The oldest birotule-bearing freshwater sponges (Porifera: Palaeospongillidae) from the Upper Cretaceous-lower Palaeocene Deccan volcanic-associated sediments of India

Abstract

A new fossil occurrence of freshwater sponges (Porifera: Demospongiae) is reported from the Deccan volcanic associated
Naskal intertrappean locality, deposited in an interval of

Full text

Acta Palaeontol. Pol. 68 (1): 167–174, 2023 https://doi.org/10.4202/app.01040.2022
The oldest birotule-bearing freshwater sponges
from the Upper Cretaceous–lower Paleocene Deccan
volcanic-associated sediments of India
BANDANA SAMANT, ROBERTO PRONZATO, DHANANJAY MAHENDRAKUMAR MOHABEY,
TIZIANA CUBEDDU, GIACINTA ANGELA STOCCHINO, KRUTIKA JANGALE,
PRANAY THALAL, ANUP DHOBALE, and RENATA MANCONI
Samant, B., Pronzato, R., Mohabey, D.M., Cubeddu, T., Stocchino, G.A., Jangale, K., Thalal, P., Dhobale, A., and
Manconi, R. 2023. The oldest birotule-bearing freshwater sponges from the Upper Cretaceous–lower Paleocene Deccan
volcanic-associated sediments of India. Acta Palaeontologica Polonica 68 (1): 167–174.
A new fossil occurrence of freshwater sponges (Porifera: Demospongiae) is reported from the Deccan volcanic associ-
ated Naskal intertrappean locality, deposited in an interval of <100 kyr across the Cretaceous/Paleogene (K/Pg) bound-
ary. This is the oldest record of siliceous fossil birotule spicules (gemmuloscleres) belonging to asexual resting stages
typical of the order Spongillida. The analysis supports the ascription of these fossils to the family Palaeospongillidae.
The diagnosis and description of Longibirotula Pronzato and Manconi gen. nov. and its type species Longibirotula
antiqua Manconi and Samant sp. nov. from the Naskal intertrappean is based on skeletal and gemmular spicular mor-
photraits. The findings have provided evidence of the presence of diversified groups of freshwater sponges during the
Late Cretaceous on the Indian subcontinent and Gondwanaland. From the biogeographic context, the findings track the
evolutionary trends of the oldest continental sponges in the Asian and Australasian/Insular Pacific regions.
Key words: Porifera, Palaeospongillidae, inland water sponges, conservative morphotraits, gemmules, siliceous
skeleton, palaeobiodiversity, palaeogeography, Cretaceous, Paleocene, Deccan traps.
Bandana Samant [bandanabhu@gmail.com, ORCID: https://orcid.org/0000-0001-8257-452 ], Dhananjay Mahen-
drakumar Mohabey [dinomohabey@yahoo.com; ORCID: https://orcid.org.0000-0003-4538-6907 ], Krutika Jangale
[krutismile1999@gmail.com; ORCID: https://orcid.org/0000-0002-1452-3545 ],
Pranay Thalal [ppt52525.pt.pt@gmail.com; ORCID: https://orcid.org/0000-0003-3768-1519 ] and Anup Dhobale
[anupdhobale@gmail.com; ORCID: https://orcid.org/0000-0001-9496-0069 ], Postgraduate Department of Geology,
RTM Nagpur University, Nagpur, Maharashtra 440001, India.
Roberto Pronzato [roberto.pronzato@unige.it; ORCID: https://orcid.org/0000-0003-1355-0936 ], Dipartimento di Sci-
enze della Terra, dell’Ambiente e della Vita (DISTAV), Università di Genova, Genova, Italy.
Renata Manconi [rmanconi@uniss.it; ORCID: https://orcid.org/0000-0002-7619-8493 ], Tiziana Cubeddu [tcubeddu
@uniss.it; ORCID: https://orcid.org/0000-0001-7522-8200 ], and Giacinta Angela Stocchino [stocchin@uniss.it; OR-
CID: https://orcid.org/0000-0002-7005-208X ], Dipartimento di Medicina Veterinaria, Laboratorio Zoologia, Univer-
sità di Sassari, Sassari, Italy.
Received 30 October 2022, accepted 25 January 2023, available online 8 March 2023.
Copyright © 2023 B. Samant et al. This is an open-access article distributed under the terms of the Creative Commons
Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original author and source are credited.
Introduction
The fossil freshwater sponges (Porifera: Demospongiae),
known from both Laurasia and Gondwana, belong to the
order Spongillida Manconi and Pronzato, 2002, which com-
prises three families with fossil taxa, i.e., Palaeospongillidae
Volkmer-Ribeiro and Reitner, 1991, Potamolepidae Brien,
1967, and Spongillidae Gray, 1867. The oldest Spongillida
fossil records (only spicules) dates back to the Permo-
Carboniferous (Schindler et al. 2008), and the Late Jurassic
(Spongilla purbeckensis Hinde, 1883). Unfortunately, fossil
records of inland water sponges are very scarce, due to their
fragile skeletal siliceous architecture (Pronzato et al. 2017).
Most fossil Spongillida are hard to identify to the family,
genus, or species level because they lack gemmuloscleres,
which are one of the most important diagnostic morpho-

168 ACTA PALAEONTOLOGICA POLONICA 68 (1), 2023
traits for systematics and phylogeny. Till date, the oldest
known entire gemmule is of Palaeospongilla chubutensis
Volkmer-Ribeiro and Reitner, 1991 (Palaeospongillidae)
from the Lower Cretaceous of Patagonia (Volkmer-Ribeiro
and Reitner 1991).
The gemmuloscleres remains of other fossil species of
the families Spongillidae and Potamolepidae share simi-
lar morphotraits with living genera, e.g., Anheteromeyenia
Schröder, 1927; Corvospongilla Annandale, 1911; Ephydatia
Lamouroux, 1816; Eunapius Gray, 1867; Radiospongilla
Penney and Racek, 1968; Spongilla Linnaeus, 1759; Troch o -
spongilla Vejdovsky, 1883; Oncosclera Volkmer-Ribeiro,
1970, and Potamophloios Brien, 1970 (Pisera 2006; Pisera
et al. 2013, 2016; Pronzato et al. 2017).
From the Deccan volcanic province, some unidenti-
fied spicules of freshwater sponges have been recorded
from the Deccan infra (Lameta Formation) and intertrap-
pean sediments of central India and the intertrappean beds
of northwestern India (Samant and Mohabey 2009, 2014;
Samant et al. 2014), but definite record of fossil freshwater
sponges of Spongillida is scarce. Recently, the new genus
Palaeocorvospongilla Pronzato and Manconi, 2021, of the
family Palaeospongillida was described from the Deccan
intertrappean sediments of India (Maastrichtian, Upper
Cretaceous) with the new species Palaeocorvospongilla cre-
tacea Manconi and Samant, 2021 (Samant et al. 2021).
The present study records the oldest fossil birotule-bear-
ing freshwater sponge (Porifera: Demospongiae) of a new
genus and species of Palaeospongillidae from the Naskal
inter trappean deposit in the south-eastern part of the Deccan
volcanic province.
Nomenclatural acts.—This published work and the nomen-
clatural acts it contains have been registered in ZooBank:
urn:lsid:zoobank.org:pub:7B9F49CE-E5E9-46CD-BC9A-
17C204352BC1.
Institutional abbreviations.—PGDG, Museum Postgraduate
Department of Geology, Nagpur University, India; PGNU,
Postgraduate Department of Geology, Nagpur Uni versity,
India.
Other abbreviations.—NSKQ, Naskal Quarry section.
Geological setting
The Upper Cretaceous–lower Paleocene Deccan volca nic-
associated sediment covers an area of ~500,000 km2 in the
south, western and central parts of India. The Deccan vol-
canic flows are associated with infratrappean (deposited
below the volcanic flows) and intertrappean sediments (de-
posited in-between the two volcanic flows). Vertebrates,
invertebrates, micro and megaflora have been discovered in
both infratrappean and intertrappean sediments (Mohabey
1996; Mohabey and Udhoji 2000; Khosla and Sahni 2003).
The Naskal section is located at 17°14’21” N, 77°53’16” E,
in the Ranga Reddy District in the state of Telangana, India
(Fig. 1A). This section has a thickness of <3 m and a lateral
exposure of <15 m and an aerial distribution of <5 m2. It oc-
curs between Flow-3 and Flow-4 at an elevation of 624 m. In
the Naskal locality there are three geographically separated
sections designated as Naskal-A, Naskal-B, and Naskal GSI
Quarry (Wilson Mantilla et al. 2022: fig. 3). Based on Ar40/
Ar39 plagioclase dating of the flows, it is indicated that the
permissible age range of the Naskal intertrappean sedi-
ments is between 66.136 and 66.056 Ma at 68% confidence
(Wilson Mantilla et al. 2022), thus deposited close to the
Cretaceous/Paleogene boundary.
All the three Naskal sections were targeted for the sam-
pling but the sponge spicules were recovered only from the
Naskal GSI Quarry section (Fig. 1). The sponge spicule
bearing section has a thickness of 120 cm comprises black to
grey cherty limestone, hard yellowish shaly mudstone, loose
shaly to carbonate mudstone to marlstone, white mudstone,
and dark clay with sandy lenses in ascending stratigraphic
order (Fig. 1B). Naskal B (Fig. 1C) which is ~7 m west of the
Naskal GSI Quarry section yielded only palynomorphs (for
details see Wilson Mantilla et al. 2022).
Palaeoecology and age of the Naskal Intertrappean.—T he
Naskal intertrappean beds have yielded a rich mammal re-
cord (summarised in Wilson Mantilla et al. 2022), including
the first Indian record of a Cretaceous mammal (Prasad and
Sahni 1988). In addition to mammals, it has other vertebra-
tes such as fish, anurans, squamates, sphenodontian, turt les,
and crocodilians (Prasad and Sahni 1988; Prasad 2012).
Palyno flora of Naskal is represented by marker Maastrich-
tian taxa, i.e., Crybelosporites intertrappea, Maas tri ch tian–
Paleo cene taxa, i.e., Gabonisporis vigourouxii, M u l l e r i -
pollis bol puren sis, and Paleocene taxa, i.e., Stria col porites
striatus and Echistephanocolpites meghalayensis. Overall,
palynoassemblage indicates the presence of transitional
flora where Maastrichtian palynomorphs were depleted and
Paleocene flora was dominating.
Material and methods
Samples of shale, carbonate mudstone, and marlstone (100 g
each) were treated with 5% hydrochloric acid (HCl) until
the effervescence subsided. This was followed by thorough
washing with distilled water and treatment with 5% hydro-
fluoric acid (HF) for 5 minutes to remove the thin siliceous
secondary coating on the fossils. To remove organic mat-
ter, samples were sometimes treated with dilute nitric acid
(HNO3) and 5% potassium hydroxide (KOH). Every chemi-
cal treatment was followed by washing the sample with dis-
tilled water. After chemical treatments, the sieved samples
were divided into two aliquots, one for the preparation of
slides for Light Microscopy (LM) using polyvinyl alcohol
and Canada balsam, and the other for the preparation of
stubs for Scanning Electron Microscopy (SEM).

SAMANT ET AL.—CRETACEOUS/PALEOGENE FRESHWATER SPONGES FROM INDIA 169
All the slides were studied under a transmitted light
Olympus BX51 (Japan) microscope, and the photographs
were taken with a DP 20 (Olympus, Japan) camera. A to-
tal of 25 spicules from each category were studied and
measured under the LM microscope. For SEM studies, the
sponge spicules bearing sample was spread over a glass
slide and the slide was scanned under a microscope using
a 10X objective. The spicules were picked up with the help
of hair attached to the dissecting needle and placed over
the SEM stub. Later, the stub was coated with gold/palla-
dium, and SEM observations were carried out under the Jeol
microscope (Japan) at the Jawaharlal Nehru Aluminium
Research Development and Design Centre (JNARDDC),
Nagpur, India.
The holotype and paratype material, including the
rock samples, studied slides and SEM stubs, are housed in
the Museum of the Postgraduate Department of Geology
(PGDG), RTM Nagpur University, Nagpur, India. The ac-
ronym followed by sample number of the (a) holotype rock
sample is PGDG/NSKQ/2019; (b) slides (SL) followed by
numbers are PGNU/NSKQ/SL-1 to 13; (c) SEM stubs (ST)
followed by numbers are PGNU/NSKQ/ST-1, 2.
Terminology of morphotraits follows Manconi and Pron-
zato (2002). The systematic status of taxa was checked in
the World Porifera Database (Voogd et al. 2023).
Systematic palaeontology
Phylum Porifera Grant, 1836
Class Demospongiae Sollas, 1885
Subclass Heteroscleromorpha Cárdenas, Perez, and
Boury-Esnault, 2012
Order Spongillida Manconi and Pronzato, 2002
Family Palaeospongillidae Volkmer-Ribeiro and
Reitner, 1991
Fig. 1. Map of India showing Deccan volcanic province (green area). A. Location of Naskal intertrappean, Naskal B (white star) and Naskal GSI Quarry
sections (red star); map modified after Ahluwalia (1990) and Wilson Mantilla et al. (2022). B. Sponge spicule and diatom bearing horizon in Naskal GSI
Quarry section. C. Palynomorph bearing Naskal B section (modified after Wilson Mantilla et al. 2022).

170 ACTA PALAEONTOLOGICA POLONICA 68 (1), 2023
Genus Longibirotula Pronzato and Manconi nov.
Zoobank LSID: urn:lsid:zoobank.org:act:9A389B3D-118E-42AA-AD
9E-1AF484CE7681
Type species: Longibirotula antiqua Manconi and Samant, by mono-
typy.
Etymology: In reference to the long shaft of birotules gemmuloscleres,
gender feminine.
Diagnosis.—Longibirotula is characterised by gemmulo-
scleres slender birotules with very long spiny shaft, and two
types of skeletal megascleres, i.e., long slim oxeas and short
acanthoxeas.
Longibirotula antiqua Manconi and Samant sp. nov.
Figs. 1–5.
Zoobank LSID: urn:lsid:zoobank.org:act:A55B4EB4-F32C-41D0-B20
C-C1609F643060
Etymology: From Latin antiqua, old.
Type material: Holotype, rock sample PGDG/NSKQ/2019, slides
PGNU/NSKQ/SL-1–13, stubs PGNU/NSK/ST-1, 2.
Type locality: Naskal GSI Quarry, 17°14’21” N, 77°53’16” E, Ranga
Reddy District, Telangana State, India.
Type horizon: White marl/carbonate layers of Naskal GSI Quarry, Naskal
intertrappean beds (Upper Cretaceous–lower Paleo cene).
Fig. 3. Megascleres of palaeospongillid sponge Longibirotula antiqua gen. et sp. nov. from Upper Cretaceous–lower of Paleocene of Naskal GSI Quarry
(India). A–H. Acanthoxeas (slides PGNU/NSKQ/SL-1–13) with large spines. Diagenetic processes affect all spicules to various degree. Scale bars 20 µm.
Fig. 2. Megascleres of palaeospongillid sponge Longibirotula antiqua gen. et sp. nov. from Upper Cretaceous–lower Paleocene of Naskal GSI Quarry (India).
A–I. Oxeas (slides PGNU/NSKQ/SL-1–13) slim to stout with variably pointed tips. Diagenetic processes affect all spicules to various degree. Scale bars 20 µm.
{fig. will be greyscale in printed version}

SAMANT ET AL.—CRETACEOUS/PALEOGENE FRESHWATER SPONGES FROM INDIA 171
Diagnosis.—Longibirotula antiqua sp. nov. is character-
ized by slender gemmuloscleres birotules with long spiny
shaft, straight to slightly curved, with large scattered spines
variably dense and numerous. Rotules flat with indented
margins. Skeletal megascleres of two types, slim oxeas mi-
crospiny to smooth and short, stout acanthoxeas.
Description.—Skeletal megascleres monaxial with two
morphotypes. Slim, long, microspiny to smooth oxeas (142–
425 × 6–13 µm) rarely sinuous. Shorter, stouter acanthoxeas
(71–104 × 6–9 µm) straight to slightly curve with dense,
large spines. Microscleres absent. Entire gemmules are not
found because organic matter associated with gemmular
theca is not preserved. Fossil remains are only represented
by siliceous spicules. Gemmuloscleres birotules with very
long, straight to slightly curved, spiny shaft (47–76 µm in
length, 3–6 µm in thickness), with spines large, numer-
ous, variably dense, and rotules flat with indented margins
(12–19 µm in diameter).
Remarks.—Spicule deposits suggest that the Naskal in-
tertrappean palaeolake was inhabited by a population of
sponges with resting stages at the time of deposition. In
addition, centric diatoms (Aulacoseira spp.) and aquatic to
semiaquatic flora were also part of the biotic community.
Stratigraphic and geographic range.—Type locality and
horizon only.
Discussion
The association of centric diatoms Aulacoseira sp. with
sponge spicules in the uppermost part of the intertrappean
indicates the development of eutrophic conditions in the
lake, likely due to volcanogenic input in the palaeolake. The
presence of resting stages in the sponge remains (gemmulo-
scleres) suggests possible seasonal variations of water level.
Gemmulation processes, synchronised with the local
long-term seasonal rhythm, represent the most successful
evolutionary strategy in the life cycle of sponges to colo-
nize inland water; gemmules are asexual propagules able
to survive in unfavourable climatic/environmental critical
phases and to perform dispersal and defensive roles to per-
sist in situ; architecture of gemmules and their morphotraits
show a wide adaptive radiation worldwide (Manconi and
Pronzato 2002, 2008, 2009, 2015, 2016a, b; Manconi 2008).
Although entire gemmules have not been found in the
Naskal fossil remains, Longibirotula antiqua Manconi and
Samant sp. nov. is well distinguished from the rest of the
Fig. 4. Gemmuloscleres of palaeospongillid sponge Longibirotula antiqua gen. et sp. non. from Upper Cretaceous–lower Paleocene of Naskal GSI Quarry
(India). A–O. Birotules (slides PGNU/NSKQ/SL-1–13) slender, spiny, with long shaft. Diagenetic processes affect all spicules to various degree. Scale
bars 20 µm.

172 ACTA PALAEONTOLOGICA POLONICA 68 (1), 2023
known fossil taxa by a unique spicular complement lacking
skeletal microscleres, and composed of skeletal monaxons
(long slim oxeas, shorter acanthoxeas) and long gemmulo-
scleres birotules. The combination of these three spicular
morphs of Longibirotula antiqua Manconi and Samant sp.
nov. partially resembles with that of some fossil and living
taxa of the order Spongillida.
Gemmuloscleres morphologies and morphometries of
Longibirotula antiqua Manconi and Samant sp. nov. in-
dicate divergence from fossils birotule-bearing species of
the worldwide reported genus Ephydatia Lamouroux, 1816
(Pronzato et al. 2017), i.e., (i) Ephydatia fossilis Traxler,
1894 (Miocene in age, western Palaearctic, Romania) with
stout, spiny birotules (41–69 μm in length) and rotules
with indented margins (Traxler 1894); (ii) Ephydatia kai-
seri Rauff, 1926 (pre-middle Eocene in age, south-west-
ern Afrotropical, Namibia) with entire gemmules bearing
smooth birotules (44–65 μm in length) and non-incised
margins of flat rotules (Rauff 1926); (iii) Ephydatia chil-
eana Pisera and Sáez, 2003 (Late Miocene in age, south-
western Neotropical, Atacama Region, Chile) with birotules
(39–45 μm in length) bearing spiny shaft and rotules with
irregular margins, often deeply incised (Pisera and Saez
2003), and (iv) Ephydatia cf. facunda Weltner, 1895 (mid-
dle Eocene in age, Northern Nearctic, Canada) with stout
birotules (26–57 μm in length) with spiny shaft and strongly
incised margins of both rotules (Pisera et al. 2016).
As for extant birotules-bearing Spongillida, Longi-
birotula antiqua Manconi and Samant sp. nov. slightly re-
sembles some taxa, but gemmulosclere outline and/or rot-
ules morphs differ from species of e.g., Anheteromeyenia
Schröder, 1927; Corvoheteromeyenia Ezcurra de Drago,
1979; Corvomeyenia Weltner, 1913; Dosilia Gray, 1867;
Heteromeyenia Potts, 1881; Racekiela Bass and Volkmer-
Ribeiro, 1998, and Umborotula Penney and Racek, 1968
(Penney and Racek 1968; Manconi and Pronzato 2002,
2015, 2016b; Pronzato and Manconi 2019a).
The fossil Longibirotula antiqua Manconi and Samant
sp. nov. partly shares gemmuloscleres outline and length
(47–76 µm) with extant species of Ephydatia (e.g., 45–50
µm, Ephydatia robusta Potts, 1887), and Heteromeyenia (e.g.,
75–88 µm, Heteromeyenia stepanowi Dybowsky, 1884). The
Longibirotula antiqua Manconi and Samant sp. nov. biro-
tules morphs and length particularly resembles the slender
birotules known from the species of the genus Heterorotula
Penney and Racek, 1968 e.g., Heterorotula kakahuensis
(Traxler, 1896) (38–44 μm from New Zealand), Heterorotula
multidentata (Weltner, 1895) (64–84 µm from Australia,
Penney and Racek 1968; 32–48 μm from New Caledonia,
Rützler 1968) and Heterorotula caledonensis (Rützler, 1968)
(30–100 μm from New Caledonia) (Penney and Racek 1968;
Rützler 1968; Racek 1969; Pronzato and Manconi 2019b).
As for megascleres, smooth to microspiny long oxeas of
Longibirotula antiqua Manconi and Samant sp. nov. (142–
425 × 6–13 µm) partly show similarity with the megascleres
morphometric values in the species of Heterorotula (Penney
and Racek 1968; Racek 1969; Volkmer-Ribeiro and Motta
1995; Rützler 1968; Pronzato and Manconi 2002, 2019b),
namely (i) the Australian endemic lineage, e.g., Heterorotula
capewelli (Bowerbank, 1863) (195–330 × 13–18 µm, type
species), Heterorotula nigra (Lendenfeld, 1887)(224–360 ×
7–13 μm), Heterorotula multidentata (284–320 × 10–18 μm),
Heterorotula multiformis (Weltner, 1910) (330–420 × 13–
20 μm), (ii) the New Zealand–New Caledonia endemic lin-
Fig. 5. Spicular complement of skeleton and gemmules of palaeospongillid
sponge Longibirotula antiqua gen. et sp. nov. from Upper Cretaceous–lower
Paleocene of Naskal GSI Quarry (India) (slides PGNU/NSKQ/ST-1, 2).
A, B. Acan thoxeas short with dense spines. C, D. Oxeas fusiform, long and
with acute tips. E, F. Birotules with long shaft. Diagenetic processes affect
all spicules to various degree. Scale bars 20 µm.

SAMANT ET AL.—CRETACEOUS/PALEOGENE FRESHWATER SPONGES FROM INDIA 173
eage with Heterorotula kakahuensis (170–288 × 8–22 μm)
and Heterorotula caledonensis (100–150 × 8–19 μm) and,
in addition, with (iii) the Neotropical endemic lineage
Heterorotula fistula Volkmer-Ribeiro and Motta, 1995 (196–
361 × 16–31 μm).
Furthermore, short acanthoxeas of Longibirotula antiqua
(71–104 × 6–9 μm, here interpreted as gemmular cage com-
ponents) partly show similarity with those of Heterorotula,
which were reported as a short skeletal megascleres and/
or belonging to the gemmular cage, particularly evident in
H. kakahuensis endemic to New Zealand (Pronzato and
Manconi 2019b: fig. 7C, D) and H. caledonensis endemic to
New Caledonia (Pronzato and Manconi 2019b: fig. 4A, B)
In conclusion, the gemmuloscleres and megascleres
morphotraits of Longibirotula antiqua gen. et sp. nov. are
similar to that of some species of Heterorotula known from
living populations in Australia, New Zealand, and New
Cale donia and from subequatorial Brazil fossils remains,
suggesting a Gondwanan track which is not seen in the
Afrotropical Region (Manconi and Pronzato 2009).
The old evolutionary history and radiation of Spongillida
seem to be written in the gemmular architecture of coe val fos-
sil remains. The Longibirotula antiqua Manconi and Samant
sp. nov. from the Deccan intertrappean sediments close to the
Cretaceous/Paleogene boundary, presumably displays a ra-
dial arrangement of birotule spicules in the gemmular theca.
In contrast, the Deccan Upper Cretaceous (Maastrichtian,
Chron 30N) intertrappean sediments of Malwa Group were
inhabited by Palaeocorvospongilla cretacea characterized
by a totally different spicular complement of spiny stout ox-
eas/strongyles/strongyloxeas as gemmuloscleres and pseu-
dobirotules as skeletal microscleres (Samant et al. 2021).
The gemmules of the Early Cretaceous Palaeospongilla
chubutensis Ott and Volkmer, 1972, from Patagonia (Volk-
mer- Ribeiro and Reitner 1991) has oxeas gemmuloscleres
which are almost radially, irregularly arranged in the gem-
mular theca. On the basis of these record, the family Palaeo-
spongillidae is enlarged and now comprises five monotypic
genera: Eospongilla (Eospongilla morrisonensis Dunagan,
1999), Longibirotula Pronzato and Manconi gen. nov., Lute-
tio spongilla (Lutetiospongilla heili Richter and Wuttke,
1999), Palaeocorvospongilla Pronzato and Man coni, 2021,
and Palaeospongilla Ott and Volkmer, 1972.
The discovery of the new species Longibirotula antiqua
confirm that the Deccan volcanic province in India is a
favourable area to study the natural history of freshwater
sponges adaptive processes. The data give us more infor-
mation about how the anatomy of Spongillida have changed
or persisted over time and how they have always been able
to drive a morphological diversification by a successful
evolution of resistant bodies, i.e., gemmules (Manconi and
Pronzato 2002, 2015, 2016a). Importantly, the long lasting
structural conservative trend of freshwater sponges (Pisera
2006; Pronzato et al. 2017; Samant et al. 2021) seems to be
confirmed by Deccan spicule morphologies that remained
almost unchanged through tens of millions of years.
Acknowledgements
BS and DMM are thankful to Kirtikumar R. Randive (Department
of Geology, RTM Nagpur University, India), for providing working
facilities. We are also thankful to the Jawaharlal Nehru Aluminium
Research Development and Design Centre, Nagpur, India for help in
SEM study. Special thanks to Laura Negretti (DISTAV, Genova, Italy)
for her kind efforts in technical cooperation. We are also thankful to the
anonymous reviewer and Joachim Reitner (Georg-August-Universität
Göttingen, Germany) for useful comments and suggestions that sig-
nificantly improved the manuscript. BS and DMM are thankful to
the Science and Engineering Research Board, New Delhi under grant
(CRG/2020/001339) and National Science Foundation (EAR-1736787)
for financial assistance. RM research was co-funded by the Fondazione
di Sardegna (grant FdS/RAS-2016/CUP J86C1800082005), Regione
Autonoma Sardegna (grant RAS2012-LR7/2007-CRP-60215), Parco
Nazionale dell’Asinara (grant PNA2016), and Sassari University
(grant 2019-2021).
References
Ahluwalia, A.D. 1990. Detailed geological map and field observations in
the Cretaceous mammal bearing locality, Hyderabad District, Andhra
Pradesh, India. In: A. Sahni and A. Jolly (eds.), Cretaceous Event Stra-
tigraphy and the Correlation of the Indian Nonmarine Strata. Proceed-
ings Seminar cum Workshop IGCP 216 and 245, 120–122. Chandigarh.
Khosla, A. and Sahni, A. 2003. Biodiversity during the Deccan volcanic
eruptive episode. Journal of Asian Earth Sciences 21: 895–908.
Manconi, R. 2008. The genus Ephydatia (Spongillina: Spongillidae) in Af-
rica: a case of Mediterranean vs. southern Africa disjunct distribution.
Biogeographia. The Journal of Integrative Biogeography 29: 19–28.
Manconi, R. and Pronzato, R. 2002. Spongillina new suborder, Lubomir-
skiidae, Malawispongiidae n. fam., Metaniidae, Metschnikowiidae,
Palaeo spongillidae, Potamolepidae, Spongillidae. In: H.J.N Hooper
and R.W.M. Van Soest (eds.), Systema Porifera: A Guide to the Clas-
sification of Sponges 1, 921–1019. Kluwer Academic/Plenum Publish-
ers, New York.
Manconi, R. and Pronzato, R. 2008. Global diversity of sponges (Porifera:
Spongillina) in freshwater. Hydrobiologia 595: 27–33.
Manconi, R. and Pronzato, R. 2009. Atlas of African freshwater sponges.
Studies in Afrotropical Zoology 295: 1–214.
Manconi, R. and Pronzato, R. 2015. Phylum Porifera. In: J.H. Thorp and
D.C. Rogers (eds.), Ecology and General Biology: Thorp and Covich’s
Freshwater Invertebrates, vol 1. 4th Edition, 133–157. Academic Press,
Elsevier, London.
Manconi, R. and Pronzato, R. 2016a. How to survive and persist in tem-
porary freshwater? Adaptive traits of sponges (Porifera, Spongillida).
A review. Hydrobiologia 782: 11–22.
Manconi, R. and Pronzato, R. 2016b. Phylum Porifera. In: J.H. Thorp
and D.C. Rogers (eds.), Keys to Nearctic Fauna: Thorp and Covich’s
Freshwater Invertebrates, vol 2, 4th Edition, 39–83. Academic Press,
Elsevier, San Diego.
Mohabey, D.M. 1996. Depositional environments of Lameta Formation
(Late Cretaceous) of Nand-Dongargaon inland basin, Maharashtra:
The fossil and lithological evidence. Memoirs Geological Society of
India 37: 363–386.
Mohabey, D.M. and Udhoji, S.G. 2000. Vertebrate Fauna of Late Creta-
ceous dinosaur bearing Lameta Formation of Nand-Dongargaon in-
land basin, Maharashtra and KT boundary implications. Memoirs Geo-
logical Society of India 46: 295–322.
Penney, J.T. and Racek, A.A. 1968. Comprehensive revision of a world-
wide collection of freshwater sponges (Porifera, Spongillidae). Bulle-
tin of the United States National Museum 272: 1–184.

174 ACTA PALAEONTOLOGICA POLONICA 68 (1), 2023
Pisera, A. 2006. Palaeontology of sponges—a review. Canadian Journal
of Zoology 84: 242–261.
Pisera, A. and Saez, A. 2003. Paleoenvironmental significance of a new
species of freshwater sponge from the Late Miocene Quillagua Forma-
tion (N Chile). Journal of South American Earth Studies 15: 847–852.
Pisera, A., Manconi, R., Siver, P., and Wolfe, A. 2016. The sponge genus
Ephydatia from the Middle Eocene: environmental and evolutionary
significance. Paläontologische Zeitschrift 90: 673–680.
Pisera, A., Siver, P.A., and Wolfe, A.P. 2013. A first account of freshwater
potamolepid sponges (Demospongiae, Spongillina, Potamolepidae)
from the middle Eocene: biogeographic and paleoclimatic implica-
tions. Journal of Paleontology 87: 373–378.
Prasad, G.V.R. 2012. Vertebrate biodiversity of the Deccan volcanic prov-
ince of India: a review. Bulletin de la Société géologique de France 183:
597–610.
Prasad, G.V.R. and Sahni, A. 1988. First Cretaceous mammal from India.
Nature 332: 638–640.
Pronzato, R. and Manconi, R. 2019a. An overview on the freshwater sponge
fauna (Demospongiae: Spongillida) of New Zealand and New Caledo-
nia with new insights into Heterorotula from deep thermal vents of the
Lake Taupo. Journal of Natural History 53: 2207–2229.
Pronzato, R. and Manconi, R. 2019b. Phylum Porifera. In: J.H. Thorp and
D.C. Rogers (eds.), Keys to Palaearctic Fauna. Thorp and Covich’s
Fresh water Invertebrates. 4th Edition, 39–83. Academic Press, London.
Pronzato, R. and Manconi, R. (in press). Phylum Porifera. In: J.H. Thorp
and D.C. Rogers (eds.), Keys to Australasian Fauna. Thorp and
Covich’s Freshwater Invertebrates. 4th Edition. Academic Press, El-
sevier, London.
Pronzato, R., Pisera, A., and Manconi, R. 2017. Fossil freshwater sponges:
Taxonomy, geographic distribution, and critical review. Acta Palaeon-
tologica Polonica 62: 467–495.
Rauff, H. 1926. Überprämitteleozäne fossilfuhrende Süswasser-Hornstein-
eaus der Namib. In: E. Kaiser (ed.), Die Diamantenwuste Südwest-
Afrikas 2: 160–166.
Racek, A.A. 1969. The freshwater sponges of Australia (Porifera: Spongil-
lidae). Marine and Freshwater Research 20: 267–310.
Rützler, K. 1968. Fresh-water sponges from New Caledonia. Cahiers
ORSTOM (Office de la Recherche Scientifique et Technique Outre-
Mer), Series Hydrobiologie 2: 57–66.
Samant, B. and Mohabey, D.M. 2009. Palynoflora from Deccan volcano-
sedimentary sequence (Cretaceous–Paleocene transition) of central
India: implications for spatio-temporal correlation. Journal of Bio-
sciences 34: 811–823.
Samant, B. and Mohabey, D.M. 2014. Deccan volcanic eruptions and their
impact on flora: palynological evidence. Geological Society of Ame-
rica, Special Papers 505: 171–191.
Samant, B., Mohabey, D.M., Srivastava, P., and Thakre, D. 2014. Paleo-
cene palynoflora from the intertrappean sediments of Saurashtra, Gu-
jarat: age and paleoenvironments. Journal of Earth System Science
123: 219–232.
Samant, B., Pronzato, R., Mohabey, D.M., Kumar, D., Dhobale, A., Pizal,
P., and Manconi, R. 2021. Insight into the evolutionary history of fresh-
water sponges: A new genus and new species of Spongillida (Porifera:
Demospongiae) from Upper Cretaceous (Maastrichtian) Deccan inter-
trappean lacustrine deposits of the Malwa Group, Central India. Creta-
ceous Research 126: 104851.
Schindler, T., Wuttke, M.P., and Poschmann, M. 2008. Oldest record
of freshwater sponges (Porifera: Spongillina) spiculite finds in the
Permo- Carboniferous of Europe. Paläontologische Zeitschrift 82:
373–384.
Traxler, L. 1894. Ephydatia fossilis, eine neu Art der fossilen Spongilliden.
Földatni Közlöny 24: 234–237.
Volkmer-Ribeiro, C. and Motta, J.F.M. 1995. Esponjas formador as de es-
pongilitos em lagoas no Triângulo Mineiro e ajacências com indicaçao
de preservaçao de habitat. Biociencias 3: 183–205.
Volkmer-Ribeiro, C. and Reitner, J. 1991. Renewed study of the type ma-
terial of Palaeospongilla chubutensis Ott and Volkheimer (1972). In:
J. Reitner and H. Keupp (eds.), Fossil and Recent Sponges, 121–133.
Springer, Berlin.
Voogd, N.J. de, Alvarez, B., Boury-Esnault, N., Carballo, J.L., Cárdenas, P.,
Díaz, M.-C., Dohrmann, M., Downey, R., Hajdu, E., Hooper, J.N.A.,
Kelly, M., Klautau, M., Manconi, R., Morrow, C.C., Pisera, A.B., Ríos,
P., Rützler, K., Schönberg, C., Vacelet, J., and van Soest, R.W.M. 2023.
World Porifera Database [available online, https://www.marinespecies.
org/porifera/porifera.php?p=taxdetails&id=845456 on 2023-03-06].
Wilson Mantilla, G.P., Renne, P.R., Samant, B., Mohabey, D.M., Dhobale,
A., Tholt, A.J., Tobin, T.S., Widdowson, M., Anantharaman, S., Das-
sarma, D.C., and Wilson Mantilla, J.A. 2022. New mammals from the
Naskal intertrappean site and the age of India’s earliest eutherians.
Palaeo geography, Palaeoclimatology, Palaeoecology 591: 110857.