Abstract
The environmental background to the Neolithic occupation of the Rio Sizandro, western Portugal is elucidated via the ostracod assemblages and sedimentology of a well-dated part of borehole COU_14 from near the village of Coutada. Results from this location, under greater marine influence than sites previously studied, confirm changes from fluvial to brackish estuarine and lagoonal conditions driven by the interaction of changing eustatic sea level, barrier lagoon formation and valley infill from erosion related to climate change and possible human activity.
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Acknowledgements
We thank the Deutsches Archäologisches Institut in Madrid for financial support. Moreover, we are grateful to Heinrich Thiemeyer (Institut für Physische Geographie, Frankfurt am Main) and Nico Herrmann (Institut für Geographie, Hildesheim) as well as to students Ina Charlotte Haase, Christian Sänger and Stefan Sylla (formerly Institut für Physische Geographie, Frankfurt am Main) for field assistance. Claudia Franz (Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main) and Telmo Nunes (Unidade de Microscopia, Faculty of Sciences, University of Lisbon) took the SEM images, for which we thank them. We are grateful to Hans-Peter Stika (Institut für Botanik, Stuttgart-Hohenheim) who analysed the terrestrial macro remains suitable for radiocarbon dating, Holger Rittweger (Waldbrunn/Westerwald) for analyses of bivalve and gastropod shells and Arie Joop Kalis (Boxum, The Netherlands), Wim van Leeuwaarden (Monchique, Algarve, Portugal) and Astrid Stobbe (Institut für Archäologische Wissenschaften, Frankfurt am Main) for pollen analyses. We also thank Doris Bergmann-Dörr and Dagmar Schneider (Institut für Physische Geographie, Frankfurt am Main) for geochemical analyses of sediment samples. Figures 6, 7 and 8 and Fig. 5 were kindly prepared respectively by Isabel Loureiro (Centro de Geologia, Faculty of Sciences, University of Lisbon) and Vera Lopes (Departament of Geology, Faculty of Sciences, University of Lisbon). Ilaria Mazzini (Roma) and Peter Frenzel (Jena) carefully reviewed this manuscript and provided valuable suggestions.
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Appendices
Appendix 1
List of ostracod taxa found in Borehole COU_14, Coutada, Rio Sizandro.
The species of the genera Neonesidea, Paradoxostoma, Sclerochilus and Xestoleberis, all of them generally uncommon, were rarely identified due to their scarcity and dimensions (juveniles) and were considered together as spp. for the counts (Fig. 5). Aurila convexa and Aurila woutersi were identified when present as adults, as most of the specimens were juveniles they were counted together (Fig. 5). Identifications were mainly based on Athersuch et al. (1989), Bonaduce et al. (1976), Meisch (2000) and Cabral and Loureiro (2013). Systematics according to Horne et al. (2002).
Class Ostracoda Latreille, 1802
Subclass Podocopa Sars, 1866
Order Podocopida Sars, 1866
Suborder Cytherocopina Gründel, 1967
Superfamily Cytheroidea Baird, 1850
Family Bythocytheridae Sars, 1866
Genus Bythocythere Sars, 1866
Bythocythere bradyi Sars, 1926 - Fig. 6a
Genus Pseudocythere Sars, 1866
Pseudocythere caudata Sars, 1866
Genus Sclerochilus Sars, 1866
Sclerochilus spp. - Fig. 6b
Family Cuneocytheridae Mandelstam, 1959
Genus Cuneocythere Lienenklaus, 1894
Cuneocythere semipunctata (Brady, 1868)
Family Cushmanideidae Puri, 1974
Genus Pontocythere Dubowsky, 1939
Pontocythere elongata (Brady, 1868) - Fig. 6c
Family Cytherideidae Sars, 1925
Genus Cyprideis Jones, 1857
Cyprideis torosa (Jones, 1850) - Fig. 6d–e
Family Cytheruridae G. W. Müller, 1894
Genus Cytheropteron Sars, 1866
Cytheropteron dorsocostatum Whatley and Masson, 1980 - Fig. 7b
Genus Hemicytherura Elofson, 1941
Hemicytherura aff. defiorei Ruggieri, 1953 - Fig. 6f
Genus Microcytherura G. W. Müller, 1894
Microcytherura fulva (Brady and Robertson, 1874) - Fig. 6g
Genus Pseudocytherura Dubowsky, 1939
Pseudocytherura cf. calcarata (Seguenza, 1880) - Fig. 6h
Genus Semicytherura Wagner, 1957
Semicytherura acuminata (G.W.Müller, 1894) - Fig. 6i
Semicytherura acuta (G.W.Müller, 1912) - Fig. 6j–k
Semicytherura acuticostata ventricosa (Sars, 1866) - Fig. 6l
Semicytherura arcachonensis Yassini, 1969 - Fig. 6m
Semicytherura robertsi Whittaker and Horne, 2009 - Fig. 6n–o
Semicytherura sella (Sars, 1866) - Fig. 5p
Semicytherura cf. stilifera Bonaduce, Ciampo and Masoli, 1976 - Fig. 6q
Semicytherura sulcata (G. W. Müller, 1894) - Figs. 6r and 7a
Genus Tetracytherura Ruggieri, 1952
Tetracytherura angulosa (Seguenza, 1880)
Family Eucytheridae Puri, 1954
Genus Eucythere Brady, 1868
Eucythere prava Brady and Robertson, 1869 - Fig. 7c
Family Hemicytheridae Puri, 1953
Genus Aurila Pokorny, 1955
Aurila arborescens (Brady, 1865)
Aurila convexa (Baird, 1850) - Fig. 7d
Aurila woutersi Horne, 1986 - Fig. 7e
Genus Caudites Coryell and Fields, 1937
Caudites calceolatus (O. G. Costa, 1853) - Fig. 7f
Genus Hemicythere Sars, 1925
Hemicythere rubida (Brady, 1868)
Genus Heterocythereis Elofson, 1941
Heterocythereis albomaculata (Baird, 1838) - Fig. 7g–i
Genus Urocythereis Ruggieri, 1950
Urocythereis britannica Athersuch, 1977 - Fig. 7k
Family Leptocytheridae Hanai, 1957
Genus Callistocythere Ruggieri, 1953
Callistocythere badia (Norman, 1862) - Fig. 7l
Callistocythere curryi Horne, Lord, Robinson and Whittaker, 1990 - Fig. 7m
Callistocythere littoralis (G. W. Müller, 1894)
Callistocythere murrayi Whittaker, 1978 - Fig. 7n
Genus Leptocythere Sars, 1928
Leptocythere castanea (Sars, 1866) - Fig. 7o
Leptocythere cribrosa (Brady, Crosskey and Robertson, 1874)
Leptocythere lacertosa (Hirschmann, 1912)
Leptocythere macallana (Brady and Robertson, 1869)
Leptocythere fabaeformis (G. W. Müller, 1894) - Fig. 7p–r
Leptocythere pellucida (Baird, 1850) - Fig. 8a–b
Leptocythere porcellanea (Brady, 1869) - Fig. 8c
Family Limnocytheridae Klie, 1938
Limnocythere s. str. Brady, 1867
Limnocythere inopinata (Baird, 1845) - Fig. 8r
Family Loxoconchidae Sars, 1925
Genus Elofsonia Wagner, 1957
Elofsonia pusilla (Brady and Robertson, 1870)
Genus Loxoconcha Sars, 1866
Loxoconcha elliptica Brady, 1868 - Fig. 8d
Loxoconcha malcomsoni Horne and Robinson, 1985
Loxoconcha rhomboidea (Fischer, 1855) - Fig. 8e
Genus Palmoconcha Swain and Gilby, 1974
Palmoconcha laevata (Norman, 1865)
Genus Roundstonia Neale, 1973
Roundstonia robertsoni (Brady, 1868)
Genus Sagmatocythere Athersuch, 1976
Sagmatocythere napoliana (Puri, 1963) - Fig. 8f
Family Neocytherideidae Puri, 1957
Genus Neocytherideis Puri, 1957
Neocytherideis subulata (Brady, 1868) - Fig. 8g
Genus Procytherideis Ruggieri, 1978
Procytherideis aff. subspiralis (Brady, Crosskey and Robertson, 1874)
Genus Sahnicythere Athersuch, 1982
Sahnicythere retroflexa (Klie, 1936)
Family Paracytherideidae Puri, 1957
Genus Paracytheridea G. W. Müller, 1894
Paracytheridea sp.
Family Paradoxostomatidae Brady and Norman, 1889
Genus Cytherois G. W. Müller, 1884
Cytherois fischeri (Sars, 1866) - Fig. 8h
Genus Paradoxostoma Fischer, 1855
Paradoxostoma spp.
Family Thaerocytheridae Hazel, 1967
Genus Thaerocythere Hazel, 1967
Thaerocythere hoptonensis (Brady, Crosskey and Robertson, 1874) - Fig. 7j
Family Trachyleberididae Sylvester-Bradley, 1948
Genus Basslerites, Teichert, 1937
Basslerites teres (Brady, 1869) - Fig. 8i
Genus Carinocythereis Ruggieri, 1956
Carinocythereis whitei (Baird, 1850)
Genus Costa Neviani, 1928
Costa runcinata (Baird, 1850)
Genus Hiltermannicythere Bassiouni, 1970
Hiltermannicythere emaciata (Brady, 1867)
Family Xestoleberididae Sars, 1928
Genus Xestoleberis Sars, 1866
Xestoleberis rubens Whittaker, 1978 - Fig. 8j
Xestoleberis spp.
Suborder Bairdiocopina Gründel, 1967
Superfamily Bairdioidea Sars, 1888
Family Bairdiidae Sars, 1888
Genus Bairdia McCoy, 1844
‘Bairdia’ subcircinata (Brady and Norman, 1869) - Fig. 8k
Genus Neonesidea Maddocks, 1969
Neonesidea sp. 2 (Bonaduce, Ciampo and Masoli, 1976) - Fig. 8l
Neonesidea spp.
Suborder Cypridocopina Gründel, 1967
Superfamily Cypridoidea Baird, 1845
Family Cyprididae Baird, 1845
Genus Cypridopsis Brady, 1867
Cypridopsis vidua (O.F. Müller, 1776) - Fig. 8m
Genus Sarscypridopsis McKenzie, 1977
Sarscypridopsis aculeata (Costa, 1847)
Family Ilyocyprididae Kaufmann, 1900
Genus Ilyocypris Brady and Norman, 1889
Ilyocypris bradyi Sars, 1890 - Fig. 8n–o
Ilyocypris inermis Kaufmann, 1900 - Fig. 8p–q
Appendix 2
Main ecological groups of the ostracod species collected in Borehole COU_14, Coutada, Rio Sizandro, with remarks on environmental significance of the most important species. Ecological characteristics especially based on Athersuch et al. (1989), Bonaduce et al. (1976) and Cabral and Loureiro (2013).
Group 1: brackish water species.
Cyprideis torosa (Jones, 1850) - a widespread species (Europe, Asia, Africa), highly euryhaline, found from inland lakes to marginal marine settings; prefers a muddy or sandy mud substrate, also found on algae (Athersuch et al. 1989). In Portugal found alive from the tidal flat to the low marsh in estuaries and lagoons (Cabral and Loureiro 2013).
Loxoconcha elliptica Brady, 1868 - a very common species found from NW Europe to the Mediterranean, usually associated with algae and mud, in estuaries, lagoons and pools (Athersuch et al. 1989). In Portugal, highly euryhaline, found alive from the tidal flat to the high marsh in estuaries and lagoons (Cabral and Loureiro 2013).
Cytherois fischeri (Sars, 1866) - a common highly euryhaline species, found from the Mediterranean to western Europe, usually associated with sand and algae, often found close to river mouths (Athersuch et al. 1989). In Portugal found alive especially on muddy substrates, in marshes of several estuaries, from the tidal flat to the lower part of the low marsh (Cabral and Loureiro, 2013).
Leptocythere porcellanea (Brady, 1869) - a western European species, usually found in mud substrates in sheltered creeks (Athersuch et al. 1989). In Portugal, highly euryhaline, found alive especially on muddy substrates, in marshes of several estuaries, from the tidal flat to the lower part of the high marsh (Cabral and Loureiro 2013).
Leptocythere lacertosa (Hirschmann, 1912) - an estuarine western European species, usually found on mud or fine sand (Athersuch et al. 1989). In Portugal found alive on muddy and sandy muddy substrates, in marshes of several estuaries, from the tidal flat to the lower part of the high marsh (Cabral and Loureiro 2013).
Callistocythere murrayi Whittaker, 1978 - a western European species, usually associated with algae (Athersuch et al. 1989), found alive in Portugal on muddy and sandy mud substrates, in marshes of several estuaries, from the tidal flat to the low marsh (Cabral and Loureiro 2013).
Loxoconcha malcomsoni Horne and Robinson, 1985 - a rare species, until now only known living in outer estuaries of British Isles and mainland Portugal (probable record in Azores – Meireles et al. 2014), in mud and mud-sand substrates, in salt marshes, close to the low marsh (Horne and Boomer 2000; Loureiro et al. 2009; Cabral and Loureiro 2013).
Group 2: marine littoral to sublittoral species, prefering sandy to silty sandy or muddy substrates, frequently with algae (phytal forms).
Loxoconcha rhomboidea (Fischer, 1855) - a phytal marine species, living in western Europe and Mediterranean, generally on littoral and shallow sublittoral zones, also found in outer estuaries (Athersuch et al. 1989; Athersuch and Whittaker 1976). In Portugal, found alive from the low to the high marsh in rare estuaries (Cabral and Loureiro 2013).
Heterocythereis albomaculata (Baird, 1838) - a phytal, littoral and sublittoral marine species, often abundant in rock pools, living in western Europe and Mediterranean (Athersuch et al. 1989). In Portugal found alive associated with green algae and sediment, in outer estuaries and intertidal rock pools and therefore having some tolerance of salinity variation (Cabral and Loureiro 2013).
Leptocythere fabaeformis (G.W. Müller, 1894) - a phytal/littoral marine species, euryhaline, tolerating a very wide salinity range, 13 to 33 (Yassini 1969), known from the Mediterranean to western Europe, until France. In Portugal found alive on mud substrates in the tidal flat of only one estuary (Cabral and Loureiro 2013).
Aurila convexa (Baird, 1850) - known from the Mediterranean to western Europe, until France/southern Britain, is a common species, living amongst algae, algal debris or on sand, silty sand and silt at different depths in continental shelf (up to 122 m in the Mediterranean) and in littoral zones (Athersuch et al. 1989; Bonaduce et al. 1976). In Portugal, until now, not found alive, but empty valves and carapaces are frequently abundant in Holocene and Recent marginal marine settings and in Recent littoral to sublittoral (until the continental slope) sediments (Cabral and Loureiro 2013).
Aurila woutersi Horne, 1986 – a phytal, littoral and shallow sublittoral species, living in western Europe (surely found in Great Britain - Athersuch et al., 1989). In Portugal, until now, never found alive but empty valves and carapaces were found in Holocene and Recent marginal marine sediments (Cabral and Loureiro 2013).
Aurila arborescens (Brady, 1865) - known from the Mediterranean, as Aurila woodwardi (Brady, 1868) to western Europe, until southern Britain, it is a littoral? to sublittoral marine species living in the Mediterranean at depths not exceeding 20 m (Athersuch et al. 1989; Bonaduce et al. 1976). In Portugal, until now, not found alive, but carapaces and empty valves were found in Holocene marginal marine sediments (Cearreta et al. 2003; Cabral et al. 2006; Cabral et al. 2011a; Cabral and Loureiro 2013).
Pontocythere elongata (Brady, 1868) - a western European species generally living on sand substrates, in marine and outer estuarine conditions (Athersuch et al. 1989). In Portugal, until now, not found alive, but empty valves and carapaces are frequent in marginal marine Holocene and Recent sediments (Cabral and Loureiro 2013).
Urocythereis britannica Athersuch, 1977 - a western European species generally living on sand substrates, in marine littoral and shallow sublittoral conditions (Athersuch et al. 1989). In Portugal, until now, not found alive, but empty valves and carapaces are frequent in Holocene and Recent marginal marine settings and in Recent littoral to sublittoral (until the continental slope) sediments (Cabral and Loureiro 2013).
Thaerocythere hoptonensis (Brady, Crosskey and Robertson, 1874) - a western European species, with records in Recent sediments from the United Kingdom, Spain, northern Morocco (continental shelf) and Portugal (Wood and Whatley 1997). In Portugal, until now, not found alive, but empty valves and carapaces are frequent in marginal marine Holocene and Recent sediments and in the western Algarve continental shelf, until 104 m depth (Cabral et al. 2011b; Cabral and Loureiro 2013).
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Cabral, M.C., Lord, A.R., Dambeck, R. et al. Ostracod evidence for the Neolithic environment of Rio Sizandro, Portugal: Part 2. Palaeobio Palaeoenv 96, 541–557 (2016). https://doi.org/10.1007/s12549-016-0240-5
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DOI: https://doi.org/10.1007/s12549-016-0240-5