Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology Official Journal of the Societa Botanica Italiana ISSN: 1126-3504 (Print) 1724-5575 (Online) Journal homepage: https://www.tandfonline.com/loi/tplb20 Splitting Asperula (Rubiaceae): a proposal for consistency purposes within sections Cynanchicae, Thliphthisa and Hexaphylla E. Del Guacchio & P. Caputo To cite this article: E. Del Guacchio & P. Caputo (2020): Splitting Asperula (Rubiaceae): a proposal for consistency purposes within sections Cynanchicae, Thliphthisa and Hexaphylla, Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology, DOI: 10.1080/11263504.2020.1804008 To link to this article: https://doi.org/10.1080/11263504.2020.1804008 Accepted author version posted online: 03 Aug 2020. Submit your article to this journal View related articles View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=tplb20 Splitting Asperula (Rubiaceae): a proposal for consistency purposes within sections Cynanchicae, Thliphthisa and Hexaphylla E. Del Guacchio1and P. Caputo1,2 1 Orto botanico, University of Naples “Federico II”, via Foria 223, I-80139, Naples, Italy 2 Department of Biology, University of Naples “Federico II”, Cupa Nuova Cinthia, I-80126, Naples, Italy. ip t Corresponding author E. Del Guacchio edelgua@email.it cr ABSTRACT us Authors report 139 new nomenclatural combinations in the sections Cynanchicae, Thliphthisa and Hexaphylla of the genus Asperula L. (Rubiaceae). These changes were deemed necessary as a consequence of numerous molecular phylogenetic studies which have provided abundant evidence on the polyphyletic nature of the genus. As the complete species-level phylogeny of the tribe Rubieae, and especially, of the genus Galium L. is not available as yet, the authors preferred to instigate changes only in the sections of Asperula whose phylogenetic position is deemed stable. an Keywords: Cynanchica; Molecular phylogenetics; Monophyly; Rubieae; Sherardia M Introduction Ac ce pt e d Retention of plesiomorphic features or homoplasy in morphological traits (especially those related to floral biology) have often obscured phylogenetic relationships in angiosperms. As mere examples, the understanding of the phylogeny of Caprifoliaceae s. l. (Bell et al. 2001; Donoghue et al. 2001; Winkworth et al. 2008), Liliales and Liliaceae (Dahlgren et al. 1985; Chase et al. 1995; Kim and Kim 2018), Scrophulariaceae and Lamiales (Olmstead and Reeves 1995; Schäferhoff et al. 2010), was greatly improved when molecular data or detailed anatomical studies showed that the traditional concepts of these taxa were often a blatant misrepresentation of the patterns of descent. Symplesiomorphies or homoplasies misconstrued as indications of relationships have been an almost widespread source of confusion also at lower circumscriptions as, for example, among genera of Dipsacoideae (Mayer and Ehrendorfer 1999; Avino et al. 2009; Carlson et al. 2009), Orchidinae (Aceto et al. 1999; Jin et al. 2014), Amaranthaceae (Iamonico 2015, Sánchez et al. 2016) or Rubieae (Natali et al. 1996; Soza and Olmstead 2010a; Yang et al. 2018). Obviously, any new insight on the phylogeny of a group is prone to beget nomenclatural changes, as it is for example evident by examination of the recent Italian checklists (Galasso et al. 2018; Bartolucci et al. 2018). More broadly, it raises the question as how to reconcile patterns of descent with the taxonomic hierarchy (De Queiroz 1997; Holt and Jønsson 2014) and, in particular, whether to deflect from the requirement of strict monophyly in circumscribing taxa. This last issue in the last twenty years has found some strenuous advocates on both sides (Sosef 1997; Nordal and Stedje 2005; Ebach et al. 2006; Hörandl 2007; Wyk 2007; Albach 2008; Hörandl and Stuessy 2010; Schmidt‐Lebuhn 2011; Brummit 2014; Stuessy and Hörandl 2014; Hörandl 2014), who base their contrasting arguments on reasons ranging from nomenclatural stability or other practical matters to evolutionary, generally biological or epistemological issues. However, no one of the aforementioned scientists seems to defend completely artificial taxa, if their polyphyletic status is evident beyond doubt. ip t Tribe Rubieae has been object of several phylogenetic investigations in the recent past (Ehrendorfer et al. 1994; Manen et al. 1994; Natali et al. 1995, 1996; Manen 2000; Nie et al. 2005; Soza and Olmstead 2010 a, 2010b; Ehrendorfer and Barfuss 2014; Jeong et al. 2016; Yang et al. 2016, 2018; Ehrendorfer et al. 2018). All these studies provided a generally congruent picture of the evolution in the tribe, by which, with the exceptions of the archaic and strongly disjunct genus Kelloggia Torr. ex Benth. & Hook.f., of Didymaea Hook.f. (little more than ten New World species), of Rubia L. (about 80 Eurasian and African taxa) and of the newly described genus Pseudogalium L.-E.Yang, Z.-L.Nie & H.Sun, the remaining Rubieae (about 900 species) may be described as a largely resolved monophyletic assemblage in which, however, the genus Galium L. is paraphyletic whereas Asperula L. is definitely polyphyletic (Soza and Olmstead 2010a; Ehrendorfer and Barfuss 2014; Ehrendorfer et al. 2018); Yang et al. 2018). What Yang et al. (2018) indicated as “Clade III” (fig. 1, p. 224), and Ehrendorfer et al. (2018) as the clade including the “Asperula Clade” and the “Galium Clade” (fig. 1, p. 6/26) is one of the most stable within Rubieae, as it was recovered (to the extent of the species employed) also in the investigations by Soza and Olmstead (2010a) and Ehrendorfer and Barfuss (2014). ce pt e d M an us cr The just mentioned “Clade III” (Yang et al. 2018) includes several subclades (which authors of the said paper label as “Clade III-a” to “Clade III-e”). “Clade III-a” is divided into two large monophyletic groups. One of them includes Callipeltis Steven, Crucianella L., Phuopsis (Griseb.) Benth. & Hook.f. and representatives of Asperula sect. Cruciana Griseb. (two species). The other includes Galium paniculatum (Bunge) Pobed., Sherardia arvensis L., one representative of Asperula sect. Hexaphylla Ehrend., in a sister group relationship with a clade including two species in representation of Asperula sect. Thliphthisa (Griseb.) Ehrend. and ten other taxa in representation of Asperula sect. Cynanchicae (DC.) Boiss. This phylogenetic outline is largely confirmed by Ehrendorfer et al. (2018). In particular, also these authors report a clade (“Clade VI” in their fig. 1) including Sherardia L., A. sect. Hexaphylla, A. sect. Cynanchicae and A. sect. Thliphthisa, employing four to seven species per section (in the absence of G. paniculatum, not taken into account in their analysis). Similarly to Yang et al. (2018), Ehrendorfer et al. (2018) also indicate that the three just mentioned sections of Asperula are monophyletic, even if their mutual relationships are not resolved. Ac The detailed topology for the other sections of Asperula and for the remaining Rubieae recovered in the analyses by Ehrendorfer et al. (2018) and Yang et al. (2018) is not entirely comparable. However, in neither investigation is Asperula sect. Glabella Griseb. monophyletic (paraphyletic in the former study, polyphyletic in the latter) and Asperula sect. Asperula (including A. arvensis L., which is the type of the generic name) has close relationships with sect. Hexaphylla, sect. Thliphthisa, and sect. Cynanchicae. All recent investigations on Rubieae clearly indicate that several evident morphological characters which have traditionally been employed to separate genera (e.g., the number of leafy appendages per whorl, the rotate, funnel-shaped or tubular corolla, and the organization of the inflorescences) are homoplasious (Soza and Olmstead 2010a; Ehrendorfer and Barfuss 2014; Yang et al. 2018; Ehrendorfer et al. 2018). However, the relationships of members of some infrageneric taxa, which may represent different evolutionary lineages, are still unknown (e.g., A. sect. Oppositifoliae Schischk. ex Schönb.-Tem. and the monotypic A. sect. Crucianelloides Boiss.), even if Ehrendorfer and Barfuss (2014) give a sketch of their phylogeny based on unpublished molecular results. As a consequence of this, modifying the nomenclature to make it congruent with the phylogeny of the whole Rubieae would be definitely premature. However, Ehrendorfer and Barfuss (2014) discussed at some length the possible strategy to employ in renaming Rubieae, and most reasonably concluded that “the first and obviously necessary step in this direction is the elimination of clearly polyphyletic taxa as Asperula s.l.” and that, when choosing between lumping and splitting the second should be preferred as “it makes relationships within the speciose Rubieae better visible on the binomial level and can be achieved with fewer nomenclatural changes” (Ehrendorfer and Barfuss 2014: 85–86). In this paper we particularly focus on Asperula sect. Cynanchicae, sect. Thliphthisa and sect. Hexaphylla. an us cr ip t Each one of these sections has been verified to be monophyletic (Gargiulo et al. 2015, Yang et al. 2018, Ehrendorfer et al. 2018), at least to the extent of the taxa employed. In addition, at least the sect. Cynanchicae has been investigated in a fairly greater detail (Gargiulo et al. 2015), in a study including 49 species (out of the nearly 80 recognized) and nearly 70 separate accessions. Molecular information is available for 4 species of the sect. Hexaphylla (out of 12 species presently recognized) and other 4 within the sect. Thliphthisa (22 species). However, these sections are generally very closely-knit units from a morphological point of view (especially as narrowly defined at present), for which differential retention of plesiomorphic features or homoplasy may be regarded as less likely. Besides, as already mentioned, these taxa belong to a monophyletic clade, together with Sherardia, both in Yang et al. (2018) (together with Galium paniculatum) and, with a less robust support, in Ehrendorfer et al. (2018). However, in Yang et al. (2018) A. sect. Thliphthisa is sister group to sect. Cynanchicae; this clade, in turn, is sister group to a clade including Galium paniculatum and a subclade constituted by A. hirta Ramond (sect. Hexaphylla) and Sherardia arvensis L. On the contrary, in Ehrendorfer et al. (2018), the clades representing respectively A. sect. Cynanchicae, sect. Thliphthisa, sect. Hexaphylla and Sherardia, even if well supported, are not reciprocally resolved. M We here propose a new monophyletic classification of the taxa belonging to the above-mentioned sections, attempting at a reconciliation between the recent molecular findings and the morphological observations. ce pt e d This contribution is to be intended as a preliminary contribution for the ongoing taxonomic revision of the genus Asperula in Italy (see e.g., Del Guacchio and Caputo 2005, 2013a; Gargiulo et al. 2015; Del Guacchio et al. 2016). MATERIAL AND METHODS Ac Our proposal mostly relies on the studies of Sosa and Olmstead (2010a, 2010b), Gargiulo et al. (2015), Jeong et al. (2016), Yang et al. (2018), and Ehrendorfer et al. (2018), integrated with morphological observations. A consensus tree based on the above cited contributions and limited to the taxa of our interest is shown in fig. 1. It serves as a base for our discussion. For nomenclatural purposes, i.e. to trace synonymies and establish priority, we checked the following databases: Marhold (2011+), IPNI (2019), POWO (2019), Tropicos (2019), and Govaerts (2019). Especially on this latter comprehensive and updated work we relied for the taxonomic treatment, except where otherwise specified. In addition, several monographs and standards floras where consulted. Even if a full taxonomic revision is beyond the purposes of the present work, numerous specimens from Europe and western Asia where examined, kept in BC, FI, G, M, NAP, P, RO, VAL, W (herbarum codes according to Thiers, 2018). Further specimens from B, BM and P were examined online. The protologues of all the basionyms where consulted and corrections and amendments to the above cited databases are reported in the notes. The articles cited throughout text follow the Shenzhen Code (Turland et al. 2018), ICN hereafter. RESULTS AND DISCUSSION One hundred and thirty-nine new combinations are here proposed, encompassing, to the best of our knowledge, all the taxa currently recognized by the monographers of the groups in study. These taxa have been accommodated in four genera, as indicated in the next paragraph. Concerning these genera, one name was already available at generic level; two new genera were named by changing the status available sectional names; a new genus is described. In addition, a nomen novum is proposed, mainly to avoid a tautonym. Overall, a total of 154 nomenclatural novelties are proposed. The reasons of our nomenclatural choices are given here. t Asperula sect. Cynanchicae, as circumscribed by modern authors (e.g., Ehrendorfer and Krendl 1976) is homogeneous from a morphological standpoint (see below), and this is also strongly supported by molecular studies, which have proved that no group of lesser rank can be convincingly recognized. Indeed, at the infrasectional level, geography seems to overcome any taxonomic consideration (Gargiulo et al. 2015). We employed this well studied section as a benchmark for the construction of a coherent taxonomic proposal. M an us cr ip Splitting Asperula into segregate monophyletic taxa necessarily implies including all the member of sect. Cynanchicae in a single taxon, which could be indeed recognized as an autonomous genus or alternatively included with sect. Thliphthisa and sect. Hexaphylla in Sherardia (this Linnaean name would clearly claim priority at the generic rank). The following considerations strongly suggest to keep Asperula sect. Cynanchicae, sect. Thliphthisa, sect. Hexaphylla and Sherardia as separate taxa at the same rank. The presently monospecific genus Sherardia, with S. arvensis, shows at least three notable autapomorphies: (1) annual habit, (2) well developed calyx teeth, growing in fruit, (3) six teeth. In addition, even if Sherardia definitely resembles taxa of A. sect. Hexaphylla much more than those of A. sect. Cynanchicae or sect. Thliphthisa, molecular results by Ehrendorfer et al. (2018) do not authorize the inclusion of A. sect. Hexaphylla in Sherardia, as the consensus tree by Yang et al. (2018) may suggest. ce pt e d On the other hand, a very large genus Sherardia, including also Asperula sections Cynanchicae and Thliphthisa (without mentioning Galium paniculatum), would imply that, for the sake of consistency, further very comprehensive genera should be recognized throughout Rubieae. This would cause nomenclatural disappearance of some small but well characterized clades and would obscure the phyletic lineages in Asperula s. l.mo, with excessive taxonomic flattening, and, indeed, without any nomenclatural advantage (as all the taxa of Asperula sections Cynanchica and Thliphthisa would be moved to Sherardia). Indeed, a genus Sherardia so conceived would result too heterogeneous, and should be necessarily divided into several subgenera and sections. Ac Our proposal, therefore, is to raise to generic rank the three sections of Asperula which are the object of the present study. As a consequence, this choice implies numerous new combinations: a disadvantage which is almost impossible to avoid, or even to minimize following alternative taxonomic treatments. In fact, the generitype of Asperula L., nom. cons. is A. arvensis L., typ. cons.; and sect. Arvensis in no case could include the taxa of our interest without phagocytizing many welldistinct, largely accepted and indeed monophyletic genera, in addition to several sections of Galium. Paradoxically, this would require in turn a much larger number of new combinations. According to Yang et al. (2018), A. sect. Cynanchicae, A. sect. Thliphthisa, and A. sect. Sherardia belong to a well-supported clade with Galium paniculatum, an aberrant central-Asian species, alternatively regarded as belonging to Asperula or Galium by different authors (Chen and Ehrendorfer, 2011: 131–132). A satisfactory taxonomic assessment in this case would require further studies. Similarly, we are not aware of any published molecular information on the West-Asian A. sect. Oppositifoliae Schischk. ex Schönb.-Tem., which appears as very close to polymorphic A. sect. Cynanchicae (Ehrendorfer et al., 2018), and, in our opinion, probably to be included in it. In fact, so far as we can argue by the examination of descriptions and exsiccata, the morphological traits which could help in discrimination towards sect. Cynanchicae are rather weak and sometimes largely subjective. For example, the most relevant feature in sect. Oppositifoliae is the absence of welldeveloped stipules and therefore the arrangement of the leaves in pairs; this, however, also occurs in some representatives of sect. Cynanchicae. For this reason, it is preferable to wait for further studies before attempting at a nomenclatural proposal concerning this group. TAXONOMIC TREATMENT AND NOMENCLATURAL IMPLICATIONS In the following summary, we list synonyms down to the subspecies rank; the most relevant basionyms are reported as well. ip t Cynanchica P.Caputo & Del Guacchio, gen. nov. us cr Description: Perennial plants, herbaceous or rarely suffruticose, sometimes rhizomatose or stoloniferous, with whorls with 4 lanceolate or linear elements (sometimes the stipules are very reduced upwards), bracts and bracteoles, partial inflorescences from capitate to spiciform, with hypocrateriform or infundibuliform corollas, with 4 lobes as a rule, calyx teeth inconspicuous, stamens inserted and subequal to the capitate stigma, fruits ovate, never spiny. an Type: Asperula cynanchica L., Sp. Pl.: 104. 1753 [≡ Cynanchica pyrenaica subsp. cynanchica (L.) P.Caputo & Del Guacchio]. Lectotype of the name Asperula cynanchica: Herb. Burser XIX: 8 (UPSBURSER), designated by Ferrer-Gallego (2015, p. 100). ce pt e d M Etym: The name derives from ancient Greek and is an adjective referring to an affection of the throat or larynx (possibly a tonsillitis) against which ancient medicinal use of Cynanchica (and similar plants) is recorded. In the protologue of A. cynanchica, Linnaeus (1753) cited several authors, and in particular Bauhin for the use of word of our interest (as in the synonym “Rubia cynanchica Bauh. Pin. 333, Bauh. Hist. 3. p. 723.”). However, Bauhin (1623) in turn referred to Dalechamp (1586), who wrote “Synanchica” and regarded it as a “genus”. Curiously, his “Synanchica” cannot be referred to any Cynanchica species, on account of its whorls of 5–6 elements. Ac Distribution: The genus includes about 100 taxa (83 species in our treatment), widespread in western Palearctic, and especially differentiated throughout the Mediterranean Basin and around the Black Sea, where it occurs with several narrow endemics (Klokov 1958; Ehrendorfer & Krendl 1976). ≡ Asperula [unranked] Cynanchicae DC., Prodr. 4: 582. 1830 p. p., limit. Ehrendorfer (1975) ≡ Asperula sect. Cynanchicae (DC.) Boiss., Diagn. Pl. Orient., ser. 1, 3: 31. 1843 [“Asperula sect. Cynanchica (DC.) Griseb.”, Spic. Fl. Rumel. 2: 164, 1846 [Jan, “1844”] (Art. 41.4), isonym (Art. 6.3, Note 2)]. Type automatically determined according to Art. 10.8 (cf. Ehrendorfer 1975, p. 2) = Asperula sect. Rubeola Lange in Willkomm & Lange, Prodr. Fl. Hisp. 2: 301. 1868 p. p., non Griseb. Type: Asperula aristata L. f., Suppl. Pl.: 120. 1794 [≡ Cynanchica aristata (L.f.) P.Caputo & Del Guacchio], designated by Ehrendorfer (1975, p. 2) – “Cynanchica Fourr.” in Ann. Soc. Linn. Lyon n.s. 16: 398. 1868, nom. nud. cr ip t Notes: Fourreau (1868, p. 398) published the generic name “Cynanchica” only listing two species, i.e. “C. tinctoria (L.)”, an intended new combination based on A. tinctoria L.; and “C. tenuifolia”, an intended replacement name for A. cynanchica, which he cited and whose combination under Cynanchica would cause a tautonym (Art. 23.4 of the ICN). Fourreau did not provide any description or potentially validating synonym so, even if his intent was to base his generic name on Asperula [unranked] Cynanchicae DC., as his name did not otherwise meet the conditions for valid publicatiom, Art. 41.4 cannot be applied and his name is not validly published. Besides, it should to be noted that Fourreau had a different circumscription for his genus than that of the our new genus Cynanchica, as A. tinctoria appears to be quite removed from it (Ehrendorfer et al. 2018; Yang et al. 2018). We might have published Cynanchica as a new name with new status based on Asperula [unranked] Cynanchicae DC., but as it would entail a change from a plural adjective to a singular noun, it seemed simpler to describe it as a new genus, homotypic with the name of the generic subdivision. Another name which was taken into consideration is ×Asperugalium (Asperula × Galium), P.Fourn., Quatre Fl. France: 883. 1938. Asperugalium includes a representative of the new genus Cynanchica, i.e. Asperugalium ×occidentale (Rouy) P.Fourn. (Asperula occidentalis Rouy) (thought by Fournier to be Asperula cynanchica × Galium arenarium Loisel.). By chance, this latter taxon is not in fact a hybrid (Ehrendorfer & Krendl 1976). However, Asperugalium is not validly published as a generic name, but only as a nothogeneric name (a condensed formula), by means of a statement of the parent genera (Art. H.9.1); as such it a later synonym of ×Galiasperula Ronniger (Repert. Spec. Nov. Regni Veg. 29: 143. 1931). d M an us The group “Cynanchicae” as conceived by Candolle (1830) remained very heterogeneous for a long time, as it also included relatively distant species such as A. taurina L., A. arvensis and A. tinctoria (e.g., Grisebach 1844, Lange 1870, Schumann 1891), but it was progressively limited by Ehrendorfer (1958, 1975). For the new genus Cynanchica, we adopt the circumscription illustrated in Ehrendorfer and Krendl (1976) and Ehrendorfer and Schönbeck-Temesy (1982), because it fully complies with the recent molecular findings. The most relevant synapomorphy of the group is represented by the whorls of 4 elements. Even if the Spanish endemic A. paui Viv. is sometimes reported as bearing leaves in pairs (Ortega Oliviencia and Devesa 2007: p. 46–47, fig. 10), almost all the numerous specimens that we examined show at least the lower internodes with two stipules (even if often very reduced). ce pt e Cynanchica abbreviata (Halácsy) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lutea var. abbreviata Halácsy, Consp. Fl. Graec. 1: 737. 1901 [basion.] ≡ Asperula abbreviata (Halácsy) Rech.f., Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 105(2; 1): 132. 1943. [n.v.] Ac Cynanchica abchasica (V.I.Krecz.) P. Caputo & Del Guacchio, comb. nov. ≡ Asperula abchasica V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 27. 1934 [basion.] Cynanchica accrescens (Klokov) P. Caputo & Del Guacchio, comb. nov. ≡ Asperula accrescens Klokov in Komarov, Fl. URSS 23: 704. 1958 [basion.] Cynanchica affinis (Boiss. & A.Huet) P. Caputo & Del Guacchio, comb. nov. ≡ Asperula affinis Boiss. & A.Huet in Boissier, Diagn. Pl. Orient., ser. 2, 2: 110. 1856 [basion.] ≡ A. cynanchica var. affinis (Boiss. & A.Huet) Boiss., Fl. Orient. 3: 41. 1875 = A. dolichophylla Klokov in Komarov, Fl. URSS 23: 706. 1958 Cynanchica aristata (L.f.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula aristata L.f., Suppl. Pl.: 120. 1794 [basion.] ≡ A. cynanchica subsp. aristata (L.f.) Bég. in Fiori, Fl. Italia 3: 122. 1903 – “Galium aristulatum F.Herm.”, Fl. N. Mitteleur.: 942. 1956, comb. inval. Note: Hermann (1956) cited A. aristata L.f. as basionym. The adoption of the epithet “aristulatum” instead of “aristatum” cannot be interpreted as a lapsus. In fact, Hermann was aware that the adjective aristatum was not available in Galium, due to the existence of Galium aristatum L. (cf. Hermann 1956: 946). The name Galium aristulatum must therefore be intended as a replacing name; however, it is not validly published, lacking a full direct reference to the replaced synonym (Art. 41.5). ip Cynanchica aristata subsp. calabra (Fiori) P.Caputo & Del Guacchio, comb. nov. t Cynanchica aristata subsp. aristata us cr ≡ A. cynanchica var. calabra Fiori, Nuov. Fl. Italia 2: 503. 1927 [basion.] ≡ A. calabra (Fiori) Gavioli in Arch. Bot. Sist. 8: 61. 1932 ≡ A. aristata subsp. calabra (Fiori) Del Guacchio & P.Caputo in Pl. Biosystems 151: 358. 2016 an Note: According to our previous treatment (Del Guacchio et al., 2016), the presumed hybrid A. ×portae Peruzzi (in Inform. Bot. Ital. 42: 530. 2010), is a transitional morph between subsp. calabra and subsp. scabra, which does not deserve taxonomic recognition and therefore any new combination under Cynanchica. M Cynanchica aristata subsp. condensata (Heldr. ex Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula longiflora var. condensata Heldr. ex Boiss., Fl. Orient., Suppl.: 281. 1888 [basion.] ce pt e d ≡ A. aristata subsp. condensata (Heldr. ex Boiss.) Ehrend. & Krendl, Bot. J. Linn. Soc. 68: 268. 1974 Cynanchica aristata subsp. nestia (Rech.f.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula nestia Rech.f., Bot. Jahrb. Syst. 69: 510. 1939 [basion.] ≡ A. aristata subsp. nestia (Rech.f.) Ehrend. & Krendl, Bot. J. Linn. Soc. 68: 268. 1974 Ac Cynanchica aristata subsp. scabra (Nyman) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula aristata subsp. scabra Nyman, Consp. Fl. Eur.: 334. 1879 [basion.] = A. cynanchica var. oreophila Briq. in Schinz & Keller, Fl. Schweiz: 484. 1900 = A. longiflora Waldst. & Kit., Descr. Icon. Pl. Hung. 2: 162. 1804 = A. umbellulata Reut., Compt.-Rend. Trav. Soc. Hallér. 1854–1856: 143. 1854 = A. exaristata Lacaita, Bull. Orto Bot. Regia Univ. Napoli 6: 170. 1921 Note: For the intricate nomenclature of this taxon, see also Del Guacchio & Caputo (2013b). Cynanchica aristata subsp. thessala (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula thessala Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., ser. 2, 2: 111. 1856 [basion.] ≡ A. aristata subsp. thessala (Boiss. & Heldr.) Hayek, Repert. Spec. Nov. Regni Veg. Beih. 30(2): 452. 1930 = A. peristeriensis Halácsy ex Maire & Petitm., Matér. Étude Fl. Géogr. Bot. Orient. 4: 107. 1908 ip t Note: The name of the new taxon and its diagnosis were written “in litteris” by Halácsy, who, however, added “basing on the few specimens, I do not dare to propose, as usual, a new species” (Maire and Petitmengin 1908). The latter authors attributed the new species to Halácsy, even with a doubt derived from his apparently provisional choice: “Hal. nov. sp. ad interim?”. Interestingly, later Hayek (1930) described the taxon again basing on the contribution of Halácsy, but recognizing the subspecific rank and with a different spelling (peristeraea), without citing Maire and Petitmengin (1908). It is questionable whether A. peristeriensis should be ascribed only to Maire and Petitmengin (cf. Art. 36.1, Art. 46.2). us Cynanchica beckiana (Degen) P.Caputo & Del Guacchio, comb. nov. cr = A. aristata subsp. peristeraea Halácsy ex Hayek, Repert. Spec. Nov. Regni Veg. Beih. 30(2): 452. 1930 ≡ Asperula beckiana Degen, Magyar Bot. Lapok 7: 105. 1908 [basion.] an Cynanchica biebersteinii (V.I.Krecz.) P.Caputo & Del Guacchio, comb. nov. M ≡ Asperula biebersteinii V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 26. 1934 [basion.] Cynanchica boissieri (Heldr. ex Boiss.) P.Caputo & Del Guacchio, comb. nov. d = Asperula boissieri Heldr. ex Boiss., Fl. Orient., Suppl.: 281. 1888 [basion.] ce pt e Cynanchica borbasiana (Korica) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula woloszczakii subsp. borbasiana Korica, Phyton (Horn) 17: 146. 1975 [basion.] ≡ A. borbasiana (Korica) Korica, Bot. Jahrb. Syst. 102: 340. 1981 Cynanchica bornmuelleri (Velen.) P.Caputo & Del Guacchio, comb. nov. Ac ≡ Asperula bornmuelleri Velen., Fl. Bulg., Suppl.: 143. 1898 [basion.] Note: The authorship of Asperula bornmuelleri is sometimes given as “Velenovský ex Bornmüller” (Govaerts 2019). However, despite Bornmüller (1931) regarding the name published by Velenovský (1898) as a “nomen subnudum” [i.e., a name “almost” without description], this latter author doubtless validly published it by a diagnosis and a description, with indication of syntypes and locality. = A. refracta Czeczott, Acta Soc. Bot. Poloniae 9: 39. 1932 Cynanchica brachyphylla (Trigas & Iatroú) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula brachyphylla Trigas & Iatroú, Phyton (Horn) 43: 30. 2003 [basion.] Cynanchica breviflora (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula breviflora Boiss., Diagn. Pl. Orient., ser. 1, 10: 63. 1849 [basion.] Cynanchica bryoides (Stapf) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula bryoides Stapf, Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 50(2): 106. 1885 [basion.] Cynanchica capitellata (Hausskn. & Bornm. ex Bornm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula capitellata Hausskn. & Bornm. ex Bornm., Repert. Spec. Nov. Regni Veg. Beih. 89: 296. 1941 [basion.] Cynanchica carpatica (Morariu) P.Caputo & Del Guacchio, comb. nov. an = A. tomentosa Ten., Prodr. Fl. Napol. 1: XII. 1811 cr ≡ Asperula crassifolia L., Mant. Pl.: 37. 1767 [basion.] us Cynanchica crassifolia (L.) P.Caputo & Del Guacchio, comb. nov. ip t ≡ Asperula carpatica Morariu, Stud. Cercet. Biol. (Bucharest) Ser. Biol. Veg. 31(2): 88. 1979 [basion.] Cynanchica cristata (Sommier & Levier), P.Caputo & Del Guacchio, comb. nov. M ≡ Asperula cynanchica var. cristata Sommier & Levier, Trudy Imp. S.-Peterburgsk. Bot. Sada 16: 204. 1900 [basion.] ≡ A. cristata (Sommier & Levier) V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 26. 1934. ce pt e d Cynanchica daphneola (O.Schwarz) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula daphneola O.Schwarz, Repert. Spec. Nov. Regni Veg. 36: 139. 1934 [basion.] Cynanchica deficiens (Viv.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula deficiens Viv., Ann. Storia Nat. 4: 231. 1830 [basion.] Ac Cynanchica diminuta (Klokov) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula diminuta Klokov in Komarov, Fl. URSS 23: 707. 1958 [basion.] Cynanchica garganica (Huter, Porta & Rigo ex Ehrend. & Krendl) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula garganica Huter, Porta & Rigo ex Ehrend. & Krendl, Bot. J. Linn. Soc. 68: 268. 1974 [basion.] Cynanchica glareosa (Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula glareosa Ehrend., Notes Roy. Bot. Gard. Edinburgh 22: 342. 1958 [basion.] Cynanchica graveolens (M.Bieb. ex Schult. & Schult.f.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula graveolens M.Bieb. ex Schult. & Schult.f., Mant. 3: 376. 1827 [basion.] Cynanchica graveolens subsp. danilewskiana (Basiner) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula danilewskiana Basiner, Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 2: 202. 1844 [basion.] ≡ A. graveolens subsp. danilewskiana (Basiner) Pyatunina, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 100(4): 77. 1995 = A. laevissima Klokov in Komarov, Fl. URSS 23: 708. 1958 ip t Note: Govaerts (2019) refers this synonym to C. graveolens subsp. graveolens, but see Klokov (1958), Pobedimova (1978), Czerepanov (1995), and Elenevsky and Pjatunina (1995). Note that Ehrendorfer and Krendl (1976) accept A. savranica, A. setulosa, A. lejograveolens and A. diminuta as valid species. Cynanchica graveolens subsp. graveolens = A. savranica Klokov in Kom., Fl. URSS 23: 707. 1958 us cr = A. pseudograveolens Popov & Chrshan., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 50(5– 6): 94. 1946 an Cynanchica graveolens subsp. leiograveolens (Popov & Chrshan.) P.Caputo & Del Guacchio, comb. nov. M ≡ Asperula leiograveolens Popov & Chrshan., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 50(5-6): 96. 1946 [basion.] ≡ A. graveolens subsp. leiograveolens (Popov & Chrshan.) Pjatunina, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 100(4): 76. 1995 ce pt e d Cynanchica gussonei (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula gussonei Boiss., Diagn. Pl. Orient., ser. 1, 10: 63. 1849 [basion.] = A. gussoneana Boiss. ex Ces., Pass. & Gibelli, Comp. Fl. Ital.: 559. 1886 Ac Note: Originally spelled as “gussoniana”, derived from “Gussonius” (latinized form of “Gussone”], but see Art. 60.0, Ex. 31. Cynanchica ×jordanii (E.P.Perrier & Songeon) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula ×jordanii E.P.Perrier & Songeon, Bull. Soc. Hist. Nat. Savoie 1853: 196. 1854 [basion.] Cynanchica icarica (Ehrend. & Schönb.-Tem.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula icarica Ehrend. & Schönb.-Tem., Pl. Syst. Evol. 134: 133. 1980 [basion.] Cynanchica idaea (Halácsy) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula idaea Halácsy, Consp. Fl. Graec. 1: 737. 1901 [basion.] Cynanchica inopinata (Schönb.-Tem.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula inopinata Schönb.-Tem., Fl. Iranica 276: 145. 2005 [basion.] Cynanchica intersita (Klokov) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula intersita Klokov in Komarov, Fl. URSS 23: 705. 1958 [basion.] Cynanchica kemulariae (Manden.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula kemulariae Manden., Zametki Sist. Geogr. Rast. 17: 58. 1953 [n.v.] [basion.] Cynanchica lactea (Brullo & al.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lactea Brullo, Gargano, N.G.Passal. & Peruzzi, Phytotaxa 196: 24. 2015 [basion.] ip t ≡ A. montana Waldst. & Kit. ex Willd. var. lactea Porta ex Galasso, Phytotaxa 196: 24. 2015 us cr Note: The previous two names were simultaneously published at different rank by different authors for special reasons (Peruzzi et al. 2015), but see Bartolucci et al. (2019), and Govaerts (2019). In any case, in our opinion, the distinctness of this taxon with respect to A. aristata subsp. aristata deserves further study, but it cannot be included in A. cynanchica, whose presence in southern Italy is very doubtful (Del Guacchio et al. 2016). an Cynanchica lilaciflora (Boiss.) P.Caputo & Del Guacchio, comb. nov. = A. acerosa K.Koch, Linnaea 19: 31. 1846 M ≡ Asperula lilaciflora Boiss., Diagn. Pl. Orient. 3: 33. 1843 [basion.] d Cynanchica lilaciflora subsp. coa (Rech. f.) P.Caputo & Del Guacchio, comb. nov. ce pt e ≡ Asperula coa Rech.f., Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 105: 566. 1943 [basion.] = A. lilaciflora subsp. coa (Rech.f.) Ehrend., Notes Roy. Bot. Gard. Edinburgh 37: 245. 1979 Cynanchica lilaciflora subsp. lilaciflora Ac Cynanchica lilaciflora subsp. mutensis (Schönb.-Tem.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lilaciflora subsp. mutensis Schönb.-Tem., Notes Roy. Bot. Gard. Edinburgh 37: 245. 1979 [basion.] Cynanchica lilaciflora subsp. phrygia (Bornm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula stricta var. phrygia Bornm., Beih. Bot. Centralbl. 24: 467. 1909 [basion.] ≡ A. lilaciflora subsp. phrygia (Bornm.) Schönb.-Tem., Notes Roy. Bot. Gard. Edinburgh 37: 245. 1979 Cynanchica lilaciflora subsp. runemarkii (Ehrend. & Schönb.-Tem.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lilaciflora subsp. runemarkii Ehrend. & Schönb.-Tem., Pl. Syst. Evol. 134: 134. 1980 [basion.] Cynanchica lipskyana (V.I.Krecz.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lipskyana V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 26. 1934 [basion.] Cynanchica litardierei (Humbert) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula litardierei Humbert, Bull. Soc. Hist. Nat. Afrique N. 18: 147. 1927 [basion.] Cynanchica littoralis (Sm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula littoralis Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 89. 1806 [basion.] ip ≡ Asperula lutea Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 88. 1806 [basion.] t Cynanchica lutea (Sm.) P.Caputo & Del Guacchio, comb. nov. cr Cynanchica lutea subsp. euboea (Ehrend.) P.Caputo & Del Guacchio, comb. nov. us ≡ Asperula lutea subsp. euboea Ehrend., Bot. Jahrb. Syst. 80: 403. 1961 [basion.] an Cynanchica lutea subsp. griseola (Greuter) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lutea subsp. griseola Greuter, Bocconea 25: 107. 2012 [basion.] M Cynanchica lutea subsp. lutea Cynanchica lycia (Stapf) P.Caputo & Del Guacchio, comb. nov. ce pt e d ≡ Asperula lycia Stapf, Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 50(2): 105. 1885 [basion.] Cynanchica malevonensis (Ehrend. & Schönb.-Tem.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula malevonensis Ehrend. & Schönb.-Tem. in Strid & Kit Tan (eds.), Mount. Fl. Greece 2: 294. 1991 [basion.] Ac = A. boissieri var. transiens Halácsy, Oesterr. Bot. Z. 45: 340. 1895 Cynanchica markothensis (Klokov) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula markothensis Klokov in Komarov, Fl. URSS 23: 700. 1958 [basion.] Cynanchica mungieri (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula mungieri Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., ser. 1, 10: 61. 1849 [basion.] ≡ A. lutea subsp. mungieri (Boiss. & Heldr.) Ehrend. & Krendl, Bot. J. Linn. Soc. 68: 269. 1974 Cynanchica naufraga (Ehrend. & Gutermann) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula naufraga Ehrend. & Gutermann, Bot. Chron. 13: 62. 2000 [basion.] Cynanchica neilreichii (Beck) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula neilreichii Beck, Verh. Zool.-Bot. Ges. Wien 32: 182. 1883 [basion.], nom. nov. pro A. cynanchica var. alpina Neilr., Fl. Nieder-Österreich: 464. 1859 (Art. 6.11, Ex. 14) – “Galium neilreichii (Beck) F.Herm.”, Fl. N. Mitteleur.: 942. 1956, comb. inval. (Art. 41.5) Cynanchica nitida (Sm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula nitida Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 89. 1806 [basion.] Cynanchica nitida subsp. hirtella (Boiss.) P.Caputo & Del Guacchio, comb. nov. ip t ≡ Asperula nitida var. hirtella Boiss., Fl. Orient. 3: 39. 1875 [basion.] ≡ A. nitida subsp. hirtella (Boiss.) Ehrend., Notes Roy. Bot. Gard. Edinburgh 37: 245. 1979 cr Cynanchica nitida subsp. mytilinica (Ehrend.) P.Caputo & Del Guacchio, comb. nov. us ≡ Asperula nitida subsp. mytilinica Ehrend., Notes Roy. Bot. Gard. Edinburgh 37: 244. 1979 [basion.] an Cynanchica nitida subsp. nitida Cynanchica nitida subsp. subcapitellata (Ehrend.) P.Caputo & Del Guacchio, comb. nov. M ≡ Asperula nitida subsp. subcapitellata Ehrend., Notes Roy. Bot. Gard. Edinburgh 37: 245. 1979 [basion.] d Cynanchica oetaea (Boiss.) P.Caputo & Del Guacchio, comb. nov. ce pt e ≡ Asperula lutea var. oetaea Boiss., Fl. Orient. Suppl.: 280. 1888 [basion.] ≡ A. oetaea (Boiss.) Heldr. ex Halácsy, Consp. Fl. Graec. 1: 739. 1901 ≡ A. lutea subsp. oetaea (Boiss.) Nyman, Consp. Fl. Eur. Suppl. 2: 156. 1889 Note: Nyman (1889) did not cite the basionym, but see Art. 41.3. Ac Cynanchica ophiolitica (Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula ophiolithica Ehrend. in Pl. Syst. Evol. 123: 153. 1975 [basion.] Cynanchica paui (Font Quer) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula paui Font Quer, Butl. Inst. Catalana Hist. Nat. 20: 188. 1920 [basion.] Cynanchica paui subsp. dianensis (Font Quer) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula paui var. dianensis Font Quer, Butl. Inst. Catalana Hist. Nat. 26: 57. 1926 [basion.] ≡ A. paui subsp. dianensis (Font Quer) De la Torre, Alcaraz & M.B.Crespo, Lazaroa 16 (1995): 154. 1996 Cynanchica paui subsp. paui Cynanchica pedicellata (Klokov) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pedicellata Klokov in Komarov, Fl. URSS 23: 703. 1958 [basion.] Cynanchica peloritana (C.Brullo, Brullo, Giusso & Scuderi) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula peloritana C.Brullo, Brullo, Giusso & Scuderi, Anales Jard. Bot. Madrid 66: 86. 2009 [basion.] Cynanchica pestalozzae (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pestalozzae Boiss., Diagn. Pl. Orient., ser. 2, 2: 112. 1856 [basion.] t Cynanchica pinifolia (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pontica Boiss., Fl. Orient. 3: 39. 1875 [basion.] us Cynanchica pontica (Boiss.) P.Caputo & Del Guacchio, comb. nov. cr ip ≡ Asperula lutea var. pinifolia Boiss., Fl. Orient. Suppl.: 280. 1888 [basion.] ≡ A. pinifolia (Boiss.) Ehrend. & Schönb.-Tem. in Strid & Kit Tan (eds.), Mount. Fl. Greece 2: 293. 1991 an Cynanchica tenuifolia (Boiss.) P.Caputo & Del Guacchio, comb. nov. M ≡ A. tenuifolia Boiss., Diagn. Pl. Orient., ser. 1, 3: 32. 1843 [basion.] = Asperula stricta subsp. pruinosa Ehrend., Notes Roy. Bot. Gard. Edinburgh 22: 337. 1958 Ac ce pt e d Note: As Fourreau’s (1868) “C. tenuifolia” is not validly published, that by Boissier can serve as basionym. In addition, there is some confusion about the correct synonymy of A. stricta subsp. pruinosa. Ehrendorfer and Krendl (1976) included it under A. ophiolitica, endemic to Evvoia (Greece), followed by Marhold (2011). However, already Ehrendorfer & SchönbeckTemesy (1975) stated, in the protologue of A. ophiolitica, that A. stricta subsp. pruinosa should be included in the western Anatolian A. tenuifolia, A. ophiolitica being a different species. This latter treatment was later restated by the same authors (Ehrendofer and Schönbeck-Temesy 1982). On one hand, Govaerts (2019) reported A. stricta subsp. pruinosa among the synonyms of A. stricta subsp. stricta; on the other hand, he listed the same subspecies under A. tenuifolia, as “Asperula scabra subsp. pruinosa Ehrend., Notes Roy. Bot. Gard. Edinburgh 22: 337 (1958)” (sic!). Cynanchica puberula (Halácsy & Sint.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula puberula Halácsy & Sint., Oesterr. Bot. Z. 40: 38. 1890 [basion.] Cynanchica pulvinaris (Heldr. ex Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pulvinaris Heldr. ex Boiss., Fl. Orient. Suppl.: 281. 1888 [basion.] Cynanchica pumila (Moris) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pumila Moris, Mem. Reale Accad. Sci. Torino 38: 26. 1835 [basion.] Note: “Asperula neglecta Moris” and “Asperula scabra Moris” are not illegitimate names (cf. Govaerts 2019), but simply do not exist. In fact, Moris (1828: 4; 1829) misapplied the names by Presl and Gussone to the Sardinian plant that he later recognized as a distinct species. Cynanchica pyrenaica (L.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pyrenaica L., Sp. Pl.: 104. 1753 [basion.] ≡ A. cynanchica subsp. pyrenaica (L.) Nyman, Consp. Fl. Eur.: 333. 1879 ip t Note: As noted above, the use of the epithet “cynanchica” under this genus would cause a tautonym. If A. cynanchica L., A. pyrenaica L., A. neglecta Guss. and A. occidentalis Rouy (and obviously their synonyms) are included in a single species, as modern authors mostly do (see below), the name wirh priority within Cynanchica is the Linnaean A. pyrenaica (Art. 11.4). Cynanchica pyrenaica subsp. cynanchica (L.) P.Caputo & Del Guacchio, comb. nov. us cr ≡ Asperula cynanchica L., Sp. Pl.: 104. 1753 [basion.] ≡ Galium cynanchicum (L.) Scop., Fl. Carniol., ed. 2, 1: 101. 1771 an Note: The name A. cynanchica subsp. cynanchica claims priority from 1879, when Nyman, publishing A. cynanchica subsp. pyrenaica (L.) Nyman, automatically establishing the autonym A. cynanchica subsp. cynanchica (Art. 26.3). M = Asperula rubeola Lam., Fl. Franç. 3: 375. 1779, nom. illeg. d Note: Lamarck (1779) incorrectly included in A. rubeola three Linnaean species published earlier, so making his name illegitimate (Art. 52.2). The reference in Govaerts (2019) must be amended. ce pt e = A. saxatilis Lam., Tabl. Encycl. 1: 139. 1791? Ac Note: This name is surprisingly regarded as a synonym of Cynanchica nitida by Govaerts (2019, sub A. nitida), in which case it would claim priority, while Ortega Oliviencia and Devesa (2007) reported it among the synonyms of A. cynanchica subsp. cynanchica var. cynanchica. Actually, the protologue described a mountain plant of France, citing as a possible synonym the preceeding A. pyrenaica (Lamarck 1791, “An Asperula Pyrenaica Lin.”). = A. collina Salisb., Prodr. Stirp. Chap. Allerton: 58. 1796, nom. illeg. (Art. 52.2) = A. multiflora Lapeyr., Hist. Pl. Pyrénées: 62. 1813 = A. alpina M.Bieb., Fl. Taur.-Caucas. Suppl.: 103. 1819 = A. minor Gray, Nat. Arr. Brit. Pl. 2: 481. 1821, nom. Illeg. (Art. 52.2) = A. rubeola Gratel., Bull. Soc. Nat. Hist. Bord. 1: 311. 1826 = A. tenuiflora Jord., Mém. Acad. Natl. Sci. Lyon, Sect. Sci., sér. 2, 1: 288. 1851 = A. macroclada A.Huet, Ann. Sci. Nat., Bot., sér. 3, 19: 253. 1853 = A. arenicola Reut., Cat. Pl. Vasc. Genève, ed. 2: 101. 1861 = A. cynanchica var. capillacea Lange in Willkomm & Lange, Prodr. Fl. Hispan. 2: 303. 1868 ≡ A. cynanchica subsp. capillacea (Lange) Rouy, Ill. Pl. Eur. 17: 133. 1902 ≡ A. capillacea (Lange) R.Vilm. ex Kerguélen, in Lejeunia, n.s. 120: 53. 1987 [n. v.] Note: The combination at the subspecific rank was first published by Rouy (1902) in the place reported above; while the name “ASPERULA CAPILLACEA” appears below the plate n. 404 of the same work (cf. Kerguélen 1999; Ortega Oliviencia and Devesa 2007; Govaerts 2019). However, in the Illustrationes (Rouy 1902), the rank accepted by Rouy appears only in the note to each binomial (es. “NEPETA MALLOPHORA”, p. 137) (cf. Art. 36.3). = A. heteroclada Hausskn., Mitth. Thüring. Bot. Vereins, n.f. 6: 31. 1894 cr = A. trabutii Sennen, Butl. Inst. Catalana Hist. Nat. 32: 98. 1932 ip t = A. bazargiciensis Prodan, Fl. Român. 1: 967. 1923 [n. v.] us = A. semiamicta Klokov in Komarov, Fl. URSS 23: 694. 1958 – “Asperula subalpina Schur”, Enum. Pl. Transsilv.: 277. 1866 (Art. 36.1) (cf. Govaerts 2019) an – “Cynanchica tenuifolia Fourr.”, Ann. Soc. Linn. Lyon n.s. 16: 398. 1868, nom inval. (Art. 35.1) M – “Asperula kerneri Procupiu ex O.Deg.”, Fl. Veleb. 3: 65. 1938, nom. inval. (Art. 36.1) [n. v.] – “Galium cynanchicum (L.) F.Herm.”, Fl. N. Mitteleur.: 942. 1956, comb. inval. (see Art. 6.3, Note 2, Art. 41.5) ce pt e d Cynanchica pyrenaica subsp. neglecta (Guss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula neglecta Guss., Pl. Rar.: 69. 1826 [basion.] ≡ A. cynanchica subsp. neglecta (Guss.) Arcang., Comp. Fl. Ital.: 308. 1882 Ac Note: For the treatment of this taxon we follow Del Guacchio and Caputo (2005). According to these authors, Asperula nitens Guss. is an intermediate form between C. pyrenaica subsp. neglecta and C. cynanchica subsp. cynanchica. Cynanchica pyrenaica subsp. occidentalis (Rouy) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula occidentalis Rouy in Rouy & Foucaud, Fl. France 8: 60. 1903 [pro hybr.] [basion.] ≡ Asperugalium × occidentale (Rouy) P.Fourn., Quatre Fl. France: 883. 1938 ≡ Asperula cynanchica subsp. occidentalis (Rouy) Stace, Watsonia 17: 444. 1989 Cynanchica pyrenaica subsp. pyrenaica Cynanchica rigidula (Halácsy) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula lutea var. rigidula Halácsy, Oesterr. Bot. Z. 45: 382. 1895 [basion.] (the name first occurs at page 339) ≡ A. rigidula (Halácsy) Halácsy, Consp. Fl. Graec. 1: 738. 1900. Cynanchica rupicola (Jord.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula rupicola Jord., Mém. Acad. Sci. Natl. Lyon, Sect. Sci. 1: 287. 1851 [basion.] Cynanchica rumelica (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula rumelica Boiss., Diagn. Pl. Orient., ser. 2, 2: 113. 1856 [basion.] Note: Asperula montana Waldst. & Kit. ex Willd. (described from Hungary) is usually regarded as a possible synonym of A. rumelica (e.g., Ehrendorfer & Krendl 1976, Govaerts 2019), as a tetraploid race of A. cynanchica (Barksay 1957), and even as a good species (Marhold 2011). However, Willdenow (1809) reported it as very similar to A. tinctoria (but bearing the lower leaves in whorl of six elements). We did not examine original material, but the latter hypothesis, in the light of the results of the molecular investigation by Gargiulo et al. (2015) seems unlikely; however, the correct interpretation of the name requires further study. ip t = A. divergens Boiss., Diagn. Pl. Orient., ser. 2, 6: 90. 1859 cr = A. graniticola Klokov in Komarov, Fl. URSS 23: 699. 1958 = A. hypanica Klokov in Komarov, Fl. URSS 23: 698. 1958 us Note: Accepted by Marhold (2011). an Note: Regarded as a synonym of A. tenella by Elenevsky and Pjatunina (1995); accepted as a valid species by Marhold (2011). M incl. A. barthae Pénzes, Ann. Mus. Hist. Nat. Hung. Bot. 31: 113. 1937 d Note: Ehrendorfer and Krendl (1976) report this name as a possible synonym of A. rumelica; while, according to Marhold (2011), it would be a distinct species. ce pt e incl. A. attenuata Klokov in Komarov, Fl. URSS 23: 698. 1958 Note: Accepted as a distinct species by Pobedimova (1978) and Czerepanov (1995), but reported as a synonym of A. tenella by Elenevsky and Pjatunina (1995) and of A. rumelica by Govaerts (2019). According to Ehrendorfer and Krendl (1976), it would represent an intermediate between A. tenella and A. rumelica. Ac Cynanchica samia (D.H.Christ & Goeriadis) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula samia D.H.Christ & Goeriadis, Willdenowia 13(2): 341. 1984 [basion.] Cynanchica setulosa (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ A. setulosa Boiss., Diagn. Pl. Orient., ser. 1, 10: 61. 1849 [basion.] Note: The taxonomic value of this species requires further study. According to some authors (Elenevsky and Pjatunina 1995; Govaerts 2019), it would be a synonym of A. graveolens subsp. graveolens. Klokov (1958) reported intermediate populations between A. graveolens and A. setulosa. However, this latter taxon is mostly regarded as more similar to A. littoralis (Ehrendorfer and Krendl 1976; Pobedimova 1978; Czerepanov 1995), from which, regardless, it seems sufficiently distinct. Cynanchica sintenisii (Asch. ex Bornm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula sintenisii Asch. ex Bornm., Repert. Spec. Nov. Regni Veg. Beih. 89: 299. 1941 [basion.] Note: The presumed name “Asperula sintenisii Halácsy” reported by Govaerts (2019) does not exist, as Halácsy (1890, p. 36) reported it only as a misapplied name, employed in herbarium and indeed never published. Cynanchica staliana (Vis.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula staliana Vis., Fl. Dalmat. 3: 11. 1850 [basion.] t Cynanchica staliana subsp. arenaria (Korica) P.Caputo & Del Guacchio, comb. nov. ip ≡ Asperula staliana subsp. arenaria Korica, Bot. Jahrb. Syst. 102: 347. 1981 [basion.] us cr Cynanchica staliana subsp. diomedea (Korica, Lausi & Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula staliana subsp. diomedea Korica, Lausi & Ehrend., Fl. Medit. 2: 68. 1992 [basion.] an Cynanchica staliana subsp. issaea (Korica) P.Caputo & Del Guacchio, comb. nov. M ≡ Asperula staliana subsp. issaea Korica, Bot. Jahrb. Syst. 102: 349. 1981 [basion.] Cynanchica staliana subsp. staliana d Cynanchica stricta (Boiss.) P.Caputo & Del Guacchio, comb. nov. ce pt e ≡ Asperula stricta Boiss., Diagn. Pl. Orient., ser. 1, 3: 33. 1843 [basion.] Cynanchica stricta subsp. elmaliensis (Schönb.-Tem) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula stricta subsp. elmaliensis Schönb.-Tem., Notes Roy. Bot. Gard. Edinburgh 37: 246. 1979 [basion.] Ac Cynanchica stricta subsp. grandiflora (Schönb.-Tem.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula stricta subsp. grandiflora Schönb.-Tem., Notes Roy. Bot. Gard. Edinburgh 37: 246. 1979 [basion.] Cynanchica stricta subsp. latibracteata (Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula stricta var. latibracteata Boiss., Fl. Orient. 3: 35. 1875 [basion.] ≡ A. stricta subsp. latibracteata (Boiss.) Ehrend., Notes Roy. Bot. Gard. Edinburgh 22. 337. 1958 = A. cappadocica Hausskn. ex Bornm., Repert. Spec. Nov. Regni Veg. Beih. 89: 296. 1941 = A. macrantha Hausskn. ex Bornm., Repert. Spec. Nov. Regni Veg. Beih. 89: 296. 1941 Cynanchica stricta subsp. monticola (Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula stricta subsp. monticola Ehrend., Notes Roy. Bot. Gard. Edinburgh 37: 246. 1979 [basion.] Cynanchica stricta subsp. stricta = Asperula fasciculata Boiss., Diagn. Pl. Orient., ser .1, 10: 60. 1849 Cynanchica suberosa (Sm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula suberosa Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 89. 1806 [basion.] = A. athoa Boiss., Diagn. Pl. Orient., ser. 1, 10: 62. 1849 ip t = A. pirinica Stoj. & Acht., Izv. Tsarsk. Prir. Inst. Sofiya 12: 185. 1939 Cynanchica suffruticosa (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. us Cynanchica supina (M.Bieb.) P.Caputo & Del Guacchio, comb. nov. cr ≡ Asperula suffruticosa Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., ser. 2 2: 111. 1856 [basion.] an ≡ Asperula supina M.Bieb., Fl. Taur.-Caucas. 1: 101. 1808 [basion.] = A. praevestita Klokov in Komarov, Fl. URSS 23: 701. 1958 M Note: Accepted by Pobedimova (1978) and Czerepanov (1995). = A. kotovii Klokov, Fl. RSS Ucr. 10: 457. 1961 ce pt e d Note: Accepted by Pobedimova (1978) and Czerepanov (1995). Cynanchica supina subsp. caespitans (Juz.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula caespitans Juz., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 14: 35. 1951 [basion.] [n.v.] ≡ A. supina subsp. caespitans (Juz.) Pjatunina, Bull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 100(4): 75. 1995 Ac = A. aemulans V.I.Krecz. ex Klokov in Komarov, Fl. URSS 23: 700. 1958 = A. tranzshelii Klokov in Komarov, Fl. URSS 23: 697. 1958 Note: This subspecies is not accepted by Marhold (2011). Cynanchica supina subsp. supina = Asperula vestita V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 25. 1934 Note: Accepted by Klokov (1958). According to Marhold (2011), it would be a synonym of A. cretacea. = A. cimmerica V.I.Krecz. ex Klokov in Komarov, Fl. URSS 23: 702. 1958 Note: It was accepted as a distinct species by Pobedimova (1978) and Czerepanov (1995). = A. praepilosa V.I.Krecz. ex Klokov in Komarov, Fl. URSS 23: 702. 1958 Note: Accepted by Pobedimova (1978) and Czerepanov (1995). Cynanchica tenella (Heuff. ex Degen) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula tenella Heuff. ex Degen, Oesterr. Bot. Z. 47: 196. 1897 [basion.] = A. stevenii V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 25. 1934 Note: Accepted by Klokov (1958) and Pobedimova (1978). ip t = A. maeotica Popov & Chrshan., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 50(5-6): 97. 1946 cr = A. bidentata Klokov in Komarov, Fl. URSS 23: 697. 1958 us – “Galium danubiale F.Herm.”, Fl. N. Mitteleur.: 941. 1956, nom. inval. an Note: Probably an intended replacing name for A. tenella (due to the existence of Galium tenellum Jord.), but not validly published (Art. 41.5). M Cynanchica tephrocarpa (Popov & Chrshan.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula tephrocarpa Czern. ex Popov & Chrshan., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 50(5-6): 96. 1946 [basion.] ce pt e d – “Asperula exasperata subsp. tephrocarpa (Czern. ex Popov & Chrshan.) Pjatunina”, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 100(4): 72. 1995, comb. inval. Ac Note: Asperula exasperata and A. petraea are regarded as distinct species by most authors (e.g., Ehrendorfer and Krendl 1976; Pobedimova 1978; Czerepanov 1995; Marhold 2011). According to us, at the present state of knowledge, a more parsimonious treatment is to join A. tephrocarpa, A. exasperata and A. petraea in a single taxon within 3 subspecies. This treatment was first proposed by Elenevsky and Pjatunina (1995), who, however, did not subordinate A. tephrocarpa under the later A. exasperata (Artt. 11.4, 25.1). Cynanchica tephrocarpa subsp. exasperata (Klokov) P.Caputo & Del Guacchio, comb. nov. Note: The combination that we propose here under Cynanchica is not a status novus, because the previous A. exasperata subsp. exasperata (Elenevsky and Pjatunina 1995) is valid. ≡ A. exasperata V.I.Krecz. ex Klokov in Komarov, Fl. URSS 23: 695. 1958 Cynanchica tephrocarpa subsp. petraea (V.I.Krecz. ex Klokov) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula petraea V.I.Krecz. ex Klokov in Komarov, Fl. URSS 23: 696. 1958 [basion.] ≡ A. exasperata subsp. petraea (Klokov) Pjatunina, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 100(4): 73. 1995 Cynanchica tephrocarpa subsp. tephrocarpa = A. creticola Klokov in Komarov, Fl. URSS 23: 694. 1958 Cynanchica tragacanthoides (Brullo) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula tragacanthoides Brullo, Bot. Not. 132: 291. 1979 [basion.] Cynanchica visianii (Korica) P.Caputo & Del Guacchio, comb. nov. ip Cynanchica wettsteinii (Adamovic) P.Caputo & Del Guacchio, comb. nov. t ≡ Asperula visianii Korica, Pl. Syst. Evol. 133: 72. 1979 [basion.] cr ≡ Asperula wettsteinii Adamovic, Deutsche Bot. Monatsschr. 7: 117. 1889 [basion.] us Cynanchica woloszczakii (Korica) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula woloszczakii Korica, Phyton (Horn) 17: 143. 1975 [basion.] an Cynanchica woronowii (V.I.Krecz.) P.Caputo & Del Guacchio, comb. nov. M ≡ Asperula woronowii V.I.Krecz. in Grossheim, Fl. Kavkaza 4: 25. 1934 [basion.] Note: In Russian language, but see art. 39.1. d Thliphthisa (Griseb.) P.Caputo & Del Guacchio, stat. nov. ce pt e ≡ Galium sect. Thliphthisa Griseb., Spic. Fl. Rumel. 2: 161. 1846 [Jan, “1844”] [basion.] ≡ Asperula sect. Thliphthisa (Griseb.) Ehrend., in J. Linn. Soc. Bot. 70: 3. 1975. Type: Galium rupestre Vis., Flora 12(1 Erg.): 5. 1829 [≡ Thliphtisa rupestris (Vis.) P.Caputo & Del Guacchio], designated by Ehrendorfer (1975: 3) Ac = Galium [unranked] Erythrogalia DC., Prodr. 4: 603. 1830. Type: Galium purpureum L., Sp. Pl.: 107. 1753 designated by Ehrendorfer (1975: 3) Description: Herbaceous or suffruticose perennial plants, with woody taproot, at least some whorls with 6-8 elliptic to linear elements, 1-veined, bracts and bracteoles, partial inflorescences leafy, never capitate, with rotate to infundibuliform corollas, with 4 lobes, calyx teeth inconspicuous, stamens inserted by rule and subequal to the oblong or globose stigma, fruits truncate, tuberculate or smooth, but never spiny. Etym: From ancient Greek ϑλιϕϑείσα (from the verb ϑλίβω); it may mean either “compressed”, “flattened” (alluding to the truncated fruit), or, with reference to the protologue, “squeezed”, “rubbing” (alluding to the root, buried in the rocks). Distribution: The genus is centered in the eastern Mediterranean, where is mostly represented by narrow endemics (Minareci and Yıldız 2010). Notes: The best known representative of the genus, i.e. Thliphthisa purpurea, was initially described under Galium, on account of its rotate corolla. Ehrendorfer (1958) first hypothesized that Galium purpureum was to be united to similar Asperula species with long tube in a distinct genus, and this is fully confirmed by recent molecular studies. The latter, however, do not support an intermediate position of Thliphthisa between Galium and Asperula (Ehrendorfer 1958). Clearly, corolla shape, which induced the first scholars to include these species time by time in Galium or in Asperula, changed more than once during evolution of Rubieae and therefore does not appear to have great phylogenetic value (e.g., Soza and Olmstead 2010a). Thliphthisa antalyensis (Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula antalyensis Ehrend., Notes Roy. Bot. Gard. Edinburgh 22: 352. 1958 [basion.] ≡ A. pseudochlorantha var. antalyensis (Ehrend.) Minareci & K.Yildiz, Ann. Bot. Fenn. 47: 122. 2010 us Thliphthisa apuana (Fiori) P.Caputo & Del Guacchio, comb. nov. cr ip t Note: The occurrence in the same places of two taxa so similar, but well distinct on account of few characters (length and colour of the corolla) without any reported intermediate morph (Minareci and Yildiz 2010), is difficult to explain, in our opinion, with a treatment at an infraspecific rank. an ≡ Galium purpureum var. apuanum Fiori, Nuov. Fl. Italia 2: 491. 1927 [basion.] ≡ Asperula purpurea subsp. apuana (Fiori) Bechi & Garbari, Fl. Medit. 4: 223. 1994 ≡ A. apuana (Fiori) Arrigoni, in Webbia 67: 42. 2012 M Thliphthisa baenitzii (Heldr. ex Boiss.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula baenitzii Heldr. ex Boiss., Fl. Orient. Suppl.: 280. 1888 [basion.] ce pt e d Thliphthisa baldaccii (Halácsy) P.Caputo & Del Guacchio, comb. nov. ≡ Galium baldaccii Halácsy, in Oesterr. Bot. Z. 40: 165. 1890 [basion.] ≡ Asperula baldaccii (Halácsy) Ehrend., Bot. J. Linn. Soc. 68: 269. 1974. Thliphthisa boryana (Walp.) P.Caputo & Del Guacchio, comb. nov. Ac ≡ Galium boryanum Walp., in Repert. Bot. Syst. 2: 454. 1843 [basion.] ≡ Asperula boryana (Walp.) Ehrend., Bot. J. Linn. Soc. 68: 269. 1974 = Sherardia pusilla Bory & Chaub., Nouv. Fl. Pélop.: 10. 1838 Thliphthisa brevifolia (Vent.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula brevifolia Vent., Descr. Pl. Nouv.: t. 63. 1802 [basion] – “Asperula brunnea Boiss.”, Fl. Orient. 3: 32. 1875, nom. inval. [Art. 36.1(b)] Thliphthisa chlorantha (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula chlorantha Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., ser. 2, 6: 90. 1859 [basion.] Thliphthisa crassula (Greuter & Zaffran) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula crassula Greuter & Zaffran, Willdenowia 14: 289. 1985 [“1984”] [basion.] Thliphthisa elonea (Iatroú & Goergiadis) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula elonea Iatroú & Goergiadis, Willdenowia 14: 55. 1984 [basion.] Thliphthisa gorganica (Schönb.-Tem. & Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula gorganica Schönb.-Tem. & Ehrend., Bot. Jahrb. Syst. 10: 80. 1985 [basion.] Thliphthisa mazanderanica (Ehrend.) P.Caputo & Del Guacchio, comb. nov. cr Thliphthisa microphylla (Boiss.) P.Caputo & Del Guacchio, comb. nov. ip t ≡ Asperula mazanderanica Ehrend., Ann. Naturhist. Hofmus. 56: 225. 1948 [basion.] us ≡ Asperula microphylla Boiss., Diagn. Pl. Orient., ser .1, 3: 34. 1843 [basion.] Thliphthisa muscosa (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. an ≡ Asperula muscosa Boiss. & Heldr. in Boiss., Diagn. Pl. Orient., ser. 2, 2: 109. 1856 [basion.] M Thliphthisa pseudochlorantha (Ehrend.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula pseudochlorantha Ehrend., Notes Roy. Bot. Gard. Edinburgh 22: 351. 1958 [basion.] d Thliphthisa purpurea (L.) P.Caputo & Del Guacchio, comb. nov. ce pt e ≡ Galium purpureum L., Sp. Pl.: 107. 1753 [basion.] ≡ Asperula purpurea (L.) Ehrend., Oesterr. Bot. Z. 122: 260. 1973 Note: Govaerts (2019) reports also “Crucianella purpurea (L.) Honck.” among the synonyms, but the identification of the annual plant described by Honckeny (1782, p. 431) seems to indicate Galium spurium L. or an affine species. Ac Thliphthisa purpurea subsp. apiculata (Sm.) P.Caputo & Del Guacchio, comb. nov. ≡ Galium apiculatum Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 91. 1806 [basion.] ≡ Asperula purpurea subsp. apiculata (Sm.) Ehrend., Bot. J. Linn. Soc. 68: 269. 1974 = Galium purpureum var. luteoviride Rohlena, Sitzungsber. Königl. Böhm. Ges. Wiss., Math.Naturwiss. Cl. 1912: 54. 1913 Thliphthisa purpurea subsp. purpurea Thliphthisa rigida (Sm.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula rigida Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 89. 1806 [basion.] = A. platygona Gand., Bull. Soc. Bot. France 62: 154. 1916 [“1915”] = A. kritsensis Coustur. & Gand., Bull. Soc. Bot. France 63: 9, 14. 1917 [“1916”] Thliphthisa rupestris (Vis.) P.Caputo & Del Guacchio, comb. nov. ≡ Galium rupestre Vis., Flora 12(1 Erg.): 5. 1829 [basion.] ≡ Asperula rupestris (Vis.) Reich., Fl. Germ. Excurs.: 205. 1831, nom. illeg. ≡ Asperula scutellaris Vis., Sem. Rar. Hort. Patav. Coll. MDCCCXXXVI: 8. 1836 ip t Note: The name Galium rupestre is not a later homonym of Galium rupestre Candolle (1830), as Visiani (1836) himself believed; nevertheless, it cannot be employed in Asperula because of the existence, since 1827, of Asperula rupestris Tineo, so that the combination A. rupestris (Vis.) Rchb. (Reichenbach 1831) is illegitimate and the correct name under Asperula is A. scutellaris. ≡ Asperula saxicola Ehrend., Bot. J. Linn. Soc. 68: 269. 1974 [basion.] cr Thliphthisa saxicola (Ehrend.) P.Caputo & Del Guacchio, comb. nov. us Thliphthisa semanensis (Schönb.-Tem. & Ehrend.) P.Caputo & Del Guacchio, comb. nov. an ≡ Asperula semanensis Schönb.-Tem. & Ehrend., Bot. Jahrb. Syst. 107: 81. 1985 [basion.] Thliphthisa serotina (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. M ≡ Galium serotinum Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., ser. 1, 10: 71. 1849 [basion.] ≡ Asperula serotina (Boiss. & Heldr.) Ehrend. in Davis (Ed.), Fl. Turkey 7: 766. 1982 d Thliphthisa suberosa (Sm.) P.Caputo & Del Guacchio, comb. nov. ce pt e ≡ Asperula cypria Ehrend., Pl. Syst. Evol. 126: 406. 1977, nom. nov. pro Galium suberosum Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 91. 1806 [basion.] Thliphthisa tournefortii (Sieber ex Spreng.) P.Caputo & Del Guacchio, comb. nov. Ac ≡ Asperula tournefortii Sieber ex Spreng., Syst. Veg. 1: 395. 1824 [basion.] ≡ Crucianella tournefortii (Sieber ex Spreng.) Tausch, Flora 18: 359. 1835 = Asperula majori Barbey, Bull. Herb. Boissier 2: 243. 1894 Hexaphylla (Klokov) P.Caputo & Del Guacchio, stat. nov. ≡ Asperula sect. Hexaphylla Klokov, Fl. RSS Ucr. 10: 458. 1961 [basion.]. Type: Asperula hexaphylla All. [≡ Hexaphylla allionii P.Caputo & Del Guacchio], automatically determined according to Art. 10.8 (cf. Klokov 1961: 458) = Asperula sect. Cynanchicae subsect. Capitatae Degen, in Oesterr. Bot. Z. 40: 14. 1890 – “Asperula sect. Hexaphylla Ehrend.”, Bot. J. Linn. Soc. 70: 2. 1975, isonym (Art. 6.3, Note 2) Description: Perennial herbs, sometimes stoloniferous or suffruticose, with leafy elements typically in whorls of 6, broadly lanceolate to linear, 1-veined, without a distinct hyaline apex; inflorescence pyramidal to corymbiform with partial inflorescences subcapitate and subtended by bracts; flowers 4-merous, with hypocrateriform to infundibuliform corolla, anthers and stigma mainly exserted and ovoid ovary and fruit. Etym: The name from Greek means “with six leaves” and alludes to one of the relevant distinctive features of the genus. Distribution: From the Mediterranean Basin to W-Asia (desumed from Govaerts 2019). ip t Notes: The subsectional name “Capitatae” (= with capitate inflorescence), if raised to generic rank as “Capitata” could be regarded as a technical Latin term, and therefore not validly published (Art. 20.2). Degen (1890) described this group at pag. 14 as “Capitatae”, but only in the table at pag. 16 he clarified the intended rank (“subsect. Capitae” [sic!]). Klovok (1961) reported “Sect. Hexaphyllae” (not “Hexaphylla”) already at p. 164, but with a description in Ukrainian. Hexaphylla allionii P.Caputo & Del Guacchio, nom. nov. us cr ≡ A. hexaphylla All., Fl. Pedem. 1: 12. 1785 ≡ A. allionii Baumg., Enum. Stirp. Transsilv. 1: 80. 1816, nom. illeg. (see the note immediately below). ce pt e d M an Note: The prior name Asperula hexaphylla cannot be employed as basionym, because this would cause a tautonym (Art. 23.4). The final epithet of the next earliest legitimate name at the same rank must be used (Art. 11.4). The plant described by Baumgarten (1816) as A. allionii is different from the authentic A. hexaphylla, which is endemic to France and Italy. Probably, Baumgarten described the plant until now called A. capitata (Degen 1890). Neverthless, as Baumgarten (1816) cited A. hexaphylla in synonymy, A. allionii is a superfluous name and then illegitimate (Art. 52), with the same type of A. hexaphylla. The legitimate A. umbellata Willd. ex DC. is not a synonym of A. hexaphylla, as reported by Govaerts (2019). Candolle (1830) listed this species among those not sufficiently known. The original material (B-W!) is to be ascribed to at least two different species. Indeed, the locus classicus (Hungary) and the diagnosis by Candolle (1830) (“it differs from A. hexaphylla for the much shorter flowers”) seem to be conclusive in rejecting such a synonymy (cf. Hexaphylla capitata). Therefore, we decided to re-use the epithet of the illegitimate name A. allionii at the same rank, without excluding the type of A. hexaphylla, because the epithet hexaphylla is not available for use in Hexaphylla (Art. 58.1, Note 1, Ex. 7). The epithet allionii pays tribute to the first to describe the species. Ac Hexaphylla arcadiensis (Sims) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula arcadiensis Sims, Bot. Mag. 47: n. 2146. 1820 [basion.] Note: The presumed synonym A. mollis Boissier (1843: 31) (Govaerts 2019) deserves further study. Hexaphylla capitata (Kit. ex Schult.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula capitata Kit. ex Schult., Oestr. Fl., ed. 2, 1: 312. 1814 [basion.] ≡ A. hexaphylla subsp. capitata (Kit. ex Schult.) Nyman, Consp. Fl. Eur.: 333. 1879 – “Galium capitatum (Kit. ex Schult.) F.Herm.”, Fl. N. Mitteleur.: 941. 1956, comb. inval. (Art. 41.5) Hexaphylla cretacea (Willd. ex Roem. & Schult.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula cretacea Willd. ex Roem. & Schult., Syst. Veg., ed. 15 bis 3: 529. 1818 [basion.] = A. infracta Klokov in Kom., Fl. URSS 23: 701. 1958 = A. tauroscythica Klokov, Fl. RSS Ucr. 10: 458. 1961 (“tauro-scythica”, Art. 60.11) Note: Ehrendorfer and Krendl (1976), presumably misapplying the name, refer this taxon to sect. Cynanchicae; however, it belongs without doubt to sect. Hexaphylla (e.g., Klokov 1958; Elenevsky and Pjatunina 1995). Hexaphylla cyrenaica (E.A.Durand & Barratte) P.Caputo & Del Guacchio, comb. nov. ip t ≡ Asperula hirsuta var. cyrenaica E.A.Durand & Barratte, Fl. Libyc. Prodr.: 116. 1910 [basion.] ≡ A. cyrenaica (E.A.Durand & Barratte) Pamp., Arch. Bot. (Forlì) 12: 44. 1936 Hexaphylla doerfleri (Wettst.) P.Caputo & Del Guacchio, comb. nov. cr ≡ Asperula doerfleri Wettst., Biblioth. Bot. 5(26): 59. 1892 (“Dörfleri”) [basion.] us Hexaphylla hercegovina (Degen) P.Caputo & Del Guacchio, comb. nov. an ≡ Asperula hercegovina Degen, Oesterr. Bot. Z. 40: 15. 1890 [basion.] ≡ A. capitata subsp. hercegovina (Degen) Hayek, Repert. Spec. Nov. Regni Veg. 30(2): 445 M = A. hexaphylla var. pilosa Beck, Ann. K. K. Naturhist. Hofmus. 2: 153. 1887 ≡ A. pilosa (Beck) Degen, Oesterr. Bot. Z. 40: 15. 1890 ce pt e d Note: Listed as “Asperula pilosa Degen” by Govaerts (2019), but Degen (1890) clearly reported the basionym by Beck. For the priority of the name A. hercegovina against A. pilosa, see Art. 11.5. Hexaphylla hirsuta (Desf.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula hirsuta Desf., Fl. Atlant. 1: 127. 1798 [basion.] = A. repens Brot., Fl. Lusit. 1: 152. 1804 Ac = A. algerica Pers., Syn. Pl. 1: 124. 1805, nom. illeg. (Art. 52.2) Note: The presumed synonym “Asperula denudata Vahl ex Willl. & Lange” (Govaerts 2019) has no nomenclatural status, because cited by Wilkomm and Lange (1868) as a mere synonym of A. hirsuta (Art. 36.1) occurring in herbaria. Hexaphylla hirta (Ramond) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula hirta Ramond, Bull. Sci. Soc. Philom. Paris 2: 131. 1800 [basion.] Hexaphylla pubescens (Willd.) P.Caputo & Del Guacchio, comb. nov. ≡ Crucianella pubescens Willd., Sp. Pl. 1: 602. 1798 [basion.] ≡ Asperula pubescens (Willd.) Ehrend. & Schönb.-Tem. in Strid & Kit Tan (eds.), Mount. Fl. Greece 2: 295. 1991 = A. incana Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 1: 88. 1806 Hexaphylla rupestris (Tineo) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula rupestris Tineo, Cat. Pl. Hort. Panorm. 276. 1827 [basion.] ≡ A. hirsuta subsp. rupestris (Tineo) Nyman, Consp. Fl. Eur.: 334. 1879 Hexaphylla taygetea (Boiss. & Heldr.) P.Caputo & Del Guacchio, comb. nov. ≡ Asperula taygetea Boiss. & Heldr. in Boissier, Diagn. Pl. Orient., ser. 1, 10: 60. 1849 [basion.] cr ≡ Rubeola Hill, Brit. Herb.: 396. 1756, nom. illeg., non Mill. (Art. 53.1) ip t Sherardia L., Sp. Pl. 1: 102. 1753 ≡ Asperula sect. Sherardia (L.) Baill., Hist. Pl. 4: 1873. Type: Sherardia arvensis L., designated by Britton & Brown (1918. 3: 266), affirmed (Art. 10.5 & Rec. 10A.2) by Hitchcock (in Sprague 1929: 123). For a lectotypification of the species name, see Natali in Jarvis et al. (1993, p. 88). us ≡ Dillenia Heist. ex Fabr., Enum., ed. 2: 57. 1763, nom. illeg., non L. (Art. 53.1) an ≡ Hexodontocarpus Dulac, Fl. Hautes-Pyrénées: 467. 1867, nom. illeg. (Art. 52.2) M Description: Annual plants, with leafy elements typically (4-)6 per whorl, obovate to lanceolate, mucronate, 1-veined, flowers in terminal capitula, subtended by an involucre of connate elements; calyx 4- to 6-toothed, persistent in fruit, corolla infundibuliform and 4-lobed, style filiform and bifid with capitate stigma, fruit dry and scabrid of 2 mericarps, each 1-seeded. ce pt e d Notes: The relevant autapomorphies of this taxon are convincing for retention of Sherardia as a separate genus. In the main clade including Asperula, Thliphtisa, and Hexaphylla, this is the only taxon with annual habit and a calyx with showy and persistent theeth. The genus is nowadays regarded as monotypic and western-Palaeotemperate, but largely naturalized elsewhere. Sherardia arvensis L., Sp. Pl. 1: 102. 1753 ≡ Asterophyllum scherardianum K.F.Schimp. & Spenn. in Spenner, Fl. Friburg. 3: 1077. 1829 Ac ≡ Hexodontocarpus arvensis (L.) Dulac, Fl. Hautes-Pyrénées: 467. 1867. Note: This combination is not illegitimate (Art. 55.1), even if Hexodontocarpus is illegitimate. ≡ Asperula sherardia Hallier in W.D.J.Koch, Syn. Deut. Schweiz. Fl.: 1199. 1893, nom. illeg. [n.v.] ≡ Galium sherardia (Hallier) E.H.L.Krause in Sturm, Deutschl. Fl. (Sturm), ed. 2, 12: 187. 1904. Note: As Krause cited as synonyms both S. arvensis and Asperula sherardia (“Asperula Sherardi”), we consider his name as a new combination based on A. sherardia Hallier. = S. arvensis var. maritima Griseb., Spic. Fl. Rumel. 2: 169. 1846 ≡ S. maritima (Griseb.) Borbás, Magyar Bot. Lapok 2: 302. 1903 ≡ S. arvensis subsp. maritima (Griseb.) Jáv., Magyar Fl.: 1035. 1925 = S. affinis Gand., Contr. Fl. Terr. Slav. Merid. 1: 14. 1883 = S. pantocsekii Gand., Contr. Fl. Terr. Slav. 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