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

Review of Palaeobotany and Palynology 281 (2020) 104281

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Review of Palaeobotany and Palynology

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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,

2005; Petit et al., 2003). The combination of geomorphological, paleocli- 4. Sampling and chronological framework matic and paleobotanical characteristics of the Iberian Peninsula suggest 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). The due to their anthropozoogenic nature (Bottema, 1975; Burjachs et al., Araña Caves were divided into 7 karst systems and the rock shelters 2003; Carrión, 1992b; López-Sáez et al., 2003). The relative value of present were numbered. In 1981 and 1983, Julián Ramos carried out the excluded palynomorphs has been calculated with respect to the the first excavation campaigns in Abrigo 3 del Complejo del Humo P.S. Figs. 5 and 6 show the pollen diagrams elaborated with TILIA and that confirmed the presence of the Middle Paleolithic in the deepest TGView software (Grimm, 1992, 2004). All samples, except for three levels 26 to 21 and Upper Paleolithic at the highest levels 15 to top. (10, 18 and 23), were polleniferous. A synthetic diagram was prepared Levels 17 to 20 show difficulties to be assigned to an archeological con- with the main taxa and ecological groups (Fig. 6). text. In a recent work, it has been proposed that the level 17 also contain Mousterian remains (Ramos Fernández et al., 2012). 6. Palynological results

3. Modern-day vegetation and climate The Middle Paleolithic levels 26-24 show predominance of herbaceous taxa (ca. 45–65%) such as Poaceae, Artemisia, Fabaceae, Liliaceae, Abrigo 3 del Complejo del Humo is located in an area of warm and Amaranthaceae, Cichorioideae, Aster and Anthemis. The tree component temperate oceanic bioclimate, in the lower limits of the European Med- accounts for 15–20%, with mainly pines (Pinus pinea type and Pinus iterranean Region next to the Ibero-Maghrebian-Atlantic Super Prov- sylvestris/nigra type) and to a lesser extent, deciduous and evergreen ince of Northern Africa (Fig. 1). The nearby meteorological station, in Quercus. Pollen frequencies of shrubs are noticeable (ca. 30–40%). Over- Málaga city, shows an annual average temperature of 18.4 °C and an an- all, the pollen spectra suggest heliophilic vegetation dominated by Arte- nual precipitation of 507 mm. The area is located in the lower misia, Poaceae and Amaranthaceae, with xerothermic influence such as thermomediterranean belt with upper dry ombroclimate. Biogeograp- suggested by Cistaceae, Erica, Lamiaceae, Myrtus, Olea europaea, hically, it belongs to the Baetic Province, Malacitano-Axarquiense sector Phillyrea and Pistacia lentiscus. These inferred arid conditions fit with (Montes de Málaga Unit) (Pérez Latorre et al., 2008; Rivas-Martínez, the low frequencies of alder (Alnus)(Figs. 5, 6). 1987; Rivas-Martínez et al., 1997, 2002). The territory is characterized The levels 23 and 22, within Middle Paleolithic context, continue to by strong human impact (agriculture, industrial exploitation and be dominated by herbaceous plants (ca. 40–45%) with slightly lower urban planning). The vegetation present in the study area is dominated amounts of anthropogenic pollen indicators (Cichorioideae, Aster, by Ibero-Maghrebian and Mediterranean scrub with scattered pines Anthemis). It is equally noteworthy the incidence of Amaranthaceae (Pinus halepensis), patchily altered by overgrazing, fires, and extreme and Artemisia. However, the arboreal elements are now more impor- weather events (heat waves, droughts or windstorms). tant, reaching ca. 25%, especially with Pinus and Quercus, higher Alnus J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281 3

Fig. 1. Location of Abrigo 3 del Complejo del Humo in southern Spain. The three top images have been redrawn from Google Earth.

and the presence of Abies (5–8%). The shrub layer keeps its values with and to a lesser extent Quercus, while a decline is observed in Alnus and respect to the previous levels, with the majority of the aforementioned Abies disappears. The scrub elements, however, fall significantly below taxa being present, with the first appearance of Ephedra fragilis and the 20%, and although most of them maintain some number, others disap- continuity of Viburnum.ThepresenceofGlomus and Pseudoschizaea pear (Ephedra fragilis or Myrtus), or keep presence only in a few samples circula is again noticeable (Figs. 5, 6). (Erica arborea, Maytenus, Viburnum). Glomus and Pseudoschizaea circula The most recent Middle Paleolithic levels 21-19 indicate a significant still appear suggesting the existence of erosive processes. In level 19 the change in the vegetation landscape. The herbaceous component con- parasithic fungus Diporotheca is present (Figs. 5, 6). tinues to be predominant with a maximum of 50% at level 21 and a min- The Middle Paleolithic (level 17) and the Upper Paleolithic (level imum of 27% in level 19. The tree cover is higher (ca. 30%) with Pinus, 16) show a substantial change in the pollen spectra, now with a 4 J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281

Fig. 2. Geological map of part of the Complejo del Humo where Abrigo 3 del Complejo del Humo appears.

codominance of arboreal (32–45%) and herbaceous (30–40%) there is a somewhat higher presence of anthropogenic and anth- vegetation, but above all, a signiﬁcant rise in scrub (28–37%). It is ropozoogenic elements such as Aster, Boraginaceae, Polygonum worth mentioning the reappearance of Abies and the increase of aviculare and Sordaria. Vegetation during this period suggests more Alnus. Pines are about 15–20%, and both deciduous and evergreen humid and warm conditions. Finally, the Upper Paleolithic, levels Quercus are important. The shrubs are mainly represented by Ephe- 16 and 15, show similar vegetation landscape with a progressive re- dra fragilis, Myrtus, Olea, Phillyrea, Viburnum and Pistacia lentiscus. gression of the thermophilic maquia and some increase of herbs. It is noticeable the presence of the arto (Maytenus senegalensis). Abies disappears, while pines and oaks continue to be important The herbaceous layer remains similar species composition, although (Figs. 5, 6).

Fig. 3. Stratigraﬁcal levels and proﬁle sampled for pollen analysis in Abrigo 3 del Complejo del Humo. Redrawn from Ramos Fernández et al. (2012). J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281 5

Fig. 4. Bayesian model containing the available chronometric data from Abrigo 3 del Complejo del Humo. Redrawn from Ramos Fernández et al. (2012).

7. Vegetation history and paleobotanical remarks of these species (Cabezudo and Pérez Latorre, 2001; Pérez Latorre et al., 2006, 2008) as well as and its pollen dispersion and representa- Although some uncertainties persist about the exact chronology of tiveness (López-Sáez et al., 2013; Broothaerts et al., 2018). This picture the deposits (Ferrer et al., 2005; Ortega Ruiz and Ramos Fernández, of Pleistocene vegetation has been widely discussed before in reference 2015; Ramos Fernández et al., 2012), the palynological analysis pre- to other territories (Carrión, 1992a, 1992b, 2001, 2002; Carrión et al., sented here allows visualizing the full-glacial vegetation of the Upper 1995a, 1999, 2000, 2003b, 2008, 2018, 2019a, 2019b, 2019c; Pleistocene in a territory on the southern coastline of Spain (Fig. 4). González-Ramón et al., 2012; López-Sáez et al., 2007; Ochando et al., While noticing some fluctuation between the aridity plant indicators, 2019; Zilhão et al., 2016). it bears emphasis that even during the most arid periods, a diversity of Oaks are fairly common in the southern Iberian Pleistocene such as thermophilous taxa occur continuously in the pollen sequence. These in Bajondillo, Cueva Perneras, Sima de las Palomas, Gorham´s and Van- results therefore confirm research so far in the Málaga province guard Caves (Carrión et al., 1995a, 2003b, 2008, 2018; López-Sáez (Bajondillo: López-Sáez et al., 2007; Cortés-Sánchez et al., 2007, 2008, et al., 2007). Nowadays, the nearest populations of Quercus coccifera, 2011, 2019), southwards in Gibraltar (Gorham and Vanguard sites: Q. ilex, Q. faginea and Q. suber are located in the northern area of Montes Finlayson et al., 2006; Carrión et al., 2008, 2018) and northwards in de Málaga. Blanco-Castro et al. (1997) consider Betic holm oak (Quercus Murcia (Sima de las Palomas and Perneras sites: Carrión and Dupré, ilex) as a formation restricted to montane formation. However, the steep 1994; Carrión et al., 1995a) suggesting the existence of glacial refugia relief of some areas of the Andalusian coast allows the oaks to reach po- for temperate, Mediterranean and Ibero-Maghrebian plant taxa (Arpe sitions quite close to the coastline (for example, in La Axarquía of Má- et al., 2011; Carrión et al., 2013, 2015; González-Sampériz et al., 2010; laga). Quercus ilex must have been present during the Upper Leroy and Arpe, 2007; Magri et al., 2017; Manzano et al., 2017; Pleistocene in the coast of Málaga, as Bajondillo site indicates (López- Postigo-Mijarra et al., 2008, 2010). Sáez et al., 2007). As for the deciduous oak species, the most common The Pleistocene flora of these territories includes pines, junipers, and regionally are Quercus canariensis and Q. faginea.Thecorkoak,Quercus xerophytes such as Artemisia, Ephedra, Asphodelus,Cistaceae,Lamiaceae suber, has not been palynologically considered in this study, but it is and Amaranthaceae which may well have formed part of halophytic plausible that it was also in the vicinity of Abrigo 3 del Complejo del coastal communities (Lendínez, 2010; Lendínez et al., 2012; Salazar Humo taking into account the current potential of the species in the et al., 2014), but it is worth emphasizing the continuous occurrence of study area (Montoya, 1981, 1987). both deciduous and evergreen oaks, most likely involving Quercus It is worth mentioning the presence of pinsapo fir(Abies faginea, Q. rotundifolia and Q. coccifera. The occurrence of the alder pinsapo), whose first appearance in the pollen diagram occurs on (Alnus) and pinsapo fir(Abies pinsapo) are also pointing to an important level 23. It continues on level 22, during the Middle Paleolithic, probably patchily tree cover (Alba-Sánchez et al., 2019). What is more and then reappears again in levels 17 and 16 in the transition be- relevant is however the maintainance of the woody assemblage of tween the ending of the Middle Paleolithic and the beginning of Maytenus, Myrtus, Olea, Phillyrea and Pistacia, demonstrating the exis- the Upper Paleolithic. In all cases, its pollen percentages are suffi- tence of enclaves where winter frosts would have been absent. The ciently high – given the low dispersal capacity of this species – to pine pollen signal found in this record suggests the occurrence of coastal sustain a relative proximity of the “pinsapar” (Abies pinsapo com- platforms with Pinus pinea,confirmed by the presence of macroremains munities) to the study area (Alba-Sánchez et al., 2010). Abies in Nerja (Badal, 1990, 1998, 2001), and higher in altitude, Pinus pinsapo, is currently present in the south of the Iberian Peninsula. halepensis and Pinus nigra if we take into account the current ecology This tree is endemic to the western Baetics, with three populations: 6 .Ohnoe l eiwo aaooayadPlnlg 8 22)104281 (2020) 281 Palynology and Palaeobotany of Review / al. et Ochando J.

Fig. 5. Pollen diagram of Abrigo 3 del Complejo del Humo including trees types, shrubs types, herbs types, morphotypes of hydro-hygrophytes plants, and non-pollen microfossils.Cichorioideae,Aster type, hydro-hygrophytes taxa and non-pollen microfossils (NPPs) are out from the total pollen sum. J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281 7

Fig. 6. Synthetic pollen diagram of Abrigo 3 del Complejo del Humo including ecological groups and the main pollen contributors. Pinus include Pinus pinea type and Pinus sylvestris/nigra type. Quercus includes Quercus evergreen and Quercus deciduous (including Q. suber palynotypes). Deciduous Trees include Alnus and Quercus deciduous. Thermophytes include Quercus evergreen, Cistaceae, Ephedra fragilis, Maytenus senegalensis, Myrtus, Olea, Phillyrea and Pistacia.

Sierra de Grazalema, in Cádiz; and Sierra de las Nieves and 1986). It reaches southwestern European border on the edges of the Reales of Sierra Bermeja, in Málaga (Alba-Sánchez et al., the Irabia swamp (Irati, Navarra), while the Montseny massif (Bar- 2018). It is noteworthy the presence in the Iberian Peninsula of celona), in the Coastal-Catalan mountain range, constitutes its the two most diverse morphologically taxa (Abies alba and A. Spanish southeast boundary. On the other hand, Abies pinsapo is lo- pinsapo), whose geographical proximity constitutes a remarkable cated within the group of south-eastern fir trees. Its Iberian distri- corological–paleobotanical curiosity (Asensi et al., 1980). Although bution is restricted to some specific points in the Serranía de the period of appearance, as well as the place of origin and subse- Ronda, in the westernmost side of the Betic mountains and whose quent migrations of the different circum-Mediterranean species of populations have been greatly reduced because of the progressive the Abies genus, remains a controversial issue as there is insufficient aridification of the climate during the Quaternary (Blanco-Castro evidences (Linares, 2011). Several authors (Gaussen, 1964; Quézel, et al., 1997; Alba-Sánchez and López-Sáez, 2013). 1978) place the origin of Mediterranean firs in the Oligocene, and Along the Andalusian Mediterranean coast, the pinsapo fir is also their subsequent diversification in the Miocene. Abies alba must represented in the pollen study of Bajondillo (López-Sáez et al., 2007): have been relatively abundant throughout Europe, a fact confirmed it appears for the first time in the Middle Paleolithic (ca. 50–42 ka), by many pollen records since Abies pollen has been found during the and reappears in the transition to the Upper Paleolithic (ca. 40–38 ka), early Pleistocene samples in some sites (Gaussen, 1944-1968; being relatively abundant during interstadial periods. In Bajondillo, López-Sáez and López-García, 1999; González-Sampériz et al., the decline of Abies pinsapo forests in the most arid climatic periods (Au- 2010). When the cold became very intense Abies took refuge in rignacian to Solutrean) is visible, even though it is still present in the more temperate areas such as Italy, southern Balkan Peninsula or area between 35 and 20 ka. In the pollen sequence of Boquete de Iberian Peninsula. It is unlikely, then, that Abies pollen occurring in Zafarraya (Lebreton et al., 2003) the pinsapo fir appears testimonially Abrigo 3 del Complejo del Humo belong to Abies alba (García in some Middle Paleolithic levels dated before 46 ka. In summary, the López and Allué Camacho, 2005; Alba-Sánchez et al., 2010). In the documentation of pinsapo fir pollen in the Abrigo del Humo comes to Iberian Peninsula, Abies alba occupies almost exclusively the great indicate a new Pleistocene location for this species in the south of the mountain complex of the Pyrenees (Ceballos, 1958; Bresset, Iberian Peninsula. 8 J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281

8. Persistence of Ibero-North African taxa during Late Quaternary Messinian (Quézel, 1985). Many of these elements with tropical origin (Coriaria, Osyris, Smilax, Tribulus, Vitex, Ziziphus) are currently The ocurrence of the arto in the area (Maytenus senegalensis)is restricted to areas in the south and southeast of Spain, mainly through- worth discussing (Plate I). The arto is a Tertiary relict, threatened spe- out the thermomediterranean thermoclimate and arid, semi-arid and cies, whose communities (artales) are especially abundant in Málaga dry ombroclimates (Díez-Garretas et al., 2005). The species is not cur- province (Mendoza-Fernández et al., 2015; Pérez Latorre et al., 2006). rently growing in Gibraltar (Galán de Mera et al., 2000) but it does Díez-Garretas et al. (2005) describe communities with this species in along the southeast coast of the Iberian Peninsula (Cabezudo and the coastal areas of Málaga, mainly dispersed on limestone slopes and Pérez Latorre, 2001). cliffs near the sea (Rincón de la Victoria, La Araña), where the arto is ac- The oldest pollen records of Maytenus in the Iberian Pleistocene ap- companied by Aristolochia baetica, Asparagus albus, A. horridus, Pistacia pear in Sima de las Palomas (Murcia) between 125 and 40–60 ka lentiscus, Rhamnus fontquerianus, R. oleoides, Chamaerops humilis and (Carrión et al., 2003b; Walker et al., 1999, 2008)(Fig. 7). Although the Olea europaea var. sylvestris. These associations develop today in the chronology of both records in Gorham´s and Vanguard Caves has neither dry thermomediterranean bioclimatic belt, apparently as a degradation been finely nor definitively established, it is clear that a great part of the stage of Quercus ilex formations (Pérez Latorre et al., 2006). Maytenus MIS 4-2 dry-cold interval is captured (Carrión et al., 2008, 2018; occurs in the pollen diagram of Abrigo del Humo sporadically during Doerschner et al., 2019), and it includes the survival of Maytenus along the Middle Paleolithic (levels 26, 20 and 17) and continuously since with other typical elements of the xerothermic scrub. Carrión et al. the transition between Middle and Upper Paleolithic layers (levels 17 (2003a) show that the favorable climatic conditions in Sierra de Gádor to 15) (Figs. 5, 6). allowed the coexistence of mesothermophytes elements such as The presence of Maytenus senegalensis in the flora of southeastern Maytenus senegalensis, Myrtus communis, Buxus balearica and Iberia, along with other elements indicating of semi-desert areas such B. sempervirens together with deciduous oak forests during mid- as Tetraclinis or Ziziphus, could have its origin in the Lower Cretaceous Holocene. Similarly, in the Mazarrón sequence (ca. 7617–1569 cal yr in connection with the paleogeographical history of the Mediterranean B.P.) (Carrión et al., 2018), Maytenus abounds in the lower part of the di- (Quézel, 1985). They co-occur with Tertiary taxa such as Fagonia, Lycium agram. The Mayteno-Periplocetum formations could have expanded and Cneorum (Axelrod and Raven, 1978; Quézel, 1983) as well as taxa of more as a consequence of human activities than as a direct cause of bio- xerophytic character (Aristida, Caralluma, Periploca, Zygophyllum)which climatic factors (Carrión et al., 1995b, 2007). Charcoal analyses in Punta represent the African eremitic element related to the arid phases of the de los Gavilanes (García-Martínez, 2009), suggest the presence of

Plate I. Maytenus senegalensis plant. a. Leaves details and spiny stem, b. Flowers details, c. Maytenus senegalensis populations. Pollen grains of Maytenus senegalensis. d. Polar vision, e, f. Equatorial vision. J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281 9

Fig. 7. Iberian Pleistocene and Holocene records of Withania frutescens, Maytenus senegalensis and Periploca laevigata.

Maytenus in the southeastern coast during ca. 4080–2456 cal yr B.P. appearance of Maytenus, Withania and Periploca is restricted to a few (Fig. 7). sites (García-Martínez, 2009; Carrión et al., 2018)(Fig. 7). Although absent from Abrigo del Humo pollen record, other taxa The occurrence of warmth-loving plant communities during the should be mentioned in this paleoecological scenario. Withania Middle-Upper Paleolithic transition in the extreme south of Iberia is frutescens appears along with other thermophytes elements (Myrtus, supported by the Mediterranean pollen sequences of Bajondillo Erica arborea, Pistacia lentiscus, Buxus, Periploca, Maytenus, Osyris, (Torremolinos, Málaga) (López-Sáez et al., 2007), Gorham`s (Gibraltar) Lycium, Calicotome, Ephedra fragilis, Cosentinia vellea, Selaginella (Carrión et al., 2008), and Vanguard (Gibraltar) (Carrión et al., 2018). denticulata, Ruta) in the Pleistocene sequences of Vanguard (Carrión This last shows Pinus, Poaceae, Chenopodiaceae, and evergreen and de- et al., 2018), Sima de las Palomas (Carrión et al., 2003b), Bajondillo ciduous Quercus accompanied by Alnus, Betula, Castanea sativa, Corylus (López-Sáez et al., 2007) and Cueva Perneras (Carrión et al., 1995a) avellana, Juglans regia, Fraxinus, Salix, Ulmus, Olea europaea, Phillyrea, (Fig. 7). With respect to the Pleistocene records of Periploca angustifolia, Buxus, Coriaria, Myrica, Rhamnus, and signiﬁcantly, the thermophytes they are exclusive to the southeast of the Iberian Peninsula at the sites of Maytenus senegalensis, Withania frutescens, Calicotome, Pistacia lentiscus Sima de las Palomas (Carrión et al., 2003b) and Cueva Perneras (Carrión and Myrtus communis. On the southeastern Spanish coast, pollen spectra et al., 1995a), apart from the occurrence of Asclepiadaceae at of mesothermophilous taxa have been recovered from Middle Paleo- Almizaraque, Almería (Mariscal, 1992) during the mid-Holocene, lithic deposits of Cueva Perneras (Carrión et al., 1995a), and Sima de which should correspond to Periploca. For its part, in the Holocene, the las Palomas (Walker, 2001; Carrión et al., 2003b). These show abundant 10 J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281

Quercus and Oleaceae, and frequent presence of thermophytes (Myrtus, Furthermore, the appearance of remains of other carnivore taxa in Erica arborea, Pistacia, Buxus, Periploca, Maytenus, Osyris, Withania, Zafarraya, such as Cuon alpinus, Vulpes vulpes, Crocuta crocuta, Panthera Lycium, Calicotome, Ephedra fragilis, Cosentinia vellea, Selaginella pardus, Mustela erminea and Mustela nivalis, could be considered more denticulata, Ruta). or less ubiquitous when it comes to the occupation of their habitats (Monclova Bohórquez et al., 2012). 9. Neanderthals surviving in coastal southern Iberia Several archeozoological studies from Middle Paleolithic sites across the Iberian Peninsula suggest that Neanderthals obtained a wide variety The palynological record of Abrigo 3 del Complejo del Humo of animal resources, similar from the pattern observed in Iberian early emerges as an optimun thermic refuge during Late Quaternary in Upper Paleolithic modern humans (Delpech and Grayson, 2007; the southern Iberian Peninsula coastline. Thus, the studied pollen se- Patou-Mathis, 2011). The use of resources of coastal origin (mainly ma- quence can be contextualized with the Neanderthal occupation in rine molluscs) during the Middle-Upper Paleolithic is notorious in de- Abrigo 3 del Complejo del Humo. The important connections be- posits such as Gorham´s, Vanguard, Bajondillo, Abrigo 3 and 4 del tween past human groups and their ecosystems can be also consid- Complejo del Humo, and Sima de las Palomas (Barton, 2000; Cáceres ered. It is clear now that the southern Iberian Neanderthals used Cuello de Oro et al., 2001; Cortés-Sánchez, 2000; Cortés-Sánchez et al., extensively the present-day thermomediterranean belts and the 2008, 2011; Fa et al., 2016; Finlayson et al., 2006; Ramos-Fernández lower mesomediterranean, since the deposits with stratigraphic se- et al., 2005; Ramos Fernández et al., 2012; Stringer et al., 2008; Walker quences of the Middle Paleolithic in the south of the Iberian Penin- et al., 2012). In any case, the consumption of shellfish seems to be a sula are contained between both bioclimatic belts (Cortés-Sánchez, complement in the diet of Neanderthal populations in the southern Ibe- 2006; Finlayson and Giles-Pacheco, 1999). The continuous coastal rian Peninsula. Thus, the meat consumption use would be determined resources exploitation is reflected mainly in the records of the Bays by the biotope in which the site is located, as happens in Boquete de of Malaga and Algeciras, where in caves there is enough evidence Zafarraya, finding species of different sizes such as the auroch (Bos to confirm a prolonged persistence of humans (Barton, 2000; primigenius), horse (Equus ferus), red deer (Cervus elaphus), ibex Cáceres Cuello de Oro et al., 2001; Cortés-Sánchez, 2000; Cortés- (Capra pyrenaica), rabbit (Oryctolagus cuniculus) and tortoise (Testudo Sánchez et al., 2008; Fa et al., 2016; Finlayson et al., 2006; Ramos- hermanni), among others (Barroso, 2003; Cortés-Sánchez et al., 1996; Fernández et al., 2005; Ramos Fernández et al., 2012; Stringer Monclova Bohórquez et al., 2012). et al., 2008). The late survival of Neanderthals in this region has Salazar-García et al. (2013) suggest that Neanderthals consumed a been associated to the persistence of these mild climatic conditions wide variety of vegetal resources. The paleoenvironmental and paleo- (Finlayson, 2008; Finlayson et al., 2006; Sepulchre et al., 2007). Nu- ecological traits of the Mediterranean in the south of the Iberian Penin- merous deposits with sequences of long-term occupation such as sula have been reconstructed by pollen studies at cave sites occupied by Gorham´s (Finlayson, 2004; Finlayson et al., 2000), Bajondillo Neanderthals, both inland and near the coast (Carrión et al., 2003b, (Cortés-Sánchez et al., 2004),Abrigo3and4delComplejodel 2013, 2015). Currently, not much is known about Neanderthal con- Humo (Ferre et al., 2004)andCarihuela(Carrión et al., 2019a)reaf- sumption of these foods, in spite of the environmental reconstructions firm these climatic conditions. of southeastern Iberia (Carrión et al., 2003b). It precisely in this context Abrigo 3 del Complejo del Humo site emerge as a remarkable where the paleobotanical findings become more relevant, because we paleofloristic record showing Paleolithic in connection with the Nean- can add now to the Neanderthal diet the possibility of a broad spectrum derthal ecology, highlighted by the existence of a forested habitat simi- of edible plants that likely grew in the vicinity of Abrigo 3 del Complejo lar to other sites of the southern Iberian Peninsula (Carrión et al., 2003b, del Humo, notably the holly oak (Quercus ilex), wild olive (Olea europaea 2008, 2018; González-Sampériz et al., 2010). This forested environment var. sylvestris), and the Stone pine (Pinus pinea). Additionally, more re- could have favored Neanderthal hunting strategies (Finlayson and cent studies have also highlighted how relevant plants were for Nean- Carrión, 2007; Rosas, 2016; Stewart et al., 2019). It seems likely that derthal diets not only in southernmost parts such as at Gorham’sCave this particular environment provided a mosaic of localities where rain- in Gibraltar (Barton et al., 1999), or the east of Iberian Peninsula in fall and warm conditions preserved resource-rich habitats (Jennings Bolomor Cave (Ochando et al., 2019), but even in northern parts of et al., 2011), in contrast with other areas of Europe where Neanderthals Spain (Hardy et al., 2012). In consequence, it is possible to say that may have not had access to supply, particularly during cold periods. For plant foods may have been particularly important in Neanderthal diets instance, archeological works in Abrigo 3 del Complejo del Humo have in south Iberian Peninsula refugia. shown a collection of molluscs close to 1000 specimens on the rocky It cannot be stressed enough that the Pleistocene vegetation land- substrates, formed mainly by mussels and limpets, among others scapes of these coastal territories were sharply different to those ob- (Ramos-Fernández et al., 2005; Ramos Fernández et al., 2012). served today, which are dominated by thermophilous scrub with From a paleoenvironmental perspective, the archeozoological re- scattered pines. Thus, the new data presented in Abrigo 3 del Complejo cords in Paleolithic sites in the coast of Málaga are limited to compare del Humo show that, as in Gibraltar (Carrión et al., 2008, 2018; with the microbotanical data from Abrigo 3 del Complejo del Humo. Finlayson and Carrión, 2007), the last Neanderthals and the first "mod- However, the proximity of Zafarraya and the chronology of the levels ern humans" lived for thousands of years in a propitious environment falling within MIS 3 (42–34 ka) can be used to establish certain for the development and permanence of diversity. This coincides with paleoenvironmental correlation. Zafarraya Cave was mainly inhabited the consideration that the appearance of evolutionary developments by large mammal species (Capra pyrenaica and Rupicapra pyrenaica) within hominins and the long permanence of populations is concen- (Monclova Bohórquez et al., 2012). The presence of Cervus elaphus, trated in the regions of greatest biological diversity in the world Ursus arctos, Lynx pardina and Felis silvestris, exclusive of forest environ- (Carrión et al., 2011). The late Pleistocene coasts of the south of the Ibe- ments, strongly suggests that the region was forested (Saarma et al., rian Peninsula are outstanding in showing the coexistence of thermo-, 2007), which fits the relatively high arboreal pollen percentages in meso-, and supramediterranean plant and animal including dry and Abrigo 3 del Complejo del Humo (Figs. 5, 6). The predominance of her- humid, wooded and treeless biotopes (Carrión et al., 2018). In addition, baceous formations (mainly Amaranthaceae, Fabaceae and Poaceae) the most thermophytic Maytenus, Calicotome, Withania, Periploca, and to some lesser extent shrubs (Erica, Juniperus, Lamiaceae and Osyris, Olea and Pistacia only coexist in the southernmost fossil sites of Olea), mainly in the lower levels of Middle Paleolithic, suggests the pres- the coastal areas that extend from Murcia to Gibraltar suggesting a ence of open areas covered with grasslands. The few remains found of coastal platform as a favorable territory for the survival of the greatest Equus caballus and E. hydruntinus in Zafarraya, possibly due to its higher diversity of environments in the Iberian Peninsula during the MIS 3 in altitude (Bignon and Eisenmann, 2006), supports this interpretation. which the late survival and extinction of the Neanderthals took place. J. Ochando et al. / Review of Palaeobotany and Palynology 281 (2020) 104281 11

Neanderthals lived here in a uniquely diverse environment, and one Bignon, O., Eisenmann, V., 2006. Western European Late Glacial horse diversity and its ecological implications, Equids in Time and Space. In: Mashkour, M. (Ed.), 9th ICAZ that is markedly different to that present landscape. Conference, Durham 2002, pp. 161–171. Birks, H.J.B., 2005. Fifty years of Quaternary pollen analysis in Fennoscandia 1954-2004. Grana 44, 1–22. Blanco-Castro, E., Casado-González, M.A., Costa-Tenorio, M., Escribano-Bombín, R., García- Declaration of Competing Interest Antón, M., Génova-Fuster, M., Gómez-Manzaneque, F., Moreno-Saiz, J.C., Morla- Juaristi, C., Regato-Pajares, P., Sainz-Ollero, H., 1997. Los bosques ibéricos. Una The authors declare that they have no known competing financial interpretación geobotánica, Planeta, S.A, p. 572. fl Bonnefille, R., 2010. Cenozoic vegetation, climate changes and hominin evolution in Trop- interests or personal relationships that could have appeared to in u- ical Africa. Global Planet. Change 72, 390–411. ence the work reported in this paper. Bottema, S., 1975. The interpretation of pollen spectra from prehistoric settlements (with special attention to liguliflorae). Palaeohistoria 17, 17–35. Acknowledgements Bresset, V., 1986. Contribution a l´étude phyto-écologique des sapinières oriento- pyrénéennes. Dissertation. Universidad de Niza. Broothaerts, N., Robles, S., Abel-Schaad, D., Pérez-Díaz, S., Alba-Sánchez, F., Luelmo- The research of this work was funded by the projects MED-REFUGIA- Lautenschlaeger, R., Glais, A., López-Sáez, J.A., 2018. Reconstructing past arboreal RTI2018-101714-B-I00 (Plan Nacional I+D+I, Spanish Ministry of cover based on modern and fossil pollen data: A statistical approach for the Gredos Range (Central Spain). Rev. Palaeobot. Palynol. 255, 1–13. 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