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Introduction
The genus Cistus contains 21 species, most of them with Mediterranean distribution or confined to Canary Islands (Guzmán et al., 2009). The highest diversity with 14 species occurs in the western Mediterranean: Iberian Peninsula and northwestern Africa (Guzmán & Vargas, 2005). In general, Cistus species are divided into two phylogenetic lineages: white-flowered species (12) which consist the subgenus and purple-flowered species (9). The groups of Cistus salviifolius (10 species) and C. clusii (2 species) make the white-flowered lineage (Guzmán et al., 2009). Sometimes these two groups are treated as subgenera Leucocistus Willk. and Halimiodes (Willk.) Demoly & P. Monts while purple-flowered species consist the third subgenus Cistus L. (Barrajón-Catalán et al., 2011; Lukas et al., 2021). Lately, Halimium (Dunal) Spach species were also included in the genus Cistus (von Raab-Straube, 2018) following the proven monophily of the Cistus-Halimium complex (Guzmán et al., 2009).
Cistus salviifolius radiated 0.82 ± 0.20 mya (million years ago) within the white-flowered Cistus lineage which started to appear 1.046 ± 0.25 mya and this diversification likely occurred after the development of Mediterranean climate 2.8 mya (Guzmán et al. 2009).
C. salviifolius is the most widespread species of Cistus genus and it is usually part of the understorey in forests and woodlands inhabiting many other habitats (Fernández-Mazuecos & Vargas, 2010). It spreads around the Mediterranean Basin, from Portugal to Greece, Turkey and Israel, extending to North Africa, though it may be locally disjunct and forming isolated population (Farley & McNeilly, 2004; Fernández-Mazuecos & Vargas, 2010). This species is insect-pollinated (entomophilous) (Moretti et al., 2020). The studies on the seed dispersal of Cistus salviifolius revealed no apparent mechanism of distant dispersal (Skourou & Arianoutsou, 2004; Tavsanoglu, 2010).
The genus Cistus was represented so far by only one taxon in the flora of North Macedonia –Cistus creticus L. ssp. eriocephalus (Viv.) Greuter & Burdet (=Cistus tauricus C. Presl.) which in the monograph of the flora of North Macedonia was presented sub. C. incanus L. (Micevski, 1995), a species with purple flowers. This taxon is distributed mostly in south-eastern parts of the country (Strumica, Dojran, Valandovo, Gevgelija and Demir Kapija) with additional occurrence in the areas of Mariovo, Prilep Kavadarci (Drenovo) and Jablanica (Micevski, 1995; Teofilovski, 2017). This species is also known from the Greek part of Prespa watershed (Strid et al., 2020). This species is characteristic and diagnostic of the association Diantho–Cistetum incani Micevski et Matevski ex Čarni, Matevski et Šilc, 2010 distributed in North Macedonia, Bulgaria and Greece (Čarni et al., 2010).
The presence of another Cistus species besides C. creticus ssp. eriocephalus in North Macedonia was expected when analysing the distribution maps of the other Cistus species in Albania and Greece (Barina et al., 2016; Strid, 2024). The aim of this paper is to present the first record of Cistus salviifolius in North Macedonia (Ohrid valley) as well as to describe its plant community, preference to geologic substrates and soils and discuss its conservation status.
Site description
The hills of Trojani are situated in the southwest part of North Macedonia, in the Ohrid valley, north of Lake Ohrid. It belongs to the Ohridsko Pole area -30834b according to the division of Melovski et al. (2013). Galičica mountain to the east and Jablanica mountain to the west surround the Ohrid valley.
The hilly massif of Trojani was named after a historic village Trojani that was situated above village Dolno Lakočerej. The highest peak is Gorenička Krasta also known as Bajrak (1203m). The lowest part of the Trojani hill massif is at altitude of 725 m which represents the floodplain formed by Lake Ohrid.
Lake sediments are present in the floodplain, while phylitic schists are dominant in the hilly area with smaller outcrops of alluvial or diluvial sediments, dolomites or marbled limestones (Dumurdzhanov & Ivanovski, 1972). The pedology is mainly represented by complex mix of cambisols, leptosols and regosols developing on phyllitic schists (MASIS, 2015).
The climate of Ohrid valley is influenced by its altitude (above 700 m), the presence of large Lake Ohrid and the corridor through created by the river Crn Drim. Rainfall in the Ohrid valley is determined by the Mediterranean pluviometric regime (Lazarevski, 1993).
Materials and Methods
Fieldwork at the locality Trojani (v. Dolno Lakočerej, Ohrid) was conducted on several short visits on 22.07.2023, 01.08.2023, 11.05.2024 and 18.05.2024. Photographs of the plants were taken by digital camera. The composition of the plant community was noted during the fieldwork on 18.05.2024 and some specimens were collected and later on deposited in the Macedonian National Herbarium at the Institute of Biology, Faculty of Natural Sciences and Mathematics, Ss. Cyril and Methodius University, Skopje, North Macedonia (Index Herbariorum code: MKNH) with the following accession numbers: 071912, 071913, 071914, 071915 and 071916.
The following is description of the specimens from Trojani (Figure 1) which is in accordance to the description in different floras. Shrubs, up to 1m high, procumbent. Twigs and branches reddish. Leaves ovate to elliptical, flat and woolly when young, somewhat undulate when old; petioles furrowed on the upper side. Cymes with 1–2 flowers. Flowers 3–5 cm, with long pedicels; petals white; sepals 5, the two outer sepals cordate at base. All of the aboveground parts have stellate hairs: twigs and branches, both sides of the leaves, petioles, pedicels and sepals.
Figure 1:
Cistus salviifolius L. (Sageleaf Rockrose), Trojani area. A – Habitus, B – Branch, C – Twig, D – leaves (upper side), E – leaf (lower side), F – Flowers, G – Flowers (side view), H – Flower with sepals, I – Fruit (from above), J – Fruit (side view), 22. 02 & 01. 08. 2023, 11. 05. & 18. 05. 2024 (photos: S. Hristovski).
Slika 1:Cistus salviifolius L. (kaduljelistni brškin), območje Trojani. A – habitus, B – veja, C – vejica, D – listi (zgornja stran), E – list (spodnja stran), F – cvetovi, G – cvetovi (stranski pogled), H – cvet s čašnimi listi, I – plod (pogled od zgoraj), J – plod (stranski pogled), 22.02 & 01.08.2023, 11.05. & 18.05.2024 (fotografije: S. Hristovski).
Locality
Specimens of Cistus salviifolius were recorded during the field visits (Figure 2):
Trojani, v. Dolno Lakočerej, Ohrid, N41.1618133°, E20.7922617°, 780 m a.s.l., Scrubland on eroded soil, 23.07.2023, leg. Slavčo Hristovski, Iskra Hristovska & Slobodan Hristovski.
Trojani, v. Dolno Lakočerej, Ohrid, N41.1641233°, E20.7948917°, 745 m a.s.l., degraded Quercetum frainetto-cerridis, 11.05.2024, leg. Slavčo Hristovski & Slobodan Hristovski.
Trojani, v. Dolno Lakočerej, Ohrid, N41.1622113°, E20.785548°, 745 m a.s.l., degraded Quercetum frainetto-cerridis, 11.05.2024, leg. leg. Slavčo Hristovski, Slobodan Hristovski, Marjan Komnenov, Renata Ćušterevska, Vlado Matevski.
Trojani, v. Dolno Lakočerej, Ohrid, N41.1643202°, E20.7852465°, 880 m a.s.l., degraded Quercetum frainetto-cerridis, 18.05.2024, leg. Slavčo Hristovski, Slobodan Hristovski, Marjan Komnenov, Renata Ćušterevska, Vlado Matevski.
Figure 2:
Distribution of Cistus salviifolius in North Macedonia ().
Slika 2: Razširjenost vrste Cistus salviifolius v Severni Makedoniji ().
Plant community
The community of Cistus salviifolius develops on silicate bedrock i.e. the majority of records are on phyllitic schists (Figure 3). Actually, silicate bedrock is the ancestral soil type of C. salviifolius lineage. Nevertheless, C. salviifolius is also known as a calcicolous species (Guzmán et al., 2009).
Figure 3:
Distribution of Cistus salviifolius at locality Trojani and preference of geologic substrate.
Slika 3: Razširjenost vrste Cistus salviifolius na območju Trojani in prevladujoča geološka podlaga.
Almost all of the records were on the soil complex of cambisols, leptosols and regosols according to the pedologic map of North Macedonia (MASIS, 2015). Actually, from the field visit we can conclude that the majority of Cistus individuals developed on eroded regosols (Figure 4).
Figure 4:
Distribution of Cistus salviifolius at locality Trojani and soil preference.
Slika 4: Razširjenost vrste Cistus salviifolius na območju Trojani in prevladujoči tip tal.
Figure 5:
Community of Cistus salviifolius at locality Trojani, 30.07.2024 (photo: S. Hristovski).
Slika 5: Združba z vrsto Cistus salviifolius na območju Trojani, 30. 07. 2024 (fotografija: S. Hristovski).
Cistus salviifolius is a representative species of typical Mediterranean habitats as garrigues or open marquises (Mucina et al., 2016). It is a name giving taxon to numerous phytocoenoses, most recently described by (Hardy et al., 2024). Depending on the geological substrate on which they develop (silicate or limestone), the corresponding plant communities are syntaxonomically separated into different syntaxonomic categories (Cisto-Lavanduletea stoechadis Br.-Bl., Ononido-Rosmarinetea Br.-Bl. etc.) (Mucina et al., 2016).
According to the research of Čarni et al. (2010), referring to the Mediterranean Cistus creticus subsp. creticus L. (=Cistus incanus subsp. creticus (L.) Heywood) – dominated community in Northern Greece (Palia Leptokaria, Krania, Halkidiki, Kavala), the species Cistus salviifolius is registered as a diagnostic species in the association Calicotomo villosae-Cistetum cretici [Hyperico olympici-Cistion cretici (Oberd. 1954) R. Jahn et Bergmeier in Mucina et al. 2009, Cisto-Micromerietalia julianae Oberd. 1954, Ononido-Rosmarinetea Br.-Bl. in A. Bolos y Vayreda 1950]. The composition of the community is dominated by numerous Mediterranean, semi-shrubby and herbaceous plants such as Calycotome villosa, Quercus coccifera, Cistus creticus subsp. creticus, Cytinus hypocystis subsp. clusii, Erica arborea, Hymenocarpus circinnatus, and others.
The flora and vegetation associated with Cistus salviifolius in the region of Ohrid (Trojani area), are significantly different from the flora and vegetation in which this species thrives in Northern Greece (Čarni et al., 2010). These are dry grasslands (all. Trifolion cherleri) that develop on a silicate substrate. They are in the process of being regrown from the adjacent forest phytocoenosis dominated by Quercus frainetto, Quercus pubescens, Carpinus orientalis, Juniperus deltoides, Ligustrum vulgare and others. The following taxa were registered in the investigated Cistus salviifolius population: Achillea coarctata, Agrostis castellana, Aira elegantissima, Carpinus orientalis, Centaurea grisebachii, Clinopodium alpinum subsp. hungaricum, Dorycnium pentaphyllum subsp. herbaceum, Eryngium campestre, Festuca valesiaca, Filago germanica, Fumana procumbens, Galium divaricatum, Hypericum olympicum, Hypericum perforatum, Juniperus deltoides, Lathyrus cicera, Lembotropis nigricans, Ligustrum vulgare, Linaria pelisseriana, Lonicera etrusca, Micropyrum tenellum, Myosotis stricta, Ornithopus compressus, Pilosella bauhini, Poa bulbosa, Psilurus incurvus, Quercus frainetto, Quercus pubescens, Rumex acetosella, Sanguisorba minor subsp. balearica, Sedum amplexicaule, Silene sp., Stachys angustifolia, Thymus longicaulis subsp. longicaulis (=Thymus rohlenae), Thymus sibthorpii, Trifolium arvense, Trifolium campestre, Trifolium hirtum, Veronica chamaedrys, Vicia lathyroides, Vicia villosa aggr.
This region is influenced by a mild continental, i.e. modified (sub)Mediterranean climate which favours development of a large number of Mediterranean and sub-Mediterranean plant species and plant communities (Em et al., 1985; Matevski & Kostadinovski, 1996). This is also the case with the finding of the species Cistus salviifolius in this area, which joins the large number of Mediterranean and sub-Mediterranean plant species in the wider area around lakes Ohrid and Prespa, such as Asparagus acutifolius, Biarum tenuifolium, Buxus sempervirens, Clematis viticella, Convolvulus althaeoides subsp. tenuissimus, Hyssopus officinalis var. pilifer, Jasminum fruticans, Juniperus excelsa, Ephedra fragilis subsp. campylopoda, Phyllirea latifolia, Pistacia terebinthus, Sternbergia colchiciflora and others (Matevski & Kostadinovski, 1996; Matevski et al., 2011).
Distribution
Cistus salviifolius shows low genetic differentiation with the same haplotype lineages in Europe and Africa, which is due to at least three intercontinental colonizations (even though the plant is apparently unable to spread over long distances). A recent (Pleistocene) European route of colonization between Iberia and Anatolia through France and Italy was proposed in order to explain the low haplotype diversity (Fernández-Mazuecos & Vargas, 2010). Moretti et al. (2020) showed that marginal or peripheral populations of Cistus salviifolius have similar genetic diversity as central populations.
In Bulgaria, Cistus salviifolius is distributed only in Strandzha at the border with Turkiye (Stoyanov et al., 2021). From the Balkan countries it is absent from Serbia and Romania. In Albania the species is distributed along the Adriatic and Ionian coasts with few inland areas in the central and southern parts of the country, at altitudes between the sea level and 800 m (Barina et al., 2016). The two closest populations in Albania in west and southwest direction (Barina et al., 2016) are more than 60 km away from Trojani area.
Thus, the finding of the primarily Mediterranean species Cistus salviifolius in the Ohrid area is very intriguing. However, other species with (Sub)Mediterranean distribution are known to occur in the Ohrid valley i.e. the upper watershed of river Crn Drim in North Macedonia, such as Moltkia petraea, Salvia officinalis, Cistus creticus ssp. eriocephalus, etc. (Em et al., 1985; Matevski, 2013; Teofilovski, 2017). In fact, the area of river Crn Drim valley is part of the Ohrid-Prespa refugial region (Em et al., 1985). This region is under the influence of modified sub-Mediterranean climate (Melovski et al., 2013) which provides conditions for development of rich flora and vegetation with presence of many relict and endemic species (Micevski & Matevski, 1987; Matevski & Kostadinovski, 1996). It is obvious that new findings of rare species in the south-western parts of North Macedonia (Galičica and Jablanica mountains with the Ohrid valley in between) improve our knowledge of this flora with predominantly Eastern Mediterranean distribution (Aegean-Ionian-Adriatic part). Additional phytogeopgrapical research combined with genetic approach will shed light on the pathways of distribution of these species such as river valley of Crn Drim as well as present and past ecological conditions (e.g. general climate, glaciation).
Conservation status
The area of Trojani has never been a subject of floristic investigation which explains why the species was not recorded so far in North Macedonia. The population of Cistus salviifolius in Trojani is obviously isolated from the main distribution of the species, but we believe it has natural origin.
In Albania and Greece, the species is not included in their respective red lists. However, in Bulgaria is was assessed as Endangered [EN B1ab(iii)+2ab(iii)] (Assyov & Denchev, 2015).
The population in Trojani area represents the only known locality in North Macedonia. Its extent of occurrence is about 62 ha (=0.62 km2). The main threat to the species is probably the afforestation with non-native species, possible expansion of the agricultural land (vineyards) and construction of forest roads. Thus, its status can be assessed as Endangered [EN B1a+2a] according to the IUCN Red List categories and criteria (IUCN Species Survival Commission, 2012).
Orcid-Profil
