Anacamptis × dafnii nothosubsp. solanoi Serra & López Esp., nothosubsp. nov. is a hybrid of A. collina (Banks & Sol. ex Russell) R.M.Bateman & al. (Orchis collina Banks & Sol. ex Russell, bason.) and A. papilionacea subsp. grandiflora (Boiss.) Kreutz (Orchis papilionacea var. grandiflora Boiss., bason.). Anacamptis collina is a species that occurs along the coasts of the Mediterranean reaching, along the Tigris and Euphrates rivers, the Persian Gulf and Azerbaijan. It has stems 10–50 cm tall with 3–20 flowers; lateral petals 5.8–10.7 mm long with one nerve; labellum 7.6–11.8 × 7.8–12.3 mm without macules, with scarce papillae; spur 4.6–7.8 × 3–4.7 mm, sac-shaped. This species exhibits little variability (Aedo 2005; Kretzschmar & al. 2007; Delforge 2016), except for a variety with yellowish flower, which must be referred as Orchis collina var. flavescens Soó, and some specimens with a somewhat laxer inflorescence recently described in Badajoz as Anacamptis collina f. laxi-spicata F.M.Vázquez (Vázquez Pardo 2009).

Anacamptis papilionacea (L.) R.M.Bateman & al. (Orchis papilionacea L., bason.) is a species that also inhabits the Mediterranean, reaching the east of Asia Minor and the Caspian Sea; in the Iberian Peninsula and northern Africa it reaches the Atlantic coast. It has stems 11–55 cm tall with 6–22 flowers; lateral petals 10–17.2 mm long with 3–4 nerves; labellum (9)13.7–21(26) × (7)14.3–23.6(27) mm, with pink stripes or macules, with abundant papillae; spur 8.7–13.5 × 1.4–2,5 mm, cylindrical. It shows a great variability, on which consensus has not yet been achieved. It is discussed whether it is a single taxon across the Mediterranean and the Near East (Aedo 2005) or it groups several entities (Baumann 1986; Baumann & al. 2007; Kretzschmar & al. 2007; Delforge 2016). A genetic analysis performed few years ago revealed a low variability, but it relied exclusively on material collected in the central Mediterranean, just where the typical subspecies is abundant (Arduino & al. 1995). We consider here the subspecific rank as the most appropriate for these taxa, because the morphological variations are associated to particular geographic areas (Kretzschmar & al. 2007), and because when two of these entities cohabit no transition blurring their traits is observed (Scopece & al. 2009).

Thus, there are various opinions about the number of taxa that A. papilionacea includes —four in Baumann (1986), five in Baumann & al. (2007), six in Kretzschmar & al. (2007), and up to eight in Delforge (2016)—, and also about their taxonomic rank, since Baumann (1986), Baumann & al. (2007), and Kretzschmar & al. (2007) consider them as subspecies, while Delforge (2016) considers them as varieties. These different views regarding the number of entities in the eastern Mediterranean, where the diversity of the group is greater.

However, there is agreement in separating at least four entities, one inhabiting the western Mediterranean, another one the central Mediterranean, and two more the eastern Mediterranean. Two of them have a lip 13–18(19.5) × (12.7)13–25 mm, wide, more or less rounded, flabellate [A. papilionacea subsp. grandiflora in the western Mediterranean, and A. papilionacea subsp. heroica (E.D.Clarke) Kreutz in the eastern Mediterranean] and two others with lip 8.5–13 × 7–13(14) mm, narrow, more or less cuneate [A. papilionacea subsp. papilionacea in the central Mediterranean and A. papilionacea subsp. schirwanica (Woronow) H.Kretzschmar & al. in the eastern Mediterranean]. This matches the criteria of Baumann (1986), which is the most conservative one. To these four taxa, we should add another subspecies of the eastern Mediterranean with small lip, A. papilionacea subsp. palaestina (H.Baumann & R.Lorenz) H.Kretzschmar & al., close to the «papilionacea-schirwanica group», but with obovoid lip with rounded apex, and with lines or points that do not appear in the last group.

We follow here the criterion of Bateman & al. (2003), including Orchis papilionacea and Orchis collina in Anacamptis Rich. as in several recent general studies (Akbaç 2012; Tison & al. 2014; Claessens & Kleynen 2016; GIROS 2016). The concept of the genus Orchis L. has changed over time; a detailed study of its evolution can be found in Kretzschmar & al. (2007). We apply here a cladistic monophyletic criterion to separate genera, so that Orchis papilionacea and Orchis collina are placed in Anacamptis, as is shown in molecular biology studies on the ITS region of ribosomal DNA (Bateman & al. 1997; Bateman & al., 2003), and in a more recent research that also includes the mitochondrial cox1 intron and the plastid rp116 intron (Inda & al. 2012) to nrITS. Likewise, the ornamentation of seeds supports this approach (Gamarra & al. 2012). After the reorganization of Orchis, other studies have separated out the genus even further. Thus, Tyteca & Klein (2008) created a new genus (Herorchis D.Tyteca & E.Klein), where they included the plants that we consider here in Anacamptis, whereas Delforge (2009) also considered different genera for Orchis s.l., although he left Anacamptis as a monospecific genus and recovered the genus Vermeulenia A.Löve & D.Löve (Löve & Löve 1972) for A. papilionacea and A. collina.

Regarding the described hybrids of A. collina and A. papilionacea s.l., the first of them, which was described from a Syrian specimen, was Orchis × dueluekae Hautz. (Hautzinger 1976: 52, 1978: 69); for this hybrid A. papilionacea s.l. should have been ascribed to A. papilionacea subsp. palaestina, since this subspecies lives in Syria, while A. papilionacea subsp. papilionacea does not (Baumann & Lorenz 2005; Kretzschmar & al. 2007: 148). Some years later, Luz & Schmidt (1981) revised the type —deposited in the herbarium W— of Orchis × dueluekae, and concluded that it was in fact Orchis collina. In their paper, they provided a monochrome image of the type, which actually has the appearance of Orchis collina: despite of the bad quality of the image, the spur and the labellum are typical for this taxon. Unfortunately, we have recently looked for that sheet in W without success —Ernst Vitek, pers. comm.—. In their description of the hybrid, Luz & Schmidt (1981) proposed Orchis collina and Orchis caspia Trautv. as parents, where the second taxon is not, apparently, a synonym of A. papilionacea subsp. palaestina, a subspecies not present in the area, but of A. papilionacea subsp. schirwanica. The type was collected in Galilee —Israel— and deposited in the herbarium STU. They included two images of the parents and the hybrid, where the intermediate traits of the Israeli specimen can indeed be observed. They described it as Orchis × dafnii W.Schmidt & Luz.

The other hybrid that has been described so far in the group comes from the Italian Peninsula and is a hybrid of the nominal subspecies of A. papilionacea and A. collina (Kohlmüller 1993). It was collected in Mount Gargano, where only the nominal subspecies is present (Kretzschmar & al. 2007: 148), so that there is no doubt about the parents. A former reference must be assigned to the same hybrid, although as O. × dueluekae, from Surbo —Lecce, NE Italy— (D’Emerico & al. 1989).

The finding of a group of deviant specimens of A. papilionacea by Pedro Solano, as well as a solitary individual a few kilometers away from those by Juan Monpeán later, gave us an indication to consider them as belonging to a hybrid between A. collina and A. papilionacea subsp. grandiflora, two early flowering species which cohabit in the area.

Positioning data have been taken with a GARMIN GPS device, using DATUM ETRS 89. Thirty-two morphological characters were considered, comprising sixteen qualitative characters, sixteen quantitative characters, and one phenological character. The morphological data of the parents are based on previously published works (Aedo 2005; Kretzschmar & al. 2007) and on data taken from alive individuals and herbarium specimens from Murcia (see table 1). The RAL colour palette (Ral Colours 2017) has been used to define the colours of bracts and flowers, in order to objectify a trait that can be relevant in orchids (Serra & Soler 2012). The separation between the previously described hybrids and the new one are summarized in the table 2. The nomenclature, synonymy and types of the involved taxa are summarized in the appendix 1. The scarce number of specimens of populations mentioned in Additional material studied prevent us to collect them. Thus, to support such records, photographs of specimens of the different populations have been provided in the appendix 2.

Anacamptis × dafnii nothosubsp. solanoi Serra & López Esp., nothosubsp. nov. [A. collina (Banks & Sol. ex Russell) R.M.Bateman, Pridgeon & M.W.Chase × A. papilionacea subsp. grandiflora (Boiss.) Kreutz]. Type: Spain, Murcia, Cartagena, Colada del Cedacero, 30SXG7465, 50 m a.s.l., ubi inter parentes, 20–II–2016, L. Serra, P. Solano, J. A. López Espinosa & A. Bort s.n. (holo-: VAL 232771!). Figs. 1, 2b, 3.

LISD: urn:lsid:ipni.org:names:77174191-1

It differs from A. collina by its bigger lateral petals and the presence of 3-4 nerves; connivent sepals; labellum wider, without macules or with very few, more intense colour; spur longer, less sac-shaped but flattened; papillae of labellum a little longer and more abundant. It differs from the second parent —A. papilionacea subsp. grandiflora— by its broadly lanceolate leaves, cylindrical inflorescence, elongated, never compact nor subglobous; clearly smaller lateral sepals; labellum somewhat smaller, purple, with hardly any lines or macules and flattened, sac-shaped spur; papillae of labellum a little smaller and more scarce.

Stems 17–29(48) cm, glabrous, with 2–5 basal leaves 4.8–6 × 1.6–2,4 cm, broadly lanceolate, in rosette, glabrous, without macules, smooth; top stem leaves 4, without macules, the upper ones similar to the bracts. Inflorescence 7.5–10.5(24) cm, cylindrical, with 10–15(19) flowers, sessile, opening from the base to the apex. Bract of the basal flower 24–30 × 5–10 mm, signal violet in colour —RAL 4008—, lanceolate, glabrous. Sepals free, glabrous, signal violet in colour —RAL 4008—, with 3 nerves, lanceolate, more or less connivent, the lateral sepals 10 × 3–4 mm, the central sepals 12–14 × 3–4 mm. Lateral petals 14–15 × 5–6 mm, with 3–4 nerves, lanceolate, glabrous, traffic purple in colour —RAL 4006—. Labellum 13–18 × 10–17 mm, more or less flat, and almost rounded, with margin whole or slightly sawed, traffic purple in colour —RAL 4006—, without macules or with very few in the throat, this one white. Spur 9–10 × 3 mm, slightly sac-shaped but flattened, arched down, light pink in colour —RAL 3015—, with a green or pink line in its ventral part. Gynostegium 3–4 mm. Ovary 12–14 mm, glabrous.

Anacamptis × dafnii nothosubsp. solanoi Serra & López Esp., nothosubsp. nov. differs from A. × dafnii nothosubsp. dafnii by its bigger bracts, fewer nerves in the sepals, lateral petals and labellum wider and bigger, and labellum with no macules or very few at the mouth of the throat. It differs from A. × dafnii nothosubsp. camparonensis (Kohlmüller) H.Kretzschmar & al. for its higher number of flowers per inflorescence, smaller lateral sepals, and wider and bigger lateral petals and labellum (see table 2).

It is currently acknowledged that hybridization is one of the main factors for speciation (Abbott & al. 2013), especially in plants (Whitman & al. 1999: 426; Mallet 2005), but when the hybrid cohabits with any of the parents and these are legally protected, the risk of disappearance of the parental species due to genetic dilution with the hybrid places decision makers with a complex dilemma, the alternatives being: (i) to intervene in the hybridization process and the possible consolidation of a taxon; (ii) to eradicate the hybrid in order to preserve the protected population of the scarce species (Fay & al. 2007).

The taxonomic complexity of some Mediterranean orchid genera, and the existence of active evolutionary processes with the participation of hybrids, complicate the application of protection regulations conceived for groups which are more evolutionarily stable or have a clearer taxonomy. Therefore, the management of populations affected by hybridization/speciation processes demands a preservation bias (Vereecken & al. 2010: 235; Serra & Soler 2012: 241). Some countries are developing guidance for the management of hybrids with preservation value. In those procedures, the origin of the hybrid (natural/anthropogenic) and the conservation status of the parental species are considered (Jackiw & al. 2015).

The case we are dealing with is a small population of a hybrid cohabiting with large populations of its parents, which are not endangered. In such circumstances we consider of interest the conservation of the hybrid, as is the case with Narcissus × perez-larae Font Quer (Marques & Draper 2006) in the Valencian Community: the hybrid, very scarce, does not pose any risk to the survival of any the parental species, and has been protected.

In the case of Mediterranean orchids, the strategy of presenting a spur without nectar to deceive pollinators is effective only if these can find food in other taxa of the surroundings. This is the case of A. papilionacea subsp. palaestina, A. israelitica (H.Baumann & Dafni) R.M.Bateman & al. and their hybrid A. × feinbruniae (H.Baumann & Dafni) H.Kretzschmar & al. (Vereecken & al. 2010: 233), studied in Israel, in which nectar producing species, like Asphodelus ramosus L. (Asphodelus microcarpus Viv.), live near those orchids. In our case it is highly likely that the abundant presence of Asphodelus cerasiferus J.Gay cohabiting with A. papilionacea subsp. grandiflora has the same effect, increasing the numbers of pollinators of the hybrid.

For all these reasons, it would be necessary to take some in situ conservation measures in the areas where these processes are occurring. Specifically, a flora micro reserve could be established in the classic locality where the hybrid appears, following the example of the Valencian Community (Laguna & al. 2004). Furthermore, it should be noted that the risk of disappearance of these species and the ecological and evolutionary processes in place is high, as part of the hillocks in which the new hybrid lives are considered as building land by the urban regulations of Cartagena.