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A New Pollen Sequence from Southern Iberia Suggesting Coastal Pleistocene Phytodiversity Hotspot

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A New Pollen Sequence from Southern Iberia Suggesting Coastal Pleistocene Phytodiversity Hotspot Review of Palaeobotany and Palynology 281 (2020) 104281 Contents lists available at ScienceDirect Review of Palaeobotany and Palynology journal homepage: www.elsevier.com/locate/revpalbo A new pollen sequence from southern Iberia suggesting coastal Pleistocene phytodiversity hotspot Juan Ochando a,⁎, José Antonio López-Sáez b, Sebastián Pérez-Díaz c, Julián Ramos-Fernández d, Manuel Munuera e, Santiago Fernández a, Federico B. Galacho-Jiménez f, Reyes Luelmo-Lautenschlaeger b,h,JoséS.Carrióna,g a Department of Plant Biology (Botany Area), Faculty of Biology, University of Murcia, Campus de Espinardo, 30100 Murcia, Spain b Environmental Archaeology Research Group, Institute of History, CSIC, CCHS, Albasanz 26-28, 28037 Madrid, Spain c Grupo de Investigación Pasado y Presente Geográfico del Relieve, la Vegetación y el Paisaje (GEOREVE), Departamento de Geografía, Urbanismo y Ordenación del Territorio, Universidad de Cantabria, E.T.S. de Ingenieros de Caminos, Canales y Puertos. Avenida de los Castros, 44, 39005 Santander, Spain d Parque Arqueológico de la Araña, 29004 Málaga, Spain e Department of Agricultural Engineering, Polytechnic University of Cartagena, 30203 Cartagena, Spain f Department of Geography, University of Málaga, Avda. de Cervantes 2, 29071 Málaga, Spain g Evolutionary Studies Institute, University of Witwatersrand, Witwatersrand, South Africa h Department of Geography, University Autonoma of Madrid, Calle Francisco Tomás y Valiente, 1, 28049 Madrid, Spain article info abstract Article history: This paper presents a palynological study of the archeological layers from the Neanderthal site Abrigo 3 del Received 6 March 2020 Complejo del Humo, in southern Spain (Málaga), with the aim of reconstructing the environmental conditions Received in revised form 14 July 2020 in the vicinity of this hominin site. The Upper Pleistocene vegetation and its variability are described, revealing Accepted 18 July 2020 a high diversity of thermophilous plant taxa throughout the cold dry phases, together with a long-term persis- Available online 22 July 2020 tence of woody taxa, including Mediterranean, mesophytes, xerothermics and conifers. With the pollen records of Maytenus senegalensis as an outstanding finding, this study demonstrates the co-existence of temperate, Med- Keywords: Palynology iterranean and Ibero-Maghrebian angiosperms on the southern coastal plains of the Iberian Pleistocene where Paleoecology Neanderthals survived for a long time. It is therefore clear that Neanderthals and early Upper Paleolithic modern Neanderthals humans lived in a litoral refugium, which was a propitious environment for maintaining a high biodiversity, in- Glacial refugia cluding potentially edible plant species. Besides, this coastal refugium offers broad possibilities for hunting, and interpopulational relationships through coastal platforms. © 2020 Published by Elsevier B.V. 1. Introduction data for human evolution studies have been provided by archeological palynology (Bonnefille, 2010; Burjachs and Julià, 1994; Burjachs et al., A vast majority of paleopalynological investigations in continental 2003; Carrión et al., 2003b, 2008, 2011, 2018, 2019a, 2019b, 2019c; context are based on lakes and bogs sequences (Bennett and Willis, González-Sampériz et al., 2010; Lechterbeck and Jensen, 2020; 2001; Birks, 2005; Ritchie, 1995). However, in Western Europe only a Ochando et al., 2019; Scott et al., 2003, 2019; Val-Peón et al., 2019), few of them extend to the Pleistocene, or they are generally not located and they represent a background which should not be undervalued. in the vicinity of hominin and/or archeological sites. This means, there- In connection with the former, the persistence of glacial refugia for fore, that in the case of Neanderthals, the reconstruction of their habitats temperate trees during the Pleistocene is important to note. Paleoeco- must be based on the palynological reconstruction of the sediments de- logical and phylogeographic studies have highlighted how important posited in archeological caves and rockshelters, in spite of the method- for biodiversity patterns in middle and high latitudes the existence of ological limitations broadly analyzed (Bottema, 1975; Carrión, 2002; plant refugia has been (Bhagwat and Willis, 2008; Taberlet et al., Coles and Gilbertson, 1994; Carrión et al., 1999, 2009; Davis, 1990; 1998; Willis and Whittaker, 2000). Refugia are also essential for species Fiacconi and Hunt, 2015, 2017; Hunt and Fiacconi, 2018; Navarro diversity and the level of endemism (Jansson, 2003). Furthermore, gla- et al., 2000, 2001; Sánchez-Goñi, 1993; Sánchez-Goñi et al., 2005). Re- ciations have built historical patterns traces still perceptible that par- gardless of the difficulties described by these authors, relevant habitat tially explain the distribution and richness of forest formations in Europe and North America (Svenning and Skov, 2007). Indeed, those fl ⁎ Corresponding author. Pleistocene refugia seem to have had a powerful in uence on current E-mail address: [email protected] (J. Ochando). genetic diversity of temperate forests in Europe (Hampe and Petit, https://doi.org/10.1016/j.revpalbo.2020.104281 0034-6667/© 2020 Published by Elsevier B.V. 2 J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281 2005; Petit et al., 2003). The combination of geomorphological, paleocli- 4. Sampling and chronological framework matic and paleobotanical characteristics of the Iberian Peninsula sug- gest it played a strategic role in the survival of tree taxa during the A total of 26 palynological samples were collected in Abrigo 3 Quaternary (Carrión and Leroy, 2010; Leroy and Arpe, 2007; Ochando (Figs. 2, 3), where a complete sedimentary column includes 26 levels. et al., 2019; Postigo-Mijarra et al., 2008, 2010), with coastal plains The uppermost samples come from the Upper Paleolithic levels 15 being of crucial importance (Carrión et al., 2003b, 2008, 2018; Yll and (samples 1 and 2) and 16 (samples 3 to 7). No pollen samples were Carrión, 2003; Ochando et al., 2019). This is the case with the taken from the overlying levels 14 and 13, corresponding to the same archeopalynological analysis presented here in Abrigo 3 del Complejo cultural period, because they were above the stratigraphic cut, although del Humo. a TL dating of level 13 was >19 ka. Level 15 is dated by TL at >24 ka (Ramos Fernández et al., 2012). The rest of the samples, up to the base 2. Physical setting and archeological remarks of the sequence, are all assigned to stratigraphic levels of the Middle Pa- leolithic: 8 and 9 (level 17), 10 (level 18), 11 (level 19), 12 (level 20), 13 Abrigo 3 del Humo (4° 20′ 42″ W, 36° 42′ 52″ N, 5 m a.s.l.) is an (level 21), 14 and 15 (level 22), 16 (level 23, dated by TL at 35 ka), 17 to archeological site lying in the Complejo del Humo (Cuevas de la 22 (level 24), 23 (level 25), and finally 24 to 26 (level 26) (Fig. 4) Araña), east from Málaga Bay in southern Spain (Fig. 1). The main en- (Ramos Fernández et al., 2012). trance, southernly, is in the lowermost paleocliff of the territory. Abrigo 3 del Complejo del Humo is a sea balma filled with about 7 m of sedi- 5. Palynological methods ments and attributed mainly to the Upper Pleistocene (Ferre et al., 2004). The Humo cave, which gives name to the complex, opens to The laboratory treatment was carried out in the Archaeobiology Lab- Abrigo 3 and is composed of a series of galleries whose dimensions oratory (CSIC) using the method proposed by Faegri and Iversen (1989) are not too large but difficult to transit. Its origin was due to the rainwa- with densimetric separation of microfossils (Goeury and de Beaulieu, ter dissolution of Jurassic limestones and karstification processes, com- 1979). The pollen morphotypes have been identified according to bined with the detachment of large blocks from the site's roof (Ortega Faegri and Iversen (1989), Moore et al. (1991), Reille (1992, 1995) Ruiz and Ramos Fernández, 2015). and Valdés et al. (1987). In the distinction of the Plantago, references Fieldwork in this locality was first carried out by Miguel Such in from Ubera et al. (1988) were used, Renault-Miskovsky et al. (1976) Hoyo de la Mina Cave in 1917 (Such, 1920). In 1942 José Sánchez was taken as guidance for Oleaceae, and Burgaz et al. (1994) for Pistacia Montes excavated the Humo Cave access, finding tools that he attrib- lentiscus. The Maytenus senegalensis morphotype was established ac- uted to the Aurignacian, and other materials possibly of Mousterian as- cording to Perveen and Qaiser (2008). Non-pollen microfossils were ba- cription (Sánchez Montes, 1947). After him, Giménez Reyna prospected sically identified following van Geel (1986, 2001) and van Geel et al. the area, and assigned the archeological levels of Hoyo de la Mina to (1986, 1989). deep Upper Paleolithic (Giménez Reyna, 1946). Abrigo 3 del Complejo Whenever a valid sample has been given, the number of pollen del Humo was first excavated by F. Gálvez-Pacheco in 1965, who iden- grains counted or pollen sum (P.S.) has exceeded 200 grains of terres- tified in a stratigraphic profile Levallois Mousterian lithics associated trial plants, also housing a minimum taxonomical diversity of 20 types with abundant seashells and few bones remains. In the upper sections, (López-Sáez et al., 2003; Sánchez-Goñi, 1994). In the calculation of the this author found Aurignacian lithics (Gálvez-Pacheco, 1975). By these percentages, hydro-hygrophytes taxa and non-pollen palynomorphs dates Julián Ramos took over the fieldwork. He prospected the area (NPPs), – sometimes considered local and others extra-local – have and organized the chaotic and sometimes imprecise nomenclature of been excluded from the P.S. because they are usually overrepresented some deposits, such as Hoyo de la Mina and Cueva de Navarro 4, (López-Sáez et al., 1998, 2000, 2003; Wright and Patten, 1963). In addi- which was assigned to La Cala del Moral (Rincón de la Victoria), when tion, Cichorioideae and Aster type have been also excluded from the P.S. in fact they belonged to the neighborhood of La Araña (Málaga).
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