M. Erben, C. S. Christodoulou, R. Hand & K. Kefalas Limonium ammochostianum , L. karpasiticum and L. paralimniticum ( Plumbaginaceae ), three new endemic species from the eastern part of Cyprus

Erben, M., Christodoulou, C. S., Hand, R. & Kefalas, K.: Limonium ammochostianum , L. karpasiticum and L. paralimniticum ( Plumbaginaceae ), three new endemic species from the eastern part of Cyprus. — Fl. Medit. 32: 35-45. 2022. — ISSN: 1120-4052 printed, 2240-4538 online. Limonium ammochostianum , L. karpasiticum and L. paralimniticum ( Plumbaginaceae ) are described as new to science. The former species was long time known in Cyprus under the mis-applied name L. ocymifolium subsp. bellidifolium . The three species occur in a few coastal stretches in the east of the island. The new taxa show strong morphological similarities and can be considered as segregates of a L. ammochostianum species group. All hitherto known populations of the species group make up about 6000 individuals. L. ammochostianum and L. paralimniticum should be classified as “Endangered”, whereas L. karpasiticum as a “Data Deficient” taxon.


Introduction
The Limonium flora of Cyprus, the third largest island of the Mediterranean, is rather poor in species compared to areas with similar size located more to the west of the basin. Currently, eight species are known to occur, two of them, Limonium cyprium (Meikle) Hand & Buttler and L. mucronulatum (H. Lindb.) Greuter & Burdet, are endemic to the island (Hand & al. 2021). Karyological research considering the majority of taxa has been dealt with recently by Kouzali & al. (2012). At the southeast tip of the island, known as Kavo Gkreko (or Cavo Greco, Cape Greco, Akrotirio Gkreko), a rare taxon was first found in 1958 by Macdonald, which was named as L. ocymifolium subsp. bellidifolium (Sm.) Meikle in the "Flora of Cyprus" (Meikle 1985). Nowadays, most authors prefer the heterotypic species name L. aucheri (Girard) Greuter & Burdet. Few authors, e. g. Strid (2016), advocate lumping L. aucheri with L. ocymifolium (Poir.) Kuntze. There is also a short summary about the nomenclature of this L. aucheri by Kouzali & al. (2012). However, both names used in Cyprus are based on material collected in the Aegean region. Furthermore, not much is known about the phylogenetic relationship of all discussed taxa. L. aucheri and L. ocymifolium have not been included into the analysis by Malekmohammadi & al. (2017) but were integrated in the study by Koutroumpa & al. (2018). The latter study used accessions of L. aucheri and L. ocymifolium from Greece and Turkey respectively, which, according to the results, proved to be surprisingly remotely related. A recent paper by Koutroumpa & al. (2021) adds general aspects of origin and diversification of Mediterranean Limonium taxa.
Intensive field and herbarium studies revealed that the Cypriot plants differ considerably from plants collected in Greece as well as from all known taxa of the wider East Mediterranean region. As a result, we describe a new species group comprising three microspecies.

Material and methods
The current study is based on fieldwork at all known sites of the new taxa, as well as on studies of herbarium material in various collections (B, CYP, M).
Chromosome numbers have been counted according to the protocol described by Brullo & Erben (2011: 11). Taxonomy and nomenclature of taxa mentioned in the text follow Hand & al. (2021). Georeferenced data for all locations is stored in the database on endemic and endangered vascular plants of Cyprus kept at the Department of Forests (Lefkosia).

Results
The three new species share many morphological characters. A diagnosis for the species group is therefore first given to help distinguish it from other E Mediterranean taxa.

Limonium karpasiticum
Phenology: L. ammochostianum flowers earlier than the other two species, usually from (March-) April to early June; L. paralimniticum and L. karpasiticum flower from (April-) May to July.
Etymology: The epithet of the new species L. ammochostianum refers to Ammochostos district in the eastern part of Cyprus. The eponymous district capital, a few kilometers away from the type locality, is also well known under its colonial name Famagusta. The species L. paralimniticum is named after Paralimni municipality where its distribution area is confined. The name of L. karpasiticum is based on the Karpasia peninsula, the characteristic "panhandle" in the northeast of Cyprus.
Chromosome numbers: Kouzali & al. (2012) published three chromosome counts based on Cypriot material, two from the type region of L. paralimniticum in the Protaras area (Paralimni municipality) and one from Kavo Gkreko area, the type locality of the species L. ammochostianum. In all three cases they revealed the pentaploid number 2n=43. We have not seen the corresponding specimens, so that the identity with the newly circumscribed taxa needs to be clarified, especially as regards the latter case. However, our own countings based on plants grown from seeds of the type specimens (performed by ME) confirmed the pentaploid number 2n=43 for L. paralimniticum. In the other cases plants proved to be hexaploids with 2n=51 (type material of L. ammochostianum and L. karpasiticum, also plants from Galounia near Davlos).  All three new taxa are rare species of Cyprus, known so far from four small coastal stretches in the eastern part of the island (Fig. 11). In the past two decades, the coastline of Cyprus has been studied relatively intensively regarding the occurrence and distribution of Limonium. This resulted in the mapping of many new sites of L. cyprium, formerly thought to be rare along the northern coast (summary of records in Hand & al. 2021). Hitherto, the southeast coast of Cyprus has been intensively studied. L. ammochostianum is restricted to the coastline of Kavo Gkreko from "Palatia" sea caves to "Agioi Anargyroi" chapel, where the taxon has been known since more than 60 years. L. paralimniticum is known to occur along the Protaras coastline (Paralimni municipality). Because of the presence of extensive stretches of sandy beaches, no additional occurrences are to be expected in the Ammochostos gulf between Protaras and Bogazi. In the Karpasia peninsula, which follows to the northeast, L. karpasiticum has been found on the northern coast near the tip (Mora Psaria) and another location at the base of the peninsula (Galounia, Davlos), where also L. ammochostianum occurs sympatrically. In one case L. karpasiticum material has also been identified from Kavo Gkreko area.

Habitat
All three species inhabit maritime calcareous rocks, which correspond to habitat type 1240: Vegetated sea cliffs of the Mediterranean coasts with endemic Limonium sp. pl. (sensu Annex I of the EU Directive 92/43/EEC). Other accompanying species are Limonium virgatum (Willd.) Fourr., Frankenia hirsuta L., Crithmum maritimum L., Silene sedoides Poir., Atriplex portulacoides L. and sporadically Tetraena alba (L. f.) Beier & Thulin. In the wider Kavo Gkreko area, L. ammochostianum is mainly found on Quaternary marine terrace calcarenite and to a lesser extent on Quaternary marine terrace calcarenite with Tera Member limestone in places; L. paralimniticum, along Protaras coasts, grows on Quaternary marine terrace calcarenite as well as on Pachna Formation chalk and marl, in some places topped by Quaternary marine terrace calcarenite. In the north coast of Karpasia peninsula, L. karpasiticum colonises Kythrea formation sandstones and Quaternary marine terrace calcarenite (Geological Survey Department 1995).

Conservation
Undoubtedly, the L. ammochostianum species group is a rare taxon encompassing three species with restricted and fragmented distribution, small population and threatened habitat. The estimated population of the species group is around 6000 individuals (L. ammochostianum 4000, L. paralimniticum 1700 and L. karpasiticum 200 individuals respectively). Considering the current knowledge, the L. ammochostianum species group could be assessed as Vulnerable: B1ab(iii+v)&2ab(iii+v), because the Extent of Occurrence (EOO) is 116 km 2 , the Area of Occupancy (AOO) is 21 km 2 and the taxa of the species group occur at 6 locations; also, the quality of habitat has been degraded, at least at three of the locations, and as a result the population declined.
However, L. ammochostianum can be classified as Endangered: B1ab(iii+v)&2ab(iii+v) due to the number of locations (three), EOO (116 km 2 ), AOO (11 km 2 ) and the quality of habitat, which has been degraded at one location resulting in reduction of the population. Similarly, in the "Red Data Book of the Flora of Cyprus" (Tsintides & al. 2007), L. ammochostianum (= L. aucheri, including part of the population of L. paralimniticum) has been assessed as Endangered: B1ab(iii+v)&2ab(iii+v); this classification is still valid, although L. ammochostianum occurs almost entirely within a protected area, since Kavo Gkreko is a National Forest Park as well as a designated Natura 2000 site. L. paralimniticum could be also classified as Endangered: (B1ab(iii+v)&2ab(iii+v) based on the EOO (13 km 2 ), the AOO (7 km 2 ), the number of locations (2) and the continuing decline in area/extent and quality of habitat, resulting also in population decline; habitat degradation is due to extensive touristic development along the coastal zone of Protaras (Paralimni municipality). L. karpasiticum is hitherto known from two locations with a population of about 200 individuals, thus it could be also characterised as Endangered under criterion D (number of mature individuals <250) (IUCN Standards and Petitions Subcommittee 2014). However, owing to its recent discovery, the insufficient survey and one collection without exact locality, we believe it is more appropriate to be classified as Data Deficient.

Discussion
The new species are similar in habit, leaves and inflorescences to L. aucheri, a species which is frequent on many islands of the central and southern Aegean region and rare in Attika and Peloponnese (Greece; see also fig. 4). They differ from L. aucheri first of all by having fleshy, canaliculate in cross-section leaves (vs. flat leaves), larger (vs. 5.5-6.3 mm), more remotely arranged spikelets, greater bracts (vs. inner bract 4.3-5.2 × 3.2-3.9 mm) and often longer calyces (vs. 4.3-5.4 mm).
The known populations of the 3 new species differ slightly in morphological characters from each other but also in their flowering periods. Furthermore, in one of the four known coastal stretches, where the species occur, transitional plants have been identified. Most probably, the new taxa represent cases of rather recent speciation events. The differences between the three taxa are small, but they are easily diagnosable. Also, because of the almost parapatric distribution, classification as subspecies would be a viable solution, but in view of the current preferred practice in the Mediterranean area, we have opted for microspecies. Additional future fieldwork in some remote coastal areas of the Karpasia peninsula may improve the chorological knowledge as will do additional karyologial work. The somewhat complicated known distribution patterns, e. g. in L. ammochostianum which "brackets" the other species occurrences as well as the occurrence of transitional plants with or without the postulated parental taxa may be explained by relictual sites or long-distance dispersal by coastal currents in the past.