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Cyclocrinitids from the Lower Palaeozoic Tethyan sequence of Spiti, India

Husain Shabbar a,b , Anju Saxena a,∗ , Kamal Jeet Singh a , Shreerup Goswami b
aBirbal Sahni Institute of Palaeosciences, Lucknow 226007, Uttar Pradesh, India
b Department of Earth Sciences, Sambalpur University, Sambalpur 768019, Odisha, India
Received 7 March 2019; received in revised form 17 June 2019; accepted 15 July 2019

Abstract
Well preserved cyclocrinitids (calcareous green algae) are reported from early Palaeozoic deposits of Spiti, Himachal Pradesh, India. Casts of the
algal fossils are preserved in calcareous grey siltstone unit of the Takche Formation (Late Ordovician–Early Silurian) which appears rusty/earthy
brown due to weathering. The assemblage includes Cyclocrinites favus, Cyclocrinites pyriformis, Cyclocrinites cf. welleri, Cyclocrinites sp. and
Cyclocrinites globosus. Both Cyclocrinites cf. welleri and Cyclocrinites globosus are reported for the first time from the entire Tethyan Himalaya,
India. The described algal flora basically denotes relatively shallow marine depositional setting and low to moderate hydrodynamic conditions.
© 2019 Elsevier Ireland Ltd Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. Published by Elsevier B.V. All rights reserved.

Keywords: Cyclocrinitids; Ordovician; Spiti; Takche Formation; Tethyan Himalaya

1. Introduction by Kato et al. (1987), Maithy et al. (1999), Hubmann and

Spiti Valley is a remote part of Himachal Pradesh in the
North-Western Himalaya and offers one of the most complete
and spectacular rock exposures in the world. This basin along
with Zanskar Basin constitutes the largest marine sub-basin of
the Tethyan Himalayan Master Basin (Fig. 1) and preserves an
excellent and complete succession of rocks from Neoproterozoic
to Cretaceous ages. The contemporaneous Palaeozoic sedi-
mentary successions are exposed in the Ladakh–Kashmir, and
Kinnaur–Uttarakhand Himalayas. The Spiti–Zanskar sequences
have been extensively studied by a number of workers for their
geological setting, depositional history, and faunal biodiversity
(Stoliczka, 1865; Griesbach, 1889; Hayden, 1904; Reed, 1910,
1912; Srikantia et al., 1977; Srikantia, 1981; Bhargava and Bassi,
1998; Draganits et al., 2002; Bhargava, 2008).
Although, a number of studies have recorded faunal biodiver-
sity from the Early Palaeozoic sequences (Cambrian–Devonian)
of Spiti; the studies pertaining to plant remains (whether
marine or terrestrial) are meagre. So far, the plant remains
of marine affinity (algae) from Spiti Valley are reported
∗ Johnson and Konishi (1959), Osgood and Fischer (1960) and
Corresponding author.
E-mail address: anju saxena@bsip.res.in (A. Saxena).
Suttner (2007), Pandey and Parcha (2018), and very recently,
warm-water dasycladaceae algae are reported from the Takche
Formation, Gechang village, Parahio Valley (Chaubey et al.,
2019). Algal remains of marine affinity are also reported from
Ordovician–Silurian sequences of Kinnaur–Uttarakhand Basin
(Bhargava and Bassi, 1987; Sinha and Trampisch, 2013).
The cyclocrinitids, an extinct algal tribe, has a long history
of its doubtful affiliation with the sponges, foraminifers, gas-
tropod eggs, bryozoans, corals, tunicates, crinoids, or cystoids,
which are now widely considered as calcareous green algae
(Beadle, 1988, 1991). The first cyclocrinitids were reported by
Von Eichwald (1840) as Cyclocrinites spaskii, and its systematic
position was not defined until the resemblance of cyclocrinitids
were established with dasycladacean green algae, documented
by Stolley in 1896; later Pia (1920, 1927) reviewed and cate-
gorised them as a dasycladacean tribe. Thereafter, cyclocrinitids
have been placed in the Order Dasycladales (Pia, 1920, 1927;
Egerod, 1952; Johnson, 1961; Wray, 1977; Bassoullet et al.,
1979). Although European palaeontologists accepted Stolley’s
idea; North American workers considered them as aberrant
sponges for so many years. Twenhofel (1928), an American
palaeontologist regarded Pasceolus identical with Cyclocrinites.
Johnson (1961) also described cyclocrinitids as dasycladaceans,

https://doi.org/10.1016/j.palwor.2019.07.007
1871-174X/© 2019 Elsevier Ireland Ltd Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. Published by Elsevier B.V. All rights reserved.

Please cite this article in press as: Shabbar, H., et al., Cyclocrinitids from the Lower Palaeozoic Tethyan sequence of Spiti, India. Palaeoworld
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Fig. 1. General map of northern India and surrounding countries, showing the major structural units of the Himalaya from Nanga Parbat to Namcha Barwa and
south of the Indus-Tsangpo suture zone. The inset shows the Indian part of the Tethyan Belt with location of four sub-basins with Palaeozoic sedimentary sequences
belonging to Tethyan Himalaya (after Gansser, 1964).

notwithstanding some workers classified them as receptaculi- 2. Study area

tids (Nitecki and Toomey, 1979; Fisher and Nitecki, 1982;
Nitecki, 1986). Moreover, some algologists doubted on the
taxonomic assignment proposed by Pia (1920) and Bassoullet
et al. (1979) and proposed that cyclocrinitids belong to the
order Cyclocrinales, a possible sister order of Dasycladales
(Nitecki, 1986; Spjeldnæs and Nitecki, 1990a, 1990b; Nitecki
and Spjeldnæs, 1992). Even though typically rare as fossils,
they may be locally occurring in the company of more familiar
shallow marine Palaeozoic organisms such as corals and bra-
chiopods. They are index fossils for several North American
and European rock units, and a number of marker beds have
been designated for cyclocrinites or an associated genus. The
cyclocrinitids, a small tribe of macrofossils, first appeared in
lower middle Ordovician (Nitecki, 1970), boomed during the
Caradoc (lower late Ordovician) (Beadle, 1991), and were osten-
sibly wiped out at the beginning of the Wenlock time (Middle
Silurian) (Beadle and Johnson, 1986).
Here we present, the taxonomic study of cyclocrinitids recov-
ered from the Spiti, Himachal Pradesh, India. The algal remains
are recorded from the Takche Formation of the Sanugba Group
exposed along the right bank of the Spiti River near the Takche
locality in the Spiti Valley. Based on our collection compris-
ing thirty-five specimens, we have identified four different taxa
namely, Cyclocrinites favus, C. pyriformis, C. globosus, and C.
cf. welleri.
2.1. General geology
Spiti valley lies parallel to the general Himalayan trend
NW-SE (Bagati et al., 1991). The geology and lithostratigra-
phy of the basin have been studied and discussed by many
workers (Srikantia, 1981; Bhargava et al., 1991; Bhargava
and Bassi, 1998; Bhargava, 2008 and references therein). The
Early Palaeozoic successions of Spiti valley comprise of Kun-
zum La (Cambrian), Thango (Early Ordovician), Takche (Late
Ordovician–Early Silurian), and Muth (Late Devonian) forma-
tions. Two contrasting lithological successions exist between the
underlying fossiliferous member of the Kunzum La Formation
of Cambrian age and overlying Muth Formation of Devonian
age; of these the lower succession is recognizable by its deep
crimson color and rugged topography, designated as Thango
Formation (Srikantia, 1981; Bhargava, 2008). This formation
rests over the Kunzam La Formation with a sharp and angular
contact and is conformably overlain by the upper succession,
the Takche Formation. The Takche Formation underlies the
Muth Formation unconformably (Bhargava, 2008). It appears
earthy brown/rusty brown on the weathered surface. The geo-
logical map of Spiti Basin along with the lithology of Takche
Formation of Takche section is shown in Figs. 2, 3 respec-
tively.

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Fig. 2. Geological Map of the Spiti Area, showing different Palaeozoic formations (after Bhargava et al., 1991).

2.2. Age of Takche formation
The age and lithological association and nomenclature of
the Takche sequence have been debatable since its pioneer
description by Stoliczka (1865), and thereafter by many workers
(Griesbach, 1889; Hayden, 1904). Goel and Nair (1977) pro-
pounded the name Pin Limestone for this succession whereas the
name Takche Formation was proposed by Srikantia (1977, 1981
and Bhargava (2008) for this Ordovician–Silurian sequence.
Here we follow Bhargava and Bassi (1998) and prefer to use
Takche Formation instead of Pin Formation (Goel and Nair,
1977; redefined by Suttner, 2007). The build-out of carbonate
bands in the upper part of the Thango Formation is regarded to
demarcate starting of the Takche Formation (Bhargava, 2008).
The formation contains limestone, nodular limestone, marl,
shale, well-bedded limestone, siltstone, dolomite, calcareous
sandstone and is characterised by a minor development of coral
with little algal input (Ranga Rao et al., 1984; Bhargava and
Bassi, 1987; Suttner et al., 2005). The succession is exposed at
Gechang in Parahio Valley, at Muth in Pin Valley and at Takche
Locality of Spiti Basin. The sequence attains the maximum
thickness of about 272 m at Gechang and has been categorised
into three members namely A, B and C (Mehrotra et al., 1982).
Srikantia (1981) ascribed an Upper Silurian to Lower Devo-
nian age to the entire sequence. However, Mehrotra et al. (1982)
assigned an Ordovician age to A and B members of the sequence
which was suggested by Hayden (1904) and Reed (1912) as well.
Mehrotra et al. (1982) also considered that the boundary between
Ordovician and Silurian exists within the basal section of the C
member and not at the upper part of the B member as envisaged
by Hayden (1904) and Reed (1912). Bhargava and Bassi (1986)
advocated an early to middle Silurian age to reefal portion of
the sequence. On the other hand, Kato et al. (1987) suggested an
Ordovician age to their Pin Limestone (equivalent to Takche For-
mation) on the findings of algal remains. However, Bhargava and
Bassi (1998), advocated an age spanning from Late Ordovician
to Middle/early late Silurian owing to the occurrence of coral and
algal fossils. Suttner (2003) has subdivided the Pin Formation
(equivalent to Takche Formation) in to 17 litho-units viz., P1-P17
that are exposed near Farka Muth village in the Pin Valley. Fur-
thermore, in the year 2007, Suttner has proposed three members
of the Pin Formation exposed near Farka Muth village viz., Farka
Muth Member (P1-P6), Takche Member (P7-P13) and Mikim
Member (P14-P17). Recently, Myrow et al. (2018) has assigned

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Fig. 3. (A) Photograph of studied section of Takche Formation, Takche locality. (B) Closer view of the studied section showing fossils locality. (C) Lithology of a
part of Takche Formation exposed near Takche locality, Spiti India; part 2 of lithology conformably overlies the part 1 of litholog.

Silurian (more specifically Llandovery) age to uppermost mem-
ber of the Pin Formation exposed in Gechang, Parahio Valley
on the basis of the occurrence of ozarkodinid fauna. Our stud-
ied sequence, which has yielded the Cyclocrinitids, corroborates
with the units (P1-P6) of Farka Muth Member. The age of the
overlying units of P7-P13 have been assigned mid to late Katian
(Amorphognathus ordovicicus Zone) on the basis of presence of
conodonts by Suttner (2003) and Suttner et al. (2007), whereas
Early Silurian (?Llandovery) age is suggested to the upper part,
i.e., units P14-P17, however, the definite age for the lowermost
units P1-P6 remains uncertain.
3. Material and methods
The algal fossils reported and described in this article were
collected from an outcrop section of Takche Formation, Takche
Locality (32◦ 27.024 N, 77◦ 41.767 E), 7 km from Losar Village
towards Kunzum La, Spiti Basin (Fig. 3). The fossils were recov-
ered from calcareous grey siltstone unit of Takche Formation
which appears rusty brown due to weathering. The algal fossils
are recovered along with some brachiopods and trace fossils.
The fossils include only impressions of complete or fragments
of thalli. Thirty-five fossil impressions are studied using low
power Leica microscope (MZ6) to document different morpho-
logical characters such as the arrangement of lateral heads and
facets, shape and size of thalli, the outline of facets and lateral
heads and identified up to genus and species level. Specimens
are photographed using a Sony (DSC-HX400 V) digital cam-
era and Nikon (DS-Fi1) digital camera mounted on a low power
Leica microscope (MZ6); measured using ImageJ software. The
systematics of these algal remains is given in the followings.
Repository: All the macrofossil specimens documented in
this article are stored in the repository of Birbal Sahni Institute
of Palaeosciences, Lucknow. The specimens are registered vide
statement number 1512 and museum specimen numbers 41681-
41686.

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4. Systematic palaeobotany
Division Chlorophyta (Pascher, 1914) Papenfuss, 1946
Order Cyclocrinales Nitecki et Toomey
Family Cyclocrinaceae Maslov, 1956
Tribe Cyclocriniteae (Pia, 1920) Bassoullet et al., 1979
Genus Cyclocrinites (Von Eichwald, 1840) Stolley, 1896
Cyclocrinites favus Salter, 1851
(Fig. 4F–K)
1912 Pasceolus mellifluus Salter – Reed, pl. 16, figs. 13–16.
1912 Pasceolus shianensis Reed – Reed, pl. 16, fig. 17.
1987 Palaeodictyon sp. – Kumar and Kashkari, pl. 1, figs. 4, 5.
1999 Cyclocrinites favus Salter – Maithy et al., pl. 1, figs. 1–5.
Material: Six specimens in the collection.
Description: The specimens of this species were preserved as
impressions of the thalli, more or less complete, spherical to
ovate in shape, preserved along the bedding plane. Size of thalli
varies from 29.03 mm to 32.92 mm in length and 18.16 mm
to 23.25 mm in width. The impressions of lateral heads are
observed in the form of protrusions of facets upon the surface.
Lateral heads are six-sided and are the extension of the walls of
the facets. Typically each facet adjoins four to six other facets.
Lateral heads vary in size from 0.38 mm to 0.89 mm. The total
number of lateral heads ranges from 280 to 350. No attachment
mechanism is observed.
Comparison and remarks: Our specimens are comparable
with the holotype specimen of Cyclocrinites favus described
by Nitecki (1970) and also resemble with the specimen reported
by Beadle and Johnson (1986, fig. 2f) in overall shape and mor-
phological characteristics. Thalli also show resemblance with
C. favus figured by Beadle (1991, fig. 3c, d) as well as C. favus
figured and described by Maithy et al. (1999, pl. 1, figs. 1–5).
Cyclocrinites cf. welleri Nitecki, 1970
(Fig. 4D, E)
1963 Mastopora? sp. – Greife and Langenheim, pl. 63, fig. 4.
1970 Cyclocrinites welleri – Nitecki, figs. 4, 5, 7D, 21E, 41.
Type species: Mastopora? sp. Greife and Langenheim, 1963,
subsequent designation by Nitecki (1970), Cyclocrinites welleri.
Middle Ordovician, California.
Material: Two specimens in the collection.
Description: The specimens of this species were preserved as
thalli impressions along the bedding plane, nearly spherical in
shape, regular in outline. Specimens range from 27.435 mm to
32.610 mm in length and 17.229 mm to 19.907 mm in width.
Oval-shaped cavity observed in the centre of the thallus. Hexag-
onal facets are preserved as impressions and are interconnected
with six other facets. Walls of the facets are rather thick as
compared to other species; occasionally up to 0.657 mm. Few
lateral heads are observed. Facets vary in size from 0.585 mm
to 1.089 mm.
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5
Comparison and remarks: The preserved specimens closely
resemble with the Holotype specimen of Cyclocrinites welleri
described by Nitecki (1970) from Mazourka Formation, Mid-
dle Ordovician, California, in having nearly spherical shape,
thick-walled of facets, and characteristic arrangement of facets.
Though, our specimens are comparatively larger than the Holo-
type specimen while other characterises are very similar.
Cyclocrinites globosus Billings, 1857
(Fig. 4L, M)
1857 Pasceolus globosus – Billings, p. 343.
1970 Cyclocrinites globosus Billings – Nitecki, figs. 3B, 7C, 21D, 32, 37.
(For detail synonymy see Nitecki, 1970 and references therein.)
Type species: Pasceolus globosus Billings, 1857, subsequent
designation by Nitecki (1970), Cyclocrinites globosus. Middle
Ordovician, Ontario.
Material: Three specimens in the collection.
Description: The specimens were preserved as thalli impres-
sions, almost spherical in shape with regular outline. The
diameter of the thalli ranges from 18.5 mm to 22.545 mm. Bul-
bous protrusions are seen in the centre of the specimens. Lateral
heads are few and nearly round. Facets are concave, and each
facet commonly communicates with six adjacent facets. The
facets are mostly annular, occasionally polygonal and rarely
hexagonal, with size from 0.316 mm to 0.336 mm.
Comparison and remarks: The specimens described herein
closely resemble in their morphology with the holotype of
Cyclocrinites globosus figured and described by Nitecki (1970,
fig. 37) from Middle Ordovician, Ottawa Formation, Ontario,
Canada.
Cyclocrinites pyriformis Bassler, 1915
(Fig. 4A–C)
1896 Cyclocrnius pyriformis – Stolley, text-figs. 20–27.
1909 Nidulites – Bassler, pl. 7, fig. 11.
1909 Nidulites cf. favus – Bassler, p. 59.
1970 Cyclocrinites pyriformis Bassler – Nitecki, figs. 49, 50.
1999 Cyclocrinites pyriformis Bassler – Maithy et al., pl. 1, fig. 6.
Type species: Nidulites pyriformis Bassler, 1915, subsequent
designation by Nitecki (1970), Cyclocrinites pyriformis. Mid-
dle Ordovician, Virginia.
Material: Four specimens in the collection.
Description: The specimens were preserved as impressions
of thalli, more or less complete, nearly claviform to ovoid
and regular in outline. The preserved thalli vary and are
13.061 mm–30.935 mm in length and 8.467 mm–18.077 mm in
width. Facets are regular and hexagonal in outlines. The largest
is 1.359 mm across. Facets are relatively deep. The depth of
the facets is typically annulus; however, walls are hexagonal
and form well-ordered lines on the surface of the thalli. The
maximum thickness of the facet’s wall reaches up to 0.173 mm.
From the preserved interior of the facets, the heads appear to
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Fig. 4. Photographs of different types of Cyclocrinitids recorded from the Takche Formation, Takche locality, Spiti. (A–C) Cyclocrinites pyriformis Bassler; (A)
BSIP Museum Specimen No. 41684/A; (B) enlarged view of (A), showing depth of facets and hexagonal outline; (C) BSIP Museum Specimen No. 41684/D. (D, E)
Cyclocrinites cf. welleri Nitecki; (D) BSIP Museum Specimen No. 41683/A; (E) enlarged view of (D), showing depths of facets, thickness of wall and hexagonal
outline. (F–K) Cyclocrinites favus Salter; (F) BSIP Museum Specimen No. 41682/B; (G) enlarged view of (F), showing lateral heads; (H) BSIP Museum Specimen
No. 41682/C; (I) enlarged view of (H), showing lateral heads; (J) BSIP Museum Specimen No. 41681/A; (K) enlarged view of (J), showing lateral heads, the distal
tips of lateral branches. (L, M) Cyclocrinites globosus Billings; (L) BSIP Museum Specimen No. 41684/C; (M) BSIP Museum Specimen No. 41686/A. (N, O)
Cyclocrinites sp.; (N) BSIP Museum Specimen No. 41682/A; (O) BSIP Museum Specimen No. 41684/B. Scale bar: 5 mm for (A, D, F, H, J, L, M); 1 mm for (B, C,
E, G, I, K, N, O).

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Fig. 5. Palaeogeography of the northeastern Gondwana and adjacent peri-Gondwanan areas in early to mid-Ordovician (Arenig) times at 480 Ma. Ab, Alborz terrane;
Afg, Afghan terrane; Lut, Lut terrane; M, Madagascar; Qiang, Qiang terrane; San, Sanand terrane; Tau, Taurides terrane (after Torsvik and Cocks, 2009). Star
indicates the position of our fossil locality.

be relatively deep. Very few lateral heads are preserved. Actual
attachment mechanism could not be observed.
Comparison and remarks: In appearance and pattern of facets
and lateral heads of thalli, the specimens are comparable with
the holotype of Cyclocrinites pyriformis as described by Nitecki
(1970, figs. 49, 50) and they also resemble with the specimens
reported by Beadle (1991, fig. 2a-d) as well as by Maithy et al.
(1999, pl. 1, fig. 6).
Cyclocrinites sp.
(Fig. 4N, O)
Material: Twenty specimens in the collection.
Description: Impressions of spheroidal to ovate thalli and
fragments of thalli, poorly preserved with negligible surface
features observed. Lateral heads and deep facets observed in
some specimens. Closely packed facets generally manifest reg-
ular hexagonal appearances; however, most of the specimens
are slightly irregular in shape. Specimens can not be assigned
to species level because of ill-preserved and fragmentary
nature, however by the presence and arrangement of polygo-
nal heads and deep facets they are identified as the Cyclocrinites
thalli.
5. Discussion
Cyclocrinitids are extinct group of small macrofossil remains,
of general receptaculitid type, whose affinity to plant remains
or animals, has been debatable and doubtful for quite long
time (Nitecki, 1970). They have been grouped with Protozoans,
Porifera, Corals, Bryozoans, molluscs and algae. Stratigraph-
ically, they range from lower Middle Ordovician to Lower
Silurian. The structure of the Cyclocrinitids is very akin to that
of modern calcareous algae (family Dasycladaceae).
In India, the report of Cyclorinitids dates back to 1912,
when Reed documented a few curious objects, of somewhat
problematic nature, from the Ordovician sequence of Tethyan
Himalayan Basin. He placed them under Incertae sedis and
collated with genera Ischadites, Pasceolus and Apidium. The
definite biological association was not known, until Pia (1927),
Johnson and Konishi (1959), Johnson (1961) and Kesling and
Graham (1962) cogently characterised them as floral remains of
marine Dasycladacean algae. A macroscopic fragment resem-
bling honeycomb in appearance was reported by Sahni (1953),
and he compared his fragmental specimen with Pasceolus mel-
lifluus (Salter) documented by Reed (1912). There was a gap
in the algal studies in the Tethyan Himalaya until Kato et al.

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(1987) reported and illustrated two taxa of calcareous green
algae namely Apidium indicum and Coelosphaeridium shia-
nense from the Ordovician sections of Pin Valley and Parahio
Valley of Spiti. In the same year, Kumar and Kashkari (1987)
reckoned the honeycomb-shaped macrofossils as ichnofossils
from the Gechang (Parahio Valley) and Muth (Pin Valley) and
described them as Palaeodictyon sp. which was later recognised
as cyclocrinites (Maithy et al., 1999). Furthermore, Bhargava
(2008) cast doubt on both the identifications and summarised the
fossil to be a remains of Palaeofavosites/Favosites colony (coral
remains). However, the occurrence of Cyclocrinitids is recently
reported from the Takche Formation of Gechang village, Parahio
Valley, Spiti (Chaubey et al., 2019). Anatomy and morphology
of dasycladacean algae have been discussed in detail by Egerod
(1952), Valet (1968), Nitecki (1970) and Beadle (1988). Accord-
ing to the above-mentioned researchers, “Dasycladacean algae
have thin, siphonous median axes, which are attached to the sub-
strate. The lateral branches are arranged on a central axis in the
spiralling pattern at regular intervals and these typically furcate
distally to secondary branches”. In recent genera Neomeris and
Bornetella, bulbous lateral heads are grown on the distal tips of
secondary branches, sub-hexagonal/hexagonal in form and are
densely packed together forming a faceted outer cortex which is
claviform to spherical in shape (Beadle, 1988).
In modern dasycladaceans the main controlling factors in
thriving, dispersal and abundance are depth, water energy, clar-
ity of the water, light intensity, normal salinity, depth of light
penetration and substrate’s quality (Nitecki, 1970; Lee and
Caldwell, 1977; Hammann and Serpagli, 2003). In clear Pacific
and Caribbean seas, they are most prevalent up to the depth of
5 to 6 m from low tide level; however, they may occur up to
the depth of 90 m (Taylor, 1960; Johnson, 1961; Nitecki, 1970).
Furthermore, they can not extend beyond 20 m depths in more
turbid waters like the Persian Gulf (Clarke and Keij, 1973). So,
they are primarily confined to water depths extending up to about
100 m in warm temperate waters in the equatorial region and the
shallowest of tropical (Lee and Caldwell, 1977).
Cyclocrinitids may have been quite sensitive to water energy,
depth and latitude (Nitecki, 1970; Mørk and Worsley, 1980;
Beadle and Johnson, 1986). Extracellular calcification and
aragonitic nature of cyclocrinitids suggest that they also had
low latitude and warm water distribution (Beadle, 1988). It
is inferred from palaeogeographic reconstructions that mostly
all cyclocrinitids inhabited within 30◦ south and north of the
palaeoequator (Beadle and Johnson, 1986; Beadle, 1988), except
occurrence of a few as far as 45◦ of the south (Beadle and
Johnson, 1986). Maithy et al. (1999) opined that cyclocrini-
tids seem to be good indicators of low palaeolatitudes and they
also can be reliable relative depth indicators. Very recently,
Chaubey et al. (2019) have also argued that large accumula-
tion of cyclocrinitids recovered from the Takche Formation near
Gechang village, Parahio valley Spiti, might have been formed
during a storm event which generated currents that penetrated
into normally quiet water and swept thalli together. This event is
further substantiated with the occurrence of present cyclocrini-
tids assemblage from the Takche Formation, Takche village
locality. It also suggests that Takche Formation indicates that
Please cite this article in press as: Shabbar, H., et al., Cyclocrinitids from the Lower Palaeozoic Tethyan sequence of Spiti, India. Palaeoworld
(2019), https://doi.org/10.1016/j.palwor.2019.07.007
the Spiti region of the northwestern Himalaya must have been
located in low palaeolatitudes during the Late Ordovician and
early Silurian (Chaubey et al., 2019) (Fig. 5).
Cyclocrinitids are known from North America (USA and
Canada), South China (Sichuan), Baltoscandia (Norway, Rus-
sia, Estonia, glacial erratics of northern Germany and Poland),
Eastern Kazakhstan and Central Himalayas–India (Hammann
and Serpagli, 2003). Modern calcareous green algae rarely grow
on muddy or sandy ocean bottoms (Taylor, 1960), since these
materials do not provide an adequate and firm foundation for
anchoring. On this ground, it is inferred by Nitecki (1970) that
cyclocrinitids are lithophytic. Therefore, they avoided very soft
muddy or sandy sea bottoms and typical reef environments
(Hammann and Serpagli, 2003). Cyclocrinitids typically occur
in fine-grained rocks such as siltstones, micritic limestones and
also in shales that form a quite resistant substrate on which
they were attached (Nitecki, 1970; Mørk and Worsley, 1980;
Beadle and Johnson, 1986; Beadle, 1988, 1991). According to
Hammann and Serpagli (2003), they also appear to have avoided
very shallowly, stirred bathymetric regions and probably inhab-
ited below normal wave base due to frangible nature.
They were quite vulnerable to compaction and specimens
in shales are frequently quite flattened (Beadle, 1991). Speci-
mens have different diagenetic histories (Osgood and Fischer,
1960; Nitecki and Johnson, 1978; Mørk and Worsley, 1980).
Consequently, their preservation is entirely protean, even within
a single species (Nitecki, 1970). Cyclocrinitids are usually
encrusted by a thin layer of calcium carbonate; however, they
are not as completely preserved as fossil dasycladaceans (Lee
and Caldwell, 1977). Even though cyclocrinitids is usually
associated with common marine invertebrates of the shallow
environment, they also occur in a marine environment restricted
to high salinity (Johnson and Campbell, 1980).
Cyclocrinitids had a medial main axis on which thin lateral
branches grew in whorl extending outwards in all directions,
and bulbous hexagonal/sub-hexagonal lateral heads grew on
the distal point of lateral branches; cyclocrinitids vary in out-
line from spherical to claviform (Nitecki, 1970; Beadle, 1988,
1991). All cyclocrinitids belonging to Silurian time had gen-
erally spherical thalli (Beadle and Johnson, 1986). They were
confined to small sizes, mostly ranging from 1.5 to 3.4 cm in
length (Beadle, 1988) and the largest reported cyclocrinitid is
C. halli thallus 5.86 cm long from Ellis Bay Formation, Quebec,
Canada from Upper Ordovician (Nitecki, 1970). The emergence
of cyclocrinitids concurs with major dasyclads radiation dur-
ing middle Ordovician (Chuvashov and Riding, 1984; Beadle
and Johnson, 1986). Their global range began to shrink steadily
during Late Ordovician (Beadle, 1988). Only two species of
cyclocrinitids viz., C. favus and C. gregarius were present
by early Llandovery and confined to Avalonia, Laurentia and
Baltica regions (Beadle and Johnson, 1986). Cyclocrinites pyri-
formis remained present almost throughout the entire middle
Champlainian series–middle Ordovician (Darriwilian) in North
America, and it is used there as an index fossil (Nitecki, 1970).
Our specimens range from 18.5 mm to 32.91 mm in length and
except Cyclocrinites favus all other specimens described herein
are known from Middle Ordovician. However, for our studied
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H. Shabbar et al. / Palaeoworld xxx (2019) xxx–xxx
assemblage no definite age can be assigned for want of more pre-
cise index fossils. Though, present Cyclocrinites assemblage is
recovered from the units which are correlatable with units P1-P6
(Farka Muth Member) of Pin Formation (= Takche Formation)
for which definite age remains uncertain. The age of the overly-
ing units of P7-P13 have been assigned mid to late Katian (Upper
Ordovician) on the basis of presence of conodonts (Amorphog-
nathus ordovicicus Zone) (Suttner, 2007). Hence, the age of
Cyclorinites yielding units may be traced back to? early late
Ordovician in age.
6. Conclusion
I. Four taxa of the genus Cyclocrinites namely Cyclocrinites
pyriformis, C. favus, C. welleri and C. globosus discussed
in this article are found as flattened thalli in grey calcareous
siltstone unit of Takche Formation.
II. They probably colonized in relatively shallow, less turbid
and low to moderate hydrodynamic conditions within photic
zone.
III. Both C. welleri (Nitecki) and C. globosus (Billings) are
reported and described here for the first time from Tethyan
Himalaya of India.
IV. Cyclocrinites favus and C. pyriformis are also reported
from United Kingdom (Wales and Scotland), North America
(USA and Canada), Baltoscandia (Norway and Germany)
and NW Himalaya in India while Cyclocrinites globosus is
found in North America (USA and Canada) and Germany.
Cyclocrinites welleri is only reported from California, USA.
Acknowledgements
The authors are thankful to the Director, Birbal Sahni Insti-
tute of Palaeosciences for granting permission to publish this
paper and providing necessary facilities to carry out this work.
HS is grateful to UGC for (MANF-F1-17.1/2015-16/MANF-
2015-17-UTT-52912) for providing Senior Research fellowship
grant to facilitate this work. AS and KJS also acknowledged the
financial support given by SERB, DST, Government of India
in the form of sponsored project (EMR/0006042/2016) to carry
out this work. Mr. Suyash Gupta is also acknowledged for his
help in preparation of figures. The authors are also indebted to
reviewers Dr. Olev Vinn and Dr. Thomas J. Suttner and also
to the editor Si-Wei Chen for their constructive comments and
suggestions that significantly improved this manuscript.
References
Bagati, T.N., Kumar, R., Ghosh, S.K., 1991. Regressive-transgressive sedimen-
tation in the Ordovician sequence of the Spiti (Tethys) basin, Himachal
Pradesh. India. Sedimentary Geology 73 (1–2), 171–184.
Bassler, R.S., 1909. The cement resources of Virginia west of the Blue Ridge.
Virginia Geological Survey Bulletin (2), 309.
Bassler, R.S., 1915. Bibliographic index of American Ordovician and Silurian
fossils. Bulletin of the United States National Museum 92 (2), 1–1512.
Bassoullet, J.P., Bernier, P., Deloffre, R., Génot, P., Jaffrezo, M., Vachard, D.,
1979. Essai de classification des Dasycladales in tribus. Bulletin des Centres
de Recherches Exploration-Production Elf-Aquitaine 3, 429–442.
Please cite this article in press as: Shabbar, H., et al., Cyclocrinitids from the Lower Palaeozoic Tethyan sequence of Spiti, India. Palaeoworld
(2019), https://doi.org/10.1016/j.palwor.2019.07.007
9
Beadle, S.C., 1988. Dasyclads, cyclocrinitids and receptaculitids: comparative
morphology and paleoecology. Lethaia 21 (1), 1–12.
Beadle, S.C., 1991. Cyclocrinitids. In: Riding, R. (Ed.), Calcareous Algae and
Stromatolites. Springer, Berlin, Heidelberg, pp. 114–124.
Beadle, S.C., Johnson, M.E., 1986. Palaeoecology of Silurian cyclocrinitid
algae. Palaeontology 29 (3), 585–601.
Bhargava, O.N., 2008. An updated introduction to the Spiti geology. Journal of
the Palaeontological Society of India 53 (2), 113–128.
Bhargava, O.N., Bassi, U.K., 1986. Silurian reefal buildups: Spiti-Kinnaur,
Himachal Himalaya, India. Facies 15 (1), 35.
Bhargava, O.N., Bassi, U.K., 1987. Algae from the Silurian Manchap Formation,
Tidong Valley (Kinnaur), Himachal Himalaya. Journal of the Geological
Society of India 29 (5), 500–502.
Bhargava, O.N., Bassi, U.K., 1998. Geology of Spiti-Kinnaur Himachal
Himalaya. Memoirs of the Geological Survey of India 124, 1–210.
Bhargava, O.N., Srivastava, R.N., Gadhoke, S.K., 1991. Proterozoic–Palaeozoic
Spiti Sedimentary Basin. In: Tandon, S.K., Pant, C.C., Casshyap, S.M.
(Eds.), Sedimentary Basins of India: Tectonic Context. Gyanodaya
Prakashan, Nainital, pp. 236–260.
Billings, E., 1857. Report for the year 1856, of E. Billings Esq., palaeontolo-
gist addressed to Sir William E. Logan, provincial geologist. Geological
Survey of Canada, Report of Progress (for the years 1853-54-55-56),
247-345.
Chaubey, R.S., Vinn, O., Singh, B.P., Bhargava, O.N., Prasad, S.K., Kishore,
N., 2019. Warm-water Dasycladaceae algae from the Late Ordovician of
the Parahio Valley, Spiti, India. Estonian Journal of Earth Sciences 68 (1),
45–53.
Chuvashov, B., Riding, R., 1984. Principal floras of Palaeozoic marine calcare-
ous algae. Palaeontology 27 (3), 487–500.
Clarke, M.H., Keij, A.J., 1973. Organisms as producers of carbonate sediment
and indicators of environment in the southern Persian Gulf. In: Purser, B.H.
(Ed.), The Persian Gulf. Springer, Berlin, Heidelberg, pp. 33–56.
Draganits, E., Mawson, R., Talent, J.A., Krystyn, L., 2002. Lithostratigraphy,
conodont biostratigraphy and depositional environment of the Middle Devo-
nian (Givetian) to Early Carboniferous (Tournaisian) Lipak Formation in the
Pin valley of Spiti (NW India). Rivista Italiana di Paleontologia e Stratigrafia
108 (1), 7–36.
Egerod, L.E., 1952. An analysis of the siphonous Chlorophy-cophyta
with special reference to the Siphonocladales, Siphonales, and Dasy-
cladales of Hawaii. University of California Publications in Botany 25,
325–454.
Fisher, D.C., Nitecki, M.H., 1982. Standardization of the anatomical orientation
of receptaculitids. Journal of Paleontology 56 (S13), 1–40.
Gansser, A., 1964. Geology of the Himalaya. Interscience Publishers, London,
289 pp.
Greife, J.L., Langenheim Jr., R.L., 1963. Sponges and brachiopods from the Mid-
dle Ordovician Mazourka Formation, Independence Quadrangle, California.
Journal of Paleontology 37 (3), 564–574.
Goel, R.K., Nair, N.G.K., 1977. The Spiti Ordovician–Silurian succession. Jour-
nal of the Geological Society of India 18, 47–48.
Griesbach, C.L., 1889. Geological notes — a sequence of formations in Spiti.
Records of the Geological Survey of India 22, 158–167.
Hammann, W., Serpagli, E., 2003. The algal genera Ischadites Murchison, 1839
and Cyclocrinites Eichwald, 1840 from the Upper Ordovician Portixeddu
Formation of SW Sardinia. Bollettino-Societa Paleontologica Italiana 42
(1/2), 1–30.
Hayden, H.H., 1904. The geology of Spiti with parts of Bashahr and Rupshu.
Memoirs of the Geological Survey of India 36, 1–129.
Hubmann, B., Suttner, T.J., 2007. Calcimicrobes and calcareous algae of the Pin
Formation (NW Himalayas, India): a contribution to the Ordovician Flora.
Acta Palaeontologica Sinica 46 (Suppl), 188–198.
Johnson, J.H., 1961. Review of Ordovician algae. Colorado School of Mines
Quarterly 56 (2), 1–101.
Johnson, J.H., Konishi, K., 1959. A review of Silurian (Gotlandian) algae.
Colorado School of Mines Quarterly 55 (1), 1–114.
Johnson, M.E., Campbell, G.T., 1980. Recurrent carbonate environments in the
Lower Silurian of northern Michigan and their inter-regional correlation.
Journal of Paleontology 54 (5), 1041–1057.
+Model
PALWOR-529; No. of Pages 10 ARTICLE IN PRESS
10 H. Shabbar et al. / Palaeoworld xxx (2019) xxx–xxx
Kato, M., Goel, R.K., Srivastava, S.S., 1987. Ordovician algae from Spiti, India.
Journal of the Faculty of Science, Hokkaido University, Series 4, Geology
and Mineralogy 22 (2), 313–323.
Kesling, R.V., Graham, A., 1962. Ischadites is a dasycladacean alga. Journal of
Paleontology 36 (5), 943–952.
Kumar, G., Kashkari, R.L., 1987. Palaeodictyon and Chondritesfrom the Takche:
Formation (Ordovician–?Lower Devonian) Spiti Valley, Himachal Pradesh,
India. Journal of the Palaeontological Society of India 32, 47–52.
Lee, D.G., Caldwell, W.G.E., 1977. A new dasycladacean alga associated with
the ‘Arctic Ordovician’ fauna on Cornwallis Island. Canadian Journal of
Botany 55 (1), 52–60.
Maithy, P.K., Ghosh, A.K., Kumar, G., 1999. First report of dasycladacean alga-
Cyclocrinites from the Ordovician of Takche Formation, Spiti, India. Journal
of the Geological Society of India 54 (4), 379–385.
Maslov, V.P., 1956. Fossil calcareous algae of the USSR. Trudy Instituta Geo-
logicheskh Nauk, Akademii Nauk SSSR 160, 1–301.
Mehrotra, P.C., Singh, G., Kumar, G., Ahluwalia, A.D., 1982. Chitinozoa from
Lower Palaeozoic sequence of Spiti, Himachal Pradesh, India. Geophytology
12 (1), 111–116.
Mørk, A., Worsley, D., 1980. The environmental significance of algae in the mi
Llandovery succession of the central Oslo Region. Lethaia 13 (4), 339–346.
Myrow, P.M., Fike, D.A., Malmskog, E., Leslie, S.A., Zhang, T., Singh, B.P.,
Prasad, S.K., 2018. Ordovician–Silurian boundary strata of the Indian
Himalaya: Record of the latest Ordovician Boda event. Geological Survey
of America Bulletin 131 (5–6), 881–898.
Nitecki, M.H., 1970. North American cyclocrinitid algae. Fieldiana Geology,
Publications of Field Museum of Natural History 21, 1–192.
Nitecki, M.H., 1986. Receptaculitids and their relationship to other problematic
fossils. In: Hoffman, A., Nitecki, M.H. (Eds.), Problematic Fossil Taxa.
Oxford University Press, New York, pp. 27–34.
Nitecki, M.H., Johnson, M.E., 1978. Internal structures of Cyclocrinites dac-
tioloides, a receptaculitid alga from the lower Silurian of Iowa. Fieldiana
Geology, Publications of Field Museum of Natural History 39 (1), 1–15.
Nitecki, M.H., Spjeldnæs, N., 1992. Cyclocrinites Spaskii: A Model of
Cyclocrinitid Morphology. Intern Skriftserie 63. Institutt for Geologi, Uni-
versitetet i Oslo, 69 pp.
Nitecki, M.H., Toomey, D.F., 1979. Nature and classification of receptaculitids.
Bulletin des Centres de Recherches Exploration-Production Elf-Aquitaine
3, 725–732.
Osgood Jr., R.G., Fischer, A.G., 1960. Structure and preservation of Mastopora
pyriformis, an Ordovician dasycladacean alga. Journal of Paleontology 34
(5), 896–902.
Pandey, S., Parcha, S.K., 2018. Calcareous algae from the Ordovician succes-
sion (Thango Formation) of the Spiti Basin, Tethys Himalaya. India. Acta
Palaeobotanica 58 (2), 97–106.
Papenfuss, G.F., 1946. Proposed names for the phyla of algae. Bulletin of the
Torrey Botanical Club 73, 217–218.
Pascher, A., 1914. Uber Flagellaten und Algen. Berichte der Deutschen Botanis-
chen Gesellschaft 32, 136–160.
Pia, J., 1920. Die Siphoneae verticillatae vom Karbonbiszur Kreide. Abhand-
lungen der Zoologisch-Botanischen Gesellschaft in Wien 11 (2), 1–259.
Pia, J., 1927. Thallophyta I. Abteilung. In: Hirmer, M. (Ed.), Handbuch der
Paläobotanik. Oldenbourg Munich Berlin Poncet J (1987) Paléobiogéogra-
phie du genre Vermiporella (algue verte calcaire) á l’Ordovicien moyen et
supérieur. Annales de la Société Géologique du Nord 106 (3), 279–283.
Ranga Rao, A., Dhar, C.I., Jokhan, R., Rao, S.V., Shah, S.K., 1984. Contribution
to stratigraphy of Spiti. Himalayan Geology 12, 98–115.
Reed F.R.C., 1910. The Cambrian fossils of Spiti. Geological Survey of India,
Palaeontologia Indica, Series 15, 7 (1), 1–70.
Reed F.R.C., 1912. Ordovician and Silurian fossils from the central Himalayas.
Geological Survey of India, Palaeontologia Indica, Series 15, 7 (2), 1–168.
Sahni, B., 1953. Note on some possible psilophyte remains from Spiti, North-
West Himalayas. The Palaeobotanist 2, 1–4.
Please cite this article in press as: Shabbar, H., et al., Cyclocrinitids from the Lower Palaeozoic Tethyan sequence of Spiti, India. Palaeoworld
(2019), https://doi.org/10.1016/j.palwor.2019.07.007
Salter, J.W., 1851. List of some of the Silurian fossils of Ayrshire. Quarterly
Journal of the Geological Society of London 7, 170–178.
Sinha, H.N., Trampisch, C., 2013. Calcareous algal association from the Lower
Paleozoic Tethyan sedimentary sequence of the Shiala Formation, Indian
Gondwana. Journal of the Geological Society of India 82 (4), 339–350.
Spjeldnæs, N., Nitecki, M.H., 1990a. Coelosphaeridium: An Ordovician Alga
from Norway. Intern Skriftserie 59. Institutt for Geologi, Universitetet i Oslo,
53 pp.
Spjeldnæs, N., Nitecki, M.H., 1990b. Anatomy and Relationship of the Ordovi-
cian Algal Genus Apidium. Intern Skriftserie 61. Institutt for Geologi,
Universitetet i Oslo, 37 pp.
Srikantia, S.V., 1977. Sedimentary cycle in the Himalaya and their significance
on the orogenic evolution of the mountain belt. Colloques Internationaux
Centre National de la Recherché Scientifique 268, 395–407.
Srikantia, S.V., 1981. The lithostratigraphy, sedimentation and structure of
Proterozoic–Phanerozoic formations of Spiti Basin in the Higher Himalaya
of Himachal Pradesh, India. In: Sinha, A.K. (Ed.), Contemporary Geoscien-
tific Researches in Himalaya. Bishen Singh Mahendra Pal Singh, Dehradun,
pp. 31–48.
Srikantia, S.V., Ganesh, T.M., Sinha, P.K., Tirky, B., 1977. A note on the sig-
nificance of granite clasts in a diamictite of the Thango Formation (Lower
Silurian) Zanskar valley, Ladakh Himalaya. Journal of the Geological Soci-
ety of India 18, 509–511.
Stoliczka, F., 1865. Geological section across the Himalayan Mountains, from
Wangtu bridge on the River Sutlej to Sungdo on the Indus: with an account
of the formations of Spiti, accompanied by a revision of all known fossils
from that district. Memoirs of the Geological Survey of India 5, 1–173.
Stolley, E., 1896. Untersuchungenüber Coelosphaeridium, Cyclocrinus, Masto-
pora und verwandte Genera des Silur. Archive für Anthropologie und
Geologie Schleswig-Holsteins und der Benachbarten Gebiete 1, 177–282.
Suttner, T.J., 2003. Pin Formation (Lower Palaeozoic) at Muth, Spiti (Indian
Himalaya); Stratigraphy and Facies. Doctoral Dissertation, Diploma Thesis.
Vienna University, 80 pp.
Suttner, T.J., 2007. The Upper Ordovician to Lower Silurian Pin Formation
(Farka Muth, Pin Valley, North India) — a formal discussion and redefinition
of its controversial type-section. Acta Palaeontologica Sinica 46 (Suppl),
460–465.
Suttner, T.J., Hubmann, B., Bhargava, O.N., 2005. Late Ordovician calcimi-
crobes and green algae from Spiti (Himalayas, N-India). Berichte des
Institutes für Erdwissenschaften der Karl-Franzens-Universität Graz 10,
130–131.
Suttner, T.J., Lehnert, O., Joachimski, M., Buggisch, W., 2007. Recognition of
the Boda event in the Pin Formation of northern India based on new ␦13 C
and conodont data. Acta Palaeontologica Sinica 46 (Suppl), 466–470.
Taylor, W.R., 1960. Marine Algae of the Eastern Tropical and Subtropical Coasts
of the Americas. Scientific Series 21. University of Michigan Press, Ann
Arbor, 870 pp.
Torsvik, T.H., Cocks, L.R.M., 2009. The Lower Palaeozoic palaeogeographi-
cal evolution of the northeastern and eastern peri-Gondwanan margin from
Turkey to New Zealand. Geological Society, London, Special Publications
325 (1), 3–21.
Twenhofel, W.H., 1928. Geology of Anticosti Island. Memoir Geological Survey
of Canada 154, 1–481.
Valet, G., 1968. Contribution à l’étude des Dasycladales. 1. Morphogenése. Nova
Hedwigia. Zeitschrift für Kryptogamnekunde 16, 21–82.
Von Eichwald, C.E., 1840. Über das silurische Schichten system in Esthland.
Zeitschrift für Nature- und Heilkunde der Medizinischen Akademie zu St.
Petersburg 1 (2), 1–210.
Wray, J.L., 1977. Calcareous Algae, Developments in Paleontology and Stratig-
raphy, Vol. 4. Elsevier Scientific Publishing Co., Amsterdam, New York,
185 pp.