Teucrium salaminium Hadjik. & Hand, spec. nova (Fig. 1–6)

Holotype: Cyprus. Division 7 (sensu Meikle, 1977): Ypsarovounos forest, c. 2.5 km southwest of Mandres, steep to even gypsum slopes dissected by streams, phrygana vegetation, with scattered Pinus brutia and Cupressus sempervirens, alt. c. 265 m, 27.VI.2007, Hadjikyriakou 7071 (holo-: B; iso-: B, CYP, G, STU, herb. Hadjikyriakou).

Differt a Teucrio micropodioide caulibus (sub)erectis (non procurrentibus), foliis olivaceis vel viridis (non glaucoviridis), inflorescentiis obpyramidalibus vel obconicis (non globosis plus minusve), nuculis 1.1–1.3 mm (non 1.8 mm) longis.

Shrub, 15–40 cm high, with erect or suberect stems. Indumentum consists out of the following types of trichomes: (1) short and long clavate glandular trichomes, mostly with 2–3 stalk cells, (2) subsessile one-celled glandular trichomes, (3) non-glandular, unbranched, thick-walled trichomes, with 2 to several stalk cells. Old wood greyish-brown, fissured; current year's shoots straight, terete, erect to suberect, simple or variously branched, lengthening up to 20(-30) cm; lowermost part of the shoots remaining alive throughout the year becomes leafless; the upper part bearing the flower heads dries out; indumentum consists of dense, white, crisped or adpressed eglandular, unbranched hairs, intermixed with spreading, long or short glandular ones. Leaves subsessile or shortly petiolate, narrowly to broadly oblanceolate or elliptical, 12–22 × 3–6 mm when fully open, olive-green to green on both sides, becoming yellowish-green with age; margins slightly revolute, the lower ½–¾ entire or somewhat sinuate-undulate, the upper ¼-½ with an apical tooth and (0-)2-4(-7) crenations on either side; leaves appearing from late spring onwards become progressively smaller, narrowly oblong to linear, tufted on top of the sterile shoots, 4–11(-19) × 1.5-2(-2.5) mm when fully opened, subsessile or sessile, olive-green to green on both sides; margins strongly revolute, entire except for an apical tooth, or the upper ¼-½ with 2–4 crenations on each side and an apical tooth; both surfaces of the leaves with sessile glands, at first with lax to dense, white, unbranched, spreading or subappressed, short or long glandular and eglandular hairs, becoming appressed and crisped with age. Flower heads obpyramidal or obconical, flattopped or slightly rounded, by the end of the flowering period many slightly elongated and ovoid, (6-)8-16 × (6-)8-16 mm, arising on top of long shoots or on lateral, long or short branches, altogether giving the appearance of a lax panicle; sometimes the heads on top of the long shoots are subtended by 2–4 smaller, subsessile ones. Bracts foliaceous, greenish, for most of the flowering period, equal or exceeding the flowers, all more or less reaching to the same height, 4.5–7.5(-2.4) × 0.5–1.3 mm, linear, narrowly oblong or oblanceolate, with revolute margins, entire, except for an apical tooth. Bracteoles absent. Flowers 4–15(-20) per head, sessile or shortly petiolate, sometimes 1–2 of the central buds never open. Calyx tubular, 4–6.1 mm; tube curved, 3.2–4.3 × 1.8–2 mm, 5-nerved, reticulately veined, externally with sessile glands and lax or dense white, spreading or adpressed, crisped, short or long glandular and eglandular hairs and internally with spreading or shortly subadpressed, eglandular and glandular hairs; teeth somewhat unequal; the abaxial deltoid, somewhat acuminate, 1–1.8 × 0.5–0.7 mm, the adaxial about 0.8–1.3 × 0.8–1 mm. Corolla tube 2.5–5 × 0.8–1.3 mm, slightly curved, glabrous to subglabrous towards the base externally, glabrous towards the base internally, pubescent at the region of the throat; limb white flashed with pink-violet on the lower part or pinkish-violet all over, 4–6 mm long; basal lobes oblong, 1.2–2 × 0.5–0.8 mm, lateral lobes spreading, oblong, 1.1–1.3(-1.4) × 0.3–0.6 mm, median lobe nearly circular, 1.6–2.5 × 1.2–2.1 mm, cucullate; external surface thinly pubescent and with sessile glands; internal surface gland-dotted, hirsute at the region of the basal and lateral lobes, glabrous at the lateral lobes. Stamens inserted at or slightly below the mouth of corolla tube, exserted; filaments pinkish-violet above, thinly pilose, the adaxial 1.3–3.7 mm long, the abaxial 2.5–5.3 mm long; anthers pinkish-violet, 0.5–1 mm long. Ovary glabrous, about 0.5–0.8 mm long; style thinly pubescent, 4.5–6 mm long; stigmatic lobes subulate, unequal, 0.5–0.9 mm long. Nutlets somewhat cochleariform to obovate, 1.1–1.3 × 0.7–0.9 mm, rounded at apex, wedge-shaped at the base, attachment area flat, suborbicular, 0.6–0.7 mm in diameter, adjacent area convex; dark brown to blackish, granulose, slightly reticulately veined and foveolate.

Eponymy. — The specific epithet is the Latinized form of salamínios, which refers to somebody or something that comes from Salamis, an ancient city of Cyprus, situated about 17 km SE of the locality of collection. Salamis was founded by the Trojan hero Teucer, after whom the genus Teucrium is most likely named.

Flowering period. — Teucrium salaminium and T. micropodioides grow side by side in this area, but the former flowers from late June to end of September and the latter from May to late June; observations show that there is a period of 5–10 days overlapping. There were no indications of interbreeding or intermediacy.

Distribution and habitat. — Teucrium salaminium is confined chiefly within Ypsarovounos forest (ýpsaros = gypsum, vounón = mountain) and is found in the vicinity of the villages Mandres, Agios Iakovos and Platani (main occurrence at 35°19′N 33°47′E). Smaller groups have been located on gypsum hills which are situated at the northeast boundaries of the forest. It is generally found at an altitude between 200–380 m. The area is dissected by small streams, and it is characterized by steep to even slopes, cliffs and scattered, small flattish areas. The new species is restricted to Kalavasos geological formation (Upper Miocene) which is composed of gypsum alternating with chalky marls and marly chalks, in an area of about 5 km². This formation is mainly surrounded by Terrace Deposits (Pleistocene), but in a few places it adjoins outcrops of Alluvium-Colluvium, Pachna and Kythrea Formation (Middle Miocene) characterized by, e.g., several types of sands, silts, gravels, marls and conglomerates.

The precipitation rate in three nearby stations (Akanthou, Lefkoniko, Trikomo; 1951–1980; Meteorological Service, 1985) ranges from 316–513 mm per year. Because of the topo graphical situation, a rate of about 400 mm seems probable in the Ypsarovounos area.

Teucrium salaminium grows chiefly in habitat type 1520 Gypsum steppes - priority habitat type (sensu Directive 92/43 EEC), which covers extensive areas, or it is found in forest openings. It also occurs in other habitat types such as 5220 Matoral with Ziziphus (priority habitat type), 6220 Pseudosteppe with grasses and annuals (priority habitat type), 9290 forests of Cupressus sempervirens, and 9540 Mediterranean pine forests with endemic Mesogean pines, Pinus brutia. Characteristic taxa of these habitats are mostly gypsovags, e.g., Asparagus horridus L., Asphodelus ramosus L., Calycotome villosa (Poir.) Link, Cistus creticus L., Cistus salviifolius L., Cupressus sempervirens L., Cynara cornigera Lindl., Drimia aphylla (Forssk.) J. C. Manning & Goldblatt, Ferula communis L., Fumana arabica (L.) Spach, Fumana thymifolia (L.) Verl., Helichrysum stoechas subsp. barrelieri (Ten.) Nyman, Hyparrhenia hirta (L.) Stapf, Juniperus phoenicea L., Pistacia lentiscus L., Pistacia terebinthus L., Rhamnus lycioides subsp. graeca (Boiss. & Reut.) Tutin, Sarcopoterium spinosum (L.) Spach, Teucrium micropodioides, Thimbra capitata (L.) Cav., and Ziziphus lotus (L.) Lam. The complete set of Cypriot gypsophiles is also found there: Campanula fastigiata Schult., Gypsophila linearifolia (Fisch. & C. A. Mey.) Boiss., and Herniaria hemistemon J. Gay.

Three of the Cypriot taxa of sect. Polium in Cyprus (T. micropodioides, T. cyprium and T. karpasiticum) grow in a variety of geological substrates and T. kyreniae grows in crevices of limestone cliffs, whereas repeated investigations show that T. salaminium is restricted exclusively to gypsum substrate. No plants have been found on the surrounding formations. Further investigations were carried out on extensive gypsum outcrops in the vicinity of Gastria village 15 km east of the type locality, in an area of 3 km², and on a smaller gypsum outcrop in the vicinity of Gypsou village, 6 km south of the type locality - in both areas without success.

Conservation. — Teucrium salaminium seems to be a rare endemic of Cyprus and according to the IUCN Red List Categories and Criteria (Version 3.1) is categorized as VU (Vulnerable): D2. This means that the area of occupancy is less than 20 km² and it occurs in only one location. Potential threats are forest fires, grazing, quarriyng of gypsum, road construction, agriculture and forestry operations. The largest part of its population is found within Ypsarovounos Main State Forest. Additionally, this forest is included in a proposed Natura 2000 site, and the habitat type 1520 (Gypsum steppes) is a priority habitat.

Systematic relationships. — Obviously, the new taxon is an endemic of the island. Probably the closest relatives of T. salaminium in Cyprus are T. micropodioides and T. kyreniae. The former grows together with T. salaminium, whereas the latter is a chasmophyte growing on limestone cliffs 5–7 km northeast of the type locality.

Teucrium salaminium and the mentioned taxa are distinguishable by the characters summarized in (Table 1) (see also Meikle, 1985 and Hadjikyriakou & Hand, 2008 for further details). From a morphological point of view, T. micropodioides shares many characters with T. salaminium. Nonetheless, the two taxa are easily separable by, e.g., different growth form, colouration of leaves and flowers and structure of inflorescences.

The quality of indumentum is very similar in all five Cypriot taxa of sect. Polium. They all belong to subsect. Rotundifolia, which is characterized by, e.g., the lack of branched trichomes, whereas subsect. Polium has branched hairs (see also Hadjikyriakou & Hand, 2008). The former taxon is widespread in the Mediterranean and the Near East; its members occur primarily in microclimatically hot habitats, on cliffs, rocks, especially on serpentine, as well as in maritime environments, avoided by other taxa of the genus (see Navarro, 1995; Navarro & el Oualidi, 2000a). Teucrium salaminium seems to be the only endemic gypsophile of Cyprus.

A detailed phylogenetic and ecological study of the Cypriot members of sect. Polium is planned for the near future.

Specimens seen (division sensu Meikle, 1977; herb. GH = herb. Hadjikyriakou). — Cyprus. Division 7: Ypsarovounos forest, 2.5 km southwest of Mandres village, Ammochostos district, opening in pine forest on gypsum, alt. c. 250 m, 19.XII. 2008, Hadjikyriakou 7041 (B, herb. GH); ibid., 25.III.2009, Hadjikyriakou 7044 (B, herb. GH); ibid., 5.VII.2009, Hadjikyriakou 7073 & 7074 (B, herb. GH); ibid, 25.VII.2009, Hadjikyriakou 7075 & 7076 (B, herb. GH); ibid., 29.VIII.2009, Hadjikyriakou 7100 (B, herb. GH); ibid., 11.IX.2009, Hadjikyriakou 7124 (B, herb. GH); Kakotris hill Mandres Ammochostou, phrygana on gypsum, alt. c. 200 m, 1.VIII.2009, Hadjikyriakou 7077 (B, herb. GH); 2 km E of Platani Village in Ypsarovounos forest, phrygana on gypsum, alt. c. 200 m, 16.VIII.2009, Hadjikyriakou 7079 & 7080 (B, herb. GH); Agios Iakovos, c. 1.5 km W of village, lower slopes of gypsum hill, on N side close to the ascending track, open gypsum phrygana, alt. c. 350 m, 11.V.2009, Hand 5475, Christo doulou & Hadjikyriakou (B).