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Plant Biosystems - An International Journal Dealing with all Aspects of Biology: Official Journal of the Click for updates Societa Botanica Italiana Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tplb20 Are an indicator of open habitats and pastoralism in current and past vegetation studies? A. Florenzanoa, M. Marignanib, L. Rosatic, S. Fascettic & A. M. Mercuria a Laboratorio di Palinologia e Paleobotanica, Dipartimento di Scienze della Vita, Università di Modena e Reggio Emilia, Modena, Italy b Dipartimento di Scienze della Vita e dell'Ambiente, Macrosezione Botanica ed Orto Botanico, Università di Cagliari, Cagliari, Italy c Scuola di Scienze Agrarie, Forestali, Alimentari ed Ambientali, Università degli Studi della Basilicata, Potenza, Italy Accepted author version posted online: 15 Dec 2014.Published online: 22 Jan 2015.

To cite this article: A. Florenzano, M. Marignani, L. Rosati, S. Fascetti & A. M. Mercuri (2015): Are Cichorieae an indicator of open habitats and pastoralism in current and past vegetation studies?, Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana, DOI: 10.1080/11263504.2014.998311 To link to this article: http://dx.doi.org/10.1080/11263504.2014.998311

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ORIGINAL ARTICLE

Are Cichorieae an indicator of open habitats and pastoralism in current and past vegetation studies?

A. FLORENZANO1, M. MARIGNANI2, L. ROSATI3, S. FASCETTI3, & A. M. MERCURI1

1Laboratorio di Palinologia e Paleobotanica, Dipartimento di Scienze della Vita, Universita` di Modena e Reggio Emilia, Modena, Italy; 2Dipartimento di Scienze della Vita e dell’Ambiente, Macrosezione Botanica ed Orto Botanico, Universita` di Cagliari, Cagliari, Italy and 3Scuola di Scienze Agrarie, Forestali, Alimentari ed Ambientali, Universita` degli Studi della Basilicata, Potenza, Italy

Abstract Cichorieae, one of the six tribes of the sub-family Cichorioideae (), produces a well-recognisable fenestrate pollen type. In the Mediterranean area, the significance of high percentages of Cichorieae pollen from archaeological layers is still questioned. We assessed the presence of Cichorieae as indicators of open habitats and pasturelands in current plant communities by comparing data on vegetation composition with pollen spectra from two Hellenistic sites of Basilicata (southern Italy): Difesa San Biagio in the low valley of the river Bradano and Torre di Satriano in the Lucanian Apennines. We also analysed the pollen morphology bringing to the discrimination of size classes within the fenestrate type of Cichorieae. Pollen spectra from the considered archaeological sites have low forest cover (7% on average); Asteraceae and Poaceae are prevalent; Cichorieae account to ca. 23%; coprophilous fungal spores are varied and present high concentrations. In surface soil samples collected near the sites, Cichorieae pollen is about 12%. In current vegetation types, an increasing abundance of Cichorieae was observed from salt marshes, forests and shrublands to open habitats and grasslands. This is coherent with the actual land cover around the study sites and the findings of the archaeological sample that point to an open landscape dominated by pastures and cultivated fields. Our integrated approach confirmed that today Cichorieae are common in secondary pastures and in some types of primary open habitats of southern Italy: hence, high percentages of this pollen can be considered a good indicator of these habitats even in past environment reconstructions.

Keywords: Archaeological sites, Basilicata, Mediterranean, palaeoecology, phytosociological database, pollen morphotypes

Introduction effect of browsing on soil and plant communities (e.g. Bakker 1998; Blasi et al. 2009; Farris et al. Land-use and climate shape vegetation and land- 2010b; Mercuri et al. 2010; Papanikolaou et al. Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 scape. This is especially evident in the Mediterranean basin where a cultural-ecological continuum con- 2011; Hillbrand et al. 2014). tributed to the development of agricultural practices In general, land exploitation by humans produced that has been visible during the last millennia changes that are visible in the archaeobotanical (Mercuri & Sadori 2014). records studied from Eastern (Bakker et al. 2012; Besides crop cultivation, forest management and Willcox et al. 2012), Southern (Giraudi et al. 2013; clearance, animal breeding and pastoralism are Zapata et al. 2013), Western (Zapata et al. 2004; among the primary agents that have transformed Ejarque et al. 2010) and Central Mediterranean area the Mediterranean landscape. (Branch et al. 2005; Sadori et al. 2011, 2014; Mercuri Past evidence of such ancient practices is found in 2014). In Italy, anthropogenic pollen indicator curves palaeoenvironmental records and in organic deposits increase during the last three millennia when the trend of archaeological sites (Lowe et al. 1994; Branch & of Cichorieae follows Poaceae, Plantago and Urtica in Marini 2013; di Lernia et al. 2013; Orengo et al. off-site (e.g. marine and terrestrial cores) and on-site 2014). Most of the traces left by animals are in the (archaeological layers) diagrams (Mercuri et al. 2012: chemical elements that enriched sediments or in the p. 358).

Correspondence: A. Florenzano, Laboratorio di Palinologia e Paleobotanica, Dipartimento di Scienze della Vita, Universita` di Modena e Reggio Emilia, Modena,Italy.Tel:þ39 059 2056016. Email: assunta.fl[email protected]

q 2015 Societa` Botanica Italiana 2 A. Florenzano et al.

Cichorieae, one of the six tribes of the sub-family despite the occurrence of more deep land-use Cichorioideae (Asteraceae family; ICN 2014), transformations during the last two centuries produces a well-recognisable pollen type that many (Morano 1994;Pontrandolfi1999). Moreover, palaeoecologists consider a good marker of the action Basilicata displays a high diversity of environmental of herbivores on vegetation (Hjelle 1999; Mazier features and vegetation types, belonging either to et al. 2006; Ejarque et al. 2011). Mediterranean and Temperate biochores (e.g. Blasi The significance of Cichorieae in pollen records 2010; Fascetti et al. 2013). from Mediterranean context is still controversial. The The aim of this paper is to assess if a higher high percentage of Cichorieae is usually explained by incidence of Cichorieae species is present in modern two different interpretations: (a) selective corrosion grassland communities linked to human activities (Bottema 1975; Dimbleby 1985; Di Rita & Magri and pastoralism, a hypothesis already proposed for 2009; Triantaphyllou et al. 2010) resulting in the the interpretation of pollen deposits from archae- overrepresentation of pollen with high-resistant exines ological sites but never tested on the current in little conservative, alkaline and poor sediments, like vegetation of southern Italy. This paper proposes some layers from archaeological sites; (b) presence of an integrated discussion of the main results obtained pastures (Behre 1986; Birks et al. 1989)inwhich from palaeoecological and ecological studies carried these pollen grains are abundant as they are linked to out in the region: the presence of Cichorieae in pollen herbivore action and animal browsing leading to a and vegetation spectra is evaluated according to the certain selection of . For example, Cichorieae joint interpretation of the past and current data. pollen is included in the Local Pastoral Pollen Indicators by Mazier et al. (2009), and in the Pollen Disturbance Index by Kouli (2014). Materials and methods In 2010, two papers have demonstrated that the Study area (Figure 1) over-representation of Cichorieae cannot rely only on selective corrosion and oxidation of exine: (i) Lebreton The Basilicata region lies in the southern Apennines et al. (2010) proved through an experimental approach between the Ofanto river in the north and the Mt. that oxidation does not generate any artificial over- Pollino (2248 m a.s.l.) massif to the south. Most of the representation of a specific pollen taxon, including territory is mountainous (47%) or hilly (45%). The Cichorieae; (ii) Mercuri et al. (2010)compared region also has a short coastline to the southwest on the Cichorieae and indeterminable pollen percentages, Tyrrhenian Sea and along the Ionian Sea to the and calculated that an increase of Cichorieae does not southeast. Mediterranean bioclimate prevails along the correspond to an increase of deteriorated pollen. They coast, whereas the inner areas display a temperate both concluded that the presence of Cichorieae can be climate with continental features. Rainfall has a typical reasonably interpreted as pasture indicators. Mediterranean regime, with rainy winters and dry In Mediterranean and arid zones, Cichorieae summers; minimum annual values are recorded in the pollen is found in records describing environments Metaponto coastal plain (532 mm) close to the Ionian that had developed under a long-time human Sea, and maximum ones (up to 2000 mm) are found in pressure. Besides the above mentioned anthropo- the mountain area along the Tyrrhenian Sea. Phytocli-

Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 genic pollen indicators, indicating the spreading of matic belts range from the Dry Thermo-Mediterra- human-influenced environments, their significance nean zone along the Ionian coast to the Hyperhumid as indicators of pasturelands is suggested by the co- Orotemperate zone (Blasi & Michetti 2007)ofthe presence of coprophilous fungal spores in the same highest mountains. Broadleaved deciduous forests are samples (Kouli et al. 2009; Mercuri et al. 2010). the dominant Potential Natural Vegetation (PNV, sensu This paper will appraise the value of Cichorieae Farris et al. 2010a), spanning from xeric white oak as indicator of open habitats and pasturelands by Mediterranean communities (e.g. Oleo-Quercetum assessing the incidence of Cichorieae in current plant virgilianae) to Temperate oceanic microthermal beech communities of southern Italy. This will contribute forests (Ranunculo brutii-Fagetum sylvaticae); however, to clarify the meaning of high rates of Cichorieae most of the territory is occupied by the Mesotemperate pollen found in Mediterranean records including Turkey oak vegetation series of Lathyro digitati- archaeological sites. In particular, the central Quercetum cerridis (see also Di Pietro et al. 2010). Mediterranean region of Basilicata was selected for The region has a heterogeneous land cover, with a comparison of past and current contexts due to two large forests and secondary pastures in the moun- main reasons: (i) the availability of recently published tains, and cereal crops and olive orchards in the hilly pollen analyses from several archaeological sites landscape. Due to mild temperature regime and (Mercuri et al. 2010; Florenzano & Mercuri 2012); water availability, the Ionian coastal plain is (ii) the features of this region that preserve a fairly characterised by intensive fruit farming and intact evidence of ancient agricultural practices horticulture. Cichorieae in current and past vegetation studies 3 Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 Figure 1. Location map of the Basilicata region, including the two archaeological sites (Torre di Satriano and Difesa San Biagio).

The archaeological context and studied sites both microscopic grazing indicators and archaeolo- gical evidence, environmental reconstructions stress The central and eastern parts of the Basilicata region the importance of pastoralism in the past economy of have well-preserved archaeological sites belonging to these lands (Florenzano & Mercuri 2012; Floren- different chronological phases (Bronze age, Helle- zano et al. 2013). Two archaeological sites, Torre di nistic, Roman and Medieval periods) that have been Satriano (TS) and Difesa San Biagio (DSB) covering investigated by archaeologists since the 1970s. Pollen from the 6th to the 1st century BC, were selected for and plant macroremains have been analysed with the this study (Figure 1; see Annex point 1). aim of obtaining reconstructions of plant landscape, land-use and diet (Costantini 1983; Novellis 2009; Florenzano 2013; Roubis et al. 2014). The palyno- Palynology (see Annex point 2) logical investigations have contributed to palaeoen- vironmental reconstructions providing details for the Pollen morphology of Cichorieae. Cichorieae have vegetation history recorded from terrestrial off-site fenestrate pollen (Faegri & Iversen 1989;syn. cores on a supraregional scale (e.g. Allen et al. 2002; echinolophate). This is the only case in the subfamily Joannin et al. 2012; Sadori et al. 2013). Based on of Cichorioideae, whose other pollen grains are 4 A. Florenzano et al.

indeed tricolporate and echinate like those occurring by a high presence of Cichorieae in Basilicata. in the Asteraceae family. This pollen type was We used the Basilicata vegetation database (Rosati described by Wodehouse (1935) as comprising an et al. 2012) that to date includes data from about 2000 outer exine that is raised in a pattern of echinate releve´s, carried out over a period of time ranging from ridges (lophae) surrounding depressions (lacunae). 1972 to 2013 (published and unpublished data). Within the tribe, Scorzonera humilis type has echinate We considered for inclusion in the database phytoso- pollen (lacunae absent; Blackmore 1984), and it is ciological releve´s or other vegetation plots containing not included in the fenestrate pollen type. Pollen records of vascular plants species composition and an morphology bringing to the discrimination of estimate of species cover. Vascular plants nomencla- morphotypes within the fenestrate type of Cichorieae ture follows Conti et al. (2005). A releve´s classifi- is described on the basis of lacunae (Chester & Raine cation was obtained by using a modified two-way 2001), dimensions (Florenzano et al. 2012) and on indicator species analysis (TWINSPAN) included in the combination of the two previous characters JUICE software (Tichy´ 2002; Rolecˇek et al. 2009). (Blackmore 1984; Wang et al. 2009). In the analyses, we set four pseudo species cut levels Cichorieae are easy to identify even when they are (0%, 5%, 25%, 50%) and total inertia as a measure of broken or fragmented. However, there is hardly any heterogeneity. For each vegetation type, we calculated species distinction within the subtribe. The longest the weighted cover percentage at family level (and to axis ranges between 15 and 62 mm (Blackmore 1984; subtribe only for Cichorieae). De Leonardis et al. 1992; Osman 2006; Wang et al. The analyses on current vegetation provided 2009, and reference collection), and some sub- information on floristic composition of the veg- categories were proposed as the whole grains may be etation types for the area spanning from Basilicata to easily measured during routine analyses (Florenzano Apulia, south Campania and north Calabria, in et al. 2012; Table I). A minimum of 50 well- southern Italy. preserved and whole pollen grains of Cichorieae were measured at 400 £ magnification in the samples from the two sites here studied. Results The incidence of Cichorieae in pollen spectra Current vegetation data and methods Cichorieae are significant in the past and current To achieve more detailed palaeoenvironmental pollen spectra analysed (Figures 2 and 3). reconstructions, we tried to link the Cichorieae pollen In archaeological sites, they represent one quarter morphology to current vegetation types characterised of the total pollen (22% in TS, and 26% in DSB, on

Table I. Pollen morphotypes of Cichorieae according to different authors; the emphasis is on the dimension.

Florenzano et al. (2012); Blackmore (1984); Osman (2006); Chester and Raine (2001) Wang et al. (2009) De Leonardis et al. (1992) Included taxa

Hieracium type – Hieracium type (,18 mm) Hieracium p.p., Tolpis p.p. Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 – – Small pollen grains (18–25 mm) Andryala, Hieracium p.p., Tolpis p.p.

Catananche, Chondrilla, p.p., Crepis, Geropogon, Hedypnois, Hyoseris, Hypochaeris, Lactuca, Lactuca type Lactuca sativa type Lapsana, Leontodon, Picris, Sonchus type Sonchus oleracerus type Prenanthes, Reichardia, Taraxacum type – Medium pollen grains (26–44 mm) Rhagadiolus, Scorzonera villosa Cichorium type Cichorium intybus type subsp. columnae (De Leonardis et al. 1992), Sonchus, Taraxacum, Urospermum; Cicerbita, Launaea and Willemetia (not in the flora of Basilicata)

Scolymus – Scorzonera laciniata type Scorzonera laciniata type Cichorium p.p., Scolymus, Scorzonera hispanica type Scorzonera humilis Scolymus type ($45 mm) Scorzonera (excl. S. humulis), Tragopogon Tragopogon pratensis type Tragopogon

– Arnoseris minima ––

Note: The last column reports the genera that are described in relevant literature. Cichorieae in current and past vegetation studies 5

average), and in the modern samples, they are half of ceae, which include plants common in saline this value (12%). According to the size groups (Table or alkaline soils of badlands, are high in DSB I), classes of 18–25 and 26–44 mm are prevalent. (11%) and absent in TS samples. The incidence of the smallest size, ,18 mm, is quite (4) Among the cultivated herb plants, cereals are variable but relatively low being around 10% in well attested at TS (6%), and only in traces surface soil and DSB samples. On the opposite side, at DSB. Fabaceae, which can include fodder the highest size, $45 mm, is similarly low and with plants, are common in both sites, but their comparable values (always ,2%) in past and surface amounts are more significant in TS (4%) soil samples. than in DSB spectra (0.5%). Pollen spectra from the archaeological sites (5) Coprophilous fungal spores are varied and have (Figure 4), which are relevant to understand the high concentrations; in particular, the ascos- main palaeoenvironmental context, are also marked pores of Sordaria type and Chaetomium are by other features besides Cichorieae: known to be quite common in the dung of herbivores (van Geel et al. 2003), and (1) Forest cover is very low (7% on average): Sporormiella type includes obligate coprophi- woody plants include low percentages of lous fungi (Ahmed & Cain 1972). Coupled with Pinus, broadleaved trees (deciduous Quercus, the archaeological and pollen data, they are Corylus;andCarpinus betulus found only at evidence of local grazing or breeding activities. TS), and elements of the Mediterranean maquis (e.g. Quercus ilex type, Olea and Cichorieae rate in current vegetation Phillyrea). There are some interesting traces of Castanea in TS, while Alnus is found in a By means of TWINSPAN clustering, we identified in few samples of DSB. the dataset 20 main vegetation types in the Basilicata (2) Wet environments are only reported in TS, dataset (Table II). These approximately correspond with the presence of Cyperaceae (5%) and to class/order level of the syntaxonomical classifi- other hygro-hydrophytes (Phragmites cf. aus- cation, ranging from subalpine shrublands to marine tralis, Typha/Sparganium, Sagittaria, Nym- dunes vegetation. phaea cf. alba). Data analyses show that the presence of (3) Asteraceae and Poaceae – indicating open Cichorieae is highly variable in current vegetation. habitats – are prevalent (35%), Plantago is a A very low presence (,1%) is found in forest and significant evidence of trampled areas, while shrubby vegetation (except for the MRS type the nithrophilous Urtica is rare. Chenopodia- discussed below); a low presence (,2%) is registered in wetlands, cliffs and dune vegetation. High values are observed in open, secondary or naturally disturbed habitats, such as the Mediterranean river bed communities. The highest values (,7–8%) are observed in pastures and open vegetation types. Apparently, an exception is represented by Medi- terranean riparian shrubland on braided river (Nerio- Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 Tamaricetea)–MRS,showingalmost5%of Cichorieae, but it can be explained by the fact that those communities naturally grow interspersed between nitrophilous vegetation of Mediterranean muddy river banks (NVM) and Mediterranean river bed vegetation on gravels (MRG). Cichorieae species reach .3% only in six vegetation types (MRG, TPS, NVM, MRS, BXV, MDG) in modern releve´s. The pollen of some of the species that characterise those six vegetation types are described in literature: this permits to infer more precisely some taxa within the Figure 2. Incidence of four size classes of Cichorieae in past and routine identification carried out during pollen current pollen rain from archaeological (TS, DSB) and surface soil (Modern) samples, respectively. The histograms show the analyses (Table III). percentage of each Cichorieae pollen class (broken and not measurable pollen grains are not reported) considering the Total Cichorieae Sum as 100%. The last group of histograms shows the Discussion mean value of Cichorieae in the same spectra calculated as percentage on the Pollen Sum (Tot on PS). Broken and not In past pollen spectra (Figure 4), Cichorieae are measurable pollen grains are only included in the Tot on PS. significant in open landscapes. The main features of 6 A. Florenzano et al.

Figure 3. Incidence of four size classes of Cichorieae (see the text) from 15 samples of the two archaeological sites (DSB, 11 samples; TS, 4 samples): the histograms show the mean percentage of each Cichorieae class calculated on the Total Pollen Sum.

the palaeoenvironmental reconstruction of the Cichorieae and pollen morphology of these plants is two archaeological sites that are reported in Annex proposed. (point 3) help to understand the archaeobotanical contexts. Data interpretation is based on the multi- Cichorieae in current and past vegetation disciplinary archaeological studies and pollen ana- lyses discussed by Florenzano and Mercuri (2013: The analysis of the vegetation database regarding TS) and Mercuri et al. (2010: DSB). the Basilicata territory shows a gradient of abun- Then the main features of vegetation types dance of Cichorieae from saltmarsh environments, to currently present in the region are reported. forests and shrubby and open grassland vegetation A comparison between the vegetation types with (Table II). Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015

Figure 4. Percentage pollen diagram from the archaeological sites Difesa San Biagio (DSB, 11 samples) and Torre di Satriano (TS, 4 samples). Selected pollen taxa were grouped into sums useful for land-use reconstructions: (a) Mixed oakwood (Carpinus betulus, Corylus, Ostrya/Carpinus orientalis, deciduous Quercus, Ulmus); (b) Mediterranean plants (Cistus, Helianthemum, Olea, Phillyrea, Pistacia, Quercus ilex type); (c) Hydro-hygrophytes (Alnus, Cyperaceae, Nymphaea, Phragmites, Ranunculus macrophyllus, Sagittaria, Typha); (d) Cultivated plants (Avena/Triticum group, Hordeum group, Castanea, Olea, Juglans); (e) Pastoral indicators (Asteraceae including Cichorieae, Plantago, Ranunculaceae, Urtica). NPP concentrations are calculated as NPP per gram (shown as npp/g/100). Cichorieae in current and past vegetation studies 7

Table II. Composition of the identified vegetation types, showing families that usually indicate open habitats.

Poaceae Other wild Vegetation types Cichorieae Asteraceae grasses Plantaginaceae Urticaceae Fabaceae Chenopodiaceae

MRG – Mediterranean river bed vegetation on 8.20 23.07 19.44 2.45 0 17.30 0 gravels (Cisto-Micromerietea /Helianthemetea) TPS – Temperate pastures and fallow fields on 7.12 8.08 33.69 6.27 0 15.60 0 siliceous substrata (Molinio-Arrhenateretea, Artemisietea) NVM – Nitrophilous vegetation of 6.08 6.99 14.25 2.25 0 19.40 0.40 Mediterranean muddy river banks (Bidentetea) MRS – Mediterranean riparian shrubland on 4.96 0.82 15.18 0.72 0.01 3.70 0 braided rivers (Nerio-Tamaricetea) BXV – Badlands xeric vegetation 4.93 3.68 38.35 2.56 0 20.70 15.00 (Lygeo-Stipetea) MDG – Mediterranean dry grasslands on 3.03 5.88 60.15 0.29 0 4.90 0 limestone (Helianthemetea, Lygeo-Stipetea, Scorzonero-Crysopogonetalia) WV – Wetland vegetation 1.88 1.51 18.10 0.01 0 17.40 0 (Phragmito-Magnocaricetea) TGL – Temperate dry grasslands on limestone 1.45 3.26 37.40 3.01 0 12.70 0 (Brometalia erecti) HSV – Hygrophilous shrub vegetation of 0.78 3.23 37.35 0 0 0 8.10 badlands gullies (Nerio-Tamaricetea) SQF – Submediterranean/temperate Quercus 0.68 1.49 9.65 0.11 0.001 7.20 0 cerris, Q. pubescens and Q. frainetto forests (Teucrio-Quercion cerridis) MCL – Mediterranean chasmophytic vegetation 0.42 1.84 6.82 0.17 6.62 0.37 0 on limestone cliffs (Asplenietea) TMF – Temperate mesophilous Quercus cerris or 0.29 1.32 3.70 0 0.0006 1.60 0 Fagus sylvatica forests (Fagetalia sylvaticae) MQF – Mediterranean Quercus virgiliana/ 0.10 0.32 5.66 0 0.10 0.10 0 Quercus trojana forests (Quercetalia pubescentis) MDV – Marine dune vegetation 0.08 14.94 37.48 0 0 7.00 0.17 (Ammophiletea) MMF – Mediterranean mixed forests 0.06 0.06 4.28 0.09 0.002 2.90 0 (Quercetalia ilicis; Quercetalia pubescentis) MCV – Mediterranean chamaephytic vegetation 0.05 0.12 9.20 0.01 0 24.00 0 (Cisto-Micromerietea) SS – Subalpine shrublands (Pino-Juniperetea) 0.02 0.75 5.61 0.007 0.44 2.00 0.02 MM – Mediterranean maquis 0.02 0.82 5.79 0.04 0 8.70 0.10 (Oleo-Ceratonion)

Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 TSF – Temperate swamp forests 0.01 0.16 2.23 0.00 1.72 0 0 (Alnetea glutinosae) SV – Saltmarsh vegetation (Salicornietalia, 0 2.56 29.78 0.07 0 5.30 24.20 Limonietalia, Juncetalia)

Notes: Vegetation types are sorted according to the percentage of Cichorieae. Data are expressed in percentage values.

The mean percentage of Cichorieae obtained Lygeo-Stipetea) are frequent on clayey slopes or from releve´s of current vegetation points to a badlands and garrigues on gravels and conglomerates preference of these plants for open environments (Cisto-Micromerietea) around DSB. and pastures, in agreement with the general Cichorieae does not seem to be linked only to the palaeoenvironmental interpretation. Cichorieae in human effects but they could be attributed also to modern pollen samples reflect some specific veg- natural distribution of specific plant communities. etation types (Figure 2): i.e. secondary mesophilous This is the case of the three primary vegetation types pastures (Molinio-Arrhenatheretea), dry grasslands linked to the Mediterranean river environment (Festuco-Brometea) and sub-nitrophilous vegetation MRG, NVM and MRS (Mediterranean riparian of fallow fields (Artemisietea vulgaris) are widespread shrubland on braided rivers-Nerio-Tamaricetea, Table around TS; Mediterranean therophytic or hemicrip- II) growing in environments frequently disturbed by tophytic grasslands (e.g. Helianthemetea guttati, the action of flooding water. 8 A. Florenzano et al.

Table III. Vegetation types (labels refer to Table II) showing the highest values of Cichorieae (.3%), and description of pollen morphology of Cichorieae species identified in each vegetation type according to different authors.

Phytosociological releve´s Pollen morphology

Vegetation Pollen type by size types Cichorieae species Cichorieae pollen (Florenzano et al. 2012)

Chondrilla juncea Chondrilla juncea range size: P ¼ 24 mm; E ¼ 32 mm (De Leonardis et al. 1992) Crepis range size: P ¼ 29–39 mm; E ¼ 32–48 mm(Blackmore1984). Crepis neglecta Cichorium intybus group (within Cichorium intybus type) – MRG Echinolophatae, and narrow equatorial ridges with one row of echinae acicular, much shorter than the height of the ridges. Polar areawith 4– Medium pollen grains 15 isolated central echinae. Total of 15 lacunae (Blackmore 1984) (26–44 mm) Hypochaeris range size: P ¼ 27–36 mm; E ¼ 32–44 mm Hypochaeris achyrophorus (Blackmore 1984). Taraxacum officinalis group (within Cichorium Hypochaeris radicata intybus type) – Echinolophatae, and narrow equatorial ridges with one row of echinae acicular, much shorter than the height of the ridges. Polar area with 4–15 isolated central echinae. Total of 15 lacunae (Blackmore 1984)

Hypochaeris achyrophorus See above Hypochaeris achyrophorus Medium pollen grains (26–44 mm) Cichorium intybus range size: P ¼ 41–48 mm; E ¼ 51–59 mm (Blackmore 1984). Cichorium type – Ectoapertures colpi, each TPS divided into 1 poral and 2 abporal lacunae by 2 pairs of short Scolymus type Cichorium intybus continuous ridges. Paraporal lacunae 2 per mesocolpium. Polar ($45 mm) lacunae absent, polar area ranging from a triradiate ridge to an extensive echinate region; total of 15 lacunae. Polar area small with 1 ^ (2) ^ 4 isolated central echinae (Chester & Raine 2001).

Chondrilla juncea See above Chondrilla juncea Picris echioides range size: P ¼ 31–44 mm; E ¼ 37–45 mm Helminthotheca echioides (syn. (Blackmore 1984). Taraxacum officinalis group (within Cichorium Picris echioides, Crepis echioides) intybus type) – Echinolophatae, and narrow equatorial ridges with one row of echinae acicular, much shorter than the height of the ridges. Polar area with 4–15 isolated central echinae. Total of 15 lacunae (Blackmore 1984) Picris hieracioides Picris hieracioides range size: P ¼ 31–36 mm; E ¼ 36–41 mm Medium pollen grains NVM (Blackmore 1984). See above Helminthotheca echioides (26–44 mm) Sonchus range size: P ¼ 28 mm; E ¼ 43 mm (Blackmore 1984). Sonchus type – Ectoapertures colpi, often divided into lacunae by Sonchus asper constrictions or short paired ridges. Colpi each divided into 2 abporal lacunae by a central constriction or pair of ridges. Two equatorial lacunae per mesocolpium. Poral lacunae 3 in each of the Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 extensive polar areas; total of 21 lacunae (Chester & Raine 2001) Cichorium intybus See above Cichorium intybus Scolymus type ($45 mm)

Lactuca serriola range size: P ¼ 30–37 mm; E ¼ 35–43 mm (Blackmore 1984). Lactuca type – Ectoapertures colpi, each divided into 1 poral and 2 abporal lacunae by 2 pairs of short continuous Medium pollen grains ridges. Paraporal lacunae 2 per mesocolpium. Polar lacunae absent, (26–44 mm) MRS Lactuca serriola polar area ranging from a triradiate ridge to an extensive echinate region; total of 15 lacunae. Polar area not more than a triradiate ridge usually with one row of echinae, abporal and paraporal lacunae extending almost to the poles (Chester & Raine 2001)

Geropogon glaber range size: P ¼ 35–40 mm; E ¼ 40–46 mm (De Geropogon glaber Leonardis et al. 1992). Geropogon type – Ectoapertures colpi each divided into 1 poral and 2 abporal lacunae by 2 pairs of short continuous ridges. Total of 15 lacunae (Osman 2006) Medium pollen grains Sonchus oleraceus range size: P ¼ 32–41 mm; E ¼ 32–46 mm (26–44 mm) BXV Sonchus oleraceus (Blackmore 1984). See above Sonchus asper Cichorieae in current and past vegetation studies 9

Scorzonera range size: P ¼ 42–58 mm; E ¼ 44–62 mm(Blackmore Scorzonera hispanica glastifolia 1984). Scorzonera hispanica type – Ectoapertures colpi, each divided into 2 abporal lacunae by a central constriction or pair of ridges. Two equatorial lacunae per mesocolpium. Abporal lacunae extending to an expanded spiny polar area, or to a polar lacuna. Polar lacunae 1 per pole, large, hexagonal; total of 20 lacunae (Chester & Raine 2001) Scolymus type ¼ m ¼ m Scorzonera laciniata range size: P 42–58 m; E 44–62 m ($45 mm) (Blackmore 1984). Scorzonera laciniata type – Ectoapertures colpi, Scorzonera laciniata often divided into lacunae by constrictions or short paired ridges. Colpi each divided into 2 abporal lacunae by a central constriction or pair of ridges. Two equatorial lacunae per mesocolpium. Abporal lacunae extending to an expanded spiny polar area. Polar lacunae absent, extensive echinate polar areas present; total of 18 lacunae (Chester & Raine 2001)

Crepis corymbosa See above Crepis neglecta Medium pollen grains (26–44 mm) MDG Hypochaeris achyrophorus See above Hypochaeris achyrophorus Scorzonera villosa columnae Scorzonera villosa columnae range size: P ¼ 35–42 mm; E ¼ 39– Medium pollen grains 44 mm (De Leonardis et al. 1992). See above Scorzonera laciniata (26–44 mm)

Notes: Pollen size range is determined by measuring the maximum diameter length. P, polar view; E, equatorial view.

At DSB, the archaeopalynological dataset meadows (Ejarque et al. 2011). In phytosociology, it describesanopenlandscapealongwithhigh is considered a species of pioneer ruderal and Cichorieae (up to 40%); this feature suggests a long- nitrophilous sunny communities, growing on rich term local grazing or breeding activities. In fact, this soils in Mediterranean and temperate territories (i.e. may have produced the clearing of vegetation and Artemisietea vulgaris; Rivas-Martı´nez et al. 2002). increased the effects of the fluvial dynamics on the Within the vegetation types listed in Table III, the riparian vegetation. Also the good values of Cheno- coenoses characterised by Cichorium intybus, cer- podiaceae (11%), a clear indicator of badland tainly related to grazing/breeding activities are TPS vegetation due to the common presence of Atriplex, (Molinio-Arrhenateretea, Artemisietea)andNVM Beta, Camphorosma, Suaeda, point out that sheep and (Bidentetea). cattle grazing was probably facilitated by the natural presence of open habitats in the site. Conclusions Cichorieae are common in open habitats, mainly Pollen morphology and vegetation types pasturelands, in current plant communities. There- Although the pollen identification is difficult, it may fore, in pollen-based environmental reconstructions,

Downloaded by [Univ Studi Basilicata] at 02:06 27 January 2015 be useful to link the Cichorieae pollen morphology to Cichorieae (the tribe of Cichorioideae used as specific vegetation types (Table III). The identifi- synonym of fenestrate pollen) can be considered cation never reaches the plant species level except for one of the most important pastureland or grazing Cichorium intybus (in TPS, NVM, see below). indicators. Their pollen grains are often abundant in The smallest Hieracium pollen type may refer to the spectra from central and southern , their species H. pilosella and H. pseudopilosella,thatare over-representation being properly linked to grazing frequent in the TGL – Temperate dry grasslands on activity rather than selective deterioration. limestone (Brometalia erecti), and in the SS – subalpine The discrimination of size classes within the vegetation (Pino-Juniperetea/Seslerion apenninae). Cichorieae pollen may help to refine palaeoenviron- The most represented Cichorieae pollen types mental reconstructions. In particular, in the present belong to Taraxacum officinalis group and C. intybus study we assess that the large size pollen ($45 mm) group within the C. intybus type (Blackmore 1984). belongs to Cichorium intybus, which is mainly Though having a very similar exine sculpture, associated with fallow fields and sub-nitrophilous C. intybus is easily noticeable thanks to its large size pastures and was rare in the studied sites. The ($45 mm–Scolymus type; Florenzano et al. 2012). medium size class includes most of the species in Cichorium intybus is considered a characteristic modern vegetation types; they actually prevail in past species of synanthropic vegetation, being mainly and current pollen spectra but result a low- associated to fallow fields (included in TPS, Table informative general group. Nevertheless, the Cichor- II), sub-nitrophilous pastures (NVM) and to grazed ieae medium-size pollen is a good indicator of 10 A. Florenzano et al.

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