GeolCarp_Vol68_No5_464

GEOLOGICA CARPATHICA

, OCTOBER 2017, 68, 5, 464–478

doi: 10.1515/geoca-2017-0030

www.geologicacarpathica.com

Asteriacites and other trace fossils from the Po Formation

(Visean–Serpukhovian), Ganmachidam Hill, Spiti Valley

(Himalaya) and its paleoenvironmental significance

BIRENDRA P. SINGH

, OM N. BHARGAVA

, RADEK MIKULÁŠ



, SUBHAY K. PRASAD

GARRY SINGLA

and RAMANPREET KAUR

Center of Advanced Study in Geology (CAS), Panjab University, Chandigarh 160014, India

Honorary Scientist (Indian National Science Academy), 103, Sector-7, Panchkula (Haryana), India

Czech Academy of Sciences, Institute of Geology, Rozvojová 269, Prague, Czech Republic;



mikulas@gli.cas.cz

(Manuscript received November 16, 2016; accepted in revised form June 9, 2017)

Abstract: An assemblage of trace fossils comprising Asteriacites stelliformis, A. quinquefolius, Biformites insolitus,

Helminthoidichnites? isp., Lingulichnus isp., Lockeia siliquaria, Palaeophycus tubularis, Planolites isp., Protovirgularia

isp. A, Protovirgularia isp. B, Protovirgularia isp. C, Psammichnites isp., Rusophycus isp., and Treptichnus isp. from the

Po Formation (Visean–Serpukhovian) exposed along the base of Ganmachidam Hill near the village of Chichong, Spiti

Valley in the Himalaya, is described. Storm beds (tempestites) are highly bioturbated. Sedimentary structures such as

hummocky cross-stratification (HCS), low-angle planar and trough cross beds, and shallow, slightly asymmetrical gutter

casts are observed. The overall trace fossil assemblage indicates the presence of upper shoreface to lower shoreface

Cruziana ichnofacies of an open shelf.

Keywords: Po Formation, Visean–Serpukhovian, Spiti, Trace fossils, Cruziana ichnofacies.

Introduction

The trace fossil assemblage reported here derives from the

Po Formation (Visean–Serpukhovian) cropping out at the base

of Ganmachidam Hill, close to the Chichong village in the

Spiti Valley of the Tethyan Himalaya (Figs. 1, 2). The Po

Formation is exposed in the northwestern (Losar section),

northern (Lingti Valley) and northeastern (Poh–Kaurik

section) parts of the Spiti Valley, forming an arch-like pattern.

It is not deve loped in the central and southern parts of the

Spiti Valley and is entirely missing in the adjacent Kinnaur

region.

The Po Formation constitutes a part of the Kanawar Group

(Hayden 1904; Bhargava & Bassi 1998), exposed at various

sites at Poh (old spelling Po), Thabo, Ganmachidam Hill

(Chichong village, SW of Losar), Takche Nala, Kabjiama

Nala, upper Lahaul and Zanskar valleys (Hayden 1904;

Srikantia l981; Gaetani et al. 1986; Bagati 1990; Vannay 1993;

Garzanti et al. 1994; Bhargava & Bassi 1998). It conformably

overlies the siliciclastic-carbonate Lipak Formation (Givetian

to Tournaisian; Draganits et al. 2002) which includes gypsum

beds in its upper part, particularly at Dhuna Dangse (in the

vicinity of Takche) and Hurling–Shalkar stretch in the lower

Spiti Valley (Hayden 1904; Bhargava & Bassi 1998). The Po

and the overlying Ganmachidam formations are intimately

related and nowhere developed independently; they are

either both present or both absent. The absence of the Po–

Ganmachidam Formations in most parts of the Himalaya is

related to shallowing/regression of the sea initiated during the

deposition of gypsum in the upper part of the underlying Lipak

Formation (Bhargava & Bassi 1998). As a consequence, the

sedimentation of the Po Formation was confined to the distal

and deeper parts of the erstwhile basin. The southern and

central parts not only remained a topographically positive area

but were also uplifted during the late Carboniferous and con-

tributed to diamictons of the Ganmachidam Formation (?early

Pennsylvanian–early Asselian). In some parts even the under-

lying Lipak Formation (Givetian–Tournaisian) was also

eroded; as a result, the Gechang Formation (Asselian–

Sakmarian) and Gungri Formation (Wuchiapingian–early

Changhsingian) were deposited over the Devonian Muth

Formation during the Permian transgression (Bhargava &

Bassi 1998; Fig. 1b). However, Gaetani & Garzanti (1991) and

Garzanti et al. (1994, 1996) considered that the selective

development of the Lipak Formation results from the origin of

complex half-graben structures related to the opening of the

Neo-Tethys.

The Po Formation in the vicinity of its type section at Poh

village is 600 m thick and contains plant fossils, nautiloids,

crinoids, bryozoans and other invertebrate fauna (Hayden

1904; Ranga Rao et al. 1984; Bhargava & Bassi 1998). At

Thabo (old spelling Tabo), the basal part of the Po Formation

contains plant fossils which were named as the “Tabo Plant

Stage” (Hayden 1904). The Po Formation, on the basis of

brachiopod and associated fauna, is considered to range from

Visean to Serpukhovian (Bhargava & Bassi 1998).

A wide range of depositional settings have been suggested

for the Po Formation, including shallow marine (Hayden

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1904; Ranga Rao et al. 1984), shallow inner shelf to

shoreface with high sediment input (Shanker et al. 1993),

tide- dominated coastal setting to middle shelf (Garzanti et

al. 1996), and middle shelf to upper shoreface (Bhargava &

Bassi 1998).

Previous workers also reported trace fossils including

Asteriacites, Aulichnites, Gyrochorte, Phycodes, Planolites,

Rusophycus, Rhizocorallium and Skolithos from the Po

Formation (Bhargava & Bassi 1998) but the trace fossils have

never been subjected to ichnotaxonomic considerations or

analysed in terms of their paleoenvironmental implications.

This study presents a fresh collection of trace fossils from

the Po Formation and includes documentation of 11 ichno-

genera comprising 14 ichnospecies, including resting traces

of asteroids and ophiuroids (ichnogenus Asteriacites von

Schlotheim, 1820) found in the Po Formation (Visean–

Serpukhovian) exposed along the slope of Ganmachidam Hill,

Spiti region of northwestern Himalaya.

Stratigraphic section and lithological details

Along the Chichong village (the present study section),

the upper part of the Po Formation (up to 163 m thick) is

exposed at the base of Ganmachidam Hill (Figs. 1, 2). It com-

prises dark blackish shale, fine-grained siltstone and massive

to bedded sandstone beds. We measured the lowermost 65 m

of the section at the base of Ganmachidam Hill which is exten-

sively bioturbated in three specific levels (Fig. 3). The measu-

red section consists of black shales, fine-grained siltstones/

sandstones and medium-grained thickly bedded to massive

sandstone beds (Fig. 3). The lithological contact between the

Po Formation and the underlying Lipak Formation is fully

covered by debris at the base of the measured section. The Po

Formation is conformably overlain by the Ganmachidam

Diamictite Formation (?Bashkirian–early Asselian).

The measured section is subdivided into seven vertically

stacked upwards-coarsening parasequences (PA to PG),

separated by transgressive surfaces (TS1

to TS6) (Fig. 3). An ideal parasequence

is sub

divided into three units (A–C)

(Fig. 2b). Unit A is represented by trans-

gressive dark shale, B by strongly biotur-

bated alternation of fine-grained sand stone,

shale and siltstone intervals, and C by

unbioturbated medium-grained thickly

bedded sandstone beds. However, in

TS 2, 3 and 6, Unit A is directly overlain

by Unit C, indicating a rapid shallowing

and an abrupt influx of coarser material.

The parasequences (PA) comprise tem-

pestites composed of prominent shale–

siltstone-thinly bedded fine-grained

sand stone, and preserve the maximum

number and diversity of trace fossils.

TS 6 to 7 are 2.7 to 23 m thick, respec-

tively; the thickness of the parasequences

decreases in the middle part of the section

and then increases upward. Bioturbate

structures were recorded in parasequences

PA, PE and PG, conspicuously in the tem-

pestite parts. Overall, the frequency and

diversity of trace fossils decrease upward

in these parasequences.

Unit A is interpreted to have been depo-

sited in a low-energy setting (offshore).

Fine-grained sandstone and siltstone beds

(Unit B) in the lower 12.2 to 22.7 m of

the section exhibit hummocky cross-

strati fication (HCS) (Fig. 4d,c), low-angle

planar and trough cross beds (Fig. 4f, g)

and a shallow, slightly asymmetrical gutter

cast at the base of the unit (Fig. 4d, e).

The gutter cast is square-shaped (i.e., flat-

based), 56 mm deep and 148 mm wide.

Ripple cross-lamination is the dominant

Fig. 1. a — simplified geological map of the study area near the Chichong village and

Ganmachidam Hill in the NW part of the Spiti Basin (modified after Bhargava & Bassi

1998); b — lithostratigraphic classification scheme of the Po Formation in Spiti region of the

Tethyan Himalaya (after Bhargava & Bassi 1998).

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GEOLOGICA CARPATHICA

, 2017, 68, 5, 464–478

internal sedimentary structure in fine-grained sandstone and

siltstone, although low-angle cross bedding and oscillatory

cross-lamination are also present (Fig. 4f, g). The transition

from offshore mudstone (Unit A) to the hummocky cross-

strati fied fine-grained sandstone–siltstone interval (Unit B)

indicates a deposition in storm-dominated shoreface. This unit

is strongly bioturbated in the lowermost parasequence (PA)

and partially bioturbated in other parasequences (PE and PG).

The recorded trace fossils include Asteriacites stelliformis,

A. quinquefolius, Biformites insolitus, Helminthoidichnites?

isp., Lingulichnus isp., Lockeia siliquaria, Palaeophycus

tubularis, Planolites isp., Protovirgularia isp. A, Proto

virgularia isp. B, Protovirgularia isp. C, Psammichnites isp.,

Rusophycus isp. and Treptichnus isp. Microbial mat structures

(MISS) found in the lower part of the section are confined

to interfaces of fine-grained siltstone and sandstone within

Unit B.

Unit C represents the upper shoreface. Upward in the sec-

tion, the cycles are thinner with a higher proportion of sand

together with a decrease in the frequency and abundance of

HCS beds and bioturbation. The abundance of trace fossils in

the tempestite sequence suggests that the storm flushed not

only silt/sand, but also nutritive material creating ideal condi-

tions for proliferation of fauna. These parasequences record

shoreline progradation (cf. van Wagoner et al. 1990) and

reflect a progressive increase in hydraulic energy, sand content

and mobility of the substrate which, in turn, controlled the

distribution of trace fossils in the stratal packages of the

Po Formation.

Systematic ichnology

The collected specimens that are described in the following

chapter are housed in the collection of the Centre of Advanced

Study in Geology, Panjab University, Chandigarh, India.

Asteriacites von Schlotheim, 1820

Star-shaped burrows consisting of five arms departing from

a central discoid area and tapering towards the tips, preserved

as convex hyporelief at the base of hummocky cross-stratified

sandstone and siltstone interval in the lower and middle part of

the studied section. High-density (per m

) occurrences are not

Fig. 2. a — field photograph of the section measured at the base of the Ganmachidam Hill near Chichong village, Spiti region of Northwest

Himalaya; b — an ideal upward-coarsening parasequence with sub-environments (Unit: A–C); c — highly bioturbated unit B of the para-

sequence PA; d — transition from parasequence PA to PB.

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observed but isolated specimens are randomly distributed on

the sole of HCS beds. Two species of Asteriacites, namely

A. stelliformis and A. quinquefolius, were identified based on

morphometric analysis (Fig. 5) (maximum width vs. maxi-

mum length of arm) suggested by Knaust & Neumann (2016).

These two ichnospecies differ from A. lumbricalis which is

restricted to imprints with slender and elongated arms

emerging from a discoid centre (produced by an ophiuroid).

Asteriacites is generally interpreted as a resting trace of

asterozoans, such as sea stars (Asteroidea) and brittle stars

(Ophiuroidea) and has been reported from shallow marginal-

marine to deeper sea from the Cambrian to Recent (Seilacher

1953; Crimes & Zhiwen 1986; West & Ward 1990; Mikuláš

1992; Mángano et al. 1999; Chen & McNamara 2006; Schatz

et al. 2013; Baucon & Carvalho 2016). It was originally

described as a body fossil (Walch 1773). Recently, Schlirf

(2012) considered Asteriacites as a nomen dubium (conside-

ring the type material of the type ichnospecies is lost) and

recommended its discontinuation in favour of Heliophycus.

Therefore, the taxonomical attri-

butes of the Asteriacites were

disputed until recently (Knaust

2012; Schlirf 2012; Paranjape

et al. 2013; Gurav et al. 2014).

Rediscovery of the type material

in the von Schlotheim collection

and an efficient revision of

the ichnogenus Asteriacites by

Knaust & Neumann (2016)

justified the validity of Asteria

cites von Schlotheim, 1820 as

an ichno genus.

Asteriacites stelliformis

(Miller and Dyer, 1878)

Fig. 6a, d; Fig. 7b, c, e

Material: Approximately one

third from the overall number of

76 specimens of Asteriacites

ispp. documented in the outcrop

(29 specimens of Asteriacites isp.

collected) belong to A. stelli

formis. The remaining two thirds

of Asteriacites isp. belong to

A. quinquefolius or cannot be

determined on the ichnospecies

level.

Star-shaped imprints (convex

hyporeliefs) consisting of rela-

tively short arms. Most speci-

mens bear five arms, but only

a few specimens show approxi-

mately even angles between the

neighbouring ones. For a regular

pentagram, the angle of 72° is

determined, which is close to that in some specimens (e.g.,

Fig. 6d). Angles measured on the figured specimens (Figs. 6a, d;

7b, c, e) range from 26° (Fig. 7e) to 120° (Fig. 6a); the diffe-

rence in angles can be understood by movements of the trace-

maker rather than by water currents. The length of arms varies

between 10 and 50 mm.

Asteriacites arms sometimes exhibit broad U-shaped or

chevron-like imbricate ornamentation. Such features are

attributed to the tube feet / ambulacra activity during the

burrowing process (Seilacher 1953).

Asteriacites quinquefolius Quenstedt, 1876

Fig. 6 b–c, e–f; Fig. 7a, d, f

Material: See the listing of specimens of A. stelliformis.

Star-shaped imprints (convex hyporeliefs) composed of

typi cally five wide arms; width:length ratio of the arms is less

than 1:2. The ground plan of the specimens varies from

a nearly regular pentagram (Fig. 7a) to asymmetrical

Fig. 3. Lithocolumn of the Po

Formation measured at the

Chichong village (base

of the Ganmachidam Hill)

showing coarsening upward

parasequences (PA–PG) and

sub-environments (Unit A–C),

and distribution of observed

and recorded trace fossils and

sedimentary structures.

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structures still consisting of five arms (Figs. 6e, 7d). The sur-

face of the arms is typically ornamented by fine chevron-like

(Fig. 6e) imbricate (Fig. 7d) structures, similarly to that of

A. stelliformis (described above). In certain specimens,

some arms (typically one arm) show the width:length ratio

higher than 1:2 but most arms have the ratio less than 1:2.

In such cases (Fig. 7d, f), we determine these specimens as

A. quinquefolius.

Biformites Linck, 1949

The ichnogenus of Biformites Linck, 1949 was recently

revised by Schlirf (2012) and Knaust & Neumann (2016) as

narrow, bedding-parallel, vermiform, hook-shaped or sinuous

imprints with slightly tapering terminations, unbranched or

with secondary successive branching, with or without orna-

ment. Knaust & Neumann (2016) erected the ichnofamily

Biformitidae for imprints resulting from locomotion using

asterozoan arms, selected Biformites Linck, 1949 as the

type ichnogenus, and included the other ichnogenera Arcichnus

Sutcliffe, 1997, Harpichnus Vallon et al., 2015 and

Pentichnus Maerz et al., 1976 under this family. They also

stated that ichnogenera Zhadaichnus Yang and Song, 1985

and Ophioichnus Bell, 2004 are junior synonyms of

Biformites.

Biformites insolitus Linck, 1949

Fig. 8f

Material: Three specimens; all collected.

Narrow (0.2–0.6 cm), short (0.5–2.6 cm), vermiform,

slightly tapering, straight to curved, bisymmetrically arranged

sets of transverse, elongate ridges or protuberances preserved

on lower bedding planes. They can be, considering all aspects

of morphology, determined as Biformites insolitus; this

trace fossil is interpreted as imprints of arms of walking

ophiuroids (cf. Boyer 1979; Schlirf 2012; Knaust & Neumann

2016).

Helminthoidichnites Fitch, 1850

Simple, horizontal, small, thin, unbranched, non-meande-

ring, straight or curved, more rarely circular trails or burrows

that commonly display overlap between specimens but lack

self-overcrossing (Buatois et al. 1998). Helminthoidichnites is

regarded as a grazing trace probably produced by vermiform

animals (Buatois et al. 1998).

Helminthoidichnites isp.

Fig. 8a

Material: Ten specimens observed in the outcrop; three

collected.

Small, mostly straight to slightly curved, smoothly horizontal

trails displaying some angular turns in their path and preserved

as positive hyporelief. No lateral grooves or levees have been

observed. One end of the path of trail displays circular mounds.

Diameter of the trace varies from 0.5 to 1.2 mm and the

figured/collected trail is 47 mm long. The speci men also

shows close morphological resemblance to Haplotichnus

indianensis Miller, 1889 recorded from the lower

Pennsylvanian of Orange County, Indiana (Rindsberg &

Kopaska-Merkel 2005). However, recently Getty & Bush

(2017) convincingly illustrated that Haplotichnus indianensis

Miller, 1889 exhibits bifurcating projections at the bends sim-

ilar to Treptichnus bifurcus; therefore, they synonymized

Haplo tichnus with the ichnogenus Treptichnus. Demircan &

Uchman (2016) considered Haplotichnus indianensis Miller,

1889 as Gordia indianensis. Our specimens do not exhibit the

ichnogeneric character of either Treptichnus or Gordia;

hence they were grouped under the ichnogenus

Helminthoidichnites.

Lingulichnus Hakes, 1976

Vertically to obliquely orientated sediment-filled tubes

with elliptical to sub-circular cross sections (Zonneveld &

Pemberton 2003). Lingulichnus has been interpreted as

a dwelling trace of a lingulid brachiopod (Hakes 1976; Szmuc

et al. 1976; Zonneveld & Pemberton 2003; Zonneveld et al.

2007; Alonso-Muruaga et al. 2013).

Lingulichnus isp.

Fig. 9j

Material: Two bedding planes with dozens of specimens

observed in the outcrop and collected.

Circular to elliptical, endichnial, vertically to near-vertically

orientated sediment-filled structures preserved in full relief.

They occur as dense assemblages of specimens tightly

arranged (ca. 10 specimens per 2×2 cm; Fig. 9j). The determi-

nation of these trace fossils as Lingulichnus follows the

description of Zonneveld & Pemberton (2003).

Fig. 5. Morphometric analysis of maximum arm length and width of

the 19 specimens of Asteriacites von Schlotheim, 1820 recovered

from basal part of the Po Formation, showing grouping of A. stelli

formis (As) and A. quinquefolius (Aq). Measurement method adopted

from Knaust & Neumann (2016).

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Lockeia James, 1879

Lockeia is a cubichnion produced mostly by bivalves

(Seilacher & Seilacher 1994) in various environments. Lockeia

is known to occur in a wide range of mostly marine environ-

ments from shallow to deep marine and even in non-marine

continental deposits (Seilacher 1953; Pickerill 1977; Bromley

& Asgaard 1979; Crimes et al. 1981; Archer & Maples 1984)

and considered as a resting trace of bivalves (Seilacher 1953;

Osgood 1970; Pickerill 1977; Vossler & Pemberton 1988;

Seilacher & Seilacher 1994) or a dwelling trace (domichnion)

of suspension feeders (Mángano et al. 2002a).

Lockeia siliquaria James, 1879

Fig. 9 e, f, h, k; Fig. 8c

Material: 15 specimens observed in the outcrop; 7 of them

were collected.

Bilaterally symmetrical, almond -shaped, sandstone-filled

bodies tapering at both ends to a sharp point, variously orien-

tated, den sely aggregated (Fig. 9 e), preserved as convex hypo-

reliefs. They are 0.8–1.5 cm long and 0.5–0.8 cm wide. The

central median ridge or crest is poorly developed or missing.

Palaeophycus Hall, 1847

Trace fossils of Palaeophycus are open burrows, horizontal,

slightly sinuous. They are both smooth and ornamented bur-

rows, variable in diameter; the crucial determination features

are a prominent wall lining and a passive fill (Osgood 1970;

Alpert 1975; Pemberton & Frey 1982; Fillion & Pickerill

1990; Stanley & Pickerill 1994; Keighley & Pickerill 1995;

Jensen 1997). Palaeophycus is interpreted as dwelling struc-

tures of suspension feeders or preda tors (Osgood 1970;

Pemberton & Frey 1982; Uchman 1995; Tchoumatchenco &

Uchman 2001; Mángano & Buatois 2003).

Palaeophycus tubularis Hall, 1847

Fig. 9k; Fig. 8a, b, d

Material: About 20 specimens observed in the outcrop;

5 of them collected.

Straight to slightly curved, horizontal, undulose, essentially

horizontal, thinly lined, smooth-walled cylindrical to subcy-

lindrical open burrows preserved as full relief in fine grained

sandstone and siltstone. Burrow diameters and lengths vary

from 2 to 8 mm and 30 to 90 mm, respectively. The burrow fill

is similar to the host rock, typically massive. The burrows

occur very frequently in most of the rocks. Palaeophycus was

found in association with Lockeia siliquaria (Fig. 9 k) as well

as with Helminthoidichnites-type traces (Fig. 8a). One speci-

men of Palaeophycus is preserved as a convex epirelief and

shows a burrow collapse (Fig. 8a).

Planolites Nicholson, 1873

Straight or moderately curved, smooth, exceptionally

branching tunnels of circular outline, parallel to bedding,

Fig. 6. Resting traces Asteriacites von Schlotheim, 1820 burrowed by sea stars (Asteroidea); basal part of the Po Formation at Chichong

(Ganmachidam Hill) section (Spiti region, Himalaya), scale bar =1 cm. a — Asteriacites

stelliformis, CAS/LO/2016/-PO-4; b — A. quinque

folius, CAS/LO/ 2016/-PO-3; c — A. quinque folius, CAS/LO/2016/-PO-15; d — A. stelliformis, CAS/LO/2016/-PO-6; e — A. quinquefolius,

CAS/LO/2016/-PO-9; f — A. quinquefolius, CAS/LO/ 2016/-PO-7.

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filled with material differing from the host rock (

Häntzschel

1975;

Pemberton & Frey 1982).

Planolites is a eurybathic,

extremely facies-crossing ichnogenus and has been interpreted

as a product of vermiform deposit-feeders (Häntzschel 1975;

Pemberton & Frey 1982; Frey & Howard 1985, 1990; Fillion

& Pickerill 1990; Uchman 1995).

Planolites isp.

Fig. 9e, g; Fig. 8e

Material: 16 specimens observed in the outcrop; 5 of them

collected.

Small, simple, unlined, horizontal to slightly inclined,

rarely branched, straight to curved burrows without a wall.

The burrows are 6–35 mm long and 1–4 mm wide. One

of the specimens shows faintly developed transverse

annulae and irregular morphology with thinning at one end

(Fig.

8g). This specimen shows some morphological

aspects of Planolites reinecki (Stanley & Pickerill 1994;

Uchman 1998; Mángano & Buatois 2003) but differs

in the absence of both transverse annulae and longitudinal

striae.

Protovirgularia M’Coy, 1850

The ichnogenus Protovirgularia is described as horizontal

or subhorizontal cylindrical burrows, trapezoidal, almond-

shaped, or triangular in cross section, distinctly or indistinctly

bilobated, occasionally with oval, mound-like terminations of

the trace (Pickerill & Narbonne 1995; Uchman 1998; Uchman

& Gazdzicki 2006; Fernandez et al. 2010). Protovirgularia is

interpreted as being produced by bivalves (Han & Pickerill

1994; Seilacher & Seilacher 1994). Several preservational

variants of Protovirgularia are known (Carmona et al. 2010);

they are attributed to substrate conditions (Maples & West

1989; Mángano et al. 1998; Carmona et al. 2010).

Protovirgularia isp. A

Fig. 9b

Material: About a dozen specimens of Protovirgularia isp.

were observed in the outcrop; collected specimens include

2 specimens of Protovirgularia isp. A, 2 specimens of

Proto virgularia isp. B, and one specimen of Protovirgularia

isp. C.

Fig. 7. Resting trace of Asteriacites von Schlotheim, 1820 burrowed by sea star (Asteroidea); basal part of the Po Formation at Chichong

(Ganmachidam Hill) section (Spiti, Himalaya), scale bar = 1 cm. a — A. quinquefolius, CAS/LO/2016/-PO-8; b — A. stelliformis,

CAS/LO/2016/-PO-1; c — A. stelliformis, CAS/LO/2016/-PO-14; d — A. quinquefolius, CAS/LO/2016/-PO-2; e — A. stelliformis,

CAS/LO/2016/-PO-13; f — A. quinquefolius, CAS/LO/2016/-PO-19.

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This slightly curved, unbranched, horizontal structure con-

sists of a central furrow flanked by regular closely spaced oval

lobes or pads, which are well-preserved only on one side of the

trace (Fig. 9b). The outer margins of some lobes show fine

striations. The specimen is grouped here under a common

epirelief preservational variant of Protovirgularia. A similar

trace from the Vallès-Penedès Basin in NE Spain was described

as Protovirgularia dichotoma (Gibert & Domènech 2008).

Protovirgularia isp. B

Fig. 9h

An elongated and slightly curved trace, 85 mm long and

5–6 mm wide, preserved as a positive hyporelief. Chevron-

like lamellar ornamentation is developed but reduced towards

one of the extremes where Protovirgularia is terminated by

a Lockeia-like body. Lockeia siliquaria is almond-shaped,

smooth, and lacks a well-developed keel and ornamentation.

It shows a compound trace fossil of the resting trace of

Lockeia siliquaria and the locomotion trace of Protovirgularia

(cf. Han & Pickerill 1994). Protovirgularia specimens that

terminate with Lockeia siliquaria are assigned to

Protovirgularia rugosa. Ekdale & Bromley (2001) stated that

L. siliquaria represents the place where the bivalve tempo-

rarily stopped for feeding.

Protovirgularia isp. C

Fig. 8c

A poorly preserved Protovirgularia isp. with effaced upper

surface of the specimen. The trace is 13 mm long and 8 mm

wide, with prominent chevron markings on one side.

Psammichnites Torell, 1870

The ichnogenus Psammichnites Torell 1870 includes a wide

variety of predominantly horizontal, sinuous to looped, back-

filled traces, characterized by a distinctive median dorsal

Fig. 8. Trace fossils from basal part of the Po Formation (Visean-Serpukhovian), Spiti Himalaya, India. a — (He) Helminthoidichnites isp., and

(Pt) Palaeophycus tubularis, CAS/LO/2016/-PO-24; b — (Pal) Palaeophycus tubularis, CAS/LO/2016/-PO-25; c — (Pv) Proto virgularia isp. C,

and (Ls) Lockeia siliquaria, CAS/LO/2016/-PO-28; d — (Pals) Palaeophycus tubularis; e — (Pl) Planolites isp., CAS/LO/2016/-PO-42;

f — (Bi) Biformites insolitus, CAS/LO/2016/-PO-48.

474

SINGH, BHARGAVA, MIKULÁŠ, PRASAD, SINGLA and KAUR

GEOLOGICA CARPATHICA

, 2017, 68, 5, 464–478

attributed to three ichnospecies, namely, P. plummeri (Fenton

& Fenton 1937a), P. grumula (Romano & Meléndez 1985),

and P. implexus (Rindsberg 1994). Psammichnites plummeri is

the most common Carboniferous ichnospecies and is charac-

terized by a relatively straight, continuous dorsal ridge/groove,

fine transverse ridges, a wider size range, and a non-looping

geometric pattern (Mangano et al. 2002b). Psammichnites

grumula differs from the other ichnospecies of Psammichnites

by having median dorsal holes or protruding mounds.

Psammichnites implexus is characterized by its consistently

narrower size range, subtle backfill structure, and tendency to

scribble. Psammichnites records the feeding activities of

a subsurface animal using a siphon-like device.

Psammichnites isp.

Fig. 9i

Material: Two specimens observed in the field; one of them

was collected and measured.

A poorly preserved, nearly straight, smooth, unbranched,

bilobed ridge with a faintly developed median furrow, pre-

served as a convex epirelief. The trace is 5.1 cm long and

1.0 cm wide. The specimen is weathered and shows a faint

meniscate structure; however, the median furrow is clearly

visible along one part of the specimen. No internal backfill

structural pattern is visible. The specimen morphologically

closely resembles Psammichnites plummeri (Fenton & Fenton

1937c) in its pattern of straight ridges but lacks fine, closely

spaced transverse ridges (Mángano et al. 2002b). The speci-

men differs from the other bilobate trace of Archaeonassa in

lacking a broad central flat area between the lobes (Jensen

1997).

Rusophycus Hall, 1852

The ichnogenus of Rusophycus Hall, 1852 refers to a short,

bilobate, coffee-bean shaped trace fossil, usually with

a median furrow. The lobes are transversely wrinkled by ante-

rolaterally orientated fine striae (Häntzschel 1975). Paleozoic

Rusophycus indicates nesting or resting behaviours of trilo-

bites (Osgood 1970). Rusophycus is furthermore interpreted as

resting traces of arthropods (Bergström 1973; Jensen 1990)

and notostracan branchiopods (Bromley & Asgaard 1979).

Rusophycus isp.

Fig. 9a

Material: Three specimens observed in the field; two of

them were collected.

A butterfly-shaped (anterior and posterior, broad openings),

relatively shallow, horizontal, bilobate structure with a distinct

cleft separating two strongly striated lobes. The bilobate struc-

ture is 10 mm wide and 14 mm long, and preserved as positive

hyporelief in fine- to very fine-grained sandstone. Lobes are

strongly convex; median furrow deepens and widens towards

the anterior end of the trace. The scratchings seem to be clearly

clustered in groups and these groups open (or expand) towards

the axial area.

Treptichnus Miller, 1889

Treptichnus Miller, 1889 is a burrow consisting of segments

connected at their ends, each to the next one, characteristically

but not invariably in a zigzag pattern. Ichnotaxonomy of

Treptichnus has been discussed in detail by Maples & Archer

(1987), and Buatois & Mángano (1993). The ichnogenera of

Plangtichnus and Haplotichnus are junior synonyms of

Treptichnus (Buatois & Mángano 1993; Getty & Bush 2017).

Treptichnus is interpreted as feeding structures (fodinichnia)

produced by vermiform animals or insect larvae (Buatois &

Mángano 1993; Getty & Bush 2017).

Treptichnus isp.

Fig. 9d

Material: A depicted and described specimen only.

A burrow system consisting of more-or-less straight six to

seven segments, joined to each other at very blunt angles

(approximately 160°); other segments are orientated at

an oblique angle, namely, 20–40°, to the main axis of the trace.

The trace is 87 mm long; segments are 10–22 mm long and

4–6 mm wide, preserved in hyporelief. The fill is identical to

the host rock.

Microbially induced sedimentary structures (MISS)

Wrinkle structures consisting of ridges surrounding elon-

gated depressions (Fig. 9c) were recorded on the upper bed-

ding plane of a fine-grained sandstone bed in the lower part of

the section. The ridges are flat and horizontal, slightly curved,

and interconnected at some places. Microbially induced sedi-

mentary structures (MISS) result from the response of micro-

bial mats to physical sediment dynamics (Lowe 1980; Walter

et al. 1980; Byerly et al. 1986; Hofmann et al. 1999; Allwood

et al. 2006, 2007, 2009; Noffke 2009). Wrinkle structures

(MISS) are found primarily in wave-dominated, low-energy

shoreface settings (Calner & Eriksson 2011) but also occur in

a wide range of environmental settings (Hagadorn & Bottjer

1997, 1999; Pﬂüger 1999; Bottjer et al. 2000; Noffke et al.

2002, 2006; Noffke 2009).

Paleoenvironmental significance

The recorded trace fossils of Asteriacites stelliformis,

A. quinquefolius, Biformites insolitus, Helminthoidichnites?

isp., Lingulichnus isp., Lockeia siliquaria, Palaeophycus

tubularis, Planolites isp., Protovirgularia isp. A, Proto

virgularia isp. B, Protovirgularia isp. C, Psammichnites isp.,

Rusophycus isp., and Treptichnus isp. are characteristic for the

Cruziana ichnofacies.

475

THE ASTERIACITES FROM THE PO FORMATION IN SPITY VALLEY, HIMALAYA

GEOLOGICA CARPATHICA

, 2017, 68, 5, 464–478

This ichnofacies is characterized by the dominance of hori-

zontal traces of mobile organisms and subordinate presence of

vertical and inclined permanent structures, a wide variety of

ethologic categories, the dominance of deposit- and detritus-

feeding traces, a limited participation of suspension feeders

and predators, and a high ichnodiversity and abundance

(cf.

MacEachern & Pemberton 1992; MacEachern et al. 1999;

Buatois & Mángano 2011). The shallowing-upward, hetero-

lithic interval of interbedded fine-grained sandstone and silt-

stone of the parasequences, highly bioturbated in the lower

part of the section, is interpreted as deposits from a lower

shoreface environment. The heterolithic successions of fine-

grained sandstone and siltstone provided a favourable setting

for the preservation of trace fossils. The dark shale interval at

the base of the parasequences indicates a transgressive phase

(offshore setting), followed by a deposition of heterolithic suc-

cessions of fine-grained siltstone and sandstone (lower shore-

face). Higher up in the section, it is followed by coarser,

thick-bedded sandstone beds indicating an upper shoreface

setting. The high sediment supply can be related to tectonic

uplift in adjacent areas.

Baucon & Carvalho (2016) suggested that Asteriacites-

bearing beds may be considered ichnological proxies for

marine settings, low bioturbation intensity, shallow tiering,

high sedimentation rate and/or event-bed deposition, signifi-

cant levels of hydraulic energy, and low predation pressure.

The presence of hummocky cross stratification, planar and

trough cross bedding, and traces of Cruziana ichnofacies indi-

cate a paleoenvironment slightly above the fair-weather wave

base to the storm wave base, in a zone ranging from the lower

shoreface to the lower offshore setting.

Acknowledgements: BPS is thankful to the University Grant

Commission (UGC, New Delhi) for providing financial sup-

port within grant No. F.20-1/2012(BSR)/20-8(12)/2012(BSR).

ONB acknowledges the Indian National Science Academy

(No. 567-3. 2015) for its ﬁnancial assistance. The work of RM

was supported by the RVO67985831 Project of the Institute of

Geology of the Czech Academy of Sciences. We are thankful

to the reviewers Gabriela Mángano, Alfred Uchman and

Lothar Vallon for constructive reviews of the manuscript.

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