The circumscription and distribution of Limonium cornarianum

In the area of Moni Kapsa (Fig. 1), the narrow endemic Limonium cornarianum was originally described by Kypriotakis & Artelari (1998) from the calcareous rocks below the Kapsa monastery and the nearby gorge (named Moni Kapsa or Perivolakia gorge) that has an opening to the coast. Along the gorge, the species grows in crevices of steep cliffs (Fig. 2A) up to a distance of 1.5 km from the seashore (Kypriotakis & Artelari 1998). Later collectors (see e.g. specimens in Brullo & Erben 2016 and photos in Kypriotakis & Artelari 2009) have assigned also Limonium plants from the coastal habitats of Moni Kapsa to L. cornarianum. During our fieldwork, we observed that the individuals growing along the coast (Fig. 3) were morphologically different from the ones inside the gorge. The coastal populations fit the description of L. hierapetrae Rech. f. that has a wider distribution in SE Crete (Rechinger 1943; Artelari 1989c). These observations were verified by (1) morphologically examining and comparing specimens from Moni Kapsa and the surrounding areas, including the type material of L. cornarianum and L. hierapetrae, and (2) analysing samples from the Moni Kapsa gorge and the coast (Kapsa beach) in a phylogenomic study that revealed their placement in different highly supported clades (Koutroumpa 2020). The sample representing the coastal population is nested in the well-supported L. hierapetrae clade (Koutroumpa 2020). The confusion of the L. hierapetrae samples from Kapsa beach with L. cornarianum explains the significant differences between the species description given by Brullo & Erben (2016: 101) and the protologue of L. cornarianum by Kypriotakis & Artelari (1998: 144). Brullo & Erben (2016) did not see the holotype of L. cornarianum and assigned further populations growing along the coast from Ierapetra to Moni Kapsa to L. cornarianum (see, e.g., B [ B 10 1189131]), which explains the much wider geographical distribution given by these authors for L. cornarianum that clearly overlaps with the known range of L. hierapetrae. In addition, the chromosome number 2n = 43 given for L. cornarianum (Brullo & Erben 2016) from material collected in Moni Kapsa matches the chromosome number of the pentaploid L. hierapetrae (2n = 43, Artelari 1989c), whereas L. cornarianum is triploid according to our initial studies (see below). The material used for the chromosome counts by Brullo & Erben (2016) was most probably collected from the coast and belongs to L. hierapetrae (see B [ B 10 1132915]). Taken together, in the area of Moni Kapsa both L. cornarianum and L. hierapetrae co-occur (Fig. 1) but occupy rather distinct habitats: L. cornarianum is restricted to crevices of calcareous rocks (Fig. 2A) and L. hierapetrae grows on the nearby pebbly beach (Fig. 3) and in the maritime calcareous cliffs next to the beach. The two species can be easily distinguished by their different leaf shape (Fig. 4B, C) and the smaller spikelets, spikelet bracts and calyces of L. cornarianum compared to L. hierapetrae (Table 1).

Fig. 3.

Limonium hierapetrae growing on gravely soil at the coastal area of Moni Kapsa (Crete), habit and inflorescence. – All photographs taken by and © Konstantina Koutroumpa.

Until now, Limonium cornarianum was known only from its locus classicus (Moni Kapsa) in SE Crete, where it is rare, comprising only few individuals (Kypriotakis & Artelari 1998, 2009). The fieldwork in Crete revealed a new population of L. cornarianum in the NE part of the island (Fig. 1, 2B–D). The population is geographically and ecologically distinct from the one in Moni Kapsa and located at the E edge of Pacheia Ammos cove in a dry salt-marsh habitat (Fig. 1, 2B). Only few individuals (c. 10) were found, occupying a small area that seems to be seasonally flooded and is the mouth of a stream that runs through the gorge of Cha, which is located about three kilometres from the seashore (Fig. 2B). Because L. cornarianum is a chasmophyte of steep, usually inaccessible cliffs free of other vegetation in its locus classicus, it could be assumed that the species might also occur inside the Cha gorge and that the population in Pacheia Ammos was formed by seeds washed down the stream that runs through the gorge. Cha is a narrow inaccessible gorge with high vertical cliffs and chasmophytic vegetation that is botanically not well explored because its passage requires canyoning expertise. The individuals of the newly found population in Pacheia Ammos undoubtfully represent L. cornarianum (Fig. 2C, D). They are morphologically indistinguishable and form a well-supported monophyletic group in the phylogeny with individuals from the type locality (Koutroumpa 2020). In addition, preliminary flow cytometric data of L. cornarianum from Pacheia Ammos indicate that the species is likely a triploid. Specifically, the 1C genome size of L. cornarianum is 2.37 pg, which is comparable to the genome size of the triploid L. virgatum (1C = 2.61 pg) and considerably smaller than the tetraploid L. aegaeum Erben & Brullo (1C = 3.15 pg) and the pentaploid L. sitiacum Rech. f. (1C = 3.52 pg). A triploid chromosome count is also found in L. aphroditae R. Artelari & Georgiou, an endemic in Kythira island that represents the closest morphological relative of L. cornarianum (Artelari & Georgiou 1999). Both species constitute a distinct morphological group compared to the other Aegean Limonium species, characterized by its small spikelet traits and sharing similarities with the C Mediterranean L. minutiflorum group that includes triploid and tetraploid taxa (Artelari & Georgiou 1999). The close relationship of L. cornarianum and L. aphroditae was further supported by phylogenomic data (Koutroumpa 2020).

The circumscription of Limonium hierapetrae

Limonium hierapetrae was described by Rechinger (1943) from a sandy beach next to Ierapetra in SE Crete. This endemic species was later found in a further five localities between Makrygialos and Nea Myrtos in the same wider area by Artelari (1989c), who studied the bio-systematics of the species, recorded its morphological variation and provided a detailed species description. Brullo & Erben (2016) described L. minoicum Erben & Brullo as a new species from SE Crete occurring between Tertsa and Tsoutsouros, a few kilometres W of the then known distribution of L. hierapetrae. According to the authors, the new species differs from L. hierapetrae by its shorter floral spikelets, inner bracts and calyces. However, Brullo & Erben (2016) gave a different description for L. hierapetrae than the one previously published by Artelari (1989c). Specifically, the measurements given for the sizes of leaves, spikes, spikelets, bracts and calyces by Brullo & Erben (2016) were only a subset of the size ranges given for L. hierapetrae by Artelari (1989c), namely the upper size ranges were assigned to L. hierapetrae and the lower size ranges were assigned to the newly described L. minoicum, for all the characters mentioned above. When Brullo & Erben's (2016) descriptions were employed to identify specimens collected along the coast between Moni Kapsa and Dermatos-Tsoutsouros (representing the entire range of L. hierapetrae and L. minoicum), several of them belonging to previously identified L. hierapetrae populations were assigned to L. minoicum or L. cornarianum sensu Brullo & Erben (2016), and many other specimens could not be unambiguously identified with either of them, but all perfectly match Artelari's (1989c) concept of L. hierapetrae. The phylogenomic analysis confirms that samples of L. hierapetrae and L. minoicum form one well-supported lineage, with L. minoicum specimens from the type locality and the population in Dermatos (bank of the river Anapodaris) intermingled with L. hierapetrae individuals from Moni Kapsa beach, Ferma and Ierapetra (i.e. the two species were not reciprocally monophyletic; Koutroumpa 2020). Additionally, chromosome counts of L. minoicum material with provenance from its locus classicus, M. Erben Li-1708 (FR [ FR-0128958], B [ B 10 1009652]), share the same chromosome number (2n = 43) with L. hierapetrae (Brullo & Erben 2016). Combining the morphological, molecular and chromosomal evidence, it becomes clear that L. minoicum is conspecific with L. hierapetrae. Also, L. chrisianum Brullo & Guarino, described from the islet of Chrysi S of Ierapetra, is conspecific with L. hierapetrae and represents just a small form of it, because the morphological features previously used to differentiate it from L. hierapetrae (e.g. smaller size and leaf length, dense arrangement of spikelets and larger middle bract width, Brullo & Guarino 2000) clearly fall within the range of variation of the latter species.

Fig. 4.

Leaf shapes of the three closely related Limonium species in SE Crete – A: L. artelariae (K. Koutroumpa 299, UPA); B: L. cornarianum (K. Koutroumpa 751, Z [Z-000278454]); C: L. hierapetrae (K. Koutroumpa 269, Z [Z-000278465]). – Scale bar = 10 mm.

A new endemic Limonium species in SE Crete

During the fieldwork in summer of 2014, an impressive population of Limonium was found in SE Crete, in the area between Myrtos and Tertsa, comprising tall plants up to 110 cm, bearing large, broadly spathulate rosette leaves (Fig. 5). Resembling L. hierapetrae in terms of habit and inflorescence, its leaves are of a different shape, resembling more those of L. cornarianum (Fig. 4A, B). However, plants of such height and with leaves of such size found in that population are unique among the Limonium species in Greece belonging to the “Mediterranean lineage”, i.e. all Greek Limonium species apart from eight belonging to the morphologically and phylogenetically distinct L. sect. Limonium (L. vulgare group), L. sect. Nephrophyllum Rech. f. s.l. and L. sect. Pterocladus (Spach) Bokhari (Koutroumpa & al. 2018). Three representatives sampled in a phylogenomic study formed a well-supported monophyletic group in close evolutionary relationship with L. cornarianum and L. hierapetrae but clearly distinct from them (Koutroumpa 2020). Apart from the large population found during the fieldwork, a careful study of previously collected herbarium specimens revealed that material collected from Cape Theophilos (Brullo & Giusso s.n., herb. Erben) at the E edge of Sidonia beach, which is located to the W of Tertsa, also belong to the new species. In addition, plants grown from seeds of the Cape Theophilos collection were studied karyologically by M. Erben (Li-1721) who counted the pentaploid chromosome number 2n = 43 in a cultivated individual (voucher deposited in FR [ FR-0128957]). The pentaploid chromosome number of the new species agrees with the preliminary flow cytometric analysis of material collected from Vatos and Kallikovrechtis, two localities between Myrtos and Tertsa. Specifically, the 1C genome size of the new species is 3.67–3.69 pg, which is comparable to the genome size of the pentaploid (2n = 43) L. sitiacum (1C = 3.52 pg) but larger than the genome sizes of the tetraploid (2n = 34) L. aegaeum (1C = 3.15 pg) and the triploid (2n = 27) L. virgatum (1C = 2.61 pg) and smaller than the genome size of the hexaploid L. creticum (1C = c. 4 pg; this an approximate count because this genome size was partially overlapping with the Pisum sativum reference genome, 1C = 4.42 pg). In the Taxonomic treatment below, the new species is described and compared with the closely related L. cornarianum and L. hierapetrae, for which emended species descriptions are provided based on the plethora of material seen and examined, including the type specimens.

Fig. 5.

Limonium artelariae growing on fine marls of coastal cliffs (A), habit (B) and inflorescence (C). – All photographs taken by and © Konstantina Koutroumpa.

Limonium cythereum newly recorded for Crete

The endemic Limonium cythereum R. Artelari & Georgiou was until now known only from the islands of Kythira and Antikythira (Artelari & Georgiou 1999; Brullo & Erben 2016), which are located SE of Peloponnese and between Kythira and NW Crete, respectively (Fig. 6). Apart from the three known populations of L. cythereum on the S and E coasts of Kythira (Chalkos, Vrouleas, and Avlemon) and the single population on the N coast of Antikythira (Potamos), the fieldwork revealed the occurrence of L. cythereum in two localities in NW Crete (briefly mentioned in Koutroumpa 2020). The species was recorded from a small port at the E edge of the city of Chania (see, e.g., B [ B 10 1163863]) and Kalathas beach, which is few kilometres NE of Chania (see, e.g., B [ B 10 1163868]). Limonium cythereum is easily distinguished from the closely related L. sieberi Kuntze, which also occurs in NW Crete and co-occurs with L. cythereum at Kalathas beach, by its shorter spikelets, the hairy and shorter inner spikelet bracts and the shorter calyces. The current distribution of L. cythereum reflects the phytogeographical connections of Crete and Kythira and the paleogeography of the region during the Pleistocene (c. 0.8 Mya) when low sea levels during the glacial period reconnected islands with each other and some islands with the mainland (see maps in Valakos 2008).

Limonium fragile was erroneously reported for Crete

Limonium fragile Erben & Brullo is a small species, up to 15 cm tall, bearing delicate branches, and has been reported from only few localities in Karpathos, Crete and Antikythira islands by Brullo & Erben (2016). Its occurrence in Crete is based on a specimen collected from the saline habitats of Olous near Elounda in NE Crete (Hansen 1066, PAL-Gr!; Brullo & Erben 2016). The specimen was originally identified as L. virgatum by W. Greuter in 1974, but Brullo & Erben (2016) later assigned it to L. fragile possibly on the basis of its small size. Hybrids between L. fragile and L. virgatum have also been reported from Crete by Brullo & Erben (2016), based on material collected from the same locality (i.e. salt marshes in Olous, H. Kalheber 99-192, herb. Kalheber!). The latter specimen comprises immature individuals collected in early spring. Olous and the surrounding areas in Elounda were visited twice during the fieldwork, but an extensive search failed to find representatives of L. fragile. However, a very large population of L. virgatum was observed with plants of different sizes, some of them very small, growing on the saline basins that were seasonally flooded. All the material collected from this region clearly represents L. virgatum. Field observations and the examination of several specimens from Elounda, including the Cretan L. fragile specimens seen by Brullo & Erben (2016), support the view that small and immature individuals of L. virgatum from Olous were misidentified as L. fragile or L. fragile × L. virgatum by Brullo & Erben (2016). Therefore, L. fragile is to be regarded as absent from Crete and plants growing in Olous (Elounda) in NE Crete belong to L. virgatum.

Fig. 6.

Distribution map of Limonium cythereum in Kythira, Antikythira and Crete.

Taxonomic treatment

Limonium artelariae Koutr., sp. nov. – Fig. 4A, 5, 7, 8. Holotype: Greece, Crete, prefecture of Lasithi, beach of the village Vatos, between Myrtos and Tertsa, 34.99686°N, 25.55936°E, rocks and sand, steep, Aug 2014, K. Koutroumpa 300 (Z [Z-000278447]; isotype: B [ B 10 1161811]).

Diagnosis — Limonium artelariae is distinguished from its closely related species in SE Crete, L. cornarianum and L. hierapetrae, by its broadly spathulate, rounded leaves and broadly obovate and wider inner spikelet bracts. Limonium artelariae is rather unique among all other Aegean Limonium endemics for its predominantly tall habit combined with longer and wider, spathulate leaves.

Description — Plants perennial, 48–110 cm tall, glabrous, with several erect stems. Caudices 2–5 cm long, branched with age and then plant with numerous leaf rosettes. Rosette leaves fleshy, light green to glaucous, flat or irregularly folded, rugose to verrucose, (39–)50–165(–210) × 12–43 mm, broadly spathulate, more or less abruptly tapering into petiole, with petiole usually much longer than lamina, margin cartilaginous, sometimes undulate about 0.2 mm wide, apex rounded, mucronate, mucro 0.3–1 mm long, curved backward; 1 central nerve with 2 lateral nerves in large leaves. Stems numerous, erect, usually robust, rugose, branched usually above lower 1/4–1/3; branches very rarely proliferous. Inflorescence trullate to obtrullate in outline; sterile branches absent or 1–3 at lower part of stem, 15–80 mm long, straight; fertile branches straight or slightly arched, erect, directed obliquely upward or spreading. Spikes (12–)15–55(–73) mm long, straight to slightly arched. Spikelets 5.5–6.8 mm long, densely arranged, 3 or 4(or 5) per cm, composed of 2–4(–8) flowers. Outer bract (1.8–)2–2.6 × (1.6–)1.8–2.1(–2.3) mm, glabrous, ovate-triangular, margin broadly membranous, to 0.6 mm wide, apex acute; central part herbaceous, forming a short point. Middle bract 2.2–2.5 × 1.5–2 mm, glabrous, membranous, oblong-elliptic, apex rounded or sometimes emarginate. Inner bract 4.3–5.5 × 3.8–4.5 mm, glabrous, broadly obovate, margin membranous, 0.6–1.2 mm wide, apex rounded; central part herbaceous, forming a short point to 0.5 mm long, indistinct in mature spikelets. Calyx 4.5–5.5 mm long; calyx tube more or less densely long hairy; calyx lobes 0.5–0.6 mm long, ovate-triangular; tube ribs ending usually above base of lobes. Corolla lilac.

Note — Plant height and leaf size (length and width) measurements were taken from several individuals in the field, at Vatos and Kallikovrechtis (Crete), though not all these individuals were collected as herbarium specimens.

Chromosome number and ploidy — 2n = 43 (chromosome count by M. Erben (Li-1721) on a cultivated plant of wild source; voucher: FR [ FR-0128957]); ploidy level: 5x (inferred from preliminary flow cytometric results of genome size).

Breeding system — Apomixis is inferred from the monomorphic population with self-incompatible pollen-stigma combination B. In addition, the species is of high ploidy level with an odd chromosome number further supporting the view of an apomictic reproduction, following Erben (1978, 1979).

Distribution — Endemic to SE Crete, along the coast between the villages of Myrtos and Tertsa (replaced by Limonium hierapetrae in Tertsa), and at Cape Theophilos at the E edge of Sidonia beach (Fig. 1).

Habitat — Maritime cliffs of marly, calcareous, conglomerate and sandy substrates.

Eponymy — The specific epithet “artelariae” honours Dr. Rea Artelari, a former Associate Professor of the University of Patras, Greece, for her invaluable contributions to the systematics of Limonium in Greece and Cyprus throughout her academic career.

Taxonomic notes — The new species is morphologically most closely related to Limonium hierapetrae, which grows in the same area of SE Crete. However, L. artelariae has large, broadly spathulate, rounded leaves compared to the smaller, oblanceolate to oblong-obovate, mostly obtuse leaves of L. hierapetrae. The individuals of the latter are also smaller and have usually narrower, elliptic to oblong-obovate inner spikelet bracts (vs broadly obovate in L. artelariae; Table 1). The dense leaf rosettes with spathulate, sometimes irregularly folded leaves, the densely arranged inflorescences and the obovate shape of the inner spikelet bracts morphologically link L. artelariae to L. cornarianum, which also grows in SE Crete. Further, L. cornarianum is clearly distinguished from L. artelariae by its smaller spikelets and spikelet bracts, and leaves with an acute apex (Table 1, Fig. 4).

Limonium artelariae is also related to other representatives of the informal taxonomic “L. sieberi” and “L. ocymifolium” groups sensu Brullo & Erben (2016). Specifically, L. artelariae shows similarities in habit, absence of or very low number of sterile branches, shape of leaves and inflorescences with L. aucheri (Girard) Greuter & Burdet, L. creticum, L. cythereum and L. sieberi also occurring in Crete, and also with L. pigadiense (Rech. f.) Rech. f. from Karpathos and L. ocymifolium (Poir.) Kuntze from the Kyklades. However, several morphological characters clearly separate L. artelariae from these allied species. Specifically, L. aucheri, which has a broad Aegean distribution but occurs in a single locality in NW Crete, is considerably smaller (10–40 cm tall), with narrower leaves (5–10 mm wide) and obovate, narrower inner spikelet bracts (3.2–3.9(–4.1) mm wide; Brullo & Erben 2016). The Cretan endemic L. creticum has partially smaller (20–85 × 5–20 mm), obovate-spathulate, obtuse leaves, longer spikelets (6.5–7 mm long) with longer but narrower (6–6.8 × 3.4–3.7 mm) elliptic inner bracts and longer calyces (5.6–6.2 mm long; Artelari 1989b). Limonium cythereum which occurs in Kythira, Antikythira and is newly found in NW Crete, is clearly differentiated by its hairy spikelet bracts (Artelari & Georgiou 1999; Brullo & Erben 2016). Limonium sieberi is smaller (20–50 cm tall) and has longer spikelets (6.5–9 mm long), inner spikelet bracts (5.2–7 mm long) and calyces (5.2–6.8 mm long; Brullo & Erben 2016). The Karpathos endemic L. pigadiense is also smaller (20–50 cm tall), with narrower (5–15 mm wide), oblanceolate-spathulate leaves, longer spikelets (6.6–7.3 mm long), inner spikelet bracts (5.6–6 mm long) and calyces (5.2–6.5 mm long; Brullo & Erben 2016). And finally, the Kykladean endemic L. ocymifolium is significantly smaller (15–35 cm tall) bearing shorter and somewhat narrower leaves (25–50 × 9–20 mm), mostly longer spikelets (6.5–7.5 mm long), inner spikelet bracts (5.1–5.9 mm long) and calyces (5.2–6 mm long), and significantly wider outer spikelet bracts (2.5–3 mm wide; Brullo & Erben 2016).

Limonium artelariae has the same ploidy level and chromosome number (2n = 5x = 43) as L. aucheri (Brullo & Erben 2016), L. hierapetrae (Artelari 1989c; Brullo & Erben 2016), L. ocymifolium (Artelari 1989a; Brullo & Erben 2016), L. pigadiense (Brullo & Erben 2016) and L. sieberi (Artelari & Georgiou 2003; Brullo & Erben 2016), while L. cornarianum is probably triploid (see above) and both L. creticum and L. cythereum are hexaploids but with different chromosome numbers (2n = 6x = 51 and 2n = 6x = 52, respectively; Artelari 1989b, 1992; Artelari & Georgiou 2003).

Additional specimens seen — GREECE, CRETE, PREFECTURE OF IRAKLIO: Akro Theophilou, calcareniti conglomeratiche, 27 Jun 2002, Brullo & Giusso s.n. (herb. Erben); cultivated from wild source, Akro Theophilou, calcareniti conglomeratiche, 27 Jun 2002, Brullo & Giusso s.n., M. Erben Li-1721 (B [ B 10 0710890], FR [ FR-0128957]); between Kallikovrechtis and Tertsa, 34.98966°N, 25.54750°E, marls and sand, Aug 2014, K. Koutroumpa 307 (Z [Z-000278446]), K. Koutroumpa 308 (B [ B 10 1161812,  B 10 1243313], Z [Z-000278441]). — PREFECTURE OF LASITHI: beach of village Vatos, between Myrtos and Tertsa, 34.99686°N, 25.55936°E, rocks and sand, steep, Aug 2014, K. Koutroumpa 299 (UPA), K. Koutroumpa 301 (ATH); Vatos, 34.99675°N, 25.55969°E, habitat with Tamarix, Poaceae, Pancratium maritimum and Pistacia lentiscus, on sand-marls, on a ‘carpet’ of dried Posidonia, Aug 2014, K. Koutroumpa 754 (Z [Z-000278445]), K. Koutroumpa 755 (Z [Z-000278442, Z-000278443, Z-000278444]).

Fig. 7.

Holotype of Limonium artelariae, K. Koutroumpa 300 (Z [Z-000278447]). – Image by United Herbaria Z+ZT /CC BY 4.0.

Table 1.

Morphological diagnostic features of Limonium artelariae, L. cornarianum and L. hierapetrae.

Limonium hierapetrae Rech. f. in Denkschr. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 105(2, 1): 104. 1943. – Lectotype (designated by Brullo & Erben 2016: 99): Greece, Creta, distr. Hierapetra: in litore arenoso a porto Hierapetra occidentem versus, 18 May 1942, K. H. Rechinger 13080 (W [accession no.  W1960-0012705]!; isolectotypes: G [ G00440217]!, M!). – Fig. 3, 4C.

= Limonium chrisianum Brullo & Guarino in Fl. Medit. 10: 269. 2000. – Holotype: Hrisi (Ierapetra, Creta), 25 Aug 1996, Brullo & Guarino s.n. (CAT; isotype: FR [ FR-0030972]!).

= Limonium minoicum Erben & Brullo in Phytotaxa 240: 107. 2016. – Holotype: Creta, Tertsa (Ierapetra), rupi calcarenitiche, 27 Jun 2002, Brullo & Giusso s.n. (MSB [MSB-164247]!; isotypes: B [ B 10 1189106]!, CAT, FR [ FR-0030969]!).

Description — Plants perennial, (10–)20–45(–80) cm tall, glabrous, with several erect stems. Caudices 1–7 cm long, branched with age and then plant with numerous leaf rosettes. Rosette leaves fleshy, pale grey-green, flat, coriaceous, verrucose to rugose, (8–)15–80(–100) × 3–16(–18) mm, oblanceolate to oblong-obovate, gradually tapering into petiole, with petiole longer than or equal to lamina, margin cartilaginous, sometimes undulate, c. 0.3 mm wide, apex obtuse (sometimes almost rounded) to slightly acute, mucronate, mucro 0.2–0.7(–1) mm long; 1 central nerve with 2 lateral nerves in large leaves. Stems numerous, erect, verrucose-rugose, branched usually above lower 1/4–1/3. Inflorescence paniculate, trullate or obtrullate in outline; sterile branches absent or 1–4 at lower part of stem, 10–70 mm long, straight; fertile branches straight or slightly arched, directed obliquely upward. Spikes (6–)15–120 mm long, straight to slightly arched. Spikelets 4.8–7 mm long, more or less laxly arranged, 1–5 per cm, composed of 1–4(–6) flowers. Outer bract (1.4–)1.5–2.6(–3) × 1.5–2.1 mm, glabrous, ovate-triangular, margin membranous, apex acute; central part herbaceous, forming a short point. Middle bract 1.7–2.9 × 1.4–1.9 mm, glabrous, membranous, oblong-elliptic, apex rounded or sometimes emarginate. Inner bract (4–)4.2–5.8(–6) × 2.7–4 mm, glabrous, elliptic to oblong-obovate, margin membranous, c. 1 mm wide, apex obtuse to rounded; central part herbaceous, forming a short point 0.5–0.7 mm long. Calyx 4–5.6 mm long; calyx tube usually unilaterally long hairy; calyx lobes 0.4–0.6 mm long, ovate; tube ribs ending at or above base of lobes. Corolla bluish-lilac.

Fig. 8.

Paratype of Limonium artelariae, K. Koutroumpa 754 (Z [Z-000278445]). – Image by United Herbaria Z+ZT /CC BY 4.0.

Chromosome number and ploidy — 2n = 43; ploidy level: 5x (Artelari 1989c).

Breeding system — Apomixis was inferred from the monomorphic populations with self-incompatible pollen-stigma combination B, low pollen fertility, high seed production, high ploidy level and odd chromosome number (Artelari 1989c).

Distribution — Endemic to SE Crete, growing along the coast between Moni Kapsa and Dermatos and on the neighbouring islets of Chrysi and Fotia (Fig. 1).

Habitat — Maritime calcareous, conglomerate and marly rocks, sandy and pebbly substrates.

Additional specimens seen — GREECE, CRETE, PREFECTURE OF IRAKLIO: Eparchia Viannou, Anapodaris-Küstenhof, 34°59′03″N, 25°19′30″E, 1–2 m ü.N.N, offenes Lycium schweinfurthii-Atriplex halimus-Tamarix-Gebüsch der Küste, 15 Oct 1998, N. Böhling 9019 (B [ B 10 0193536]); Ep[archia]. Viannos, Mündung des Anapodharis, 34°59′N, 25°19′30″E, c. 1 m, Flussbett mit Sand- und Kieselflächen, 19 Sep 1984, H. Risse 1517 (B [ B 10 1155993]); Dermatos village, at bank of river Anapodaris, 34.98328°N, 25.33094°E, Aug 2014, K. Koutroumpa 318 (B [ B 10 1164294], Z [Z-000278449]), K. Koutroumpa 319 (Z [Z-000278458]); presso la foce del fiume Anapodaris, depressione limoso-argillosa, 22 Aug 1996, Brullo & Guarino s.n. (CAT); Tertsa, on calcareous maritime rocks, 34.99146°N, 25.52927°E, Aug 2014, K. Koutroumpa 312 (Z [Z-000278450]), K. Koutroumpa 313 (B [ B 10 1163946], Z [Z-000223471]), K. Koutroumpa 314 (Z [Z-000278451]); cultivated from wild source, Tertsa (Ierapetra), rupi calcarenitiche, Brullo & Giusso s.n., M. Erben Li-1708 (B [ B 10 1009652], FR [ FR-0128958]). — PREFECTURE OF LASITHI: Strand bei Myrtos westlich von Ierapetra, Südküste, 9 Apr 1971, A. Hansen 307 (C); Nea Myrtos (after Ierapetra), 10 m from sign of “Nea Myrtos” (end), 7 May 1986, Tiniakou 1350 (UPA); Prov. Ierapetra, Neos Myrtos, 10 m, S-facing, sandy slope, 22 Apr 1994, P. Wind 94-291 (C); from Ierapetra toward Myrtos, c. 1 km after village Nea Anatoli, Nea Myrtos, at edges of road with Ebenus cretica, 31 May 1988, Artelari & Chondropoulos 1012 (UPA); cultivated from wild source, 1 km W of Nea Anatoli, R. Artelari 1019 & V. Chondropoulos, M. Erben Li-1202 (B [ B 10 0710892]); between Nea Anatoli and Ammoudares, in marls, hardly accessible spot, 35.00585°N, 25.65085°E, Aug 2014, K. Koutroumpa 287 (B [ B 10 1163942], Z [Z-000223477]), K. Koutroumpa 290 (B [ B 10 1163933], Z [Z-000278462]); Nea Anatoli, steep marls, Artelari & Chondropoulos 1019 (B [ B 10 0710893], UPA); 1 km E of Ierapetra, together with Tamarix and several Poaceae, sandy beach, 31 May 1988, Artelari & Chondropoulos 1013 (UPA); 3.2 km E of Ierapetra, “Nicolas Apartments”, on way down from stairs built on rocks, growing along and in both sides of stairs, hardly accessible spot, vertical rocks, 35.00018°N, 25.77360°E, Aug 2014, K. Koutroumpa 281 (Z [Z-000278463]); 3.4 km E of Ierapetra, conglomerate rocks, sandy beach (E side), Artelari & Chrondropoulos 1055 (UPA); c. 1 km W of village Ferma (Ierapetra district), maritime calcareous rocks, plants under Pistacia, 1 Jun 1988, Artelari & Chondropoulos 1017 (UPA); c. 1 km W of village Ferma (Ierapetra district), coffee-bar Lilo, 31 May 1988, Artelari & Chondropoulos 1009 (UPA); c. 1 km W of Ferma, on rocky peninsula close to ancient fish ponds (coffee-bar Lilo), rock crevices, 6 May 1986, Tiniakou 1349 (UPA); Ferma beach, 8 Sep 2013, K. Koutroumpa & al. 2013.09.08-11 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-12 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-13 (herb. University of Crete); Ferma, next to little port close to ancient fish pools, on cement floor, 35.01467°N, 25.84797°E, Aug 2014, K. Koutroumpa 264 (Z [Z-000278464]), K. Koutroumpa 265 (B [ B 10 1133542], Z [Z-000278466]), K. Koutroumpa 269 (Z [Z-000278465]); Distr. Hierapetrae, in saxosis litoreis insulae Photia, 22 May 1942, H. Stubbe 13267 (W [accession no.  W1960-0012700]); Makrygialos (toward Ierapetra), S coast, bushes on beach with conglomerate rocks, at edge of cliff, 6 May 1986, Tiniakou 1348a (UPA); Makrygialos, conglomerate rocks, in inaccessible spot, under Roman settlement, Artelari & Chondropoulos 1018 (UPA); Makrygialos, conglomerate, inaccessible rocks below Roman settlement, 35.03499°N, 25.97258°E, Aug 2014, K. Koutroumpa 252 (Z [Z-000278470]), K. Koutroumpa 256 (B [ B 10 1164323], Z [Z-000278448]), K. Koutroumpa 257 (B [ B 10 1163952], Z [Z-000278468]); Kalo Nero, 27 Mar 1998, N. Böhling 7283 (UPA, B [ B 10 0203903]); Kalo Nero, 10 Jun 2000, Brullo & Giusso s.n. (B [ B 10 1189126, B 10 1189131]); Kalo Nero, 24 Aug 1996, Brullo & Guarino s.n. (B [ B 10 1189129]); Kalo Nero, beach Staousas, on rocks at NE side of beach, 35.01763°N, 26.03379°E, Aug 2014, K. Koutroumpa 235 (Z [Z-000278455]), K. Koutroumpa 236 (Z [Z-000278456]), K. Koutroumpa 237 (B [ B 10 1164105], Z [Z-000278457]); Eparchia Sitias, Kalo Nero, 35°00′55″N, 26°01′32″E, 5 m ü.N.N, Limonium hierapetrae-L. pigadiense [sic]-Lygeum-Flur auf Konglomerat-Kalksandstein, mit Schoenus nig[ricans]. (rar), WSW-exp., mittel alkal., stark lehmiger Sand, 5 May 1997, N. Böhling 5458b (B [ B 10 0199664]); cultivated from wild source, Kalo Nero, Brullo & Giusso s.n., M. Erben Li-1705 (B [ B 10 1009651]); Moni Kapsa, 8 Sep 2013, K. Koutroumpa & al. 2013.09.08-14 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-15 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-16 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-17 (herb. University of Crete); Moni Kapsa in front of Perivolakia, 8 Sep 2013, K. Koutroumpa & al. 2013.09.08-18 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-19 (herb. University of Crete), K. Koutroumpa & al. 2013.09.08-20 (herb. University of Crete); Moni Kapsa, 10 Jun 2000, Brullo & Giusso s.n. (B [ B 10 1189130]); Moni Kapsa, next to monastery in gorge which ends to sea, 35.01973°N, 26.05157°E, Aug 2014, K. Koutroumpa 227 (Z [Z-000278467]), K. Koutroumpa 228 (Z [Z-000278469]); Eparchia Sitias, M[oni]. Kapsa, 35°01′01″N, 26°03′01″E, 5 m ü.N.N., Konglomeratfelsküste, 16 Oct 1997, N. Böhling 6734 (B [ B 10 0192510]); cultivated from wild source, Moni Kapsa, S. Brullo & Giusso s.n., M. Erben Li-1719 (B [ B 10 1132915]); cultivated from wild source, Südost-Küste, Kloster Kapsa, Kattari s.n., M. Erben Li-1727 (B [ B 10 1009650]).

Limonium cornarianum Kypr. & R. Artelari in Phyton (Horn) 38: 144. 1998. – Holotype: Greece, Crete, Nomos Lasithiou, Ep[archia]. Sitias, Moni Kapsa, 35°01′N, 26°03′E, on calcareous maritime rocks, 20 Sep 1994, Z. Kypriotakis 8534 (UPA!). – Fig. 2, 4B.

Description — Plants perennial, 17–63 cm tall, glabrous, with several erect stems. Caudices 2–12 cm long, branched with age and then plant with numerous leaf rosettes. Rosette leaves fleshy, pale green, flat or more usually irregularly folded, rugose, 25–90 × 6–20 mm, oblanceolate-spathulate, abruptly tapering into petiole, with petiole longer than lamina, margin cartilaginous, sometimes undulate, c. 0.1 mm wide, apex acute, mucronate to cuspidate, mucro 0.3–1.2 mm long; 1 central nerve with 2 lateral nerves in large leaves. Stems several, usually slender, erect, rugose, branched usually above lower 1/3. Inflorescence rhombic to trullate, rarely obtrullate in outline; sterile branches absent or 1–3 at lower part of stem, straight; fertile branches straight or slightly arched, directed obliquely upward. Spikes 8–37 mm long, more or less straight. Spikelets 3.8–4.5 mm long, very densely arranged, 3–11 per cm, mainly distichous, composed of 2–7 flowers. Outer bract 1.1–1.7 × 1–1.4 mm, glabrous, ovate-triangular, margin membranous, apex acute; central part, herbaceous forming a short point. Middle bract 1.2–1.9 × 0.9–1.3 mm, glabrous, membranous, oblong-elliptic, apex rounded. Inner bract 2.8–3.8 × 2.7–3.2 mm, glabrous, obovate, margin membranous, 0.5–0.8 mm wide, apex obtuse to rounded; central part herbaceous, forming a short point 0.4–0.5 mm long. Calyx 3.2–3.9 mm long; calyx tube usually unilaterally long hairy; calyx lobes 0.3–0.7 mm long, ovate; tube ribs ending at or above base of lobes. Corolla lilac.

Note — Plant leaf size (length and width) measurements were taken from several individuals in the field, at Pacheia Ammos (Crete), though not all these individuals were collected as herbarium specimens.

Chromosome number and ploidy — Ploidy level: 3x (inferred from preliminary flow cytometric results of genome size).

Breeding system — Apomixis was inferred from the monomorphic population in the locus classicus with self-incompatible floral combination B (Artelari & Georgiou 1999) and triploidy.

Distribution — Endemic, restricted to the Moni Kapsa gorge (SE Crete) and the E edge of Pacheia Ammos cove (NE Crete) (Fig. 1).

Habitat — Steep, calcareous rocks and dried salt marsh near the coast.

Additional specimens seen — GREECE, CRETE, PREFECTURE OF LASITHI: Pacheia Ammos, at E side of coast, on salt marshes between little church Agia Foteini and a small port, marshes with Tamarix, 35.11060°N, 25.81533°E, Aug 2014, K. Koutroumpa 99 (Z [Z-000278459]), K. Koutroumpa 100 (Z [Z-000278452]); Sitia, 3 km SSW of Kato Perivolakia, lower part of Moni Kapsas gorge, alt. 15 m, crevice of E-facing calcareous cliff, 26 Jun 1994, N. Turland 777 (BM, UPA); Moni Kapsa gorge on lower part toward sea, alt. 15–20 m, 35.02189°N, 26.05044°E, Aug 2014, K. Koutroumpa 731 (B [ B 10 1164150], Z [Z-000278453]), K. Koutroumpa 751 (B [ B 10 1164136], Z [Z-000278454]); Eparchia Sitias, M[oni]. Kapsa, Schlucht, 35°01′03″N, 26°03′05″E, 50–100 m ü.N.N., Felswände (Kalk), 16 Oct 1997, N. Böhling 6738 (B [ B 10 0192506]); Exp. E der Kapsa-Schlucht, Ostkreta, 20 m, Kalkfelsen, 15 Oct 1998, R. Ulrich (B [ B 10 0189672]).