New Reports of Orchidaceae Family in Southern Calabria (Italy): Distribution and Conservation

Abstract

The Orchidaceae family in Calabria has been scarcely investigated during the 21st century, and available knowledge remains fragmentary, particularly for the rarer taxa. The last comprehensive study dates back to 2002, whereas subsequent checklists, limited to restricted areas of the region, provide incomplete or taxonomically uncertain data. Considering that the family is protected at global (CITES, Bern Convention, IUCN) and national (Italian Red List) levels, broader and more systematic attention is required. In this work, focused on the southern sector of the Metropolitan City of Reggio Calabria, with special reference to the mountain and foothill areas of the Aspromonte massif and in the adjacent districts, we describe four new hybrids for science, each assessed as Critically Endangered (CR), and report four previously unrecorded taxa for the region, evaluated as VU/CR. In addition, two hybrids, newly recorded for the Calabrian flora, were likewise assigned a CR conservation status. The study also provides confirmation of historical records of Ophrys speculum and identifies the southernmost stations in continental Italy for both O. speculum and Orchis branciforti Standardized floral and labellar morphometric traits were measured on representative individuals from each population, including the parental species in the case of hybrids. Conservation status was evaluated following IUCN criteria and GeoCAT-derived AOO values, complemented by field observations on population size, habitat conditions, and site-specific threats. Hybrid names comply with ICN provisions. These findings enhance understanding of orchid biodiversity in Southern Italy and provide new insights for regional conservation efforts.

1. Introduction

The Orchidaceae represent one of the most species-rich families worldwide and are regarded as one of the most fascinating and diverse groups among angiosperms [1]. They are characterized by remarkable floral morphological heterogeneity and an exceptionally broad geographic distribution, occupying a wide range of habitats with the exception of polar regions and deserts and occurring on all continents except Antarctica, with the greatest richness in tropical and subtropical regions [2]. Life forms include epiphytic, terrestrial, and subterranean taxa [3,4], accounting for approximately 8% of all vascular plant species [5,6]. Their distinctive ecological requirements, reproductive strategies, striking colors and scents, along with their high conservation value, have long attracted the attention of both researchers and enthusiasts [1,7,8]. Globally, Orchidaceae comprise ~28,000 taxa assigned to ~763 genera [4,9,10], many of which are endemic, range-restricted, naturally rare, and often threatened [11,12,13,14].

Charles Darwin identified orchids as an ideal model for studying evolution, given their extraordinary diversity and species richness [15]. He highlighted their complex pollination systems as key adaptations promoting cross-fertilization, central to his theory of natural selection [16]. Over time, orchids have evolved highly sophisticated mechanisms to offset the rarity of pollinator visits, combining chemical, visual, and tactile cues [17,18,19,20,21,22,23,24]. They display a wide array of strategies, from producing nectar, lipids, and fragrances to employing diverse deceptive tactics [25,26]. Deceptive orchids represent about 25% of species [27], rising to 33% (“nectarless” sensu van der Pijl & Dodson [25] and up to 36% in recent estimates [21]. “Food deception”, typical of Orchis L., Anacamptis Rich., and Dactylorhiza Neck. ex Nevski, involves showy, nectarless flowers that mimic rewarding species [28,29,30,31,32]. “Sexual deception”, characteristic of Ophrys L., relies on floral mimicry of female pollinators to trigger pseudocopulation [18,30,33,34]. In Serapias L., the flowers mimic small cavities or shelters used by insects for resting, which may not involve true deception since the pollinator obtains the expected reward (i.e., shelter), whereas in Ophrys helenae the mimicry remains primarily deceptive, relying on attraction rather than reward [35,36]. Despite their evolutionary success, Orchidaceae rank among the most threatened plant families worldwide [1,7,8,10,37,38], mainly due to their high ecological specialization, which depends on obligate interactions with pollinators and mycorrhizal fungi, essential for their dispersal and survival [3,4,31,39,40,41]. The main threats are represented by climate change, which negatively affects orchid habitats and reproductive capacities [41,42,43,44,45,46]; by climate-driven range shifts and plant–pollinator mismatches [47,48,49]; by anthropogenic pressure [50]; and by indiscriminate collection and trade, both legal and illegal, because of their demand for ornamental purposes and as raw materials for the production of cosmetic, medicinal and food [51]. Recent assessments indicate that 21.6% of 1855 orchid species analyzed according to IUCN Red List criteria are classified as Endangered (EN), while 12% are Critically Endangered (CR) [39,52,53,54,55]. Due to the high levels of rarity and endemism within the family, all Orchidaceae taxa are listed under Appendix II of the CITES Convention (Bern Convention) [51,56]. Most Orchidaceae taxa are also included in national [57,58,59] and international [60] red lists. In Europe, orchids are a widespread floristic component, occurring in nearly all major ecosystems. However, the highest level of species diversity is observed in southern Europe, particularly in the Mediterranean area, which is regarded as the center of origin and diversification for certain genera, such as Ophrys and Serapias [46]. The Mediterranean Basin represents one of the most important global biodiversity hotspots [61], second only to tropical regions in terms of floristic richness and endemism [62]. Within this biogeographical context, orchids play a prominent role: the Euro-Mediterranean flora includes more than 640 species distributed across 35 genera according to Delforge [63]. From a taxonomic and morphological perspective, Mediterranean orchids are divided into three subfamilies according to pollinia structure: Cypripedioideae, characterized by free pollen grains; Epidendroideae, with compact and indivisible pollinia, also referred to as “waxy” or “hard”; and Orchidoideae, with friable, granular pollinia easily separable into subunits known as massulae [64,65]. Italy represents one of the main Mediterranean hotspots of orchid biodiversity, with 28 genera and approximately 259 species and subspecies, excluding hybrids; despite this global significance, regional knowledge in Southern Italy remains incomplete, particularly in terms of hybridization dynamics and updated taxonomic resolution [66]. Within this framework, the orchid flora of Calabria, although still relatively underexplored, is of particular interest due to the richness and diversity of its taxa. The earliest records of orchids from Calabria are attributed to Tenore [67,68,69,70,71], who provided the oldest detailed observations, later incorporated into the renowned Flore italiane of Bertoloni [72] and Parlatore [73]. Subsequent studies expanded the floristic knowledge of Calabria through regional catalogues and focused surveys [74,75,76,77,78]. In recent decades, research has advanced markedly within national and continental frameworks [79,80,81], culminating in a comprehensive monograph reporting about 80 taxa for the region [82], many involved in ecological interactions and spontaneous hybridization [83]. More recent works have also refined the understanding of spring-flowering species [84]. Since the last comprehensive floristic contribution published in 2003, there has been a notable lack of updated and systematic studies addressing the flora of Calabria. In particular, the Metropolitan City of Reggio Calabria remains poorly investigated, with only scattered or site-specific data available.

Prior to the present study, 78 orchid species were known from the region, including hybrids [85]. The discovery of 11 additional taxa raises the total number of orchid species recorded to 89, thereby confirming the area as one of the richest orchid regions in Southern Italy [66].

In light of these considerations, our contribution provides novel evidence that expands the current knowledge of the Italian orchid flora, with particular reference to Calabria. These findings confirm the role of Orchidaceae as a key group for investigating diversification processes in the Mediterranean region and highlight their relevance for the development of effective biodiversity conservation strategies.

2. Materials and Methods

2.1. Study Area

This study was conducted in Southern Calabria (Southern Italy), focusing on specific areas of the Metropolitan City of Reggio Calabria, with particular attention to the foothill and mountain sectors of the Aspromonte massif (Figure 1).

Research activities took place between 2022 and 2025, through field surveys carried out mainly during the spring and summer seasons. The investigated area is characterized by a Mediterranean macro-bioclimate, with hot and dry summers and maximum rainfall occurring in autumn and winter, showing a pronounced coastal-to-mountain gradient in both temperature and precipitation [89]. According to the bioclimatic classification of Rivas-Martínez applied to the Italian peninsula, dry to subhumid thermo-Mediterranean conditions prevail along the Ionian coast, while on the Tyrrhenian side, more exposed to moist westerly winds, subhumid to humid ombrotypes are observed [89,90]. Moving inland, between 300 and 900 m a.s.l., conditions shift to subhumid–humid meso-Mediterranean, with marked differences between the drier Ionian slopes and the wetter Tyrrhenian slopes [89,91]. Above 900 m, in the mountain sectors of the Aspromonte, humid supra-Mediterranean conditions prevail, while above 1500 m a.s.l. humid to hyper-humid oro-Mediterranean climates occur, characterized by lower temperatures and frequent winter snowfalls [89]. Such climatic variability strongly influences vegetation and soil distribution, resulting in a complex mosaic of habitats ranging from coastal thermo-xerophilous communities to meso- and supra-Mediterranean forest formations at higher elevations [92]. From a geolithological perspective, Southern Calabria, located at the extremity of the Calabrian–Peloritan Arc, exhibits a complex geology characterized by Paleozoic metamorphic rocks, such as gneiss and mica schists, together with Hercynian granitic intrusions outcropping in the Aspromonte massif, shaped over time by subduction, collision, and extensional processes between the African and Eurasian plates [93]. Quaternary uplift events have led to the formation of marine terraces, coastal basins, and a dense network of ephemeral streams (fiumare), while flysch-like successions of the Stilo–Capo d’Orlando formation are also highly relevant [94,95]. This morphological and lithological variety is reflected in the high pedological diversity, with Leptosols and Cambisols on mountain slopes, Arenosols and Calcisols on terraces and dune ridges, and Fluvisols and Vertisols in alluvial plains [96]. As the studied taxa are protected, the geographical coordinates of the stations were generalized for conservation purposes. Points were rounded to 0.02° latitude/longitude (≈2.2 km) and reported with an uncertainty of ±2.2 km; the original coordinates are kept in the authors’ archives and are available upon justified request for scientific purposes. The choice of the generalization level follows the guidelines for the publication of sensitive data, according to Chapman [97].

2.2. Morphometric Data

For each taxon, the following morphometric traits were recorded: flower color and size; shape, color, and dimensions of the perianth elements (petals and sepals); labellum features including callosities, macula, basal field, stigmatic cavity, pseudo-eyes, gynostemium, and apiculum, following the criteria of Vladimirov et al. [98] and Eccarius [99]. The selected characters correspond to those considered taxonomically stable and diagnostically relevant in the Orchidaceae literature, and were chosen because they are easily and consistently measurable on both fresh material and standardized photographic documentation. For morphometric analyses, 15 individuals were measured for the most numerous populations, while all available individuals were analyzed in the case of rare or sparsely represented taxa; for hybrids, measurements were also performed on the parental individuals [100]. To ensure data reliability, measurement error was evaluated through repeated measurements on a subset of samples taken at different times, following the same protocol, with minimal variation between replicates (≤0.2 mm for linear traits), confirming the precision and repeatability of the adopted method. All data were compiled into a summary table. For documentation purposes, no specimens were collected; instead, high-resolution photographic documentation was produced for all taxa studied. The complete photographic database, together with the corresponding georeferenced data, is archived at the Herbarium of the Mediterranean University of Reggio Calabria (REGGIO) [101]. Given the particular complexity of the species concept in Orchidaceae [102,103], nomenclature was first defined according to Bartolucci et al. [66] and subsequently verified and harmonized following the criteria adopted by Plants of the World Online [104] and International Plant Names Index [105]. In addition, the distribution and ecological preferences of the observed taxa were analyzed.

2.3. Conservation Status Assessment

The conservation status of the newly described hybrids was evaluated according to the IUCN Red List Categories and Criteria [106] and the Guidelines for Using the IUCN Red List Categories and Criteria [107]. The analysis was performed using GeoCAT (Geospatial Conservation Assessment Tool) [108] to calculate the Area of Occupancy (AOO) on a 2 × 2 km grid. The final assessment was based on the AOO values obtained through GeoCAT and field data concerning the number of mature individuals and habitat conditions observed for each population. The main threats affecting the habitat of the studied taxa were identified according to the IUCN Threats Classification Scheme [109], which provides a standardized hierarchical system for recording and reporting threat factors impacting species and habitats. Each threat was assigned to its corresponding category and code, and the descriptions were adapted to the local ecological context observed during fieldwork. In addition, the level of threat was also assessed directly in the field, based on the environmental conditions and specific pressures recorded at the sites where each taxon was observed. The conservation assessment was carried out for the newly described hybrids at the global level, and for the other taxa at the regional level (Calabria), since not all georeferenced data for the Sicilian region are currently available; however, these will be implemented and integrated in a future update.

2.4. Hybrids Naming

All hybrid names used in this study follow the rules of the International Code of Nomenclature for algae, fungi, and plants (ICN) [110]. In particular, the nomenclature of hybrids involving different parental species fully complies with the provisions of the Code. In the case of hybrids derived from taxa that differ only at the subspecific level, the nomenclatural treatment adopted is also consistent with the ICN. According to Turland et al. [110] (Arts. H.10–H.12), hybrids between different subspecies may be recognized as distinct nothotaxa when the parental subspecies are morphologically or geographically differentiated.

3. Results

Floristic investigations conducted in the Reggio Calabria area revealed four new orchid hybrids (nov. hybr.) here proposed for the first time in science, along with six taxa newly recorded for the flora of Calabria (nov. rec.), including one species, three subspecies, and two hybrids. In addition, two further taxa were documented at the limit of their distribution range, thus enriching the known diversity of orchid diversity in Southern Italy (

Table 1. List of newly described orchid hybrids and new records for the flora of Calabria (Southern Italy). For each hybrid, the parental species are reported, as determined by morphological traits and field observations.

Hybrids Newly Proposed for Science
New hybrids	Parental species
Dactylorhiza × aspromontana	D. gervasiana × D. sambucina
Ophrys × montis-stellae	O. bertolonii × O. exaltata
Ophrys × spampinatii	O. exaltata × O. incubacea subsp. incubacea
Ophrys × stilarensis	O. exaltata × O. incubacea subsp. brutia
New records for Calabria
Ophrys subfusca (Rchb.f.) Hausskn. subsp. flammeola (P.Delforge) Kreutz
Ophrys subfusca (Rchb.f.) Hausskn. subsp. archimedea (P.Delforge & M.Walravens) Kreutz
Ophrys fusca Link subsp. obaesa (Lojac.) E.G.Camus & A.Camus
Ophrys lunulata Parl.
Ophrys × gelana H.Baumann & Künkele	O. incubacea × O. oxyrrhynchos
Ophrys × mirellae (D’Alonzo & Perilli) Soca	O. fusca subsp. forestieri × O. incubacea subsp. incubacea
Rare species at the southernmost limit of continental Italy
Ophrys speculum Link
Orchis brancifortii Biv.

).

Morphological descriptions of each hybrid examined, together with their parental species, are presented here based on our observations, measurements, and literature data. Morphological descriptions of the newly recorded taxa are likewise provided according to our observations, measurements, and published sources.

The examined individuals exhibited a consistent morphology with only minor phenotypic variation among them, as confirmed by the narrow variability ranges recorded for the main floral characters (mean ± SD), thus supporting a uniform hybrid generation. For instance, hybrid populations showed limited dispersion in key diagnostic traits, such as sepal length, petal dimensions, and labellum measurements, with all mean ± SD values falling within tight intervals characteristic of morphologically homogeneous hybrid cohorts [111].

Among identified hybrids, Ophrys × spampinatii and Ophrys × stilarensis involve the same parental species but different subspecies of one parent. In accordance with the ICN (See above: Section 2.4), both taxa were treated as distinct nothotaxa. Despite their close parentage, they are clearly distinguishable by stable and diagnostic morphological traits and by their occurrence in geographically separated populations, each corresponding to the distribution range of the respective parental subspecies.

3.1. New Hybrids

• Dactylorhiza × aspromontana V.L.A. Laface & L. Torino hybr. nov.

Descriptio: Planta robusta, 25–60 cm alta, habitu inter species parentales mediocri. Folia 4–7, lanceolata ad lineari-lanceolata, viridia, pagina superiore leviter maculata; inferiora 8–15 × 2–4 cm, ultimum (superius) folium bracteiforme efformat. Bracteae virides vel leviter purpurascĕntes, lanceolatae, saepe floribus longiores, praesertim superiores. Inflorescentia densa, conico-cylindrica, 6–15 cm longa, 15–30 floribus mediocribus. Sepala 9–13 × 4–6 mm, erecta, marginibus leviter reflexis, rosea cum striis violaceis. Petala similia, sed parum minora, colore roseo uniforme. Flores rosei; labellum ellipticum, dilatatum, 8–11 × 9–15 mm, subintegerrimum vel leviter trilobatum, zona stigmatica albido-flavescente suffusa; maculae labelli roseo-obscurae ad purpureo-rosaceas variant. Calcar crassum, subconicum, inferne recurvum, 10–14 × 2.5–4 mm, ovario subaequale. Floret mense Iunio (Figure 2b;

Table 2. Comparative morphological traits of Dactylorhiza sambucina, D. × aspromontana, and D. gervasiana.

Morphological Character	D. gervasiana	Dactylorhiza × aspromontana	Dactylorhiza sambucina
Flowers	Violet-purple, larger, often with spotted sepals	Pink, medium-sized, with whitish to yellowish tinges, variable among individuals	Yellow or purplish, medium-sized, color uniform within the same individual
Bracts	Long, especially the lower ones, protruding beyond the inflorescence	Green or purplish-tinged, often exceeding the flower	Green, lanceolate, as long as the flower
Inflorescence	Long, dense, up to 22 cm, cylindrical	Dense, conical-cylindrical, 6–15 cm long	Dense, short, ovoid to subcylindrical
Sepals	7–14 mm, sometimes spotted	9–13 × 4–6 mm, erect, with reflexed margins, pink with violet streaks	8–13 × 4–5.5 mm, ovate, erect or spreading
Petals	Similar, slightly smaller	Uniformly pink, similar to sepals but smaller	Similar to sepals, slightly smaller
Labellum	7–11 × 8.5–16 mm, broad, distinctly trilobed	8–11 × 9–15 mm, elliptic, subentire or trilobed, with dark-pink dots	7.5–11 × 11–17 mm, elliptic, subentire to trilobed, with reddish apices
Spur	9–15 × 2–4.5 mm, thick, subconical, longer than the ovary	10–14 × 2.5–4 mm, subconical to saccate, nearly equalling the ovary	10–15 × 3 mm, thick, subcylindrical, as long as the ovary
Flowering period *	VI–VIII	VI	IV–VI

* The parent species are the only Dactylorhiza taxa occurring in the area and in bloom during the flowering time of the hybrid.

).

Description (English): Sturdy plant, 25–60 cm tall, with intermediate habit between the parental species. Leaves 4–7, lanceolate to linear-lanceolate, green with faint spotting on the upper surface; lower leaves 8–15 × 2–4 cm, the uppermost reduced to a bract-like form. Bracts green or slightly purplish, lanceolate, often longer than the flower, especially the upper ones. Inflorescence dense, conical-cylindrical, 6–15 cm long, bearing 15–30 medium-sized flowers. Sepals 9–13 × 4–6 mm, erect, with slightly reflexed margins, light pink with violet striations. Petals similar but slightly smaller, uniformly pink. Flowers pink; labellum broad-elliptical, 8–11 × 9–15 mm, subentire to slightly trilobed, with pale white-yellow tinge around the stigmatic cavity; spotting on the labellum dark pink to pinkish-purple. Spur thick, subconical, curved downward, 10–14 × 2.5–4 mm, approximately equal to the ovary. Flowering in June (Figure 2b; Table 2).

Type locality: Italia, Calabria, Tre Limiti, Roccaforte del Greco (RC), (WGS84: 38.14° N–15.86° E ± 2.2 km) 1588 m s.l.m., in beech forest.

Iconotype: [2024-06-20], leg. et det. V.L.A. Laface, L. Torino s.n., photographic documentation preserved at REGGIO.

Etymology: The specific epithet refers to Aspromonte massif in Calabria (Southern Italy), where the hybrid was naturally discovered.

Conservation status proposed

Assessed globally as Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with only five genets currently observed in the wild.

Threats: Forestry operations (5.3 Logging & wood harvesting), trampling (6.1 Recreational activities), hydrological alteration (7.2 Dams & water management/use), recurrent wildfires (7.1 Fire & fire suppression), grazing (2.3 Livestock farming & ranching), and occasional plant collection (5.2 Gathering terrestrial plants).

Rationale for the assessment: The hybrid is known only from a single locality in the Aspromonte massif (Southern Calabria), where five mature genets were recorded within a very restricted area (AOO = 4 km²). The habitat, consisting of montane wet meadows and forest margins, is experiencing progressive degradation as a consequence of forestry operations (5.3 Logging & wood harvesting), grazing (2.3 Livestock farming & ranching), trampling from recreational activities (6.1 Recreational activities), and local hydrological alteration (7.2 Dams & water management/use). In addition, the occasional collection of plants by visitors (5.2 Gathering terrestrial plants), attracted by their aesthetic appearance, directly removes reproductive individuals from the population, contributing to its decline. Given the extremely limited distribution, the very small population size, and the continuing decline in both habitat quality and number of mature individuals, the taxon qualifies as Critically Endangered (CR; B2ab(iii,v) + D1) according to the IUCN Red List Categories and Criteria [107].

Previous assessment: No previous assessment is available for this hybrid; this represents the first evaluation both at the regional level (Calabria) and globally, as the taxon is newly described for science.

Conservation actions: The hybrid is currently unprotected by national or regional legislation. It occurs within the boundaries of the Aspromonte National Park, where conservation measures should follow the general guidelines applied to other Orchidaceae. These include the in situ protection of the known population, the restriction of trampling and plant collection, and the long-term monitoring of flowering individuals and habitat conditions.

Parental species description

Dactylorhiza gervasiana (Tod.) H. Baumann & Künkele 1981

Habit and morphology: Plant more robust than D. fuchsii, reaching up to 90 cm in height; leaves 10–20 × 2–6 mm; bracts longer than the flowers, especially the lower ones, which protrude beyond the spike. Inflorescence much longer than in D. fuchsii, up to 22 cm in height; flowers slightly larger, particularly the sepals (7–14 mm, sometimes spotted), and the labellum (7–11 × 8.5–16 mm). Spur thick, subconical to almost sacciform, longer than the ovary (9–15 × 2–4.5 mm). Pollinia greenish; ovary sessile, green, sometimes with purplish tinges; capsule elongate with pronounced ribs; seeds reddish. Chromosome number: 2n = 40.

Distribution: It is an Italic subendemic species, in its typical forms widespread mainly in Sicily and Southern Italy, where it is more frequent than indicated in the distribution map. Its occurrence gradually decreases northward into central Italy, becoming very rare in Tuscany, Umbria, Marche, and Emilia-Romagna, where it is almost always influenced by Dactylorhiza fuchsii. Also reported in Corsica; absent from Sardinia and Liguria (Figure 2a; Table 2).

Etymology: From the Greek δακτυλοϛ (dáctylos) “finger” and ῥίζα (rhíza) “root”, referring to the divided roots resembling fingers; “gervasiana” is dedicated to its discoverer, the pharmacist from Palermo Nicolò Gervasi (1632–1681).

Dactylorhiza sambucina (L.) Soó

Habit and morphology: Robust and relatively short plant (rarely exceeding 30 cm). Stem with 4–7 leaves, light green; lower leaves lanceolate (5–12 × 3–5 cm), upper leaves linear-lanceolate to bract-like. Bracts green, lanceolate, as long as or slightly longer than the flowers. Inflorescence dense, not very elongated (ovoid to subcylindrical), with medium-sized flowers, yellow or purplish-violet; usually each plant bears either entirely yellow or entirely red flowers, even within the same populations. Sepals ovate (8–13 × 4–5.5 mm), lateral ones erect or spreading; petals subequal or slightly smaller. Labellum broadly elliptic (7.5–11 × 11–17 mm), subentire to subtrilobed, often with wavy or denticulate margins, reddish-spotted in the lighter central area (yellowish in red flowers); sometimes slightly convex, with lateral lobes reflexed. Spur thick, subcylindrical, directed downwards, 10–15 × 3 mm, about as long as the ovary and parallel to it. Chromosome number: 2n = 40 (Figure 2c; Table 2).

Distribution: European–Caucasian, absent from much of central-northern and Atlantic Europe. In Italy it occurs in the mountainous areas of northern and central regions, very rare and localized in Southern Italy and Sicily, absent from Sardinia.

Etymology: From the Greek δακτυλοϛ (dáctylos) “finger” and ῥίζα (rhíza) “root”, referring to the divided roots resembling the fingers of a hand. Sambucina: from the genus Sambucus, referring to the elder-like scent emitted by the flowers.

Morphometric measurements of Dactylorhiza × aspromontana based on the examined hybrid individuals (N = 5) revealed limited variability. Inflorescence length (mean ± SD = 10.38 ± 3.06 cm), sepal length (10.7 ± 1.48 mm) and width (5.2 ± 0.57 mm), labellum length (9.6 ± 0.96 mm) and width (13.0 ± 1.58 mm), as well as spur length (12.1 ± 1.43 mm) and width (3.46 ± 0.38 mm), all showed narrow variation ranges. These reduced dispersion patterns support the interpretation of a morphologically homogeneous hybrid population.

2.

Ophrys × montis-stellae V.L.A. Laface & L. Torino hybr. nov.

Descriptio: Planta mediocris (30–45 cm alta), robustior quam O. bertolonii, minus gracilis quam O. exaltata. Inflorescentia 4–10 flores mediocres ad magnos ferens. Bracteae ovario longiores. Sepala ovato-lanceolata (12–16 × 6–8 mm), albido-rosacea, nervis viridibus notata; dorsale erectum vel leviter recurvatum, angulum obtusum cum lateralibus efformans. Petala angusta (8–11 × 2.5–4 mm), rosacea vel purpurea, sepalis intensiora, margine leviter undulato. Labellum integrum vel leviter lobatum (11–17 × 10–15 mm), forma ovali-elongata; margines saepe subrevoluti; color brunneo-nigrans cum reflexibus rubescentibus; pilositas marginalis conspicua sed rarior quam in O. bertolonii, pilositas centralis nulla. Macula in duabus tertiis basalibus labelli sita, ad margines cavitatis stigmaticae extensa, scutiformis et nitida. Campus basalis indistinctus, labello concolor vel obscurior. Pseudoochii magni, griseo-virides. Apiculum in incisura levi insertum, parvum, antrorsum leviter sursumque directum. Gynostemium rostro brevi, angulum rectum cum labello formans. Floret mense Martio–Aprili (Figure 3b;

Table 3. Comparative morphological traits of Ophrys bertolonii, O. × montis-stellae, and O. exaltata.

Morphological Character	Ophrys bertolonii	Ophrys × montis-stellae	Ophrys exaltata
Flowers	2–8, large	4–10, medium to large	4–15, medium-sized
Bracts	Usually shorter than or equal to the ovary	Longer than the ovary	Longer than the ovary
Inflorescence	Few-flowered (pauciflora)	Moderately dense	Dense, multiflorous
Sepals	13–18 × 5–8 mm, whitish-pink, often recurved	12–16 × 6–8 mm, whitish-pink with green veins	Ovate-lanceolate, 10–15 × 5.5–9 mm, whitish-green to lilac
Petals *,1	8–12 × 2–4.5 mm, purplish, ciliate, subparallel	8–11 × 2.5–4 mm, pinkish-purple, with undulate margins	7–11.5 × 2–3.5 mm, dark, with straight or slightly undulate margins
Labellum *,2	13–19 mm, rectangular, dark brown to blackish, saddle-shaped	11–17 × 10–15 mm, oval to elongate, dark brown, with slightly revolute margins	9–15 × 10–16 mm, ovoid-rhombic, brown to reddish-brown
Lateral swellings (gibbosities)	Absent	Absent	Strongly reduced
Macula *,3	Entire, shield- to horseshoe-shaped, shiny	Shield-shaped, glossy, central, extending towards the stigmatic cavity	Simple, H- or X-shaped, sometimes elaborated
Basal field	Dark, not well delimited	Weakly distinct, concolorous or slightly darker	Small, concolorous with the labellum
Stigmatic cavity	Quadrangular, higher than wide	Intermediate, broad, proportionate	Narrowed at base, small
Pseudo-eyes	Blackish, globose, prominent	Grey-greenish, clearly visible	Large, grey with pale ring
Gynostemium	Long and acute beak, at a right angle to the labellum	Short rostrum, forming a right angle with the labellum	Elongated beak, forming an obtuse angle with the labellum
Apiculus	Thick, greenish-yellow, directed upward and forward	Small, slightly directed forward and upward	Very reduced, directed downward
Flowering period	III–V	III–IV	III–IV

* Other sympatric taxa (O. passionis, O. tenthredinifera) were excluded as potential parents based on the shape of the petals ¹ and labellum ² (O. passionis) and the extended macula ³ (O. passionis, O. tenthredinifera).

).

Description (English): Intermediate-sized plant (30–45 cm tall), more robust than O. bertolonii but less slender than O. exaltata. Inflorescence with 4–10 medium to large flowers. Bracts longer than the ovary. Sepals ovate-lanceolate (12–16 × 6–8 mm), whitish-pink with distinct green veins; dorsal sepal erect or slightly recurved, forming an obtuse angle with the laterals. Petals relatively narrow (8–11 × 2.5–4 mm), pink to purplish, darker than the sepals, with slightly undulate margins. Labellum entire or weakly lobed (11–17 × 10–15 mm), elongate-oval, with somewhat revolute margins; dark brown to blackish with reddish tinges; marginal pilosity evident but less dense than in O. bertolonii; central pilosity absent. Macula occupying the basal two-thirds of the labellum, centrally positioned and extending to the edges of the stigmatic cavity, scutiform and shiny as in O. bertolonii. Basal field poorly defined, concolorous with or darker than the labellum. Pseudo-eyes prominent, grey-green. Apiculum inserted in a shallow notch, small, directed slightly upwards and forwards. Gynostemium with short beak, forming a right angle with the labellum. Flowering period: March–April (Figure 3b; Table 3).

Type locality: Italia, Calabria, Santuario Madonna della Stella, Pazzano (RC), (WGS84: 38.48° N–16.46° E ± 2.2 km), 726 m s.l.m., in a meadow near pine plantations.

Iconotype: [2025-03-29], leg. et det. V.L.A. Laface, L. Torino s.n., photographic documentation preserved at REGGIO.

Etymology: the name originates from the place of discovery, namely from the Sanctuary of Monte Stella near the town of Pazzano (Metropolitan City of Reggio Calabria, Southern Italy).

Conservation status proposed

Assessed globally as Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with only five genets currently observed in the wild.

Threats: Forestry operations (5.3 Logging & wood harvesting), trampling (6.1 Recreational activities), hydrological alteration (7.2 Dams & water management/use), recurrent wildfires (7.1 Fire & fire suppression), grazing (2.3 Livestock farming & ranching), and occasional plant collection (5.2 Gathering terrestrial plants).

Rationale for the assessment: The hybrid is known only from its type locality at the Santuario Madonna della Stella near Pazzano (Reggio Calabria Province, Southern Italy), at approximately 726 m a.s.l. The population consists of only five mature genets (fewer than 50 mature individuals in total), confined to a small montane meadow near pine plantations (AOO = 4 km²). The habitat, characterized by open mesophilous grasslands and garrigue patches, is undergoing progressive degradation due to forestry operations, grazing, trampling by visitors to the sanctuary, and local hydrological alteration. The occasional collection of flowering plants by visitors, attracted by their showy appearance, further reduces the number of reproductive individuals. Given the extremely limited distribution, the very small population size, and the continuing decline in both habitat quality and number of mature individuals, the taxon qualifies as Critically Endangered (CR; B2ab(iii,v) + D1) according to the IUCN Red List Categories and Criteria [107].

Previous assessment: No previous assessment is available for this hybrid; this represents the first evaluation both at the regional level (Calabria) and globally, as the taxon is newly proposed for science.

Conservation actions: The hybrid is not currently protected by specific national or regional legislation. Although its population is limited to a single locality, no formal conservation measures have yet been implemented. Actions consistent with those applied to other Orchidaceae are recommended, including in situ protection of the known site, control of trampling and plant collection near the area, regulation of grazing and small-scale forestry activities, and long-term monitoring of flowering individuals and habitat conditions to prevent further decline.

Parental species description

Ophrys bertolonii Moretti

Habit and morphology: Slender plant with an erect habit, usually attaining 25–30 cm in height. Inflorescence pauciflorous, with 2–8 relatively large flowers. Sepals ovate-lanceolate (13–18 × 5–8 mm), whitish-pink to rose, occasionally with greenish tinges but almost always marked by green veins, typically recurved backwards. Petals narrowly lanceolate (8–12 × 2–4.5 mm), reddish-purple and more intensely colored than the sepals; acute, with straight (rarely slightly undulate) margins, slightly ciliate, arranged nearly parallel. Labellum entire, elongate (13–19 mm), sub-rectangular, blackish, generally with a shallow saddle-like depression at the level of the macula and with revolute margins; surface covered by dense, dark hairs, sometimes with reddish hues. Macula glossy, located in the median to distal portion of the labellum, scutiform or horseshoe-shaped, brightly colored from blue to reddish-violet. Basal field concolorous with the labellum, not clearly delimited. Stigmatic cavity higher than wide, quadrangular, constricted above the pseudo-eyes, which are globose, blackish, and prominent, positioned towards the base of the labellum rather than on the cavity walls. Apiculum robust, inserted into a deep notch, greenish-yellow, obtuse, directed upwards or forwards. Gynostemium elongated, acute, beak-shaped, arranged at a right angle to the labellum.

Distribution: Italic endemic, occurring throughout the peninsula south of the Po River and extending to Sicily (Figure 3a; Table 3).

Etymology: Ophrys, from the Greek ὀφρύς (ophrýs), meaning “eyelash” or “eyebrow”; bertolonii, dedicated to the Bolognese botanist Antonio Bertoloni (1775–1869).

Ophrys exaltata Ten.

Habit and morphology: Vigorous plant with an erect and slender habit, reaching 50–60 cm in height. Inflorescence bearing 4–15 medium to large flowers; bracts longer than the ovary. Sepals ovate-lanceolate (10–15 × 5.5–9 mm), varying in color from whitish-green to pink or lilac, sometimes with more intense tones and greenish veins or suffusions; the dorsal sepal is nearly erect and forms an obtuse angle (>110°) with the lateral ones. Petals relatively long and narrow (7–11.5 × 2–3.5 mm), glabrous, darker than the sepals, occasionally with reddish tinges, with margins straight or slightly undulate. Labellum subentire, transversely convex, relatively small (9–15 × 10–16 mm), ovate to rhomboid in shape, reddish-brown; callosities weakly developed; surface densely hairy in a submarginal band, sometimes interrupted by a paler glabrous border. Macula usually simple, basal, H or X shaped, occasionally with whitish ornamentation or more complex patterns, extending from the base to the middle or two-thirds of the labellum. Basal field small and concolorous with the labellum, as is the stigmatic cavity, which has a more or less constricted base. Pseudo-eyes large, grey, often surrounded by a pale halo. Apiculum very reduced, inserted into a deep notch, directed downwards (Figure 3c; Table 3).

Distribution: Southern Italian subendemic, mainly occurring in Sicily and Calabria; further north it is only sporadically reported as far as Abruzzo.

Etymology: Ophrys, from the Greek δακτυλοϛ (dáctylos) “finger” and ῥίζα (rhíza) “root”; exaltata, from the Latin “elevata”, referring to the erect and slender habit of the plant.

Morphometric measurements of Ophrys × montis-stellae, based on the examined hybrid individuals (N = 5), revealed limited variability. Flower number (mean ± SD = 7.2 ± 2.39), sepal length (14.26 ± 1.25 mm) and width (7.08 ± 0.62 mm), petal length (9.66 ± 1.07 mm) and width (3.30 ± 0.52 mm), and labellum length (14.2 ± 2.07 mm) and width (12.6 ± 1.79 mm) all showed relatively narrow variation ranges. These reduced dispersion patterns support the interpretation of a morphologically homogeneous hybrid population.

3.

Ophrys × spampinatii V.L.A. Laface & L. Torino hybr. nov.

Descriptio: Planta habitu intermedio inter species parentales, gracilis, 30–40 cm alta. Inflorescentia mediocriter pauciflora, 3–10 flores magnos gerens, compacta. Bracteae lanceolatae. Sepala 12–15 × 3–7 mm, ovato-lanceolata, viridia vel albida cum nervis viridibus, leviter antrorsum curvata. Petala 7–10 × 2–4 mm, angusta et elongata, viridi-flavescentia, marginibus leviter undulatis, intensiora. Labellum 10–14 × 10–15 mm, rectangulare, integrum vel subtrilobum, marginibus retroflexis, pilis submarginalibus brunneo-rubescentibus, mediocriter densis. Gibbae fere absentes. Macula variabilis, forma H/X vel duabus fasciis circum campum basalem, interdum ornata zona clariore. Campus basalis obscurior quam labellum. Cavitas stigmatica rectangulata. Pseudo-ocelli viridi-grisei, annulo viridi pallido cincti. Apiculus minimus vel nullus, in incisura parva insertus. Gynostemium rostro mediocri. Floret a fine Martii ad Aprilem (Figure 4b;

Table 4. Comparative morphological traits of Ophrys exaltata, O. × spampinatii, and O. incubacea subsp. incubacea.

Morphological character	Ophrys exaltata	Ophrys × spampinatii	Ophrys incubacea subsp. incubacea
Flowers	4–15, medium-sized	3–10, large	3–8, large
Bracts	Longer than the ovary	Lanceolate	Not always longer than the ovary
Inflorescence	Dense, multiflorous	Moderately few-flowered, compact	Robust, slender
Sepals	Ovate-lanceolate, 10–15 × 5.5–9 mm, whitish-green to lilac	12–15 × 3–7 mm, ovate-lanceolate, light green to whitish, with greenish veins, slightly recurved	10–15.5 × 4–7.5 mm, light green, sometimes reddish
Petals *,1	7–11.5 × 2–3.5 mm, dark, with straight or slightly undulate margins	7–10 × 2–4 mm, relatively long and narrow, yellow-greenish, margin weakly undulate	6–9.5 × 2.5–5 mm, ovate-oblong, margin always undulate
Labellum *,2	9–15 × 10–16 mm, ovoid-rhombic, brown to reddish-brown	10–14 × 10–15 mm, rectangular, entire or subtrilobed, with reflexed margins, brown-reddish submarginal pilosity	13–14 × 10–14.5 mm, large, ovoid-orbicular, dark brown to blackish, with diffuse brown-reddish pilosity
Lateral swellings (gibbosities)	Nearly absent	Strongly reduced	Well developed
Macula	Simple, H- or X-shaped, sometimes elaborated	H/X-shaped with two bands sometimes with pale ornamentation, not reaching the gibbosities	Two parallel, grey-violet bands, extending to the gibbosities
Basal field	Small, concolorous with the labellum	Darker than the labellum	Same color as the labellum
Stigmatic cavity	Narrowed at base, small	Rectangular	Very narrow, dark, with a central white spot
Pseudo-eyes *,3	Large, grey with pale ring	Grey-greenish, bordered with light green	Small, blackish, bordered with white
Gynostemium	Elongated beak, forming an obtuse angle with the labellum	Intermediate in length	Short, acute rostrum
Apiculus	Very reduced, directed downward	Reduced or absent, forming a small incision	Extremely reduced, often recurved
Flowering period	III–IV	III–IV	III–V

* The overall morphology (long and narrow petals ¹, strongly developed labellar pilosity ², ex. O. passionis) and the presence of pseudo-eyes ³ exclude the other sympatric species (Ophrys incubacea, Ophrys fusca subsp. forestieri, O. sicula).

).

Description (English): Plant of intermediate habit between the parental species, usually slender, 30–40 cm tall. Inflorescence moderately few-flowered (3–10 large flowers), compact. Bracts lanceolate. Sepals 12–15 × 3–7 mm, ovate-lanceolate, color ranging from light green to whitish with greenish veins, slightly incurved. Petals 7–10 × 2–4 mm, relatively long and narrow, greenish-yellow, margins weakly undulate with more intense tones. Labellum 10–14 × 10–15 mm, rectangular, entire or subtrilobed, margins reflexed; submarginal hairs present and often consistent but not dense, reddish-brown in color. Calli nearly absent. Basal macula variable, H/X-shaped with two bands surrounding the basal field, sometimes with pale ornaments. Basal field darker than the lip. Stigmatic cavity rectangular. Pseudo-eyes greyish-green, bordered with pale green. Apicule reduced or absent, set in a small notch. Gynostemium with rostrum of intermediate length. Flowering: late March—April (Figure 4b; Table 4).

Type locality: Italia, Calabria, Scariglione, Motta San Giovanni (RC), (WGS84: 38.00° N–15.74° E ± 2.2 km), 839 m s.l.m.,at the edge of a pine forest.

Iconotype: [2024-03-23], leg. et det. V.L.A. Laface, L. Torino s.n., photographic documentation preserved at REGGIO.

Etymology: The specific name is dedicated to the botanist Professor Giovanni Spampinato, Full Professor at the Faculty of Agriculture of the Mediterranean University of Reggio Calabria, and teacher and mentor of the author of this work.

Conservation status proposed

Assessed globally as Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with about ten genets currently observed in the wild.

Threats: Shrub encroachment due to land abandonment (7.3 Other ecosystem modifications), recurrent fires (7.1 Fire & fire suppression), and occasional collection of plants by visitors (5.2 Gathering terrestrial plants) attracted by their aesthetic appearance.

Rationale for the assessment: The hybrid is known only from its type locality at Motta San Giovanni (Reggio Calabria Province, Southern Italy), at approximately 839 m a.s.l., located at the edge of a pine forest. The population consists of about ten mature genets (fewer than 50 mature individuals in total) confined to a small area (AOO = 4 km²). The habitat, consisting of xeric calcareous slopes and ecotonal grasslands bordering reforested areas, is undergoing progressive degradation due to grazing, shrub encroachment following land abandonment and the occurrence of recurrent wildfires. The occasional collection of flowering plants by visitors, attracted by their distinctive appearance, represents an additional threat that directly reduces the number of reproductive individuals. Given the extremely limited distribution, the very small population size, and the continuing decline in both habitat quality and number of mature individuals, the taxon qualifies as Critically Endangered (CR; B2ab(iii,v) + D1) according to the IUCN Red List Categories and Criteria [107].

Previous assessment: No previous assessment is available for this hybrid; this represents the first evaluation both regionally (Calabria) and globally, as the taxon is newly proposed for science.

Conservation actions: The hybrid is not currently protected by specific national or regional legislation. Although its population is restricted to a single locality, no formal conservation measures have been implemented to date. Actions consistent with those applied to other Orchidaceae are recommended, including in situ protection of the known site, regulation of grazing, prevention of plant collection, and long-term monitoring of flowering individuals and habitat conditions to prevent further decline.

Parental species description

Ophrys exaltata Ten.: See above (Section 3.2, Table 3).

Ophrys incubacea Bianca subsp. incubacea

Habit and morphology: Robust, slender plant, with stems reaching up to 50 cm in height. Inflorescence bearing 3–8 large flowers. Sepals ovate-lanceolate (10–15.5 × 4–7.5 mm), light green in color, sometimes with reddish tinges and, more rarely, pinkish (var. dianensis). Petals relatively large (about 2/3 the length of the sepals: 6–9.5 × 2.5–5 mm), ovate-oblong, green, ochraceous, or reddish-brown, consistently darker than the sepals, with markedly undulate margins. Labellum large, ovate-orbicular (13–14 × 10–14.5 mm), directed obliquely forward and downward; dark brown to blackish, occasionally reddish (var. septentrionalis), with recurved margins and a dense, long, reddish-brown submarginal indumentum. Callosities well developed, glabrous on the inner surface and hairy externally. Basal field concolorous with the labellum. Macula usually composed of two elongated, parallel, glossy bands, grayish-violet to bluish, rarely with a pale border, extending from the base to the middle or two-thirds of the labellum. Apiculum very reduced, often bent backwards, inserted in a small but distinct notch. Stigmatic cavity narrow at the base, showing a strong contrast between the white walls and the basal white spot and the otherwise dark labellum. Pseudo-eyes small, blackish, surrounded by a whitish ring. Pollinial chambers often reddish externally. Gynostemium short, with an acute beak (Figure 4c; Table 4).

Distribution: Central-western Mediterranean element, ranging from the Iberian Peninsula to the Adriatic countries. In Italy, mainly found in the major islands, the southern and Tyrrhenian regions, and western Liguria, while rare in northern and Adriatic areas.

Etymology: Ophrys, from the Greek ὀφρύς (ophrýs), meaning “eyelash” or “eyebrow.” The epithet incubacea derives from the Latin incubus “nightmare”, referring to the dark coloration of the labellum reminiscent of nocturnal terrors (the synonym atrata conveys the same meaning).

Morphometric measurements based on the examined individuals (N = 10) showed limited variability in Ophrys × spampinatii. Flower number (mean ± SD = 7.00 ± 2.45), sepal length (13.90 ± 0.96 mm), sepal width (5.58 ± 1.18 mm), petal length (9.12 ± 0.99 mm), petal width (3.34 ± 0.59 mm), labellum length (12.95 ± 1.32 mm) and labellum width (13.55 ± 1.51 mm) all exhibited narrow variation ranges. The reduced dispersion around intermediate values between the parental species supports the interpretation of a morphologically homogeneous hybrid population.

4.

Ophrys × stilarensis V.L.A. Laface & L. Torino hybr. nov.

Descriptio: Planta robusta sed gracilis, 35–55 cm alta. Inflorescentia mediocriter densa, 3–10 floribus mediocribus ad magnis instructa. Bracteae ovario longiores. Sepala ovato-lanceolata (8–13 × 4–7 mm), plerumque viridia pallida nervis viridibus leviter signata; sepalum dorsale antrorsum inflexum, angulum rectum cum lateralibus efficiens. Petala satis magna (6–10 × 2–4 mm), ovato-oblonga, sepalis obscuriora, viridia vel brunneo-rubentia, margine undulato. Labellum subintegrum, transversim convexum, ovaliforme (10–12 × 11–13 mm), brunneo-rubrum vel fusco-brunneum, marginibus revolutis et pilis submarginalibus conspicuis; gibbis carens. Campus basalis et cavitas stigmatica paulo obscuriores quam labellum, rubro-brunnei, cavitate ad basin relative angustata. Macula simplex vel moderate composita, saepe in fasciis vel forma H/X, nitida, griseo-violacea ad cyanea. Apiculum valde reductum vel nullum, in incisura parva non profunda insertum, deorsum directum vel antrorsum leviter inflexum. Pseudo-oculi virides vel griseo-nigricantes, plerumque annulo claro cincti. Florescit: exeunte Martio–Aprili (Figure 5b;

Table 5. Comparative morphological traits of Comparative morphological traits of Ophrys exaltata, O. × stilarensis, and O. incubacea subsp. brutia.

Morphological character	Ophrys exaltata	Ophrys × stilarensis	Ophrys incubacea subsp. brutia
Flowers	4–15, medium-sized	3–10, medium-sized	3–8, large
Bracts	Longer than the ovary	Longer than the ovary	About as long as the ovary
Inflorescence	Dense, multiflorous	Moderately dense	Few-flowered, moderately compact
Sepals	Ovate-lanceolate, 10–15 × 5.5–9 mm, whitish-green to lilac	8–13 × 4–7 mm, light green, with faint greenish veins	7–12 × 3–6 mm, green to whitish or pinkish
Petals	7–11.5 × 2–3.5 mm, dark, with straight or slightly undulate margins	6–10 × 2–4 mm, ovate-oblong, green or reddish-brown, margin undulate	5–7 × 2.5–5 mm, ovate-oblong, green, ochre or reddish-brown, margin undulate
Labellum *,1	9–15 × 10–16 mm, ovoid-rhombic, brown to reddish-brown	10–12 × 11–13 mm, ovate, dark brown to reddish-brown, recurved margins with submarginal pilosity	10–14 × 11–15 mm, subequal, brownish, with recurved margins and evident submarginal hairs
Lateral swellings (gibbosities)	Strongly reduced	Absent	Absent or faint
Macula	Simple, H- or X-shaped, sometimes elaborated	Simple or moderately complex, banded or H/X-shaped	Simple, with two parallel grey-violet or bluish bands
Basal field	Small, concolorous with the labellum	Slightly darker than the labellum, reddish-brown	Lighter than the labellum, yellow-orange
Stigmatic cavity	Narrowed at base, small	Relatively narrow at the base	Very narrow at the base
Pseudo-eyes *,2	Large, grey with pale ring	Green or grey-blackish, bordered with pale ring	Dark green, often bordered with pale ring
Gynostemium	Elongated beak, forming an obtuse angle with the labellum	Intermediate between parental taxa	Robust, typical of the group
Apiculus *,3	Very reduced, directed downward	Very reduced or absent, in a shallow notch, directed downward	Extremely reduced, directed downward and/or forward
Flowering period *,4	III–IV	III–IV	III–IV

* Other Ophrys species recorded in the area were excluded as possible parents because of the hybrid’s distinctive labellar pilosity ¹ (O. apulica), the presence of pseudo-eyes ² (O. bombyliflora, O. sicula), the almost absent apiculum ³ and the flowering time ⁴ (O. apulica).

).

Description (English): Plant robust yet slender, 35–55 cm tall. Inflorescence moderately dense, with 3–10 medium to large flowers. Bracts longer than the ovary. Sepals ovate-lanceolate (8–13 × 4–7 mm), usually pale green with faint greenish veins; dorsal sepal bent forward, forming a right angle with the laterals. Petals relatively large (6–10 × 2–4 mm), ovate-oblong, darker than the sepals, green or reddish-brown, with undulate margins. Lip subentire, transversely convex, oval in shape (10–12 × 11–13 mm), reddish-brown to dark brown, with recurved margins and conspicuous submarginal pubescence; swellings absent. Basal field and stigmatic cavity slightly darker than the lip, reddish-brown, the cavity rather narrow at the base. Macula simple or moderately complex, often band-like or forming an H/X pattern, glossy, greyish-violet to bluish. Apicule very reduced or absent, set in a shallow and narrow notch, directed downward or slightly forward. Pseudo-eyes green or greyish-black, usually encircled by a pale halo. Flowering: late March–April (Figure 5b; Table 5).

Tipe locality: Italia, Calabria, Bavigliardo, Camini (RC), (WGS84: 38.44° N–16.50° E ± 2.2 km), 137 m s.l.m., in garrigue habitat. Ophrys × stilarensis occurs within a Habitat Directive type, corresponding to habitat 5330 “Thermo-Mediterranean and pre-desert scrub”, subtype 32.23—Garrigues dominated by Ampelodesmos mauritanicus (sensu Annex I of Directive 92/43/EEC).

Iconotype: [2025-03-30], leg. et det. V.L.A. Laface, L. Torino s.n., photographic documentation preserved at REGGIO.

Etymology: The name derives from the Stilaro river, which flows near the towns of Stilo and Bivongi (Metropolitan City of Reggio Calabria, Southern Italy).

Conservation status proposed

Assessed globally as Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with about seven genets currently observed in the wild.

Threats: Occasional plant collection by visitors (5.2 Gathering terrestrial plants) attracted by their ornamental appearance, grazing (2.3 Livestock farming & ranching), recurrent fires (7.1 Fire & fire suppression), and minor infrastructure expansion (1.3 Tourism & recreation areas).

Rationale for the assessment: This natural hybrid is restricted to a single locality at Bavigliardo, Camini (Reggio Calabria Province, Southern Italy), at approximately 137 m a.s.l. Only seven mature genets have been observed, confined to a small garrigue habitat within a Mediterranean shrubland landscape (AOO = 4 km²). The site is subject to multiple pressures, including the occasional collection of plants by visitors (5.2), attracted by their showy appearance, grazing by livestock (2.3), recurrent fires (7.1), and localized infrastructure development associated with tourism (1.3). These factors collectively contribute to a progressive decline in habitat quality and in the number of mature individuals. Given the extremely restricted distribution, the small population size, and the ongoing decline in both habitat condition and population size, the taxon qualifies as Critically Endangered (CR; B2ab(iii,v) + D1) according to the IUCN Red List Categories and Criteria [107].

Previous assessment: No previous assessment is available for this hybrid; this represents the first evaluation both regionally (Calabria) and globally, as the taxon is newly proposed for science.

Conservation actions: The hybrid is not currently covered by any specific national or regional protection measures. Despite being confined to a single locality, no dedicated conservation actions have yet been implemented. Management strategies similar to those adopted for other Orchidaceae are recommended, including in situ protection of the known site, regulation of grazing and tourist activities, prevention of plant collection, and long-term monitoring of flowering individuals and habitat conditions to prevent further decline.

Parental species description

Ophrys exaltata Ten.: See above (Section 3.2, Table 3)

Ophrys incubacea Bianca subsp. brutia (P.Delforge) Kreutz

Habit and morphology: Species related to Ophrys incubacea. Robust and slender plant, erect stem up to 50 cm, bearing 3–8 large flowers. Sepals ovate-lanceolate (7–12 × 3–6 mm), green, whitish or pink, sometimes with reddish tinges. Petals relatively large (5–7 × 2.5–5 mm), ovate-oblong, green-ochre or reddish-brown, darker than sepals, margins undulate. Lip subequal (10–14 × 11–15 mm), directed obliquely forward and downward, brownish with recurved margins and conspicuous submarginal pubescence, reddish-brown; swellings weak or scarcely visible. Basal field and stigmatic cavity lighter than lip, yellow-orange; cavity very narrow at base. Macula simple, often consisting of two parallel elongated bands, glossy, grey-violet or bluish, broader than long. Apicule very reduced, often recurved, inserted in a small but distinct notch. Pseudo-eyes dark green, often encircled by a pale ring (Figure 5c; Table 5).

Distribution: Southern Italian subendemic, mainly occurring in Calabria.

Etymology: Ophrys, from the Greek δακτυλοϛ (dáctylos) “finger” and ῥίζα (rhíza) “root”; brutia, from the Bruttii, an ancient pre-Roman people of Calabria.

Morphometric measurements of Ophrys × stilarensis, based on the examined hybrid individuals (N = 7), revealed limited variability. Flower number (mean ± SD = 6.14 ± 2.41), sepal length (10.83 ± 1.68 mm) and width (5.56 ± 1.01 mm), petal length (8.21 ± 1.36 mm) and width (3.01 ± 0.67 mm), and labellum length (11.20 ± 0.71 mm) and width (12.26 ± 0.68 mm) all exhibited narrow variation ranges. These reduced dispersion patterns support the interpretation of a morphologically homogeneous hybrid population.

3.2. New Records for Calabria

Floristic investigations conducted in Southern Calabria have also led to the identification of new records of spontaneous orchids for the regional flora. For the first time, Ophrys subfusca subsp. flammeola, O. subfusca subsp. archimedea, O. fusca subsp. obaesa, and O. lunulata have been recorded, together with the hybrids Ophrys × gelana and O. × mirellae (Supplementary Materials: Table S1; Figure S1).

Ophrys subfusca (Rchb.f.) Hausskn. subsp. flammeola (P.Delforge) Kreutz

+CAL: Scariglione, Motta San Giovanni (Reggio Calabria), (WGS84: 38.02° N–15.74° E ± 2.2 km) Pinus sp. pl. plantation, 847 m a.s.l., 2 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO).—Species new for the flora of Calabria.

Note: Species regarded as a Sicilian endemic [66]. The observed individuals occur in relatively dense populations, mostly in the vicinity of Pinus sp.pl. plantations, flowering from late March to late April.

Conservation status proposed

Regional assessment proposed for Calabria region, according to IUCN [107] criteria:

Vulnerable (VU; B2ab(iii) + D1 + D2) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality.
• Criterion D1: Population composed of fewer than 250 mature individuals, approximately 100 mature individuals observed in the wild, concentrated within a relatively small area;
• Criterion D2: Occurrence restricted to a single locality with an AOO < 20 km² and exposure to plausible threats, which could rapidly drive the taxon to a higher threat category.

Threats: Grazing (2.3 Livestock farming & ranching), trampling from recreational activities (6.1 Recreational activities), recurrent fires (7.1 Fire & fire suppression), and the occasional collection of flowering individuals (5.2 Gathering terrestrial plants) by visitors attracted by their ornamental appearance.

Rationale for the assessment: This subspecies, endemic of Sicily, is known in Calabria from a single locality at Motta San Giovanni (Reggio Calabria Province, Southern Italy), at approximately 847 m a.s.l., in the vicinity of Pinus sp. pl. plantations. About 100 mature individuals were recorded within a relatively small area (AOO = 4 km²). The population occupies Mediterranean grasslands and open shrublands influenced by human disturbance and forestry contexts. The main pressures include grazing, trampling, and recurrent fires, which contribute to the degradation of the habitat. The occasional collection of flowering individuals further reduces the number of reproductive plants. Although the Area of Occupancy (AOO = 4 km²) formally falls within the threshold for higher threat categories, the overall level of threat and population size do not indicate an imminent risk of extinction. Given its limited distribution, restricted AOO, relatively small population size, and the inferred continuing decline in habitat quality, the taxon is best assessed as Vulnerable (VU; B2ab(iii) + D1) at the regional level (Calabria), according to the IUCN Red List Categories and Criteria [107].

Previous assessment: In the Lista Rossa della Flora Italiana. 2. Endemismi e altre specie minacciate [58], the taxon is listed as Ophrys flammeola P. Delforge, a synonym of Ophrys subfusca (Rchb.f.) Hausskn. subsp. flammeola (P. Delforge) Kreutz, and is assessed as Least Concern (LC). At present, comprehensive data required for a global assessment, including the recently discovered Calabrian populations, are not available; therefore, we here provide only a regional assessment for Calabria.

Conservation actions: The subspecies is currently protected by specific national and regional legislation only for Sicily [58]. In Calabria, its occurrence near managed pine plantations and Mediterranean grasslands suggests potential vulnerability to human activities and fire. Conservation actions should focus on in situ protection of the known site, limitation of grazing pressure and recreational trampling, prevention of flower collection, and periodic monitoring of the population to track demographic trends and habitat conditions.

Ophrys subfusca (Rchb.f.) Hausskn. subsp. archimedea (P.Delforge & M.Walravens) Kreutz

+CAL: Monte Pietrerosse, Montebello Ionico (Reggio Calabria), (WGS84: 38.00° N–15.74° E ± 2.2 km), open areas within a Pinus sp. pl. plantation, 780 m a.s.l., 2 April 2025, leg. et det. V. L.A. Laface, L. Torino (REGGIO).—Species new for the flora of Calabria.

Note: Sicilian endemic subspecies [66], probably also derived from hybridization events between the Fuscae group and the Luteae group. The Calabrian populations are rather localized and tend to form dense but spatially restricted clusters, with a preferential distribution in anthropogenic or reforested habitats, particularly in the vicinity of Pinus sp. pl. plantations. Flowering occurs mainly in April, when individuals are easily recognizable and observable in the field.

Conservation status proposed

Regional assessment proposed for Calabria region, according to IUCN [107] criteria:

Vulnerable (VU; B2ab(iii) + D1 + D2) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality.
• Criterion D1: Population composed of fewer than 250 mature individuals, approximately 100 mature individuals observed in the wild, concentrated within a relatively small area;
• Criterion D2: Occurrence restricted to a single locality with an AOO < 20 km² and exposure to plausible threats, which could rapidly drive the taxon to a higher threat category.

Threats: Grazing by livestock (2.3 Livestock farming & ranching), recurrent wildfires (7.1 Fire & fire suppression), and the occasional collection of flowering plants by visitors (5.2 Gathering terrestrial plants) attracted by their ornamental appearance.

Rationale for the assessment: In Calabria, this subspecies is known from a single locality at Monte Pietrerosse, Montebello Ionico (Reggio Calabria Province, Southern Italy), at approximately 780 m a.s.l., in open areas within a Pinus sp. pl. plantation. The population comprises around 100 mature genets, concentrated in a limited area (AOO = 4 km²). The species grows in semi-open Mediterranean shrublands associated with reforested or anthropogenic habitats, subject to grazing, fire, and collection pressure. These threats are expected to cause a continuing decline in both habitat quality. Although the AOO value formally falls within the threshold for higher threat categories, the overall level of risk and population size do not indicate imminent extinction. Given its restricted distribution, small population size, and exposure to plausible threats that could quickly affect its survival, the taxon is best assessed as Vulnerable (VU; B2ab(iii) + D1 + D2) at the regional level (Calabria), according to the IUCN Red List Categories and Criteria [107].

Previous assessment: According to Rossi et al. [59], the taxon is listed as Ophrys archimedea P.Delforge & M.Walravens, a synonym of Ophrys subfusca (Rchb.f.) Hausskn. subsp. archimedea (P.Delforge & M.Walravens) Kreutz, and is assessed as Least Concern (LC). At present, comprehensive data required for a global assessment, including the recently discovered Calabrian populations, are not available; therefore, we here provide only a regional assessment for Calabria.

Conservation actions: The subspecies is currently protected by specific national and regional legislation only for Sicily [59]. In Calabria, the population occurs within semi-anthropogenic habitats near managed Pinus plantations and Mediterranean grasslands, where it shows a strong localization in small, dense clusters. Given its restricted distribution and the potential vulnerability to human disturbance, fire, and grazing, targeted conservation measures are required. Actions consistent with those applied to other Orchidaceae are recommended, including in situ protection of the known site, limitation of grazing pressure and recreational trampling, prevention of plant and flower collection, and monitoring of flowering individuals and habitat conditions to detect early signs of population decline. Where feasible, awareness-raising initiatives should also be promoted among local visitors and land managers to prevent unintentional damage to the small existing population.

Ophrys fusca Link subsp. obaesa (Lojac.) E.G.Camus & A.Camus

+CAL: Contrada Genovese, Motta San Giovanni (Reggio Calabria), (WGS84: 38.02° N–15.72° E ± 2.2 km), grazed meadow, 691 m a.s.l., 27 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO).—Species new for the flora of Calabria.

Note: Taxon belonging to the Ophrys funerea group, considered a Sicilian endemic subspecies [66]. It occurs in grassland habitats near some artificial basins in March, coinciding with the end of the flowering period of Ophrys fusca subsp. lupercalis.

Conservation status proposed

Regional assessment proposed for Calabria region, according to IUCN [107] criteria:

Vulnerable (VU; B2ab(iii) + D1 + D2) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality.
• Criterion D1: Population composed of fewer than 250 mature individuals, approximately 50–70 mature individuals observed in the wild, concentrated within a relatively small area;
• Criterion D2: Occurrence restricted to a single locality with an AOO < 20 km² and exposure to plausible threats, which could rapidly drive the taxon to a higher threat category.

Threats: Grazing by livestock (2.3 Livestock farming & ranching), recurrent wildfires (7.1 Fire & fire suppression), and the occasional collection of flowering plants by visitors (5.2 Gathering terrestrial plants) attracted by their ornamental appearance.

Rationale for the assessment: In Calabria, this subspecies is known from a single locality at Motta San Giovanni (Reggio Calabria Province, Southern Italy), at approximately 691 m a.s.l., within grazed meadows near small artificial basins. The population consists of about 50–70 mature genets, concentrated in a restricted area (AOO = 4 km²). Ophrys fusca subsp. obaesa, is a Sicilian endemic subspecies. Its occurrence in Calabria represents the first record for the mainland, extending its known range. The site is exposed to multiple pressures, including livestock grazing, occasional fires, and flower collection by visitors. These factors are expected to cause a continuing decline in habitat quality and population viability. Although the AOO formally falls within the thresholds of higher threat categories, the overall risk and population size do not currently indicate imminent extinction. Given the restricted distribution, small population, and plausible threats that could quickly affect its survival, the taxon is best assessed as Vulnerable (VU; B2ab(iii) + D1 + D2) at the regional level (Calabria), according to the IUCN Red List Categories and Criteria [107].

Previous assessment: At the national level, the taxon was reported as Ophrys obaesa Lojac. in Rossi et al. [59], where it was assessed as Least Concern (LC). At present, there are insufficient georeferenced data to allow a comprehensive global assessment. Therefore, the current evaluation is limited to the regional context of Calabria, where the species is reported for the first time.

Conservation actions: The subspecies is currently protected by specific national and regional legislation [59]. In Calabria, it grows in semi-natural grazed meadows near artificial basins and managed Pinus plantations, forming small, dense clusters within a limited area. Considering its restricted distribution and the potential vulnerability to grazing, fire, and human disturbance, the implementation of targeted conservation measures is strongly recommended. These should include the in situ protection of the known population, the regulation of grazing pressure, and the prevention of flower collection, accompanied by regular monitoring of flowering individuals and habitat quality. Furthermore, awareness-raising initiatives aimed at local visitors and land managers would help reduce unintentional impacts and contribute to the long-term preservation of this newly discovered population.

Ophrys lunulata Parl.

+CAL: Scariglione, Motta San Giovanni (Reggio Calabria), (WGS84: 38.02° N–15.74° E), rock formations in a Pinus sp. pl. plantation, 809 m a.s.l., 13 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO).—Species new for the flora of Calabria.

Note: Sicilian endemic species [66], occurring within its historical range in two disjunct areas in the south-eastern and western parts of the island, with only scattered records elsewhere. Reported in this study for the first time from peninsular Italy/continental Europe.

Conservation status proposed

Regional assessment proposed for Calabria region, according to IUCN (2022) criteria:

Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with only two genets recorded at the single known site.

Threats: Habitat degradation from forestry management (5.3 Logging & wood harvesting), trampling from recreational activities (6.1 Recreational activities), grazing (2.3 Livestock farming & ranching), and potential collection of plants (5.2 Gathering terrestrial plants) due to their ornamental appeal.

Rationale for the assessment: Ophrys lunulata is a Sicilian endemic species. This study documents the first occurrence of the species in continental Italy (Calabria) and Europe, at Motta San Giovanni (Reggio Calabria Province), approximately 809 m a.s.l., on rocky outcrops within a Pinus sp. pl. plantation. Only two mature genets were observed within a restricted microhabitat, suggesting a very small and isolated population. The site is potentially affected by forestry operations, trampling, and grazing, which could lead to habitat disturbance and reduced reproductive success. Given its extremely limited distribution, tiny population size, and ongoing pressures on habitat quality, the species is assessed as Critically Endangered (CR; B2ab(iii,v) + D1) at the regional level (Calabria), according to the IUCN Red List Categories and Criteria (Version 3.1, Second Edition, 2022).

Previous assessment: At the national level, the species was listed as Lower Risk (LR) by Conti et al. [57] and later assessed as Least Concern (LC) by Rossi et al. [58]. The Calabrian population reported here represents the first record for mainland Italy. Due to the lack of updated and comprehensive data on the Sicilian populations, the taxon is currently assessed only at the regional level (Calabria).

Conservation actions: The species is legally protected at the national level as part of the family Orchidaceae [58]. In Calabria, the known population occurs in a reforested area near managed Pinus sp.pl. plantations, making it potentially vulnerable to human disturbance and forestry activities. Conservation priorities should include strict in situ protection of the site, limitation of forestry operations and trampling, prevention of flower collection, and regular monitoring of the population and habitat dynamics. Given its extremely small population, ex situ conservation measures (seed banking or symbiotic germination trials) should also be considered to preserve genetic material and support potential future reinforcement actions.

New hybrids

Ophrys × gelana H.Baumann & Künkele

+CAL: Contrada Genovese, Motta San Giovanni (Reggio Calabria), (WGS84: 38.017369° N–15.725959° E), grazed meadow, 797 m a.s.l., 2 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO).—Hybrid new for the flora of Calabria.

Note: The hybrid taxon between O. incubacea × O. oxyrrhynchos [105], hitherto known only from Sicily [112], was found in April in grassland habitats near artificial basins, concomitant with the flowering of the parental species. Such secondary and mosaic habitats appear to favour contact between the parent taxa and hybridization, facilitated by their co-phenology. Its presence has been continuously documented since 2019.

Conservation status proposed

Regional assessment proposed for Calabria region, according to IUCN (2022) criteria:

Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with only five genets recorded at the single known site.

Threats: Grazing (2.3 Livestock farming & ranching), recurrent wildfires (7.1 Fire & fire suppression), recreational trampling (6.1 Recreational activities), and occasional plant collection (5.2 Gathering terrestrial plants) due to ornamental appeal; locally, management of nearby water bodies may alter microhabitat conditions (7.2 Dams & water management/use, where applicable).

Rationale for the assessment: In Calabria, O. × gelana is known from a single locality within a restricted grazed meadow near artificial basins, the hybrid was first recorded in 2019, and subsequent monitoring over 7 years confirmed the persistence of only five mature genets confined to a very small area (AOO = 4 km²). The hybridization context (secondary mosaics, co-phenology of parental taxa) facilitates origin but does not ensure persistence, given intense local pressures (grazing, trampling, fires, plant collection) that plausibly drive a continuing decline in habitat quality and in the number of mature individuals. With one location, tiny population size, and documented threats, the taxon qualifies as CR (B2ab(iii,v) + D1) at the regional level (Calabria) under IUCN [107].

Previous assessment: No national IUCN assessment is available for this hybrid; hybrid taxa are generally not evaluated in Italian Red Lists. This represents the first regional assessment for Calabria.

Conservation actions: Priorities include in situ protection of the known site; regulation of grazing pressure and visitor trampling; prevention of plant/flower collection through on-site awareness; and long-term monitoring of flowering individuals and habitat condition. Given the very small population, consider ex situ safeguards (seed banking, symbiotic germination trials) as a precautionary measure.

Ophrys × mirellae (D’Alonzo & Perilli) Soca

CAL: Monte Pietrerosse, Montebello Ionico (Reggio Calabria), (WGS84: 37.998201° N–15.736820° E), Pinus sp. pl. plantation, 701 m a.s.l., 2 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO).—Hybrid new for the flora of Calabria.

Note: The hybrid between Ophrys fusca subsp. forestieri and Ophrys incubacea subsp. incubacea [105,113] was found in the vicinity of Pinus sp. pl. plantations, in marginal habitats characterized by grasslands and secondary clearings. The co-occurrence of the two parental taxa during the same phenological period favoured reproductive contact and the formation of the hybrid. This represents the first documented record for Calabria.

Conservation status proposed

Regional assessment proposed for Calabria region, according to IUCN [107] criteria:

Critically Endangered (CR; B2ab(iii,v) + D1) (Table S1).

Criteria Applied:

• Criterion B: AOO = 4 km², calculated in GeoCAT [108] using a 2 × 2 km grid.
  (a) Number of locations = 1.
  (b) Continuing decline observed in: (iii) habitat quality; (v) number of mature individuals.
• Criterion D1: Population composed of fewer than 50 mature individuals, with only five genets recorded at the single known site.

Threats: Habitat degradation linked to forestry operations (5.3 Logging & wood harvesting), grazing (2.3 Livestock farming & ranching), recurrent wildfires (7.1 Fire & fire suppression), recreational trampling (6.1 Recreational activities), and occasional plant collection (5.2 Gathering terrestrial plants). The mosaic of secondary and anthropogenic habitats increases exposure to disturbance.

Rationale for the assessment: In Calabria, Ophrys × mirellae is known from a single site within a Pinus plantation at Motta San Giovanni (701 m a.s.l.), where only three mature genets were recorded within a restricted area (AOO = 4 km²). The species occupies secondary grasslands and clearings at forest edges, environments subject to grazing, fire, forestry management, and occasional collection, all of which contribute to a decline in habitat quality and reproductive individuals. Given the extremely limited distribution, very small population size, and ongoing threats, the hybrid qualifies as Critically Endangered (CR; B2ab(iii,v) + D1) at the regional level (Calabria), in accordance with the IUCN Red List Categories and Criteria [107].

Previous assessment: No national or regional IUCN assessment is currently available for this hybrid, which was previously known only from Sicily. This represents the first evaluation and first confirmed record for Calabria.

Conservation actions: Targeted conservation measures should include in situ protection of the known site, limitation of grazing pressure and trampling, restriction of forestry operations during the flowering period, and prevention of plant collection. A long-term monitoring program of flowering individuals and habitat conditions is strongly recommended. Awareness activities involving local stakeholders and forestry managers would help to prevent unintentional disturbance and promote conservation of this rare hybrid taxon.

3.3. Rare Species at the Southernmost Limit of Continental Italy

Ophrys speculum Link

Confirm historical reports and southernmost record from continental Italy: Serro di Corico, Motta San Giovanni (Reggio Calabria), (WGS84: 38.007083° N–15.733081° E), open areas within a Pinus sp. pl. plantation, 818 m a.s.l., 16 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO); Maracani, Motta San Giovanni (Reggio Calabria), (WGS84: 38.015078° N–15.741481° E), Pinus sp. pl. plantation, 866 m a.s.l., 22 April 2024, leg. et det. V.L.A. Laface, L. Torino (REGGIO); Piano di Sant’Elia, Metropolitan City of Reggio Calabria (Reggio Calabria), (WGS84: 38.115099° N–15.732537° E), Pinus sp. pl. plantation, 684 m a.s.l., 23 April 2024, leg. et det. V.L.A. Laface, L. Torino (REGGIO)—Southernmost station of peninsular Italy.

Note: This is a species with a steno-Mediterranean distribution [104], widely distributed in the major Italian islands but rare in the rest of the peninsula. In Calabria, its presence was reported for the province of Cosenza in 2010 by Rosati et al. [114]. Our record, however, represents the first documented evidence for the province of Reggio Calabria, corroborating ancient historical data. Indeed, the species had already been mentioned by Tenore [115] under the name Ophrys ciliata Biv. in the Flora Napolitana, where he indicated as typical locality the hills surrounding the city of Reggio Calabria (“Nasce ne’ colli intorno Reggio; fiorisce in maggio; è perenne”); this presence was later reiterated in the Sylloge plantarum vascularium florae neapolitanae [71]. Subsequently, Parlatore [116] again reported the species for the Reggio area. Our observation therefore constitutes the first confirmation for the province after as many as 167 years and, at the same time, documents the southernmost station of peninsular Italy.

Orchis brancifortii Biv.

CAL: Scariglione, Motta San Giovanni (Reggio Calabria), (WGS84: 38.011522° N–15.732068° E), rock formations in a Pinus sp. pl. plantation, 811 m a.s.l., 13 April 2025, leg. et det. V.L.A. Laface, L. Torino (REGGIO), southernmost station of peninsular Italy.

Note: Orchis brancifortii represents an endemic taxon with a relatively restricted distribution, limited to Sicily, Sardinia, and Calabria [66]. In the latter region, the species has so far been reported only from very restricted areas of the province of Reggio Calabria, particularly in the Stilo district [117], where populations appear localized and of small size. The occurrence of this taxon in the southernmost sector of the Italian peninsula therefore has significant biogeographical relevance, as it documents the extreme continental limit of the species’ distribution, otherwise known mainly from the major islands.

4. Discussion

Floristic and orchidological investigations carried out in the Reggio Calabria area substantially enrich current knowledge on the diversity of Orchidaceae in Southern Italy, documenting four hybrids new to science within the genera Dactylorhiza and Ophrys, as well as two additional hybrids newly recorded for the regional flora. Moreover, four taxa never previously reported from Calabria were identified (Ophrys subfusca subsp. flammeola, O. subfusca subsp. archimedea, O. fusca subsp. obaesa, O. lunulata), together with the current confirmation of O. speculum and the evidence of the southernmost peninsular station for Orchis brancifortii.

With the inclusion of the newly recorded taxa, the total number of orchid species known from Reggio Calabria area rises to 89. Within the genus Ophrys, which represents a major component of the orchid flora, the number of recorded species increases from 31 [85] to 41 following the present findings. This remarkable concentration of taxa within a relatively limited area highlights the exceptional contribution of the Metropolitan City of Reggio Calabria to the regional orchid diversity [66].

The study area can be interpreted as a major contact zone between peninsular and insular floristic elements, a pattern already documented in other Mediterranean taxa such as Quercus and Cistus [118,119]. The heterogeneous topography and habitat mosaic of the Aspromonte massif function as a micro-refugium, favouring the survival and diversification of orchids, as similarly observed for relict or thermophilous taxa such as Fagonia cretica and Lavandula multifida [91,120].

Within this context, the coexistence of different evolutionary lineages, combined with habitat fragmentation, heterogeneity and complex pollination dynamics, has historically promoted, and continues to promote, the formation of new hybrids, while enabling the persistence of small peripheral populations [91,111,121,122,123,124]. Similar hybridization patterns in Sicily, Apulia and the Aegean region further support the interpretation of Southern Calabria as a comparable Mediterranean hotspot where the ecological heterogeneity of limestone landscapes favours repeated hybrid formation [63]. The presence in Calabria of taxa traditionally considered Sicilian endemics (e.g., O. lunulata, O. subfusca subsp. flammeola, O. fusca subsp. obaesa, O. subfusca subsp. archimedea) extends their known distribution to the continental side of the Strait of Messina and reinforces the long-recognised biogeographical bridge role of this region [82,125]. This evidence fits within the broader Calabrian–Peloritan biogeographical framework, which identifies the area as a key transition node between southern Apennine and Sicilian floristic lineages [91,126]. This insular influence can be interpreted in the context of Quaternary climatic oscillations and post-glacial migration routes, during which the Strait of Messina acted as a connection corridor rather than a barrier, favouring the persistence of relict populations and the bidirectional dispersal of floristic elements [121,127,128]. This selective bridge role is supported by the strong chorological and phytogeographical affinities between the Peloritani and the Aspromonte, shaped by successive phases of colonization and isolation [125,129]. In this context, the concept of Mediterranean micro-refugia [130] aptly describes the Aspromonte and the Ionian ranges, where topographic and climatic complexity has allowed the persistence of taxa with disjunct or fragmented distributions.

The new hybrids (hybr. nov.) show consistent and repeatable morphological patterns, with traits that are intermediate between the parental species.

In Dactylorhiza, the most distinctive characters involve the size and proportions of the inflorescence and the spur. Hybridization between D. sambucina and D. gervasiana appears to be favoured by a partial ecological overlap in terms of altitude and soil requirements, with contacts occurring between montane meadows and mesophilous–ombrophilous clearings and more open, light-exposed patches. Added to this is the morphological plasticity of the spur (ranging from subconical to saccate), of the bracts, and of the overall size of the inflorescence, which in hybrid individuals tends to stabilize at intermediate values. These constant and easily recognizable traits represent a recurrent diagnostic feature in the D. fuchsii s.l. × D. sambucina hybrid complexes already documented in the literature [63].

In Ophrys, the main diagnostic traits involve the architecture of the labellum, the pattern of submarginal pilosity, the shape and sheen of the macula, and the configuration of both the apiculum and the basal field. When these morphological characters are considered together with phenological and microenvironmental data, it becomes possible to reliably distinguish the different parental combinations and to infer the likely direction of introgressive flows. These findings are consistent with the classical literature on the morphology and pollination of the genus [63,131] and meet the pragmatic criteria for taxonomic recognition pending genetic confirmation [132].

Hybridization of Ophrys species is frequently promoted by phenological overlap and by a partial convergence of the odor bouquets that regulate the mechanisms of “sexual deception” in pollinators [133,134]. The co-occurrence of O. exaltata and O. incubacea (incl. subsp. brutia) in xerothermophilous ecotones and at the margins of reforestations creates contact zones where even slight variations in visual and chemical signals may induce cross-pollen transfer and pollinator-mediated interference. The morphological evidence observed in the three Ophrys hybrids described here is consistent with this scenario, showing on the one hand the retention of distinctive parental traits (e.g., the extension of the macula in combinations with O. bertolonii, or the submarginal villous band and the typical “H/X” patterns of O. incubacea), and on the other hand a clearly intermediate structural morphology (such as the angle of the dorsal sepal, the development of the rostrum of the gynostemium, or the degree of revolute margins of the labellum). These traits represent classical indicators of recent hybridogenesis in the genus [63] and confirm the crucial role of ecological interactions and pollinator-mediated signals in the diversification processes of Ophrys [133,134].

The formation of the recorded hybrids seems to be driven by multiple interacting factors, including anthropogenic disturbance, habitat overlap, and pollinator sharing. These forces act together across fragmented Mediterranean landscapes. Human-induced alterations, such as reforestation, grazing, and forest-edge management, create secondary contact zones and increase both spatial and phenological proximity among closely related taxa [131,135]. In such disturbed or mosaic environments, the breakdown of ecological barriers can facilitate opportunistic hybridization, raising the probability of cross-pollination events [136]. Furthermore, the partial overlap in pollinator assemblages and floral scent patterns, well documented in Ophrys and other sexually deceptive orchids, promotes hybrid formation by increasing pollinator-mediated reproductive interference [137]. Taken together, these elements indicate that hybridization in the study area is not accidental. Instead, it appears to represent an adaptive response to environmental heterogeneity and to anthropogenic modification of Mediterranean ecosystems.

A recurring pattern observed in several hybrids and in three of the new regional records is their frequent association with artificial Pinus sp. pl. plantations. Although these stands are not natural ecosystems, they create a mosaic of clearings, edges, and nutrient-poor or poorly developed soils (skeletal or weakly acidic on detrital substrates), which increase microenvironmental heterogeneity and bring ecological niches into closer spatial proximity [138]. Such conditions favour secondary contact zones and promote opportunistic hybridization. These environments also act as corridors for xerophilous species with early phenology (late March–April), increasing the chances of co-flowering and cross-visits by pollinators [139]. While these settings facilitate contact and hybrid formation, they also raise extinction risk due to forest management, trampling, and recurrent wildfires. Comparable situations have been documented in other Mediterranean areas, where artificial reforestations and secondary grasslands host residual populations of terrestrial orchids that remain vulnerable to management practices and anthropogenic disturbance [38,140].

These findings also underline the conservation relevance of the area, which includes several Natura 2000 sites [141]. The occurrence of endemic or rare taxa, listed in Annexes II and IV of the Habitats Directive and in CITES, provides an additional criterion for management priority [142]. In particular, one of the newly described hybrids, Ophrys × montis-stellae, occurs within the SAC IT9350136 “Vallata dello Stilaro”, further emphasizing the floristic and conservation importance of the site.

Among the recorded taxa, Ophrys × stilarensis is the only one occurring within a recognized EU Habitats Directive type, specifically habitat 5330 “Thermo-Mediterranean and pre-desert scrub”, subtype 32.23—Garrigues dominated by Ampelodesmos mauritanicus (sensu 92/43/EEC, Annex I [143]). This association highlights the ecological specificity of the hybrid, confined to xeric garrigue environments, and reinforces the conservation relevance of Mediterranean scrub habitats for the maintenance of local orchid diversity [144,145].

From a conservation perspective, the taxa described and recorded here are characterized by extremely limited distributions and small populations, often restricted to single localities (AOO = 4 km²) and subject to ongoing pressures. The four newly described hybrids (Dactylorhiza × aspromontana, Ophrys × montis-stellae, O. × spampinatii, O. × stilarensis) have been assessed as Critically Endangered (CR; B2ab(iii,v) + D1), while among the new records, Ophrys lunulata also qualifies as CR due to the presence of only two mature genets. The remaining taxa (O. subfusca subsp. flammeola, O. subfusca subsp. archimedea, and O. fusca subsp. obaesa) are best assessed as Vulnerable (VU; B2ab(iii) + D1 + D2), given their small populations (50–100 individuals) and exposure to plausible threats such as grazing, fires, trampling, and collection.

Priority conservation measures should include the in situ protection of known sites, the seasonal regulation of forestry operations, the management of visitor access to limit trampling and opportunistic collection, and the adaptive control of grazing pressure. Evidence from European and Mediterranean case studies demonstrate that controlled grazing and forest management can significantly contribute to maintaining floristic and microhabitat diversity, thereby supporting the conservation of Orchidaceae communities [139,146,147]. In this regard, projects such as LIFE Xero-grazing have shown that extensive and well-regulated grazing can favour the persistence of xeric grasslands and orchid-rich meadows, counteracting shrub encroachment and the loss of open habitats [148]. All these actions should be integrated into long-term monitoring programmes of flowering individuals and habitat conditions, in order to evaluate the effectiveness of the implemented measures and the population response over time [140].

Considering the extremely small size of certain populations, particularly Ophrys lunulata and several hybrids, precautionary ex situ interventions, including seed banking and symbiotic germination trials, are advisable to preserve genetic material and support potential future reinforcement actions [149,150]. Such approaches have proven effective in securing germplasm of rare European orchids and mitigating the risk of genetic erosion in micro-populations subject to stochastic events. In the sites falling within the Natura 2000 network, management should be aligned with the safeguards provided by Annexes II and IV of the Habitats Directive [143] and with the CITES provisions for Orchidaceae [151]. In addition, the integration of adaptive management and local stakeholder engagement has been shown to enhance long-term outcomes for Mediterranean orchid conservation [38,139].

The rediscovery of O. speculum in the province of Reggio Calabria holds considerable historical significance, as the species, already reported by Tenore [71,115] and later by Parlatore [116], had lacked confirmation for more than a century and a half. The observations presented here fill this knowledge gap, documenting the currently southernmost stations of peninsular Italy and at the same time confirming the steno-Mediterranean nature of the taxon [104]. Moreover, field evidence indicates the occurrence of vegetative propagation whereas spontaneous self-pollination has not been observed so far, suggesting that the studied plants are not self-fertile. A comparable biogeographical relevance is found in Orchis brancifortii, a subendemic taxon also occurring in Calabria at the southern limits of the peninsula, reported only in the last two decades from the area around Stilo [63].

Although the hybrid diagnoses presented in this study are supported by consistent morphological, phenological, and ecological evidence, the absence of molecular data represents a limitation. It is hoped that future studies will integrate DNA barcoding or population-level genomic analyses (e.g., ITS, nrDNA, plastid markers) to confirm parental combinations and clarify introgressive dynamics. The combination of molecular tools and field-based ecological observations will be essential to fully understand the evolutionary and conservation significance of these hybrid complexes.

5. Conclusions

Investigations carried out in Southern Calabria have significantly expanded knowledge of the regional orchid flora, with the description of four hybrids new to science, the identification of two additional hybrids and four taxa never before reported for the region, together with the current confirmation of Ophrys speculum and the documentation of the southernmost peninsular station of Orchis brancifortii. These results confirm the role of the Aspromonte and the Ionian ranges as a biogeographical hub of primary importance and as a hotspot of diversity, where microenvironmental heterogeneity, contact zones, and peculiar pollination dynamics promote both recent hybridization processes and the persistence of localized populations at the margin of their range. Taken together, these data highlight the need to consider such environments as priority areas for conservation, requiring continuous monitoring and integrated approaches (morphometric, genetic, and ecological) aimed at clarifying diversification mechanisms and defining targeted management strategies for the long-term protection of Mediterranean orchid biodiversity.