky, 1966). SMNH X3698 (A–C), SMNH
X3699 (D), SMNH X3700 (E–F), SMNH X3701 (G), SMNH X3702 (H–I), SMNH X3703 (J–K), SMNH X3704 (L–P). A, phosphatised theca. B, close-up
of A showing outer surface of the outer phosphatic crust with growth lines. C, view of the apertural end; sample 11/16.05. D, internal mould with
remains of phosphatised wall; sample 11/16.05. I, internal mould with phosphatised wall. H, close-up of I; sample 11/16.05. J, internal mould
with phosphatised wall. K, close-up of J showing fibrous composition of phosphatised wall and outer phosphatic crust (arrow); sample 11/16.1.
L–M, phosphatised theca with preserved aperture (M, view of the concave side). N–O, views of the apertural end with a flange. P, detail of N;
sample 11/16.05. F, internal mould, lateral view. E, close-up of F with replicas of the wall; sample 106b. G, internal mould with the outer
phosphatic crust (arrow); sample 106. Scale bar: A, F, G, I, J, L, M = 500 μm; B, D = 200 μm; C, O = 300 μm; E, H, K, N, P = 100 μm.

MATERIAL. Two specimens from samples 106 and 106b
(SMNH X3701 & X3700); 4 from 11/16.05 (SMNH X3698,
X3699, X3702 & X3704); 1 from 11/16.1 (SMNH X3703).
OCCURRENCE. Nochorojcyathus sunnaginicus and Dokido-
cyathus regularis Biozones of the Tommotian Stage of the
Siberian Platform; reported also from the Meishucunian
Stage of the South China Platform and from the lower part
of the Lower Cambrian of Avalonia.
verse profile rounded triangular or ovoid, extended in the
direction of the convex side (Fig. 37C, O). Growth lines and
aperture straight. In one case a flange is preserved close to
the aperture (Fig. 37L–O). The wall apparently consisted of
fibres orientated longitudinally (Fig. 37D–E, H-K, N, P).
DIAGNOSIS. Anabarites species combining a curvature in
one plane and a rounded-triangular cross-section extended in
the direction of the convex side (see Fig. 2R).

DESCRIPTION. Internal moulds and thecae curved in one REMARKS. Anabarites korobovi was first described as a hy-
plane. Sides of moulds flattened or slightly convex. Trans- olith, either as Semielliptotheca korobovi (in Rozanov &
282 Artem Kouchinsky et al.

Figure 38 A–D, H–K, O, Anabarites cf. korobovi (Semielliptotheca korobovi Missarzhevsky, in Rozanov & Missarzhevsky, 1966). A–B, internal
mould with phosphatised remains of the wall (A, view of the convex side). C, view of the apertural end. D, close-up of the apertural end of B;
sample 1487. H–I, internal mould (I, view of the concave side); sample 882. J, internal mould, lateral view. K, view of the apertural end; sample
882. O, internal mould with growth lines arched towards the aperture; sample 882. E–G, L–N, P–R, Anabarites biplicatus (Kotuites biplicatus
Missarzhevsky, 1989). F–G, internal mould (G, view of the convex side). E, initial part enlarged from F; sample 92–2/21. L, internal mould, lateral
view. M–N, close-ups of the initial part showing the apical end (N) and a cancellate texture on the surface of internal mould; sample M321/26. P,
internal mould, lateral view; sample 92–2/21. R, internal mould, lateral view. Q, close-up of the initial part of the mould (R); sample 92-2/21.
SMNH X3706 (A–C), SMNH X3707 (E–G), SMNH X3708 (H–I), SMNH X3709 (J–K), SMNH X3710 (L–N), SMNH X3711 (O), SMNH X3712 (P), SMNH
X3713 (Q–R). Scale bar: A, B, F–L, O, P, R = 500 μm; C = 300 μm; D = 100 μm; E, M, N, Q = 50 μm.

Missarzhevsky, 1966) or Tiksitheca korobovi (transferred
with question to a new genus Tiksitheca Missarzhevsky, in
Rozanov et al. 1969: 114–115). The species and Tiksitheca
in general were later reinterpreted as anabaritids (Missar-
zhevsky 1982). Qian & Bengtson (1989) placed Tiksitheca
in synonymy with Anabarites.
Investigation of internal moulds with parts of the wall
preserved in phosphate from the type locality (samples
106 and 106b) of Anabarites korobovi (Missarzhevsky, in
Rozanov et al., 1966; Fig. 37E–G) shows a close similarity to
forms recovered from the basal Erkeket Formation (samples
11/16.05 and 11/16.1; Fig. 37A–D, H–P). Both localities are
within the Dokidocyathus regularis Biozone.
Similar internal moulds of A. cf. korobovi are known
throughout the Siberian Platform, e.g. from older beds of
the upper Kessyusa Formation (samples 882 and 1487;
Fig. 38A–D, H–K, O). Internal moulds with a similar mor-
phology are found at localities in the Medvezhin Formation
 Systematics of Lower Cambrian fossil Anabaritids
Figure 39 Anabarites? volutus (Kugdatheca voluta Missarzhevsky,
in Rozanov et al., 1969). A–C, internal mould; sample GIN 3593/ 560
from the Kotuj River, with locality not specified; the specimen was
figured in Missarzhevsky (1989: pl. 12, fig. 11). A, lateral view. B, view
of the apertural end. C, view of the apical end. D, holotype of
Kugdatheca voluta (considered herein as Anabarites? volutus) with
the wall preserved; sample M311/3. E, fragment of Anabarites? licis
(Tiksitheca licis Missarzhevsky, in Rozanov et al., 1969) having its wall
with flanges preserved in celestite and barite; specimen SMNH X3410,
sample 92–2/26 (see more pictures in Kouchinsky & Bengtson 2002).
Scale bar: A–C = 500 μm; D = 3,3 mm; E = 200 μm.
along the Kotuj River, and have been described as Kotu-
ites biplicatus Missarzhevsky, 1989 (Fig. 38E–G, L–N, P–
R). The difference between K. biplicatus Missarzhevsky
and Tiksitheca korobovi (Misarzhevsky, in Rozanov & Mis-
sarzhevsky, 1966) is minimal and is probably preserva-
tional. The former is more flattened laterally, may show
two shallow and wide longitudinal depressions (Fig. 38G,
P), and has a convex lobe that is apparently more exten-
ded than in T. korobovi. At the early developmental stages
internal moulds of K. biplicatus have three weak grooves and
thus show a regular tri-fold symmetry, similar to that seen in
other species of Anabarites (Missarzhevsky, 1989: 194). At a
later stage K. biplicatus acquires an ovoid aperture, with one
of the lobes being more extended than the other two sides
283
Figure 40 Sample 1282. A–C, Kotujkanites vallatus Bokova, 1985;
holotype specimen. B–C, internal mould with a longitudinal twisted
ridge. A, apertural view. D, E, Kotujkanites sulcatus Bokova, 1985;
holotype specimen. E, internal mould with a single longitudinal
groove. D, apical view. Scale bar: A = 300 μm; B–E = 500 μm.
and curved in one plane. This variation may be sufficient to
recognise separate species, especially given the older occur-
rence of the type material of K. biplicatus. There appears to
be little reason, however, to assign these two forms to differ-
ent genera, since their bilateral symmetry can be interpreted
as a species-level distinction within the genus Anabarites
Missarzhevsky. A possible synonymy would favour Anabar-
ites korobovi (Missarzhevsky, in Rozanov & Missarzhevsky,
1966) as the senior synonym. Tiksitheca licis Missarzhevsky,
in Rozanov et al., 1969 differs in having a rounded triangular,
radially symmetrical cross-section, and in being irregularly
curved or almost straight, with the wall having straight and
rather prominent flanges and intervening growth lines (see
also remarks on genus Anabarites and Fig. 39E).
Kotuites biplicatus Missarzhevsky, 1989, is syn-
onymised herein with Anabarites bisulcatus Fedorov, in
Pel’man et al., 1990 (see Fig. 12C, F), from the southeastern
part of the Siberian Platform (Pel’man et al. 1990: 25–26).
Bilaterally symmetrical features are also known from other
anabaritids. These include a fourth dotted line of impres-
sions in Jakutiochrea portentosa Val’kov, 1987 (see remarks
on Anabarites tristichus and Fig. 28C–E), and a secondary
groove on one of the three lobes of Angustiochrea exima
Val’kov, 1989 [nomen nudum]. It appears that in the latter
two cases the bilateral symmetry merely represents intraspe-
cific variation. Two other forms with bilateral symmetry, but
no traces of the tri-fold symmetry, were formally described as
the anabaritids Kotujkanites sulcatus and K. vallatus (Fig. 40)
284 Artem Kouchinsky et al.

Figure 41 Anabarites rectus Vasil’eva, in Rudavskaya & Vasil’eva, 1984; internal moulds with phosphatised tubular remains of other
organisms inside; sample M302/1-2. SMNH X3714 (A–F) and SMNH X3715 (G–I). A, D–E, lateral views. B, close-up showing a possible aperture.
C, view of the cross-section at the end opposite to the probable aperture. F, close-up of E. H, lateral view. G, close-up of the upper part of H. I,
close-up of G. Scale bar: A, C–E, H = 500 μm; B = 200 μm; F = 100 μm; G = 300 μm; I = 150 μm.

by Bokova (1985). They do not seem to represent preserva-
tional variants, because several specimens are known from
the type locality. Their taxonomic position remains uncer-
tain, however. Forms described by Qian 1989 as Anabar-
ites sulcatus (Bokova, 1985) bear three longitudinal grooves,
making their identification in China problematical (those
are placed with question in synonymy with A. trisulcatus
herein).
Anabarites rectus Vasil’eva, in Rudavskaya &
Vasil’eva 1984 (Fig. 41)
1984 Anabarites rectus Vasil’eva; Rudavskaya & Vasil’eva:
1455, fig. 1.
1989 Anabarites rectus Vasil’eva (non sp. n.); Missar-
zhevsky: pl. 13(18).
MATERIAL. Two specimens from sample 302/1-2 (SMNH
X3714 & X3715).
OCCURRENCE. Dokidocyathus regularis Biozone of the
Tommotian Stage of the Siberian Platform.
DIAGNOSIS. Anabarites species characterised by internal
moulds with almost circular transverse profile and three con-
sistently preserved longitudinal bands of phosphatic material
attached.
DESCRIPTION. Straight internal moulds with rounded trian-
gular, almost circular cross-section (Fig. 41C). Three lon-
gitudinal bands of phosphatic material are attached to the
moulds (Fig. 41D–F). The funnel-shaped structure at the
end of one of the specimens (Fig. 41A–E) may represent an
aperture.
 Systematics of Lower Cambrian fossil Anabaritids 285

Figure 42 A–I, Cambrotubulus ex gr. decurvatus Missarzhevsky, in Rozanov et al., 1969. SMNH X3716 (A–B), SMNH X3717 (C–D), SMNH
X3718-X3722 (E–I). A, internal mould. B, close–up of the initial part; sample M321/34. D, internal mould of a juvenile specimen. C, close–up of
the initial part; sample 355a. E–G, internal moulds from sample 355a. H, internal mould from sample 907. I, internal mould from sample B-489.
J, holotype of Cambrotubulus conicus Missarzhevsky, 1989 (considered herein Cambrotubulus ex gr. decurvatus Missarzhevsky, in Rozanov
et al., 1969); sample M71-2/66. Scale bar: A, D–I = 500 μm; B, C = 200 μm; J = 1 mm.

Genus CAMBROTUBULUS Missarzhevsky, in
Rozanov et al., 1969
1967 Cambrotubulus Missarzhevsky: 20 [nomen nudum].
1969 Cambrotubulus Missarzhevsky; Rozanov et al.: 159.
?1979 Cambrotubulus Missarzhevsky; Qian et al.: 217.
1982 Cambrotubulus Missarzhevsky; Val’kov: 71–72.
1989 Cambrotubulus Missarzhevsky; Missarzhevsky: 189.
1996 Cambrotubulus Missarzhevsky; Esakova & Zhegallo:
94–95.
TYPE SPECIES. Cambrotubulus decurvatus Missarzhevsky,
in Rozanov et al. 1969 (pp. 160, pl. 7(5–7, 10)).
DIAGNOSIS. Tubes, or their internal moulds, variable in
shape and expansion rate, different from other anabaritids
in having a circular cross-section at every stage of growth,
but a similar microstructure in the wall and a tapered initial
part with open apex.
OCCURRENCE. Manykayan (Nemakit–Daldynian) and Tom-
motian Stages of the Siberian Platform; fossils assigned to
Cambrotubulus are reported from the Meishucunian and
Qiongzhusian Stages of the South China Platform, Lower
Cambrian of Western Gondwana, and lower Lower Cam-
brian of Western Mongolia.
REMARKS. Although a three-fold symmetry is missing, the
similarity of wall microstructures (Kouchinsky & Bengtson
2002) and initial parts (Missarzhevsky 1989; Fig. 42) to other
anabaritids supports the assignment of Cambrotubulus Mis-
sarzhevsky, in Rozanov et al., 1969 to this group. V.V. Mis-
sarzhevsky (in Rozanov et al. 1969: 159) described Cambro-
tubulus as irregularly curved tubes with a straight aperture
and circular cross-section. In addition (pp. 160) the tubes
show considerable morphological variation, including relat-
ively straight, slightly and strongly curved forms. All such
anabaritids with circular cross-sections are regarded herein as
Cambrotubulus, but they vary considerably in curvature, size
and expansion rate (samples 907, 355a, M71-2/62, M321/31,
B-489 and 1326). There are always transitional morphologies
in these samples, from straight and narrow to wider, twisted
and curved varieties (see Fig. 44, below).
Cambrotubulus and Conotheca Missarzhevsky, in Roz-
anov et al., 1969 can commonly be confused, especially when
their apical parts are not preserved (e.g. Qian & Bengtson
1989: fig. 85; Steiner et al. 2004, fig. 3(17)). Conotheca is
arbitrarily regarded within hyoliths (Rozanov et al. 1969:
112–113) and was originally described as a calcareous tu-
bular shell with a circular transverse cross-section, straight
aperture perpendicular to the longitudinal axis of the tube,
no internal septa and blunt initial part. Otherwise similar
tubes of Cambrotubulus have a tapered and open initial part.
Tubes of Conotheca circumflexa Missarzhevsky, in Rozanov
et al., 1969 from the type locality collected by V.V. Mis-
sarzhevsky have a wall microstructure that is different from
that in Cambrotubulus (see Fig. 45 herein; Kouchinsky 2000;
Feng et al. 2001; Kouchinsky & Bengtson 2002). The fossil
extracted using diluted acetic acid from carbonate sample
M31/46 demonstrates a relatively thick wall consisting of
two layers (see Fig. 45). The elongated units of the inner
layer run around the tube in a manner similar to that in
fossils described as hyoliths by Kouchinsky (2000) and Feng
et al. (2001) and not found in Cambrotubulus or other ana-
baritids (Kouchinsky & Bengtson 2002).
286 Artem Kouchinsky et al.
Figure 43 Cambrotubulus ex gr. decurvatus Missarzhevsky, in
Rozanov et al., 1969; sample 92-2/25. B, overall view of a theca
replaced by celestite and barite. A, close-up of the aperture. SMNH
X3723. C, another specimen preserved in celestite–barite (specimen
SMNH X3411, see Kouchinsky & Bengtson 2002). Scale bar:
A = 500 μm; B, C = 1 mm.
Cambrotubulus ex gr. decurvatus Missarzhevsky,
in Rozanov et al., 1969 (Figs 12B, 14M, 42–44)
1967 Cambrotubulus decurvatus Missarzhevsky: 20
[nomen nudum].
?1968 Platysolenites sibirica Val’kov: 116–117, figs 2–5.
1969 Cambrotubulus decurvatus Missarzhevsky; Rozanov
et al.: 160, pl. 7(5–7, 10).
1975 Cambrotubulus decurvatus Missarzhevsky; Mat-
thews & Missarzhevsky: pl. 2(6).
?1975 Cambrotubulus sibiricus (Val’kov); Val’kov: pl.
14(2–5).
1979 Cambrotubulus decurvatus Missarzhevsky; Qian
et al.: 217, pl. 2(13–16).
1982 Cambrotubulus decurvatus Missarzhevsky; Val’kov:
72, pl. 11(1–12).
?1982 Cambrotubulus sibiricus (Val’kov); Val’kov: 72–73.
1983 Cambrotubulus decurvatus Missarzhevsky; Sokolov
& Zhuravleva: 160, pl. 51(3–4).
1984 Cambrotubulus decurvatus Missarzhevsky; Chen:
56, pl. 1(2).
?1987 Cambrotubulus plicativus Val’kov: 110–111, pl.
13(19–21).
1989 Cambrotubulus decurvatus Missarzhevsky: pl. 13(9–
10).
1989 Cambrotubulus conicus Missarzhevsky: pl. 12(7).
1989 Cambrotubulus decurvatus Missarzhevsky; Hamdi
et al.: fig. 3e.
1990 Cambrotubulus crassus Fedorov; Pel’man et al.: 25,
pl. 2(1).
2002 Cambrotubulus conicus Missarzhevsky; Kouchinsky
& Bengtson: fig. 8A–D.
MATERIAL. Nine specimens from sample 907 (SMNH
X3721, X3740-X3747); 11 specimens from sample 355
(SMNH X3717-X3720, X3754-X3760); 6 specimens from
sample M71-2/62 (SMNH X3748-X3753); 5 specimens from
sample M321/31 (SMNH X3735-X3739); 1 specimen from
sample M321/34 (SMNH X3716); 6 specimens from sample
1326 (SMNH X3724-X3729); 2 specimens from sample 92-
2/25 (SMNH X3411 & X3723); 4 specimens designated as
holotypes of C. conicus Missarzhevsky, 1989, C. crassus
Fedorov, in Pel’man et al., 1990, C. plicativus Val’kov, 1987
and C. sibiricus (Val’kov, 1968) from samples and localities
indicated in Appendix 1 of Supplementary Material.
OCCURRENCE. As for the genus.
DESCRIPTION. Variably and irregularly curved internal
moulds (Fig. 44) with thin straight growth lines (Fig. 42).
Preserved thecae have circular straight aperture, tapered open
initial part and fine, straight growth lines (Figs 42 & 43).
Genus SELINDEOCHREA Val’kov, 1982
1982 Selindeochrea Val’kov: 68.
1989 Selindeochera Val’kov; Missarzhevsky: 192.
1989 Anabarites Missarzhevsky; Qian & Bengtson: 125
(pars).
1991 Anabaritellus Missarzhevsky; Landing & Murphy:
392 (pars).
1996 Anabarites Missarzhevsky; Esakova & Zhegallo: 90–
91 (pars).
TYPE SPECIES. Anabarites tricarinatus Missarzhevsky, in
Rozanov et al., 1969 (157–158, pls 8(13) and 16(5)).
DIAGNOSIS. Anabaritids characterised by internal moulds
and thecae with three prominent lobes. Prominent longitud-
inal keels produced by the walls extend from distal portions
of the lobes (see Fig. 2Y).
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
REMARKS. Selindeochrea differs from other anabaritids in
having diagnostic prominent longitudinal keels produced by
the wall at distal portions of the lobes. Those are rarely
preserved (see Fig. 48C) and better observed in thin sections
(Abaimova 1978; Val’kov 1987: pl. 13 (4–13); see Fig. 48H).
In mainland Nova Scotia limonitic internal moulds with
a variable number of longitudinal folds (or ridges) were
 Systematics of Lower Cambrian fossil Anabaritids 287

Figure 44 Cambrotubulus ex gr. decurvatus Missarzhevsky, in Rozanov et al., 1969; variation of internal moulds. SMNH X3724–X3760,
respectively, excluding the holotype, DG. AA–AF, sample 1326. BA–BE, sample B-489. CA–CE, sample M321/31. DA–DI, sample 907; DG,
holotype of Cambrotubulus plicativus Val’kov, 1987. EA–EF, sample M71–2/62. FA–FG, sample 355a. Scale bar = 1 mm.

attributed to Coleoloides trigeminatus Missarzhevsky, in
Rozanov et al., 1969 (Landing & Murphy 1991). Similar
moulds were found also in the Upper Member of the Arbuckle
Brook Formation, but referred to Anabaritellus tricarinatus.
The latter was synonymised by Landing & Murphy (1991)
with Anabaritellus hexasulcatus, Anabarites isiticus, Ana-
barites ternarius, Coleoloides trigeminatus and Selindeo-
chrea tecta. Except for such a doubtful report from Ava-
lonia (Landing & Murphy 1991), provisionally reinterpreted
herein as Coleoloides trigeminatus Missarzhevsky, no forms
with keels similar to Selindeochrea are known beyond the
Siberian Platform.
Coleoloides trigeminatus internal moulds have a circu-
lar transverse profile at the initial growth stage, but during the
following growth they acquire consequitively two, three, and
more, bifurcating longitudinal ridges (up to 16 in the Siberian
material) commonly running spirally around the longitudinal
axis of mould (Qian & Bengtson 1989; Landing & Murphy
1991; A.K., unpublished observations). Although Missar-
zhevsky (1982, 1989) indicated C. trigeminatus among
288 Artem Kouchinsky et al.
Figure 45 Conotheca circumflexa Missarzhevsky, in Rozanov et al.,
1969; sample M31/46, GIN 3481/75; locality M31 (Rozanov et al.
1969), lower reaches of the Lena River, near Chekurovka Village and
the mouth of the Biskeebit River; Middle Sub-Formation of the Tyuser
Formation, Member 7. Atdabanian Stage. A, general view. B, close-up
showing the outer and inner layers of the wall. C, close-up showing
elongated units of the inner layer running around the conch. Scale
bar: A = 400 μm; B = 60 μm; C = 30 μm.
anabaritids, it is not considered to be an anabaritid herein, due
to the unstable number of longitudinal ridges and lack of ra-
dial symmetry. The affinity of Coleoloides to any other group
of the earliest tubular fossils is problematic. Brasier (1984:
234) and Qian & Bengtson (1989: 130–131) synonymised C.
trigeminatus Missarzhevsky, 1969 with Coleoloides typicalis
Walcott, 1890, but this question awaits detailed investigation
of morphological variability.
Selindeochrea tecta Val’kov, 1982 (Fig. 46)
1982 Selindeochrea tecta Val’kov: 68, pls 8(18–27) and
9(1–2).
MATERIAL. Two specimens from sample 355 (SMNH
X3761 & X3762) and holotype.
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
DIAGNOSIS. Selindeochrea species characterised by internal
moulds that can be twisted through several revolutions, and
V-shaped grooves between lobes narrowing distally.
DESCRIPTION. Internal moulds twisted anticlockwise, hav-
ing three lobes with elongated transverse profiles narrowing
distally (Fig. 46A, C, E). The lobes are separated by grooves
that are V-shaped transversally (Fig. 46A, C) and become
less distinct towards the initial part (Fig. 46B). In larger spe-
cimens the grooves are W-shaped in cross-section, with a
central ridge carrying a median narrow groove (Fig. 46E).
Between the lobes growth lines are arched and produce acute
angles pointed towards the aperture (Fig. 46G).
REMARKS. In samples 96-5a/34.5 and 96-5a/34.75 glaucon-
itic internal moulds of S. cf. tecta occur (Fig. 47). They are
twisted clockwise through several revolutions in the oppos-
ite direction to that seen in S. tecta. In sample 806/3-4 S.
cf. tecta is also encountered with A. missarzhevsky and may,
therefore, represents a species different from S. tecta in be-
ing twisted clockwise. Moulds of S. tricarinata from other
samples are also twisted clockwise, but much less so than
in S. tecta. Microstructure and configuration of growth lines
in S. tecta, S. cf. tecta and S. tricarinata are otherwise very
similar (Figs 46–48).
Selindeochrea tricarinata (Missarzhevsky, in
Rozanov et al., 1969) (Fig. 48)
1967 Anabarites tricarinatus Missarzhevsky: 20 [nomen
nudum].
1969 Anabarites tricarinatus Missarzhevsky: Rozanov
et al., 1969 157–158, pls 8(13) and 16(5).
1982 Anabarites tricarinatus Missarzhevsky; Val’kov: 76,
pl. 12(22–28).
1989 Selindeochrea tricarinata (Missarzhevsky); Missar-
zhevsky: pl. 14(5–6).
2002 Anabarites tricarinatus Missarzhevsky; Kouchinsky
& Bengtson: fig. 9A, F–G.
MATERIAL. Three specimens, from sample 92-2/25 (SMNH
X3412, X3763 & X3764).
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
DIAGNOSIS. Selindeochrea species characterised by almost
straight or slightly twisted internal moulds with rounded
lobes and flattened depressions in between them.
DESCRIPTION. Internal moulds slightly twisted clockwise,
with triangular tri-lobate cross-sections (Fig. 48F, H)).
Grooves (depressions) between extended lobes are wide and
shallow with a flattened middle part (Fig. 48F–H). There is a
median narrow ridge on the flattened depressions between the
lobes (Fig. 48B, D–E). In the flattened depressions, growth
lines are arched and pointed at an acute angle towards the
aperture (Fig. 48B, E). In preserved thecae, the lobes are ex-
tended distally by prominent keels (Fig. 48C, H). The keels
undulate, with folds running towards the aperture at ca. 45◦
(Fig. 48D–E). Lobes of the internal moulds are rounded or
somewhat swelled and almost straight (Fig. 48F).
REMARKS. Selindeochrea tricarinata (Missarzhevsky), re-
ported from the lower part of the Emyaksin Formation of
northern Siberian Platform, was placed within the Dokido-
cyathus regularis Biozone of the Tommotian Stage (Roz-
anov et al. 1992: 100–101). Nevertheless, a pre-Tommotian
age (Manykayan Stage) for the lower Emyaksin Formation
is suggested by the biostratigraphy (Val’kov 1987) and con-
firmed by the chemostratigraphy (Kouchinsky et al. 2001;
see Fig. 4).
Problematic fossils resembling S. tricarinata (Missar-
zhevsky) from the upper Lower Cambrian of England and
misidentified as Anabarites compositus Missarzhevsky (Hinz
1987: 37–38, pl. 14) have not been examined herein and their
 Systematics of Lower Cambrian fossil Anabaritids 289

Figure 46 Selindeochrea tecta Val’kov, 1982; sample 355a. B, D, internal moulds, lateral views. A, C, apertural views. SMNH X3761 (A–B) and
SMNH X3762 (C–D). E–F, holotype specimen, internal mould. G, counterpart (cast of the outer wall) of the holotype (included in the original
description of the holotype by Val’kov (1982)). Scale bar: A, C, E = 300 μm; B, D, F = 500 μm; G = 200 μm.

affinity with anabaritids needs verification (see also Brasier
1989: 135).
Genus ACULEOCHREA Val’kov & Sysoev, 1970
1970 Aculeochrea Val’kov & Sysoev: 99.
1970 Longiochrea Val’kov & Sysoev: 95.
1982 Longiochrea Val’kov & Sysoev; Val’kov: 64–65.
1982 Aculeochrea Val’kov & Sysoev; Val’kov: 79.
1989 Longiochrea Val’kov & Sysoev; Missarzhevsky:
190.
1989 Aculeochrea Val’kov & Sysoev; Missarzhevsky:
190.
1989 Anabarites Missarzhevsky; Qian & Bengtson: 125
(pars).
TYPE SPECIES. Aculeochrea ornata Val’kov & Sysoev, 1970
(p. 99, pl. 1(10)).
DIAGNOSIS. Anabaritids characterised by internal moulds
with rounded triangular cross-sections, composed of seg-
ments produced by concentric folds. One or several tubercles
occur along lower margins of the folds. Each segment is sub-
divided by wide and shallow longitudinal grooves into three
parts.
OCCURRENCE. Lower Manykayan Stage of the Siberian
Platform.
REMARKS. Aculeochrea may be endemic to the Siberian
Platform and has not been reported from anywhere else.
Aculeochrea is known as internal moulds, sometimes with
remains of the wall. Well-preserved internal moulds bear
distinct concentric folds, producing overhangs with single or
multiple tubercles at their margins (Figs 49 & 50). Aculeo-
chrea ornata and A. rugosa (Val’kov & Sysoev: see below)
were figured upside down (i.e. opposite to the current recon-
struction of the direction of growth) in Val’kov & Sysoev
(1970: pl. 1(1 & 10)) and Val’kov (1975: pl. 13(1 & 10)).
290 Artem Kouchinsky et al.

Figure 47 Selindeochrea cf. tecta Val’kov, 1982; sample 96–5a/34.75. SMNH X3778 (A–E), SMNH X3779 (F–G), SMNH X3780 (H–I). C, internal
mould with remains of phosphatised wall. B, D, E, close-ups of C showing fibrous composition of the phosphatised wall. A, view of the apertural
end. G, internal mould. F, view of the apertural end. H, internal mould. I, view of the apical end of the mould. Scale bar: A, F, I = 300 μm;
B = 50 μm; C, G, H = 500 μm; D, E = 200 μm.

The only difference between the monospecific Longiochrea
Val’kov & Sysoev, 1970 and Aculeochrea Val’kov & Sysoev,
1970 appears to be the larger size of the former (as described
in Val’kov & Sysoev, 1970) and a single tubercle in Aculeo-
chrea against several on the lobes of Longiochrea (see be-
low). These features are considered insufficient to separate
two genera and, therefore, we have chosen to synonymise
Longiochrea with Aculeochrea.
Aculeochrea ornata Val’kov & Sysoev, 1970
(Figs 14I–L & 49)
1970 Aculeochrea ornata Val’kov & Sysoev: 99, pl.
1(10).
1975 Aculeochrea ornata Val’kov & Sysoev; Val’kov: pl.
13(10).
1982 Aculeochrea ornata Val’kov & Sysoev; Val’kov: 79,
pl. 13(22–23).
1989 Aculeochrea ornata Val’kov & Sysoev; Missar-
zhevsky: pl. 13(12).
MATERIAL. Three specimens from sample B-489 (SMNH
X3765-X3767) and holotype.
OCCURRENCE. As for the genus.
DIAGNOSIS. Aculeochrea species having a single tubercle
in the lower middle part of every fold on each lobe
(Fig. 49).
DESCRIPTION. Straight internal moulds, rounded triangular
in cross-section. A succession of folds runs around the mould
and three longitudinal broad depressions subdivide the mould
into three parts (lobes) (Fig. 49B). Each of the folds bears a
single tubercle in the lower middle part of every fold on each
lobe (Fig. 49A, C–E). In the plan view the folds may not be
strictly perpendicular to the axis (Fig. 49A, D). Thin casts of
growth lines occur on their surface parallel to the edges of
the rings (Fig. 49C, E).
Aculeochrea rugosa (Val’kov & Sysoev, 1970)
(Fig. 50)
1970 Longiochrea rugosa Val’kov & Sysoev: 95, pl. 1(1).
1975 Longiochrea rugosa Val’kov & Sysoev; Val’kov: pl.
13(1).
1982 Longiochrea rugosa Val’kov & Sysoev; Val’kov:
65.
 Systematics of Lower Cambrian fossil Anabaritids 291

Figure 48 Selindeochrea tricarinata (Anabarites tricarinatus Missarzhevsky, in Rozanov et al., 1969); sample 92–2/25. C, internal mould with
remains of the outer phosphatic crust (arrow), which covered the now-dissolved theca and prominent longitudinal keels. D, counterpart of the
same specimen, cast of the outer surface. B, E, close-ups of D showing growth lines curved upward, towards the aperture. A, close-up of B.
SMNH X3412. F, cross-section of an internal mould, SMNH X3763. G, lateral view of the same internal mould. H, polished cross-section of
possibly the same species from sample M321/31, SMNH X3764. Scale bar: A = 50 μm; B, E = 200 μm; C, D, F, G = 500 μm; H = 100 μm.

1989 Longiochrea rugosa Val’kov & Sysoev; Missar-
zhevsky: pl. 13(15).
MATERIAL. Two specimens from sample 1404 (SMNH
X3769 & X3772), 2 specimens from sample 96-4/4 (SMNH
X3768 & X3773) and 2 specimens from sample 1326
(SMNH X3770 & X3771).
pressions of internal tooth-like protrusions in three shal-
low grooves between the lobes, one per fold per groove
(Fig. 50B, D-E).
REMARKS. Some weathered internal moulds of A. rugosa
have notches in the longitudinal depressions (Fig. 50E),
similar to those in Anabarites tristichus Missarzhevsky.

OCCURRENCE. As for the genus.

Genus MARIOCHREA Val’kov, 1982
DIAGNOSIS. Aculeochrea species having multiple tubercles
along lower margin of every fold (Fig. 50).
1982 Mariochrea Val’kov: 69.
DESCRIPTION. Almost straight internal moulds, rounded tri- 1982 Gastreochrea Val’kov: 70.
angular in cross-section. A succession of folds runs around 1989 Mariochrea Val’kov; Missarzhevsky: 190.
the mould and three longitudinal broad depressions sub- 1989 Anabarites Missarzhevsky; Qian & Bengtson: 125
divide the mould into three parts (lobes) (Fig. 50D, G). (pars).
Each of the folds bears several tubercles along the lower
margin (Fig. 50A–D, G). In the plan view the folds may TYPE SPECIES. Mariochrea sinuosa Val’kov, 1982: 69–70,
not be strictly perpendicular to the axis (Fig. 50A–B, D, pls 10(1–2), 14(1–2), 15(1).
G). Thin casts of growth lines occur on their surface (Fig.
50D). No distinct folds are found at the apical portion of DIAGNOSIS. Anabaritids characterised by internal moulds
smaller specimens (Fig. 50A–B, G). There may be im- subdivided into stacked segments with rounded triangular
292 Artem Kouchinsky et al.

Figure 49 Aculeochrea ornata Val’kov & Sysoev, 1970; sample B-489. SMNH X3765 (A–C), SMNH X3766 (D), SMNH X3767 (E–F). A–B, different
views on the same internal mould. C, upper part of A enlarged. D, lateral view. E, lateral view of a specimen with remains of recrystallised wall,
enlarged in F (framed). Scale bar: A, D, E = 500 μm; B = 300 μm; C, F = 200 μm.

transverse profile and edges perpendicular to the growth axis.
The upper face of each segment shows a cast of a concentric
invagination of the wall with a central rounded-triangular or
clover-leaf-shaped aperture (Figs 2Z & 51).
OCCURRENCE. Upper Manykayan Stage of the Siberian
Platform.
an approach to a hexasulcate shape (Val’kov, 1982). Mario-
chrea is assigned to anabaritids on account of its three-fold
apertural opening. The appearance of the internal moulds in
comparison with other anabaritids is very distinctive. Mario-
chrea may be endemic and has not been reported beyond the
Siberian Platform.

REMARKS. The larger specimens referred to as Gastreochrea
viva Val’kov, 1982 are more likely to be later developmental
stages of M. sinuosa. These two forms are otherwise insigni-
ficantly different in terms of the shape of the apertural open-
ing, which shows a secondary invagination of the lobes and
Mariochrea sinuosa Val’kov, 1982 (Fig. 51)
1982 Mariochrea sinuosa Val’kov: 69–70, pls 10(1–2),
14(1–2), 15(1).
1982 Gastreochrea viva Val’kov: 70–71, pls 10(9), 14(1).
 Systematics of Lower Cambrian fossil Anabaritids 293

Figure 50 Aculeochrea rugosa (Longiochrea rugosa Val’kov & Sysoev, 1970); internal moulds in lateral views. SMNH X3768 (A–B), SMNH
X3769-X3773 (C–G respectively). A, internal mould with remains of the outer phosphatic crust (arrowed). B, close-up of A with notches (arrows);
sample 96–4/4. C, fragment of an internal mould; sample 1404. D, fragment of an internal mould with ring-like folds and notches; sample 1326.
E, fragment of an internal mould with notches; sample 1326. F, internal mould with remains of phosphatic outer crust (arrows); sample 1404. G,
internal mould; sample 96–4/4. Scale bar: A, C–G = 500 μm; B = 200 μm.

MATERIAL. Five specimens from sample 355 (SMNH Acknowledgements

X3774-X3777).
OCCURRENCE. As for the genus.
DESCRIPTION. Internal moulds and thecae straight or
slightly curved with rounded triangular profiles. The moulds
consist of segments (or rings) with edges perpendicular to
the growth axis and ornamented with tubercles along their
upper edges (Fig. 51A–B, F–I). The face of the upper ring
reflects invagination of the wall with a rounded triangular or
clover-leaf-like elevated opening in the middle (Fig. 51A–
B, F–G). The segments are covered with thin growth lines
parallel to the edges (Fig. 51B, H). Smaller phosphatised
thecae are composed of stacked cone-in-cone structures rep-
resenting successive growth stages of the apertural flange
(Fig. 51C–E). Unlike bigger specimens, they do not
display an invagination of the wall at the aperture
(Fig. 51C).
Artem Kouchinsky’s work was initially supported by his PhD
stipendium from the Swedish Academy of Science in 1997–
2001 and later from the NASA Astrobiology Institute and
post-doctoral grant (623-2003-207) from the Swedish Re-
search Council. A. Kouchinsky is currently supported by
the NordCEE (Nordic Center for Earth Evolution) project
(Danish National Research Foundation (Danmarks Grund-
forskningsfond)) grant to Prof. Donald Canfield. Stefan
Bengtson was supported by the Swedish Research Coun-
cil (grant no. 621-2001-1751). Feng Weimin acknowledges
support for the SEM work on the specimens from China
by the Chinese Major Basic Research Projects of Ministry
of Science and Technology, China (G2000077700). The au-
thors also acknowledge use of material collected by Anna
Bokova, Vladimir Missarzhevsky and Alexander Fedorov,
as well as kind help in obtaining specimens and valuable in-
formation from Igor Korovnikov, Olaf Eliki and Christian
294 Artem Kouchinsky et al.

Figure 51 Mariochrea sinuosa Val’kov, 1982; sample 355a. SMNH X3774 (A–B), SMNH X3775 (C–D), SMNH X3776 (E), SMNH X3777 (F–G). A,
F, views of the apertural ends of internal moulds B and G, respectively. D–E, smaller phosphatised thecae. C, apertural view of D. H, internal
mould with remains of phosphatised wall at the apical end (arrowed); holotype specimen figured in Val’kov 1982: pl. 10, figs. 1–2; pl. 14, fig. 1. I,
enlargement of the arrowed area. Scale bar: A–H = 500 μm; I = 200 μm.

Skovsted. We also wish to thank three anonymous re-
viewers, who contributed much to the improvement of the
manuscript.
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 APPENDIX 1

The appendix contains a list of species and genera assigned to anabaritids in the present study
with the originally given name and occurrence, as well as available information on other
forms of anabaritids reported from the same sample. Sample numbers from localities listed in
Appendix 2 are given in bold, and in this case the localities are not additionally specified
herein. Institutional names and abbreviations for the material discussed are: Institute of
Diamond and Precious Metal Geology, Russian Academy of Sciences (Siberian Branch),
Yakutsk, under either of the following three original abbreviations (in Russian) “ЯИГН”
(Yakutian Institute of Geological Sciences) replaced herein for convenience with YaIGN,
“ЯФАН” (Yakutian Filial of the Academy of Sciences) replaced with YaFAN, and “ЯНЦ”
(Yakutian Science Centre) replaced with YaSC; Museum of Geology of the Central Siberia,
Siberian Branch of the Russian Academy of Sciences, Novosibirsk, with “ЦСГМ” substituted
by CSGM; Siberian Institute of Geology, Geophysics and Mineral Resources, Novosibirsk,
with “СНИИГГиМС” referred herein as SNIIGGiMS; All-Russia Petroleum Research
Exploration Institute, St. Petersburg, with “ВНИГРИ” replaced by VNIGRI; and Geological
Institute of the Russian Academy of Sciences, Moscow, with original “ГИН” changed to
GIN. Some of the specimens are stored at the Swedish Museum of Natural History,
Stockholm (SMNH) and can be labeled with SMNH numbers. Other places of storage are
discussed in the following text. If a holotype is not found anywhere, it is referred to as
“holotype missing”.

Aculeochrea Val’kov & Sysoev, 1970.

Type species - Aculeochrea ornata Val’kov & Sysoev, 1970 (p.99, pl. 1(10)).

A. ornata Val’kov & Sysoev, 1970.

YaIGN 147/10, sample 1326. Forms reported from the sample: Aculeochrea ornata,
Anabarites tristichus, A. trisulcatus, Angustiochrea lata, Cambrotubulus decurvatus,
Jakutiochrea convexa, Lobiochrea natella, Longiochrea rugosa (Val’kov, 1975, p. 18-19 and
AK personal observations).

Anabaritellus Missarzhevsky, 1974.

Type species - Anabaritellus hexasulcatus Missarzhevsky, 1974 (p.187, pl. 23(6-7)).
A. cylindriculus Qian & Ding, 1988 in Ding & Qian 1988.
Specimen 84960, Lower Member (“Nisha Member”) of the Yangjiaping Formation,
Yangjiaping, Shimen, Hunan, China; Protohertzina unguliformis – Kaiyangites multispinatus
assemblage zone of the Lower Meishucunian Stage.

A. hariolus Val’kov, 1987.

YaIGN 164/39, sample 862, Erkeket Formation, Allatheca anabarica Biozone, Olenyok
River, left bank, at the mouth of the Bejenchime Brook. Holotype represented by thin section
stored in Yakutsk (Russia) has not been found (RK personal observation).

A. hexasulcatus Missarzhevsky, 1974.

GIN 4287/8, sample M71-2/69 (indicated in V.V. Missarzhevsky’s collection as M74/92).
Stored at SMNH. Forms reported from the sample: Anabaritellus hexasulcatus, Anabarites
kelleri, A. tricarinatus, A. tripartitus, A. tristichus, Angustiochrea magna [nomen nudum],
Cambrotubulus decurvatus (AK personal observations).

A. xishuiensis Qian, 1977.

Specimen 33767 from sample Z001, Yuhucun Formation, Meishucunian Stage, Xishui,
Guizhou Province, China. Holotype is housed at the Nanjing Institute of Geology and
Palaeontology, Chinese Academy of Sciences.

Anabarites Missarzhevsky, 1969 in Voronova & Missarzhevsky 1969 (also indicated as gen.
nov. in Rozanov et al. 1969, p. 155).
Type species - Anabarites trisulcatus Missarzhevsky, 1969 (p. 210, figs 1, 8, 9).

A. acuminatus Chen & Xiong, 1984 in Xing et al. 1984.

Tianzhushan Member of Dengying Formation, Anabarites – Circotheca – Protohertzina
assemblage zone, Tianzhushan of Yichang, Hubei Province, China.

A. alvenosulcatus Bokova, 1992 [nomen nudum].

YaIGN 169/23-5a, sample B-337. Stored at SMNH. Forms reported from the sample:
Anabarites alvenosulcatus [nomen nudum], Angustiochrea magna [nomen nudum],
Cambrotubulus decurvatus, C. plicativus, Jakutiochrea valkovi, Tiksitheca korobovi, T. licis
(Bokova, 1992 and personal observations).
A. bisulcatus Fedorov, 1990 in Pel’man et al. 1990.
CSGM 907/31, sample 3-2d'''' (=3-2д'''' in Russian), Pestrotsvet Formation, beds with
Archaeospira ornata - Bemella jacutica; Dzhanda River, right bank, 100 m downstream the
Kybyty Brook, section 3. Forms reported from the sample: Anabarites bisulcatus, A. signatus,
A. ternarius, A. trisulcatus, Cambrotubulus crassus, C. decurvatus, Tiksitheca sp., Udzhaites
sp. (Pel’man et al. 1990, p. 7).

A. compositus Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/126, sample M314/12. Holotype is missing. Forms reported from the sample:
Anabarites compositus, A. ternarius, A. tricarinatus, A. tristichus (personal observations).

A. gracilis Chen, 1984.

Meishucunian Stage, upper part; Jijiapo, Yichang, West Hubei, China.

A. grandis Val’kov, 1982.

YaFAN 159/33, Ust’-Yudoma Formation, Anabarites trisulcatus Biozone (indicated as
Angustiochrea lata Biozone in Val’kov, 1982); Allakh-Yun’ River, opposite to the mouth of
the Malaya Sakhara River, Uchur-Maya region, southeastern Siberian Platform. Holotype is
missing (RK personal observation).

A. imperceptus Bokova, 1992 [nomen nudum].

YaIGN 160/24-2a, sample B-102, Pestrotsvet Formation, “beds with Anabarites tribaculatus”
[nomen nudum] in Bokova, 1992; Selinde River (upper reaches of the Uchur River),
southeastern Siberian Platform, collected by A. Bokova. Stored at SMNH. Forms reported
from the sample: Anabarites imperceptus [nomen nudum], A. tricarinatus, Cambrotubulus
plicativus, Mariochrea sp. (Bokova, 1992 and AK personal observations).

A. isiticus Missarzhevsky, 1974.

GIN 4287/7 (indicated as GIN 3593/561 in Missarzhevsky, 1989), sample 2088/41 (locality
indicated as 2028 in Rozanov et al. 1969), Pestrotzvet Formation, Dokidocyathus regularis
Biozone (Lapworthella tortuosa Subzone), Lena River, middle reaches, opposite the village
Isit’. Holotype is missing.
A. kelleri Missarzhevsky, 1989.
GIN 3593/563, sample M421/94, Medvezhin Formation, Aldanella crassa Biozone; Kotuj
River, at the mouth of the Kotujkan River. Holotype is missing. Forms reported from the
sample: Anabarites kelleri, Cambrotubulus conicus, Cambrotubulus cf. decurvatus,
Tiksitheca licis (AK personal observations).

A. longissimoides Chen & Xiong, 1984 in Xing et al. 1984.

Upper part of Tianzhushan Member of Dengying Formation, Anabarites – Circotheca –
Protohertzina assemblage zone, Taishanmiao of Yichang, Hubei Province, China.

A. modestus Bokova, 1985.

YaFAN 166/5a, sample 1282. Stored at SMNH. Forms reported from the sample: Anabarites
modestus, A. signatus, A. tripartitus, A. trisulcatus, Angustiochrea lata, Cambrotubulus
decurvatus, C. sibiricus, Kotujkanites sulcatus, K. vallatus, Tiksitheca sp. (Bokova, 1985, p.
14 and AK personal observations).

A. obliquasulcatus Qian, 1978.

Specimen 33692 from sample Ningkuan-1, Kuanchuangpu Formation, Circotheca –
Tiksitheca – Anabarites – Protohertzina assemblage zone of the Meishucunian Stage,
Ningqiang, Shaanxi Province, China. Holotype is housed at the Nanjing Institute of Geology
and Palaeontology, Chinese Academy of Sciences.

A. primitivus Qian & Jiang, 1980 in Luo et al. 1982 (p. 172, pl. 14: 10).
Base of the Xiaowaitoushan Member of the Yuhucun Formation, Anabarites primitivus
Biozone, Beideng of Anning, Yunnan Province, China. Holotype is housed at the Yunnan
Institute of Geological Sciences.

A. rectus Vasil’eva, 1984 in Rudavskaya & Vasil’eva, 1984.

VNIGRI 732/13, Tyusser Formation, Dokidocyathus regularis Biozone, Lapworthella
tortuosa Subzone; lower Lena River, left bank, 4 km downstream the Chekurovka village,
mouth of the Biskeebit Brook. Specimens from sample 302/1-2 were indicated by mistake
“sp. n.” in Missarzhevsky (1989, p. 227), with specimen number GIN 3593/508.
A. rotundum Qian, 1977.
Specimen 33770 from sample Ningkuan-1, Kuanchuangpu Formation, Meishucunian Stage,
Ningqiang, Shaanxi Province, China. Holotype is housed at the Nanjing Institute of Geology
and Palaeontology, Chinese Academy of Sciences.

A. signatus Mambetov, 1981 in Missarzhevsky & Mambetov, 1981.

Collection number 22/1, sample M572, Chulaktau Formation, Karatau Member,
Pseudorthotheca costata Biozone; Berkuty section, Maly Karatau Range. Holotype is stored,
as indicated by Missarzhevsky & Mambetov (1981), at the Geological Institute of the
Academy of Sciences of Kirgiz SSR (presently National Academy of Sciences of Kirgizstan).

A. sexalox Conway Morris & Bengtson, 1990 in Bengtson et al. 1990.

SAMP30823 from sample UNEL1856, Parara Limestone, Horse Gully, South Australia.
Forms reported from the sample: Anabarites sexalox, A. trymatus (Bengtson et al. 1990, p.
195).

A. sulcoconvex Qian, 1978.

Specimen 33696, Kuanchuanpu Formation, Circotheca – Tiksitheca – Anabarites –
Protohertzina Biozone; Kuanchuanpu of Ningqiang, Shaanxi Province, China. Holotype is
housed at the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences.

A. tenuistriatus Qian, 1989.

Specimen 89424, Huangshadong Member, Tongying Formation, Qiaowan, Shipai, Yichang,
Hubei Province, China. Holotype is housed at the Nanjing Institute of Geology and
Palaeontology, Chinese Academy of Sciences.

A. ternarius Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/127, sample M314/7. Holotype is missing. Forms reported from the sample:
Anabarites ternarius, A. tristichus (AK personal observations).

A. tribaculatus Bokova, 1992 [nomen nudum].

YaIGN 169/19-8a, sample B-335. The holotype is missing, but a possible fragment of it has
been found in sample B-335b. Forms reported from the sample: Anabaritellus hexasulcatus,
Anabarites imperceptus [nomen nudum], A. tribaculatus [nomen nudum], Angustiochrea
aspera, A. magna [nomen nudum], Cambrotubulus decurvatus, C. plicativus, C. sibiricus,
Jakutiochrea solita, J. valkovi, Tiksitheca korobovi, T. licis, Udzhaites missarzhevskyi
(Bokova, 1992, p. 27 and AK personal observations).

A. tricarinatus Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/122, sample M321/31. Holotype is missing. Forms reported from the sample: A.
kelleri, Anabarites ternarius, A. tricarinatus, A. tripartitus, A. tristichus, Cambrotubulus
conicus, C. decurvatus (Rozanov et al. 1969 and AK personal observations).

A. tripartitus Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/123, sample 316/4. Holotype is missing.

A. tristichus Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/119, sample M321/31. Holotype is missing. Forms reported from the sample: A.
kelleri, A. tricarinatus, A. tripartitus, A. tristichus, Cambrotubulus conicus, C. decurvatus,
Tiksitheca cf. licis (Rozanov et al. 1969 and AK personal observations).

A. trisulcatus Missarzhevsky, 1969 in Voronova & Missarzhevsky 1969.

GIN 3481/405 (indicated as 3593/116 in Rozanov et al. 1969), sample M419/12.
Holotype is missing.

A. trymatus Conway Morris & Bengtson, 1990, in Bengtson et al. 1990.

SAMP30816 from sample UNEL1852, Parara Limestone, Curramulka, South Australia.
Forms reported from the sample: Anabarites sexalox, A. trymatus (Bengtson et al. 1990, p.
195).

A. undulatus Qian, 1978.

Specimen 33698, the Phosphate Bed of the Meishucun Formation, Circotheca – Tiksitheca –
Anabarites – Protohertzina Biozone. Kunyang, Yunnan Province, China. Holotype is housed
at the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences.

A. valkovi Fedorov, 1990 in Pel’man et al. 1990.

CSGM 907/34, sample 17-3, Ust’-Yudoma Formation, “Beds with abundant
angustiochreids”, Dzhanda River, right bank, 2.2 km upstream the Uesya-Kymystan Brook.
Forms reported from the sample: Anabarites bisulcatus, A. signatus, A. tristichus, A.
trisulcatus, A. valkovi, Angustiochrea lata, Cambrotubulus decurvatus, C. plicativus,
?Jakutiochrea convexa, Lobiochrea sp., Longiochrea sp., Sexangulatus denudatus (Pel’man
et al. 1990, p. 13).

Angustiochrea Val’kov & Sysoev, 1970.

Type species - A. lata Val’kov & Sysoev, 1970 (p. 98, pl.1(6-7)).

A. aspera Vasil’eva, 1990 in Bokova & Vasil’eva 1990.

VNIGRI 732/96-1, Kessyusa Formation, lower Allatheca cana Biozone (Bokova & Vasil’eva
1990); Olenyok River, 6 km downstream the Chuskuna Brook.

A. exima Val’kov, 1989 [nomen nudum].

Upper Kessyusa Formation, Anabarella plana Biozone, Olenyok River, 8 km upstream of the
Boroulakh Brook.

A. lata Val’kov & Sysoev, 1970.

YaFAN 147/6, sample 1326. Forms reported from the sample: Aculeochrea ornata,
Anabarites tristichus, A. trisulcatus, Angustiochrea lata, Cambrotubulus decurvatus,
Jakutiochrea convexa, Lobiochrea natella, Longiochrea rugosa (Val’kov, 1975, p. 18-19 and
AK personal observations).

A. magna Bokova, 1992 [nomen nudum].

YaIGN 169/25-1a, sample B-339. Stored at SMNH. Forms reported from the sample:
Angustiochrea magna [nomen nudum], Cambrotubulus decurvatus, Jakutiochrea sp.,
Tiksitheca korobovi, T. licis (AK personal observations).

A. rara Fedorov, 1984

Collection № 1603/5, SNIIGGiMS specimen 1, sample ShB-182/II-18 (=ШБ-182/II-18, in
Russian), Nemakit-Daldyn Formation, 11,5 m below the top, upper Nemakit-Daldynian
Horizon; Kotujkan River, right bank, 3 km upstream the mouth. Presence of the holotype in
the indicated place of storage is not confirmed. Forms reported from the sample:
“Angustiochreidae fam. indet.” (Fedorov, 1984).
Angustites Bokova, 1990 [nomen nudum].
Type species – A. trapezialis Bokova, 1992 (p. 96-97, pl.2(1-2)).

A. orbicularis Bokova, 1990 [nomen nudum].

YaIGN 172/9-13a, sample B-489. Stored at SMNH. Forms reported from the sample:
Aculeochrea ornata, Anabaritellus sp., Anabarites signatus, A. trisulcatus, Angustiochrea
lata, A. magna [nomen nudum], Angustites orbicularis [nomen nudum], A. trapezialis [nomen
nudum], Cambrotubulus decurvatus, Lobiochrea formosa [nomen nudum] (Bokova, 1992, p.
13 and AK personal observations).

A. trapezialis Bokova, 1990 [nomen nudum].

YaIGN 172/9-14a, sample B-489. Stored at SMNH. Forms reported from the sample:
Aculeochrea ornata, Anabaritellus sp., Anabarites signatus, A. trisulcatus, Angustiochrea
lata, A. magna [nomen nudum], Angustites orbicularis [nomen nudum], A. trapezialis [nomen
nudum], Cambrotubulus decurvatus, Lobiochrea formosa [nomen nudum] (Bokova 1992, p.
13 and AK personal observations).

Cambrotubulus Missarzhevsky, 1969.

Type species - Cambrotubulus decurvatus Missarzhevsky, 1969 (p. 160, pl. 7(5-7, 10)).

C. boroulachi Val’kov, 1989 [nomen nudum].

Kessyusa Formation, Anabarella plana Biozone, Olenyok River, right bank, 7 km upstream
of the Boroulakh Brook.

C. conicus Missarzhevsky, 1989.

GIN 3593/507 (indicated by V.V. Missarzhevsky in collection as GIN3593/566), sample
M71-2/66. Forms reported from the sample: Anabaritellus hexasulcatus, Cambrotubulus
conicus, Jakutiochrea sp., Tiksitheca cf. licis (AK personal observations).

C. corniformis Elicki, 1994.

FG 410/6, Ludwigsdorf Member, Ludwigsdorf quarry, Görlitz Syncline, Germany. Holotype
is stored at the Geological Institute of the Freiberg University. Forms reported from the
sample: Cambrotubulus decurvatus, C. corniformis, Tiksitheca korobovi (Elicki, 1994 and
personal communication by O. Elicki in 2002).
C. crassus Fedorov, 1990 in Pel’man et al. 1990.
CSGM 907/30, sample 3-3a, Pestrotzvet Formation, beds with Archaeospira ornata - Bemella
jacutica, Dzhanda River, right bank, 100 m downstream the Kybyty Brook. Forms reported
from the sample: Anabaritellus hexasulcatus, Anabarites bisulcatus, A. cf. tricarinatus, A.
tripartitus, A. trisulcatus, Cambrotubulus crassus, C. decurvatus, C. plicativus, Udzhaithes
missarzhevskyi (Pel’man et al. 1990, p. 7).

C. decurvatus Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/128, sample M410/73, Medvezhin Formation, Ajacicyathus sunnaginicus-
Tiksitheca licis Biozone, Kotuj River, mouth of the right tributary Ary-Mas-Yuryakh,
northwestern Anabar Uplift, Siberian Platform. Holotype is missing. Forms reported from the
sample: Cambrotubulus conicus, C. decurvatus, Tiksitheca licis (AK personal observations).

C. plicativus Val’kov, 1987.

YaFAN 164/44, sample 907. Forms reported from the sample: Anabaritellus hexasulcatus,
Anabarites ternarius, A. tricarinatus, A. tripartitus, A. tristichus, A. trisulcatus,
Angustiochrea magna, Cambrotubulus conicus, C. decurvatus, C. plicativus, C. sibiricus,
Jakutiochrea lenta, Tiksitheca cf. licis (Val’kov 1987, p. 32-33 and AK personal
observations).

C. sibiricus (Val’kov, 1968) (=Platysolenites sibiricus Val’kov, 1968).

YaFAN 82/16, Kessyusa Formation, Oelandiella korobkovi - Anabarella plana Biozone
(Val’kov, 1982); Olenyok River, left bank, 6 km downstream the Chuskuna Brook.

Gastreochrea Val’kov, 1982.

Type species - Gastreochrea viva Val’kov, 1982 (p. 70-71, pls 10 (9) and 16 (1)).

G. viva Val’kov, 1982.

YaFAN 159/111, sample 355. Holotype is represented by a polished section. Stored at
SMNH. Forms reported from the sample: Anabaritellus hexasulcatus, Anabarites cf. kelleri,
A. ternarius, A. cf. tricarinatus, A. tripartitus, Cambrotubulus cf. conicus, C. decurvatus,
Gastreochrea viva, Jakutiochrea solita, Mariochrea sinuosa, Selindeochrea tecta (Val’kov,
1982 and AK personal observations).
Jakutiochrea Val’kov & Sysoev, 1970.
Type species – Jakutiochrea convexa Val’kov & Sysoev, 1970 (p. 98-99, pl. 1(8)).

J. convexa Val’kov & Sysoev, 1970.

YaFAN 147/8, sample 1326. Forms reported from the sample: Aculeochrea ornata,
Anabarites tristichus, A. trisulcatus, Angustiochrea lata, Cambrotubulus decurvatus,
Jakutiochrea convexa, Lobiochrea natella, Longiochrea rugosa (Val’kov, 1975, p. 18-19 and
AK personal observations).

J. lenta Bokova & Val’kov, 1987 in Val’kov, 1987.

YaFAN 164/61, sample 907. Forms reported from the sample: Anabaritellus hexasulcatus,
Anabarites ternarius, A. tricarinatus, A. tripartitus, A. tristichus, A. trisulcatus,
Angustiochrea cf. lata, Cambrotubulus conicus, C. decurvatus, C. plicativus, C. sibiricus,
Jakutiochrea lenta, Tiksitheca cf. licis (Val’kov, 1987, p. 32-33 and AK personal
observations).

J. portentosa Bokova & Val’kov, 1987 in Val’kov, 1987.

YaFAN 164/60, sample 769. Forms reported from the sample: Anabarites kelleri, A.
tricarinatus, Cambrotubulus conicus, Jakutiochrea portentosa, J. solita, Tiksitheca licis (AK
personal observations).

J. solita Val’kov, 1987.

YaFAN 164/51, sample 769. Forms reported from the sample: Anabarites kelleri, A.
tricarinatus, Cambrotubulus conicus, Jakutiochrea portentosa, J. solita, Tiksitheca licis (AK
personal observations).

J. valkovi Bokova, 1990 in Bokova & Vasil’eva 1990.

YaSC 169/9-13a, sample B-335. Holotype is missing. Forms reported from the sample:
Anabaritellus hexasulcatus, Anabarites imperceptus, A. tribaculatus [nomen nudum],
Angustiochrea aspera, A. magna [nomen nudum], Cambrotubulus decurvatus, C. plicativus,
C. sibiricus, Jakutiochrea solita, J. valkovi, Tiksitheca korobovi, T. licis, Udzhaites
missarzhevskyi (Bokova, 1992, p. 27 and AK personal observations).
Kotujkanites Bokova, 1985 (misspelt ”Kotyikanites” in Bokova, 1985).
Type species – K. sulcatus Bokova, 1985 (p. 16-17, pl. 1(2-3)).

K. sulcatus Bokova, 1985.

YaFAN 166/7a, sample 1282. Stored at SMNH. Forms reported from the sample: Anabarites
modestus, A. signatus, A. tripartitus, A. trisulcatus, Angustiochrea lata, Cambrotubulus
decurvatus, C. sibiricus, Kotujkanites sulcatus, K. vallatus, Tiksitheca sp. (Bokova, 1985, p.
14 and AK personal observations).

K. vallatus Bokova, 1985.

YaFAN 166/9a, sample 1282. Stored at SMNH. Forms reported from the sample: Anabarites
modestus, A. signatus, A. tripartitus, A. trisulcatus, Angustiochrea lata, Cambrotubulus
decurvatus, C. sibiricus, Kotujkanites sulcatus, K. vallatus, Tiksitheca sp. (Bokova, 1985, p.
14 and AK personal observations).

Kotuites Missarzhevsky, 1989.

Type species - K. biplicatus Missarzhevsky, 1989 (p.194, pl.13(11)).

K. biplicatus Missarzhevsky, 1989.

GIN 3593/521 - in text (Missarzhevsky, 1989), but GIN 3593/561 - in explanations to figures;
Medvezhin Formation, Anabarella plana Biozone; Kotuj River, locality not indicated.
Holotype is missing.

Kugdatheca Missarzhevsky, 1969.

Type species - K. voluta Missarzhevsky, 1969 in Rozanov et al. 1969 (p. 111-112, pls 1(1 and
10), 12(13), 14(1), and 16(2b)).

K. voluta Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/18, sample M311/3 (in caption to pl. 1 in Rozanov et al. 1969 (English edition) and
marked at the specimen in collection), Medvezhin Formation; age is indicated as “A.
sunnaginicus – T. licis Biozone”; Fomich River (no further information on locality M311 in
Rozanov et al. 1969). In description of the species the sample number is however M314/5-10
(Rozanov et al. 1969 (English edition), p. 134). Stored at SMNH.
Lobiochrea Val’kov & Sysoev, 1970.
Type species – L. natella Val’kov & Sysoev, 1970 (p.96, pl. 1 (2)).

L. formosa Bokova, 1992 [nomen nudum].

YaIGN 172/5-6a, sample B-483. Stored at SMNH. Forms reported from the sample:
Anabarites trisulcatus, Angustiochrea lata, Cambrotubulus decurvatus, Lobiochrea formosa
(Bokova, 1992 and AK personal observations).

L. natella Val’kov & Sysoev, 1970.

YaFAN 147/2, sample 1326. Forms reported from the sample: Aculeochrea ornata,
Anabarites tristichus, A. trisulcatus, Angustiochrea lata, Cambrotubulus decurvatus,
Jakutiochrea convexa, Lobiochrea natella, Longiochrea rugosa (Val’kov, 1975, p. 18-19 and
AK personal observations).

L. trialata Val’kov, 1989 [nomen nudum].

Pestrotsvet Formation, Allatheca cana Biozone; Selinde River (upper reaches of the Uchur
River), southeastern Siberian Platform, collected by A.K. Val’kov.

Longiochrea Val’kov & Sysoev, 1970.

Type species - Longiochrea rugosa Val’kov & Sysoev, 1970 (p. 95, pl. 1 (1)).

L. rugosa Val’kov & Sysoev, 1970.

YaFAN 149/1, sample 1326. Holotype is missing. Forms reported from the sample:
Aculeochrea ornata, Anabarites tristichus, A. trisulcatus, Angustiochrea lata, Cambrotubulus
decurvatus, Jakutiochrea convexa, Lobiochrea natella, Longiochrea rugosa (Val’kov, 1975,
p. 18-19 and AK personal observations).

Mariochrea Val’kov, 1982.

Type species - Mariochrea sinuosa Val’kov, 1982 (p. 69-70, pls 10 (1-2), 14 (1-2) and 15
(1)).

M. sinuosa Val’kov, 1982.

YaFAN 159/109, sample 355. Forms reported from the sample: Anabaritellus hexasulcatus,
Anabarites cf. kelleri, A. ternarius, A. cf. tricarinatus, A. tripartitus, Cambrotubulus cf.
conicus, C. decurvatus, Gastreochrea viva, Jakutiochrea solita, Mariochrea sinuosa,
Selindeochrea tecta (Val’kov, 1982, 1987 and AK personal observations).

Selindeochrea Val’kov, 1982.

Type species - Selindeochrea tecta Val’kov, 1982 (p. 68, pls 8 (18-27) and 9 (1-2)).

S. tecta Val’kov, 1982

YaFAN 159/114, sample 355. Forms reported from the sample: Anabaritellus hexasulcatus,
Anabarites cf. kelleri, A. ternarius, A. cf. tricarinatus, A. tripartitus, Cambrotubulus cf.
conicus, C. decurvatus, Gastreochrea viva, Jakutiochrea solita, Mariochrea sinuosa,
Selindeochrea tecta (Val’kov, 1982, 1987 and AK personal observations).

Sexangulatus Fedorov, 1990.

Type species - Sexangulatus denudatus Fedorov, 1990 in Pel’man et al. 1990 (p. 27, pl. 2 (9)).

S. denudatus Fedorov, 1990 in Pel’man et al. 1990.

CSGM 907/38, sample 17-3, Ust’-Yudoma Formation, “beds with abundant angustiochreids”;
Dzhanda River, right bank, 2.2 km upstream the Uesya-Kymystan Brook. Forms reported
from the sample: Anabarites bisulcatus, A. signatus, A. tristichus, A. trisulcatus, A. valkovi,
Angustiochrea lata, Cambrotubulus decurvatus, C. plicativus, Jakutiochrea? convexa,
Lobiochrea sp., Longiochrea sp., Sexangulatus denudatus (Pel’man et al. 1990, p. 13).

Tiksitheca Missarzhevsky, 1969.

Type species - Tiksitheca licis Missarzhevsky, 1969 (Rozanov et al. 1969, p. 114, pls 2(10b)
and 8(6 and 22)).

T. curvata Fedorov, 1990 in Pel’man et al. 1990.

CSGM 907/36, sample 3-2g (=3-2г), Pestrotzvet Formation, beds with Archaeospira ornata-
Bemella jacutica, Dzhanda River, right bank, 100 m downstream the Kybyty Brook. Forms
reported from the sample: Anabarites bisulcatus, A. signatus, A. tripartitus, A. tristichus, A.
trisulcatus, Cambrotubulus crassus, C. decurvatus, C. plicativus, Jakutiochrea lenta,
Tiksitheca curvata, T. cf. licis (Pel’man et al. 1990, p. 6).

T. fangxianensis Duan, 1983.

Specimen SH1050, Xihaoping Formation, Shennongjia District, Hubei Province, China.
Holotype is housed at the Tianjing Institute of Geology and Mineral Resources, Chinese
Academy of Geological Sciences.

T. huangshandongensis Qian, Chen & Chen, 1979 in Qian et al. 1979.

Specimen 51728, Huangshandong Formation, Paleosulcachites – Lenatheca – Zhijinites-
Heraultipegma- Sachites - Zeugites – Lapworthella Biozone; Yichang County, Hubei
Province, China. Holotype is housed at the Nanjing Institute of Geology and Palaeontology,
Chinese Academy of Sciences.

T. irregularis Qian & Ding, 1988 in Ding & Qian, 1988.

Specimen 84971, Lower Member (“Nisha Member”) of the Yangjiaping Formation,
Protohertzina unguliformis – Kaiyangites multispinatus assemblage zone, Yangjiaping,
Shimen, Hunan, China.

T. korobovi (Missarzhevsky, 1966) (=Semielliptotheca korobovi Missarzhevsky, 1966) in

Rozanov & Missarzhevsky 1966.
GIN 3470/66, sample 106 (collected by M.N.Korobov). Holotype is missing.

T. licis Missarzhevsky, 1969 in Rozanov et al. 1969.

GIN 3593/26, sample M423/13. Holotype is missing. Forms reported from the sample:
Anabarites kelleri, Cambrotubulus decurvatus, Tiksitheca licis (Rozanov et al. 1969, p. 41
and AK personal observations).

T. minuta Yue, 1984 in Xing et al. 1984 (p. 155, pl. 25, figs 36-37).
Specimen 81134, Dengying Formation, Member 7, Paragloborilus – Tiksitheca assemblage
zone; Yuanjiaping section, southwestern Shaanxi Province, China.

Trisulcatheca [nomen nudum] Zhong, 1977 (p. 119).

Udzhaites Vasil’eva, 1986.

Type species - U. missarzhevskyi Vasil'eva, 1986 (p.103-104, fig.1).

U. missarzhevskyi Vasil’eva, 1986.

VNIGRI 732/14, Emyaksin Formation, Anabarella plana - Nikatheca kengedeica Biozone;
Udzha River, right bank, 12.6 km downstream the Tokur-Udzha Brook. Forms reported from
the sample: Anabarites compositus, A. isiticus, A. signatus, A. ternarius, A. tripartitus,
Tiksitheca korobovi (Vasil’eva, 1986).
 APPENDIX 2

The appendix contains a list of Siberian samples examined by AK and referred to in this
work, with their localities and originally specified stratigraphic positions. The samples are
grouped into three sets representing in ascending order the lower and upper beds of the
Manykayan Stage and the Tommotian Stage, according to the authors’ present understanding
of the stratigraphy discussed in the text (see also Figs 3 & 4).

1. Lower Manykayan Stage

M419/12 and 92-1b/18

Locality M419 (Rozanov et al. 1969): right bank of the Kotujkan River, ca. 2 km from the
mouth, northwestern flanks of the Anabar uplift; M419/12 - collected by V.V. Missarzhevsky,
92-1b/18 - collected by A.V. Kouchinsky; Nemakit-Daldyn Formation, “Nemakit-Daldynian
Horizon” in Rozanov et al. 1969.

1404, 96-4/4, and 96-4/4(2)

Locality 96-4: Bol’shaya Kuonamka River, left bank, ca. 1 km upstream the mouth of the
Ulakhan-Tyulen Brook, northeastern flanks of the Anabar uplift (locality A-50 in Val’kov
1975); 1404 - collected by A.K. Val’kov; others - collected by A.V. Kouchinsky; Manykay
Formation, Angustiochrea lata Biozone in Val’kov, 1975.

1326
Malaya Kuonamka River, between the Yulegir and Chiekidimyl brooks, northern Siberian
Platform, collected by A.K. Val’kov; Manykay Formation, “Aculeochreidae” Biozone in
Val’kov & Sysoev, 1970.

B-483 and B-484

Locality 45 (Bokova 1992): Malaya Kuonamka River, 3 km straight from the mouth of the
Luchakan Brook, northern Siberian Platform, collected by A.R. Bokova; Manykay Formation,
“beds with Lobiochrea formosa” in Bokova, 1992.

B-489
Locality 47 (Bokova, 1992): Malaya Kuonamka River, 2.5 km straight from the mouth of the
Ongkhoj Brook, northern Siberian Platform, collected by A.R. Bokova; Manykay Formation,
“beds with Lobiochrea formosa” in Bokova, 1992.

A-380k
Left bank of the Aim River, downstream the Malyj Aim River; Ust’-Yudoma Formation,
Angustiochrea lata Biozone in Val’kov, 1987 (p. 36-37).

2. Upper Manykayan Stage

1282
Locality M419 (Rozanov et al. 1969; locality E-8 in Val’kov, 1975): right bank of the
Kotujkan River, ca. 2.5 km from the mouth, northwestern flanks of the Anabar uplift,
collected by A.K. Val’kov; Nemakit-Daldyn Formation, Angustiochrea lata Biozone in
Val’kov, 1975.

M423/13, 92-2/21, 92-2/25, and 92-2/26

Locality M423 (Rozanov et al. 1969): mouth of the Kugda Brook, on the right bank of the
Kotuj River, northeastern flanks of the Anabar uplift; M423/13 - collected by V.V.
Missarzhevsky; other samples - by A.V. Kouchinsky; Medvezhin Formation, Ajacicyathus
sunnaginicus - Tiksitheca licis Biozone in Rozanov et al. 1969.

M321/26, M321/31, and M321/34

Locality M321 (Rozanov et al. 1969): left bank of the Eriechka River, 1.5 km downstream the
Nemakit-Daldyn River, northwestern flanks of the Anabar uplift, collected by V.V.
Missarzhevsky; Medvezhin Formation, Ajacicyathus sunnaginicus - Tiksitheca licis Biozone
in Rozanov et al. 1969.

M314/5-10, M314/7, and M314/12

Locality M314 (Rozanov et al. 1969): right bank of the Fomich River, ca. 6 km upstream the
Afanas’evskie lakes, northwestern flanks of the Anabar Uplift (collected by V.V.
Missarzhevsky); Medvezhin Formation, Ajacicyathus sunnaginicus - Tiksitheca licis Biozone
in Rozanov et al. 1969.
769
Kengede River (left tributary of the Arga-Sala River), left bank, 1.8 km downstream the
Kharyjalakh Brook, collected by A.K. Val’kov; Emyaksin Formation, Allatheca cana
Biozone in Val’kov, 1987 (p. 24).

B-335, B-335b, B-337, and B-339

Locality 24 (Bokova 1992): Olenyok River, left bank, 6 km downstream the Chuskuna Brook,
collected by A.R. Bokova; Kessyusa Formation, beds with Anabarites tribaculatus in
Bokova, 1992.

907
Olenyok River, left bank, 6 km downstream the Chuskuna Brook, collected by A.K. Val’kov;
Kessyusa Formation, Anabarella plana Biozone in Val’kov 1987 (p. 32).

M71-2/62, M71-2/66, and M71-2/69

Locality M71-2: left bank of the Olenyok River, 6 km downstream the Chuskuna Brook,
collected by V.V. Missarzhevsky; Kessyusa Formation, Aldanocyathus sunnaginicus -
Tiksitheca licis Biozone in Missarzhevsky, 1974, but Anabarella plana Biozone in
Missarzhevsky, 1989.

882 and 1487

Olenyok River, between Torkukuj and Boroulakh rivers, 3.6 km upstream the Dyogurdaakh
River (section V-9 (=B-9) in Val’kov, 1975), collected by A.K. Val’kov; Kessyusa
Formation, Oelandiella korobkovi – Anabarella plana Biozone in Val’kov, 1975, but
Spinulitheca rotunda Biozone in Val’kov, 1987.

355 and 369

Selinde River (upper reaches of the Uchur River), southeastern Siberian Platform, collected
by A.K. Val’kov; Pestrotsvet Formation, Anabarella plana Biozone in Val’kov, 1982.

806/3-4
Locality 806 (Vasil’eva, 1985): Udzha River, right bank, 12.6 km downstream the Tokur-
Udzha Brook, northern Siberian Platform, collected by N.I. Vasil’eva; Emyaksin Formation,
Anabarella plana - Nikatheca kengedeica Biozone in Vasil’eva, 1986.
96-5a/34.5 and 96-5a/34.75
Locality 96-5a: right side of the mouth of the Ulakhan-Tyulen Brook, at the right bank of the
Bol’shaya Kuonamka River, northeastern flanks of the Anabar uplift, collected by A.V.
Kouchinsky; Emyaksin Formation, uppermost Allatheca cana Biozone of Val’kov, 1975,
1987.

? Upper Manykayan Stage

1733
Right bank of the Dzhanda River, 2 km downstream the Olom River, collected by A.K.
Val’kov; basalmost Pestrotsvet Formation, Anabarella plana Biozone (after A.K. Val’kov in
Khomentovsky, Val’kov & Karlova 1990).

316/4
Maya River, upstream of the mouth of the Inikan River, collected by M.A. Semikhatov; basal
Pestrotsvet Formation, Ajacicyathus sunnaginicus - Tiksitheca licis Biozone in Rozanov et al.
1969.

1571/2
Locality at the mouth of the right tributary of the Leningradskaya River, opposite the mouth
of the Stepanovo Uschel’e Brook, northeastern Tajmyr, collected by V.M. Lopatin in 1989;
2.5-4.5 m above the top of the Kolosovskaya Formation, Lower Cambrian.

3. Tommotian Stage

11/16.05 and 11/16.1

Khorbusuonka River, right bank, right of the mouth of the Yuesej-Yueteekh Brook, northern
Siberian Platform, collected by A.V. Kouchinsky; Erkeket Formation, lower Dokidocyathus
regularis Biozone.

106 and 106b

Lena River, lower reaches, left bank, near village Chekurovka, northern Siberian Platform,
collected by M.N. Korobov in 1962; Tyuser Formation, 0.7-1 m above the base of the Middle
Sub-Formation (the sample derives from Bed 3 of Rozanov et al. 1969, p. 34); Dokidocyathus
regularis Biozone (Rozanov et al. 1992).

M302/1-2
Locality M302 (Rozanov et al. 1969): right bank of the Lena River, to the right of the mouth
of the Tiktirikteekh Brook, collected by V.V. Missarzhevsky; Pestrotsvet Formation,
Dokidocyathus regularis Biozone (Rozanov et al. 1969).