# Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs

Marie-Pierre Ledru, Raquel Franco Cassino, Vaneicia dos Santos Gomes, Julia

Caram Sfair, Francisca Soares Araújo

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Marie-Pierre Ledru, Raquel Franco Cassino, Vaneicia dos Santos Gomes, Julia Caram Sfair, Francisca Soares Araújo. Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs. Acta Botânica Brasílica, 2022, 10.1590/1677-941x-abb-2022-0111. hal-04893635

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# Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs

Marie-Pierre Ledru<fnref n="1" />* ID, Raquel Franco Cassino<fnref n="2" /> ID, Vaneicia dos Santos Gomes<fnref n="3" />, <fnref n="4" /> ID, Julia Caram Sfair<fnref n="4" /> ID
and Francisca Soares Araújo<fnref n="4" /> ID

Received: May 01, 2022

Accepted: July 12, 2022

## ABSTRACT

The vegetation cover is the result of many biological and abiotic interactions. To identify the different factors that cause changes is crucial when defining future sustainable development and protection of natural resources. In the Brazilian semiarid region, the vegetation cover has been subjected to drastic deforestation and land use at centennial-scale that has led to desertification. Pollen analysis is an efficient tool to reconstruct the different processes of degradation of the vegetation cover over time. We built a referential data set for the vegetation cover using 48 pollen surface samples collected in the reservoirs of the Ceará. We used satellite images for comparison with the pollen signatures and defined an alteration score to express the correlation between terrestrial pollen and anthropic cover. Our results showed our surface samples to be generally representative of the vegetation cover and of the general degradation of the landscape. Our study areas can be considered as degraded as the initial categories “preserved” or “intermediary” are not reflected in the pollen assemblages, in agreement with results of botanical surveys and the historical background. The on-going process of desertification is climate-independent and was initiated many decades or centuries ago by intensive land use for agriculture and grazing.

**Keywords:** palynology, calibration, tropical dry forest, human impact, Brazilian semiarid region, degradation.

## Introduction

Modern vegetation cover is a result of many environmental and physical factors. Disentangling climatic from anthropogenic forcings on vegetation and biodiversity is a priority when planning reforestation and/or conservation policies.

Long-term reconstruction of past agricultural activity is based on anthropogenic indicators, for instance *Zea mays* pollen grains or phytolith (Piperno & McMichael 2020), found in sediment cores. Quantitative reconstruction of human-induced landscapes from fossil pollen data in recent decades has enabled the development of a theoretical framework and of models of pollen-vegetation relationships

<footnote n="1">1 Institut des Sciences de l'Evolution Montpellier, Université de Montpellier CNRS IRD EPHE, Place Eugène Bataillon cc061, 34095 Montpellier, France.</footnote>

<footnote n="2">2 Departamento de Geologia, Universidade Federal de Ouro Preto, 35400-000, Ouro Preto, MG, Brazil.</footnote>

<footnote n="3">3 Faculdade de Educação, Ciências e Letras do Sertão Central, Universidade Estadual do Ceará, 63902-098, Quixadá, CE, Brazil.</footnote>

<footnote n="4">4 Programa de Pós-Graduação em Ecologia e Recursos Naturais, Departamento de Biologia, Universidade Federal do Ceará, 60455-760, Fortaleza, CE, Brazil.</footnote>

<footnote>* Corresponding author: marie-pierre.ledru@ird.fr</footnote>
at the Holocene time scale. However, these studies have primarily focused on temperate ecosystems in the Northern Hemisphere aided by the rich archeological background (e.g. Marquer et al. 2017; Harrison et al. 2020). In the Brazilian semiarid region (BSA), the vegetation cover has been subjected to drastic deforestation and land use at centennial scale and the lack of historical archives makes it impossible to fully evaluate the successive states that led to desertification in some areas (see map of the FUNCEME in CGEE 2016). Even though we cannot ignore the possibility that hunter-gatherer societies may also have impacted the landscape, anthropogenic and paleo studies showed that they were keen on preserving their resources (Bliege et al. 2008; Morrison et al. 2022). In the BSA region, over the last three centuries, extensive agriculture has been practised for commercial purposes with little concern for the sustainability of the resources. Analyzing historical changes in land use/land cover is essential to understand current human-environment dynamics and more specifically, the different steps that lead to desertification in a poor but densely populated region. There is a drastic lack of historical or natural archives to estimate changes in the landscape at millennial or decadal scales in tropical semiarid regions (Flantua et al. 2015). The absence of historical records is due to the fact that transmission was mostly oral, but also logical given that paleoecological reconstructions require permanent lakes or swamps for the preservation of the biological indicators. Changes in the landscape in a highly anthropized region of the BSA region, Vale do Acaraú, Ceará, were recently reconstructed from sediment cores collected in reservoirs (Ledru et al. 2020). They showed that at decadal scale, droughts do not affect the vegetation due to the resilience of the ecosystem whereas public policies have had a strong impact on the composition of the vegetation and biodiversity. To calibrate the responses of the vegetation to disturbances, pollen analysts usually infer modern pollen rain studies (Chevalier et al. 2020). However, the lack of such studies in the Caatinga biome (Flantua et al. 2015), the dry deciduous forest of the BSA, limited our interpretation (Ledru et al. 2020). In addition, when a relationship exists between vegetation and its associated pollen production/deposition, such relationships are usually analyzed in areas that have undergone little disturbance (e.g. Amaral et al. 2006; Gosling et al. 2009; Montade et al. 2016). Moreover, when dealing with mosaics, different vegetation (e.g. biomes) and land use (e.g. farming) conditions in space and over time means systematic approaches cannot be used (Deza-Araujo et al. 2020).

Soil samples or samples from pollen traps are usually used to build referential data sets. However, pollen traps were not used in the BSA region because they require regular collection over several months or years before a representative data set is obtained (e.g. Portes et al. 2020). Another problem with soil sampling in semi-arid regions is

that high temperatures and low soil moisture prevent good preservation of the pollen grains in soil samples. However, in the BSA region, the large number of reservoirs built to retain the water during the long dry season created local permanent moisture conditions able to retain the pollen rain produced by the surrounding vegetation (Markgraf et al. 2002). The construction of reservoirs in Northeastern Brazil started in 1890 (Assunção & Livingstone 1993) and today there are more than 5,000 reservoirs from small to average capacity in the State of Ceará (COGERH). Before colonization, the tropical dry forest, Caatinga, covered 844,000 km², but was subsequently subjected to large-scale deforestation for wood extraction, farming and agriculture, and today a large area is under desertification (CGEE 2016; IPBES 2018). More than 40 % of the vegetation has been deforested (Overbeck et al. 2015), while protected areas account for only 9 % of the Caatinga region (Brasil 2000). Only 1.8 % of the Caatinga is fully protected (IUCN categories I to III), while 7.2 % is under sustainable use in which the direct use of natural resources is permitted according to Brazilian legislation (Brasil 2000). Given the current climate change and more specifically the predicted increase in temperature (IPCC 2018) it is urgent to characterize the degree of preservation required for the recovery and/or the protection of the Caatinga. In this context, we present a new data set built using reservoirs as potential pollen traps with the aim of defining the relation between vegetation degradation and pollen deposition and more particularly to characterize the degradation of the semi-arid vegetation using alteration scores.

Materials and methods

Modern vegetation and climate

The phytogeographical domain of the Caatinga is characterized by a hot semiarid climate (Bsh) according to Köppen's classification (Peel et al. 2007), mean annual temperature is 28 °C and mean annual precipitation ranges between 500 and 800 mm. The rainy season lasts from January to May, although with frequent inter- and intra-annual variability related to changes in the surface temperature of the tropical Atlantic and the El Niño Southern oscillation (Barbosa et al. 2006). For instance, 13 drought events have been recorded in the BSA region in the last 60 years (Marengo & Bernasconi 2015).

Two natural reserves in the crystalline Caatinga were created on ancient farming areas. Thirty-four botanical surveys were performed in the two reserves (Costa et al. 2007, Araujo et al. 2011, Moro et al. 2014). They showed the high frequency of the herbaceous layer among the plant communities. Among the 10 main herbaceous species were Melochia tomentosa (Malvaceae), Tacinga palmadora, Tacinga inamoena (Cactaceae), Mollugo verticillata (Molluginaceae),

<figure><img src="image_2.png" /></figure>
Neoglaziovia variegate (Bromeliaceae), Waltheria americana (Malvaceae), Alternanthera brasiliana (Amaranthaceae), Delilia biflora (Asteraceae), Sida galheirensis (Malvaceae), Euphorbia hyssopifolia (Euphorbiaceae). Among the main woody species are Aspidosperma pyrifolium (Apocynaceae), Commiphora leptophloeos (Burseraceae), Jatropha mollissima (Euphorbiaceae), Cereus jamacaru (Cactaceae), Myracrodruon urundeuva (Anacardiaceae), Anadenanthera colubrine (Fabaceae), Bauhinia cheilantha (Fabaceae), Mimosa tenuiflora (Fabaceae), Poincianella pyramidalis (Fabaceae), Cynophalla flexuosa (Capparaceae), Spondias tuberosa (Anacardiaceae), Piptadenia stipulacea (Fabaceae), Schinopsis brasiliensis (Anacardiaceae), Libidibia ferrea (Fabaceae), Combretum leprosum (Combretaceae), Ziziphus joazeiro (Rhamnaceae), Pilosocereus gounellei (Cactaceae), Croton sonderianus (Euphorbiaceae), Lantana camara (Verbenaceae), Croton heliotropiifolius (Euphorbiaceae), Amburana cearenses (Fabaceae) (Moro *et al.* 2014).

For this study, we selected 48 reservoirs located in the State of the Ceará, one of the driest semi-arid regions in Northeastern Brazil (Fig. 1, Tab. 1). The reservoirs are located within the crystalline basement where Caatinga vegetation grows in shallow, stony or fertile soils (Sampaio 1995). Our samples were collected in the top 2 cm of the soil-water interface in three different locations around the reservoir, at the end of the 2015 dry season. Studies performed in ponds showed that each sample represented between 5 and 20 years of pollen deposition (McLauchlan et al. 2007).

Pollen analyses

For each sample, we separated a volume of 2 cm³ for
pollen extraction following the method of Faegri & Iversen
(1989). Samples were treated with fluorhydric acid (HF
70%) for 12 hours to dissolve the silicate and silica, rinsed
with 50% HCl in a bain-marie at 80 °C to remove the silica
colloids, then with 10% KOH in a bain-marie at 80 °C to
remove the humic acids and finally acetolysis with a 9:1
preparation of acetic anhydride and sulfuric acid. A spike
(Lycopodium clavatum) was added to each sample to calculate
the concentration (Stockmarr 1971). The pollen grains
were identified and counted on microscope slides under
an optical microscope at x60 magnification. A total of 300
grains of arboreal and non-arboreal pollen were counted
in each sample, excluding aquatic and water-level related
pollen and spores taxa. Eight samples were discarded due
to their poor pollen content. Pollen grains were identified
with the help of the pollen reference collections belonging
to the Federal University of Ceará, Brazil, to the University
of Montpellier, France, pollen atlases and catalogs covering
different South American flora (e.g. Salgado-Labouriau
1973; Barth 1989; Silva 2007; Silva et al. 2016). Broken and
deformed pollen grains were classified as indeterminate.

Reconstruction of the vegetation cover

The vegetation cover at each sampling site was characterized according to Mapbiomas (3rd collection) classification where the terms forest, savanna, grassland only relate to the vegetation cover and not to other biomes. The Caatinga domain is represented by both forested and non-forested formations; the term forest implies a continuous vegetation cover, while the term non-forested means the vegetation is dominated by herbaceous species and floodable areas. At each site, the vegetation cover was described in three areas (with a radius of respectively, 300 m, 500 m, and 1 km) around the collection point, and vegetation data were converted into percentages (surface areas of the water bodies were excluded when calculating the percentage).

Statistical analyses

Pollen counts were also converted into percentages; the total pollen sum used to calculate the percentage is represented by the sum of all pollen taxa except aquatic or water-level related taxa (Agalinis, Cleome, Cuphea, Cyperaceae, Echinodorus, Ludwigia, Salvinia, Typha, Utricularia) (Matias et al. 2021). Pollen frequencies are presented in diagrams drawn with C2 software (Juggins 2007).

We defined an alteration score to express the correlation between the percentage of terrestrial pollen and the percentage of anthropic cover around the sites according to the following method: first, pollen types were classified in two groups: anthropic vegetation taxa and natural vegetation taxa (Tab. 2). Pearson correlation coefficients were then calculated between the percentage of the individual pollen taxa and the percentage of anthropic cover. Anthropic vegetation taxa that were positively correlated and natural vegetation taxa that were negatively correlated with the percentage of anthropic cover were then used to calculate the alteration score.

The alteration score was used to calculate a linear regression model to reconstruct the area of anthropic cover from pollen percentages. For model calculations, the dataset was randomly split into a training dataset containing 80 % of the samples, and a test dataset containing the remaining 20 %. Linear regression coefficients were calculated first using the training dataset; the remaining samples (20 %) were then used to test the model and to calculate prediction errors. Accuracy was measured by calculating mean absolute errors (MAE). The model was then applied to the complete dataset to compare the predicted and real percentages of land cover.

To test the application of our results to fossil pollen data, we applied the linear regression model to a 60 cm sediment core collected in the reservoir of Araras near the city of Varjota (Ceará, Brazil) (4°11'59"S, 40°27'59"W) dated between 1959 and 2002 AD. (Ledru *et al.* 2020).

<figure><img src="image_3.png" /></figure>
<figure><img src="image_4.png" /><figcaption>**Figure 1.** A. Map of South America showing the location of the State of Ceará. B. Map of the vegetation cover of the State of Ceará showing the location of the samples collected for this study. C, D, E, F, G. Zooms on the location of the samples and associated vegetation cover (Tab. 1).</figcaption></figure>
Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs

<table><caption>**Table 1.** List of the 48 samples analyzed in the present study with their location and information concerning the degree of preservation; D: degraded, P: preserved, IT: intermediate, and the nature of the surroundings.</caption><thead><tr><td>Sample number</td><td>Location</td><td>Name</td><td>State</td><td>Vegetation</td></tr></thead><tbody><tr><td>1</td><td>-3,503812<br/>-40,279428</td><td>Acaraú-mirim</td><td>IT</td><td>Neptunia sp., Cleome sp., Cryptostegia sp., Heliotropium sp., Tephrosia sp., Borreria sp., Nymphaeaceae, Angelonia sp., Attalea speciosa Mart. ex Spreng on the hilltop</td></tr><tr><td>2</td><td>-3,738653<br/>-39,720979</td><td>Saco Verde</td><td>IT</td><td>Open vegetation with abundant Prosopis juliflora (Sw.) DC. and Cordia oncocalyx Allemão</td></tr><tr><td>3</td><td>-3,740844<br/>-39,361434</td><td>Caixitoré</td><td>IT</td><td>Cultivated plants: coconut, banana, Azadirachta indica A.Juss., Cleome sp., Ipomeas sp., Chamaecrista sp., cattle breeding and aquaculture</td></tr><tr><td>4</td><td>-3,76721<br/>-39,771146</td><td>Mocó</td><td>D</td><td>Open vegetation with Cryptostegia madagascariensis Bojer ex Decne, Jatropha sp., Prosopis juliflora (Sw.) DC</td></tr><tr><td>5</td><td>-3,777572<br/>-40,500035</td><td>Jaibaras</td><td>D</td><td>Caatinga with 2 to 3 years preservation. treelets with Calotopsis procera (Aiton) W.T.Aiton, Heliotropium sp., Lamiaceae, Borreria on the margins</td></tr><tr><td>6</td><td>-3,799482<br/>-40,240663</td><td>Forquilha</td><td>D</td><td>Open vegetation<br/>Grass for fodder and beans</td></tr><tr><td>7</td><td>-3,883624<br/>-40,014361</td><td>Aracatiaçu</td><td>D</td><td>Open vegetation<br/>Cultivated area with corn, beans, grass, cattle-breeding</td></tr><tr><td>8</td><td>-3,943714<br/>-40,113189</td><td>EMASA</td><td>D</td><td>Open vegetation with Cleome sp. on the margins of the reservoirs, Senna alata (L.) Roxb.</td></tr><tr><td>9</td><td>-3,979371<br/>-40,481917</td><td>Jaibaras-Varjota</td><td>IT</td><td>Degraded Caatinga</td></tr><tr><td>10</td><td>-4,208846<br/>-40,453924</td><td>Araras</td><td>D</td><td>Open vegetation</td></tr><tr><td>11</td><td>-4,218465<br/>-40,067361</td><td>Edson Queiroz</td><td>IT</td><td>No habitation</td></tr><tr><td>12</td><td>-5,135178<br/>-40,769448</td><td>Sobradinho</td><td>D</td><td>Open vegetation with Nymphaeaceae, Trema sp., the palm Copernicia prunifera (Mill.) H.E.Moore on the margins and Licania, Anadenanthera colubrina (Vell.) Brenan, Cynophalla flexuosa (L.) J.Presl, Ziziphus joazeiro Mart., Calliandra spinosa Ducke in the landscape</td></tr><tr><td>13</td><td>-5,136582<br/>-40,766436</td><td>Crateus</td><td>D</td><td>Open vegetation</td></tr><tr><td>14</td><td>-5,382674<br/>-40,335292</td><td>Independência</td><td>D</td><td>Open vegetation</td></tr><tr><td>15</td><td>-5,408405<br/>-40,476955</td><td>Jaburu</td><td>D</td><td>Open vegetation.<br/>Libidibia ferrea (Mart. ex Tul.) L.P.Queiroz, Cnidoscolus quercifolius Pohl, Commiphora leptophloeos (Mart.) J.B.Gillett, Aspidosperma pyrifolium Mart. & Zucc., Ziziphus joazeiro Mat., Malvaceae, Jatropha sp., Prosopis juliflora (Sw.) DC., Parkinsonia aculeata L., Senna alata (L.) Roxb., Cleome sp., Piloscereus sp.</td></tr></tbody></table>
<table><caption>**Table 1.** Cont.</caption><thead><tr><td>Sample number</td><td>Location</td><td>Name</td><td>State</td><td>Vegetation</td></tr></thead><tbody><tr><td>16</td><td>-5,437099<br/>-40,15313</td><td>Independência - Tauá</td><td>D</td><td>Open vegetation, on the road margin</td></tr><tr><td>17</td><td>-5,292696<br/>-38,921363</td><td>Banabuiú</td><td>D</td><td>Open vegetation</td></tr><tr><td>18</td><td>-5,32387<br/>-38,510461</td><td>Poço Redondo</td><td>D</td><td>Inside a farm, deforested and burnt landscape</td></tr><tr><td>19</td><td>-5,383877<br/>-38,461477</td><td>Poço de Barro</td><td>D</td><td>Reservoir since 1950</td></tr><tr><td>20</td><td>-5,497965<br/>-38,450226</td><td>Castanhão. -areia</td><td>IT</td><td>Secondary vegetation</td></tr><tr><td>21</td><td>-5,555133<br/>-38,425808</td><td>Castanhão - lama</td><td>IT</td><td>Secondary vegetation</td></tr><tr><td>22</td><td>-5,610476<br/>-38,962084</td><td>Japão</td><td>P</td><td>Deciduous forest</td></tr><tr><td>23</td><td>-5,67738<br/>-38,915552</td><td>Sítio Pasta</td><td>IT</td><td>Degraded area</td></tr><tr><td>24</td><td>-5,69445<br/>-38,950995</td><td>Boqueirão 20</td><td>IT</td><td>Secondary vegetation. Reservoir built in 1915.</td></tr><tr><td>25</td><td>-5,701907<br/>-38,498357</td><td>Castanhão-BR</td><td>IT</td><td>Secondary vegetation</td></tr><tr><td>26</td><td>-5,732898<br/>-38,967316</td><td>Junco</td><td>D</td><td>On the road margins</td></tr><tr><td>27</td><td>-5,755312<br/>-38,959446</td><td>São José</td><td>D</td><td>Aquaculture. Mimosa tenuiflora</td></tr><tr><td>28</td><td>-5,814402<br/>-38,979705</td><td>Alfredo</td><td>IT</td><td>Commiphora leptophloeo s (Mart.) J.B.Gillett, Luetzelburgia auriculata (Allemão) Ducke, Mimosa sp., Cordia oncocalyx Allemão</td></tr><tr><td>29</td><td>-5,81586<br/>-38,959298</td><td>Caldeirões</td><td>IT</td><td>Secondary vegetation. Macrophytes and abundant Mimosa tenuiflora</td></tr><tr><td>30</td><td>-5,818772<br/>-38,967736</td><td>Açude 805</td><td>D</td><td></td></tr><tr><td>31</td><td>-5,824815<br/>-38,505206</td><td>Seixo</td><td>D</td><td>Invasives and Piptadenia sp.</td></tr><tr><td>32</td><td>-5,861444<br/>-38,588897</td><td>açude Estrada</td><td>D</td><td>Desert, exposed soil</td></tr><tr><td>33</td><td>-5,884668<br/>-38,877293</td><td>Jaguaribe 1</td><td>D</td><td></td></tr></tbody></table>
Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs

<table><caption>**Table 1.** Cont.</caption><thead><tr><td>Sample number</td><td>Location</td><td>Name</td><td>State</td><td>Vegetation</td></tr></thead><tbody><tr><td>34</td><td>-5,923215<br/>-38,851595</td><td>Jaguaribe 2</td><td>D</td><td></td></tr><tr><td>35</td><td>-5,953434<br/>-38,833031</td><td>Feiticeiro</td><td>D</td><td>Open vegetation. Ipomea sp., Cleome sp.</td></tr><tr><td>36</td><td>-5,954901<br/>-38,721432</td><td>Loca d'Agua</td><td>D</td><td>Open vegetation with exposed soil</td></tr><tr><td>37</td><td>-5,992088<br/>-38,629156</td><td>Guariba</td><td>D</td><td>Near the road</td></tr><tr><td>38</td><td>-6,274404<br/>-40,219854</td><td>Arneiroz</td><td>D</td><td>Open vegetation</td></tr><tr><td>39</td><td>-6,457772<br/>-40,121623</td><td>Aiuaba</td><td>D</td><td>Aspidosperma pyrifolium Mart. &amp; Zucc., Jatropha sp., Ipomea sp., Mimosa caesalpiniifolia Benth, Mimosa tenuiflora (Willd.) Poir., Helicteres sp., Cnidoscolos quercifolius Pohl.</td></tr><tr><td>40</td><td>-6,487218<br/>-40,135008</td><td>Murzela</td><td>IT</td><td>Preserved vegetation with Cnidoscolos quercifolius Pohl, Ziziphus, Anadenanthera colubrina (Vell.) Brenan, cactaceae, Licania rigida Benth., Aspidosperma pyrifolium Mart. &amp; Zucc., Crateva tapia L., Handroanthus sp., Astronium urundeuva (M.Allemão) Engl., Croton sp, Cenostigma pyramidale (Tul.) Gagnon &amp; G.P.Lewis, Cassia sp., Cynophalla flexuosa (L.) J.Presl, Clitoria sp., Pseudobombax marginatum (A.St.-Hil., Juss. &amp; Cambess.) A.Robyns</td></tr><tr><td>41</td><td>-6,515474<br/>-40,172484</td><td>Andressa</td><td>IT</td><td>One side deforested, the other side secondary vegetation with Jatropha sp., Anadenanthera colubrina (Vell.) Brenan, abundant Cyperaceae</td></tr><tr><td>42</td><td>-6,577638<br/>-40,165643</td><td>Autorama</td><td>D</td><td>Heliotropium sp., Anadenanthera colubrina (Vell.) Brenan, Aspidosperma pyrifolium Mart. &amp; Zucc., Schinopsis brasiliensis Engl., Croton sp.</td></tr><tr><td>43</td><td>-6,621212<br/>-40,126234</td><td>Tonico</td><td>P</td><td>Deciduous forest with Ipomea sp, Anadenanthera colubrina (Vell.) Brenan, Mimosa tenuiflora (Willd.) Poir., Croton sp, Ziziphus joazeiro Mart., Licania rigida Benth., Peltophorum dubium (Spreng.) Taub, Clitoria sp., Indigofera sp., Sida sp., Cleome sp.</td></tr><tr><td>44</td><td>-6,640991<br/>-40,119839</td><td>Boqueirão 45</td><td>P</td><td>Deciduous forest</td></tr><tr><td>45</td><td>-6,7268<br/>-40,08028</td><td>São Raimundo</td><td>D</td><td>30-year old reservoir</td></tr><tr><td>46</td><td>-6,728287<br/>-40,076116</td><td>Pista São Raimundo</td><td>IT</td><td>Nearby vegetation slashed and burned. On the other side of the road vegetation preserved for at least 25 years containing Ipomea sp., Cleome sp., Bidens sp.</td></tr><tr><td>47</td><td>-6,735128<br/>-40,102142</td><td>Catolezeiro</td><td>D</td><td>Secondary vegetation</td></tr><tr><td>48</td><td>-7,077822<br/>-39,495017</td><td>Cotia</td><td>D</td><td></td></tr></tbody></table>
<table><caption>**Table 2.** List of the pollen taxa used for the calculation of the alteration score.</caption><tr><td><strong>Anthropic indicators</strong></td><td>Asteraceae Liguliflorae, Mollunginaceae, Urticaceae, Amaranthaceae, <em>Acalypha</em>, <em>Cuphea</em>, Anacardiaceae, <em>Gomphrena</em>, <em>Sebastiania</em>-t, Convolvulaceae, <em>Spermacoce</em>, Acanthaceae, <em>Ipomea</em>, <em>Evolvulus</em>, Boraginaceae, <em>Agalinis</em>, <em>Orbignya</em>, Solanaceae, Gymnosperma, Lamiaceae, <em>Zea mays</em></td></tr><tr><td><strong>Natural vegetation</strong></td><td>Chloroleucon, Myracrodruon urundeuva, Cassia, Poincianella pyramidalis, Mimosa acutistipula, Pithecelobium, Cordia, Amburana cearensis, Piptadenia, Salvinia, Diodella, Mimosa caesalpinifolia, Alstroemeriaceae, <em>Ziziphus joazeiro</em>, Bignoniaceae, <em>Angelonia</em>-t, Himatanthus, <em>Copaifera</em>, Asteraceae</td></tr></table>

## Results

Modern vegetation and degree of anthropic alteration (MapBiomas data)

In 33 of the 48 sites, the percentage of anthropic cover within the 300 m radius was higher than the percentage of natural vegetation (Fig. 2). In most sites, anthropic cover was characterized by pasture and agriculture. Among the natural vegetation, savanna (intermediary vegetation) was the most abundant type of vegetation in most sites.

### Modern pollen assemblages

Among the 90 pollen taxa identified, 35 were arboreal pollen taxa (39 %); 36 non-arboreal pollen taxa (40 %); 10 indeterminate (11 %); 9 aquatic pollen taxa (10 %), all displaying good pollen preservation. The percentage of arboreal pollen taxa varied from 16 % to 67 %. The arboreal pollen taxa were dominated by Fabaceae, Anacardiaceae, Combretaceae, Apocynaceae and Euphorbiaceae. Fabaceae represented up to 64 % in sample 25, *Mimosa tenuiflora* was the dominant taxon in all the samples accounting for from 6 % to 37 %. The family Euphorbiaceae accounted for 41 % mainly by *Croton*. *Croton* was found in 19 samples at percentages of more than 1 % (between 2 % and 3 %) in samples 18, 30, 32, 33, 34, 47. Combretaceae was mainly represented by *Combretum* (5 % in sample 9), Anacardiaceae represented 10 % of pollen in sample 7 and 7 % in samples 14 and 16 (Fig. 3). Bignoniaceae (5 % in sample 47) and *Ziziphus joazeiro* in 7 samples were also observed but at lower percentages. Non-arboreal pollen taxa were present at high percentages (> 50 %) in all the samples (84.8 % in sample 36) except in samples 21 and 16 where they represented 35 %. The most abundant non-arboreal taxa were Poaceae and *Alternanthera*, which were both present in all samples at percentages ranging between 2 % (sample 4) and 30 % (sample 48), and 1 % (sample 37) and 41 % (sample 2), respectively. Other non-arboreal pollen taxa were dominated by Rubiaceae, Malvaceae and Asteraceae.

Pollen grains of Rubiaceae (*Borreria* and *Mitracarpus*) were present in all the samples except four (15, 38, 47, 41) at percentages ranging between 1 % and 28 %. Asteraceae was represented by the group of the “liguliflorae” 3 % in sample 4, and “tubiflorae” 19 % in sample 43. Malvaceae represented up to 23 % (sample 45) of non-arboreal pollen. The highest percentages of non-arboreal pollen taxa were *Alternanthera* (41 %), *Mitracarpus* (18 % in sample 18) and *Borreria* (15 % in sample 3). Convolvulaceae was represented by three taxa *Evolvulus*, *Ipomea* (6 % in sample 39) while *Jacquemontia* was only found at low percentages in a few samples. Poaceae ranged between 2 % and 29 % with some pollen grains of *Zea mays* observed in 23 samples, at percentages ranging between 1 % and 3 % (the latter in sample 43). Low percentages (0.3-4 %) of allochthonous taxa of gymnosperms (*Podocarpus* and *Pinus*) were observed in 22 samples, the highest percentages being 4 % in sample 20 and 3 % in sample 4 (Fig. 3). Among the taxa observed only in one sample were Meliaceae (1 % in sample 2), *Ipomea* (6 % in sample 39), *Jatropha* (1 % in sample 37), *Amburana cearensis* (2 % in sample 6), Boraginaceae (1 % in sample 12) and Polygalaceae (1.65 % in sample 37).

### Relationship between vegetation and pollen data

In the samples with high percentages of anthropogenic cover, pollen assemblages tended to present high frequencies of Amaranthaceae, *Acalypha*, *Gomphrena*, and Anacardiaceae (Fig. 4A-I). For Amaranthaceae, *Acalypha* and *Gomphrena*, we found a more significant correlation of pollen frequencies with the percentage of “pasture+agriculture” cover than with the total anthropic cover (i.e., including urban areas) (Fig. 4A, C, E compared with 4B, D, F). The best correlation was found between the percentage of Amaranthaceae and the percentage of “pasture+agriculture”, however, no pollen type alone was found to be a good predictor of the degree of anthropic cover. The total percentage of non-arboreal pollen, the total percentage of arboreal pollen, the AP/NAP ratio, and the Poaceae frequencies showed no correlation with the percentage of anthropic cover (Fig. 4I-L).
Estimated degradation of the Caatinga based
on modern pollen rain deposited in reservoirs

<figure><img src="image_5.png" /><figcaption>**Figure 2.** Land cover data (from Mapbiomas 2019) reconstructed at 1,000 m, 500 m, 300 m for each weir. Sites are organized according to latitude. In the legend, Forest, Savanna and Grassland refer to vegetation cover that is respectively, continuous, intermediary, and open.</figcaption></figure>
<figure><img src="image_6.png" /><figcaption>**Figure 3.** Diagram showing the percentages of the main taxa identified in the 48 surface samples. Green - arboreal pollen taxa; orange - non-arboreal pollen taxa; blue - aquatic or water-level related taxa. Sites are organized according to the degree of degradation delimited by horizontal black lines with from top to bottom: D: degraded, P: preserved, IT: intermediate.</figcaption></figure>
Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs

<figure><img src="image_7.png" /><figcaption>Figure 3. Cont.</figcaption></figure>
<figure><img src="image_8.png" /><figcaption>**Figure 4.** Relationship between pollen frequencies and the percentage of anthropic cover in the area with a radius of 300 m around the sites, considering the percentage of “pasture + agriculture” cover (A, C, F, H) or the total percentage of anthropic cover (i.e., including pasture, agriculture, and urban areas) (B, D, E, G, I, J, K, L).</figcaption></figure>

However, to predict the percentage of anthropic cover, it is possible to estimate the degree of anthropic disturbance from pollen assemblages when considering the frequencies of multiple pollen types. Hence, we listed and summarized the “anthropic indicators” and the “natural vegetation indicators” (Tab. 1).

The total anthropic indicators tended to be higher at sites with higher percentages of anthropic cover and the natural vegetation tended to be higher at sites with lower percentages of anthropic cover (Fig. 5).

Our definition of an “alteration score” is based on the following calculation: “sum (Anthropic indicators)” - “sum (Natural vegetation)”. The “alteration score” was higher at sites with higher percentages of anthropic cover (Fig. 5) and can thus be used to predict the degree of anthropic disturbance around the sites.

The correlation between the alteration score and the percentage of anthropic cover around the sites was significant (Fig. 5) and was highest for the area with a radius of 300 m. (%Anthropic cover includes “pasture”,

<figure><img src="image_9.png" /></figure>
Estimated degradation of the Caatinga based
on modern pollen rain deposited in reservoirs

<figure><img src="image_10.png" /><figcaption>**Figure 5.** Synthetic diagram showing the results obtained from Mapbiomas (Fig. 2) and from the pollen analyses (Fig. 3) with anthropic indicator taxa (orange) and natural vegetation indicator taxa (green). Sites are organized according to the degree of degradation as defined by the alteration scores. Numbers refer to Table 1.</figcaption></figure>
“agriculture”, “pasture-agriculture”, “urban areas”, “other urban areas” and “rocky outcrops” from MapBiomas):

## Reconstructing the percentage of anthropic cover using the pollen data - regression model

The alteration score was used to build a linear regression model and to reconstruct the percentage of anthropic cover in the area with a radius of 300 m from the point where the pollen was sampled. To calculate the model, the dataset was randomly split into training and test datasets with 80% and 20% of samples, respectively. Linear regression coefficients were calculated using the training dataset. The remaining samples (20%) were used to test the model and to calculate prediction errors. Accuracy was measured by calculating mean absolute errors (MAE) (Fig. 6).

The regression model equation obtained was:

$$ \text{AnthropicCov} = 3.17 \times \text{AlterationScore} + 60.04 \quad (1) $$

The model was applied to the complete dataset in order to compare the predicted and real percentages of anthropic land cover.

For 27 samples, the real percentage of anthropic cover fell within the margin of error of the predicted value (Tab. 3). For 10 samples, the real percentage of anthropic cover was less than 10% higher or lower than the predicted value (including the error margin), and for the other 10 samples, the real percentage of anthropic cover was more than 10% higher or lower than the predicted value (including the error margin) (Fig. 7). For at least some of the 10 samples whose percentage of anthropic cover was over- or underestimated by more than 10% (10% and 31%), the difference between real and predicted values might be related to the presence of large water bodies within the area with a radius of 300 m, as water surfaces were not considered in the land cover.

## Application to fossil pollen data

Application of the regression model to the ARA core (Ledru et al. 2020) indicated that in this area, the percentage of anthropic cover ranged from 62.6% to 0% (Fig. 8). Higher percentages of anthropic cover were found in the bottom part of the core that corresponded to the period when the dam was commissioned. A gradual decrease in anthropic cover was observed in the years following the building of the dam demonstrating recovery of the vegetation. Another increase in anthropic activities was clearly visible with the launching of the Araras Norte irrigation project in 1985. In the upper part of the core, natural vegetation predominated, evidence for recovery of the landscape. However, our results also revealed the limits of our surface sample dataset as

<figure><img src="image_11.png" /><figcaption>**Figure 6.** Regression curves between alteration scores and percentages of anthropic cover in areas with a radius of 300 m, 500 m and 1,000 m.</figcaption></figure>

today, the vegetation today is quite different from the original cover but is nevertheless characterized as “natural” by the alteration score. Our attempt to apply the score to the most recent decades also highlighted the need for additional surface samples from the native Caatinga to improve the calibration. However, we are confident that, once completed, our approach and the use of the alteration score will make it possible to characterize the state of the landscape in semi-arid regions.
Estimated degradation of the Caatinga based on modern pollen rain deposited in reservoirs

<table><caption>**Table 3.** Results of the regression model performed on our sample dataset.</caption><thead><tr><th colspan="4">Linear regression</th></tr><tr><th>Coefficients</th><th>Std. Error</th><th>Pr(&gt; |t|)</th><th>MAE</th></tr></thead><tbody><tr><td>Intercept</td><td>60.04</td><td>3.01</td><td>2E-16</td></tr><tr><td>Regression coef.</td><td>3.17</td><td>0.39</td><td>1.85E-09</td></tr></tbody></table>

<figure><img src="image_12.png" /><figcaption>**Figure 7.** A) Calculated values of the percentage of anthropic cover (orange) and real values of anthropic cover (red). B) Difference between real and predicted (mean) values of anthropic cover. The grey area indicates the model error margin (12 %) and red arrows indicate samples for which the difference between real and predicted anthropic cover was more than 10 % higher than the margin of error.</figcaption></figure>

<figure><img src="image_13.png" /><figcaption>**Figure 8.** Graphical representation of the alteration score calculated for the sediment core collected in the reservoir of Araras (Varjota, Ceará) (see Ledru *et al.* 2020 for more information on the age model and pollen analyses).</figcaption></figure>
Discussion

The Caatinga is one of the most degraded Brazilian
biomes due on the one hand, to colonial agriculture
organized around export-oriented production and on the
other hand, the pressure on natural resources due to the
recurrent threat of starvation in a highly populated region
with more than 25 million people representing ~15% of
Brazil's population at the beginning of the 21st century
(CGEE 2016). Our results show that even in areas where
the vegetation cover appears to be relatively well preserved
and hardly affected by anthropic activities, the alteration
score remains relatively high (Tab. 1, Fig. 5). Consequently,
we conclude that the BSA region is completely impacted
by anthropogenic activities although at different levels
depending on historical uses (Monteiro & Kurtz 2020). This
is not surprising as the poor rural population had to count
on natural resources for their survival, the most common
activities being wood harvesting, hunting, slash and burn
agriculture, and deforestation to create pasture for cattle
and goats. Few phytosociological studies measured the
herbaceous cover (Moro et al. 2016). However, when they
did, they all showed a high percentage of herbaceous species,
60% of the total species richness of the Caatinga, in line
with our results, in which non-arboreal pollen accounted for
more than 50% of all pollen. Floristic composition in areas
that were exploited for long periods of time (agriculture/
planted pasture) and left to recuperate for a relatively
short period of time (20 years) were strongly dominated
by Mimosa tenuiflora (Pereira et al. 2002). The domination
of M. tenuiflora in the pollen assemblages of all our samples
thus tends to prove that all our study area was already
cultivated and/or used for cattle. Indeed, whenever a piece of
land was not cultivated, it was used for permanent grazing,
thereby preventing natural regeneration and increasing
desertification (CGEE 2016). Among the herbaceous
species, the families with highest number of species were
Euphorbiaceae, Fabaceae, Asteraceae, and Convolvulaceae,
while 47% of the families were only represented by a single
species (Costa et al. 2007). This is in line with our results,
if we consider that Euphorbiaceae is entomophilous and is
a poor pollen producer (Barth 1989). In addition, among
the 90 pollen taxa identified in our modern samples, the
majority were represented by zero or only by single pollen
grains only found once in the dataset. Other studies indicate
that the tree species Fabaceae, Euphorbiaceae, and Cactaceae
are among the most species rich families in the Caatinga
(Moro et al. 2014). The pollen rain showed that Fabaceae is
the dominant family while Euphorbiaceae and Cactaceae,
two poor pollen producers, were scarce in our dataset.
When we included herbaceous species, other families
became floristically important, for instance, Asteraceae
and Convolvulaceae (Araújo et al. 2005). However, these

authors reported that ratios between herbaceous and tree
species and the dominant species can vary considerably
from one site to another (Moro et al. 2015; Monteiro &
Kurtz 2020). Additionally, even areas that are considered
as well-conserved show signs of past use (Guedes et al.
2012). We thus conclude that the historical background,
climate, orography, and soils should be taken into account
when defining a modern pollen rain sampling strategy in
the BSA. Nevertheless, the surface samples we collected in
the reservoirs are generally representative of the vegetation
cover and of the general degradation of the landscape. In
future studies, improvements such as increasing the number
of samples in protected areas, diversifying the data set by
sampling in all the different types of ecosystems found in
the Caatinga, including the historical background in the
metadata as well the age of the reservoirs and the successive
agricultural activities, adding a botanical survey in each
pollen sampling area. To summarize, our results show that
our study areas can be considered as degraded and that the
initial categories "preserved" or "intermediary" were not
confirmed in the pollen assemblages, in line with results of
botanical surveys and with the historical background of the
Caatinga biozone. The on-going process of desertification
is climate-independent and was initiated many decades or
even centuries ago when the cultivated crops were replaced
by grazing, thereby threatening the regional biodiversity.

Acknowledgements

This research was funded by a “sciences without boarder”
(Portuguese acronym CSF) project “Evolution of biodiversity
loss of areas in desertification processes” (CNPq
400890-2014-3), INCT IN-TREE: Interdisciplinary and
transdisciplinary studies in ecology and evolution (CNPq
CAPES FAPESBA) and covered a visiting foreign professor
fellowship for MPL at the Federal University of Ceara (UFC)
in the framework of the CSF project (2014–2019). This
research is also part of the JEAI SANA (Le Semi-aride du
Nordeste brésilien à l’Anthropocène) (IRD FUNCAP) and
the Labex-CEBA.

Authors contribution

Marie-Pierre Ledru: Supervision, Visualization,
Investigation, Conceptualization, Writing - Original Draft.

Raquel Franco Cassino: Formal analysis, Visualization,
Investigation, Writing - Review & Editing.

Vaneicia Gomes: Formal analysis, Writing - Review &
Editing.

Julia Sfair: Formal analysis, Writing - Review & Editing.

Francisca Soares Araújo: Supervision, Resources, Writing -
Review & Editing, Project administration, Funding
acquisition.

<figure><img src="image_14.png" /></figure>
Estimated degradation of the Caatinga based
on modern pollen rain deposited in reservoirs

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<figure><img src="image_15.png" /></figure>