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First report on late Miocene (Tortonian: ~ 11-10 Ma) charophyte gyrogonites

from Tapar, Kachchh District, Gujarat State, western India

Article  in  Proceedings of the Indian National Science Academy · September 2022

DOI: 10.1007/s43538-022-00102-4

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Proceedings of the Indian National Science Academy
https://doi.org/10.1007/s43538-022-00102-4

RESEARCH PAPER

First report on late Miocene (Tortonian: ~ 11–10 Ma) charophyte

gyrogonites from Tapar, Kachchh District, Gujarat State, western India
Nongmaithem Amardas Singh1 · Ningthoujam Premjit Singh2,3 · K. Milankumar Sharma1,5 · Rajeev Patnaik2 ·
R. P. Tiwari1 · Ramesh Kumar Sehgal3 · Vinay Kumar4 · Wasim Abass Wazir2 · Y. Priyananda Singh1 ·
Deepak Choudhari1,2,3,4,5,6

Received: 2 June 2022 / Accepted: 23 August 2022

© Indian National Science Academy 2022

Abstract
This paper describes the first record of charophyte gyrogonites from the late Miocene (Tortonian; ~ 11–10 Ma) Tapar local-
ity of Kachchh, Gujarat State, western India. The recovered charophyte assemblage is constituted by Chara globularis var.
aspera, C. globularis var. globularis, Lychnothamnus cf. sahnii, Lychnothamnus sp. and Nitellopsis sp. In addition, the present
article discusses the palaeoenvironment based on the recorded charophyte gyrogonites (this study) and previously known
faunal data (mainly vertebrates) from the Tapar locality, Kutch Basin, western India. Further, considering the significant
extension of the biostratigraphic range [especially the First Appearance Datum (FAD)] of fauna from the Kutch Basin (in
particular from the Tapar locality), we herein make an attempt to compare biostratigraphically the faunal assemblages from
Kutch with those from the Siwalik Group, north India.

Keywords  Algae · Kutch Basin · Neogene · Palaeoenvironment · Siwalik Group

Introduction record of Charophytes from India, a rich assemblage of cha-

rophyte gyrogonites had been previously recorded within
Charophytes (also known as Charophycean Green Algae) are the sediments belonging to the Siwalik Group, north India
a taxonomically diverse group of extant fresh water and ter- (Bhatia and Mathur 1970, 1978; Tewari and Sharma 1972;
restrial green algae that have been linked to the origin of land Lakhanpal et al. 1976; Suneja et al. 1978; Bhatia 1982). Later,
plants (Domozych et al. 2017). Fossil gyrogonites have been Bhatia (1999) revised the charophyte flora from the Neogene
recovered in non-marine deposits worldwide from the Silu- (Miocene, Pliocene) and Quaternary (Pleistocene) sediments
rian (~ 425 Ma) to the present (Feist et al. 2005; Beilby et al. of the Siwalik Group. As presently understood, 17 taxa of
2018). Considering the Neogene as well as the Quaternary charophytes are known from the Siwalik Group sediments,
north India (see, Bhatia 1999). Charophyte species Nitellopsis
* K. Milankumar Sharma (Tectochara) meriani, N. (T.) huangi and Lychnothamnus sah-
[email protected] nii were also reported from the middle Miocene of Ramnagar,
Jammu and Kashmir, north India (Singh et al. 2018). Recently,
1
Department of Geology, Central University of Punjab, Tiwari and Bhan (2021) reported a diverse assemblage of cha-
Bathinda 151401, India
rophyte species, including Chara globularis globularis, C.
2
Centre for Advanced Studies in Geology, Panjab University, globularis aspera, C. rantzieni, C. hispida, Chara sp., Hor-
Chandigarh 160014, India
nichara maslovi, Lychnothamnus breviovatus, L. barbatus,
3
Wadia Institute of Himalayan Geology, Dehradun 248001, Lychnothamnus sp., Sphaerochara tewarii, Lamprothamnium
India
papulosum, Nitellopsis (Tetochara) huangi, N.(T) meriani,
4
Department of Botany, Central University of Punjab, N. (T) megarensis from the middle Siwalik of Mohand area,
Bathinda 151401, India
Dehradun sub-basin, North-West Himalaya, India (also see,
5
Department of Geology, Central University of South Bihar, Table 1). Although many workers have recorded charophytes
Gaya 824236, India
from the Neogene & Quaternary Siwalik Group sediments of
6
Centre for Advanced Studies in Geology, Panjab University, north India, there has been no report of charophytes from the
Chandigarh, India

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 Proceedings of the Indian National Science Academy

Table 1  Distribution of Taxa Tapar Mohand Pinjor Tatrot Dhok Pathan Chinji Ramnagar
charophyte flora from the
different localities of Neogene Chara globularis var. globularis ✓ ✓ ✓ ✓ ✓
sediments (Bhatia 1999; Singh
Chara globularis var. aspera ✓ ✓ ✓ ✓
et al. 2018; Tiwari and Bhan
2021) Chara rantzieni ✓ ✓
Chara cf. C. hispida ✓ ✓
Hornichara maslovi ✓ ✓ ✓
Lychnothamnus breviovatus ✓ ✓ ✓
Lychnothamnus barbatus ✓
Lychnothamnus sahnii ✓ ✓ ✓
Lamprothamnium succinctum ✓
Lamprothamnium papulosum ✓ ✓
Nitellopsis (Tectochara) bicarinata ✓
Nitellopsis (Tectochara) helvetica ✓
Nitellopsis (Tectochara) huangi ✓ ✓ ✓
Nitellopsis (Tectochara) meriani ✓ ✓ ✓
Nitellopsis (Tectochara) megarensis ✓
Sphaerochara prolifera ✓ ✓
Sphaerochara tewarii ✓ ✓

Neogene (Miocene) interval of Kutch Basin, western India, Geological setting and age
prior to the present investigation.
The Miocene deposits of Kutch are well known for Miocene deposits of Kutch are divided into three litho-
their rich assemblage of Neogene vertebrates comprising logical units i.e. Khari Nadi, Chhasra and Sandhan for-
marine and terrestrial fauna (also see Grant 1840; Wynne mations (Biswas 1992). Khari Nadi Formation has been
1872; Lydekker 1876, 1880; Prasad 1964; Sahni and Mishra biostratigraphically dated as early Miocene (Aquitanian;
1975; Mishra 1976; Bajpai and Domning 1997; Bajpai et al. 23.03–20.43 + / − 0.05  Ma according to Gradstein et  al.
2006; Thewissen and Bajpai 2008, 2009; Patnaik et al. 2014, 2004) based on foraminiferal assemblage. Very recently,
2022). Previously, various vertebrate taxa such as elephants, Kapur et al. (2022) have also provided an Aquitanian age
rhinos, giraffids, equids, suids, crocodiles and turtles were for the Khari Nadi Formation based on nannofossils (Raju
reported from Tapar locality (Kutch) in Gujarat (Bhandari 1974). The overlying Chhasra Formation is correlatable to
et al. 2009, 20155). Later, Bhandari et al. (2018, 2021) dis- the Burdigalian (~ 20–16.2 Ma) based on the presence of
covered hominid remains of Sivapithecus as well as rodents myogypsinid assemblage (foraminifera), whereas the Sand-
from the late Miocene deposits of Tapar (Kutch), Gujarat, han Formation is not assigned to particular age due to lack of
India. Recently, a complete assemblage of vertebrates has diagnostic fauna but it tentatively assigned to Pliocene (Bis-
been reported from the Tapar locality including teleostei was 1992). Based on the mammalian assemblages, the Tapar
fishes (Singh et al. 2019), Deinotherium indicum (Singh locality has been dated to ~ 16 Ma by biostratigraphical cor-
et al. 2020), batoids (Sharma et al. 2021), lizards (Čerňanský relation with the Khari Nadi Formation (Bhandari et al.,
et al. 2021), rodents (Bhandari et al. 2021) and herpeto- 2009). However, the correlation is problematic, since the
fauna (Singh et al. 2022). A recent palaeontological inves- type area of the Khari Nadi Formation is located in western
tigation at Tapar locality of Kutch, Gujarat (India) yielded Kutch whereas the Tapar beds are in central Kutch, and the
charophyte gyrogonites associated with faunal (mammalian, deposits from both localities are not continuous. Bhandari
amphibian, reptilian and piscean) remains (also see, Fig. 5). et al. (2015) subsequently have dated this locality to the late
The present study documents a diverse assemblage of Miocene (~ 11–10 Ma) on the basis of the First Appear-
charophyte gyrogonites for the first time from the late Mio- ance Datum (FAD) of the equid (Hipparion). The fossil site
cene deposits of Tapar locality, Kutch Basin, Gujarat State, (23°15′12.93″N and 70° 8′52.17″E) is located about ~ 8 km
western India. Systematic studies of the collected charo- to the north of the Tapar village. Lithologically, the charo-
phytes have been carried out. In addition, we herein discuss phyte bearing section at Tapar is comprised of shale, sand-
the palaeoenvironmental and biostratigraphic significance of stone, silt, clay and mudclast conglomerate rock (Fig. 1).
the recovered charophytes taking into consideration previous As presently understood, known mammalian data (par-
vertebrate faunal records from the studied section at Tapar, ticularly presence of Hipparion) supports a late Miocene
Gujarat State, western India.

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Fig. 1  A Map of India showing the study area enclosed by red rec- recovered from the Tapar locality. E Enlarged photograph of the fos-
tangular box. B Geological map of the Kutch (study area indicated by sil bearing conglomerate bed. F Photograph of fossils bearing Litho-
the star mark) (Modified after, GSI, 2012). C Litholog of the fossils section of Tapar locality
bearing section of Tapar locality. D Photograph showing charophyte

(~ 11–10 Ma) age for the sedimentary succession at Tapar et al., 2021; Sharma et al. 2021; Made et al. 2022). Fur-
belonging to the Sandhan Formation (Bhandari et  al., ther details are discussed in the biostratigraphy section of
2015, 2018, 2021; Singh et al. 2019, 2020; Čerňanský this study.

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Materials and methods isopolarity index range from 132–151 μm (see Table 2).

Nine to twelve convolutions are visible in lateral views and
About 9000 kg of bulk sample of calcareous ferruginous the width of the spiral cells range from 41–78 μm. The con-
conglomerate was macerated following the methodology volutions are separated by a thick ridge and the spirals cells
provided in Sharma and Patnaik (2014). A total of 118 cha- are concave. The apical pole is rounded and the spirals cells
rophyte gyrogonite specimens were recovered in the present are slightly enlarged at the apex joint in a zigzagged line.
investigation. Out of these, 47 charophyte gyrogonite speci- The basal part is truncated with a rounded to pentagonal
mens were measured taking into account the morphometric basal pore at the centre surrounded by the tip of the spiral
parameters such as gyrogonite height (in μm), gyrogonite cell.
width (in μm), number of spiral turns observed in lateral
view and the isopolarity index (gyrogonite height/gyrogonite Remarks
width × 100). Charophyte gyrogonites were photographed
using a scanning electronic microscope (SEM) housed at The recovered gyrogonites differ from Chara globularis
the Central Instrumentation Laboratory (CIL) of Central var. globularis (Thuillier 1799) in having a sharp convex
University of Punjab, Bathinda, Punjab, India. Representa- spiral cell. In addition, having more convolutions cells
tive selection of the samples were photographed using Leica and less developed apical rosette nodes differs from Chara
M205C housed at the Department of Geology, Central Uni- rantzieni (Tewari and Sharma 1972). C. hispida (Soulié-
versity of Punjab, Bathinda, Punjab, India. The specimens Märsche and Garćia 2015) possesses enlarged spiral cells
are housed at the Central University of Punjab, Department at the apical junction with strongly protruding intercellular
of Geology (BIOPS/CUP/KT-Biostratigraphy, Palaeontol- sutures unlike the present recovered gyrogonites. Chara cf.
ogy and Sedimentology Laboratory/Central University of C. hispida reported by Bhatia (1999) from the Tatrot and
Punjab/Kutch Tapar) and Department of Geology, Panjab Pinjor formations of the Siwalik differs from the present
University, Chandigarh, India (acronym VPL/PU/KT-Ver- specimens (VPL/PU/KT 570–571, BIOPS/CUP/KT 1005,
tebrate Palaeontology Laboratory/Panjab University/Kutch). 1009, 1012, 1022) in having a conical shape that narrows
towards the base. The specimens are very similar to Chara
globularis var. aspera described from the Siwalik Group
Systematic palaeontology
(Bhatia 1999) in terms of being equisized, elongate and pro-
late shape, sharp and convex spiral cells, a rounded apex
Division Charophyta Migula 1890
and a pentagonal basal pore. Bhatia and Singh (1989) also
Class Charophyceae Smith 1938
reported this taxon from the Quaternary marls of the Indo-
Order Charales Lindley 1836
Gangetic plain.
Family Characeae Richard 1815 ex Agardh, 1824
Chara globularis var. globularis Thuillier 1799
Genus Chara Vaillant 1719
(Figs. 2 d–i).
Chara globularis var. aspera (Detharding ex Willdenow)
Referred materials
Wood 1962
(Fig. 2a, b, c).
VPL/PU/KT 572–577, BIOPS/CUP/KT 1006, 1016,
1033–1035, 1048–1052.
Referred materials
Locality and horizon
VPL/PU/KT 570–571, BIOPS/CUP/KT 1005, 1009, 1012,
1022.
Late Miocene (~ 11–10 Ma) mudclast conglomerate bed at
Tapar section, Kutch Basin, Gujarat State, western India.
Locality and horizon
Description
Late Miocene (~ 11–10 Ma) mudclast conglomerate bed at
Tapar section, Kutch Basin, Gujarat State, western India. The gyrogonites are medium sized (562–909 μm high and
382–619 μm wide), elongate, and prolate in shape, with an
Description isopolarity index range from 124–159 μm (see Table 2).
Ten to Twelve convolutions are seen in lateral view and the
The Gyrogonites are medium sized, 570–721  μm high, width of the lime spirals (as measured between the inter-
412–485  μm wide, elongate, prolate in shape, with an cellular sutures) at the equatorial axis of the gyrogonites
ranges from 47–75 μm. Spiral cells are generally thick,

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Fig. 2  a–c (VPL/PU/KT 570), Chara globularis var. aspera: a Lateral view; b Apical view; c Basal view. d–f (VPL/PU/KT 572); g–i (BIOPS/
CUP/KT 1006), Chara globularis var. globularis: d and g, Lateral view; e and h, Apical view; f and i, Basal view. (Scale 100 µm)

convex and sinistrally coiled. The apex is psilocharoid and Remarks

prominent. The basal pore is pentagonal in shape.
The recovered gyrogonites differ from Chara globularis
var. aspera in having a rounded convex spiral cell. Bhatia

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Table 2  Biometric values Specimens Gyrogonite Gyrogonite Isopolarity index Number of Width of the
i.e. gyrogonite height height (μm) width (μm) (h/w × 100) (μm) convolutions convolution
(μm); gyrogonite width (μm)
(μm); isopolarity index
(ISI = h/w × 100), number of Chara globularis var. aspera
convolutions visible in lateral
VPL/PU/KT 570 721 485 148 11 78
view of charophtes from Tapar
locality Kutch, Gujarat, India VPL/PU/KT 571 624 412 151 11 54
BIOPS/CUP/KT 1005 599 414 145 12 41
BIOPS/CUP/KT 1009 595 456 131 9 68
BIOPS/CUP/KT 1012 570 432 132 ? 65
BIOPS/CUP/KT 1022 580 416 139 10 49
Chara globularis var. globularis
VPL/PU/KT 572 620 466 133 12 53
VPL/PU/KT 573 685 521 131 ? 71
VPL/PU/KT 574 909 572 159 11 72
VPL/PU/KT 575 864 619 139 12 70
VPL/PU/KT 576 671 521 128 ? 71
VPL/PU/KT 577 750 575 130 ? 75
BIOPS/CUP/KT 1006 587 392 150 10 47
BIOPS/CUP/KT 1016 598 382 157 ? 48
BIOPS/CUP/KT 1033 620 492 126 ? 70
BIOPS/CUP/KT 1034 635 461 138 ? 64
BIOPS/CUP/KT 1035 562 449 125 ? 58
BIOPS/CUP/KT 1048 598 468 128 ? 56
BIOPS/CUP/KT 1049 570 413 138 ? 59
BIOPS/CUP/KT 1050 567 431 131 ? 58
BIOPS/CUP/KT 1051 697 485 144 ? 47
BIOPS/CUP/KT 1052 597 484 124 ? 48
Chara sp.
VPL/PU/KT 578 784 569 138 ? ?
VPL/PU/KT 579 727 527 137 ? ?
VPL/PU/KT 580 714 485 147 ? ?
VPL/PU/KT 581 757 542 140 10 65
BIOPS/CUP/KT 1017 657 472 139 ? ?
BIOPS/CUP/KT 1027 582 438 133 ? 37
BIOPS/CUP/KT 1028 627 419 150 ? ?
BIOPS/CUP/KT 1032 589 409 144 ? 51
BIOPS/CUP/KT 1047 436 309 141 ? 47
Lychnothamnus sp.
BIOPS/CUP/KT 1001 794 655 121 9 92
BIOPS/CUP/KT 1080 553 ? ? ? 60
Lychnothamnus cf. sahnii
BIOPS/CUP/KT 1002 721 523 138 8 88
BIOPS/CUP/KT 1073 813 689 118 ? 93
BIOPS/CUP/KT 1074 735 603 122 ? 97
Nitellopsis sp.
VPL/PU/KT 582 707 669 105 ? 84
VPL/PU/KT 583 600 515 116 9 72
VPL/PU/KT 584 620 500 124 8 73
VPL/PU/KT 585 792 672 117 7 120
VPL/PU/KT 586 865 818 105 7 120
VPL/PU/KT 587 685 592 115 7 107

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Proceedings of the Indian National Science Academy

(1999) described this species from the Siwalik Group and it preserved, the precise taxonomic allocation at the species
is characterised by a sharp convex spiral cell. Chara rantz- level is difficult to ascertain.
ieni reported from the Tatrot Formation (Pliocene) of the Lychnothamnus cf. sahnii Bhatia 1999.
Siwalik (Bhatia and Mathur 1978; Bhatia 1999) and the (Fig. 4a–c).
Middle Siwalik of Mohand, Dehradun (Tiwari and Bhan
2021) differs from the present specimens in having fewer Referred materials
convolutions and moderately developed apical rosette nodes.
Chara aff. microcera described from the Miocene of Zahle, BIOPS/CUP/KT 1002, 1073, 1074.
western margin of the Bekaa Valley, Lebanon (Sanjuan and
Alqudah 2018) differs from the present specimens in having Locality and horizon
a characteristic ornamentation consisting of isolated small
tubercles irregularly arranged along the spiral cells. The pre- Late Miocene (~ 11–10 Ma) mudclast conglomerate bed of
sent gyrogonites are very similar to Chara globularis var. Tapar section, Kutch basin, Gujarat, India.
globularis described from the Siwalik (Bhatia 1999) in terms
of morphological characters such as prolate in shape, hav- Description
ing a similar number of convolutions, having convex spiral
cells, the absence of small tubercles along the spiral cells, a The specimens are medium to large sized (721-813 μm high
psilocharoid apex and a pentagonal basal pore. and 523–689 μm wide), prolate to ellipsoidal in shape with
Chara sp. 8 convolutions in lateral view and an isopolarity index range
(Fig. 3a–i). from 122–138 μm (see Table 2). The spiral cells are smooth,
wide and concave with a faintly developed double suture
Referred materials (intercellular suture). The apical pole is truncated with
slightly protruding sutures and the tips of the lime spiral are
BIOPS/CUP/KT 1017, 1027, 1028, 1032, 1047 and VPL/ slightly enlarged. The basal funnel is well developed and the
PU/KT 578–581. basal pore is pentagonal in outline.

Locality and horizon Remarks

Late Miocene (~ 11–10 Ma) mudclast conglomerate bed at The three recovered gyrogonites differ from Lychnothamnus
Tapar section, Kutch Basin, Gujarat State, western India. breviotus by possessing a concave spiral cell and a well-
developed drown-out basal funnel, and they are also smaller
Description than Lychnothamnus barbatus. Bhatia (1999) described
Lychnothamnus sahnii from the Chinji Formation of the
The Gyrogonites are medium to large sized with height Lower Siwalik, India which is quite similar to the recov-
range of 436–784 μm and width range from 309–569 μm ered gyrogonites on account of sharing the morphological
wide; and specimens are prolate to ellipsoidal in shape features such as a concave spiral cell, double sutures and a
with an isopolarity index ranging from 133–150 μm (see well-developed basal funnel with pentagonal pore. Earlier,
Table 2). Spiral cells are not clearly observed due to poor Bhatia and Mathur (1978) originally referred this species
preservation. The width of the lime spirals observed in few to the genus Tectochara, and then Bhatia (1999) assigned
gyrogonites ranges from 37–65 μm. In apical view, a few it to the genus Lychnothamnus after re-examination of the
gyrogonites moderately develop an apical rosette node and topotype material from the Chinji Formation.
others have a highly eroded surface. Lychnothamnus (Ruprecht) Braun 1856.
Lychnothamnus sp.
Remarks (Fig. 4d–f).

The recovered gyrogonites are similar in size to those Referred materials

Chara sp. reported from the Upper Siwaliks (Bhatia and
Mathur 1978; Bhatia 1999) and the Middle Siwalik from the BIOPS/CUP/KT 1001, 1080.
Mohand area, Dehradun Sub-basin, North-West Himalaya
(Tiwari and Bhan 2021). Since the specimens are poorly

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Fig. 3  a–c (BIOPS/CUP/KT 1017); d–f (BIOPS/CUP/KT 1027); g–i (VPL/PU/KT 578), Chara sp.: a, d and g are in Lateral view; b, e and h are
Apical view; c, f and i are in Basal view (Scale 100 µm)

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Fig. 4  a–c (BIOPS/CUP/KT 1002), Lychnothamnus sahnii: a Lat- (VPL/PU/KT 582), Nitellopsis sp.: g Lateral view; h Apical view; i
eral view; b Apical view; c Basal view; d–e (BIOPS/CUP/KT 1001), Basal view (Scale 100 µm)
Lychnothamnus sp.: d Lateral view; e Apical view; f Basal view; g–i

Locality and horizon Description

Late Miocene (~ 11-10 Ma) mudclast conglomerate bed at The specimens are medium to large sized (553–794 μm
Tapar section, Kutch Basin, Gujarat State, western India. high and 655 μm wide; see Table 2), slightly elongated and
oval to spheroidal-rounded in shape. Nine convolutions
are visible in lateral view and the width of the spiral lime
ranges from 60–92 μm. The spiral cells are smooth, wide

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and slightly concave to flat with a faintly developed double and less convolutions (see Bhatia and Mathur 1978; Bhatia
suture (intercellular suture). In apical view, the pole is flat 1999; Tiwari and Bhan 2021). However, the main important
with the absence of a apical node. The spiral cells at the api- feature preserved in the basal pore to distinguish the species
cal junction are constant in width and thickness. The basal within the genus is not clearly seen, and allocation to a more
funnel is poorly developed with small pores. precise taxonomic level is not possible at the moment.

Remarks Faunal composition

The two gyrogonites (BIOPS/CUP/KT 1001, 1080) are mor- The charophyte bearing Tapar Section of Kutch is well
phologically similar to those described as Lychnothamnus known for its diverse fossil assemblages comprising of
barbatus from the sediments of the Ganga plain (Bhatia rich mammalian fauna including hominoids (Sivapithecus),
2006) and the Middle Siwaliks sediment at Mohand, Deh- suids, Sanitherium, tragulids, giraffids, pecorans, bovids,
radun (Tiwari and Bhan 2021) in having a smooth flat spiral Hipparion, deinotheres, rhinocerotids, rodents, chelonians,
cell, and a double suture and constant width of the spiral crocodiles, insectivores, lizards, teleosts, skates and rays,
cell at the apical junction. However, the recovered gyrogo- etc. (also see Bhandari et al. 2015, 2018, 2021; Singh et al.
nites differ from L. barbatus in having smaller size, but falls 2019, 2020; Sharma et al. 2021; Čerňanský et al. 2021)
within the range of L. breviotus. L. breviotus is distinguished (also see Fig. 5A, B). Thus, taxonomically identifiable fau-
from L. barbatus on account of its smaller size and ovoidal nal remains of the Tapar section represent mammals 55%
to nearly spheroidal shape (Tiwari and Bhan 2021). Moreo- of the total families, 66% of the genera and 80% of the total
ver, the recovered specimens are difficult to assign the spe- species; reptiles 16% of the families and 12% of the genera;
cies level due to the incomplete preservation. amphibians 6% of the families, 6% of genera and 3% of the
Genus Nitellopsis Hy 1889 species; pisces 23% of the families, 16% of the genera and
Nitellopsis sp. 17% of species (also see Fig. 5, Table 3).
(Fig. 4g–i).

Referred materials Discussion

VPL/PU/KT 582–587.
Palaeoenvironments
Locality and horizon
Based on the presence of fossil batoids including Dasyatis
Late Miocene (~ 11–10 Ma) mudclast conglomerate bed at rugosa, D. aff. probsti, Pastinachus sp., and Pristis sp., etc.
Tapar section, Kutch Basin, Gujarat State, western India. the deposition of the Tapar section is considered to have
taken place in a near-coastal environment where freshwater
Description fauna were mixed (also see, Sharma et al. 2021). The lithol-
ogy of this section comprised massive sand, cross-laminated
The Gyrogonites are medium in size (600–865 μm high and sandstone, mudclast conglomerate, silt and mudstone with
500–818 μm wide) and spheroidal to ovoidal or ellipsoidal intermittent conglomerate beds composed of calcareous nod-
in shape with an isopolarity index ranging from 105–124 μm ules, agate pebbles, and very coarse sand and mudclasts with
(Table 2). Seven to nine convolutions are visible in lateral an occasional coarsening upward sequence, suggesting a flu-
view. The lime spiral cells are flat to convex and sinistrally vial deposit (Bhandari et al. 2015). The recent identifications
coiled with the width of the lime spirals (as measured of mammalian, reptilian, and piscian fauna (Bhandari et al.
between the intercellular sutures) at the equatorial axis of 2015; Singh et al. 2019, 2020; Sharma et al. 2021) indicate
the gyrogonites ranging from 73–120 μm. The apical pole the existence of a fluvial depositional setup with estuaries
is rounded and slightly truncated in apical view. The lime under tropical to subtropical humid conditions, very close
spirals are thick at the apical periphery and form an apical to the coastal area.
rosette. The charophytes flora of the Tapar section are represented
by Chara globularis var. aspera, C. globularis var. globu-
Remarks laris, Lychnothamnus cf. sahnii, Lychnothamnus sp., and
Nitellopsis (Tectochara) sp. Among taxa, Chara globula-
The recovered specimens are assigned to the genus Nitellop- ris var. aspera and C. globularis var. globularis are most
sis on account of medium size, spheroidal to ovoidal shape often found in lacustrine environments and contradict the
palaeoenvironmental data from other associated fauna.

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Fig. 5  A Total number of fauna reported from the Tapar; B Percentage wise Pie chart of fossils reported from the Tapar site (source: Bhandari
et al. 2009, 2015, 2018, 2021; Singh et al. 2019, 2020; Sharma et al. 2021; Čerňanskýet al. 2021)

However, the lithostratigraphy of the locality includes an found in coal strip mine pools to small and large lakes
occasional coarsening upward sequence suggesting a cre- (Daily 1958). Lychnothamnus is confined to freshwater and
vasse splay deposit, with parallel to cross laminated sand- depths ranging from 2–10 m (Bhatia 2006). Deeper condi-
stone, siltstone and mudstone with intermittent conglomer- tions influence the production of large-size gyrogonites
ate beds comprised of calcareous nodules, agate pebbles, and enable the plants to remain submerged for a long time
and very coarse sand and mudclasts (also see, Bhandari providing longer periods of reproduction (Soulié-Märsche
et al. 2015). Moreover, the presence of mammals, reptiles, and Martin-Closas 2003). Here, the ecology of Nitellop-
batoids, and teleost at the Tapar locality indicates a fluvial, sis is understood based on its single living representative
near coastal environment where freshwater fauna were Nitellopsis obtusa. The modern N. obtusa favours deep
mixed. This environment is further supported by the faunal and shallow lakes, abandoned gravel pits, rivers, oxbows,
association of gastropods, bivalves, crocodiles and turtles and secondary channels at water depths of 0.5 to > 14 m
(Fig. 5A and B). (Korsch et al. 2008; Janauer et al. 2010). It preferentially
Charophytes are found to occur in deep and cold fresh- colonizes calcareous, neutral to alkaline, mesotrophic
water lakes, shallow freshwater lakes or marginal zones to eutrophic waters (Bailly et al. 2007; Hutorowicz and
of deep lakes-tuffaceous deposits originating in springs, Dziedzic 2008). Nitellopsis obtusa has also been found in
temporary ponds with either fresh or brackish water, saline brackish waters (Langangen et al. 2002) but collapses at
inland waters and also tropical lakes, depending on the salinities higher than 5‰ (Katsuhara and Tazawa 1986).
species (Soulié-Märsche 1993). The climate, hardness and In Tapar section, charophytes are limited to the mud clast
salinity of the water are the most obvious factors affecting conglomerate bearing beds. Taking into consideration the
their distribution (Corillion 1975). Chara aspera favours very poor nature of preservation of the gyrogonites and
the alkaline waters of lakes and ponds, and brackish water their scarcity among the faunal yield in the conglomeratic
downward up to 5.5 m in depth, but mainly at depths of 0 beds, it is suggested that the charophytes might have been
to 4 m (Olsen 1944). Chara globularis has been found in reworked, transported from a nearby lacustrine site. Thus,
more varied habitats throughout the world, more so than the palaeoenvironmental data from the charophyte fauna
any other charophyte (Crum 1975). This species prefers may correspond to the provenance of the mud clasts in the
large and small lakes, ponds, clay, pits, and ditches (Olsen conglomeratic bed.
1944). In India, it has an equally wide range of tolerance,

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Table 3  Faunal compositions of the hominoid bearing Tapar locality of Kutch, Gujarat India
Order Family Genus Species References

Pisces Cypriniformes Cyprinidae Indet Indet Singh et al. (2019)

Siluriformes Bagridae Indet Indet
Perciformes Indet Indet Indet
Carcharhiniformes Carcharhinidae Carcharhinus Indet Bhandari et al. (2015)
Myliobatiformes Dasyatidae Dasyatis Dasyatis rugosa Sharma et al. (2021)
Dasyatis probsti
Dasyatis sp.
Hemantura Hemantura menoni
Pastinachus Pastinachus sp.
Pristiformes Pristidae Pristis Pristis sp. Bhandari et al. (2015), Sharma
et al. (2021)
Amphibia Anura Ranidae Rana Rana sp. Singh et al. (2021a, 2021b)
Reptilia Squamata Pythonidae Python Python sp.
Colubridae Indet Indet
(“Colubrinae” + Ahaetulii- Indet Indet
nae)
Acrochordidae Acrochrodus Acrochordus dehmi
Agamidae Uromastyx cf. Uromastyx s.l. sp. Čerňanský et al. (2021)
Scincidae Indet Indet
Varanidae Varanus Varanus sp.
Crocodilia Crocodylidae Indet Indet Bhandari et al. (2009, 2015)
Mammal Rodentia Sciuridae Tamias Tamias urialis Bhandari et al. (2021)
Tamias gilaharee sp. nov Patnaik et al. (2022)
Tamiops Tamiops sp. Bhandari et al. (2021)
Ctenodactylidae Sayimys Sayimys sivalensis Bhandari et al. (2021), Patnaik
Spalacidae Prokanisamys Prokanisamys sp. et al. (2022)
Kanisamys Kanisamys indicus Bhandari et al. (2021)
Kanisamys kutchensis sp. Patnaik et al. (2022)
nov
Cricetidae Democricetodon Democricetodon fejfari Bhandari et al. (2021), Patnaik
Muridae Dakkamys Dakkamys asiaticus et al. (2022)
Progonomys Progonomys morganae
Progonomys prasadi sp. nov Patnaik et al. (2022)
Myocricetodon Myocricetodon gujaratensis
sp. nov
Primates Hominidae Sivapithecus Sivapithecus sp. Bhandari et al. (2018)
Carnivora Amphicyonidae Indet Indet Bhandari et al. (2009)
Artiodactyla Bovidae Protrogocerus Protrogocerus sp. Bhandari et al. (2015)
Gazella Gazella sp.
Suidae Kachchhchoerus Kachchhchoerus salinus
Tetraconodon Tetraconodon indicus
Listriodon Listriodon dukkar sp. nov van der Made et al. (2022)
Anthracotheriidae Sivameryx Sivameryx palaeindicus Bhandari et al., (2009)
Sanitheriidae Sanitherium Sanitherium schlagintweiti Bhandari et al. (2015)

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Table 3  (continued)
Order Family Genus Species References

Tragulidae Dorcatherium Dorcatherium minus Bhandari et al. (2009)

Dorcatherium sp.
Giraffidae Giraffokeryx Giraffokeryx punjabiensis
Giraffa Giraffa priscilla Bhandari et al. (2015)
Perissodactyla Equidae Hipparion Hipparion sp. Bhandari et al. (2009)
Rhinocerotidae Brachypotherium Brachypotherium perimense Bhandari et al. (2015)
Brachypotherium sp. Bhandari et al. (2009)
Proboscidea Gomphotheriidae Gomphotherium Gomphotherium sp.
Deinotheriidae Deinotherium Deinotherium sindiense Bhandari et al. (2009)
Deinotherium indicum Singh et al. (2020)

Age and biostratigraphy of tapar section the age range to 10.8–10 Ma for Tapar and Pasuda localities
based on the biostratigraphic ranges of Siwalik rodents. A new
The Miocene deposits of Kutch have been studied by vari- Listriodon species from Pasuda (Made et al. 2022) suggests
ous workers, but the precise geological age of these depos- an approximate age of ~ 10 Ma and suggests a simultaneous
its has remained debatable for a long period of time due to extinction of the frugivore suid across Africa, Europe and Asia
the presence of long ranging taxa and the lack of prominent during this time (also see, Fig. 6). This age assessment sup-
index fossils. Lydekker (1876) correlated the Samda (Sam- ports the correlation of the Tapar beds with the younger Sand-
bera) deposits of Kutch, Gujarat to the early Miocene, nearly han Formation, rather than the older Khari Nadi Formation.
equivalent to the Burdigalian of Europe and the Lower Siwa- Among the charophytes from Tapar locality, the present
liks of Pakistan based on the recovered deinothere teeth from species Chara globularis var. aspera is known from the Plio-
the locality. Based on the marine fossil record, (Biswas 1992) cene (Tatrot and Pinjor Formation) and Holocene (Quater-
suggested that the age of the Khari Nadi Formation is Late nary) sediments in the Siwalik, India (Bhatia and Singh 1989;
Aquitanian and the Chhasra Formation is Burdigalian, but Bhatia 1999). According to Bhatia (1999), C. globularis var.
due to the absence of index fossils in the Sandhan Formation, aspera make its first appearance in the late Pliocene Tatrot
no specific age was assigned to this formation. However, the Formation (5–2.5 Ma). However, this species is also reported
Sandhan Formation overlies Miocene strata with a prominent by Sharma et al. (2015) from the Dhok Pathan Formation
break, and a tentative Pliocene age has been assigned to this (Early Pliocene) near Polian Prohita, Himachal Pradesh
formation (Biswas 1992). Subsequently, the recovery of mam- (India). The presence of the species at the late Miocene
malian fossils from Pasuda (Bhandari et al. 2009) supported (~ 10 Ma) Tapar locality suggests that the species chronologi-
the earlier age proposed by Biswas (1992), and an Aquitanian cal range extend into the late Miocene (Fig. 6). C. globularis
age was accepted for the Khari Nadi Formation. However, the var. globularis is frequent to abundant in the Dhok Pathan,
collection of various mammalian fossils, particularly Hip- Tatrot and Pinjor formations of the Siwalik Group (Bhatia
parion, a hominoid (Sivapithecus), rodents and suids from 1999) but now the geological age of this species extends to
the two localities Tapar and Pasuda (4.5 km south east of the the late Miocene (Fig. 6). The presence of another species
Tapar location) of Kutch indicated a late Miocene age close Lychnothamnus sahnii at the study area suggests extending
to the Chinji-Nagri Boundary (Bhandari et al. 2015, 2018) the upper age limit of this species to late Miocene ~ 10 Ma
(Fig. 6). Most recently, the presence of Deinotherium indicum (see, Fig. 6). Previously, Bhatia (1999) described this species
at the Tapar locality provides a more precise age estimate of from the Lower Siwalik Chinji Formation.
10.1–8.6 Ma based on the correlation with the Haritalyangar
section (Singh et al. 2020) (also see, Fig. 6). This proboscid is
similar to that found at Haritalyangar and other similarly aged Conclusions
deposits from the Dhok Pathan Formation on the Potwar Pla-
teau in Pakistan (Barry et al. 1982; Pillans et al. 2005), further 1. The maceration of bulk sediments from the fossiliferous
indicating a late Miocene (Tortonian) age for Tapar. Kapur site, Tapar locality of Kutch, Gujarat, India has yielded
et al. (2019) also suggested the Sandhan Formation is older 135 ill preserved gyrogonite specimens. Out of these,
than previously reported. Bhandari et al. (2021) further revised only 47 specimens are assignable to the seven Siwa-

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Fig. 6  Biostratigraphic range of Siwalik Charophyte species (modi- ity (also see, Bhatia 1999, Bhandari et  al., 2015, 2018, 2021; Singh
fied after, Bhatia and Mathur 1978; Bhatia 1999; Tiwari and Bhan et al. 2020). The age of Siwalik group is synthesised after Barry et al.
2021) and Siwalik rodents and mammals occurring at Tapar local- (2013) and Patnaik (2013)

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Proceedings of the Indian National Science Academy

lik species including Chara globularis var. aspera, C. (Jura et Doubs) et lac des Rousses (Jura). Conservatoire Botanique
globularis var. globularis, Lychnothamnus cf. sahnii, National de Franche-Comté, Besançon, France (2007)
Bajpai, S., Domning, D.P.: A new dugongine sirenian from the early
Lychnothamnus sp. and Nitellopsis sp. Miocene of India. J. Vertebr. Paleontol. 17, 219–228 (1997).
2. The gyrogonites are poorly preserved and limited to the https://​doi.​org/​10.​1080/​02724​634.​1997.​10010​965
mud clast conglomerate bearing beds. The very poor Bajpai, S., Thewissen, J.G.M., Kapur, V.V., Tiwari, B.N., Sahni, A.:
nature of preservation and their scarcity and limitation to Eocene and Oligocene sirenians (Mammalia) from Kachchh.
India. J. Vertebr. Paleontol. 26, 400–410 (2006). https://​doi.​org/​
the mudclast bearing conglomeratic beds suggests that 10.​1671/​0272-​4634(2006)​26[400:​EAOSMF]​2.0.​CO;2
the gyrogonites might have been transported along with Barry, J.C., Lindsay, E.H., Jacobs, L.L.: A biostratigraphic zonation of
the mudstone clast from a nearby lacustrine site. Thus, the Middle and Upper Siwaliks of the Potwar Plateau of northern
the palaeoenvironmental data from the charophyte fauna Pakistan. Palaeogeogr. Palaeoclimatol. Palaeoecol. 37, 95–130
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may correspond to the provenance of the conglomeratic Beilby, M., Schneider, S.C., Puckacz, A., Martín-Closas, C.: Towards
bed. an integrated understanding of charophyte biology and paleobiol-
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under tropical to subtropical humid conditions. The Early Miocene mammals from central Kutch (Gujarat), Western
presence of the reworked charophytes most dominantly India: implications for geochronology, biogeography, eustacy and
found in a lacustrine condition also suggests the lacus- intercontinental dispersals. Neues Jahrb Geol Paläontol Abh 256,
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4. The records of gyrogonites from well-dated Tapar local- fauna from the Tapar and Pasuda, Kutch. Münchner Geowissen-
ity of Kutch (~ 10 Ma) extend the first appearance datum schaftliche Abhandlungen, Reihe a, Geologie Und Paläontologie
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ris var. globularis and last appearance datum (LAD) of Bhandari, A., Kay, R.F., Williams, B.A., Tiwari, B.N., Bajpai, S., Hier-
onymus, T.: First record of the Miocene hominoid Sivapithecus
Lychnothamnus sahnii. from Kutch, Gujarat state, western India. PLoS One 13(11),
e0206314 (2018). https://​doi.​org/​10.​1371/​journ​al.​pone.​02063​14
Bhandari, A., Bajpai, S., Flynn, L.J., Tiwari, B.N., Mandal, N.: First
Acknowledgements  The authors are also thankful to Prof. M.G. Miocene rodents from Kutch, western India. Hist. Biol. 33(12),
Thakkar K.S.K.V. Kachchh University, India for his help during the 3471–3479 (2021). https://d​ oi.o​ rg/1​ 0.1​ 080/0​ 89129​ 63.2​ 020.1​ 8709​
field work. Encouragement and suggestions from Prof. Ashok Sahni 70
(Panjab University, Chandigarh, India) are highly acknowledged. Bhatia, S.B.: Revision of the charophyte flora of the Siwalik Group
(Neogene-Quaternary) of the Lesser Himalaya. India. Aust. J. Bot.
Funding  KM Sharma is thankful to the Central University of Pun- 47, 459–474 (1999). https://​doi.​org/​10.​1071/​BT970​97
jab Bathinda, India for the Research Seed Money Grant (Ref. no: Bhatia, S.B.: Charophytes from the Miocene-Pliocene transition in the
CUPB/CC/16/00/13), UGC (No.F.30-4/2014 (BSR) and SERB Siwalik/Churia Group of the Himalayan Foreland Basin: Paleo-
(ECR/2016/001100) for the financial supports. R. Patnaik is thankful biogeographic and Paleoecologic implications. Acta Micropalae-
to SERB (HRR/2018/000063) for the financial support. N. Amardas ontol Sin 20, 150–155 (2003). https://​doi.​org/​10.​1111/​iar.​12408
is thankful to UGC for providing JRF. NPS and RKS thank the Direc- Bhatia, S.B.: Ecological parameters and dispersal routes of Lych-
tor, Wadia Instiute of Himalayan geology, Dehradun for the research nothamnus barbatus (Characeae) in the Early-Middle Holocene
facilities. from the Ganga plain. India. Cryptogam. Algol. 27(4), 341–356
(2006)
Bhatia, S.B., Mathur, A.K.: First record of charophyta from the Upper
Declarations  Siwaliks near Pinjor. Bull. Indian Geol. Assoc. 3, 27–28 (1970)
Bhatia, S.B., Mathur, A.K.: The Neogene charophyte flora of the
Conflict of interest  No potential conflict of interest was reported by Siwalik Group, India and its biostratigraphic significance. Geo-
the author(s). phytology 8, 79–97 (1978)
Bhatia, S. B., Singh, N.: Holocene charophytic flora of parts of Uttar
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