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Coastal vertical cliffs of the National Park of Cilento: reservoirs of endemic

ARONNE G.1*, DE MICCO V.1, SANTANGELO A.2,SANTANGELO N.3, SANTO A.4, BUONANNO M. 5

1Department of Agricultural and Food Sciences, University of Naples Federico II (UNINA), 2Department of Biology, UNINA; 3Department of Earth Sciences, Environment and Resources, UNINA; 4Department of Civil, Architectural and Environmental Engineering, UNINA; 5 Institute for Agricultural and Forest Systems in the Mediterranean, CNR-ISAFoM, *Via Università, 100, Portici, (NA), ITALY [email protected]

Abstract: - Vertical cliffs are widely distributed, are characterized by peculiar environmental characteristics and are generally acclaimed for their flora rich of phylogenetic relicts and rare species. Many vertical cliffs occur along the coastline of the National Park of Cilento (Tyrrhenian coast of Southern Italy), an area highly affected by human impact. Using a multidisciplinary approach we aimed at verifying in this habitat the occurrence of a set of features repeatedly reported for vertical cliffs worldwide. The overall analysis of information available in literature and newly elaborated data showed that geological, geomorphological and structural features of coastal cliffs of Cilento and the number of endemic species, much higher than in the surrounding areas, are consistent with those reported for cliffs in other geographical region. Biological and ecological traits of one of the relict species, Primula palinuri Petagna, highlighted that not all the sites on the cliffs are equally suitable for life requirements of the species; moreover, long term survival of the populations is at risk due to large failure in generation turnover. Coastal cliffs of Cilento act as refuge of endemic species and deserve special concern for biodiversity conservation.

Key-Words: - Vertical cliffs, cliff ecology, endemic species, Mediterranean species, Primula palinuri

1 Introduction access may be considered the main constrain limiting the amount of studies and experiments on High and very steep rock walls are generally the biotic communities of cliffs. However, referred to as cliffs or vertical cliffs [1]. Their inaccessibility is reported to have protected these extension is difficult to quantify on geographical ecosystems in historical time from direct and maps because vertical surfaces are not easily indirect impact of anthropogenic disturbances [2]. detected by aerial photographs. Nevertheless Cliffs are asserted ecological islands because vertical cliffs occur at any latitude and elevation in peculiar abiotic characteristics of their environments all geographical regions of the world. Their very are also associated to a distinct fauna and flora [1]. steep orientation affects physical factors such as growing among rocks are generally defined incident radiation, wind speed and direction, amount as saxicolous; however, those living in narrow holes of direct precipitation, soil availability and water and crevices, often associated with vertical angle of retention. Specific studies on factors such as local slope, are classified as chasmophytes. Among many temperature, light, air humidity and underground others factors, chasmophytes have to deal with low water availability, showed that these environments availability of nutrients, very limited space for root are distinct from the nearby habitats and that development and scarce shelter sites for biotic seasonal fluctuations of the physical components are recruitment. often significantly more moderate on cliffs than on Despite environmental conditions are considered the surrounding level grounds [1]. harsh for growth, an amazing phenomenon has Despite they are commonly diffuse and provide a been reported for worldwide geographical regions: distinctive habitat, cliffs have been rarely the number of phylogenetic relicts and of rare investigated from an ecological view point. Difficult species is much higher on cliffs than in the flora of

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the surrounding areas [2-9]. Geographical isolation of them verified the presence of specific features and selective pressure towards evolution of adaptive generally reported as life strategies for cliff species features to peculiar conditions of cliffs, are basic or emphasized the co-occurrence of other relict considerations to explain the occurrence of species. numerous endemic species on cliffs all over the The aim of this work is to verify if the main world [1]. Richness of rare relict species is reported features reported for vertical cliff worldwide, occur also for cliffs of Eastern Mediterranean region [2]. also for the habitat of P. palinuri. Specific aims are: The phenomenon is considered as a result of both a) to analyse geological, geomorphological and environmental and anthropogenic pressure: in structural features of coastal cliffs of Cilento addition to past climatic changes, new supporting P. palinuri; b) to verify the richness in anthropogenic factors, such as fire, grazing and relict and endemic plant species of the same habitat; changes in land use, affected much less cliffs than c) to check if the life strategy of P. palinuri is in their surrounding areas because of their conformity with that of other cliff-hangers. To inaccessibility. Among them, grazing, especially by achieve these goals, we used a multidisciplinary goats, might have determined a selective pressure approach based on critical analyses of the available exterminating the plants living at the accessible sites literature as well as on new data elaboration. Results [2]. are discussed altogether in terms of biodiversity We have focused our scientific interest on cliffs conservation. of the Cilento’s shoreline that extends about 100 km along the Tyrrhenian coast of Southern Italy. Most of this area is within the boundaries of the National 2 Geomorphological, geological and Park of Cilento (Parco Nazionale del Cilento, Vallo structural features of coastal cliffs di Diano e Alburni, PNCVDA). This area attracts a large number of tourists, especially during the The distribution of P. palinuri along the Cilento summer, and is then affected by high anthropogenic coastline shows a lithological control (Fig. 1). impact. Despite the presence of coastal cliffs carved on Our work has been based on three main sandstone (Punta Licosa), on marl (Punta del considerations. Firstly, along the Cilento coast, Telegrafo) and carbonate rocks, all but one vertical cliffs and coastal slopes are frequent. populations live on carbonate cliffs. The only Studies on geomorphological, geological and exception is the population at Palinuro (Ficocelle) structural features of the Cilento coastline are that lives on pleistocenic cemented marine sands reported in the international literature [10, 11]. They and represents also the northern limit of the focus on geological phenomena, such as rock-falls geographical range of the species. and slides, and are aimed at identifying causes of All the others, as in the case of Capo Palinuro these phenomena and foreseen their future and Marina di Camerota coast, are on cliffs made up evolution. In these studies, environmental by Jurassic limestones and dolostones. As regards parameters are taken into account but there is no the origin and age of the carbonate cliffs, several mention of biological components. Secondly, studies have been carried out in the area establishing although several works have been done on the that the present day planar outline of the coast was Cilento’s flora [12-19], floristic knowledge of this defined at the end of middle Pleistocene (around area is still inadequate due to its naturalistic 300ka) [28-30]. In this period, the last climate complexity [20]. Regarding coastal cliffs, an oscillations determined both transgressions and exhaustive flora is not available. Thirdly, north- regressions of the coast line. During the facing vertical cliffs of a narrow area of the Cilento transgressions (OIS7 and 5), beach sediments, coast are habitat for Primula palinuri Petagna, a abrasion platforms and notches formed along the sea paleoendemic species, symbol of the National Park cliffs at elevations higher than the present sea level. of Cilento. It survives in a few sites between Capo During regressive phases (OIS6 and 3), the coastline Palinuro (Campania) and Capo Scalea (Calabria), a migrated seaward, and wider and thicker debris coastal area of about 80 km. P. palinuri is reported cones and talus formed at the base of the cliffs, as endangered in the Red List of threatened species testifying for cold and dry climate conditions which compiled by the International Union for the enhanced physical degradation of the cliffs. Conservation of Nature (IUCN) [21]. Several studies on the biology and ecology of this species are reported in literature [22-27]. Nevertheless, none

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Fig. 1. Geological sketch of Cilento coastline. Sandstones (orange), clayey-calcareous flysch (dark yellow), sandy beaches (yellow), carbonate rocks (violet). Geographical distribution of P. palinuri in the PNCVDA (red). (modified from Budetta et al. 2008)

Two configurations of rocky coasts occur in the natural arch [31], deep bays and allows the geographical range of P. palinuri: a) vertical cliffs continuous evolution of the coastline. with sloping shore platforms (e.g. the Molpa promontory and the Palinuro coastal arch) (Fig. 2 a); b) plunging cliffs with no shore platform (e.g. Capo 3 Habitat of P. palinuri Palinuro cliffs) (Fig. 2 b). Presently the cliffs show different stability Coastal vertical cliffs of Cilento include two types conditions depending mainly on the rock jointing of natural habitats of community interest (Annex I degree because the rock masses may show two of the 92/43/CEE directive). The first refers to predominant features. In one case (Capo Palinuro vegetated sea cliffs of the Mediterranean coasts with cliffs and some segments of Marina di Camerota endemic Limonium spp. (code 1240). The second coast), limestones and dolostones are massive, refers to calcareous rocky slopes with chasmophytic without discontinuities and low jointed, so the cliffs vegetation (code 8210). More specifically, coastal are very high (more than 100 m) and show good vertical cliffs of Cilento are characterised by stability conditions. In other cases, the carbonate associations of the classes Crithmo-Limonietea, succession may show a very thin stratification and which include halophytic communities, and an high degree of jointing with the presence of Asplenietea trichomanis, referred to the northern significant damage zones (it is case of the Molpa exposed rocks [32, 33]. promontory, of the Palinuro coastal arch and of Vertical cliffs play an important role in the some little bays of Marina di Camerota coast). The distribution of P. palinuri and represent the rock masses exposed along these cliffs show very preferential habitat of the species. Plants of P. bad mechanical properties and their stability palinuri live in soil pockets inside rock crevices and conditions are worse than in the previous case. In on ledges of north-facing vertical cliffs (Fig. 3). In a coincidence of these “low resistance” points, the sea few cases, groups of individuals were found in erosion was greater: it caused the formation of the rocky grassland with medium inclination slope

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either on the cliff plateau, close to the edge, or on differences have been measured in the amount of the cliff talus. Their altitudinal range is between 0 solar radiation: in 2012 the daily mean and 570 m above m.s.l., with a distance from the photosynthetically active radiation (PAR) ranged coast up to 1.8 km. from 124 to 627 µmol photons m-2s-1 on the north- facing vertical cliffs at the Marinella site, and from 887 to 2337 at the horizontal areas. These data confirmed that the quantity of solar radiation reaching north-facing vertical cliffs was significantly lower and with a range much narrower than that measured on the horizontal surrounding areas. The occurrence of environmental conditions more stable on the cliffs than on surrounding areas was confirmed also by other parameters such as temperature. At Marinella site, in 2012, during the vegetative period of P. palinuri, the mean range of daily temperatures resulted 5°C on north-facing vertical cliffs and 10°C at the horizontal surrounding area.

4 Relict and endemic plant species of the coastal cliffs

A flora of the costal cliffs of Cilento has not been published. We have extracted floristic data regarding vertical carbonate cliffs (70°-90°) in the geographical range of P. palinuri within PNCVDA boundaries from the database improved during a research project promoted by PNCVDA [34, 35]. Almost 100 entities (species and subspecies) are present in such habitats; 7 species are endemics and 5 sub-endemics (Table 1). Among them, Eokochia Fig. 2. Cliffs with sloping shore platform of the Palinuro saxicola, P. palinuri and Dianthus rupicola subsp. coastal arch (a) and plunging cliffs with no shore rupicola, deserve special concern and are included platform of Capo Palinuro (b). in Annex II and IV of 92/43/CEE directive, the first one as a priority species. At accessible sites plants are affected by grazing, Prevalent life forms are represented by perennial invasive species, fire, beach tourism, and specimen species, herbaceous (hemicryptophyte), and small collection. As a consequence, human threats might shrubs (chamaephyte). Eight out of the twelve have confined the presently survived plants on the endemic or subendemic species are rocky-slopes inaccessible free-face of the cliffs. This habitat is exclusive. Therefore, vertical cliffs of the Cilento characterized by rather stable environmental coasts play an important role as reservoirs of these conditions and is affected only by sudden landslides rare species. that are catastrophic for population survival. In Although our floristic data do not represent an 2007, a rock fall affected the Palinuro costal arch exhaustive flora of the vertical cliffs of Cilento, they destroying an extended area of the local population clearly indicate that a high percentage (7%) of of P. palinuri. endemic species live in this habitat. This Microclimatic data recorded close to the plants phenomenon is more evident considering that in the of P. palinuri are in agreement with the general whole Campania region the endemic units (species trend reported for other cliffs [1]. More specifically, and subspecies) are 5.4% of the total number of our data indicate that microclimatic conditions of vascular plants [36]. This percentage is even lower the north-facing vertical cliffs are significantly on the beaches interspersed with the vertical cliffs different from surrounding level-ground areas. Main along the coastline of Cilento, where only 3

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endemic species (1%) on a total of 346 taxa are In the alpine environment, Pleistocene climatic reported [15]. cycles gave rise to a large number of new species Richness of endemic species on cliffs of Cilento through polyploidy speciation [37, 39]. During cold is then in agreement with that reported for cliffs periods, some species colonised the Italian worldwide [1] and constitutes the basis for detailed peninsula. At present, only 3 species are present local prescriptions aimed to the biodiversity along the Apennines [40]: P. auricula L. s.l., a conservation of these sites. south-european orophyte, and two narrow endemics, P. apennina Widmer (northern Appennine) and P. palinuri (the most southern). Genetic and molecular analyses allow to include P. palinuri into the western clade of sect. Auricola (sub-sect. Euauricula Pax; [38]) and to define P. palinuri most closely related to P. allionii Lois. and P. marginata Curtis (endemic to western Alps) as well as to P. latifolia Lapeyr., but independent from the other two Apennine species P. auricula L. s.l. e P. apennina Widmer [40]. This allows hypothesising the origin of P. palinuri from an ancestor which moved from the western Alps by long-distance dispersal or, more likely, colonising the Apennines during a cold period of the Pleistocene (50,000 ± 40,000 yrs b.p.) [40]. Subsequently, it went extinct during a warming period, except along the coast of Cilento. However, in P. palinuri, the presence of several morphological characters shared with the eastern sub-sect. Cyanopsis may also support the hypotesis of an earlier ancestor [38, 41], already proposed by Chiarugi [42, 43]. Nowadays, P. palinuri is the only species growing on maritime cliffs, the only species from a Mediterranean habitat, and the most southerly in the whole section [39, 40]. Fig. 3. Schematic representation of a vertical cliff with plants of P. palinuri (modified from Pizzolongo 1964).

6 Life strategy of P. palinuri 5 Phylogenetic information on P. Plants on vertical cliffs are widely spaced but the palinuri areas between them are mostly unsuitable for the establishment of new individuals. Due to Present coastal cliffs of Cilento can be considered as geographical isolation, effective gene flow between a refuge habitat for remnant populations of P. populations is considerably limited and it is very palinuri and it is difficult to guess if, in the old past, difficult for a species to extend its range. plants of this species were distributed in other A series of biological/ecological features are habitats. However, results of phylogenetic studies reported as life strategies to survive on vertical cliffs showed that the closest relatives of P. palinuri are [1]. The most recurrent include: a) long living, slow often associated to cliffs of mountain areas. growing individuals; b) production of high number More specifically, it is now generally accepted of small propagules; c) high genetic variability. that P. palinuri belongs to the section Auricula Following, we analyse the occurrence of these Duby, a group endemic to high mountain ranges in features in P. palinuri. Europe, originated between 3.6 Ma and 2.4 Ma (i.e. Plants develop long superficial rhizome with a between Pliocene and Pleistocene) from an ancestor slow growth rate (less than 1 centimetre/year) [26]. coming from the centre of distribution of the Based on the rhizome length, we estimated that in the Sino-Himalayan region [37, 38]. most individuals present on Cilento cliffs are several tens of years old.

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Biometric data on fruit- and seed-set showed In the relict populations of P. palinuri, slow- that hundreds of seeds are produced by each growing old plants are highly predominant over inflorescence (mean number of seeds in short-styled young individuals. Together with all traits of the morph=562, SD=31; in long-styled morph=579, distyly syndrome and the development of large SD=78). Each seed is small and light (mean weight amounts of light seeds, these traits support the =0.35 mg, SD=0.08). hypothesis made by Davis [2] and Larson et al. [1] Although no data are available on the genetic that survival strategies of cliff species are high diversity of P. palinuri, we can infer that the co- genetic variability, high reproductive success and a occurrence of short-styled and long-styled flower large number of small propagules. morphs, with intra-morph incompatibility avoids inbreeding in favour of genetic variability [27].

Table 1. Endemic and sub-endemic species co-occurring with P. palinuri on coastal carbonate cliffs of Cilento.

Rocky Scientific name Chorotype Life form Regions slopes exclusive Eokochia saxicola (Guss.) Freitag and G. Endem. Ch suff cam, sic x Kadereit central/southern Italy; south- Brassica incana Ten. Subendem. Ch suff est Europe Campanula fragilis Cirillo subsp. fragilis Endem. Ch suff cam, bas, cal x Centaurea cineraria L. subsp. cineraria Endem. H scap cam, bas, cal, laz x central/southern Italy; south- Centaurea deusta Ten. Subendem. H bien est Europe longiflorus Raf. 5% Subendem. G b southern Italy; Malta cam, bas, cal, sic, western Dianthus rupicola Biv. subsp. rupicola Subendem. Ch suff x Mediterranean Helichrysum litoreum Guss. Endem. Ch suff central/southern Italy Iberis semperflorens L. Subendem. Ch suff abr, cam, sic, France x Limonium remotispiculum (Lacaita) Endem. Ch suff cam, bas, cal x Pignatti Primula palinuri Petagna. Endem. G rhiz/Ch cam, bas, cal x Seseli polyphyllum Ten. Endem. H scap laz, cam x

7 General discussion and concluding during the long-lasting interactions with humans remarks (about 8000 years). Present landscapes are the results of these processes and correct interpretations of the extant situations are possible only by means The IUCN has designated the Mediterranean Basin of a multidisciplinary approach. Our work has been as one of the planet’s biodiversity hotspots, because carried out within this context. this region has a high level of endemic species in All together data on geology, flora and species threatened status. Despite the international biology of coastal cliffs of Cilento suggest that recognition of the high level of plant diversity and ecological features of these cliffs are in conformity endemism of the Mediterranean Basin, much of its with those reported worldwide. Moreover, at the flora is unprotected from anthropogenic impacts best of our knowledge, this is the first assessment associated with economic activity such as tourism for a species of coastal cliffs of Cilento of the theory and land use. on life strategies reported for other cliff-hangers by The ancient floras of the Mediterranean region Larson et al. [1]. have been subjected to the selective pressures of the Along the Cilento coast, vertical cliffs show climatic fluctuations of the Quaternary and, more significant differences regarding several features recently, the original vegetation has been altered

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such as origin, lithology, exposure, and stability Phytogeographica Suecica Vol. 53, 1968, pp. conditions. Our overall analysis of geological and 1-153. floristic data of Cilento cliffs suggests that each of [6] Morisset P. Endemism in the vascular plants of these features plays an important role for the the Gulf of St. Lawrence region. Naturaliste survival of the numerous relict species still present Canadien Vol. 98, 1971, pp. 167-177. in these sites. Our floristic data suggest that vertical [7] Crawford R.M.M., Studies in Plant Survival. cliffs are home for a number of endemic species Blackwell Scientific Publications, Oxford, UK, much higher than in other coastal areas. However, 1989. these species are not equally distributed on cliff [8] Wiser S.K., High-elevation cliffs and outcrops faces. Detailed studies on single species biology, as of the Southern Appalachians: vascular plants in the case of P. palinuri, can explain the interaction and biogeography. Castanea Vol. 59, 1994, pp. with environmental factors and reveal which 85-116. functional features constrain local distribution of the [9] Cooper A., Plant species coexistence in cliff plants. Further work is desirable to establish and habitats. Journal of Biogeography Vol. 24, quantify specific interactions between each endemic 1997, pp. 483–494. species and the abiotic components of these cliffs. [10] Budetta P., Santo A., Vivenzio F., Landslide As in other Mediterranean areas, Cilento hazard mapping along the coastline of the coastline is highly involved in human activities, Cilento region (Italy) by means of a GIS-based therefore any action aimed to protect coastal parameter rating approach, Geomorphology, habitats is a real challenge for conservation Vol. 94, 2008, pp. 340-352. managers. [11] Budetta P., Application of the Swiss Federal Both natural and anthropogenic phenomena can Guidelines on rock fall hazard: A case study in affect the fine equilibrium of cliff ecosystems. the Cilento region (Southern Italy). Landslides Sudden rock falls, slow global warming phenomena Vol. 8, No. 3, 2011, pp. 381-389. together with direct impact due to touristic activities [12] Moggi, G., Catalogo della Flora del Cilento are some of the main threats. Protection of all the (Salerno). Repertorio delle piante vascolari sites along the Cilento coastline would be a mission finora segnalate e problemi sistematici unacceptable to achieve for both social and connessi. Inform. Bot. Ital. Vol. 33 No. 3, economic reasons. Clear indication of which local (2001), 2002, pp. 1-116. sites along the coastline deserve most concern for [13] Del Guacchio E., New data for the exotic flora biodiversity richness would be of much use for of Campania. Quad. Bot. Amb. Appl., Vol. 16, sustainable conservation actions. Identification of 2005, pp. 175-180. the sites, periodic monitoring of their conservation [14] Motti R., Salerno G., La flora del complesso status and guidelines for their safeguard should be dei monti Vesole, Soprano, Sottano e Chianello soon commissioned. To achieve these goals (appennino campano, SA) – Webbia, Vol. 61, effectively, once more, a multidisciplinary approach No. 2, 2006, pp. 325-357. is unavoidable. [15] De Natale A., Strumia S., La flora della costa sabbiosa del Parco Nazionale del Cilento e Vallo di Diano (Salerno). Webbia Vol. 62, No. References: 1, 2007, pp. 53-76. [1] Larson D.W., Matthes U., Kelly P.E., Cliff [16] Azzella M.M., Burrascano S., Notulae alla Ecology: Pattern and Process in Cliff checklist della flora vascolare italiana: 6 Ecosystems. University Press, Cambridge, UK, (1481). Inform. Bot. Ital., Vol. 40 No. 2, 2008, 2000. pp. 251-252. [2] Davis P.H., Cliff vegetation in the eastern [17] De Natale A., Di Nuzzo F., Crescenzi E., Note Mediterranean. Journal of Ecology, Vol. 39, di floristica per la Penisola Sorrentina, il 1951, pp. 63-93. Massiccio del Matese e specie notevoli per la [3] Polunin N., Artic plants in the British Isles. Campania. Inform. Bot. Ital., Vol. 40, No. 2, Journal of Botany Vol. 77, 1939, pp. 371-413. 2008, pp.243-248. [4] Bunce R.G.H., An ecological study of Ysgolion [18] Rosati L., Filibeck G., De Lorenzis A., Lattanzi Duon, a Mountain Cliff in Snowdonia. Journal E., Surbera F., Fascetti S., Blasi C., La of Ecology Vol. 56, 1968, pp. 59-75. vegetazione forestale dei Monti Alburni, nel [5] Lundqvist J., Plant cover and environment of Parco Nazionale del Cilento e Vallo di Diano steep hillsides in Pite Lappmark. Acta (Campania): analisi fitosociologica e

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