Palaeobio Palaeoenv DOI 10.1007/s12549-016-0240-5

ORIGINAL PAPER

Ostracod evidence for the Neolithic environment of Rio Sizandro, Portugal: Part 2 Maria Cristina Cabral 1 & Alan R. Lord 2 & Rainer Dambeck 3 & Michael Kunst 4

Received: 21 March 2016 / Revised: 30 May 2016 / Accepted: 21 June 2016 # Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2016

Abstract The environmental background to the Neolithic occupation of the Rio Sizandro, western Portugal is elucidated via the ostracod assemblages and sedimentology of a welldated 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.

Keywords Holocene . Neolithic . Portugal . Ostracoda . Sedimentary environments . Geoarchaeology

* Alan R. Lord [email protected] Maria Cristina Cabral [email protected] Rainer Dambeck [email protected] Michael Kunst [email protected] 1

Departamento de Geologia and Instituto Dom Luiz (IDL), Faculdade de Ciências, Universidade de Lisboa, Campo Grande, C6, 4, Lisboa 1749-016, Portugal

2

Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, Frankfurt am Main 60325, Germany

3

Institut für Physische Geographie, Johann Wolfgang Goethe-Universität, Altenhöferallee 1, Frankfurt am Main 60438, Germany

4

Abteilung Madrid, Deutsches Archäologisches Institut, Serrano 159, Madrid 28002, Spain

Introduction We continue our analysis of ostracod evidence for environmental conditions during the Neolithic for Rio Sizandro, western Portugal, commenced in Lord et al. (2011). The introduction to that work continues to be relevant here, but the application of ostracods in archaeological contexts has since developed strongly (Mazzini et al. 2015 and authors therein). The work reported here arises from a long-term project by the German Archaeological Institute (Madrid Department) to investigate the settlement history of western Portugal especially of the Rio Sizandro valley (Project ‘Sizandro–Alcabrichel: Two Neolithic settlement sites in comparison’, Kunst and Trindade 1990; Kunst and Lutz 2008; Dambeck et al. 2010a; Lord et al. 2011, pp. 216–218; Dambeck et al. 2015). Numerous boreholes have been drilled along the length of Rio Sizandro (Fig. 1), and in Lord et al. (2011), we reported on two sites near the village of Benfica where ostracod evidence indicated fresh to brackish water conditions. To continue the work we have now analysed a core (COU_14; Figs. 2, 3) from the village of Coutada, midway between Benfica and the modern coastline, where the ostracods reflect greater marine influences. The sedimentary record of the investigated core encompasses the time period of the entire early Holocene up to mid-Holocene times (c. 10,000–5000 cal BP) and therefore examines in more detail the shift from an open estuary into a coastal lagoon. This is particularly important for palaeoenvironmental reconstruction of the Sizandro valley, because in previous studies (Dambeck et al. 2010a), only the period between 6700 – 6200 cal BP has been detected with temporal resolution, so that the valley development during earlier phases of the Holocene lacks sufficient time control as yet. Since the beginning of the Postglacial, the lower Sizandro valley as other coastal lowlands was affected by eustatic sea level rise (e.g. Cearreta et al. 2003; Cabral et al. 2006; Dambeck et al.

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Fig. 1 Rio Sizandro, western Portugal, with locations of Coutada (this paper) and Benfica (Lord et al. 2011) and postulated environmental conditions in the river valley (modified from Lord et al. 2011: Fig. 1)

2010a; Cabral et al. 2011a). The general trend of Lateglacial and Holocene sea level rise along the western Portuguese coast is displayed in the model curve for the Tagus valley (Vis et al. 2008). The rapid rise of sea level between 20,000 cal BP and 6500 cal BP was interrupted by the climate cooling during the Younger Dryas (c. 12,000 cal BP) and at c. 9000 cal BP (Preboreal). From about 7000 cal BP the deceleration of the rise of sea level promoted the formation of sandy barriers (Alday et al. 2006; Trog et al. 2013, 2015). In consequence, the open estuaries were separated from the sea and coastal lagoons started to develop. As in other estuarine environments of Portugal the closure of the barrier forced the rapid aggradation of the lagoonal environments on the valley floor by silting, when the river floodplains became infilled with terrestrial sediment delivered via the rivers from the upstream parts of the catchments (e.g. Cearreta et al. 2003; Cabral et al. 2006). Sediment production was triggered by external (e.g. climate) and internal factors (e.g. lateral erosion of the river channel, autogenic controls in the river system, but also deforestation and agricultural land use). Human cultures populated the Sizandro valley from prehistoric times. Paleolithic and Mesolithic sites are documented in the coastal area west of Torres Vedras (Zilhão 1984; Zilhão et al. 1987; Kunst and Trindade 1990; Araújo 1994; Zilhão 1997, vol. 2, 603–646, 787–826; Araújo et al. 2014). Despite these early traces of prehistoric humans it is likely that

anthropogenic influence on sediment production only became sensitive in later periods of the Holocene. According to Kunst and Trindade (1990), the Sizandro drainage basin has been continuously settled since the Copper Age (c. 4800–3750 cal BP). However, more recent research has provided evidence for Neolithic agriculture (Kalis and Stobbe, in Dambeck et al. 2010a; Dambeck et al. 2015). Thus, it is possible that sediment supply into the floodplain and silting of the lagoon was already affected moderately by humans in Neolithic times.

Material and methods (RD, ARL) Over 60 sites were cored during the project (Fig. 1; Dambeck et al. 2010a, b); however, most of the cored sediment is no longer available. The sediments were subsequently analysed for sedimentology (grain size, organic matter, carbonate content, electrical conductivity, etc.) and palaeoenvironmental reconstruction (pollen, macro plant-residues, molluscs and microfossils - for details see Dambeck et al. 2010a). Material studied here came from borehole COU_14 (39°04′49.7′′N, 009°21′33.7′′E, height c. 10 m above sea level, drilled September 2008. Figs. 1, 2, 3). Twenty nine samples from between 10.00 and 27.90 m below the present surface (mbs) were analysed for ostracods, of which

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Fig. 2 East to west to south-west transect of boreholes across the Rio Sizandro floodplain near Coutada, with location of borehole COU_14 in the axis of the Neolithic valley. Vertical exaggeration x 12.6

the top two samples and bottom five samples were barren. Good macrofossil remains occur at some levels: gastropods, lamellibranchs; terrestrial wood, seeds and fruit. The terrestrial botanical remains provided eight 14C –AMS dates (Table 2, Figs. 2, 3). Site COU_14 is located c. 20 m east of the modern river channel and c. 350 m southeast of Coutada village (Fig. 4). In this area, the lithology of the Sizandro alluvial plain was studied by boreholes along a 550 m long cross-section using percussion drilling (Fig. 2). In total 15 boreholes were drilled usually at lateral distances of 30 m to study the sedimentary architecture of the subsoil. The drilling depths of the individual boreholes depended on the topography of the subsurface and varied from 4 mbs on the valley slopes up to 28 mbs in the central parts of the valley floodplain (see profiles COU_13, COU_14).

The untreated sediment samples studied here for micropalaeontology ranged in weight from 87 to 250 g, were dried before treatment with sodium hexametaphosphate and water, and washed over a 63-μm sieve. The >63-μm sediment fractions were either completely picked for ostracods or statistically picked (random splits fully picked so relative proportions reflect the entire residue) and therefore the relative proportions of species between samples shown in Fig. 5 are comparable. Appendix 1 lists the ostracods taxonomically and Appendix 2 summarises their environmental preferences. See Fig. 5 for the distribution pattern and abundances of ostracods. Common and environmentally diagnostic ostracods are illustrated in Figs. 6, 7, 8. All material is deposited in Sektion Mikropaläontologie I, Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main.

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Fig. 3 Lithostratigraphy and 14C dates for COU_14

Sedimentology (RD) The valley floor is locally deeply cut by incising fluvial erosion into the regolithic marls derived from the Upper Jurassic bedrock (Fig. 2). The dissection of the subsoil resulted from the increase of the rivers gradient when the relative mean sea level as the erosional base of the Rio Sizandro temporarily

stood at between −120 and 140 m below present sea level (mbpsl) during glacial times (Dias et al. 2000; Vis et al. 2008). While sea level related to climate change was the major controlling factor, other external and internal influences on river velocity and rate of incision are difficult to resolve. Rainfall patterns are not known and tectonic uplift from local salt diapirism may have affected river gradient and flow.

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Boreholes COU_13 and COU_14 penetrated 28 m of Holocene sediments, before drilling was interrupted for technical reasons without having reached the top of the underlying Jurassic marlstone. However, in profile COU_14 weathered regolithic basement material was found from below 27.90 mbs, suggesting that bedrock was reached approximately. In both boreholes the recovered strata set is represented by a series of clastic sediments ranging in grain size from sand (partially with subordinate gravel) to silt and clay that accumulated under varying sedimentary conditions (fluvial, estuarine, lacustrine, alluvial, colluvial; Fig. 2, Table 1). In general terms, the sediments recovered from the Coutada cross-section can be subdivided into five lithological units, described as lithozones (LZ), each representing a defined phase of landscape development (Tab. 1, Fig. 2). LZ-I is the Jurassic bedrock into which the valley has been cut, and the overlying lithozones (LZ-II to LZ-IV) can be differentiated by the character of the sediments (grain size, colour, content of faunal or floral remains): & & & & &

Jurassic bedrock: partly reworked at top (weathered regolith: LZ-I); Fluvial gravel and sand (pre-Holocene river terrace: LZ-II); Estuarine brackish to lacustrine lagoonal infill (LZ-IIIa); Older alluvial infill (not differentiated: LZ-IIIb); Colluvial deposits and younger alluvial infill (not differentiated: LZ-IV).

All along the Coutada cross-section the upper sediments deposited on the floodplain are composed of younger alluvial fill

(LZ-IV) that was deposited from the Rio Sizandro in the second half of the Holocene. Depending on the distance from the floodplain the thickness of the alluvium decreases and interfingers in some positions on the slopes with colluvial layers (LZ-IV). This is usually underlain by older alluvial infill of LZ-IIIb. The widespread occurrence of gravelly sands is striking and they can be recognised at varying depths and thicknesses in the profiles COU_2 to COU_12 (Fig. 2). The geomorphological situation reflects sediments that have developed at the foot of the lower slope in areas outside the central depth line (LZ-II). The coarse-grained texture of the fluvial transported material suggests an origin as sandy-gravelly terrace sediments that may have been deposited in an early period of the Last Glacial (Weichsel/Würm). The lack of overlap with marine and lacustrine sediments in the holes COU_2 to COU_10 shows that distal positions of the buried river terrace were not affected by early Holocene sea level rise, while the marinebrackish sedimentation influenced only the proximal margins of the terrace (see profiles COU_11, COU_12). This is also indicated by the preservation of a fossil soil profile that could be detected in the profile of COU_9 at a depth below 7.60 mbs; a possible soil is also present in COU_12. The geographical situation in more highly elevated positions outside the floodplain ensured terrestrial soil formation of a welldeveloped luvisol that is preserved under a cover of older alluvial fill (LZ-IIIb). Radiocarbon dating of the COU_9 soil (humic material) provided an age of 4965-4865 1σ cal BP and therefore dates the soil in the first half of the Holocene up to the Copper Age. This is therefore the Copper Age land

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Fig. 5 Occurrence, relative abundances and salinity preferences of ostracods in COU_14

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surface, which could have been occupied by prehistoric settlers. The fossil soil formation is covered by up to 7.60 m of alluvial sediments (LZ-IIIb) in COU_9, whereas towards the valley centre in COU_11 and COU_12 this surface is overlain by the brackish marine and lacustrine sediments of LZ-IIIa. Thus, it can be stated that the sites on the lower slope were affected by the Rio Sizandro, after the valley floor had been aggraded to such an extent that periodically flooding could reach these topographic positions, whereas more elevated areas were not so affected. COU_14. At the base of the drilled sequence, a stratified series of fluvial sands with subordinate gravel and some thin intercalated silt layers (LZ-II; 27.90 to 25.00 mbs. Figs. 2, 3, Table 1) accumulated on top of the regolithic marls derived from the Upper Jurassic bedrock (LZ-I; >27.90 mbs). Upwards the sediments of LZ-IIIa (25.00 to 11.00 mbs) comprise alternating laminations of varying grain sizes, strongly dominated by silt and clay fractions. Moreover, above 24.70 mbs brackish-marine lamellibranchs (bivalve molluscs) occur sometimes in relatively high frequency in the sediments, while botanical remains (seeds, fruits) of terrestrial plants are also present. These fossil remains provide a clear criterion to distinguish the deposits from the subjacent layers which are marked by the absence of macrofaunal and floral remains. In particular, the occurrence of marine bivalves can be taken as evidence that the sedimentary conditions had shifted from fluvial to marine-influenced deposition (LZ-IIIa). Radiocarbon dating of organic macro remains (seeds, fruits of terrestrial plants) allowed good chronostratigraphical correlation of the strata with the early Holocene period between c. 9120 – 4825 cal. BP (Table 2). Formation of LZ-IIIa therefore can be clearly attributed to the marine transgression that entered the Sizandro Valley in the first half of the Holocene, whereas the overlying LZ-IIIb (11.00 to 0 mbs; <4825 cal. BP) reflects the silting up of the coastal lagoon in the subsequent periods.

Ostracod occurrence in COU_14 (MCC, ARL) The occurrence and relative abundances of ostracod species are presented in the range chart in Fig. 5, allowing five units to be recognised; these units are not strictly ostracod zones as Units V and I are barren, however, the units clearly reflect important environmental signals. Key taxa are illustrated in Figs 6, 7, 8. Unit I – samples 25.50–27.90 mbs Fluvial sand, no microfossils in residues except plant debris and seed cases. Unit II – samples 21.40–25.00 mbs samples 21.40– 25.00 mbs Fluvial/estuarine conditions. Brackish/ euryhaline ostracods common but low diversity,

with a few littoral marine ostracods probably transported. Marginal marine benthic foraminifera present, some/all transported. Unit III – samples 13.70–20.90 mbs Estuarine to marine environment – large estuary with close marine connection and salinity at or near normal marine (33 psu). Mixed ostracod assemblages with abundant brackish/euryhaline taxa, some of them typical of salt marshes, suggesting the presence of these habitats in the margins of the estuary. Marine ostracods very diverse with over 50 species present; however, only the most abundant (the first 7 species in Fig. 4, or perhaps 9 to L. pellucida), which are generally slightly euryhaline and frequently found in outer estuaries, are in-situ, the rest transported. Rare freshwater species reflect streams and springs adjacent to the river and are probably related to years/periods with increased rainfall. Marginal marine to marine benthic foraminifera with some planktonic forms. Unit IV – samples 11.80–13.00 mbs Restricted estuarine to brackish lagoonal environment. Low diversity brackish/euryhaline ostracod assemblages, largely represented by Cyprideis torosa, suggesting a very restricted environment, with very rare transported freshwater forms; no marine forms found. Some benthic foraminifera. The characteristic oligospecific ostracod assemblage represented almost only by C. torosa indicates the formation of a well developed sandy barrier which may have temporarily closed the estuary forming a lagoon. The total absence of some of the typical salt marsh species and the abundance decrease of Loxoconcha elliptica (compared with Unit III) suggest the disappearance/decline of the salt marshes, populated only by Leptocythere porcellanea, which is today the most abundant ostracod in the Portuguese salt-marshes from north to south (Cabral and Loureiro 2013). Unit V – samples 10.00–10.80 mbs Fluvial silts, no fossil remains found. The ostracods are very well represented through part of the borehole, between 25.00 and 11.80 mbs with at least 64 species, suggesting the importance of the river/estuary during the Holocene. Most of the species were marine (more than 52), but more than 39 are mainly represented by rare juvenile valves, frequently worn and surely transported inside the estuary by tidal currents and/or storms. The marine autochthonous species, with abundant and complete or almost complete populations of adults and juveniles, are littoral or sublittoral forms, some of them common in modern outer estuaries, such

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as Loxoconcha rhomboidea, Heterocythereis albomaculata, Aurila convexa, Pontocythere elongata. The most abundant species are the brackish Cyprideis torosa and Loxoconcha elliptica, both present in almost all of the levels containing ostracods, generally with abundant to very abundant populations (more than 50 individuals), with well preserved adults and juveniles, indicating that brackish conditions were present in the studied environment during most of the time. In many levels a mixture of well preserved and opaque or damaged shells, sometimes stained orange or filled with sediment is present, demonstrating the mixture of autochthonous and allochthonous microfossils of the same species, suggesting reworking and transport inside the estuary/lagoon. In one particular level (16.60 to 16.40 mbs), a very important transported/reworked assemblage of stained orange C. torosa is found. Throughout the borehole sequence, C. torosa ranged from almost smooth to punctate/slightly reticulated, always without nodes, testifying to water chemistry variations in the environment, probably in mesohaline salinity (e.g. Kilenyi 1972; Carbonel 1980; Gliozzi and Mazzini 1998), which has been verified in other Holocene Portuguese lagoons (Cabral et al. 2006; 2011a). The other brackish species (5) are generally typical of recent salt marshes. Fresh water species (5 species in total) are poorly represented in very rare levels, especially in one probably indicating a period of intense rainfall.

Fig. 6 LV left valve, RV right valve, C carapace, L length in mm, SMF Xe„ — catalogue numbers of Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany. Scale bars 100 μm. All samples from borehole Coutada 14 (COU_14), sample depths in centimeter. below surface. a Bythocythere bradyi Sars, 1926. RV, external, female, L 0.65; SMF Xe 23266; sample 1920–1930. b Sclerochilus sp. RV, external, ?female, L 0.72; SMF Xe 23267; sample 1580–1600. c Pontocythere elongata (Brady, 1868). LV, external, juvenile, L 0.70; SMF Xe 23268; sample 2310–2325. d Cyprideis torosa (Jones, 1850). LV, external, female, L 0.97; SMF Xe 23269; sample 2310–2325. e Cyprideis torosa (Jones, 1850). RV, external, male, L 0.98; SMF Xe 23270; sample 2310–2325. f Hemicytherura aff. defiorei Ruggieri, 1953. LV, external, male, L 0.36; SMF Xe 23271; sample 1410–1430. g Microcytherura fulva (Brady and Robertson, 1874). RV, external, juvenile, L 0.45; SMF Xe 23272; sample 1640–1660. h Pseudocytherura cf. calcarata (Seguenza, 1880). RV, external, juvenile, L 0.59; SMF Xe 23273; sample 1470–1490. i Semicytherura acuminata (G. W. Müller, 1894). LV, external, female, L 0.51; SMF Xe 23274; sample 1640–1660. j Semicytherura acuta (G. W. Müller, 1912). LV, external, male, L 0.43; SMF Xe 23275; sample 1410–1430. k (= j). Ornamentation detail. l Semicytherura acuticostata ventricosa (Sars, 1866). RV, external, female, L 0.45; SMF Xe 23276; sample 1410–1430. m Semicytherura arcachonensis Yassini, 1969. LV, external, male, L 0.60; SMF Xe 23277; sample 1470–1490. n Semicytherura robertsi Whittaker and Horne, 2009. RV, external, male, L 0.45; SMF Xe 23278; sample 1430–1450. o Semicytherura robertsi Whittaker and Horne, 2009. LV, external, female, L 0.41; SMF Xe 23279; sample 1640–1660. p Semicytherura sella (Sars, 1866). LV, external, male, L 0.42; SMF Xe 23280; sample 1430–1450. q Semicytherura cf. stilifera Bonaduce, Ciampo and Masoli, 1976. RV, external, female, L 0.45; SMF Xe 23281; sample 1640–1660. r Semicytherura sulcata (G. W. Müller, 1894). RV, external, female, L 0.57; SMF Xe 23282; sample 1640–1660

Correlation between Coutada and Benfica

Discussion

The sequence of ostracod assemblages from Benfica, located about 4 km eastwards up the Rio Sizandro from Coutada and about 8 km from the modern coast, was described by Lord et al. (2011). Comparison between the Coutada and Benfica boreholes is only possible in a small part of the cores, since in Benfica, we only have ostracod data corresponding to 4.5 m of the core, for the time interval from ∼6500 to 6000 yrs cal. BP. In Benfica, Lord et al. (2011) defined two ostracod units: (i) a basal one corresponding to a brackish environment with a well represented autochthonous assemblage of Loxoconcha elliptica, Cyprideis torosa and Leptocythere porcellanea, associated with rare transported specimens of freshwater ostracods; (ii) an upper unit with less abundant L. elliptica, C. torosa and freshwater forms, most of them with opaque shells indicating transport, and absence of L. porcellanea, all these data indicating a drop of salinity with strong bottom current activity. The boundary between the two units is considered to fall at around 6300 yrs cal BP. In Coutada, located around 4.5 km from the mouth of the Sizandro River, this boundary must be slightly older, corresponding to the transition from Unit III to Unit IV, dated from around 6450 yrs cal BP, when the coastal detrital barrier was formed.

Quaternary climate had a noticeable effect on shaping the landscape of the lower Sizandro Valley. At the Last Glacial Maximum (Weichselian/Würm) (c. 20,000 cal BP) sea level was about 120 to 140 m lower than present (Dias et al. 2000; Vis et al. 2008). The drop in the erosional base caused an increased vertical erosion and valley incision because the rivers had to adjust to this level. During the Quaternary the Rio Sizandro has cut deeply into the subjacent Jurassic bed rock (LZ-1; Fig. 2). This is the result of increased vertical erosion during different periods of the Pleistocene. At the beginning of the Lateglacial, the general trend of climate warming led to eustatic sea level rise that decelerated at c. 6500 cal BP before the present sea level was reached at c. 4500 cal BP. Short-term oscillations of the sea level that can be recognised in the sea-level curve during short cold fluctuations (e.g. Younger Dryas; see Vis et al. 2008) could not been identified in the Sizandro valley. However, postglacial marine transgression forced rapid aggrading of the valley floor, since the floodplain became infilled with terrestrial sediments delivered by the Rio Sizandro and its tributaries to the estuary. At Coutada this phase is documented in the sediments of LZ-IIIa, which is characterised

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ƒFig. 7

a Semicytherura sulcata (G. W. Müller, 1894). LV, external,? male, L 0.56; SMF Xe 23283; sample 1430–1450. b Cytheropteron dorsocostatum Whatley and Masson, 1980. RV, external, female, L 0.40; SMF Xe 23284; sample 1410–1430. c Eucythere prava Brady and Robertson, 1869. RV, external, female, L 0.52; SMF Xe 232845; sample 1580–1600. d Aurila convexa (Baird, 1850). LV, external, female, L 0.79; SMF Xe 23286; sample 2310–2325. e Aurila woutersi Horne, 1986. LV, external, ?male, L 0.79; SMF Xe 23287; sample 1540– 1560. f Caudites calceolatus (O. G. Costa, 1853). LV, external, juvenile, L 0.64; SMF Xe 23288; sample 1410–1430. g Heterocythereis albomaculata (Baird, 1838). LV, external, female, L 0.80; SMF Xe 23289; sample 1800–1830. h Heterocythereis albomaculata (Baird, 1838). RV, external, male, L 0.84; SMF Xe 23290; sample 1800–1830. i (= h). Detail of ornamentation. j Thaerocythere hoptonensis (Brady, Crosskey and Robertson, 1874). RV, external, L 0.73; SMF Xe 23291; sample 1430–1450. k Urocythereis britannica Athersuch, 1977. LV, external, female, L 0.84; SMF Xe 23292; sample 1640–1660. l Callistocythere badia (Norman, 1862). C, left view, female, L 0.50; SMF Xe 23293; sample 1800–1830. m Callistocythere curryi Horne, Lord, Robinson and Whittaker, 1990. LV, external, female, L 0.49; SMF Xe 23294; sample 1640–1660. n Callistocythere murrayi Whittaker, 1978. RV, external, female, L 0.45; SMF Xe 23295; sample 2070–2090. o Leptocythere castanea (Sars, 1866). LV, external, female, L 0.62; SMF Xe 23296; sample 1410–1430. p Leptocythere fabaeformis (G. W. Müller, 1894). LV, external, male, L 0.75; SMF Xe 23297; sample 2070–2090. q (= p). Detail of ornamentation. r Leptocythere fabaeformis (G. W. Müller, 1894). RV, external, female, L 0.70; SMF Xe 23298; sample 2310–2325

by the deposition of silt and clay from c. 10,130 cal BP onwards. Variations in the occurrence and relative abundance of the analysed ostracod species display the shift from a freshwater influenced estuarine environment into a nearly full marine large estuary with close connection to the ocean and high salinity, that turned into a brackish lagoon after c. 6500 cal BP, due to the formation of the closing coastal barrier. This development is in good concordance to other lowland regions located elsewhere along the Portuguese coast, though the formation and temporary or permanent closure of the barrier varies from one place to another (see Freitas et al. 2003). In addition, the results obtained in Coutada allow better understanding of the landscape development in the early Holocene. With the formation of the barrier marine conditions retreated to the modern coast line and the coastal lagoon subsequently silted up. It is likely that sediment supply to some extent resulted from self-organised fluvial processes in the river system (e.g. retrogressive erosion by upstream migration of the erosional point, due to the aggrading of the valley floor and also lateral erosion in meandering river channels). In addition, it must be assumed that fluvial accumulation was mainly triggered by the effects of land use, because the Sizandro Valley has been inhabited since the Neolithic period (Dambeck et al. 2010a,b). Temporal resolution of soil erosion or man-made-phases of enhanced fluvial sedimentation is still open, although Herrmann (2010; unpublished Diplomarbeit, Institut für Geographie, Universität Leipzig) discovered that the

carbonate rocks from the Upper Jurassic in particular are strongly affected by soil erosion, which suggests that these areas may have been used for agriculture possibly since prehistoric times. However, Sizandro valley was occupied during the Neolithic, and it can be assumed as in other regions that soil erosion caused by Neolithic agriculture and therefore also sediment supply into the floodplain occurred on a relatively small scale. The subsequent cultural periods of the Copper Age, the Iron Age and Roman Times, and also the Dark Age and more recent times are well documented in the Rio Sizandro valley (Kunst and Trindade 1990). Due to the increasing population over time, with deforestation as well as expansion of arable fields, it is likely that soil erosion and related sediment supply intensified during the second half of the Holocene. This statement is supported by the fact that the younger sedimentary infill (LZ-IIIb, LZ-IV) has been deposited within the last 5000 years (see Fig. 2).

Conclusions Ostracod assemblages from borehole COU_14 demonstrated the existence of the following environmental conditions: & & & & &

Unit I, two dates 10,130–9780 and 9120–8980 cal BP, fluvial sands without ostracods or foraminifera; Unit II, one date 7930–7800 cal BP, fluvial to estuarine; Unit III, four dates 6630–6320 to 7410–7270 cal BP, estuarine to marine; Unit IV, one date 4960–4825 cal BP, restricted estuarine to brackish; Unit V, <5000 BP, fluvial silts without ostracods or foraminifera.

The late Pleistocene and Holocene landscape development of the Sizandro valley can be described as follows: & & & &

Rio Sizandro incised Jurassic bedrock by fluvial erosion (late Pleistocene); With the beginning of the Lateglacial eustatic sea level rise caused marine transgression in the coastal lowland; Estuary became invaded by the sea, c. 9780 cal BP; Previously terrestrial environments in the floodplain were inundated (ostracod Unit I). The deposits correlated to LZ-II demonstrate the existence of a fluvial dominated phase (formation of a river terrace) that preceded the marine transgression. This is in good agreement with the situation reconstructed at Benfica, where brackish conditions succeeded overbank deposition when the estuarine environment expanded into the

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ƒFig. 8

a Leptocythere pellucida (Baird, 1850). LV, external, female, L 0.59; SMF Xe 23299; sample 2070–2090. b Leptocythere pellucida (Baird, 1850). C, left view, external, male, L 0.65; SMF Xe 23300; sample 1920–1930. c Leptocythere porcellanea (Brady, 1869). LV, external, female, L 0.48; SMF Xe 23301; sample 1430–1450. d Loxoconcha elliptica Brady, 1868. LV, external, male, L 0.65; SMF Xe 23302 sample 2310–2325. e Loxoconcha rhomboidea (Fischer, 1855). LV, external, male, L 0.63; SMF Xe 23303; sample 1800–1830. f Sagmatocythere napoliana (Puri, 1963). RV, external, male, L 0.50; SMF Xe 23304; sample 1800–1830. g Neocytherideis subulata (Brady, 1868). RV, external, L 0.72; SMF Xe 23305; sample 1410–1430. h Cytherois fischeri (Sars, 1866). RV, external, female, L 0.54; SMF Xe 23306; sample 1800–1830. i Basslerites teres (Brady, 1869). C, left view, external, L 0.50; SMF Xe 23307; sample 2070–2090. j Xestoleberis rubens Whittaker, 1978. RV, external, female, L 0.45; SMF Xe 23308; sample 1800–1830. k ‘Bairdia’ subcircinata (Brady and Norman, 1869). LV, external, juvenile, L 0.55; SMF Xe 23309; sample 1640–1660. l Neonesidea sp. 2 (Bonaduce, Ciampo and Masoli, 1976). RV, external, juvenile, L 0.81; SMF Xe 23310; sample 1540–1560. m Cypridopsis vidua (O. F. Müller, 1776). C, right view, female, L 0.68; SMF Xe 23311; sample 1800–1830. n Ilyocypris bradyi Sars, 1890. RV, external, female, L 0.94; SMF Xe 23312; sample 1800–1830. o Ilyocypris bradyi Sars, 1890. LV, external, female, L 0.87; SMF Xe 23313; sample 1800–1830. p Ilyocypris inermis Kaufmann, 1900. RV, external, female, L 0.91; SMF Xe 23314; sample 1800–1830. q Ilyocypris inermis Kaufmann, 1900. LV, internal, female, L 0.94; SMF Xe 23315; sample 1800–1830. r Limnocythere inopinata (Baird, 1843). RV, external, female, L 0.60; SMF Xe 23316; sample 1800–1830

Table 1

& & & & & & & &

investigated areas at c. 6700 cal BP (Dambeck et al. 2010a; Lord et al. 2011); The continued rise of the sea level resulted in a fully marine environment at Coutada (parts of LZ-IIIa/ostracod Units II, III; c. 7930–7410 cal BP); With the deceleration of the rate of sea level rise a sandy coastal barrier formed (c. 6250 cal BP); Development of the barrier with possible closure (temporary) caused the establishment of a coastal lagoon (upper part of LZ IIIa/ostracod unit IV); Silting of the lagoon, due to sediment supply from upstream parts of the river catchment (from c. 6200 cal BP onwards); Rapid aggrading of the valley floor by silting (LZ-IIIb/ ostracod Unit V); Silting of valley ‘supported’ by intensified land use that probably started in the Copper Age. Deforestation and agriculture resulted in an increase in sediment supply and accumulation of the sediments of the ‘older alluvial infill’ (LZ-IIIb). Modern land use ‘produced’ the sediments of the ‘younger alluvial infill’ and colluvium (LZ-IV);

Overview of core data from the Rio Sizandro Coutada borehole transect

Depth (mbs)

Lithozone (LZ)

Ostracod zone

Sedimentary environment

Varying depth (see profiles of COU_3–COU_12)

LZ-VI

–

Fluvial/terrace

Varying depth LZ-V (see profiles of COU_ 1–COU_7, COU_ 10–COU_11, COU_15) 0–11.00 LZ-IIIb

–

Colluvial

Indet.

V

Fluvial/alluvial

Indet.

11.00–13.80

IV

Estuarine/ lacustrine

4825–6630

13.80–21.00

III

Estuarine/marine

7410–7850

21.00–25.00

II

Fluvial/estuarine

7930–7800

25.00–27.90

I

Fluvial sand c. 8980–10,130 (early Holocene terrestrial phase)

27.90+

LZ-IIIa

LZ-I

regolith Indet. (weathered Upper Jurassic bedrock)

Age (kyr cal BP)

indet. (pre-Quaternary)

Common features Colour: yellowish to reddish brown. Grain size: coarse and medium sand with subordinate gravel. Other: top parts affected by formation of a luvisol that developed in early Holocene times (top soil horizon has been dated at c. 5000 cal BP). Colour: greyish brown. Grain size: loamy sand to sandy or clayey loam. Other: stones, residues of bricks. Colour: blackish grey to dark grey at the base (below 8.30 mbs), dark grey to greyish brown or reddish brown in the superimposed parts of the profile (above 8.30 mbs). Grain size: clay in the basal parts of the profile (below 8.30 mbs), alternation of clayey silt and fine to coarse sand and loamy sand or sandy or clayey loam in the superimposed parts of the profile (above 8.30 mbs). Other: gastropods, plant remains (indet.). Colour: blackish grey. Grain size: sandy clay to clay. Other: few lamellibranch shells. Colour: grey to blackish grey. Grain size: coarse to fine sand or loamy sand alternating with sandy loam and silty clay to clay. Other: lamellibranch shells, few plant remains (seeds, fruits). Colour: dark grey to blackish grey. Grain size: coarse to fine sand or loamy sand alternating with silty clay and clay. Other: frequent lamellibranchs, few wood fragments. Colour: dark grey to blackish grey. Grain size: fine to coarse sand or loamy sand, subordinate laminated layers of clayey silt or silty clay. Other: plant remains (e.g. roots), plant residues (e.g. seeds and fruits). Colour: mottled, light greenish grey to reddish brown. Grain size: predominance of clay silt. Other: absence of organic debris or macro remains.

Palaeobio Palaeoenv Table 2

14

C-AMS dates obtained from terrestrial plant material

COU_14

BP

1σ cal BC

1σ cal BP

12.70 mbs 13.90 14.90 16.60 19.30 22.60 26.50 27.70 COU_9

4290 ± 551 5670 ± 1201 6560 ± 1401 6470 ± 402 6400 ± 402 7010 ± 402 8845 ± 401 8085 ± 451 BP

12.70 mbs

4360 ± 401

3010–2,8751 4680–4,3701 5630–5,3751 5480–5,3802 5460–5,3202 5980–5,8502 8180–7,8301 7170–7,0301 1σ cal BC 3015–2,9151

4960–4,8251 6630–6,3201 7580–7,3251 7430–7,3302 7410–7,2702 7930–7,8002 10,130–9,7801 9120–8,9801 1σ cal BP 4965–4,8651

Dating laboratories: 1 Leibniz-Labor, Kiel/Germany; 2 Beta Analytics, Miami/Florida, USA

&

&

&

During the early Holocene, the river terrace identified at the lower slope and likely to be of early Last Glacial (Weichselian/Würm) age was affected by soil formation of a luvisol; in later phases of the Holocene the soil profile was covered by younger alluvial and colluvial sediments (LZ-IIIb, LZ-IV); Radiocarbon dating of the soil of the luvisol profile provided a mid-Holocene age (4965–4865 cal BP, COU_9; Table 2) that fits well with the Copper Age; fossilized paleosoil in profile COU_9 represents the land surface of the Copper Age; A maximum width of c. 250 m can be assumed for the lagoon when sea level peaked at about 6500 cal BP.

Future Work: Study the living ostracod fauna in the lower estuary of Rio Sizandro, when estuarine (winter) and when lagoonal (summer) to help calibrate the present results (Fig. 9).

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.

Compliance with Ethical Standards Conflict of Interest Funding sources are all listed in the Acknowledgements. There are no other conflicts of interest.

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

Fig. 9 Mouth of modern Rio Sizandro at Foz do Sizandro to show winter/spring (left: open channel) and summer (right: closed lagoon) conditions (images Cabral 2014)

Palaeobio Palaeoenv

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

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 Cuneocytheridae Mandelstam, 1959 Genus Cuneocythere Lienenklaus, 1894 Cuneocythere semipunctata (Brady, 1868)

Family Limnocytheridae Klie, 1938 Limnocythere s. str. Brady, 1867 Limnocythere inopinata (Baird, 1845) - Fig. 8r

Family Cushmanideidae Puri, 1974 Genus Pontocythere Dubowsky, 1939 Pontocythere elongata (Brady, 1868) - Fig. 6c

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

Order Podocopida Sars, 1866 Suborder Cytherocopina Gründel, 1967 Superfamily Cytheroidea Baird, 1850

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

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

Palaeobio Palaeoenv 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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