# Climate change and ecosystems dynamics over the last 6000 years in the Middle Atlas, Morocco

Majda Nourelbait, Ali Rhoujjati, Abdelfattah Benkaddour, Matthieu Carré, Frédérique Eynaud, Philippe Martinez, Rachid Cheddadi

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Majda Nourelbait, Ali Rhoujjati, Abdelfattah Benkaddour, Matthieu Carré, Frédérique Eynaud, et al.. Climate change and ecosystems dynamics over the last 6000 years in the Middle Atlas, Morocco. Climate of the Past, 2016, 12 (4), pp.1029-1042. 10.5194/cp-12-1029-2016. hal-03063918

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# Climate change and ecosystems dynamics over the last 6000 years in the Middle Atlas, Morocco

Majda Nourelbait<fnref n="1" />,<fnref n="2" />,<fnref n="3" />, Ali Rhoujjiat<fnref n="2" />, Abdelfattah Benkaddour<fnref n="2" />, Matthieu Carré<fnref n="3" />, Frederique Eynaud<fnref n="4" />,
Philippe Martinez<fnref n="4" />, and Rachid Cheddadi<fnref n="3" />

*Correspondence to:* Majda Nourelbait (nourelbait.m@gmail.com)

Received: 25 July 2015 – Published in Clim. Past Discuss.: 1 September 2015

Revised: 25 March 2016 – Accepted: 29 March 2016 – Published: 21 April 2016

**Abstract.** The present study aims at reconstructing past climate changes and their environmental impacts on plant ecosystems during the last 6000 years in the Middle Atlas, Morocco. Mean January temperature ($T_{jan}$), annual precipitation ($P_{ann}$), winter ($P_w$) and summer ($P_s$) precipitation, and a seasonal index (SI) have all been quantified from a fossil pollen record. Several bio- and geo-chemical elements have also been analysed to evaluate the links between past climate, landscape, and ecosystem changes.

Over the last 6000 years, climate has changed within a low temperature and precipitation range with a trend of aridity and warming towards the present. $T_{jan}$ has varied within a ca. 2 °C range, and $P_{ann}$ within less than 100 mm yr$^{-1}$. The long-term changes reconstructed in our record between 6 ka cal BP and today are consistent with the aridity trend observed in the Mediterranean basin. Despite the overall limited range of climate fluctuation, we observe major changes in the ecosystem composition, the carbon isotopic contents of organic matter ($\delta^{13}$C), the total organic carbon and nitrogen amount, and the carbon to nitrogen ratio (C / N) after ca. 3750 cal BP. The main ecosystem changes correspond to a noticeable transition in the conifer forest between the Atlas cedar, which expanded after 3750 cal BP, and the pine forest. These vegetation changes impacted the sedimentation type and its composition in the lake.

Between 5500 and 5000 cal BP, we observe an abrupt change in all proxies which is coherent with a decrease in $T_{jan}$ without a significant change in the overall amount of precipitation.

## 1 Introduction

The amplitude of climate change during the Holocene (11 700 cal BP to the present) is known to be globally less extreme than during the last glacial period (Bianchi and McCave, 1999; Bond et al., 2001; Debret et al., 2007). However, several studies have shown that there were climate fluctuations (Alley et al., 1997; Wanner et al., 2008) related to the internal variability of the climate system, solar activity, albedo (Ruddiman, 2003; Eddy, 1982; Stuiver et al., 1991), volcanic eruptions (Kelly and Sear, 1984; Sear et al., 1987; Bryson, 1989; Mann et al., 2005), ocean circulation (Manabe and Stouffer, 1988; Dansgaard et al., 1989; Lascaratos et al., 1999; Rohling et al., 2002), etc., which all have a direct impact on the terrestrial ecosystems (Davis, 1963; Emmanuel et al., 1985). Although climate changes were less pronounced during the Holocene (Andersen et al., 2004; Mayewski et al., 2004; Witt and Schumann, 2005; Frigola et al., 2007; Cheddadi and Bar-Hen, 2009), they have still been noticeable enough to be recorded by different proxies (Dorale et al.,

<footnote n="1">¹Université Chouaib Doukkali, Laboratoire Géosciences Marines et Sciences des Sols, Unité Associée CNRST (URAC45), El Jadida, Morocco</footnote>

<footnote n="2">²Université Cadi Ayyad, FST, laboratoire de Géoressources, Unité Associée CNRST (URAC42), Gueliz Marrakech, Morocco</footnote>

<footnote n="3">³Université Montpellier 2, Institut des Sciences de l'Evolution, UMR UM2-CNRS-IRD5554, Montpellier, France</footnote>

<footnote n="4">⁴University of Bordeaux, UMR EPOC5805, CS50023, 33615 Pessac, Bordeaux, France</footnote>
<figure><img src="image_3.png" /><figcaption>**Figure 1.** The study area. (a) Geographical location of the tabular and pleated Middle Atlas (MA); (b) sketch of the geological and geomorphological characteristics of the Hachlaf area (from Martin, 1973); (c) phytoecological map showing the main ecosystems and the location of Lake Hachlaf (Dayet Hachlaf) within an oak forest (from Lecompte, 1969).</figcaption></figure>

1992; Williams et al., 2002; Geiss et al., 2003, 2004). At the global scale, the Holocene climate stability allowed sustainable vegetation dynamics with long-term ecosystem changes, plant species expansions and migrations, and an increase of species diversity over all latitudes (Rohde, 1992). However, the Holocene period has also recorded some abrupt and cold events such as the one at 8.2 ka cal BP (e.g. Alley and Agustsdottir, 2005) which recorded a depletion of about 4 °C in winter temperature in the eastern Mediterranean (Weninger et al., 2009).

In Morocco, climate changes during the Holocene have also been quantified, and they show significant fluctuations (Cheddadi et al., 1998). As a matter of fact, the climate variability of the Holocene is less known than that of the last glacial (Mayewski et al., 2004) because it has a lower amplitude and is less abrupt. This statement is even more acute in the Mediterranean region where high-resolution and chronologically well-constrained Holocene records are much less numerous than in Europe or North America. The Mediterranean area is currently a hotspot of biodiversity (Myers et al., 2000) and it is one of the largest regions in the world that undergo long-lasting and pronounced droughts during the summer season (Roberts et al., 2004; Milano et al., 2013). The southern rim of the Mediterranean region is even more

arid than the northern one because of the influence of the Azores high and the Saharan winds which increase the impact of the drought effect during the summer season. Most of the winter precipitation ($P_w$) originates from the trade winds which carry moisture from the Mediterranean Sea (Martin, 1981). The amount of $P_w$ has a strong impact on the persistence of water bodies and on the lake levels in the Mediterranean area. Strong lake level fluctuations during the Holocene were observed in Lake Van, Turkey (Lemcke and Sturm, 1997); Lago Dell'Accesa and Lago di Mezzano, Italy (Magny et al., 2006); Lake Kinneret (Hazan et al., 2005) and the Dead Sea, Israel (Migowski et al., 2006); Lake Siles, Spain (Carrión, 2002); and lakes Sidi Ali and Tigalmamine in Morocco (Lamb and Van der Kaars, 1995; Märsche-Soulié et al., 2008).

The analysis of marine and continental records from the central part of the Mediterranean shows that the lake levels were high between 10 300 and 4500 cal BP due to an enhanced moisture availability during both summer and winter (Magny et al., 2013). After 5000 cal BP, pollen data from southwestern Europe show that drought increased and led to a sustained reduction of the forest cover (Roberts et al., 2001; Jalut et al., 2009; Jiménez-Moreno et al., 2015). These environmental changes show that within the long-term climate trend there were humid-arid episodes that are related to internal forcings of the climate system such as, in the case of these westernmost Mediterranean ecosystems, the centennial changes in the North Atlantic Oscillation modes (Jiménez-Moreno et al., 2015), the enhancement/weakening of the trade winds, or the increase in the coastal upwelling off northwestern Africa (McGregor et al., 2007).

Climate reconstructions from marine pollen records suggest that the Mediterranean environments may react with a reduced time lag to rapid climate changes (Fletcher et al., 2010). The response of the western Mediterranean ecosystems has even been synchronous with the North Atlantic variability during the last glacial period and the Holocene (Combourieu-Nebout et al., 2009). Changes in the pollen assemblages of a marine record from the Alboran Sea also show very synchronous fluctuations between the surrounding land ecosystem changes and the sea surface temperature fluctuations (Fletcher and Sánchez Goñi, 2008; Combourieu-Nebout et al., 2009). Pollen records from the Middle Atlas (Reille, 1976; Lamb and Van der Kaars, 1995; Cheddadi et al., 2009; Rhoujjati et al., 2010; Nour el Bait et al., 2014; Tabel et al., 2016) and the Rif Mountains (Cheddadi et al., 2016) show that the Holocene climate change had a major impact on the ecosystems composition with a clear succession of different species sensitive to winter frost, strong rainfall seasonality, and/or the total amount of annual rainfall throughout the year.

The aim of the present study is to evaluate the impacts of the climate changes on the ecosystems and the landscape of the Middle Atlas during the last 6 millennia. Our approach is multidisciplinary and based on the analysis of pollen grains,
<table><caption>**Table 1.** Radiocarbon ages for the Hach-I core. Calibrations were performed using Libib 7.1 (Stuiver and Reimer, 1986).</caption><thead><tr><th>Lab number</th><th>Depth (cm)</th><th>Material dated</th><th><sup>14</sup>C age yr BP</th><th>95.4 % (2δ) cal age ranges (BP)</th><th>Relative area under probability distribution</th><th>Median probability cal BP</th></tr></thead><tbody><tr><td>Poz-20175</td><td>60</td><td>Bulk</td><td>2535 ± 30</td><td>2494–2746</td><td>0.447</td><td>2624</td></tr><tr><td>Poz-20177</td><td>120</td><td>Bulk</td><td>3220 ± 35</td><td>3371–3509</td><td>0.936</td><td>3436</td></tr><tr><td>Poz-20178</td><td>170</td><td>Bulk</td><td>4390 ± 35</td><td>4859–5047</td><td>0.991</td><td>4949</td></tr><tr><td>Poz-20179</td><td>240</td><td>Bulk</td><td>5200 ± 40</td><td>5897–6021</td><td>0.943</td><td>5958</td></tr></tbody></table>

elemental and isotopic geochemistry, and grain size from a fossil record collected in Lake Hachlaf, Middle Atlas. Temperature and precipitation variables have been quantified. They show a moderate change which is superimposed by an aridity trend that is combined with an increase in winter temperature over the past 6000 years. We also observed some noticeable ecosystem and landscape changes with one rapid and quite abrupt climate fluctuation between 5500 and 5000 cal BP detectable in all the proxies used.

## 2 Study area

The Middle Atlas Mountains, lying in northwestern Morocco, consist of two geological sets called pleated and tabular Middle Atlas (Fig. 1a). The latter is formed by a Paleozoic basement covered by a Mesozoic thick layer and Cenozoic and Quaternary volcanic flows (Texier et al., 1985; Herbig, 1988; Harmand and Moukadiri, 1986). The Liasic limestone and dolostone are shaped by karstic mechanisms (Martin, 1981; Baali, 1998; Hinaje and Ait Brahim, 2002; Chillasse and Dakki, 2004). In this geomorphological and structural composition, there exist nowadays about 20 permanent or semi-permanent natural lakes (Chillasse and Dakki, 2004) among which we can find the studied site, Dayet (Lake) Hachlaf (33°33′20″ N; 5°0′0″ W; 1700 m a.s.l.). This small water body is located about 10 km north-east of Ifrane National Park (Fig. 1b). Available meteorological data (HCE-FLCD, 2004) at Dayet Hachlaf show an average annual rainfall of ca. 600 mm with $P_w$ and $P_s$ ca. 150 and ca. 70 mm, respectively. The mean January temperature is ca. 4 °C with ca. 90 rainy days per year and ca. 70 frosty days, among which ca. 17 are with snow precipitation. The surface area and depth of the lake change throughout the year, reaching up to respectively 14 ha and 4 m during late spring. The lake is fed by rainwater, snow, surface runoff, and groundwater and has no river inflow.

The forest cover around the site (Fig. 1c) is composed of holm oak (Q. ilex subsp. rotundifolia), which is evergreen; zeen oak (Q. canariensis), which is deciduous; and Atlas cedar (Cedrus atlantica) with occurrences of *Pinus halepensis*. Nowadays, there are some degraded populations of *Cedrus atlantica* with cultivated lands around the lake. At higher altitude (1700 to 2500 m, Fig. 1c) an

herbaceous/shrubby vegetation (Artemisia herba-alba and Poaceae) dominates the landscape.

## 3 Materials and methods

In April 2008, a 2.5 m core (33°33′2.49″ N, 4° 59′41.57″ W) was collected using a Russian corer. Each section of the core was then sub-sampled for the analysis of pollen content (30 samples), grain size (39 samples), organic matter (43 samples) and its isotopic composition (δ¹³C<sub>(org)</sub>; 46 samples), and total nitrogen and carbonates (43 samples).

Pollen grains were extracted using a standard laboratory procedure: HCl (20 %), KOH (10 %), ZnCl₂, acetolysis (CH₃CO₂O and H₂SO₄), KOH (10 %), ethanol, and glycerine. The identification and counting of pollen grains were performed with an optical microscope (Leica DM750) using a × 40 magnification (× 63 for accurate identifications). The pollen percentages were calculated on the total sum of pollen grains originating from vascular terrestrial plants. The total of pollen grains counted varies between ca. 200 and 1300. Aquatic plant percentages (including Cyperaceae and Junceaceae) were excluded from the total pollen sum. Cyperaceae were considered as aquatic plants since there are *Juncus* and *Cyperus* genera growing around the lake today.

The particle size analysis was carried out at the Laboratoire Marocain d'Agriculture (LABOMAG) and was only performed on the sediment fraction < 2 mm. The proportions of five fractions were identified as follows: coarse sand (2000–200 µm), fine sand (200–50 µm), coarse silt (50–20 µm), fine silt (20–2 µm), and clay (below 2 µm).

Organic matter amount (OM) was estimated based on the content of the organic carbon in lacustrine sediments (OC), elaborated by spectrometry (NF ISO 14235, 1998). Sediment OC was oxidized in a sulfochromic environment with an excess of potassium dichromate at 135 °C. Subsequently, the determination of chromate ions $Cr^{3+}$ formed was analysed by spectrometry.

For total nitrogen (TN), the method used was based on the Kjeldahl mineralization (ISO 11464: 1994, 2006), but the catalyst used was the titanium dioxide (TiO₂). The technique consists in assaying the total nitrogen content in the sediment as ammonium, nitrate, nitrite, and organic form.
Carbonates were measured by adding HCl to the bulk sediment to decompose all carbonates (NF ISO 10693, 1995). The volume of the carbonic gas produced was measured using a Scheibler apparatus.

Stable isotope ratio measurements of carbon were performed on a Thermo Fisher FLASH 2000 Elemental Analyzer in line with a VG Isoprime Mass Spectrometer at the University of Bordeaux. All samples were pretreated with 1N HCl to remove inorganic carbon. The analytical precision of 0.15‰ was estimated from several calibrated laboratory standards analysed along the samples. Stable isotopic ratios were reported as δ¹³C = [(¹³C / ¹²Csample / ¹³C / ¹²Cstd) – 1]×1000, where the standard used is Vienna Pee Dee Belemnita (PDB)

Besides the multi-proxy analysis, four organic samples were dated. All these dates have been done on bulk sediment. We used the BACON software (Blaauw and Christen, 2011) to compute the age-depth model (Fig. 2). The default ¹⁴C calibration curve used by BACON for terrestrial Northern Hemisphere samples is IntCal13. The AMS ¹⁴C dates were also calibrated using the “CALIB 7.1” program (Stuiver and Reimer, 1986; Table 1). The fossil record continuously encompassed the last 6000 years.

Annual precipitation ($P_{ann}$), mean January temperature ($T_{jan}$), and precipitation seasonal index (SI) assessment (Fig. 3) were based on pollen data as follows:

$$PSI(s) = \sum P_w - \sum P_s / \sqrt{P_{ann}},$$

where $PSI(s)$ is the seasonal index quantified for sample s; $P_w$ is the sum of December, January, and February precipitation; $P_s$ is the sum of June, July, and August precipitation; and $P_{ann}$ is the total annual precipitation.

The monthly mean precipitation and $T_{jan}$ were obtained using the probability density function of modern plant species (pdf method). This method is described in Chevalier et al. (2014). In order to apply it to a fossil pollen record collected in the Mediterranean area, it required a modern database of Mediterranean plant species distributions and their corresponding modern climate variables. We used a database of plant species that have been georeferenced from *Flora Europaea* (Jalas et al., 1972, 1973, 1976, 1979, 1980, 1983, 1986, 1989, 1991, 1994) and Hulten and Fries (1986). Additional geographical distributions were obtained from GBIF (2012) and personal field observations using GPS in Morocco. In order to use plant species distributions for the pollen-based climate reconstruction, we assigned pollen taxa to the most probable plant species in our plant database (Table 2). The modern climate variables were extracted from the WORLDCLIM database (Hijmans et al., 2005) and interpolated onto the species occurrences for inferring their pdfs.

## 4 Results

During the last 6000 years, the main change in the forest cover is marked by a decline of the pine populations, the expansion of Atlas cedars after 3750 cal BP, and the persistence of the evergreen oaks. Although the latter dominate today the landscape around Lake Hachlaf, the microscope identification of the fossil pollen grains that originate from deciduous or evergreen plants may often be dubious and, therefore, may not be reproducible by another pollen analyst. We have assigned all oak pollen grains to the evergreen *Quercus ilex* in the climate reconstruction since it is the species that dominates the landscape and its climate envelope encompasses that of the other evergreen species (Fig. 4). All other taxa – including trees, shrubs, and herbs – also show some changes but within a much lower range than that of the two conifer taxa, Atlas cedar and pine (Fig. 5). We have applied a constrained cluster analysis to depict the main changes in the pollen fossil record. There are four main clusters summarizing the main changes in the ecosystem composition around Lake Hachlaf over the last 6000 years (Table 3).

The grain size analysis revealed the presence of three fractions (Fig. 3) with the following proportions: clay (22.87%), silt (60.46% with 41.9% of fine silt), and sand (16.67%). The dominant silt fraction tends to increase from the bottom to the top of the core after a brief decline between ca. 5600 and 5200 cal BP. The sandy fraction follows the same pattern. Clay shows an opposite trend to both the sandy and silt fractions.

Carbonate (CaCO₃) content is high throughout the record except around 5200 cal BP (Fig. 3). They are positively correlated with silt and sand. The total organic carbon (TOC) content is also high and varies significantly between 4 and 27.4% (Fig. 3). The TN remains low throughout the record. The carbon-to-nitrogen ratio (C/N) varies between 9 and 17.4, and the δ¹³Corg between –21 and –27‰ (Fig. 3). Two origins of the organic matter are thus identified, with lake algae characterized by C/N < 11 and very depleted δ¹³Corg and terrestrial plants characterized by C/N > 11 and less depleted δ¹³Corg (Fig. 6). δ¹³Corg and C/N are positively correlated (Fig. 3). TOC and TN are highly correlated (0.99, Figs. 3 and 6) as well.

In order to interpret the different bio- and geo-chemical proxies within a climatic frame, a pairwise correlation was performed between the three climate variables and δ¹³C, C/N, TN, and TOC (Fig. 7). Although there could be no causal relationship, SI and $T_{jan}$ are well correlated with each another. They are both correlated negatively with δ¹³C and C/N and positively with TN and TOC (Fig. 7).

## 5 Discussion

The Holocene climate around the Mediterranean Sea was suitable for the expansion of human populations and their organization towards true civilizations (Kaniewski et al., 2012).
<figure><img src="image_4.png" /><figcaption>**Figure 2.** (a) Lithology of the core Hach-I and radiocarbon ¹⁴C dates; (b) age–depth model from BACON software (Blaauw and Christen, 2011).</figcaption></figure>

The persistence and longevity of many Mediterranean populations may be linked to the relative suitability and also to an overall stability of the Holocene climate. However, climatic events have been recorded within the Holocene (e.g. Rohling and Pälike, 2005) and a causal relationship has been made between some abrupt climatic events and societal changes in the Mediterranean (Berger and Guilaine, 2009; Kaniewski et al., 2008).

In the present study, we have focused on the environmental and climate changes that occurred during the last 6 millennia in the northern part of the Moroccan Middle Atlas Mountains. We have evaluated the vegetation dynamics using the palynological content of a fossil sequence and analysed its bio- and geo-chemical content to reconstruct the overall landscape changes.

The reconstructed $T_{jan}$ and $P_{ann}$ show a relatively low amplitude of change over the last 6000 years (Fig. 3). $P_{ann}$ decreases progressively by ca. 100 mm, which is in line with the aridity trend that has been observed in other fossil records (Risacher and Fritz, 1992; Brooks, 2006; Hastenrath, 1991; Anderson and Leng, 2004; Umbanhowar et al., 2006) and particularly in the Mediterranean area (Pons and Reille, 1988; Julià et al., 2001; Burjachs et al., 1997; Yll

et al., 1997; Roberts et al., 2001; Valino et al., 2002; Jalut et al., 2009) and northern Africa (Ritchie, 1984; Ballouche, 1986; Lamb et al., 1989). At a more regional scale, reconstructed $P_{ann}$ is coherent with that obtained from Lake Tigalmamine (Cheddadi et al., 1998) which shows a decreasing trend over the last ca. 5000 cal BP. The arid trend observed after ca. 5 ka cal BP is marked by a spread of Poaceae and a progressive replacement of pines by Atlas cedars, which better stand the high seasonal contrast of precipitation at the altitude of Lake Hachlaf. SI increased from 3 to 7 times over the last 6000 years (Fig. 3). A study of drought thresholds influencing the growth and photosynthesis was performed on different cedar stands and species (*C. atlantica*, *C. libani*, *C. brevifolia*, and *C. deodara*) of different origins (Aussenac and Finkelstein, 1983). This study showed that, among many conifers, cedar trees may keep a sustained photosynthesis activity even when drought is very high. Thus, a strong precipitation contrast between $P_s$ and $P_w$ (Fig. 3) may not affect the Atlas cedar overall growth as long as the total amount of rainfall is sufficient (higher than 600 mm yr⁻¹) and the winter temperature is low enough (below 6 °C) for the vegetative cycle (Aussenac et al., 1981). The Mediterranean climate is known for its strong seasonal distribution of precipitation
<figure><img src="image_5.png" /><figcaption>**Figure 3.** Diagram showing the sediment fractions (clay, silt, and sand/silt ratio), the pollen percentages of *Cedrus atlantica* and *Pinus*, geochemical elements (delta 13 C [δ¹³C‰], nitrogen-to-carbon ratio [C/N], total organic carbon [TOC], total nitrogen [NT]) and carbonates concentrations (CaCO₃), January mean temperature (*T*<sub>jan</sub>), annual precipitation (*P*<sub>ann</sub>), winter and summer precipitations (*P*<sub>w</sub> and *P*<sub>s</sub>), and precipitation seasonality index (SI). The red rectangles are pointing the values of present-day *T*<sub>jan</sub>, *P*<sub>ann</sub>, *P*<sub>w</sub>, and *P*<sub>s</sub> (HCEFLCD, 2004); the red line shows the limit 3.7 ka cal BP; and the blue rectangle shows the time interval of the cold phase 5.2 ka cal PB.</figcaption></figure>

<table><caption>**Table 2.** Pollen taxa assigned to the most probable plant species in our plant database.</caption><thead><tr><td>Pollen taxa</td><td>Plant species</td></tr></thead><tbody><tr><td>Alisma</td><td>Alisma plantago-aquatica</td></tr><tr><td>Alnus</td><td>Alnus glutinosa</td></tr><tr><td>Berberis</td><td>Berberis hispanica</td></tr><tr><td>Brassica</td><td>Brassica</td></tr><tr><td>Campanula</td><td>Campanula afra</td></tr><tr><td>Caryophyllaceae</td><td>Caryophyllaceae</td></tr><tr><td>Centaurea</td><td>Centaurea cyanus</td></tr><tr><td>Chenopodiaceae</td><td>Chenopodiaceae</td></tr><tr><td>Asteroideae</td><td>Compositae subfam. Asteroideae</td></tr><tr><td>Cichorioideae</td><td>Compositae subfam. Cichorioideae</td></tr><tr><td>Corylus</td><td>Corylus avellana</td></tr><tr><td>Cupressaceae</td><td>Cupressaceae</td></tr><tr><td>Ephedra</td><td>Ephedra fragilis</td></tr><tr><td>Euphorbia</td><td>Euphorbia characias</td></tr><tr><td>Geranium</td><td>Geranium macrorrhizum</td></tr><tr><td>Helianthemum</td><td>Helianthemum canariense</td></tr><tr><td>Ilex</td><td>Ilex aquifolium</td></tr><tr><td>Juglans</td><td>Juglans regia</td></tr><tr><td>Myriophyllum</td><td>Myriophyllum aquaticum</td></tr><tr><td>Plantago</td><td>Plantago lanceolata</td></tr><tr><td>Polygonaceae</td><td>Polygonaceae</td></tr><tr><td>Ranunculaceae</td><td>Ranunculaceae</td></tr><tr><td>Salix</td><td>Salix pedicellata</td></tr><tr><td>Saxifraga</td><td>Saxifraga</td></tr><tr><td>Taxus</td><td>Taxus baccata</td></tr><tr><td>Urtica</td><td>Urtica dioica</td></tr><tr><td>Papaveraceae</td><td>Papaveraceae</td></tr><tr><td>Pinus</td><td>Pinus halepensis</td></tr><tr><td>Olea</td><td>Olea europaea</td></tr><tr><td>Paronychia</td><td>Paronychia argentea</td></tr><tr><td>Erica</td><td>Erica arborea</td></tr><tr><td>Quercus</td><td>Quercus ilex</td></tr><tr><td>Cedrus</td><td>Cedrus atlantica</td></tr><tr><td>Artemisia</td><td>Artemisia herba-alba</td></tr></tbody></table>

throughout the year. Summers are fairly dry, and most of the annual precipitation occurs during the cold months (end of autumn and beginning of winter).

Currently, 75% of the Moroccan territory with a grassy or wooded vegetation (thus excluding the desert) records between 500 and 800 mm of annual rainfall with an SI between 5 and 8 (Fig. 8). The whole range of SI in Morocco is between -1 in areas where $P_{ann}$ is less than 100 mm with a random distribution for instance in the south of Morocco and 15 in areas where the annual rainfall is quite high (over 800 mm) and occurs mainly in the winter season such as in the Rif Mountains today (Fig. 8). SI is higher in mountainous areas. Nowadays, in the areas surrounding Lake Hachlaf (located at ca. 1600 m elevation) SI is around 5. This SI has changed over the past 1000 years as confirmed, at least between 6000 cal BP and today, by the studied fossil archive (Fig. 3). The amplitude between $P_{w}$ and $P_{s}$ precipitation has
<table><caption>**Table 3.** Pollen zones identified in the fossil record using a constrained cluster analysis. AP: arboreal pollen taxa; NAP: non-arboreal pollen taxa; DT: taxa diversity.</caption><thead><tr><th>Zones</th><th>Depth (cm)</th><th>Age (cal BP)</th><th>Pollen data description</th></tr></thead><tbody><tr><td rowspan="3">Zone I</td><td rowspan="3">250–190</td><td rowspan="3">6227–5171</td><td>AP 27–60 % – Mainly <i>Quercus</i> and <i>Olea</i>.<br/>– Peak of <i>Pinus</i> (47 %)<br/>at 6100 cal BP then decreasing.<br/>– Low percentages of <i>Cedrus atlantica</i> with<br/>initial spread around 5800 cal BP.</td></tr><tr><td>NAP 39–72 % – Herbs dominated by Poaceae (11–48 %),<br/><i>Illecebrum</i> (3–19 %), Apiaceae (2–5 %),<br/>Brassicaceae (1–5 %), Asteraceae (0–5 %),<br/>Cichorioideae (1–6 %), Chenopodiaceae (0.5–2 %)<br/>and cereals (0–1 %).</td></tr><tr><td>DT 18–26 – Rapid fluctuations.</td></tr><tr><td rowspan="3">Zone II</td><td rowspan="3">190–111</td><td rowspan="3">5171–3651</td><td>AP 28–56 % – <i>Pinus</i> dominates the pollen record but regresses<br/>at 5500 cal BP (from 44 to less than 2 %).<br/>– <i>Cedrus atlantica</i> continues to expand (0–5 %).<br/>– We observe a peak of Rosaceae (6 %).</td></tr><tr><td>NAP 43–72 % – Herbs are dominated by Poaceae, <i>Illecebrum</i>,<br/>and Asteraceae which reach their maximum<br/>(53, 20 and 10 %, respectively).<br/>– Cereals disappear.</td></tr><tr><td>DT 19–29 – Moderate to high with two peaks.</td></tr><tr><td rowspan="3">Zone III</td><td rowspan="3">111–60</td><td rowspan="3">3651–2351</td><td>AP 23–58 % – Strong expansion of <i>Cedrus atlantica</i><br/>and <i>Quercus</i>.<br/>– An abrupt decline of <i>Cedrus atlantica</i><br/>around 2653 cal BP is recorded.<br/>– <i>Pinus</i> regresses as well but shows a peak<br/>of 20 % at 3300 cal BP.</td></tr><tr><td>NAP 41–76 % – Herbs dominate the pollen record.<br/>– Sharp decline in Poaceae, Asteraceae,<br/>Chenopodiaceae, and Caryophyllaceae at 5600 cal BP.<br/>– Appearance of cereals around 2653 cal BP.</td></tr><tr><td>DT 20–31 – High.</td></tr><tr><td rowspan="3">Zone IV</td><td rowspan="3">60–5</td><td rowspan="3">2351–173</td><td>AP 23–43 % – Abundance of <i>Cedrus atlantica</i>,<br/><i>Quercus</i>, <i>Olea</i>, and Rosaceae.<br/>– Sharp decline and disappearance of <i>Pinus</i>.</td></tr><tr><td>NAP 56–76 % – Herbs continue to dominate the pollen record, with<br/>Poaceae, cereals, Brassicaceae, Chenopodiaceae, and<br/>Caryophyllaceae being the most abundant.<br/>– Asteraceae, <i>Illecebrum</i>, and Apiaceae decline.<br/>– Centaurea and Cichorioideae disappear.</td></tr><tr><td>DT 21–32 – High.</td></tr></tbody></table>

increased 2 to 3 times towards the present (Fig. 3). Since $P_{ann}$ has a decreasing trend, the opposite increased seasonality is related to a significant reduction in the amount of rainfall during the months of June, July, and August (Fig. 3). This strengthening of the contrast between $P_w$ and $P_s$ had a rather limited impact on the dominating taxa because they can with-

stand the summer drought and the overall amount of $P_w$ remained sufficient for their persistence. However, a change in the amplitude of SI has probably favoured those species best adapted to the length of the dry season, for instance evergreen oaks rather than deciduous. Pollen-based climate reconstructions from records collected in the Alboran Sea
<figure><img src="image_6.png" /><figcaption>**Figure 4.** Density plots of $T_{jan}$ for the three dominating *Quercus* species in Morocco (*Q. ilex*, *Q. coccifera*, and *Q. suber*). The median values are as follow: *Q. ilex* – 6.5 °C; *Q. coccifera* – 7.4 °C; and *Q. suber* – 7 °C.</figcaption></figure>

(Combourieu-Nebout et al., 2009) and Italy (Magny et al., 2013; Peyron et al., 2013) suggest a rather steady and low seasonal contrast between $P_w$ and $P_s$ (about 2 times) over the past 6000 years. This discrepancy between the reconstructed SI from Hachlaf and the marine record may potentially be related to the fact that marine records collect pollen grains from a much wider geographical source area than continental (mountainous) records, which probably tends to smooth the local/regional changes. The reconstructed seasonality from the Italian records (Magny et al., 2013; Peyron et al., 2013) is buffered by the less abrupt precipitation seasonal contrast at European temperate latitudes than at arid Mediterranean ones.

SI was lower than 5 before 3750 cal BP despite an amount of precipitation between 600 and 700 mm yr⁻¹ (Fig. 3). During that period, water probably persisted in the lake all throughout the year, which allowed the presence of aquatic plants (Fig. 5) flowering during late spring and summer, and algae identified in the pollen data, through the low values of $\delta^{13}C_{org}$ and the C/N ratio being greater than 11 (Figs. 3 and 5). The proportion of aquatic plants cannot be directly related to a high lake level and may not be used to state the lake level changes but only the presence of water in the site. The $\delta^{13}C_{org}$ and C/N (Fig. 6) provide information concerning the origin of the organic matter (in situ production versus input from the catchment area) but not on the lake level changes. Thus, high $\delta^{13}C_{org}$ and C/N ratios (Fig. 3) with low presence of aquatic plants (Fig. 5) may not be inconsistent in cases where there is a low terrestrial input (low sand/silt, Fig. 3) during a period when the lake level is high.

<figure><img src="image_7.png" /><figcaption>**Figure 5.** Diagram showing the percentages of the main pollen taxa identified in the Hachlaf core. Cyperaceae and Juncaceae are included within aquatic taxa. The dashed black curves show an exaggeration (×7) of the percentages of some taxa. On the right, pollen zones with their boundaries are set up using a constrained hierarchical clustering (R Development Core Team, 2013). The taxonomic diversity is computed using a rarefaction analysis. The red line shows the limit 3.7 ka cal BP.</figcaption></figure>
C3 metabolism plants

<figure><img src="image_8.png" /><figcaption>**Figure 6.** δ¹³C and C/N bi-plot (from Meyers, 1994).</figcaption></figure>

<figure><img src="image_9.png" /><figcaption>**Figure 7.** Pairwise correlation between the three climatic variables ($T_{jan}$, $P_{ann}$, and SI) and the chemical elements.</figcaption></figure>

The relationship between δ¹³Corg and the C/N ratio indicates the occurrence of two main types of organic matter mainly originating from a C3 metabolism. Lacustrine algae can be considered as dominantly autochthonous; in the lower part of the record, the organic matter, with higher C/N ratios and less depleted δ¹³Corg, corresponds to a terrestrial input. Indeed, fresh organic matter from lake algae is known to be protein-rich and cellulose-poor with molar C/N values commonly between 4 and 10, whereas vascular land plants are protein-poor and cellulose-rich, creating organic matter usually with C/N ratios of 20 and greater (Meyers, 1994, 2003). However, a C/N ratio > 11 may correspond to a mixture of both local and terrestrial organic matter (Fig. 6).

After 3750 cal BP, Atlas cedars noticeably spread around the site, while the pine populations strongly regress. A series of fossil pollen records in the Middle Atlas show that Atlas cedar populations expanded after ca. 6 ka cal BP. The sus-

<figure><img src="image_10.png" /><figcaption>**Figure 8.** Modern SI (upper panel) and $P_{ann}$ (middle panel) from the gridded WorldClim data set (Hijmans et al., 2005) over Morocco. The lower panel shows the distribution of $P_{ann}$ vs. SI: the lowest index occurs in southern Morocco, where $P_{ann}$ is lower than 200 mm yr⁻¹, and the highest index occurs in the high-altitudinal areas (Middle Atlas and Rif Mountains).</figcaption></figure>

tained expansion of Atlas cedar after ca. 3750 cal BP around Lake Hachlaf expresses its late occurrence at higher altitude. Around Lake Tigalmamine (Lamb et al., 1995), the Ras El Ma marsh (Nour El Bait et al., 2014), and the Ait Ichou marsh (Tabel et al., 2016), which are all located at about 100 to 200 m altitude below Lake Hachlaf (ca. 1700 m a.s.l.), Atlas cedar occurs much earlier. The expansion of Atlas cedar around the lake is probably related to both an upslope spread and a south-north migration.

During this ecosystem transition we observe a major change in both $P_{ann}$ and $T_{jan}$. The increase of SI after 3750 cal BP is due to a combined increase of $P_w$ and decrease of $P_s$ (Fig. 3). The expansion of cedar forests in the studied area may be related to their better adaptation to strong SI than pines at higher altitude.

Competition is another parameter that might be worth considering. After 3750 cal BP, the C/N ratio is below 11 and the δ¹³C remains below −26‰, which suggest the important primary productivity of the lake associated with low input of land-plant-derived organic matter. Atlas cedar forests have a more important growth in both height and diameter than pines, which leads to a higher biomass production. This is linked to the genetic model of growth that is very distinct between the two taxa (Kaushal et al., 1989). Thus, the ex-
pansion of the Atlas cedar population around the site may
explain the high input of OM into the lake.

Over the last 6 millennia, superimposed to the overall
climate trend, we observe one relatively abrupt event be-
tween 5500 and 5000 cal BP during which Tjan declined by
about 2°C compared to its average over 6000 years. A cli-
matic transition between 6 and 5 ka cal BP at the end of
the Holocene thermal maximum has been globally identified
(Steig, 1999; Mayewski et al., 2004; Wanner et al., 2008;
Brooks, 2012). This transition has been recorded by a wide
range of climate proxies (e.g. Kaufman et al., 2004; Jansen
et al., 2009; Seppä et al., 2009; Bartlein et al., 2011) and has
been related to different biosphere feedbacks and potentially
to a decay of the remaining Laurentide Ice Sheet (Renssen
et al., 2009). All proxies from the Hachlaf sequence as well
as the reconstructed climate variables have recorded marked
changes during that period of time. SI has the lowest value of
the record, and a succession of abrupt changes are recorded
in the C / N ratio, the grain size fractions, the δ¹³C, TN, TOC,
and CaCO₃ (Fig. 3). Carbonates, considered as a “paleo-
thermometer” (Meyers, 1994, 2003), also decrease abruptly
around 5200 cal BP (Fig. 3). The latter may be linked to a low
evaporation of the lake which may have been favoured by low
winter temperature around 5200 cal BP. The fine grain size
sediment also increased as a consequence of low seasonal
precipitation contrast and/or a continuous sediment input to
the lake. Such sustained input of clay and decreasing car-
bonate content suggest a higher lake level between 5500 and
5000 cal BP (Fig. 3). Thus, the Tjan and SI decrease may have
contributed to the higher lake level or at least to the presence
of water throughout the year (Fig. 3). At the same time, the
sand-to-silt ratio is very low, which confirms a low energy
during the sedimentation process. The major change in the
ecosystem composition around the lake is the rapid collapse
of the pine forest which has inevitably released an important
amount of terrestrial carbon (biomass) into the lake (positive
peaks in δ¹³C and C / N, Fig. 3).

6 Conclusions

This study marks a new contribution to the knowledge of past
climates and environmental history in north Africa moun-
tainous areas. The range of climate change in the Middle
Atlas, Morocco, was rather minor between 6000 cal BP and
the present. Annual precipitation and January mean temper-
ature have respectively varied within a range of 100 mm and
2 to 3 °C. However, they both show a trend towards a more
arid and warmer climate as well as a higher rainfall season-
ality. Pann became as contrasted as today after 3750 cal BP.
The aridity trend observed in Hachlaf over the last 6000
years is consistent with other climate reconstructions avail-
able from other Mediterranean fossil records. Besides these
overall climatic trends, we also observe an abrupt cold event
between 5500 and 5000 cal BP which is well marked in all

environmental proxies from our studied fossil record. The
δ¹³C and C / N ratios, which are well correlated with each
other, suggest an increase in the organic matter input from
the catchment area. Concomitantly, the pollen record indi-
cates a decline of the pine forest which may have contributed
to the organic matter input into the lake too. The marked
change in both the carbonate content and clay composition
of the record was probably related to a perennial presence of
water throughout the year. Synchronously, seasonality index
and January mean temperature were the lowest of the record
which has contributed to a reduction of the evaporation.

The increase in rainfall seasonality has probably favoured
the expansion of Atlas cedars around the studied site at the
expense of the pine forest.

**Acknowledgements.** This work was supported by the Volubilis
Programme (Programme mixte Interuniversitaire Franco-marocain,
MA/11/251), 2011; by CNRS-CNRST Convention, 2009
(ScVie07/09); and the French national programme EC2CO-
Biohefect, “Variabilité paléoclimatique et impact sur les forêts de
conifères au Maroc depuis la période glaciaire”. M. Nourelbait
received a postdoc grant from the EU Framework Programme
Erasmus Mundus EU METALIC II (2013-2442/001-001 – EMA2)
for completing this study at ISEM. We thank Claire Grandchamps
for her revision of the English version of the manuscript. This is
ISEM contribution no. 2016-020.

Edited by: N. Combourieu Nebout

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