Research Article
Research Article
Thliphthisa sapphus (Rubiaceae, Rubieae), a new species from Lefkada (Ionian Islands, Greece) and its ecological position
expand article infoWalter Gutermann, Tae-Soo Jang§, Arndt Kästner|, David Prehsler, Dieter Reich, Andreas Berger, Ruth Flatscher, Christian Gilli, Markus Hofbauer, Margarita Lachmayer, Ruth Sander, Michaela Sonnleitner, Ladislav Mucina#¤
‡ University of Vienna, Vienna, Austria
§ Chungnam National University, Daejeon, Republic of Korea
| Unaffiliated, Halle (Saale), Germany
¶ Department of Botany, Natural History Museum Vienna, Vienna, Austria
# Murdoch University, Perth, Australia
¤ Stellenbosch University, Stellenbosch, South Africa
† Deceased author
Open Access


The new species, Thliphthisa sapphus sp. nov. (Rubiaceae, Rubieae), a narrow endemic of the white cliffs of Lefkátas on the southwest coast of Lefkada (Greece) is described and illustrated and an IUCN assessment is presented. Vegetation relevés were performed at the single known locality, limestone cliffs facing the sea and revealed a new association, the Thliphthisa sapphus-Lomelosietum dallaportae. The chromosome number of Thliphthisa sapphus was determined as 2n = 4x = 44, being the single tetraploid species in the genus to date. The species also differs markedly morphologically from its morphologically closest relatives, two Greek steno-endemic oreophytes, Th. baenitzii and Th. muscosa by the following characters: densely setose mericarps and corolla, tetraploidy and by its distribution. An identification key for the Greek species of Thliphthisa is provided. Th. sapphus constitutes the westernmost outpost of a group of Greek steno-endemics, highlighting the importance of coastal habitats and their protection as refugia for poorly competitive chamaephytes.

Key words

Asperula, coastal cliff refugia, Greece, Ionian Islands, phytosociology, Rubiaceae, species nova, Thliphthisa


The recently erected genus Thliphthisa (Griseb.) P.Caputo & Del Guacchio (Rubiaceae, Rubieae, Galiinae) is centred in the eastern Mediterranean and includes 22 species, mostly narrow endemics. The systematics and taxonomy of subtribe Galiinae with up to 900 species is complex and the genus boundaries are still disputed, but the group is generally recognised by herbaceous habit with pseudo-whorled leaves and dry mericarps (Ehrendorfer et al. 2018). According to traditional understanding, the group is divided into several small (e.g. Sherardia L.) to medium-sized genera (e.g. Asperula L.) and the large, para- and polyphyletic genus Galium L. with about 650 species. As a result of extensive morphological and phylogenetic studies, Ehrendorfer and co-workers recently proposed that several sections of Asperula and Galium be elevated to generic rank, which would allow the retention of all traditional genera of Galiinae such as Cruciata Mill., Sherardia or Valantia L., a viewpoint favoured over merging the whole Galiinae resulting in a large and heterogeneous Galium (s. latiss.) (Ehrendorfer and Barfuss 2014; Ehrendorfer et al. 2018). Following these suggestions, Galiinae should comprise more than a dozen small genera, but most formal taxonomic decisions following these results are still pending.

Asperula sects. Cynanchicae (DC.) Boiss., Thliphthisa (Griseb.) Ehrend. and Hexaphylla Klokov were consistently resolved as monophyletic groups together with the monophyletic Sherardia, but the relationships of these groups are not always resolved (Sherardia clade of Natali et al. (1995); Clade IV of Soza and Olmstead (2010); Clade VI of Ehrendorfer et al. (2018)). Following Ehrendorfer and co-workers, Del Guacchio and Caputo (2020) recently proposed to raise these sections to generic rank basically upgrading the “distinguishable smaller subclades” of Clade VI (Ehrendorfer et al. 2018) to generic level. Morphologically, Thliphthisa is variable and difficult to separate from other representatives of subtribe Galiinae, one of the few consistent characters being truncate fruits (Schönbeck-Temesy and Ehrendorfer 1985; Del Guacchio and Caputo 2020). In Thliphthisa, Del Guacchio and Caputo (2020) have combined all but one taxon treated in the synopsis (conspectus, key to species and distribution maps) of Schönbeck-Temesy and Ehrendorfer (1985), although without citing this revision. Only the African A. cyrenaica (E.A.Durand & Barratte) Pamp. was included in genus Hexaphylla (Klokov) P.Caputo & Del Guacchio.

In November 1994, during floristic and vegetation studies on the island of Lefkada, Ionian Islands, Greece, a presumably undescribed rubiaceous chamaephyte was found. At the time of discovery, the plants had passed flowering and only remnants of withered corollas and aborted fruits were found. Repeated study of the population in summer 1996 as well as spring 2000 and 2011, observations on plants grown in cultivation and herbarium studies comparing all other Mediterranean species of Asperula s.l. confirmed that the enigmatic taxon is a Thliphthisa species new to science. It is here described as Thliphthisa sapphus, its affinities, cytology and ecology are detailed and an identification key for the Greek species of the genus is provided.

Materials and methods

Due to the steep cliff terrain, direct observations are limited to a small part of the single known population of this new Thliphthisa species. Vegetation relevés using a modified Braun-Blanquet cover-abundance scale (see Barkman et al. (1964) for details) have been made at the habitats of the species in autumn 1994 and in spring 1996. These relevés have been combined into a table which was further compared to known vegetation types of chasmophytic vegetation described so far from the Mediterranean to establish the syntaxonomic community identity.

In 2011, a young plant from a secondary habitat in a road embankment above the cliffs was transferred to the Botanical Garden of the University of Vienna (thereafter HBV) for cultivation and karyological investigations. Chromosome numbers were determined applying the standard Feulgen staining technique as described in Weiss-Schneeweiss et al. (2009). Actively growing root tip meristems were pre-treated with a 0.002M aqueous solution of 8-hydroxyquinoline for 2.5 h at room temperature and 2.5 h at 4 °C, fixed in ethanol:acetic acid (3:1) for at least 3 h at room temperature and stored at –20 °C until use. Fixed root meristems were hydrolysed in 5N hydrochloric acid (HCl) for 30 min at room temperature, washed with tap water and stained with Schiff’s reagent (Merck, Darmstadt, Germany) in darkness for 1 h (Jang et al. 2013). Chromosome spreads were prepared by squashing stained root tip meristems in a drop of 60% acetic acid under the cover-slip and analysed using an AxioImager M2 microscope (Carl Zeiss). All images were acquired with a CCD camera and files processed using AxioVision version 4.8 (Carl Zeiss). Karyotypes were made from these images in Corel Photo-Paint X5 (Corel Corp., Ottawa, Ontario).

For morphological comparison with the new species, specimens from the Herbaria W and WU of the other Thliphthisa taxa were examined with respect to diacritical characters as given in Schönbeck-Temesy and Ehrendorfer (1985). For a comprehensive list of analysed herbarium material from Greece, see Suppl. material 1.

Taxonomy and nomenclature in this work follows the Flora Ionica Working Group (2016 onwards).

Taxonomic treatment and discussion

Thliphthisa sapphus Gutermann, sp. nov.

Figs 1, 2, 3, 4B, C


Greece. Ionian Islands: Nom. Lefkada, Lefkada, Halbinsel Lefkátas, Hangkante über der Westküste WNW ober Aghios Nikólaos Níras (NNE Akr. Dhoukáto, ca. 8 km SSW Atháni), 38°35'43"N, 20°33'06"E, meerseitige Kalkfelsfluren, ca. 230 m elev., 24 May 2000, W. Gutermann et al. 35241 (holotype: WU 0154346; isotypes: ATH, K, M, LD, UPA, herb. Gutermann).


Generis Thliphthisae species nova: suffrutex nanus, laxe aut densius pulvinatus e caudice valde lignoso multicaulis; caulibus scabris, dense et aeque foliatis usque ad apicem; foliis verticillatis quaternis vel senis, leniter crassiusculis, anguste oblanceolatis brevissime apiculatis, modice setosis; floribus solitariis sessilibus; corollis hypocrateriformibus, albis, extus setosis; staminibus in medio corollae inserta, antheris luteis; ovariis ± dense setosis praesertim ad apicem versus, stylo manifeste bifido stigmatibus breviter ellipsoideis. – Chromosomatum numerus 2n = 44, scil. species tetraploidea. Crescit in declivibus apricis saxosis insulae Leucas Graeciae. Floret mense Maio & Junio.

Figure 1. 

Illustration of Thliphthisa sapphus, based on the type material A whole plant B whole plant with vegetative and flowering shoots C vegetative shoot D detail of stem of vegetative shoot E flowering shoot F detail of stem of flowering shoot G flower H dissected flower (drawings by A. Kästner).

Caespitose, suffruticose cushions with lignified rootstock and numerous slender ± short herbaceous shoots, up to 6 cm tall (in cultivation up to 10 cm), internodes scabrous, shorter than leaves (in cultivation somewhat longer), leaves regularly spaced, internodes with retrorse indumentum of minute setose hairs approx. 0.1 mm in length, retrose on vegetative and antrorse on flowering shoots. Leaves and leaf-like stipules in rather closely-spaced whorls of (4–)6, sessile, more or less succulent in life, linear to narrowly oblanceolate, 0.8–1(1.2) × 4–5(7) mm with short-apiculate apex, with sparsely setose margins and lower mid-ribs. Inflorescence terminal, frondose, cymes on short setose peduncles ca. 1–3 mm long, the individual cymes reduced to solitary flowers; flowers sessile, subtended by 4 setose prophylls; calyx reduced; corolla 4-merous, salverform, (crème-)white, externally ± densely setose with patent hairs 0.2–0.3 mm long, internally glabrous, the tube 2.5–3.5 mm long, the lobes ca. 1 mm long; anthers 4, yellow, glabrous, ca. 0,8 mm long, included in the tube, filaments ca. 0,8 mm long, inserted ca. 2/3 of the tube; style glabrous, bifid, the last ca. 0.5 mm free, the 2 stigmas slightly elongated to ellipsoid, included in the tube; ovary ca. 1.3 mm long, apically truncate, ± densely setose especially towards the apex, with patent hairs 0.2–0.3 mm long. Mature infructescence and fruits unknown.

Figure 2. 

Habit and flowers of Thliphthisa sapphus A old individual of Th. sapphus with a thick lignified rootstock B flowers of Th. sapphus exhibiting the characteristic indumentum on the corolla (photographs by M. Sonnleitner).

Affinities of Thliphthisa sapphus

Due to the chamaephytic habit, with leaves (and leaf-like stipules) in rather closely-spaced whorls of 4–6, the presence of prophylls at the base of the sessile 4-merous flowers and the apically truncate ovary, Thliphthisa sapphus belongs to the subclade of the "Sherardia clade" (Manen et al. 1994; Natali et al. 1995), which is now classified as genus Thliphthisa (Del Guacchio and Caputo 2020). Within this group defined by Schönbeck-Temesy and Ehrendorfer (1985), the new species belongs to a group of chamaephytic taxa with presumably plesiomorphic flower structures, sharing well-developed corolla tubes and slightly elongated (not globose) stigmata. This group comprises Th. crassula (Greuter & Zaffran) P.Caputo & Del Guacchio, Th. tournefortii (Sieber) P.Caputo & Del Guacchio (group A of the former study), Th. baenitzii (Heldr. ex Boiss.) P.Caputo & Del Guacchio and Th. muscosa (Boiss. & Heldr.) P.Caputo & Del Guacchio (group B). Other species with similar flower morphology are xeromorphic subshrubs deviating by their broomrape-like appearance.

Thliphthisa sapphus is most similar to Th. baenitzii, but is readily distinguished by mericarps and corolla being ± densely setose (vs. glabrous), by its longer corolla of approximately 3.5–4.5 mm (vs. 2.5–3.5 mm in Th. baenitzii; Ehrendorfer and Krendl (1976), own measurements; the corolla length of 3–4 mm as given in Schönbeck-Temesy and Ehrendorfer (1985) is erroneous) and, in particular, by its tetraploid chromosome number that so far is unique within Thliphthisa (see below). Thliphthisa muscosa is also similar, but is well differentiated from both by its laxer growth, longer and ± erect flowering stems and distinctly longer, linear leaves.

Specimens examined

Additional gatherings (paratypes). Same locality as holotype: 10 November 1994, W. Gutermann et al. 28787 (WU 0154344, herb. Gutermann 28787); 14 July 1996: E. Hörandl 7766 (ATHU, WU 0154343, herb. Gutermann 80000); 26 April 2011: W. Gutermann et al. 39920 (WU 0154345, herb. Gutermann 39920).

Distribution and biogeographic considerations

To our present knowledge Thliphthisa sapphus is restricted to a small area around the type locality on the western edge of the Lefkátas Peninsula in south-western Lefkada. Restricted distribution is common in the genus Thliphthisa which generally comprises eastern Mediterranean oreophytes distributed east of the Adriatic Sea and reaching the Alborz Mts. of northern Iran (see maps in Schönbeck-Temesy and Ehrendorfer (1985): figs 9–12). Only three species with more derived flower characters (shortened corolla tubes and globose stigmata) occupy larger areas, notably Th. chlorantha (Boiss. & Heldr.) P.Caputo & Del Guacchio and Th. rupestris (Vis.) P.Caputo & Del Guacchio (NW Balkan mountain ranges), and especially Th. purpurea (L.) P.Caputo & Del Guacchio (Balkan Peninsula and Apennines to southern and south-western Alps). Apart from three Iranian species, all other species are scattered around the east Mediterranean Basin and have a more restricted distribution. This is especially true for the taxa that are morphologically most similar to Th. sapphus and that are known only from single mountain stocks: Th. baenitzii on Mt. Patéras and Th. muscosa on Mt. Olympus.

These three Hellenic steno-endemics form the north-western counterpart of the South Aegean Th. tournefortii and its local Cretan relative Th. crassula (the latter two constitute group A of Schönbeck-Temesy and Ehrendorfer (1985)). Thliphthisa tournefortii is a species of peculiar distribution (map in Runemark (1969: 115); Greuter (1971)) that can be associated chorologically with a “mesogaeic” element and, consequently, with a differentiation dating back to the Miocene (Greuter 1970). The same may be assumed for the above-mentioned more northern taxa Th. baenitzii, Th. muscosa and Th. sapphus sharing the more primitive flower structures with Th. tournefortii. The connections between these “sisters” are bridged geographically by another steno-endemic, still undescribed Thliphthisa species (Gutermann et al., in prep.) of Aetolia that, in habit and foliage, approaches the Cretan Th. crassula. Thus, Th. sapphus constitutes the westernmost outpost of this cluster of Greek steno-endemics. They underline the importance of cliffs as refugial habitats for poorly-competitive old chamaephytes (informally referred to as “Old ladies of the rocks”) through the times (as e.g. discussed in Davis (1951) and Snogerup (1971)).


The epithet commemorates Sappho, the most famous poetess of the Classical Hellenic Era and refers to the traditional (though non-historical) believe she suicidally threw herself from the white cliffs of Lefkátas because of unrequited love for beautiful young Phaon. The epithet is in genitive spelling. The genitive in classical Latin also is "Sapphūs", as testified in the poem delivered as "incerti avctoris epistvla Sapphvs ad Phaonem" in Ovid’s Epistolae Heroidum no. 15.


The chromosome number of Thliphthisa sapphus is here determined as 2n = 4x = 44. The base chromosome number of x = 11 was found in all previously investigated representatives of the genus Thliphthisa [i.e. all sub Asperula: Th. chlorantha, Th. muscosa (Faure and Pietrera 1969), Th. purpurea, Th. rupestris (Moore 1982), Th. crassula, Th. tournefortii (Montmollin 1986), Th. baenitzii (Constantinidis et al. 1997), Th. antalyensis (Ehrend.) P.Caputo & Del Guacchio, Th. brevifolia (Vent.) P.Caputo & Del Guacchio, Th. pseudochlorantha (Ehrend.) P.Caputo & Del Guacchio, Th. serotina (Boiss. & Heldr.) P.Caputo & Del Guacchio (Minareci and Yildiz 2011)], as well as in most Rubieae (Kiehn and Berger 2020, 2023). However, except for Th. sapphus, all the investigated taxa of Thliphthisa are diploid. The representative karyotype of Th. sapphus is mostly composed of metacentric and submetacentric chromosomes (Fig. 3), as reported in other Turkish Thliphthisa (formally Asperula sect. Thliphthisa) taxa (Minareci and Yildiz 2011).

Figure 3. 

Mitotic chromosomes and karyotype of Thliphthisa sapphus (2n = 4x = 44). Scale bar: 5 μm.

Phenology and growth

Flowering of Thliphthisa sapphus commences in May and, by early July, mostly withering or wilted corollas were observed. At this time, mericarps were not developed even in wilted flowers, but some were conspicuously and unusually swollen and infected by parasites, possibly gall midges (Diptera, Cecidomyiidae) as found parasitising on other Rubiaceae taxa in the Mediterranean area (e.g. Simova-Tošić et al. (2000); Skuhravá and Skuhravý (2016); Skuhravá et al. 2020). The plant cultivated in the HBV never developed fertile fruits, thus suggesting self-infertility. However, we have also failed to find fully-developed fruits in situ. The propagation rate of Th. sapphus seems to be very low, as nearly all cushions observed within this area are old plants with a thick lignified rootstock and taproot (of 1 cm ∅ and more) anchored in the rock fissures (Fig. 2A).

Ecology and phytosociology

It dwells on steep to nearly vertical limestone cliffs facing the sea and is, thus, exposed to westerly winds, either in fissures in solid rock or within a thin regolith cover (Fig. 4). However, these habitats at elevations of 200–230 m a.s.l. border the slightly elevated plateau of the Peninsula and show no marked salt influence. Whereas large parts of the plateau are dominated by low Quercus coccifera L. garrigue, areas with shallower and rockier soils, especially along the upper slope of the coastal cliffs, are covered by sclerophyllous phrygana vegetation with Anthyllis hermanniae L., Erica multiflora L., Juniperus phoenicea subsp. turbinata (Guss.) Nyman or Thymbra capitata (L.) Cav. Depending on the distribution of narrow rock ledges, erosion fissures/crevices and compact rock, the phrygana forms an intricate mosaic with sparse chasmophytic vegetation. As demonstrated by the vegetation relevés (Table 1), Thliphthisa sapphus is accompanied by another long-lived and cushion-forming chasmophyte Lomelosia crenata subsp. dallaportae (Boiss.) Greuter & Burdet showing scattered occurrences along the western coasts of the southern Ionian Islands (Flora Ionica Working Group 2016 onwards) and having also few isolated populations in Apulia (Italy) on Monte Gargano (Wagensommer 2012). Usually, together with Plocama calabrica (L.f.) M.Backlund & Thulin (Rubiaceae), they form a particular plant community covering the rock fissures.

Table 1.

Thliphthisa sapphus-Lomelosietum dallaportae Mucina: vegetation relevés were performed using a modified Braun-Blanquet cover-abundance scale (see Barkman et al. (1964) for details) (10 November 1994: 1–4; 16 May 1996: 5–11).

Running number 1 2 3 4 5 6 7 8 9 10 11
Species richness 12 8 10 6 18 20 29 11 6 14 17
Altitude (m) 235 230 230 230 230 230 230 230 230 230 230
Inclination (°) 60–70 60–90 80 70–90 60 40 40–50 0–90 80 70 40
Sampled area (m2) 3 3 9 6 8 9 4 7 6 8 9
Projected cover of vegetation (%) 25 10 40 30 30 60 50 80 35 40 40
Association character-taxa
Lomelosia crenata subsp. dallaportae (Boiss.) Greuter & Burdet 2b 2a 3 2b 2b 2a 3 3 3 3 2b
Thliphthisa sapphus Gutermann 2m 2a 2a 1 2a 2a 2b 3 2a 2a 2a
Plocama calabrica (L.f.) M.Backlund & Thulin + 2m + + 2a 1 2a 1 1 + ·
Asplenietea trichomanis
Brassica cretica subsp. aegaea (Heldr. & Hal.) Snogerup, M.A.Gust. & Bothmer · · · · · r r · · · ·
Ptilostemon chamaepeuce (L.) Less.; (juv.) · · · · · r · · · · ·
Stachys ionica Halácsy · · · · · · · + · · ·
Campanula versicolor Sm. · · · · · · r · · · ·
Melica minuta subsp. minuta L. · · · · · · + · · · +
Erica multiflora L. + 2a 2b 2a 2a 2b + · · · ·
Thymbra capitata (L.) Cav. + · · · r + + · · + r
Anthyllis hermanniae L. · · · 2a + 2b + · · + 1
Lotus hirsutus L. + · · · + + 1 · · · ·
Carlina corymbosa subsp. graeca (Heldr. & Sartori) Nyman · · · · r · · 1 · + 1
Carex illegitima Ces. · · · · · 1 2a · · · ·
Convolvulus elegantissimus Mill. · · · · · · + · · + +
Micromeria graeca subsp. graeca (L.) Benth. ex Rchb. · · + · · · · · · · ·
Cistus creticus L. · · · · · · 1 · · · ·
Cistus salviifolius L. · · · · · · · · · · 1
Fumana thymifolia (L.) Webb · · · · · · + · · · ·
Cytisus lanigerus DC.; (juv.) · · · · · + · · · · ·
Quercetea ilicis
Smilax aspera L. + · · · · · + · · · ·
Lonicera implexa Aiton · · · · · · + · · · ·
Pistacia lentiscus L.; (juv.) · · · · · + · · · · ·
Phillyrea latifolia L.; (juv.) · · · · · · · · · · ·
Quercus coccifera L.; (juv.) · · · · · · r · · · ·
Cotinus coggygria Scop.; (juv.) · · · · · + · · · · ·
Catapodium rigidum (L.) C.E.Hubb. + + · · r r + 1 · +
Lysimachia linum-stellatum L. · · · · r · r · · · ·
Arenaria leptoclados subsp. leptoclados (Rchb.) Guss. · · · · + · · · · · +
Hypochaeris achyrophorus L. · · · · r · · · · · ·
Festuca incurva (Gouan) Gutermann · · · · · · + · · · ·
Valantia muralis L. · · · · · · + · · · ·
Anthemis chia L. · · · · · · r · · · ·
Biscutella didyma L. · · · · · · · · · · +
Lolium rigidum Gaudin · · · · · · + · + · ·
Bromus madritensis L. · · · · · · · · · · +
Valerianella microcarpa Loisel. · · · · · · + · · · ·
Trifolium physodes Steven ex M.Bieb. · · · · · · · · · + ·
Geranium purpureum Vill. · · + · · · · · · · ·
Avena barbata Pott ex Link · · · · · · · · · · +
Brachypodium retusum (Pers.) P.Beauv. · · · · 1 2b · 2a + 2a ·
Leontodon graecus Boiss. & Heldr. + r r · · r + + · + +
Dactylis glomerata subsp. hispanica (Roth) Nyman · · · · · · + + · · ·
Helichrysum stoechas subsp. barrelieri (Ten.) Nyman · · · · + + · · · · ·
Reichardia picroides (L.) Roth + · · · · · + · · · ·
Other taxa
Lotus cytisoides L. · · + + · + · 2a · 1 1
Iberis carnosa Willd. + r · · r · · · · · ·
Petrosedum cf. ochroleucum (Chaix) Niederle + 1 · · 2m r · · · + ·
TaraxacumScariosa] sp. · · · · · · · + r · ·
Festuca jeanpertii subsp. achaica (Markgr.-Dann.) Markgr.-Dann. · · · · · · · · · 1 r
Bromus fasciculatus C.Presl · · · · · · 1 · · · ·
Salvia verbenaca L.; (juv.) · · · · · · · + · · ·
Trifolium campestre Schreb. · · · · · · + · · · ·
Medicago lupulina L. · · · · · · · · · + ·
Bupleurum trichopodum Boiss. & Spruner + · · · · · · · · · ·
Crucianella latifolia L. · · · · + · · · · · ·
Linum corymbulosum Rchb. + · · · · · · · · · ·
Scorpiurus subvillosus L. · · · · · · · · · · +
Mosses and lichens
Tortula sp. 1 · · · · · · · · · ·
Unknown moss · · + · · · · · · · ·
Lecanora sp. · · 2m · · · · · · · ·
Figure 4. 

Habitat and habit of Thliphthisa sapphus A coastal cliffs of the Lefkátas Peninsula, the locality of Th. sapphus (photograph by M. Hofbauer) B cushion of Th. sapphus in limestone rock fissure (photograph by M. Sonnleitner) C loose cushion of Th. sapphus within regolith cover on limestone rocks, together with Lomelosia crenata subsp. dallaportae (photograph by M. Sonnleitner).

We here describe this community as a new association, the Thliphthisa sapphus-Lomelosietum dallaportae Mucina ass. nova hoc loco (holotype: relevé 9 in Table 1) and classify this association tentatively in the Caro multiflori-Aurinion megalocarpae Terzi et D’Amico 2008, an alliance of thermo-mesomediterranean chasmophytic vegetation of limestone rock crevices of the south-eastern Italian Adriatic (Apulia) and Ionian coasts (see Mucina et al. (2016)). The syntaxonomic unit floristically and ecologically most similar is the Scabiosetum dallaportae Bianco et al. 1988 (see Bianco et al. (1988): table 4; Di Pietro and Wagensommer (2008: 198), table 3), an association described from Apulia, yet classified within the Asperulion garganicae Bianco et al. 1988 (for the syntaxonomic discussion of the Caro multiflori-Aurinion megalocarpae and the Asperulion garganicae, see Di Pietro and Wagensommer (2008)).

Conservation status

In the area directly accessible to us, we counted about two dozen mature individuals of Thliphthisa sapphus. Within the range of sight, further cushions were identified in the nearly vertical cliff slopes below. The size of this single known population can be estimated at roughly 100 individuals, although more populations may exist in the inaccessible cliffs continuously extending north and south of the known population. In 2011, a few individuals were also found in Erica L. phrygana above the slopes and established on the gravelly banks along a close-by road above the primary cliff habitat. The occurrence on competition-free road banks suggests some potential of the species to colonise open anthropogenic sites, for which an occurrence in unstable gravelly habitats may have been a pre-adaptation.

Due to the restricted known geographic range, the number of individuals may be drastically reduced by natural hazards, such as land or rock slides. The population is not exposed to immediate synanthropic threats, except for goats that graze even the steepest rock declivities such as the less precipitous slopes. Due to the small number of recorded individuals distributed within a range much less than 1 km2, the species falls into IUCN criterion D. Taking into account the possible threats mentioned above, it should be assessed as “Vulnerable” D2 (see IUCN (2022)). At national or regional scale, it classifies as a range-restricted endemic (as “r” in Dimopoulos et al. (2013)) and needs to be included in the “Red Data Lists” for Greece as “Endangered”. Further monitoring of the population is certainly needed and conservation measures, such as reduction of grazing, should be taken for the subpopulation in the accessible area at the plateau above the ridge.

Key to the Greek species (and subspecies) of the genus Thliphthisa (modified from Ehrendorfer and Krendl (1976); Schönbeck-Temesy and Ehrendorfer (1985))

1 Stigma globose 2
2 Corolla cup-shaped to rotate; main axis of synflorescence slender, lower branches elongate Th. chlorantha
2* Corolla infundibuliform; main axis of synflorescence robust, branches rather short 3
3 Corolla lobes acute; caespitose mountain plant usually < 20 cm in height 4
4 Corolla orange, 1–2 mm in length, corolla lobes outside with short stiff hairs; bracts frondose, longer than pedicels Th. pusilla
4* Corolla yellowish, 0.7–1 mm in length, corolla lobes outside glabrous; bracts small, shorter than pedicels Th. saxicola
3* Corolla lobes apiculate; plants of lower elevations, stems usually 20–45 cm in height 5 Th. purpurea
5 Leaves gradually acuminate, arranged in whorls of 7–10, margins distinctly revolute; corolla purple, apex of corolla lobes distinctly apiculate Th. purpurea subsp. purpurea
5* Leaves abruptly acuminate, arranged in whorls of 6–8, margins plane or only slightly revolute; corolla yellowish-green or purplish, apex of corolla lobes weakly apiculate Th. purpurea subsp. acuminata
1* Stigma ellipsoid to elongated 6
6 All leaves elliptic to broadly ovate or suborbiculate, apex rounded 7
7 Plant green (not glaucous), stems prostrate; leaves slightly succulent, hirsute; synflorescence short-cylindrical in outline; corolla approx. 3 mm in length Th. crassula
7* Plant glaucous, stems erect (up to 30(35) cm); leaves coriaceous, glabrous; synflorescence pyramidal in outline; corolla 3–5 mm in length Th. tournefortii
6* At least upper leaves linear to (ob)lanceolate, apex acute 8
8 Subshrubs; leaves caducous 9
9 Leaves 12–20 mm in length; corolla 6–7 mm in length, lobes ending in filiform appendices 1.5–1.7 mm in length; mericarps 2–3 mm in length Th. elonea
9* Leaves up to 10 mm in length; corolla 2–5(6) mm in length, lobes ending in short-triangular appendices 0.3–0.5 mm in length; mericarps 1–1.5 mm in length 10
10 Corolla 2–3.5 mm in length, reddish or yellowish; styles 2, separate, each approx. 1 mm in length Th. rigida
10* Corolla 3.5–5(6) mm in length, greenish-yellowish or brownish-green; style 1, furcate only towards the apex, approx. 3–3.5 mm in length Th. brevifolia
8* Plant herbaceous or woody only at the base; leaves persistent 11
11 Plants with erect flowering stems up to 16(20) cm in height; leaves narrowly lanceolate to linear, 0.8–1.5 × 9–12 mm, long acuminate Th. muscosa
11* Plants forming low, caespitose cushions 2–6(10) cm in height; leaves broadly elliptical to (ob)lanceolate, 0.8–2 × 4–7 mm, minutely acuminate 12
12 Corolla 3.5–4.5 mm in length, corolla lobes outside as well as the mericarps densely setose; leaves uniformly oblanceolate Th. sapphus
12* Corolla 2.5–3.5 mm in length, corolla lobes outside glabrous to sparsely hairy, mericarps glabrous; lower leaves elliptical, upper ones broadly lanceolate Th. baenitzii


Thanks go to our engaged companions on field-trips, Andraž Čarni, Milan Chytrý, Gerald Schneeweiss, Markus Staudinger and to Elvira Hörandl and Franz Hadacek, who gathered additional data in summer 1996. Manfred Hammer kindly advised verification of the correct genitive of the name Sappho and Michael Polansky helped providing material from the Herbarium W. Fieldwork on the Ionian Islands in the years 1994/1997 was financially supported by the FWF (Austrian Science Fund, Project P10466-BIO to WG and LM). Open access funding was provided by University of Vienna. Ladislav Mucina acknowledges logistical support of the Iluka Chair in Vegetation Science and Biogeography at Murdoch University, Perth, Western Australia. We are grateful to one anonymous reviewer, to Elmar Robbrecht and the section editor Petra De Block for their comments and suggestions, that considerably helped to improve this manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


FWF (Austrian Science Fund, Project P10466-BIO to WG and LM).

Author contributions

All authors contributed to this work.

Author ORCIDs

Walter Gutermann

Tae-Soo Jang

David Prehsler

Dieter Reich

Andreas Berger

Christian Gilli

Markus Hofbauer

Margarita Lachmayer

Ruth Sander

Michaela Sonnleitner

Ladislav Mucina

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.


  • Bianco P, Brullo S, Pignatti E, Pignatti S (1988) La vegetazione delle rupi calcareae della Puglia. Braun-Blanquetia 2: 133–151.
  • Constantinidis T, Kamari G, Phitos D (1997) A cytological study of 28 phanerogams from the mountains of SE Sterea Ellas, Greece. Willdenowia 27(1–2): 121–142.
  • Montmollin B de (1986) Etude taxonomique de la flore de la Crète. III. Nombres chromosomiques. Candollea 41: 431–439.
  • Del Guacchio E, Caputo P (2020) Splitting Asperula (Rubiaceae): A proposal for consistency purposes within sections Cynanchicae, Thliphthisa and Hexaphylla. Plant Biosystems 154(5): 766–782.
  • Di Pietro R, Wagensommer RP (2008) Analisi fitosociologica su alcune specie rare e/o minacciate del Parco Nazionale del Gargano (Italia centro-meridionale) e considerazioni sintassonomiche sulle comunità casmofitiche della Puglia. Fitosociologia 45: 177–200.
  • Dimopoulos P, Raus T, Bergmeier E, Constantinidis T, Iatrou G, Kokkini S, Strid A, Tzanoudakis D (2013) Vascular plants of Greece: An annotated checklist. Botanischer Garten und Botanisches Museum Berlin-Dahlem, Berlin; Hellenic Botanical Society, Athens, 1–372. [Englera 31]
  • Ehrendorfer F, Barfuss MHJ (2014) Paraphyly and polyphyly in the worldwide tribe Rubieae (Rubiaceae): Challenges for generic delimitation. Annals of the Missouri Botanical Garden 100(1–2): 79–88.
  • Ehrendorfer F, Krendl F (1976) 4. Asperula L. In: Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, Webb DA (Eds) Flora Europaea 4. Cambridge University Press, Cambridge, 4–14.
  • Ehrendorfer F, Barfuss MHJ, Manen J-F, Schneeweiss GM (2018) Phylogeny, character evolution and spatiotemporal diversification of the species-rich and world-wide distributed tribe Rubieae (Rubiaceae). PLoS ONE 13(12): e0207615.
  • Faure J, Pietrera D (1969) Contribution à l’etude cytotaxinomique des Rubiacees de Grèce. Annales de la Faculté des Sciences de Marseille 42: 271–283.
  • Flora Ionica Working Group (2016 onwards) Flora Ionica – An inventory of ferns and flowering plants of the Ionian Islands (Greece). [Accessed February 20, 2023]
  • Greuter W (1971) Betrachtungen zur Pflanzengeographie der Südägäis. Opera Botanica 30: 49–64.
  • Jang T-S, Emadzade K, Parker J, Temsch EM, Leitch AR, Speta F, Weiss-Schneeweiss H (2013) Chromosomal diversification and karyotype evolution of diploids in the cytologically diverse genus Prospero (Hyacinthaceae). BMC Evolutionary Biology 13(1): 136.
  • Kiehn M, Berger A (2020) Neotropical Rubiaceae: Synthesis of chromosome data from Costa Rican taxa, with insights on the systematics of the family. Annals of the Missouri Botanical Garden 105(4): 423–458.
  • Kiehn M, Berger A (2023) New chromosome counts on Rubiaceae from Africa and the Western Indian Ocean islands. Nordic Journal of Botany 2023(5): 1–14, e03732.
  • Manen J-F, Natali A, Ehrendorfer F (1994) Phylogeny of Rubiaceae-Rubieae inferred from the sequence of a cpDNA intergene region. Plant Systematics and Evolution 190(3–4): 195–211.
  • Minareci E, Yildiz K (2011) Karyotype characterization of Turkish taxa of the genus Asperula L.— Section Thliphthisa (Rubiaceae). Indian Journal of Genetics and Plant Breeding 71(1): 49–54.
  • Mucina L, Bültmann H, Dierßen K, Theurillat J-P, Raus T, Čarni A, Šumberová K, Willner W, Dengler J, Gavilán García R, Chytrý M, Hájek M, Di Pietro R, Iakushenko D, Pallas J, Daniëls FJA, Bergmeier E, Santos Guerra A, Ermakov N, Valachovič M, Schaminée JHJ, Lysenko T, Didukh YP, Pignatti S, Rodwell JS, Capelo J, Weber HE, Solomeshch A, Dimopoulos P, Aguiar C, Hennekens SM, Tichý L (2016) Vegetation of Europe: Hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science 19(S1, Suppl. 1): 3–264.
  • Natali A, Manen J-F, Ehrendorfer F (1995) Phylogeny of the Rubiaceae-Rubioideae, in particular the tribe Rubieae: Evidence from a non-coding chloroplast DNA sequence. Annals of the Missouri Botanical Garden 82(3): 428–439.
  • Runemark H (1969) Reproductive drift, a neglected principle in reproductive biology. Botaniska Notiser 122: 90–129.
  • Schönbeck-Temesy E, Ehrendorfer F (1985) Asperula gorganica und A. semanensis, zwei neue Arten aus dem Orient, und die palaeo-mediterrane Sektion Thliphthisa (Griseb.) Ehrend. (Rubiaceae). Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 107: 75–93.
  • Simova-Tošić D, Skuhravá M, Skuhravý V (2000) Gall midges (Diptera: Cecidomyiidae) of Serbia. Acta Entomologica Serbica 5(1/2): 47–93.
  • Skuhravá M, Skuhravý V (2016) Gall midges (Diptera: Cecidomyiidae) of Greece – summary of investigations of 1994–2010 and zoogeographical analysis. Acta Societatis Zoologicae Bohemicae 80: 127–163.
  • Skuhravá M, Skuhravý V, Salemkour N, Tahar Chauche S (2020) Gall midges (Diptera: Cecidomyiidae) of Algeria. Acta Societatis Zoologicae Bohemicae 84: 67–92.
  • Snogerup S (1971) Evolutionary and plant geographic aspects of chasmophytic vegetation. In: Davis PH, Harper PC, Hedge IC (Eds) Plant life of South-West Asia. Botanical Society of Edinburgh, Edinburgh, 158–169.
  • Soza VL, Olmstead RG (2010) Molecular systematics of tribe Rubieae (Rubiaceae): Evolution of major clades, development of leaf–like whorls, and biogeography. Taxon 59(3): 755–771.
  • Wagensommer RP (2012) Lomelosia crenata (Cirillo) Greuter et Burdet subsp. dallaportae (Boiss.) Greuter et Burdet. Informatore Botanico Italiano 44(1): 195–256.
  • Weiss-Schneeweiss H, Stuessy TF, Villaseñor JL (2009) Chromosome numbers, karyotypes, and evolution in Melampodium (Asteraceae). International Journal of Plant Sciences 170(9): 1168–1182.

Supplementary material

Supplementary material 1 

Specimens of Greek representatives of genus Thliphthisa examined

Walter Gutermann, Tae-Soo Jang, Arndt Kästner, David Prehsler, Dieter Reich, Andreas Berger, Ruth Flatscher, Christian Gilli, Markus Hofbauer, Margarita Lachmayer, Ruth Sander, Michaela Sonnleitner, Ladislav Mucina

Data type: docx

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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