Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299

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Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

Palaeoenvironmental changes in the Padul Basin (Granada, Spain) over the last 1 Ma
based on the biomarker content
José E. Ortiz a,⁎, Trinidad. Torres a, Antonio Delgado b, J.F. Llamas a, Vicente Soler c, Maruja Valle d,
Ramón Julià e, Laura Moreno a, Arantxa Díaz-Bautista a
a
Laboratory of Biomolecular Stratigraphy, E.T.S.I. Minas, Universidad Politécnica de Madrid. C/ Ríos Rosas 21, Madrid 28003, Spain
b
Estación Experimental “El Zaidín” (C.S.I.C.). C/ Profesor Albareda 1, Granada 18008, Spain
c
Instituto de Agrobiología y Productos Naturales (C.S.I.C.). Avda Astrofísico Fco Sánchez 3, La Laguna 38206, Tenerife, Spain
d
Facultad de Ciencias, Universidad de Salamanca. Pza de la Merced s/n. Salamanca 37008, Spain
e
Instituto de Ciencias de la Tierra “Jaume Almera” (C.S.I.C.). C/ Lluís Solé i Sabarís s/n, Barcelona 08028, Spain

a r t i c l e i n f o a b s t r a c t

Article history: The n-alkane components of sediments were measured in 530 samples taken from a 107 m-long core in the
Received 9 June 2010 Padul Basin (Andalusia, Spain) covering from the Lower Pleistocene (ca. 1 Ma) to the mid-Holocene (ca.
Received in revised form 23 September 2010 4.5 ka B.P.). We show that the relative percentages of the high molecular weight n-alkanes (C27, C29 and C31)
Accepted 10 October 2010
are useful tools to reconstruct the ancient vegetation and palaeoclimatological evolution of this basin. On the
Available online 15 October 2010
basis of these proxies, we distinguished twelve long-term climatic periods linked to alternating dry/humid
Keywords:
scenarios: 6 Dry Episodes alternating with 5 Humid Episodes were interpreted in the Pleistocene record,
n-alkanes together with a final Episode covering the Holocene. Dry scenarios produced the recession of forests and the
Palaeoclimatology development of grasses, whereas the inverse pattern was interpreted during humid phases. The good
Pleistocene correlation with the Guadix-Baza Basin record showed that, in most cases, dry periods coexisted with warm
Holocene conditions, while humid ones were linked to temperate-colder conditions. Only during the last long-term Dry
Padul Basin Episode of the Pleistocene, which coincided with the end of the Last Glacial Period, did low temperatures
concur. The relative percentages of the high molecular weight n-alkanes also indicate rapid changes in
vegetation produced by short-scale arid cycles during the last Glacial Period (b 100 ka B.P.), which are linked
to the Heinrich Events and Younger Dryas. This interpretation has been confirmed by previous pollen analysis
performed in the Padul Basin. Likewise, a correlation was observed with Marine Isotope Stages (MIS). These
results show that vegetation responds to long-term cycles (orbital scale) and millennial-scale global cycles.
Therefore, climatic variations in the Padul Basin responded to global climatic changes, although with local
climatic effects. Good correspondence was also observed with elemental, isotopic and other molecular organic
proxies, thereby amplifying the palaeoclimatological knowledge of the Padul Basin during the Pleistocene,
especially in some spans that were previously unclear.
© 2010 Elsevier B.V. All rights reserved.

Palaeoenvironmental reconstructions can be achieved by a wide
range of techniques involving studies of pollen spectra, fossil as-
semblages, stable isotope analysis of inorganic compounds, and
elemental, isotopic, and molecular organic geochemical proxies,
among others. Within organic molecular geochemistry proxies, n-
alkanes are among the biomarkers most commonly used.
Hydrocarbons are less susceptible to microbial degradation during
diagenesis than most types of organic matter because they lack the
functional groups that confer chemical reactivity (Meyers et al., 1995;
Prahl and Carpenter, 1984). Moreover, they have low water solubility.
Although hydrocarbons account for a small fraction of the total
organic matter in both biota and sediments, diagenetic degradation of
⁎ Corresponding author. Tel.: +34 91 3366970; fax: +34 91 3366977.
E-mail address: joseeugenio.ortiz@upm.es (J.E. Ortiz).
other compounds may exaggerate the source signatures of biomarker
hydrocarbons (Tenzer et al., 1999). These compounds can be easily
transported short distances by eolian sources or runoff.
The n-alkanes present in sediments reflect mainly the contribution
from algae, aquatic macrophytes and land plants. These compounds
remain unaltered during passage through the digestive systems of
animals and their contribution from animals or fungi to sediments is
negligible (Ficken et al., 1998). Lake sediments receive three main
sources of biotic hydrocarbons: 1) algae and bacteria that live in the
lake; 2) aquatic macrophytes that live in the lake; and 3) vascular
plants that live around it. These sources produce distinct n-alkane
profiles.
More specifically, Schwark et al. (2002) reported the leaf wax C27, C29
and C31 n-alkane distribution of present-day plants and showed that, in
general, grasses have high concentrations of C31, while deciduous tree
assemblages are dominated by C27. They compared the C27, C29 and C31

0031-0182/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2010.10.003
 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
profiles with the pollen assemblages in Lake Steisslingen (Germany)
along the last ca. 15.5 ka B.P. and found a similar distribution pattern.
They resolved centennial and even decadal palaeoenvironmental
variations, thereby demonstrating the suitability of using this proxy in
palaeoclimatic studies. Furthermore, these profiles are a potential tool
for discriminating autochthonous from allochthonous pollen input.
Located in Andalusia in the southern part of the Iberian Peninsula
(Fig. 1), the Padul Basin has one of the best records of Pleistocene
sediments, registering a thickness of more than 100 m. Previous studies
addressing the stratigraphy and palynology (Florschütz et al., 1971;
Menéndez Amor and Florschütz, 1962, 1964; Pons and Reille, 1988;
Valle et al., 2003), and also biomarkers (Ortiz et al., 2004; del Río et al.,
1992) of this basin have been published. The 107 m-deep borehole
drilled in 1997 near the western edge the basin, where the bog reaches
its maximum depth (Nestares and Torres, 1998) (Fig. 1), allowed the
observation of two markedly different hydrogeological scenarios (Ortiz
et al., 2004). From ca. 1 Ma (meter 107) to ca. 400 ka B.P. (meter 60), a
considerable run-off recharge made the water body deeper than in the
rest of the record, thus indicating lacustrine conditions. In contrast, from
ca. 400 to 4.5 ka B.P., the basin became a peat bog (“…enclosed basins of
deposition, such as former lake basins…where plant growth and
succession results in a compact accumulation of plant debris”, Sirkin,
1978) with the main water input entering as groundwater inflow.
The concentration of organic carbon, the atomic H/C and C/N
ratios, and the δ13C and CPI values were shown to be excellent
palaeoenvironmental proxies for the study of the palaeoclimatological
and palaeohydrological evolution of the Padul peat bog (cf. Ortiz et al.,
2004). However, the global climatic changes, such as those occurring
from ca. 170 to 25 ka B.P. (from meter 33.6 to 7), for example the
global warming occurred during Marine Isotope Stage (MIS) 5e or the
Heinrich Events, did not strongly affect these palaeoenvironmental
proxies, the values of which show little variation.
Here we sought to complete the reconstruction of the palaeoen-
vironmental evolution of the southern part of the Iberian Peninsula of
the last 1 Ma by interpreting the C27, C29 and C31 n-alkane profiles of
Fig. 1. Geographical and geological setting of the Padul Basin.
287
the sediments recorded in the Padul Basin. We also compare these
results with those obtained in the nearby Guadix-Baza Basin.
1. Geographical and geological setting
The Padul Basin is located 20 km south of the city of Granada
(Andalusia, southern Spain). It was formed in a highly subsident fault-
bound tectonic realm at the foot of the Sierra Nevada, and consists of
an endorrheic basin, surrounded by mountains. It is located close to
the Betic range and is surrounded by the Albuñuelas Range and Manar
mountains, the latter belonging to the Sierra Nevada Range. With a
NW–SE longitudinal axis, the Padul Basin is situated 720 m above sea
level and some parts are permanently covered by shallow water. It has
a surface area of 4 km2 and a maximum depth of 100 m. The bedrock
consists mainly of faulted Mesozoic dolostones that caused the basin
to sink gradually.
The basin is a discharge area for the groundwater flow of
surrounding aquifers. The flow directions change from sub-horizontal
in the Mesozoic aquifers adjacent to the basin to essentially upward
discharge inside the peat-containing depression (Cañada, 1984).
Present rainfall in the area is a minor factor in the water balance of
the peat deposit and run-off input of water is estimated to contribute
only about 8% of the total (Cañada, 1984). Consequently, changes in
the water table in the peat are controlled indirectly by infiltration of
water from the surrounding mountains (artesian waters), where
accumulated snow during colder phases subsequently melts.
The climate is Mediterranean with a strong continental influence:
winters are cold and dry while summers are extremely hot, with
maximum temperatures over 40 °C. The average annual rainfall ranges
from 400 to 450 mm yr− 1, the evapotranspiration is 700–900 mm yr− 1
and the mean annual temperature 15 °C (Worldwide Bioclimatic
Classification System Index).
The Padul Basin comprises vegetation debris from Sierra Nevada,
where it is possible to distinguish a series of vegetation belts (Florschütz
et al., 1971) comprising the following from bottom to top: steppe,
288 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
Quercus ilex (more humid), Quercus pubescens (xeromorphic), Acantho-
limon-Tragacanta (Eastern Mediterranean character) and Mediterra-
nean mountain steppe.
The stratigraphy of the Padul borehole (Latitude: 37°01´01´´ N;
Longitude: 3°36´07´´ W; Elevation: 714.20 m) is shown in Fig. 2. For
dating the borehole (cf. Ortiz et al., 2004), the following four
techniques were used: conventional radiocarbon analysis, U/Th
analysis, amino acid racemization and palaeomagnetism.
We refer to the sampled horizons of the borehole by their depth, in
cm, from top to bottom (e.g. sampled level SPD-0125 is at 125 cm).
2. Methodology
A total of 530 samples were taken at approximately 15- to 20-cm
intervals along the borehole core. The lipid fraction was then extracted
from these samples and the n-alkane content of the organic matter
measured.
About 5–10 g of sediment was first ground and biomarkers were
extracted following the LEB protocol (Lucini et al., 2000). Briefly, this
procedure consists of a 24-h soxhlet extraction with dichloromethane
and methanol 2:1 (suprasolv Merck) and concentration of the isolated
bitumen using a rotor-vapor device. Three bitumen fractions were
extracted through liquid chromatography in a silica-alumina glass
column using solvents of different polarities: n-hexane, dichloro-
methane/n-hexane 4:1 and methanol. Samples were injected into an
HP 6890 gas-chromatograph equipped with selective mass detector
(HP 5973) and an ATM-5 column (250 × 0.25 mm; 0.20 μm). Helium
was the carrier gas. The oven temperature was programmed from
60 °C to 300 °C at 6 °C/min (holding time 20 min). The injector was
programmed at 275 °C. The compounds were identified with the Data
Analysis Program and the Wiley Library. n-Alkanes were calculated
from the GC/MS chromatograms of mass/charge m/z 57 from the first
bitumen fraction (the one extracted with n-hexane) and decafluor-
obiphenyl was used as an internal standard.
3. Results
Typical chromatograms of n-alkanes from selected samples are
shown in Fig. 3. All the samples in the Padul record showed an odd-
over-even carbon number predominance, with a chain length
distribution ranging mainly from C15 or C17 to C33 or C35, maximizing
at C27, C29 or C31 n-alkanes in most samples of the upper part of the
core, while the C17 n-alkane was predominant in samples from meters
107 to 60 (Fig. 2).
The n-alkane composition as relative percentages of C27, C29 and
C31 with respect to the sum C27 + C29 + C31 reveals several oscillations
in the three curves (Fig. 2). The relative percentage of C27 varied
between 61% and 6%, whereas the relative percentage of C29 ranged
between 57% and 2%, and the percentage of the C31 enantiomer
oscillated between 74% and 5%. Likewise, higher variations were
observed in the lower part of the record.
For comparison, Fig. 2 also shows the logs of other (elemental,
isotopic and molecular) organic geochemical proxies obtained by
Ortiz et al. (2004): concentration of organic carbon (%TOC), H/C and C/
N ratios, δ13C values, the carbon preference index (CPI) and the n-
alkane predominant chain. These results will complete the interpre-
tation based on the relative percentages of the higher molecular
weight n-alkanes.
4. Discussion
The predominant n-alkane chain log is interpreted as an indicator of
the origin of the organic debris input (algae, aquatic macrophytes or
land plants) (Cranwell et al., 1987; Ficken et al., 2000) and of temporal
variations of the water body depth. The predominance of n-alkanes of
low molecular weight at the bottom of the core (Nmeter 70) was
interpreted to indicate that algae and bacteria made a significant
contribution to organic matter sources (cf. Gelpi et al., 1970; Blumer
et al., 1971; Cranwell et al., 1987), thereby suggesting, as do other
palaeoenvironmental proxies (H/C, C/N, δ13C), high water-level phases
(Ortiz et al., 2004). In contrast, in the lower and middle parts of the core,
the predominance of C27, C29 and C31 n-alkanes might be related to
palustrine conditions (Fig. 2) with considerable organic matter inputs
from trees or grasses, although the presence of aquatic macrophytes
may be relevant (cf. Cranwell, 1973, 1984; Ficken et al., 2000). In fact,
vascular plants on land or around the edges of lakes contain large
proportions of high molecular weight n-alkanes, (C27, C29 and C31), in
their epicuticular waxy coatings (Cranwell et al., 1987; Eglinton and
Hamilton, 1963, 1967; Eglinton and Calvin, 1967; Rieley et al., 1991),
and the abundance of these wax hydrocarbons reflects the amount of
organic matter transported to lakes from the surrounding land. More
specific differentiations can be made: C31 is the major n-alkane in areas
where grasses dominate, while C27 and C29 are more abundant in
sediments where trees predominate (Cranwell, 1973).
In order to discern the origin of the higher molecular weight n-
alkanes the Padul Basin sediments in the lower 60 m (when it became
a peat bog s.s.- cf. Ortiz et al., 2004), that is, the potential input from
macrophytes growing in the basin vs. terrestrial plants (deciduous
trees, conifers, and grasses), we have recently analyzed leaves and
pine needles from living plants (Table 1) in the surroundings of the
Padul Basin. The results showed that the main macrophytes living in
the bog (rush and reed) maximize at C29, with low percentages of the
C31 isomer (16–22%). As most of the values of the relative percentage
of C31 with respect to the sum C27 + C29 + C31 along the Padul record
(Fig. 2) largely differ from those obtained in living macrophytes (the
same can be observed for the C27 isomer), we assume that the input of
organic matter derived from terrestrial plants was significant and,
therefore, the relative percentages of the higher molecular weight n-
alkanes can be interpreted in terms of palaeoenvironmental varia-
tions. Also, some samples showed a predominance of C33, character-
istic of Juniperus (Ficken et al., 2000) (see Table 1).
However, some exceptions were detected, such as between meters
36.0 and 33.6, in which low molecular weight n-alkanes, typical of
phytoplankton, were found to be predominant. This observation
indicates an increase in the water body level, thus corroborating the
information provided by the other geochemical proxies (cf. Ortiz et al.,
2004).
The results of that study also showed a good preservation of
organic matter and little diagenesis (removal of components) during
transport and after deposition in the Padul Basin (cf. Ortiz et al., 2004),
as interpreted from the atomic H/C ratios and CPI values. In fact,
atomic H/C ratios, which can be used to determine the preservation of
organic matter, are below 0.8 only in very few cases, i.e., H/C values
decline during diagenesis, mainly as a result of the selective removal
of unstable components during transport and after deposition by
bacterial respiration and inorganic oxidation (Talbot and Livingstone,
1989) and the preservation of unstable components is enhanced by
rapid transport and burial in an anoxic environment (Talbot, 1988).
Similarly, the CPI can be used as a proxy for the preservation po-
tential of organic matter when there is a clear predominance of waxes
from higher plant, i.e. diagenetic processes cause their CPI to gradually
decrease to 1 (Hedges and Prahl, 1993) because of bacterial degradation,
which produces the same content of odd and even chain alkanes. In the
Padul record CPI values were greater than 1.5, which suggests minimal
diagenesis of the organic matter, that is to say, very little chemical
removal of components during transport and after deposition occurred,
with scarce exceptions in the lower part of the core. However, this index
cannot be used in all cases as a preservation potential proxy but for the
contribution from terrestrial plants (cf. Cranwell, 1973; Farrimond and
Flanagan, 1996; Rieley et al., 1991; Wang et al., 2003; Zhang et al., 2004)
and a CPI of 1 may also indicate immature organic matter with little
influence from higher plants. This would explain the range of CPI values
 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
Fig. 2. Stratigraphy and chronology of Padul peat bog borehole core (modified from Ortiz et al., 2004). Datings are followed by the method used for their calculation: 14C, U/Th, aard (amino acid racemization) and pm (palaeomagnetism).
Owing to drawing limitations, some 14C results have not been represented (SPD-0213: 7220 ± 190 yr; SPD-0335: 9019 ± 241 yr; SPD-0345: 9021 ± 245 yr, SPD-0407: 11,329 ± 213 yr). Pons and Reille (1988) dated the top of the Padul record
at 4450 ± 60 yr. Concentration of organic carbon (%TOC), H/C, C/N, δ13C, predominant n-alkane chain, CPI logs (cf. Ortiz et al., 2004) together with the n-alkane composition profile as percentages of C27, C29, and C31 isomers with respect to the
sum C27 + C29 + C31 along the Padul borehole. H/C ratios are represented up to 5, although there are some greater values, especially at the bottom of the core (in some cases N100). H/C cut-off values of different groups (0.8; 1.3 and 1.7) are

289
represented.
290 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299

Fig. 3. Representative gas chromatograms of n-alkanes (m/z) 57 distributions in selected samples from the Padul record.

detected, which varied from 0.23 to 8.27. In fact, Sachse et al. (2006)
observed CPI values as low as 0.9, 1.3 or 1.8 in mosses and Cladonia
species, although with a large variability, as specimens of these taxa also
showed values of 5.0 or 7.2. Furthermore, Mügler et al. (2008) observed
CPI values ranging from 0.9 to 23.7 in macrophytes. Similarly, deciduous
trees and grasses provided notable CPI differences, even within each
species and under different climatic conditions (Mügler et al., 2008; Rao
et al., 2009, Sachse et al., 2006).
The n-alkane composition profile as percentages of the C27, C29,
and C31 isomers with respect to the sum C27 + C29 + C31 along the
Padul borehole core shows marked variations (Fig. 2).
A C31 maximum in the n-alkane distribution of grasses is well
documented (Cranwell, 1973; Maffei, 1996a), while C27 and C29 are
more abundant in trees. However, according to Herbin and Robins
(1968), and Cranwell (1973), pines also have high concentrations of C31.
Schwark et al. (2002) reported the leaf wax C27, C29 and C31
distribution of present-day plants and showed that two Artemisia
species have high concentrations of C31, followed by 40–50% of C29
and very small amounts of C27. Cold-climate pine species (Pinus nigra
and Pinus cembra) have the highest abundance of C31, while other
types of pine are enriched in C29. In general, deciduous trees (Betula)
have higher concentrations of C27. Our results of present-day plants
 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299 291

Table 1 well as the growing environments. In order to check whether the

Predominant n-alkane of leaves and pine needles of different present-day plants from relative percentages of C27, C29 and C31 in the Padul record reflect the
Spain, most of them present in the surroundings of the Padul Basin and relative
percentages of C27, C29 and C31 n-alkanes with respect to the sum C27 + C29 + C31.
presence of deciduous trees vs. grasses and, therefore whether they
could be linked to humid (and cold) vs. dry (and warm) periods
Scientific name Common name Predominant %C27 %C29 %C31 [typical environmental patterns for the Mediterranean realm- cf.
n-alkane chain
Horowitz, 1989, 2001) and Ortiz et al. (2006)], we compared our
Castanea castanea Chesnut tree 27–29 41 41 18 profiles (Fig. 2 and 4) with the pollen assemblages reported by
Quercus robur Oak 27 42 40 18
Florschütz et al. (1971) and Pons and Reille (1988) (Fig. 5). Although
Juglans regia Walnut tree 31 24 30 46
Olea europaea Olive 33 14 35 51 further discussion is provided in Section 6, good correlation was ob-
Quercus ilex Evergreen oak 29 35 36 28 served between all records, with higher percentages of C27 when
Populus nigra Poplar 27 50 44 6 arboreal pollen increased, and lower C27 values when non-arboreal
Pinus pinaster Maritime pine 29 37 46 17 pollen, especially Artemisia and Chenopodiaceae, predominated.
Pinus nigra Black pine 27 46 34 20
Therefore, as both vegetation and its n-alkane content are
Pinus halepensis Aleppo pine 29 9 60 31
Juniperus communis Common Juniper 33 10 24 66 conditioned by temperature and moisture, we assume that the relative
Ulex europaeus Gorse 31 16 31 52 percentages of C27, C29 and C31 are explained mainly by the amount of
Artemisia arborea Mugwort 31 22 37 41 precipitation and moisture (wet-dry episodes) together with temper-
Juncus maritimus Rush 29 31 53 16
ature. The enrichment in C27 can be attributed to the colonization by
Phragmites communis Reed 29 35 43 22
Stipa tenacissima Esparto grass 31 8 35 57 many species of deciduous trees and, thus, the presence of more humid
Erica mackaiana Mackay´s Heather 31 27 25 48 conditions (linked mainly to cold conditions). However, high amounts
Calluna vulgaris Scotch Heather 31 26 17 57 of C31 in sediments can be associated with dry phases, (linked to warm
Pteridium aquilinum Fern 27 56 34 10 phases), with the development of extensive grass-vegetation cover. In
Gramineae Grass 31 13 40 47
fact, the arid-climate Artemisia is found in northern Europe and in arid/
semi-arid localities of Andalusia (Southern Spain).
The high negative correlation coefficient obtained between the
show similar results (Table 1), though with some inter-species var- percentages of C27 and C31 along the Padul record (Table 2) may be
iability: we found that deciduous trees maximize at C27, while in related to wet/dry phases. As observed from our findings, while the
grasses, gorse, and Ericacea species C31 was predominant. However, percentage of C29 showed an independent behavior with respect to that
there were some exceptions as walnut tree maximized at C31, of C27, it showed a certain negative correlation with the percentage of
although its presence along the Padul record is scarce (cf. Florschütz C31. These results suggest that the contribution of C29 is attributable to
et al., 1971; Pons and Reille, 1988), and Artemisia arborea showed a variations in the density of conifers in the basin neighborhood and some
considerable content of C29 and C31 homologs. As it can be observed, kinds of grasses, not to the establishment of diverse deciduous tree
pines maximized at C29 although they may also have substantial assemblages.
concentrations of C31 (Schwark et al., 2002). Thus, the increasing Several oscillations were observed in the curves of the relative
contribution of C29 could be attributed to a diversification in conifers percentages of high molecular weight n-alkanes (Fig. 2). However, it is
(Schwark et al., 2002), but also to the establishment of more diverse difficult to describe the climate-related phases that occurred in the last
deciduous tree assemblages (Harwood and Russell, 1984; Prasad and 1 Ma. In order to better interpret the general palaeoenvironmental
Gülz, 1990; Rieley et al., 1991). trends that affected the Padul Basin, we performed a smoothing analysis
Although leaf wax n-alkanes have proved useful as chemotaxo- of the relative percentages of C27, C29 and C31 values using the linear
nomic characters for several plants (Maffei, 1994, 1996a,b; Maffei trend method, calculating each smoothed new value from the 5 closest
et al., 1997), Sachse et al. (2006) observed an increase in the average values (Fig. 4). We also perform the smoothing to avoid variations
chain length (ACL) of n-alkanes in the same species of deciduous trees linked to the sedimentological characteristics that could affect the
from a north to south transect in Europe. This observation implies that molecular profile in some levels. In fact, the n-alkane profiles exhibited
climatological conditions have a certain influence in the molecular much larger variations in the lower part of the record than in the upper
composition of plants, although the predominant n-alkane chain peaty sediments (Fig. 2). This observation is possibly due to the different
remained the same at different sites, with the sole exception of birch hydrogeological scenarios: lacustrine conditions in the lower part (107–
(Betula). Similarly, leaf wax n-alkanes from barley (Hordeum vulgare) 60 m) vs. palustrine conditions in the upper 60 m. Likewise, with this
and beech (Fagus sylvatica) in Central Europe do not exhibit exercise we diminished the effect linked to analytical artifacts.
significant changes in their relative percentages of the C27, C29 and It should be noted that this mathematical transformation may
C31 homologs over their growing seasons (Sachse et al., 2009, 2010). mask short palaeoclimatic events and, consequently, these events will
In contrast, the maple (Acer pseudoplatanus) shows changes both in not be detected in the new curves. Nevertheless, diverse palaeoenvir-
the concentration and in the relative percentages of n-alkanes (Sachse onmental periods, representing long-term episodes of relative climatic
et al., 2009). stability, were established.
On the basis of the isotopic signal (δD) of the n-alkanes, Sachse et al. Thus, on the basis of the profiles of relative percentages of C27, C29
(2006) concluded that precipitation was the main factor influencing the and C31, we distinguished several palaeoenvironmental episodes linked
characteristics of these compounds, although temperature can also to alternating drier (D) and more humid (H) conditions in the Padul
affect the n-alkane chain length in leaves on a seasonal scale, as Maffei record (Fig. 4).
et al. (1993) observed in Rosmarinus officinalis. Warmer conditions Generally speaking, the relative percentages of the smoothed C27
produce an enrichment of high molecular weight n-alkanes in de- values were lower than those of C31, except from meters 35 to 30, in
ciduous trees (higher ACL) as plants can alter the chain-lengths of their which they were similar, and between meters 50–40 and 13–7, in which
leaf waxes to minimize the evaporation of water vapor (Sachse et al., the percentage of C27 was higher. These observations indicate that grasses
2006). Thus, the C27 n-alkane content in sediments with considerable were predominant in the surroundings of the Padul Basin along 1 Ma,
input from higher plants would increase under colder and humid although the presence of deciduous trees was constant. Conifers, which
conditions, whereas drier and warmer climates would produce usually maximize at C29, had a strong presence and extensive deciduous
enrichment in the C31 homolog. tree forests were established from meters 50–40, 35–30 and 13–6.
In brief, the molecular distribution of higher plant leaf waxes is a A description of the palaeoenvironmental episodes established along
function of the composition and plant types of source vegetation as the Padul record is found below, the duration of which was calculated
292 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
Fig. 4. Smoothed n-alkane composition profile as percentages of C27, C29, and C31 isomers with respect to the sum C27 + C29 + C31 along the Padul borehole with the
palaeoenvironmental interpretation (H: Humid Episode; D: Dry Episode).
from the age model of Ortiz et al. (2004), based on 9 14C, 5 U/Th, and 4
amino acid racemization measurements, and palaeomagnetism. For
improved accuracy, the ages of the upper part of the core (15 m) were
established by the numerical datings between the intervals considered.
4.1. Dry Episode 1 (meters 107.0–99.0; Nca. 880 ka B.P.)
From the bottom of the Padul core to meter 99.0, low percentages of
C27 and parallel high percentages of C31 are observed. These findings are
interpreted as very dry conditions, with extended steppe vegetation and
low deciduous trees in the surroundings of the basin. In fact, the
percentage of C31 is around 50%. These conditions became progressively
slightly wetter through to the end of this episode, i.e., the percentage of
C27 increases while that of C31 decreases.
4.2. Humid Episode 1 (meters 99.0–84.0; ca. 880 ka B.P.–ca. 630 ka B.P.)
In this episode, high percentages of C27 appear, being almost 10%
higher than values between meters 107 to 99. This observation is
accompanied by a double decrease (lowering of ca. 20 %) in the
percentage of C31. There is also a slight increase in the percentage of
C29 (10%). These values indicate the development of forests, including
pines, and the retreat of grasses, thereby suggesting the prevalence of
more humid conditions than during Dry Episode 1.
4.3. Dry Episode 2 (meters 84.0–68.0; ca. 630 ka B.P.–ca. 450 ka B.P.)
The transition between Humid Episode 1 to this span is characterized
by a sharp lowering of the percentage of C27 accompanied by a marked
increase in that of C31. Nevertheless, in this episode some oscillations are
observed in the n-alkane profiles, thus indicating climatic instability.
Three of these are linked to dry conditions, with low C27 and high C31
percentages (meters 84–82, 80–76, and 72–68; 630–600 ka B.P., 580–
535 ka B.P., 495–460 ka B.P., respectively), which indicate a consider-
able expansion of grasses, alternating with two wetter phases (82–80,
and 76–72; 600–580 ka B.P., and 535–495 ka B.P. respectively), with
more deciduous trees. In the latter phase, there is also an increase in the
percentage of C29, which is interpreted as a minor development of
conifers. However, during the humid phases pluviosity did not reach
rates as high as in other episodes defined in the Padul record.
4.4. Humid Episode 2 (meters 68.0–62.0; ca. 450 ka B.P.—ca. 405 ka B.P.)
The transition to this short episode is reflected by an increase in
the percentage of C27, which reaches 32% and, is paralleled by a
decrease in the percentage of C31. This observation indicates an
expansion of deciduous trees, which is linked to humid conditions.
4.5. Dry Episode 3 (meters 62.0–57.0; ca. 405 ka B.P.—ca. 360 ka B.P.)
In contrast, in this interval the percentage of C27 decreases and
similarly there is a decrease in that of C31, which marks a dry episode,
with an expansion of grasses, which predominate over deciduous trees.
4.6. Humid Episode 3 (meters 57.0–40.0; ca. 360 ka B.P.—ca. 235 ka B.P.)
This is a large time-span characterized by very wet conditions,
especially at the end, which produced the expansion of deciduous trees
over grasses. In fact, increases of ca. 15–20% in the C27 profile, which
reaches 37%, are observed with respect to the previous time-span.
Similar decreases in the percentage of C31 appear, in some cases falling
below that of C27. There is, however, a slightly drier phase between
meters 52 and 50 (ca. 320–305 ka B.P.). The end of this episode (meters
50–40; ca. 305–235 ka B.P.), in which the wettest conditions along the
last 1 Ma occurred in the Padul Basin, correlate with Riss I.
 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299 293

Fig. 5. Tentative correlation between the palaeoenvironmental episodes distinguished in the Padul Basin (smoothed profile of the relative percentage of C27 alkane), and the
palynozones established by Florschütz et al. (1971) and Pons and Reille (1988) (A.P.: Arboreal pollen; N.A.P.: Non-arboreal pollen).

4.7. Dry Episode 4 (meters 40.0–30.0; ca. 235 ka B.P.–ca. 170 ka B.P.)
On the basis of lithology, we distinguished three spans within this
episode. In the first, ranging between meters 40.0 and 36.0 (ca.
235 ka B.P.–180 ka B.P.) and made of peat, the percentage of C27
decreases while that of C31 increases (Fig. 4). These observations are
interpreted as the recession of deciduous trees and the expansion of
grasses, that is to say a transition from humid to drier conditions. It is
important to remark a marked increase in the percentage of C27 at
meter 38.70, followed by a considerable decrease at meters 38.00–
37.60.
The second part of this episode comprised meters 36.0 to 33.6 (ca.
180 ka B.P.–ca. 170 ka B.P.). In this span, which consists mainly of marls,
the percentage of C27 reaches minimum values (Fig. 4). In contrast, C31
294 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
Table 2
Correlation matrix of the relative percentages of n-C27, n-C29 and n-C31 n-alkanes along
Padul peat bog core. p: significant level.
n-alkane %n-C29
%n-C27 − 0.0115
p = 0.791
%n-C29 – − 0.5125
has the highest values. This observation is interpreted as the presence of
dry conditions, which produced the recession of deciduous forests and
the spread of grasses. This interpretation is consistent with that
provided by other organic geochemical proxies. In this case, warm
conditions also occurred. In fact, the H/C, C/N, δ13Corg values indicate an
increase in the water level of the Padul Basin, which has been attributed
to a warm-temperate period that accelerated thaw in the surrounding
mountains and higher runoff and groundwater recharge of the peat bog
(cf. Ortiz et al., 2004). In spite of being warmer and more arid, this period
is paradoxically reflected in the Padul stratigraphic record by a growth in
clastic material and water input (in this case the n-alkane profiles did
not record changes in the water level). This growth is possibly asso-
ciated with soil destruction, but, more importantly, with a displacement
of the “permanent snow” line upwards, thereby allowing more liquid
water to infiltrate during the spring and summer thaw. This phase
correlates with MIS 7a.
The third span ranges from meters 33.6 to 30.0 (ca. 170 ka B.P.–ca.
160 ka B.P.) and consists of peat. The percentages of both C27 and C31
increase while C29 decreases (Fig. 4). This finding suggests smaller
numbers of pines and a greater presence of grasses and other kinds of
trees, and reflects a transition to wetter conditions.
4.8. Humid Episode 4 (meters 30.0–25.0; ca. 170 ka B.P.–ca. 135 ka B.P.)
From meter 30.0 to the end of this span, the percentages of C27 and
C31 show a negative high covariation between each other. While C27
increases, C31 decreases (this is one of the few episodes where %C27 N %
C31). There is a peak of C27 paralleled by a minimum of C31 between
meters 27 and 26.8 (ca. 150 ka B.P.). These observations suggest the
presence of more humid conditions, which correlate with MIS 6.
4.9. Dry Episode 5 (meters 25.0–18.0; ca. 135 ka B.P.–ca. 95 ka B.P.)
Between meters 25.0 and 22.5 (ca. 135 ka B.P.–ca. 125 ka B.P.) a
progressive increase in the percentage of the C31 is observed (Fig. 4),
together with a decrease in C27, thus indicating the transition to drier
conditions.
Between meters 22.5 and 19.0 (ca. 125 ka B.P.–ca. 100 ka B.P.) the
percentage of C31 reaches a maximum (Fig. 4) while, that of C27 registers
a minimum. These observations are interpreted as the establishment of
arid and warm conditions, which correlate with MIS 5.
4.10. Humid Episode 5 (meters 18.0–7.0; ca. 95 ka B.P.–ca. 25 ka B.P.)
The transition to the Early Glacial Period is reflected by a pro-
gressive increase in the percentage of C27, which reaches 40% at the
end of this episode. This is paralleled by a decrease in C31 and indicates
an expansion of deciduous trees and, to a lesser extent, conifers,
which predominate over grasses. This finding is noteworthy between
meters 11.0 and 10.0 and between meters 7.5 and 7.0. These per-
centages mark the establishment of humid conditions, which were
probably associated with temperate-cold phases.
To better observe the characteristics of the last part of the Padul
record, we analyzed in detail the uppermost 20 m through the non-
smoothed profiles (Fig. 6). Although some of these features are observed
in the smoothed curves, there is evidence of drier episodes around
meters 12. 6 (ca. 66 ka B.P.), 10.2 (ca. 48 ka B.P.), 9.0 (ca. 39 ka B.P.), 7.7
(ca. 28 ka B.P.) and 6.8 (ca. 23 ka B.P.), which can be tentatively
correlated with Heinrich Events 6, 5, 4, 3 and 2, respectively. Heinrich
%n-C31 Events 6 and 5 seem to be less pronounced than the others.
− 0.8527
p = 0.000
4.11. Dry Episode 6 (meters 7.0–4.0; ca. 25 ka B.P.–ca. 10 ka B.P.)
p = 0.000
Considerable changes linked to the Last Glacial Maximum and the
beginning of the Holocene have been detected (meters 7–4.5) in other
palaeoenvironmental proxies (cf. Ortiz et al., 2004), i.e., samples with
lower δ13Corg values coincide with high atomic C/N ratios, atomic H/C
ratios between 1.3 and 1.7 and predominant n-alkane chains of 31
carbon atoms represent cold-dry phases. These caused the recession
of temperate forests and the extension of grasses in samples from
meters 6.9–6.7; 5.9–5.6 and 4.6. After these periods, both temperature
and precipitation increased.
There is good correspondence between these results and the
oscillations observed in the relative percentages of the C27, C29 and C31
profiles. In fact, the transition from Humid Episode 5 to meters 7–4 is
characterized by an abrupt decrease in the relative concentration of
C27, accompanied by a marked increase in the percentage of C31. We
propose that these findings represent a retreat of deciduous trees and
the spread of non-arboreal vegetation, which, in turn was produced
by a sudden dryness that coexisted with the cold conditions
associated with the end of the last glacial episode.
In our view, the minima of the non-smoothed profile observed at
meters 5.9 (ca. 18 ka B.P) and 4.6 (ca. 12 ka B.P) (Fig. 6) can be
correlated with Heinrich Event 1 (Last Glacial Maximum) and the
Younger Dryas, respectively. After these short periods, both temper-
ature and precipitation recovered quickly, causing the expansion of
deciduous trees and a rise in water level, with a major production of
lacustrine algae. These latter episodes are also characterized by
samples with intermediate C/N ratios and δ13Corg values as well as by
H/C ratios characteristic of phytoplankton or trees (cf. Ortiz et al.,
2004), and predominant C27 or C29 n-alkanes. Similarly, a wetter
episode occurred (decrease in the percentage of C31 and increase of
C27) between Heinrich Events 2 and 1, at ca. 20 ka B.P. (meter 6.7),
which was also interpreted by the low C/N and high δ13Corg values (cf.
Ortiz et al., 2004).
4.12. Holocene (uppermost 4.0 meters; ca. 10 ka B.P.–ca. 4.5 ka B.P.)
According to Ortiz et al. (2004), after the Last Glacial Maximum and
the Younger Dryas, both temperature and precipitation recovered,
causing the expansion of temperate forests and a rise in water level, with
a major production of lacustrine algae. These latter episodes are
characterized by samples with intermediate C/N ratios and δ13Corg
values as well as by H/C ratios characteristic of phytoplankton (also from
trees). Similarly, the composition of n-alkanes in the uppermost 4 m (ca.
10 ka B.P. to 4.5 ka B.P.) is dominated by long-chain isomers (n-C25 to n-
C35) with odd chain-length predominance, with few exceptions (meters
2.1, 2.0, 1.6, 1.5, and 0.5) where the phytoplankton-derived C17 is the
most abundant alkane, thereby indicating periodic increases in the
water level.
On the basis of the relative percentages of high molecular weight
n-alkanes, similar conclusions are reached. The beginning of the
Holocene (uppermost 4 m) is reflected by a wetter phase (increase in
the percentage of C27 and a decrease in the percentage of C31), which
produced the recovery of deciduous trees and retreat of grasses,
followed by an expansion of conifers at the expense of the former two
vegetation types from meter 2 to the top of the Padul record (Fig. 4).
A drier episode characterized by a dramatic drop in the percentage
of C27, which occurred at meter 2.5 (ca. 7.5–8 ka B.P.), is highlighted
(Fig. 6).
 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
Fig. 6. n-Alkane composition profile as percentages of C27, C29, and C31 isomers with respect to the sum C27 + C29 + C31 along the upper 20 m of the Padul borehole with the
palaeoenvironmental interpretation (He: Heinrich Events; YD: Younger Dryas).
5. Comparison with the Guadix-Baza Basin record
Given the proximity of the Guadix-Baza (ca. 60 km westwards) to
the Padul Basins, we attempted to make a correlation between
sequences. The Guadix-Baza Basin is an intramontanous closed
depression filled by alluvial and lacustrine sediments where a high-
resolution palaeoclimatic record from a 356 m-thick section was
obtained. The fluctuations observed in the oxygen and carbon stable
isotope measurements performed on shells of the ostracode Cyprideis
torosa along this section reveal changes in temperature, the evapora-
tion/infill ratio in the water bodies, the amount of rain, and plant
biomass vegetation from ca. 2 Ma to ca. 280 ka B.P. (Ortiz et al., 2006).
There is a general good correspondence between the smoothed δ18O
curve obtained in the Guadix-Baza Basin and the smoothed profiles of
the relative percentages of C27, C29 and C31. Four Cold and Humid Great
Periods alternating with four Warm and Dry Great Periods were defined
in the Guadix-Baza record (Fig. 7), representing long-term episodes of
relative climatic stability. Below we compare the two sequences:
– The end of the 2nd Warm and Dry Great Period of the Guadix-Baza
Basin has its correspondence in the 1st Dry Period of the Padul
Basin.
– The 3rd Cold and Humid Great Period of the Guadix-Baza Basin is
correlated with the 1st Humid Period of the Padul Basin.
– The 3rd Warm and Dry Great Period of the Guadix-Baza Basin is
correlated with the 2nd Dry Period of the Padul record.
– The 4th Cold and Humid Great Period of the Guadix-Baza Basin is
correlated with the 2nd Humid Period of the Padul record.
– The 4th Warm and Dry Great Period of the Guadix-Baza Basin is
characterized by marked oscillations, and all of these have their
correspondence with both the 3rd Dry and the 3rd Humid Episodes
in the Padul record:
– The warm and dry phase linked to the first maximum in the
smoothed δ18O curve from Guadix-Baza can be correlated with
the 3rd Dry Episode of the Padul Basin.
– The cold and humid phases related to the two minima of the
smoothed δ18O profile can be correlated with the humid episodes
linked to the two minima of the smoothed % C27 profile observed
within the 3rd Humid Period of the Padul record (or the two
295
maxima of the smoothed C31 profile), dated at ca. 340 and
270 ka B.P.
– The warm and dry period linked to the last maxima of the
smoothed δ18O profile can be correlated with the more humid
conditions interpret form the last maxima of the smoothed %C27
profile observed in the 3rd Humid Period of the Padul record (or
the minima of the smoothed C31 profile).
This comparison shows that in the Padul Basin drier episodes were
accompanied by warmer conditions, while wetter episodes coexisted
with colder phases (Fig. 7). In some cases, temperate and intermediate
precipitation conditions occurred. However, during the cold Younger
Dryas and Heinrich Events aridity increased, as shown by abrupt
decreases in the percentage of C27 and marked increases in that of C31
(Fig. 6). These findings are consistent with the pollen sequences from
marine cores drilled off the coasts of Spain and Portugal (Sánchez Goñi
et al., 2002; Roucoux et al., 2005), in which the Heinrich Events and the
Younger Dryas are characterized by a contraction of temperate tree
populations and the expansion of steppe vegetation.
The terms “colder-humid” and “warmer-drier” should be under-
stood as the conditions typical of the Mediterranean region, coinciding
with those proposed by Horowitz (1989, 2001) for Israel, which are as
follows: Pluvial (Wet Mediterranean), with lower temperatures, higher
rainfall (moderate rains in winter and summer), and the development of
deciduous oak forest; Interpluvial (Dry Mediterranean), with rare
precipitation, higher temperatures and steppe vegetation; and Inter-
stadial, like present-day conditions, with a short rainy winter and a hot,
dry summer, with evergreen oaks and Mediterranean maquis.
Thus, in the Padul Basin, Interglacial and Interstadial Periods are
characterized as dry, while during Glacial and Stadial Periods humid
conditions prevailed. This palaeoenvironmental pattern differs to those
found in central and Northern Europe records. This discrepancy could be
attributed to the geographical location of the Padul Basin, which is found
in the Mediterranean realm which falls between Northern Europe and
Africa, thus making it a unique area for the study of the paleoclimatic
evolution during the Pleistocene. The influence of Mediterranean and
North Atlantic dynamics on the temperature and precipitation in
Mediterranean Iberia caused different climatic effects to those in northern
Europe (cf. Pons and Reille, 1988; Valero-Garcés et al., 1998, 2000; Millán
296 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299

Fig. 7. Tentative correlation between Padul Basin and Guadix-Baza Basin sequences and the marine oxygen isotopic record (Shackleton, 1995). The Guadix-Baza Basin record consists
on the smoothed curve of δ18O values obtained in Cyprideis torosa ostracodes with the paleoenvironmental periods identified by Ortiz et al. (2006). The data of the marine oxygen
isotope record derive from the SPECMAP stack for the 0–0.62 Ma interval and from OPD site 677 for the 0.62–2.0 Ma interval.

et al., 2005; Ortiz et al., 2006). Glacial Periods, which are characterized by
cold and dry conditions with scarce liquid water and the development of
permafrost (ever-frozen soils) at higher European latitudes, produced
humid conditions in the Mediterranean realm (Horowitz, 1989, 2001;
Zanchetta et al., 1999; Ortiz et al., 2006). In contrast, Interglacial and
Interstadial Periods produced warmer and wetter conditions in northern
Europe than in southern European latitudes.
Thus, in Central and Northern Europe, forests were established
during Interglacial and Interstadial Periods and steppe plants during
Glacial and Stadial Periods (cf. de Beaulieu and Reille, 1992; Reille
and de Beaulieu, 1995; Urban, 1997; Reille et al., 1998, 2000; Thieme,
1999) because of their proximity to the margin of the European ice
sheet. In contrast, the inverse occurred in the southern Iberian
Peninsula.
 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
The decrease in tree abundance and the expansion of grasses
during Heinrich Events reflect the presence of very cold and arid
phases in the southern Iberian Peninsula, which is in accordance with
reports by Pailler and Bard (2002), Sánchez Goñi et al. (2002) and
Roucoux et al. (2005). In fact, Heinrich Events are associated with
massive discharges of icebergs in the North Atlantic (Heinrich, 1988;
Bond et al., 1992, among others), which affected the western coast of
the Iberian Peninsula through the deposition of ice-rafted debris
produced by iceberg melting (Zahn, 1997; Cayre et al., 1999; Cacho
et al., 2001; Roucoux et al., 2001, 2005). Furthermore, there is
evidence of polar water incursions in the Mediterranean through the
strait of Gibraltar during Heinrich Events (Cacho et al., 1999, 2000;
Pérez-Folgado et al., 2003). These incursions produced decreases in
the surface temperature of the Mediterranean Sea and caused these
episodes to be much drier, as indicated by greater expansion of steppe
vegetation (Sánchez Goñi et al., 2002; Roucoux et al., 2005;
Chondrogiani et al., 2004).
Good correspondence is also observed between the Padul Basin
record and MIS (Fig. 7), with odd MIS being related to Dry Episodes
and even MIS to Humid Episodes.
6. Comparison with previous palaeoenvironmental studies of the
Padul Basin
We provide a general comparison of our data with the pollen
sequences of this basin reported by Florschütz et al. (1971) and Pons
and Reille (1988) (Fig. 5). It has to be taken into account that there is a
certain gap between the lithology of the cores (cf. Nestares and Torres,
1998). Furthermore, Pons and Reille (1988) analyzed with criticism
some of the interpretations of Florschütz et al. (1971), indicating
either inconsistency between their diagram and their interpretation,
or discrepancy between results from the two records.
Likewise, several considerations must be taken into account: the
Padul Basin acted as an integrative system of vegetation debris (plant
derived biomarkers) and lixiviates from the degradation of plant
tissue from the Sierra Nevada Mountain Range, which holds a series of
vegetation belts (Florschütz et al., 1971), comprising from bottom to
top: steppe, Quercus ilex (more humid), Quercus pubescens (xero-
morphic), Acantholimon-Tragacanta (Eastern Mediterranean charac-
ter) and Mediterranean mountain steppe.
There is good correspondence between our data and the pollen
diagrams of Pons and Reille (1988) obtained along the upper 24.9 m
sedimentary sequence that comprises the Upper Pleistocene (ca.
100 ka B.P.) and Holocene. According to their biostratigraphical
interpretation, the last Glacial cycle was generally characterized in
Padul by temperate conditions, as evidenced by the presence of
deciduous Quercus and other trees, together with Erica arborea, and
poorly shown climatic fluctuations. Likewise, the percentage of
Artemisia is low, and corresponds to the Humid Period 5 defined in
this paper. However, a number of phases of considerable aridity were
observed (especially within their palynozones P2k, P3h, and P3l),
which Pons and Reille (1988) correlated with the EoWürm (MIS 4), the
Older Dryas and the Younger Dryas, respectively. Similarly, our data
indicate that during the last ca. 100 ka B.P. temperate and humid
conditions prevailed (Humid Episode 5), although more periods
characterized by clearly marked cold and dry conditions, associated
with Heinrich Events and the Younger Dryas, appear. On the basis of
our results, we propose that n-alkane profiles reflect more oscillations
than pollen analyses (Fig. 6), three of them being correlated with the
arid palynozones of Pons and Reille (1988), corresponding palyno-
zones P2k and P3h to the Heinrich Events 6 and 1, respectively (Figs. 5
and 6).
Considering a difference of about 3 m in core length between cores
(cf. Nestares and Torres, 1998), good correlation (Fig. 5) was also found
between our sequence and the 70 m-long pollen diagram of Florschütz
et al. (1971), ranging, according to our chronological model, from ca.
297
480 ka B.P. to the Holocene and not from the Lower Pleistocene as
Menéndez Amor and Florschütz (1964) proposed. Episodes in which
the relative percentage of C27 with respect to the sum of the C27, C29
and C31 homologues is higher, indicating more humid conditions, the
percentage of arboreal pollen increases (Florschütz et al., 1971)
(Fig. 5), whereas when the relative percentage of C31, assumed to be
linked to greater presence of grasses at the expense of deciduous trees,
and therefore to drier conditions, the non-arboreal pollen together
with Artemisia reach higher percentages. We wish to highlight the
correlation between the sharp decrease of the arboreal pollen in the
palynological record of Florschütz et al. (1971) and the low percentage
of the C27 n-alkane (Dry Period 5) in our study, both indicating dry
conditions that are possibly related to MIS 5. Nevertheless, we do not
totally agree with some of their interpretations, i.e., palynozones G, I, L,
and N are characterized by the domination of steppe plants (Artemisia
and Chenopodiacea) and are correlated with Dry Episodes 2, 3, 4 and 5,
as defined in this paper (lower percentages of C27 and higher of the C31
homolog). However, Florschütz et al. (1971) attributed them to cold
conditions, while, according to our data, after the comparison with the
Guadix-Baza Basin record and those from Israel, these palynozones
could be associated with warmer temperatures.
Similarly, according to Pons and Reille (1988), palynozone Q of
Florschütz et al. (1971) should be included in the last Glacial Period
and not in the Eemian Interglacial Period, and, therefore the previous
palynozone N, which reflects steppe expansion, and the first part of
palynozone Q (with and important Artemisia content) should be
correlated with our Dry Episode 5.
Palynozone L and the upper part of palynozone K are correlated
with Dry Episode 4, whereas the lower and middle part of palynozone
K and the whole of palynozone J correspond to Humid Episode 3
(Fig. 5). Palynozone J reflects humid conditions, but the lower part of
palynozone K is characterized by the predominance of steppe plants
(Florschütz et al., 1971) and could correspond to the decrease in the
relative percentage of C27 located between 53 and 52 m of our record.
Palynozones M and H, showing a certain expansion of trees, have a
clear correspondence with Humid Episodes 2 and 4.
The gradual decrease in arboreal pollen from palynozone S to
palynozones R and Q is correlated with the continuous descent of the
relative percentage of C27 in Humid Episode 5. Palynozones X6, X4, X2, V,
and T were considered to have been produced by cold and arid
conditions as the percentage of Artemisia and Chenopodiaceae increases.
Although these palynozones are correlated with Humid Episode 5, they
coincide with some of the Heinrich Events distinguished within this
palaeoenvironmental episode in Padul (Fig. 6).
Finally, the uppermost palynozone X (especially X8) and palyno-
zone Y, which reflects arid conditions (Older and Younger Dryas) at
the end of the last Glacial Period, has a good correspondence with Dry
Episode 6. Palynozone Z is correlated with the Holocene.
7. Conclusions
Here we demonstrate that the relative percentages of the high
molecular weight n–alkanes (C27, C29 and C31) measured in a 107 m–
long core are useful to reconstruct the ancient vegetation and
palaeoclimatological evolution of the Padul Basin from the Lower
Pleistocene (ca. 1 Ma) to the mid–Holocene (ca. 4.5 ka B.P.). Thus, here
we have amplified the knowledge of the palaeoclimatological evolution
of the Padul Basin along the Pleistocene, especially in some spans where
the global climatic changes were not recorded in other organic proxies
(cf. Ortiz et al., 2004).
The variations observed in the profiles of the high molecular
weight n-alkanes provided evidence of changes in vegetation which,
in turn, were related to climate oscillations, i.e., in dry scenarios
forests receded while grasses developed. In contrast, during humid
phases trees expanded at the expense of grasses.
298 J.E. Ortiz et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 298 (2010) 286–299
Twelve main climatic periods linked to dry/humid scenarios were
distinguished: 6 Dry Episodes alternating with 5 Humid Episodes
were interpreted in the Pleistocene record, together with a final
episode covering the Holocene. On the basis of the correlation with
the Guadix-Baza Basin record, we propose that dry phases coexisted
with warm conditions, whereas humid periods were linked to
temperate-colder conditions, except during the last Dry Episode of
the Pleistocene. In this episode, which coincided with the end of the
Last Glacial Period, low temperatures concurred with dry conditions.
This palaeoclimatic pattern in the southern Iberian Peninsula
shows that environmental conditions differed from those of Northern
Europe, with Glacial Periods characterized by cold and humid
conditions in the southern Mediterranean area, while Interglacial
Periods produced warm and dry climates.
Our analysis of the last 100 ka B.P. indicates rapid changes in
vegetation produced by short-scale climatic cycles linked to Heinrich
Events and the Younger Dryas.
Our results were confirmed by comparing them with those of pollen
analyses performed in the Padul Basin by Florschütz et al. (1971) and
Pons and Reille (1988). In some cases the n-alkanes provide enhanced
information about palaeoenvironmental changes, as Heinrich Events are
better observed.
Likewise a good correspondence was observed with the MIS defined
in marine cores. Taken together, our findings indicate that vegetation in
the Padul Basin responded to long-term cycles (orbital scale) and
millennial-scale global cycles and, thus, climatic changes responded to
global climatic changes.
Acknowledgements
We are especially indebted to Mr. Garrido who allowed us to drill
the core in his property and provide many facilities. Funding was
obtained through the projects “Evidence from Quaternary Infills
Palaeohydrogeology” (European Union, F14W/CT96/0031), “Evolu-
ción Paleoclimática de la Mitad Sur de la Península Ibérica” of ENRESA
(National Company for Radioactive Waste Management, 703238) and
“Paleoclima” of ENRESA and CSN (Spanish Nuclear Safety Council).
We want to thank Prof. Shinya Yamamoto (Hokkaido University) and
an anonymous reviewer for their constructive and helpful comments.
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