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A morphological and karyological investigation of the Anthemis sect. Hiorthia representatives of Kriti (Greece) revealed that three different species are found on the island, all endemic, and each characterised by a different ploidy level based on the haploid series of x = 9. Anthemis abrotanifolia, the species with the widest distribution, is tetraploid with 2n = 4x = 36. A. samariensis, a local endemic of the Lefka Ori, was found being decaploid, with 2n = 10x = 90, the highest number ever recorded in Anthemis. The recently discovered population on Mt. Kedros (south-central Kriti) is morphologically distinct from all the Anthemis entities growing on Kriti; it also differs from the variable and widespread A. cretica group. It is here described as a new species, A. pasiphaes Goula & Constantinidis. It is a hexaploid, with 2n = 6x = 54. All chromosome numbers are reported for the first time. Polyploidy might have acted as a reproductive barrier among these perennial species, complementing isolation by spatial distance and evolutionary divergence. Further, it might have contributed adaptation advantages to these three predominately mountain species.

Anthemis rigida subsp. runemarkii Biel & Kit Tan (2020: 190) (Fig. 5: e,f) Type:― GREECE. Kiklades: island of Milos, NW of Ag. Mamas, rocky phrygana slopes above coast, 10 m, 36°41’N, 24°25’E, 25 February 2020, Biel 2020.48 (holotype C; see Biel & Tan, 2020). Small annual with prostrate to ascending, simple or sparingly branched stems, 1−20 cm long. Stems and leaves tinged with purple, densely tomentose with medifixed hairs when young, laxly tomentose to glabrescent later. Lower leaves long-petiolate, 1-pinnatisect with segments pinnatifid to pinnatisect, glandular-punctate; ultimate lobes at least 2 times longer than wide, oblong-obovate to oblong-ovate, acute to shortly mucronate; upper leaves similar but smaller. Peduncles thickened, becoming rigid and usually recurved in fruit. Capitula solitary, radiate. Involucre 5−8 × 5−7 mm, almost cylindrical, bracts usually in 2−3 rows, imbricate, unequal in length, tomentose when young, thick, rigid, and glabrescent in fruit, often tinged with purple. Outer involucral bracts narrowly lanceolate to ovate-lanceolate, 2.5−4 × ca. 1 mm, usually not more than 1/2−2/3 as long as inner bracts, with scarious margins up to 0.4 mm. Inner bracts obovate, 5−6 × ca. 1.5 mm, with broad scarious margins up to 0.5 mm. Ligulate florets with ligules oblong-ovate, toothed at apex, 5−8 × 1.7−2.5 mm, dark to pale pink, white tinged pink or white, glandular punctate, sterile; disc florets ca. 3 mm, yellow, glandular punctate. Anthers, style, and pollen yellow. Receptacular scales hyaline, obovate to oblanceolate, mucronate and becoming lacerate at apex. Achenes obconical, faintly ribbed; outer achenes ca. 2 × 1 mm, with a corona 0.5−0.9 mm long; inner achenes longer and thinner, 2−2.8 × 0.6−0.8 mm, with a corona 0.4−0.8 mm long. Distribution and habitat:―This subspecies is endemic to Milos island group and has been located so far on the following islands and islets: Milos, Kimolos, Prasonisi, Poliegos, at elevations from sea level to ca. 87 meters. It has been found in a number of different habitats such as sandy shores, rocky and gravelly places next to the sea, phrygana formations, maquis and periodically flooded areas, all on acid volcanic substrates. Notes:―Although the pink-coloured ligules is a remarkable trait observed in this subspecies, it is not characteristic of all the known populations. Ligules in various shades of pink, from deep- to pale-pink, or white tinged pink, or totally white have been recorded, even among individuals from the same locality. The morphological features that separate Anthemis rigida subsp. runemarkii from the rest of the ligulate subspecies are mainly the narrow leaf lobes, the longer and wider involucre and the broad scarious margin of the outer involucral bracts (Fig. 5, Table 5). The “adpressedpubescent” achenes that Biel & Kit Tan (2020) mention in their description raises serious questions about achene morphology for the entire genus: the only report of hairy achenes among the Anthemis species of Europe (Fernandes 1976), Turkey (Grierson & Yavin 1975), N Africa (Oberprieler 1998), Middle East (Eig 1938; Iranshahr 1986, Ghafoor & Ali 2002) and Saudi Arabia (Ghafoor 2010) concerns “a single poor specimen with somewhat hairy young achenes” that Eig (1938) mentions from Ramadi, Iraq. A thorough examination of achenes in all the specimens collected on Milos do not indicate the presence of any pubescence. As previously mentioned, Anthemis rigida subsp. runemarkii co-exist with A. rigida subsp. rigida on the island of Milos. In the areas where the two taxa meet, subpopulations with intermediate features have been recorded. Similar cases are reported by Greuter & Rechinger (1967) for populations on Kithira and Antikithira islands., Published as part of Goula, Katerina & Constantinidis, Theophanis, 2021, Taxonomic diversity and karyology of Anthemis rigida (Anthemideae, Asteraceae) in the Aegean, Greece, pp. 129-143 in Phytotaxa 484 (1) on pages 137-140, DOI: 10.11646/phytotaxa.484.1.7, http://zenodo.org/record/5421338, {"references":["Biel, B. & Tan, K. (2020) Reports 12 - 15. In: Vladimirov, V., Aybeke, M. & Tan, K. (Eds.) New floristic records in the Balkans: 42. Phytologia Balcanica 26: 187 - 216.","Strid, A. (2016 b) Atlas of the Aegean flora, Part 2: Maps. Englera 33: 1 - 878. [Berlin: Botanic Garden and Botanical Museum Berlin, Freie Universitat Berlin]","Fernandes, R. (1976) Anthemis L. In: Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. (Eds.) Flora Europaea 4. Cambridge University Press, Cambridge, pp. 145 - 159.","Grierson, A. J. C. & Yavin, Z. (1975) Anthemis L. In: Davis, P. H. (Ed.) Flora of Turkey and the East Aegean islands. Edinburgh University Press, Edinburgh, pp. 174 - 221.","Oberprieler, C. (1998) The systematics of Anthemis L. (Compositae, Anthemideae) in W and C North Africa. Bocconea 9: 5 - 328.","Eig, A. (1938) Taxonomic studies on the oriental species of the genus Anthemis. Palestine Journal of Botany 1: 161 - 224.","Iranshahr, M. (1986) Anthemis. In: Rechinger, K. H. (Ed.) Flora Iranica 158, Compositae: VI Anthemideae. Akademische Druck und Verlagsanstalt, Graz, pp. 5 - 44.","Ghafoor, A. & Ali, S. J. (2002) The genus Anthemis L. (Compositae) in Iraq: a synopsis. Compositae Newsletter 38: 1 - 41.","Ghafoor, A. (2010) The genus Anthemis L. (Compositae-Anthemideae) in Arabian Peninsula: a taxonomic study. Pakistan Journal of Botany 42: 79 - 98.","Greuter, W. & Rechinger, K. H. (1967) Chloris Kythereia. Boissiera 13: 22 - 196."]}

Goula, Katerina, Constantinidis, Theophanis (2021): Taxonomic diversity and karyology of Anthemis rigida (Anthemideae, Asteraceae) in the Aegean, Greece. Phytotaxa 484 (1): 129-143, DOI: 10.11646/phytotaxa.484.1.7, URL: http://dx.doi.org/10.11646/phytotaxa.484.1.7

Anthemis rigida subsp. ammanthiformis (Greuter & Rech.f.) Greuter (1968: 263) Type:― GREECE. Insula Antikythera: Potamos, in parte boreali insulae, in saxosis calc. litoreis, 8 May 1964, Rechinger 24367a (holotype, W!). Annual with prostrate to ascending, simple or sparingly branched stems, 1.5−25 cm long. Stems and leaves greyishgreen, densely tomentose with medifixed hairs. Lower leaves long-petiolate, 2-pinnatisect, glandular-punctate; ultimate lobes obovate to obovate-lanceolate, subacute to shortly mucronate; upper leaves gradually smaller and less divided to simple. Peduncles thickened, becoming rigid in fruit. Capitula solitary, radiate. Involucre (3−)4−8 × 3−4.5(−5.5) mm, obconical to subhemispherical, bracts usually in 2−3 rows, imbricate, unequal in length, densely tomentose, thick and rigid in fruit. Outer involucral bracts lanceolate to narrowly oblong-lanceolate, 2−3 × ca. 1 mm, usually not more than 1/2−2/3 as long as inner bracts, with scarious margins up to 0.2 mm. Inner bracts lanceolate to obovate, 3−4.5(−5.5) × ca. 1.5 mm, with scarious margins up to 0.4 mm. Ligulate florets with ligules obovate, as long as or shorter than involucre, white, glandular punctate, sterile; disc florets yellow, glandular punctate. Anthers, style, and pollen yellow. Receptacular scales hyaline, obovate to oblanceolate, becoming lacerate. Achenes obconical to obpyramidal, faintly ribbed; outer achenes 1.8−2 mm, with a corona 0.5−0.6 mm long; inner achenes somewhat longer and thinner, 2−2.2 mm, with a corona 0.3−0.5 mm long. Distribution and habitat:―This subspecies is endemic to Antikithira and the northern part of Gramvousa promontory, west Kriti, where it grows mostly on coastal habitats., Published as part of Goula, Katerina & Constantinidis, Theophanis, 2021, Taxonomic diversity and karyology of Anthemis rigida (Anthemideae, Asteraceae) in the Aegean, Greece, pp. 129-143 in Phytotaxa 484 (1) on page 140, DOI: 10.11646/phytotaxa.484.1.7, http://zenodo.org/record/5421338, {"references":["Greuter, W. (1968) Notulae nomenclaturales et bibliographicae 5 - 6. Candollea 23: 257 - 265."]}

Anthemis rigida subsp. rigida Boiss. ex Heldr. (Fig. 5: a,b) Type (lectotype designated by Greuter 1968):― CYPRUS. Sibthorp & Lindley, Flora Graeca t. 853. Annual with prostrate to ascending, simple or sparingly branched stems. Stems and leaves more or less tomentose with medifixed hairs. Lower leaves 1−5 cm, pinnatisect with pinnatifid to pinnatisect segments, ovate-oblong in outline; ultimate lobes obovate to oblanceolate, 0.5–1 mm wide, subobtuse, acute or mucronate; upper leaves similar but smaller. Peduncles thickened, becoming rigid and usually recurved in fruit. Capitula solitary, discoid. Involucre 3.5−6(−8) × 2.5−4.5 mm, ocbonical to subcylindrical, bracts imbricate, subequal in length, more or less tomentose when young, thick and rigid in fruit. Outer involucral bracts oblong-lanceolate to ovate-lanceolate, with scarious margins up to 0.2 mm. Inner involucral bracts narrowly ovate- to obovate-lanceolate, with scarious margin 0.2−0.5 mm. Disc florets 1.8−3 mm, yellow, the peripheral sometimes tinged with red. Receptacular scales hyaline, oblanceolate, acute. Achenes obconical, faintly ribbed; outer achenes 1.5−2.1 × 0.7−1 mm, with a corona 0.3−0.9 mm; inner achenes 1.6−2.3 × 0.5−0.9(−1) mm, with a corona (0.1−) 0.2−0.5 mm. Distribution and habitat:―See Strid (2016a,b, map 413)., Published as part of Goula, Katerina & Constantinidis, Theophanis, 2021, Taxonomic diversity and karyology of Anthemis rigida (Anthemideae, Asteraceae) in the Aegean, Greece, pp. 129-143 in Phytotaxa 484 (1) on pages 140-141, DOI: 10.11646/phytotaxa.484.1.7, http://zenodo.org/record/5421338, {"references":["Greuter, W. (1968) Notulae nomenclaturales et bibliographicae 5 - 6. Candollea 23: 257 - 265.","Strid, A. (2016 a) Atlas of the Aegean flora, Part 1: Texts and Plates. Englera 33: 1 - 700. [Berlin: Botanic Garden and Botanical Museum Berlin, Freie Universitat Berlin]"]}

Anthemis rigida subsp. liguliflora (Halácsy) Greuter (1968: 263) (Fig. 5: c,d) Type:― GREECE. Cerigo [Kithira]: Hafen von Hagios-Nicolaos, 6 May 1902, Sterneck 279 (WU 035428!). Annual with prostrate to ascending, simple or sparingly branched stems, 1.5−15 cm long. Stems and leaves greyishgreen, densely tomentose with medifixed hairs when young, less tomentose in fruit. Lower leaves long-petiolate, 1-pinnatisect, with narrowly pinnatifid, glandular-punctate segments; ultimate lobes obovate, usually twice as long as wide, subacute to shortly mucronate; upper leaves similar but smaller. Peduncles thickened, becoming rigid and usually recurved in fruit. Capitula solitary, radiate. Involucre 4−6(−7) × 3−4 mm, obconical to subcylindrical, bracts usually in 2−3 rows, imbricate, subequal in length, densely tomentose when young, thick, rigid, and often tinged purple in fruit. Outer involucral bracts narrowly lanceolate, 2.5−3.6 × ca. 1 mm, with scarious margins up to 0.2 mm. Inner bracts obovate, 2.7−3.8 × 1−1.2(−1.5) mm, with scarious margins (0.2−) 0.3−0.5 mm. Ligulate florets with ligules oblong-ovate, toothed at apex, 3.2−5.2 × 1−1.6 mm, white, glandular punctate, sterile; disc florets 2.5−3 mm, white, glandular punctate. Anthers, style, and pollen yellow. Receptacular scales hyaline, obovate to oblanceolate, mucronate and becoming lacerate at apex. Achenes obconical, faintly ribbed; outer achenes ca. 2 × 1 mm, with an auricle 0.5−0.9 mm long; inner achenes longer and thinner, ca. 2(−2.4) × 0.6−0.9 mm, with a corona 0.2−0.6 mm long. Distribution and habitat:―This subspecies is endemic to the island of Kithira, where it grows on coastal habitats and stony places in phrygana communities., Published as part of Goula, Katerina & Constantinidis, Theophanis, 2021, Taxonomic diversity and karyology of Anthemis rigida (Anthemideae, Asteraceae) in the Aegean, Greece, pp. 129-143 in Phytotaxa 484 (1) on page 140, DOI: 10.11646/phytotaxa.484.1.7, http://zenodo.org/record/5421338, {"references":["Greuter, W. (1968) Notulae nomenclaturales et bibliographicae 5 - 6. Candollea 23: 257 - 265."]}