Accepted Manuscript

Tianzhushania spinosa and other large acanthomorphic acritarchs of Ediacaran

Period from the Infrakrol Formation, Lesser Himalaya, India

Harshita Joshi, Meera Tiwari

PII: S0301-9268(16)30069-9
DOI: http://dx.doi.org/10.1016/j.precamres.2016.09.024
Reference: PRECAM 4588

To appear in: Precambrian Research

Received Date: 16 April 2016

Revised Date: 23 July 2016
Accepted Date: 14 September 2016

Please cite this article as: H. Joshi, M. Tiwari, Tianzhushania spinosa and other large acanthomorphic acritarchs of
Ediacaran Period from the Infrakrol Formation, Lesser Himalaya, India, Precambrian Research (2016), doi: http://
dx.doi.org/10.1016/j.precamres.2016.09.024

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Tianzhushania spinosa and other large acanthomorphic acritarchs of Ediacaran Period

from the Infrakrol Formation, Lesser Himalaya, India

Harshita Joshi*‫ ـ‬and Meera Tiwari**

Wadia Institute of Himalayan Geology, 33 General Mahadev Singh Road, Dehradun 248001,

India

* Harshitajoshi281@gmail.com

** meeratiwari2@gmail.com; mtiwari@wihg.res.in

- Geological Survey of India, Sector E, Aliganj, Lucknow 226024, India.

* Corresponding author

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Highlights:

• The occurrence of Tianzhushania in Infrakrol Formation of India, suggests its

coevality to the lower Tianzhushania assemblage of Doushantuo Formation, south
China.
• The present assemblage of Tianzhushania spinosa, T. Polysiphonia,
Papillomembrana compta, Schizofusa sp. Paratetraphycus giganteus,
Gloeodiniopsis lamellosa, Sphaerophycus medium and Unnamed forms A, B and
C is a significant addition to the record of earliest Ediacaran acritarchs from
Infrakrol Formation.
• The age of Infrakrol Formation may be bracketed within the age of Doushantuo
Formation i.e. 635.2±0.6 Ma to 551.1±0.7 Ma.

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Abstract:

Covering a time span from Ediacaran (base of Blaini pink carbonates) to Early Cambrian

(base of Tal Group), the Krol belt in the Lesser Himalaya (India), occurs as a series of

synclines from Solan, Himachal Pradesh in the north-west to Nainital, Uttarakhand in the

south-east. Various lithostratigraphic divisions of this belt reveal many palaeobiological

entities, namely cyanobacteria, algae, acritarchs, small shelly fossils and trace fossils.

Globally, large acanthomorphic acritarchs of the Ediacaran Period are used as significant

biostratigraphic tools for global correlation. In the Krol belt, reports of acanthomorphic

acritarchs from the Infrakrol and Krol ‘A’ formations of the Krol Group have further

supported this notion. This paper reports well-preserved microfossils including

acanthomorphic acritarchs, sphaeromorphic acritarchs, coccoids namely Tianzhushania

spinosa, T. polysiphonia, Papillomembrana compta, Schizofusa sp., Gloeodiniopsis

lamellosa, Sphaerophycus medium, and the unnamed forms A, B and C from the chert

nodules of the Infrakrol Formation exposed in the Nainital Syncline of the Kumaun Lesser

Himalaya. A biostratigraphic correlation based on acanthomorphic acritarchs suggests that

the Infrakrol Formation is coeval to the lower Tianzhushania assemblage zone of the

Doushantuo Formation of south China. Tianzhushania and Papillomembrana are significant

additions to the previous record of the Ediacaran acanthomorphic acritarchs from the Lesser

Himalaya of India and provide an independent evidence for construction of both biozonation

scheme and paleogeography.

Keywords: Large acanthomorphic acritarchs, Tianzhushania spinosa, Ediacaran Period,

Infrakrol Formation, Nainital Syncline, Lesser Himalaya, India

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 1. Introduction

Ediacaran Period, the terminal part of Neoproterozoic, begins with the close of the

last global glaciation of the Cryogenian Period (635 Ma) and ends with the beginning of

the Cambrian Period (542 Ma) (Amthor et al., 2003; Bowring et al., 1993; Condon et al.,

2005; Grotzinger et al., 1995). A negative excursion in the carbon isotopic records in the

beginning and in the end, records of unusual biogeochemical events in carbonate rocks,

and sedimentary organic matter are also observed during this period. The lower and

middle parts of the Ediacaran Period are characterized by cyanobacteria, microalgae,

multicellular algae and acritarchs (Grey, 2005; Moczydlowska, 2008a, b; Xiao and Dong,

2006; Zang and Walter, 1992) and the upper part by extraordinary soft bodied metazoans,

the earliest skeletal metazoans, and tubulous metazoans (Cortijo et al., 2010; Fedonkin et

al., 2007; Gehling, 1999; Germs et al., 2009; Glaessner, 1984; Grazdhankin, 2004;

Narbonne, 2005).

It has been noted worldwide that the characteristic Large Acanthomorphic Acritarchs

(LAAs) radiated and diversified in the lower and middle parts of the Ediacaran Period

(i.e. between ~635 and ~551Ma) (Grey et al., 2003, 2005; Liu et al., 2013; Zhou et al.,

2007). These LAAs are now known from South China, Australia, Eastern European

Platform, Siberia, India and Svalbard (Grey and Walter, 2004; Grey, 2005; Knoll and

Ohta, 1988; Knoll, 1992; Liu et al., 2013, 2014; Moczydlowska et al., 1993;

Moczydlowska and Nagovitsyn, 2012; Nagovitsyn et al., 2004; Sergeev et al., 2011;

Shukla and Tiwari, 2014; Tiwari and Knoll, 1994; Tiwari and Pant, 2004; Veis et al.,

2006; Vidal, 1990; Vorob’eva et al., 2009a, 2009b; Willman, 2006; Willman and

Moczydłowska, 2008,2011; Xiao, 2004; Yin and Li, 1978; Yin, 1987, 1999; Yin and Liu,

1988; Yin et al., 2011;Yuan and Hofmann, 1998; Xiao et al., 2014; Zang and Walter,

1992; Zhang et al., 1998; Zhou et al., 2001; Zhou et al., 2007). The presence of these

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 LAAs has shown their great potential in biostratigraphic zonation and global correlation

of Ediacaran strata in spite of various complications such as facies control, taxonomic

differences due to identification of the same forms in different lithologies (e.g. chert and

shale) and different processing techniques (thin sections/ or macerated slides) (Grey,

2005; Grey et al., 2003; Grey and Willman, 2009; Liu et al., 2013, 2014; Moczydlowska

and Nagovitsyn, 2012; Sergeev et al., 2011; Vorob’eva et al., 2009a, 2009b; Willman

and Moczydlowska, 2008, 2011). Such taphonomic variations are well evident in the

Ediacaran acritarch assemblages of Australia and South China (Grey, 2005; Grey and

Willman, 2009; Zhou et al., 2007).

In India, microfossils including characteristic LAAs are reported from the Krol belt of

the Lesser Himalaya. The Krol belt extends from Solan in the north-west to Nainital in

the south-east in the form of several synclines. This belt embraces Neoproterozoic to

Cambrian succession of the Baliana, Krol and Tal groups (Table 1). Various

lithostratigraphic divisions of these groups reveal several palaeobiological entities,

including small shelly fossils, trace fossils, acritarchs, cyanobacteria and algae (Shukla

and Tiwari, 2014 and references therein). The fossil reports and sequence stratigraphic

studies in the Krol Belt (Kaufman et. al., 2006 and references therein) show that the

succession that includes pink carbonate of Blaini Formation (related to Marinoan

glaciation; Jiang et al., 2003c; Condon et al., 2005; Zhang et al., 2005) and overlying

Infrakrol Formation and Krol Group is designated as Ediacaran succession. Here, in the

lower part of the Ediacaran Period in the Krol belt well-preserved microfossils from the

Baliana Group and the Krol Group were reported. These microfossils include large

acanthomorphic acritarchs, cyanobacteria and multicellular algae viz. Asterocapsoides

sinensis, Asterocapsoides sp., Cymatiosphaeroides yinii, Knollisphaeridium maximum,

Eomicrocoleus crassus, Ericiasphaera spjeldnaesii, Leiosphaeridia crassa,

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 Melanocyrillium horodyskii, Myxococcoides sp., Obruchevella magna, O. parva,

Oscillatoriopsis media, Polybessurus crassus, Polytrichoides lineatus, Siphonophycus

inornatum, S. kestron, S. punctatum, S. robustum, S. typicum, Salome hubeiensis and

Wengania globosa from Infrakrol Formation (Krol Group) (Tiwari and Azmi, 1992;

Tiwari and Knoll, 1994; Tiwari and Pant, 2004) and from Mahi Formation /Krol A (Krol

Group) include Appendisphaera fragilis, A. grandis, Asterocapsoides sp.,

Asterocapsoides sp. B, Cavaspina acuminata, C. basiconica, Eotylotopalla dactylos,

Knollisphaeridium sp., Papillomembrana sp., Weissiella cf., W. grandistella from Krol

‘A’ Formation (Shukla and Tiwari, 2014). These microfossils have been reported

exclusively from the early diagenetic chert nodules of the Infrakrol Formation in the

Pachmunda, Krol Hill synclines in the north-western part and Nainital Syncline in the

south-eastern part of the Krol Belt (Tiwari and Knoll, 1994; Tiwari and Pant, 2004) and

from chert nodules and chert bands of the Krol A Formation of the Khanog and Rajgarh

synclines, in the north-western part of the Krol belt (Shukla and Tiwari, 2014).

The present paper describes additional three dimensionally preserved microfossils

including LAAs: Tianzhushania spinosa, T. polysiphonia, Papillomembrana compta;

sphaeromorphic acritarch: Schizofusa sp., and coccoidal cyanobacteria: Paratetraphycus

giganteus, Gloeodiniopsis lamellosa, Sphaerophycus medium along with unnamed forms

A, B and C preserved within chert nodules of the Infrakrol Formation exposed in the

Takula –Khurpatal road and Hanumangarhi sections in the Nainital Syncline. This data

confirms the presence of Tianzhushania spinosa in India and also validates the potential

of Ediacaran acritarchs in biostratigraphic correlation worldwide. However, in order to

establish biozones within the Ediacaran deposits of the Krol belt, a high resolution

biostratigraphic study with documentation of more taxa of large acanthomorphic

acritarchs from different stratigraphic lithounits of the Krol Group is still required.

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 2. Geological Setting

The Krol Belt of the Lesser Himalaya, India is exposed in the form of various doubly

plunging synclines from Solan in the north-west to Nainital in the south-east (Bhargava,

1979). In the Nainital syncline, the southeastern part of the Krol belt, well-exposed

Baliana- Krol-Tal groups of rocks are present (Fig.1, 2). The Baliana Group is

represented by the Blaini and Infrakrol formations. Earlier workers included the Infrakrol

within the Blaini Formation (Auden, 1934; Bhattacharya and Niyogi, 1971; Fuchs and

Sinha, 1974; Kumar, 1984; Rupke, 1974; Valdiya, 1980). Later on Shanker et al. (1993)

grouped the Blaini and Infrakrol formations into the Baliana Group, after the type area in

the Baliana nala section of Himachal Pradesh. The Blaini Formation consists of a lower

diamictite at the base and an upper diamictite with limestone at the top; its middle part

comprises alternating sandstone and shale (Brookfield, 1987). The diamictite consists of

pebble and boulder- sized clasts of wacke, slates, phyllites, sandstones, shales and

limestone set in a grey, carbonaceous pelitic matrix. The diamictite beds bear striations

and abrasion marks and at a few localities contain distinct glacial fabrics of elongated

clasts (Jain et al., 1981). Above the diamictites lies a few meter thick pink carbonate (~5-

15 m thick) consisting of thinly laminated iron rich muddy dolomite, interbedded with

thin shale in the upper part. This pink carbonate is conformably overlain by the Infrakrol

Formation (~120- 200 m thick) that includes siltstone and dark black, bleached

carbonaceous shale with chert nodules in its upper part. The chert nodules are wrapped

around by the carbonaceous shale layers (Fig. 3). A few lensoid patches of carbonates are

also seen in some sections. The Infrakrol Formation is overlain by the Mahi (designated

as Krol A), Jarashi (Krol B) and Kauriyala (Krol C, D, E) formations (900-1200m thick)

of the Krol Group. The Mahi/Krol A Formation consists of carbonaceous shale at the

base, with calcareous limestone and minor gypsum in some places which is overlain by

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 the Jarashi/Krol B Formation comprising red and ferruginous shale interbedded with

biohermal stromatolitic dolomite. The overlying Kauriyala/Krol C,D,E Formation

includes massive dark grey and blue dolomite, rhythmic alterations of thinly bedded black

shale with marl and carbonaceous and cherty dolomite (Shanker et al., 1993; Valdiya,

1980). It is succeeded by the carbonaceous shale and chert-phosphorite of the lower Tal

Formation (Valdiya, 1980). In the Nainital syncline, it was observed that the fossiliferous

chert bearing carbonaceous shale occurs in the top part of the Infrakrol Formation. There

is a gradational contact between the Infrakrol and the overlying Mahi/Krol A Formation,

where carbonaceous shale interbedded with limestone grades into thinly bedded

limestone. Chert nodules and bands of Mahi/Krol A as observed in Pachmunda, Krol Hill,

Khanog and Rajgarh synclines, are not present in this section.

3. Material and method

Samples of chert nodules and carbonaceous shale were collected from the Takula-

Khurpatal road (N 29° 21' 37.9'': E 79° 26' 53.1'') and Hanumangarhi sections (N 29° 21'

57.9'': E 79° 27' 35.91'') in the Nainital Syncline (Fig. 1). Identification of various forms

(Fig 4 - 6) is based solely on sectional views of the three dimensionally preserved

permineralized specimens present in the chert nodules. Carbonaceous shale and chert

samples treated with standard palynological techniques did not yield any fossils. Thin

sections were studied under the Olympus BX61 microscope and microfossils were

photographed using Olympus DP71 camera and placed in the repository of the Wadia

Institute of Himalayan Geology, Dehradun (under repository nos. WIMF/A 1091-1100,

3976-3981).

It was observed that the acanthomorphic acritarchs were present both randomly and/or

parallel to the longer axis of the chert nodules. An optical examination of thin sections

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 indicates the presence of a siliceous outer rim and sporadic occurrence of euhedral calcite

crystals. Pyrite is present as scattered framboids and/or euhedral crystals. Raman

spectroscopic analysis of selected acritarchs was performed using Horiba JY-Lab Ram

HR Raman spectrometer with Argon ion laser and Labspec software. The obtained

Raman spectra are closely similar to disordered organic carbon (Fig. 7). Phosphate is also

detected in some microfossils (Fig. 7.C). The results of the Raman shift are within ±1–2

cm-1 with respect to the standard values. The thermal maturation of the organic matter is

well indicated by its dark colour which is also reflected by the Total Organic Carbon

(TOC) (0.89 to 1.20%) and δ13C values (-28.9 to -29.9) obtained from seven chert

nodules.

4. Discussion

In the Krol belt, the base of the Ediacaran Period is represented by the base of pink

carbonates (known as cap carbonate globally; Hoffman et al., 2011 and references

therein) overlying the diamictite of the Blaini Formation. The diamictite bearing Blaini

Formation overlies the metamorphosed deposits of the Simla Group (Valdiya, 1980),

traditionally referred to as the Tonian or lower Cryogenian (Chumakov, 2009). Jiang et

al. (2003) correlated the lower diamictite of the Blaini Formation to the middle

Cryogenian Jiangkou and Gucheng formations of South China on the basis of sequence-

stratigraphic analysis, and the upper diamictite to the upper Cryogenian on the basis of

bio-, litho- and sequence-stratigraphic similarities with the Nantuo Formation of South

China. Negative δ13C values are also evident in the succession. The pink carbonate and

shales of the Blaini Formation are characterized by the negative δ13C values of -2‰ to -

4‰ PDB, the Mahi/Krol A Formation by -9‰ PDB, and the top of the Kauriyala/Krol E

Formation by -12‰ PDB suggesting that these anomalies may be related relate to the

Dounce and Bace anomalies of the Doushantuo Formation of South China and the Zhuya

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 Group and the base of Nemakit-Daldyn Horizon in the Siberian Craton (Kaufmann et al.,

2006). Thus, the upper part of the Blaini Formation, which includes diamictite and pink

carbonate, can be regarded as a deposit of the late Cryogenian / Marinoan glaciation

(Kaufmann et al., 2006; Chumakov, 2009). Moreover, isotopic age of the upper part of

diamictite of the Blaini Formation (643±7Ma; Hofmann et al., 2011) suggests its

correlation to the Nuccaleena Formation of Australia (663±4 Ma), Nantuo Formation of

China (635.2±0.6 Ma), and the Ghaub Formation of Namibia (635.6±0.5) (Condon et al.,

2005; Hoffmann et al., 2004; Hoffman and Li, 2009; Jiang et al., 2003; Zhang et al.,

2008; Zhou et al., 2004).

Well- preserved LAAs from the Infrakrol Formation (Baliana Group) and Krol A

(Krol Group) suggest its potential as one of the best known Ediacaran assemblage (Tiwari

and Knoll, 1994; Tiwari and Pant, 2004; Shukla and Tiwari, 2014). The new Takula-

Khurpatal and Hanumangarhi sections in the Nainital Syncline add additional forms to the

assemblage of Ediacaran large acanthomorphic acritarchs from the chert nodules of the

Infrakrol Formation. The present assemblage consists of Tianzhushania spinosa, T.

polysiphonia, Papillomembrana compta, Schizofusa sp., Paratetraphycus giganteus,

Gloeodiniopsis lamellosa, Sphaerophycus medium, and Unnamed Forms A, B, C. This

assemblage is dominated by Tianzhushania spinosa. Xiao et al. (2014) recognized the

first appearance of Tianzhushania spinosa to be used to define the acanthomorph

biozones for regional and global biostratigraphic correlation of the lower-middle

Ediacaran succession. In the Yangtze Gorges region of south China, two

acanthomorphic acritarch assemblage zones have been established, namely, lower

Tianzhushania spinosa (lower Member II) and upper Hocosphaeridium anozos–

Hocosphaeridium scaberfacium–Tanarium conoideum assemblage (upper Member III)

(Liu et al., 2013, Liu et al., 2014; McFadden et al., 2009; Xiao et al., 2014; Yin et al.,

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 2009, Yin et al., 2011). These are the most diverse and best studied acanthomorph

assemblages of the Doushantuo Formation of South China. The Lower Assemblage Zone

is dominated by Tianzhushania spinosa which does not extend to the Upper Assemblage

Zone. Grey (2005) established five assemblage zones in Australia: 1. the Leiosphaeridia

jacutia-Leiosphaeridia crassa Assemblage Zone (also known as Ediacaran Leiosphaerid

Palynoflora (ELP); 2. the Appendisphaera barbata-Alicesphaeridium medusoidum-

Gyalosphaeridium pulchrum Assemblage Zone; 3. the Tanarium conoideum-Schizofusa

risoria-Variomargosphaeridium lithoschum Assemblge Zone; 4. Tanarium irregulare-

Ceratosphaeridium glaberosum- Multifronsphaeridium pelorium Assemblage Zone; and

5. Ceratosphaeridium mirabilis-Distosphaera australica-Apodastoides verobturatus

Assemblage Zone (zones 2-5 are also known as Ediacaran Complex Acritarch

Palynoflora (ECAP). So far, the T. spinosa biozone is not identified in Australia, and

tentatively reported from Svalbard (Knoll, 1992), though, Grey (2005) suggested that

Tianzhushania zone may lie below the ECAP. The occurrence of Tianzhushania spinosa

in the Infrakrol Formation of India, confirms its coevality to the lower Tianzhushania

spinosa biozone in the Doushantuo Formation of South China, whereas its correlation

with the Australian counterpart is not directly feasible.

A recent analysis of findings from the Krol A Formation from the Khanog and

Rajgarh synclines shows that the forms Appendisphaera grandis and Eotylotopalla

dactylos, similar to the ECAP biozone of Australia, are present and Tianzhushania

spinosa typical of the Lower biozone of South China and Hocosphaeridium anozos and

Tanarium conoideum of Upper biozone of South China are missing. It suggests that the

assemblage may lie somewhere between the Lower and Upper biozone of the Doushantuo

Formation. Further investigations for the forms typical to the upper biozone of South

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 China are still required from the Krol Group to develop a complete biozonation scheme of

the Ediacaran sequence in the Krol belt.

The lower and middle Ediacaran acritarch assemblages play a significant role in the

study of biological and environmental evolution. Among them, the acritarch

Tianzhushania spinosa has a characteristic, large (350-750µm in diameter) multilamellate

non-sculptured vesicle with processes that penetrate the wall layer to support an external

membrane (Zhang et al., 1998; Yin L. and Li, 1978). Tianzhushania is interpreted

variously as planktic copepod eggs, animal resting eggs and embryos thus indicating its

pelagic nature (Knoll, 2000; Van Waveren et al., 1993; Yin et al., 2004; Yin et al., 2007).

A non-metazoan holozoan relation has also been proposed (Huldtgren et al., 2011).

Similarly, the genus Papillomembrana compta is characterized by its large vesicle with

numerous bulbous processes. It is regarded variously as a reproductive structure of a

thallophyte and/or microphytoplankton (Vidal, 1990; Yin et al., 1999) and is considered

to have originated in the Cryogenian Interglacial epoch with FAD older than 635±18 Ma

(Vidal and Moczydłowska, 1995).

The Raman analysis of selected forms (Fig.7) reflects the presence of aromatic

organic compounds which confirms the biogenicity of the recovered forms whereas the

presence of phosphate is noted the unnamed form ‘C’ only. This phosphate may have

either formed in the shallow sea environment where progressively shallow upwelling

cells must have brought the deeper phosphorous enriched oceanic waters (Cook and

Shergold, 1986) or may have been brought by fluvial transport (Bushinski, 1969). It is

also possible that organic phosphorus released to pore waters due to decomposition/

breakdown of organic matter within the microbes formed this phosphate (Berner et al.,

1993; Froelich et al., 1988; VanCapellen and Ingall, 1994).

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 5. Conclusions

In the present study, the potential independent evidence of acanthomorphic acritarchs

has been employed as a proxy for biostratigraphic correlation and paleogeographic

reconstructions. The assemblage recovered from a new section, in Takula- Khurpatal area,

besides the Hanumangarhi and Kilbery sections, is a significant addition to the records of the

earliest Ediacaran acanthomorphic acritarchs form the Lesser Himalaya, India. The presence

of Lower Ediacaran representative taxa Tianzhushania spinosa is now confirmed. Moreover,

the spatial abundance of this form is evident in the Infrakrol Formation. The mode of

preservation and occurrence of Infrakrol acritarchs is comparable to the lower assemblage

biozone of the Doushantuo Formation in the Yangtze Gorges section of South China. It is

interpreted that the Infrakrol Formation is coeval to the Lower Tianzhushania Assemblage

Zone and can be bracketed within the same age range. Zircon U-Pb samples from the

interbedded ash beds within Doushantuo Formation have yielded ages of 635.2±0.6 Ma for

the Member I, 632.5±0.5 Ma for the Member II, and 551.1±0.7 Ma for the Member IV

(Condon et al., 2005) which are consistent with the youngest zircon age of 643 ± 7 Ma given

by Hofmann et al., (2011) for tillites of the Blaini Group. A comparison with the Australian

counterpart is not directly feasible due to absence of Tianzhushania spinosa in the Australian

assemblage zones.

During the late Neoproterozoic time a close paleogeographic connection between

South China and India was suggested (Jiang et al., 2002), which led to the deposition of

sedimentary rocks in India and South China along a passive margin with thinned continental

crust due to rifting processes on the Rodinia shelf (Jiang, 2006). The detrital zircon data from

the Blaini diamictites also supports this notion (Hofmann et al., 2011). This interpretation is

further strengthened by the palaeomagnetic data (e.g. Duan et al., 2011; Evan, 2009; Jiang et

al., 2003a; Li and Powel, 2001; Yu et al., 2008).

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 An independent evidence of acanthomorphic acritarchs also indicates that Northern

India was located near South China during the Ediacaran Period. It is also suggested that the

sediments of the Krol Group of Northern India and Doushantuo Formation of South China

were deposited in similar low latitude depositional environments which were probably most

suitable for the origin and early diversification of metazoans just after the snowball earth

period. Employing the present records along with future high resolution biostratigraphic

studies, a biozonation scheme can be proposed within the Krol belt of the Lesser Himalaya in

India.

6. Systematic Description

6.1. Group

Acritarcha Evitt, 1963

6.2. Genus

Tianzhushania Yin and Li, 1978 emend. Yin, Zhou and Yuan, 2008

6.3.Type species

Tianzhushania spinosa Yin L. and Li, 1978 emend. Yin C. in Yin C. and Liu, 1988

6.3.1. Tianzhushania spinosa [Fig. 4. (B)-(E)]

6.3.1.1. Description. Large double walled compressed vesicle, 725-860 µm in diameter

with regularly distributed hollow cylindrical processes connecting the inner and

outer walls through a fine layer. The length of processes varies from ~58 to 68 µm

and diameter from ~4 to 8 µm.

6.3.1.2. Material. Sixty two specimens including twenty well preserved ones.

6.3.1.3. Occurrence. Doushantuo Formation in the Yangtze Gorges (Awramik et al.,

1985; Zhang et al., 1998; Yin C. et al., 2001a, 2011a; Yuan et al., 2002; Yin L. et

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 al., 2007, 2008; Zhou et al., 2007; Xiao, 2008; McFadden et al., 2009; Xiao et al.,

2010, 2012; 2014; Liu et al., 2012a, 2013), Changyang (Yin L. and Li, 1978; Yin

C. and Liu, 1988; Yin C., 1990; Yin C. and Gao, 1995; Zhang et al., 1998; Yin C.

et al., 2001a), Baokang (Yin C. and Liu, 1988), Weng’an (Yin C., 2001; Yin C. et

al., 2001a, 2001b, 2003, 2004, 2007), and Zhangcunping (Liu et al., 2009; Chen et

al., 2010) area, Svalbard (Knoll, 1992), and from the Infrakrol Formation, India as

an unidentified specimen (Plate1.fig. 6 of Tiwari and Knoll, 1994).

6.3.1.4. Remarks. The specimens are similar to T. spinosa Yin and Li, 1978 having

double membranous vesicles with processes and differ from Tianzhushania

tuberifera (characterized by polygonal sculptures) in having a vesicle with longer

cylindrical processes.

6.3.2. Tianzhushania polysiphonia [Fig. 4 (A)]

6.3.2.1. Description. Double-walled vesicle with maximum diameter of ~647 µm,

possessing cylindrical processes distributed in clusters of three (or more)

processes per cluster, connecting the inner (~584 µm) and outer membranes (~647

µm) of the vesicle. Length of processes is ~60 µm.

6.3.2.2. Material. One well preserved specimen.

6.3.2.3. Occurrence. Similar forms are reported from the Doushantuo Formation of

Weng’an, Changyang, Zhangcunping, and Yangtze Gorges areas (Yin and Liu,

1988; Yin, 1990; Yin et al., 2009; Chen et al., 2010; Liu et al., 2012; Liu et al.,

2013)

6.3.2.4. Remarks. Zhang et al. (1998) regarded the characteristic clustered processes of T.

polysiphonia as an intraspecific variation of T. spinosa. Whereas, Xiao et al.

(2014) considered all the specimens of T. spinosa characterized by clustered

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 processes from the Doushantuo Formation (Zhang et al. 1998; Yin C., 2001; Yin

C. et al., 2001a, 2001b, 2004) as T. polysiphonia.

6.4. Genus

Papillomembrana Spjeldnaes, 1963, emend.

6.5. Type species

Papillomembrana compta Spjeldnaes, 1963, emend. Vidal, 1990

6.5.1. Papillomembrana compta [Fig. 5. (A, B)]

6.5.1.1. Description. Large compressed vesicle with tightly arranged hollow, cylindrical

processes. Vesicle diameter is ~ 600 µm (including processes). Length of

processes ranges from ~ 17 to 32 µm and diameter from ~6 to 9 µm at the base

and ~4 to 7 µm at the tip.

6.5.1.2. Material. Four well preserved specimens recovered.

6.5.1.3. Occurrence. Doushantuo Formation at Yangtze Gorges (Zhang et al., 1998; Yin

C., 2001), Baokang of Hubei Province (Yin C. et al., 2009b, 2011a), and Weng’an

(Yin, C., 2001; Yin C. et al., 2007), Biskopas conglomerate in South Norway

(Spjeldnaes, 1963; 1967; Vidal, 1990) and Svalbard (Knoll, 1992).

6.5.1.4. Remarks. The genus Papillomembrana is characterized by themedium to large

vesicles with large hollow and distally bulbous processes. Vidal (1990) emended

the genus including large vesicle with hollow processes that are distally bulbous.

Xiao et al. (2014) further modified Vidal’s (1990) emendation and include smaller

vesicles, shorter processes, and less process density. Xiao et al. (2014) noted that

only specimens from southern Norway (Spjeldnaes, 1963, 1967; Vidal, 1990) and

Svalbard (Knoll, 1992) have bulbous processes and specimens described from

ϭϲ

 Doushantuo Formation do not have this characteristic feature and may represent

different species.

6.6. Genus

Schizofusa Yan, 1982

6.7. Type species

Schizofusa sinica Yan, 1982

6.7.1. Schizofusa sp. [Fig. 6 (D)]

6.7.1.1. Description. Large compressed ellipsoidal vesicle ~ 110 µm long and ~50 µm

wide ruptured into two equal parts along a median axis.

6.7.1.2. Material. Two specimens with one well preserved form.

6.7.1.3. Occurrence. Similar forms are present within the Upper assemblage (Member III)

of the Doushantuo Formation in Tianjiayuanzi, Xiaofenghe and Wangfenggang

sections in China (Liu et al. 2013, 2014), Pertatataka Formation in Australia

(Grey, 2005), and the Ura Formation in Siberia; Sergeev et al. 2011).

6.7.1.4. Remarks. Ellipsoidal vesicle having a slit like aperture is the characteristic feature

of the genus Schizofusa. In general, Schizofusa resembles Leiosphaeridia, but its

mode of excystment differentiates it from the latter (Grey, 2005). Wang et al.,

(2015) considered Schizofusa as a later stage of Leiosphaeridia.

6.8. Kingdom

Eubacteria Woese & Fox, 1977

6.9. Genus

Archaeophycus Wang, Zhang, and Guo, 1983

6.10. Synonyms

ϭϳ

 Archaeophycus yunnanensis Song in Luo et al., 1982 n. comb.

Bigeminococcus grandis: synonym of A. yunnanensis

Paratetraphycus Z. Zhang, 1985, emend. Y. Zhang et al., 1998

Paratetraphycus giganteus and Tetraphycoides multa: junior synonyms of A.

Yunnanensis (Dong et al., 2009)

6.11. Type species

Archaeophycus venustus Wang, Zhang, and Guo, 1983

6.11.1. Archaeophycus sp. [Fig. 6(A)]

6.11.1.1. Description. Spheroidal to irregular clusters of cells occur as isolate, dyads,

triads or tetrads with a diameter ranging from 20 to 50 µm. Cell wall is smooth.

6.11.1.2. Material. Three well preserved specimen recovered.

6.11.1.3. Remarks. The present specimens are characterized by the large size of

individual cells (mostly 20 to 28 µm in diameter) and tight clustering of cells

similar to Tetraphycus yunnanensis Song in Luo et al. (1982) from upper

Cambrian successions in eastern Yunnan Province of South China, hence referred

to the genus Archaeophycus.

6.11.1.4. Occurrence. Similar forms are reported from the Upper Ediacarian to basal

Cambrian successions in eastern Yunnan Province of South China (Luo et al.,

1982).

6.12. Genus

Gloeodiniopsis Schopf, 1968, emend. Knoll and Golubic, 1979

6.13. Synonym

ϭϴ

 Bigeminococcus Schopf and Blacic, 1971

Eozygion Schopf and Blacic, 1971

Eotetrahedrion Schopf and Blacic, 1971

6.13.1. Type species

Gloeodiniopsis lamellosa Schopf, 1968, emend. Knoll and Golubic, 1979

6.13.2. Gloeodiniopsis lamellosa [Fig. 6(G)]

6.13.2.1. Description. Solitary spheroidal vesicles ~50- 55 µm diameter containing a

lamellated envelop of ~5-8 µm. Two cells present within the vesicle probably

indicates a binary cell division. Dark inclusions sporadically present in the

individual cells.

6.13.2.2. Material. Two specimen recovered

6.13.2.3. Occurrence. Similar forms are reported from the upper Tindir Group,

Northwest Canada and Bitter Springs Formation (Allison and Awramik, 1989;

Schopf, 1968; Sergeev et al., 2012), Doushantuo Formation, China (Yuan and

Hoffman, 1998).

6.13.2.4. Remarks. Gloeodiniopsis has characteristic multilamellate sheaths (Schopf,

1968; Schopf and Blacic, 1971; Knoll and Golubic, 1979) and their solitary and

globular nature indicates that they were of a planktonic nature. These forms are

most wide spread among the Proterozoic silicified microbiotas (Sergeev et al.,

2012).

6.14. Genus

Sphaerophycus Schopf, 1968

6.15. Synonymy

ϭϵ

 Sphaerophycus wilsonii Knoll, 1982

Tetraphycus conjunctum Lo, 1980

Gloeodiniopsis aff. gregaria Knoll and Golubic, 1979 (partim) Sergeev, 1993

6.16. Type species

Sphaerophycus medium Horodyski and Donaldson, 1980

6.16.1.1. Sphaerophycus medium [Fig. 6(I)]

6.16.1.2. Description. Isolated cells of ~2- 6 µm in diameter that are completely filled

with dark carbonaceous material.

6.16.1.3. Material. Two well preserved specimens.

6.16.1.4. Remarks. The present specimens fit well in the size range of S. medium

described by Horodyski and Donaldson, (1980).

6.16.1.5. Occurrence. Tindir Group, Northwest Canada and Bitter Springs Formation

(Allison and Awramik, 1989), Uluksan Group , Canada (Hofmann and Jackson,

1991) and Scandinavia in Northern Europe (Butterfield et al. 1994)

6.17. Unnamed Forms

6.17.1. Unnamed Form ‘A’ [Fig. 6 (C)]

6.17.1.1. Description. Compressed vesicle of 750-780 µm diameter possessing large

densely spaced long processes ~70-90 µm in length that overlap each other. Dark

particulate matter is present within the vesicle and width of base and top cannot be

measured.

6.17.1.2. Material. Two specimen with one well preserved specimen observed.

6.17.1.3. Remark. This form is similar to Tianzhushania spinosa in its vesicle size but

differs in longer size of processes and absence of outer wall in the latter.

6.17.2. Unnamed Form ‘B’ [Fig 6. (B,E,F)]

ϮϬ

 6.17.2.1. Description. Ovate to elongate structures, 150 to 600 µm in length and 100 to

600 µm in width. A thin 50-80 µm membrane connects the outer and the inner

walls. The inner wall is irregularly lined with carbonaceous granular particulate

material in some forms. In some specimens clusters of processes like structures of

~100 µm length are seen on both the ends.

6.17.2.2. Material. Eighteen well preserved specimen observed

6.17.2.3. Remarks. These forms appear to represent the transverse/tangential section of

some ellipsoidal vesicular structure. Raman spectra of their central part shows

presence of phosphatic material whereas the inner and outer membrane is

carbonaceous.

6.17.3. Unnamed form ‘C’ [Fig. 6 (H)]

6.17.3.1. Description. Large spherical vesicle, ~60 µm in diameter with smooth to

granular outer lining. Spirally arranged cells of ~10 µm diameter are present

within the vesicle.

6.17.3.2. Material. One well preserved specimen recovered

Acknowledgements

The authors thank Professor Chongyu Yin (Beijing) for his help in identification of
microbiota. The authors are grateful to Professor V.N. Sergeev and two other unknown
reviewers for their constructive comments and suggestions and thank Director, Wadia
Institute of Himalayan Geology, Dehradun for providing the necessary facilities.

Ϯϭ

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CAPTIONS OF FIGURE
Fig. 1
Geological map of the study area showing the sample locations (modified after Valdiya,

1988).

Fig. 2

Outcrop photograph of the Takula- Khurpatal section; inset shows chert nodules wrapped in

carbonaceous shale of the Infrakrol Formation. (Height of the person is 5feet; coin is shown

as scale for nodules).

Fig. 3

Lithologs. (A) Generalized litholog of Baliana and Krol Group (modified after Jiang et al.,

2002 and Hoffman et al., 2011). (B) Litholog of the Takula- Khurpatal section. (C) Litholog

of the Hanumangarhi section.

Fig. 4
Tianzhushania Yin and Li, 1978 emend. Yin, Zhou and Yuan, 2008. (A) Tianzhushania

polysiphonia Yin C. in Yin and Liu, 1988; (i-ii): magnified view of clustered processes

connecting the inner and outer membrane, WIMF/A 1091. (B-E) Tianzhushania spinosa Yin

and Li 1978 emend. Yin C. in Yin C. and Liu1988; inset in (B): magnified view of the

cylindrical processes connecting the inner and outer membrane. WIMF/A 1092-1095. (Scale

bar for A, B, C, D, E = 200 µm; for all insets = 20 µm)

Fig. 5

Papillomembrana compta Spjeldnaes, 1963, emend. Vidal, 1990. (A-B) Papillomembrana

compta; (A) inset (i): magnified view of longitudinal section of processes; inset (ii):

magnified view of transverse section of processes, WIMF/A 1096, 1097 (B) inset (i, iii)

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magnified view of hollow cylindrical processes, inset (ii): magnified view of process

emerging from the vesicle. (Scale bar for A = 50 µm; B = 100 µm; for insets is = 20 µm).

Fig. 6
(A) Archaeophycus yunnanensis Wang, Zhang, and Guo, 1983; WIMF/A 1098. (B) Unnamed

Form ‘B’, WIMF/A 1099. (C) Unnamed form ‘A’ , WIMF/A 1100; (D) Schizofusa sp.,

WIMF/A 3979. (E-F) Unnamed form ‘B’ WIMF/A 3976, 3977. (G) Gloeodiniopsis

lamellose Schopf, 1968, emend. Knoll and Golubic, 1979, WIMF/A 3980. (H) Unnamed

form ‘C’, WIMF/A 3978. (I) Sphaerophycus medium, Horodyski and Donaldson, 1980,

WIMF/A 39811. (Scale bar for A = 25 µm; B, C, F = 100 µm; D, G, H = 20 µm; E = 200 µm;

I = 10 µm).

Fig. 7
Raman spectra of Tianzhushania and Unnamed Forms: (A-B). Trend and major peaks for

Tianzhushania polysiphonia and T. spinosa (arrows mark the points analysed). (C) Trend and

major peak of Unnamed form ‘B’ (arrows mark the points analysed).

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Highlights:

• The occurrence of Tianzhushania in Infrakrol Formation of India, suggests its

coevality to the lower Tianzhushania assemblage of Doushantuo Formation, south
China.
• The present assemblage of Tianzhushania spinosa, T. Polysiphonia,
Papillomembrana compta, Schizofusa sp. Paratetraphycus giganteus,
Gloeodiniopsis lamellosa, Sphaerophycus medium and Unnamed forms A, B and
C is a significant addition to the record of earliest Ediacaran acritarchs from
Infrakrol Formation.
• The age of Infrakrol Formation may be bracketed within the age of Doushantuo
Formation i.e. 635.2±0.6 Ma to 551.1±0.7 Ma.

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