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A New Neolepadid Cirripede from a Pleistocene Cold Seep, Krishna-Godavari Basin, Offshore India

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A New Neolepadid Cirripede from a Pleistocene Cold Seep, Krishna-Godavari Basin, Offshore India A new neolepadid cirripede from a Pleistocene cold seep, Krishna-Godavari Basin, offshore India ANDY S. GALE, CRISPIN T.S. LITTLE, JOEL E. JOHNSON, and LIVIU GIOSAN Gale, A.S., Little, C.T.S, Johnson, J.E., and Giosan, L. 2020. A new neolepadid cirripede from a Pleistocene cold seep, Krishna-Godavari Basin, offshore India. Acta Palaeontologica Polonica 65 (2): 351–362. Valves of a thoracican cirripede belonging to a new species of the Neolepadidae, Ashinkailepas indica Gale sp. nov. are described from a Late Pleistocene cold seep (52.6 ka), cored in the Krishna-Godavari Basin, offshore from the eastern coast of India. This constitutes the first fossil record of the genus, and its first occurrence in the Indian Ocean. Other fossil records of the Neolepadidae (here elevated to full family status) are discussed, and it is concluded that only Stipilepas molerensis from the Eocene of Denmark, is correctly referred to the family. Cladistic analysis of the Neolepadidae sup- ports a basal position for Ashinkailepas, as deduced independently from molecular studies, and the Lower Cretaceous brachylepadid genus Pedupycnolepas is identified as sister taxon to Neolepadidae. Neolepadids are not Mesozoic relics as claimed, preserved in association with the highly specialised environments of cold seeps and hydrothermal vents, but are rather an early Cenozoic offshoot from the clade which also gave rise to the sessile cirripedes. Key words: Cirripedia, Thoracica, Neolepadidae, cold seep, Pleistocene, Indian Ocean. Andy S. Gale [[email protected]], School of Earth, Environmental and Geographical Sciences, University of Port- smouth, Burnaby Building, Burnaby Road, Portsmouth PO13QL, UK. Crispin T.S. Little [[email protected]], School of Earth and Environment, University of Leeds, Leeds LS2 9JT, UK. Joel E. Johnson [[email protected]], Department of Earth Sciences, University of New Hampshire, Durham, NH 03824, USA. Liviu Giosan [[email protected]], Geology & Geophysics, Woods Hole Oceanographic Institution, Woods Hole, MA 02543, USA. Received 23 November 2019, accepted 14 February 2020, available online 22 April 2020. Copyright © 2020 A.S. Gale 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. from the Lower Jurassic (Sinemurian–Pliensbachian) of New Introduction Caledonia, Litholepas klausreschi Nagler, Haug, Glenner, Amongst the remarkable range of invertebrates associated and Buckeridge, 2017 from the Upper Jurassic (Tithonian) of with deep sea hydrothermal vents and cold seeps are tho- Germany, and Stipilepas molerensis Carriol in Carriol et al., racican cirripedes referred to the subfamily Neolepadinae 2016, from the Eocene (Ypresian) of Denmark. None of these Newman, 1985. Species belonging to this taxon are cur- are associated with hydrothermal vents or cold seeps. rently known from the South Atlantic, Indian, Pacific, and In this paper, we describe a new species of Ashinkailepas Southern oceans (Herrera et al. 2015; Watanabe et al. 2018). from the Late Pleistocene (52.6 ka) found in a sediment core It is likely that many of these species obtain carbon through from the Krishna-Godavari (K-G) Basin, offshore India, endosymbiotic chemoautotrophic bacteria (Southward and which was associated with a contemporary methane hydrate Newman 1998). A total of eight species of living neolepadines system and methane seepage. On the basis of this new spe- have been described, and assigned to the genera Neolepas, cies we review the taxonomy, possible origin and fossil re- Ashinkailepas, Vulcanolepas, and Leucolepas. Neolepadids cord of the Neolepadinae, and undertake a cladistic analysis have been identified as relic Mesozoic forms which have of the subfamily. survived in highly specialised habitats and considered to be primitive scalpellomorphs, most closely allied to the Jurassic Institutional abbreviations.—MJML, Museum of Jurassic Archaeolepas (Newman 1979; Yamaguchi et al. 2004). Marine Life, Kimmeridge, Dorset, UK; NHMUK, Natu ral Three fossil taxa have been assigned to the Neolepadinae; History Museum, London, UK; SMNS, Stuttgart Museum ?Neolepas augurata Buckeridge and Grant-Mackie, 1985, of Natural History, Stuttgart, Germany; UIWA, National Acta Palaeontol. Pol. 65 (2): 351–362, 2020 https://doi.org/10.4202/app.00705.2019 352 ACTA PALAEONTOLOGICA POLONICA 65 (2), 2020 82°E 83°E hemipelagic clay (Collett et al. 2015; Phillips et al. 2014) containing greater or lesser quantities of organic carbon (Johnson et al. 2014), and calcareous nannofossils and for- 17°N aminifera (Flores et al. 2014). The cirripedes come from a level between 16 and 17 metres below sea floor (mbsf) in this core (Table 1) and they are associated with whole and fragmentary shells of seep-related bivalves and gastro- pods. In addition, there are numerous variably sized authi- genic carbonate nodules formed of high magnesium calcite, two of which have δ13C values of -48.23‰ and -45.33‰ (VPDB), indicative the anaerobic oxidation of methane (AOM) near the sulfate-methane transition zone (SMTZ) Krishna-Godavari (Teichert et al. 2014). The presence of the seep molluscs and 16°N Basin methane derived authigenic carbonates (MDAC) defines a palaeo-seafloor SMTZ and thus palaeo-methane seepage. Similar shell material and MDAC have been found at the same ~16 mbsf level in two nearby sediment cores: NGHP- 01-10D (15°51.8609’ N, 81°50.0749’ E, 1038 m water depth; Collett et al. 2015; Teichert et al. 2014), and a 29.9 m long sediment core collected by Giant Calypso piston corer water Bay of Bengal during cruise MD-161 of the R/V Marian Dufresne in May 2007 at 15°51.8624’ N, 81°50.0692’ E in 1033 m water depth 100 km Arabian (Mazumdar et al. 2009). An age model developed from a 15°N Sea combined dataset of radiocarbon dates from foraminifera tests, bivalve shells and MDAC from the ~16 mbsf level Fig. 1. NGHP-01 core sites (circles) and logging sites (stars) in the offshore from these three cores yields an extrapolated age of ~52.6 ka Krishna-Godavari Basin, modified from Lorenson and Collett (2018). 14 Site 12 is co-located with sites 10 and 21 (red circle). Inset shows the study C years BP. (Mazumdar et al. 2009; Connolly et al. 2009), area in Krishna-Godavari Basin is located on the eastern margin of penin- equating to the Late Pleistocene stage. Occurrences are sular India in the western Bay of Bengal. listed in Table 1. Institute of Water and Atmospheric Research, Wellington, Table 1. Details of specimens obtained from NGHP-01 Site 12 Hole A, New Zealand; SNM, Smithsonian Museum, Washington DC, Core 3H, Section 2, offshore K-G Basin, India. USA; UoA, University of Auckland, Geology Department, Sample interval (cm) Depth (mbsf) Number of specimens New Zealand. 52–57 16.38 10 50–60 16.38 5 Other abbreviations.—AOM, anaerobic oxidation of meth- 80–90 16.68 13 ane; K-G, Krishna-Godavari; mbsf, metres below sea floor; 84–86 16.68 12 MDAC, methane derived authigenic carbonates; SMTZ, sul- 90–100 16.78 2 fate-methane transition zone; VPDB, Vienna PeeDee Bele- 100–110 16.88 5 mnite. 116–118 17 11 Nomenclatural acts.—This published work and the nomen- clatural acts it contains have been registered in ZooBank: urn:lsid:zoobank.org:pub:7D8A63AC-765E-4C95-8610- B8ED6D7B7605. Systematic palaeontology (by Andy S. Gale) Geological and geographic setting Infraclass Cirripedia Burmeister, 1834 Superorder Thoracica Darwin, 1854 The studied cirripede specimens come from sediment core Clade Thoracicalcarea Gale, 2015a NGHP-01-12A-03H-2, which was drilled in 2006 by the D/V JOIDES Resolution during the Indian National Gas Diagnosis.—Thoracica in which the shell plates are com- Hydrate Program Expedition 01 (NGHP-01) in 1046 m of posed of calcite. water depth at 15°51.6335’ N, 81°50.2274’ E in the K-G Clade Unilatera Gale, 2019a Basin, on the eastern Indian continental margin (Collett et al. 2015, 2014). The sediment in the core and through- Diagnosis.—Thoracicalcarea in which only one lateral plate out the offshore K-G Basin consists of variably coloured (the paired upper latus) is present; rostrum large, triangular, GALE ET AL.—PLEISTOCENE NEOLEPADID CIRRIPEDE FROM OFFSHORE INDIA 353 A C D1 D2 E t t 2 mm c B s s c r l 1 mm 1 mm 1 mm p 1 mm Fig. 2. Fossil taxa assigned to Neolepadidae. A, B. ?Neolepas augurata Buckeridge and Grant-Mackie, 1885 (original of Buckeridge and Grant-Mackie 1985: figs. 2, 3); Sinemurian–Pliensbachian (Lower Jurassic), New Caledonia. A. University of Auckland Geology Department UoA A291a, b, external mould of tergum. B. University of Auckland Geology Department UoA A292a, b, external mould of scutum. This is here tentatively interpreted as an eolepadid. C. Toarcolepas mutans Gale and Schweigert, 2015 (SMNS 26029); Toarcian (Jurassic), Harpoceras falciferum Zone, Zell u. Aichelberg, Ger- many; reconstruction based upon type material (Gale and Schweigert 2015: fig. 5). D. Litholepas klausreschi Nagler, Haug, Glenner, and Buckeridge, 2017; Tithonian (Upper Jurassic), Hybernoticeras hybernotum Zone, Eichstatt, Germany; D2, SMNS 70388/5 (decolourised original of Nagler et al. 2017: fig. 5B). I am unable to identify the upper latus, marked “l” in their figure; D1, reconstruction (mirrored for comparison with C). E. Con cinnalepas costata (Withers, 1928) (NHMUK IC 1103); Kimmeridge Clay, 2 m beneath Freshwater Stone Band, Pectinatites pectinatus Zone, Tithonian (Upper Jurassic), Kimmeridge, Dorset, UK; fragmentary capitulum, showing scutum, tergum, and two lateral plates (based on original of Gale 2014: fig. 6l). Abbreviations: c, carina; l, upper latus; p, peduncle; r, rostrum; s, scutum; ; t, tergum. external to the scuta. Surface sculpture plesiomorphically ?Neolepas augurata Buckeridge and Grant-Mackie, composed of evenly spaced terracing.
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