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Article

Exploring the Orchid Flora of Montenegro: Ten Newly Identified Taxa

by
Boris Đoka Radak
*,
Jovan Milan Peškanov
,
Bojana Slavenko Bokić
and
Goran Timča Anačkov
Department of Biology and Ecology, Faculty of Sciences, University of Novi Sad, Trg Dositeja Obradovića 2, 21000 Novi Sad, Serbia
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(5), 337; https://doi.org/10.3390/d17050337
Submission received: 18 March 2025 / Revised: 15 April 2025 / Accepted: 17 April 2025 / Published: 6 May 2025
(This article belongs to the Section Plant Diversity)
Browse Figures
Versions Notes

Abstract

:
Montenegro, with approximately 3600 vascular plant taxa, represents one of the floristic diversity centers of the Balkan Peninsula and the Mediterranean region. According to our current knowledge, about 80 to 100 orchids, including species, subspecies, and hybrids, occur in this country. To refine the understanding of orchid diversity in Montenegro, extensive fieldwork and herbarium revisions were conducted. As a result, ten new orchid taxa were recorded in Montenegro for the first time. These include Epipactis helleborine subsp. distans, E. leptochila subsp. neglecta, Gymnadenia densiflora, Neotinea ustulata var. aestivalis, Ophrys archipelagi, O. grammica, and Pseudorchis albida subsp. tricuspis. In addition, one intergeneric hybrid, ×Serapicamptis rousii, and two intrageneric, Anacamptis ×nicodemi and A. ×olida, were found. Distribution data, ecological preferences, and population sizes for newly registered taxa in Montenegro have been given. This study underscores the great potential of the flora of Montenegro for discoveries in the orchid world.

1. Introduction

Montenegro is a country on the Balkan Peninsula, located in its central part along the coast of the Adriatic Sea. This region is influenced by the various geological, climatic, and hydrological factors, resulting in a diverse and rich plant life. The flora of the vascular plants of Montenegro is exceptionally diverse, with an estimated 3600 species and subspecies [1]. However, this number is based generally on the published data, and some areas of the country have not been entirely floristically explored yet, suggesting that the actual number is likely to be higher [2].
The study of Montenegro’s flora and vegetation has a long history, spanning two centuries and involving over 1000 researchers [3]. Among those researchers, one of the central figures is the Czech botanist Rohlena, who published the capital work “Conspectus florae Montenegrinae” in 1942, listing 2623 species and 194 subspecies of flowering plants and ferns [4]. However, despite the numerous research studies, the Flora of Montenegro has still not been published yet. The first and most significant steps in that direction were made with the publication of the Catalogue of vascular flora of Montenegro I and II [2,5]. These two publications are based on the five prominent publications [4,6,7,8,9], a synthesis of other literature data, as well as the author’s field data. Within these Catalogues, a total of 1429 species and subspecies, as well as 25 hybrid taxa of vascular plants, were included [2,5].
The family Orchidaceae is one of the most prominent families of vascular plants, comprising about 28,000 species [10]. They inhabit all continents except Antarctica and are most numerous in humid tropical and subtropical areas, especially in the cloud forests of tropical America and Asia, where epiphytic species predominate [11]. In Europe, however, only terrestrial geophytes are present [12]. The number of orchid species in Europe increases from north to south [13,14] and reaches its center of diversity in the Mediterranean region, to which Montenegro belongs. This region is noted for its exceptional diversity in specific genera, such as Ophrys [12,15], along with the genus Serapias, which has its center of evolution and speciation in the eastern Mediterranean [16]. All orchids are protected by the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES, Appendix A) [17].
Current synthetic floristic publications regarding Montenegro [2,5] have not yet addressed the Orchidaceae family. Previous research on this group has been sporadic, with most findings emerging from general botanical studies or research of specific areas of Montenegro, leaving our understanding of its diversity relatively limited. In support of this, there are discoveries of several new taxa of the Orchidaceae family in the country’s area in the last few years [18,19,20]. Based on our research and best knowledge, this family includes at least 80 to 100 species, subspecies, and hybrids in this country, which can be classified into 20 genera [21]. Among them, the genus Epipactis Zinn has five species [21], with two newly registered subspecies—E. helleborine (L.) Crantz subsp. distans (Arv.-Touv.) R. Engel and P. Quentin and E. leptochila (Godfery) Godfery subsp. neglecta Kümpel. The first of these, E. helleborine subsp. distans, was described by French botanist C. Arvet-Touvet as E. distans [22]. While some authors classify this taxon as the subspecies of the widespread E. helleborine [16,23,24,25,26,27], others maintain its original classification as a distinct species [12,28,29]. From an extreme perspective, Kühn et al. [30] included E. distans within the type subspecies of E. helleborine. On the other side of the extreme is Delforge [12], who observes E. distans and E. helleborine var. orbicularis (K. Richt.) Verm. as two separate taxa, while, without exception, all other authors (cited above) regard them as the same taxon. Molecular analyses (next-generation sequencing technology) support the position of this taxon as the subspecies of E. helleborine [31,32]. Additionally, an extensive revision of the E. helleborine group confirmed that E. helleborine subsp. distans is a well-defined taxon [33]. Another newly recorded subspecies of this genus in Montenegro is E. leptochila subsp. neglecta, which was first described by Kümpel from Thuringia, Germany. Since then, this taxon has been classified in various ways: as a variety [12], as a subspecies [25,28,30,34,35,36,37], or at the species level [23,29,38,39]. Recent research based on ITS and AFLP analyses [40] and RAD-sequences [31] has demonstrated that E. leptochila subsp. neglecta and related taxa are not separated by a sufficiently large genetic distance to deserve the species rank. Consequently, they should be regarded as subspecies of E. leptochila.
Until now, the genus Gymnadenia R. Br. in Montenegro was represented by four species—G. conopsea (L.) R. Br., G. frivaldii Hampe ex Griseb., G. odoratissima (L.) Rich., and G. rhellicani (Teppner and E. Klein) Teppner and E. Klein [21]. A newly recorded species, G. densiflora (Wahlenb.) A. Dietr., was described in Sweden [41] as Orchis conopsea L. β (=var.) densiflora Wahlenb. Since then, different authors have treated this taxon in various ways, as a variety of G. conopsea [12], as a subspecies of G. conopsea [25,34,42,43] or a distinct species [23,44,45]. However, significant divergence in ITS sequences between G. conopsea s. str. and G. densiflora, as shown by Bateman et al. [46], indicates that G. densiflora is a valid species. Moreover, Marhold et al. [47] confirmed this claim by analyzing chromosome number, ploidy level, and morphological characteristics of both floral and vegetative traits. Recently, Bateman et al. [48] came to the same conclusion about separating these taxa at the species level.
In Montenegro, the genus Neotinea Rchb. f. is represented by three species: N. maculata (Desf.) Stearn., N. tridentata (Scop.) R. M. Bateman, Pridgeon and M. W. Chase, and N. ustulata (L.) R. M. Bateman, Pridgeon and M. W. Chase [21]. The last includes two varieties, var. ustulata and var. aestivalis (Kümpel) Tali, M. F. Fay and R. M. Bateman, a new taxon for Montenegro. Authors had different perspectives on this taxon, from the extreme case as a late-flowering ecotype [12] to variety [15,30,49] or even a subspecies of the N. ustulata [23,25,50,51]. Several studies [52,53,54] have confirmed that there are morphological, visual (spectral reflectance pattern), phytochemical, and pollinator differences between var. ustulata and var. aestivalis. Therefore, these forms cannot be regarded solely as ecotypes of N. ustulata. Furthermore, while there are phenological differences between them, morphological differentiation [55] and genetic divergence are minimal [56]. Therefore, these two forms should be considered the varieties of N. ustulata.
The genus with the most taxa is Ophrys L., which includes at least 14 species and subspecies in Montenegro [21]. Among them are two newly registered species—O. archipelagi Gölz and H. R. Reinhard and O. grammica (B. Willing and E. Willing) Devillers-Tersch. and Devillers. Taxonomic status of O. archipelagi is problematic. Delforge [12] identifies it as a distinct species within the O. exaltata group of the O. sphegodes complex. Similarly, relevant floristic publications from Croatia [34,57,58] treat O. archipelagi as an independent species. Flora and Checklist of the vascular flora of Italy [29,59] regard this taxon as a subspecies of the taxonomically problematic species O. exaltata Ten. Contrary to the previous two points of view, a group of authors [30,60] places this taxon within a partially stabilized hybrid complex called O. ×arachnitiformis Gren. and M. Philippe. Ophrys grammica was described from Grammos Mt. (Greece) by B. Willing and E. Willing [61] as a subspecies of O. mammosa Desf. Delforge [12] considers it a well-defined species within the O. mammosa group of the O. argolica complex. Most researchers from Greece [16,62] also recognize O. grammica as a species, but classify it within the O. sphegodes group. In contrast, some Western European authors [30,60] synonymize a range of species, including O. grammica, under O. sphegodes Mill. subsp. mammosa (Desf.) Soó ex E. Nelson. Recently, Niketić and Djordjević [63] proposed a new combination of names for O. sphegodes subsp. mammosa, renaming it O. sphegodes subsp. taurica (Aggeenko) Soó ex Niketić and Djordjević, and included O. grammica as a synonym. However, recent preliminary morphological studies examining various taxa of the genus Ophrys have shown that there are reliable and distinct morphological characteristics that separate O. grammica from O. sphegodes subsp. taurica [64].
Pseudorchis albida (L.) Á. Löve and D. Löve is the only species representative of its genus in the territory of Montenegro [21]. The classification of newly recorded taxon P. albida subsp. tricuspis (Beck) E. Klein has changed over time. Initially considered a form [65], it was later classified as a variety [12,66,67] and a subspecies [25,68,69,70]. It is important to highlight that while P. albida and P. straminea (Fernald) Soják (syn. P. albida subsp. straminea (Fernald) Á. Löve and D. Löve) have been subjected to the molecular analyses that confirmed their separation at the species level, samples of P. albida subsp. tricuspis have not been analyzed in this manner. Consequently, the definitive taxonomic status of this orchid remains uncertain, making it inappropriate to synonymize it with P. albida subsp. albida [71].
This study presents the first findings of ten orchid taxa in Montenegro, contributing to understanding their distribution patterns, ecological preferences, and population size in Montenegro. The following scientific objectives guide this research: (i) distributional analysis—to document the distribution of newly registered orchid taxa in Montenegro, as well as their distribution pattern on the Balkan Peninsula (the distance from their nearest populations in neighboring countries), providing a baseline for future conservation measures; (ii) ecological characterization—to determine habitat preferences and ecological conditions associated with the presence of these taxa, offering insights into their ecological requirements and (iii) population assessment—to determine the exact number of ramets of the newly registered taxa at their currently known localities in Montenegro, identifying factors that may influence their long-term survival.

2. Materials and Methods

The field research was conducted throughout all vegetation seasons, from late February to November, from 2021 to 2023. The study encompassed all altitudinal zones of Montenegro, from the sea level to the peaks of high mountains (up to 2523 m asl, Bobotov Kuk Peak, Durmitor Mt.). All geographic subregions of Montenegro were included in the field research: northern (Durmitor Mt. and Stožer Mt.), eastern (Bjelasica Mt., Kučke Mts., Prokletije Mts. and Visitor Mt.), western (Njegoš Mt. and Somina Mt.), central (Moračke Mts.), Submediterranean (Kopiljsko Polje, Kopitnik Mt., Lovćen Mt., Morača Gorge, Orjen Mt., Rumija Mt., and Skadar Lake), and Mediterranean (the Adriatic coast of Montenegro, from the Albanian to the Croatian border). Site selection was planned to ensure the representation of most of Montenegro’s geomorphological units, regardless to the intensity of previous floristic investigations in those areas. The fieldwork covered a range of habitat types, including various grasslands, shrublands, and forest ecosystems, along with their respective degradation stages under differing levels of anthropogenic influence. Additionally, to integrate these findings, an extensive revision of the herbarium collection of the Orchidaceae family from the BUNS Herbarium [72] was conducted.
Taxon identification was done according to the relevant orchidological literature [12,15,16,23,30,62]. Nomenclature generally follows the Plants of the World Online database [37]. For taxonomically problematic taxa (E. helleborine subsp. distans, O. archipelagi, O. grammica, and P. albida subsp. tricuspis), for which there is no broader consensus regarding their status, the nomenclature was determined based on the previously mentioned references. One specimen of each taxon, and only the above-ground part of the plant, was collected and deposited in the Herbarium BUNS. This collection was performed under permission granted by the Environmental Protection Agency of Montenegro for scientific research and plant material sampling (permission no. 02-UPI-914/6).
Geographic coordinates and altitude were recorded using the Garmin eTrex 20x handheld GPS device in WGS84 format. The geographical regionalization of Montenegro is given according to Marković [73] and Stešević and Caković [2]. The habitat type was classified according to the EUNIS habitat classification [74]. The geological substrate was assessed using Montenegro’s 1:200,000 scale geological map [75] and direct observations in the field. A direct census of orchids presented at the locality was conducted for the newly recorded taxa. In cases where additional orchid species already documented in Montenegro were observed at the same site, a rough estimate of their population size was made in the immediate vicinity of the newly recorded taxon. During this process, the ramet was used as the unit of account [76]. The regional conservation status of the newly documented orchid taxa in Montenegro was not assessed, as only one locality was registered for each. Additional field research is needed to understand better their actual distribution in Montenegro.

3. Results

Field research conducted in the period from 2021 to 2023 in Montenegro, and the revisions of herbarium material from the BUNS Herbarium led to the registration of ten new orchid taxa in the country (Figure 1 and Table A1). Three species, three subspecies, one variety, and one intergeneric hybrid (×Serapicamptis rousii (Du Puy) J. M. H. Shaw), as well as two intrageneric hybrids (Anacamptis ×nicodemi (Cirillo ex Ten.) B. Bockand, and A. ×olida (Bréb.) H. Kretzschmar, Eccarius and H. Dietr.) are recorded. The newly recorded taxa (excluding hybrids) belong to five genera: Epipactis (E. helleborine subsp. distans and E. leptochila subsp. neglecta), Gymnadenia (G. densiflora), Neotinea (N. ustulata var. aestivalis), Ophrys (O. archipelagi and O. grammica) and Pseudorchis (P. albida subsp. tricuspis). Detailed information, including distribution, habitat and ecology, population size, and flowering time, is provided for each newly recorded taxon in Montenegro.

3.1. Species New to Montenegro

  • Gymnadenia densiflora (Wahlenb.) A. Dietr., Allg. Gartenzeit. 7(22): 170 (1839)
Distribution in Montenegro—G. densiflora (Figure S1A) has been found in Žabljak municipality, in the northern subregion of the country: Durmitor Mt., Jablan Lake, N43.16753°, E19.06601°, MGRS 34TCN42.
Habitat and ecology—In Montenegro, G. densiflora grows on limestone and dolomitic limestone under full light conditions at 1835 m asl, in a small, wet fragment of subalpine meadow (EUNIS R451 code). Alongside this orchid, hundreds of ramets of Dactylorhiza cordigera (Fr.) Soó subsp. bosniaca (Beck) Soó were also observed. Additionally, a few plants of G. rhellicani were found in drier and higher places, and a large population of G. conopsea was found at about one hundred meters.
Population size and flowering time—This species was first recorded in Montenegro on 10 July 2021, when 52 flowering ramets were observed over an area of 800 m2. At that time, majority of plants began to open their flowers.
  • Ophrys archipelagi Gölz and H. R. Reinhard, Mitteilungsbl. Arbeitskreis Heimische Orchid. Baden-Württemberg 18(4): 731 (1986)
Distribution in Montenegro—O. archipelagi (Figure 2) has been found on the territory that belongs to the capital city of Podgorica in the Submediterranean subregion of the country: Morača Gorge, Duga, near the bridge across the Morača River, N42.54113°, E19.33718°, MGRS 34TCN63.
Habitat and ecology—O. archipelagi was found along the edges of the country road, and in thinned parts of a highly degraded Quercus trojana Webb woods (EUNIS T1981 code). In addition to this species, other shrubs were present on the site: Acer monspessulanum L., Carpinus orientalis Mill., Fraxinus ornus L., Juniperus oxycedrus L., Paliurus spina-christi Mill., Phillyrea latifolia L., and Punica granatum L. Also, a population of over 200 ramets of another orchid—Anacamptis papilionacea (L.) R. M. Bateman, Pridgeon and M. W. Chase, was observed. The population of O. archipelagi was recorded on limestone, under full light conditions, or in semi-shadow at 107 m asl. Construction activities were observed in the immediate vicinity of the locality.
Population size and flowering time—The species was registered for the first time in Montenegro on 11 April 2021, when 50 flowering ramets were recorded on an area of 2800 m2. Upon revisiting the locality on 18 April 2023, a significantly higher number of flowering ramets—107—was counted, along with the presence of rosettes of immature plants.
  • Ophrys grammica (B. Willing and E. Willing) Devillers-Tersch. and Devillers, Naturalistes Belges 72(3): 101 (1991)
Distribution in Montenegro—O. grammica (Figure 3) has been found in Cetinje municipality in the Submediterranean subregion of the country: Kopitnik Mt., Velestovo, near the road to Čevo, N42.52758°, E18.94766°, MGRS 34TCN31.
Habitat and ecology—O. grammica was recorded on eroded slopes, in the highly degraded Q. pubescens Willd. woods (EUNIS T193 code), represented by individual small trees of this oak, and solitary trees of Fraxinus ornus L. and Prunus mahaleb L. The substrate was predominantly rocky, with numerous exposed rock outcrops. Another orchid, Anacamptis morio (L.) R. M. Bateman, Pridgeon and M. W. Chase subsp. caucasica (K. Koch) H. Kretzschmar, Eccarius and H. Dietr., was also found in this locality, with its population exceeding one hundred ramets. O. grammica grew on limestone, under full light conditions, or in semi-shadow at 830 m asl.
Population size and flowering time—The first record for O. grammica in Montenegro is from 30 April 2022, when 60 flowering ramets were registered in an area of 1480 m2. At that time, most of them were at the beginning of flowering, with only one to three flowers open.

3.2. Subspecies New to Montenegro

  • Epipactis helleborine (L.) Crantz subsp. distans (Arv.-Touv.) R. Engel and P. Quentin, Orchidophile (Asnières) 124: 205 (1996)
Distribution in Montenegro—E. helleborine subsp. distans (Figure 4) has been recorded in Bijelo Polje municapility, in the northern subregion of the country: Stožer Mt., Ponikvice, N43.14202°, E19.51986°, MGRS 34TCN79.
Habitat and ecology—This orchid was observed on the edge of a mountain hay meadow (EUNIS R23 code), fragmentary surrounded by Picea abies (L.) H. Karst. forests. The meadow is still maintained by mowing, although the development of spruce saplings has been observed in some of its parts. Ramets of E. helleborine subsp. distans were recorded on limestone, under full sunlight at 1226 m asl.
Population size and flowering time—E. helleborine subsp. distans was recorded for the first time in Montenegro on 27 July 2022. During this observation, six ramets were found in four separate groups across an area of about 900 m2, with distances among the groups ranging from two to ten meters. All registered ramets were at the end of flowering.
  • Epipactis leptochila (Godfery) Godfery subsp. neglecta Kümpel, Mitt. Arbeitskreises Heimische Orchideen 11: 30 (1982)
Distribution in Montenegro—E. leptochila subsp. neglecta (Figure 5) has been found in Gusinje municapility, in the eastern subregion of the country: Prokletije Mts., Vusanje, Savino Oko, N42.51170°, E19.83481°, MGRS 34TDN04.
Habitat and ecology—E. leptochila subsp. neglecta was discovered in a subalpine stand of Fagus sylvatica L. (EUNIS T17A3 code), where this tree species is almost monodominant, creating deep shade, while the herbaceous layer of the forest is highly sparse. The small population of E. leptochila subsp. neglecta was recorded on limestone and dolomitic limestone in deep shade at 1036 m asl.
Population size and flowering time—Only five ramets of E. leptochila subsp. neglecta were found on 22 July 2022 within an area of 30 m2. Among them, only two plants were flowering, while the other three were limited to vegetative structures, developed leaves, and stems only.
  • Pseudorchis albida (L.) Á. Löve and D. Löve subsp. tricuspis (Beck) E. Klein, Phyton 40(1): 143 (2000)
Distribution in Montenegro—P. albida subsp. tricuspis (Figure S1C) has been found on the territory that belongs to the capital city of Podgorica in the eastern subregion of the country: Kučke Mts., Katun Ljakovića, N42.58190°, E19.59019°, MGRS 34TCN84.
Habitat and ecology—The herbarium label provides little information, only that the plant grew on limestone at 1764 m asl. Based on personal communication with the collector (J. Lazarević), the plants grew in full light, in a meadow that was wet in some parts.
Population size and flowering time—The precise size of the population of this subspecies on Kučke Mts. is unknown, except that there were dozens of them at the site (J. Lazarević, pers. comm.). The herbarium material shows that the plant was in full bloom when collected on 4 July 2021.

3.3. Variety New to Montenegro

  • Neotinea ustulata (L.) R. M. Bateman, Pridgeon and M. W. Chase var. aestivalis (Kümpel) Tali, M. F. Fay and R. M. Bateman, Biol. J. Linn. Soc. 87(1): 23 (2006)
Distribution in Montenegro—N. ustulata var. aestivalis (Figure S1B) has been found in Gusinje municapility, in the eastern subregion of the country: Prokletije Mts., Ćaf Bora, MGRS 34TDN09.
Habitat and ecology—The herbarium label for a specimen of N. ustulata var. aestivalis provides limited information about its habitat, indicating only that this orchid was found in meadows.
Population size and flowering time—The herbarium label attached to the plant does not provide any information about the population size of this orchid. The specimen was in full bloom when it was collected on 23 July 2014.

3.4. Hybrids New to Montenegro

  • Anacamptis ×nicodemi (Cirillo ex Ten.) B. Bock, Bull. Soc. Bot. Centre-Ouest 42: 267 (2012)
Hybrid formula: A. morio (L.) R. M. Bateman, Pridgeon and M. W. Chase × A. papilionacea (L.) R. M. Bateman, Pridgeon and M. W. Chase
Distribution in Montenegro—A. ×nicodemi (Figure S2A) has been recorded in Bar municipality, in the Mediterranean subregion of the country: Krute-Velje Selo, N42.02357°, E19.21719°, MGRS 34TCM52.
Habitat and ecology—This hybrid was discovered in an enclosed pasture (EUNIS V321 code) between Krute and Velje Selo villages. Specimens of A. ×nicodemi grew on limestone, in full light at 205 m asl. At this locality, approximately 200 ramets of the first parent species (A. papilionacea) were also recorded, while the second parent species (A. morio) was observed in the neighboring pasture, about 50 m away.
Population size and flowering time—This hybrid was recorded for the first time on 7 April 2021, when five flowering ramets were observed on an area of 1050 m2.
  • Anacamptis ×olida (Bréb.) H. Kretzschmar, Eccarius and H. Dietr., Orchid Gen. Anacamptis Orchis Neotinea 428 (2007)
Hybrid formula: A. coriophora (L.) R. M. Bateman, Pridgeon and M. W. Chase × A. morio (L.) R. M. Bateman, Pridgeon and M. W. Chase
Distribution in Montenegro—A. ×olida (Figure S2B) has been found on the territory that belongs to the capital city of Podgorica in the Submediterranean subregion of the country: Kopiljsko Polje, N42.55350°, E19.24961°, MGRS 34TCN56.
Habitat and ecology—This hybrid was found in rocky grasslands with numerous suffruticose Lamiaceae shrubs (EUNIS R1D31 code), on limestone and glaciofluvial sediments, in full sunlight at 601 m asl. Alongside the parent species A. coriophora and A. morio, other orchids were also identified at this location, including A. papilionacea, Neotinea tridentata, Orchis purpurea Huds., O. quadripunctata Cirillo ex Ten., Platanthera bifolia (L.) Rich., P. chlorantha (Custer) Rchb., and Serapias bergonii E. G. Camus. The presence of cows grazing at this site has been observed.
Population size and flowering time—This orchid was recorded for the first time on 16 May 2021, when three flowering ramets were observed on an area of 250 m2.
  • ×Serapicamptis rousii (Dupuy) J. M. H. Shaw, Orchid Rev. Suppl., 113 (1264): 20 (2005)
Hybrid formula: Anacamptis laxiflora (Lam.) R. M. Bateman, Pridgeon and M. W. Chase × Serapias vomeracea (Burm. f.) Briq.
Distribution in Montenegro—×Serapicamptis rousii (Figure 6) has been recorded in Bar municipality, in the Submediterranean subregion of the country: Skadar Lake, Gornja Seoca-Marstijepovići, N42.20085°, E19.16018°, MGRS 34TCM48.
Habitat and ecology—×Serapicamptis rousii grew along the road’s edges in its cracks (EUNIS J4.2 code). Above the road, the limestone slopes were primarily populated by Salvia officinalis L., Euphorbia characias L. subsp. wulfenii (Hoppe ex W. D. J. Koch) Radcl.-Sm., Teucrium capitatum L., Satureja subspicata Bartl. ex Vis., and Micromeria juliana (L.) Benth. ex Rchb. Both parent species, A. laxiflora and S. vomeracea, were recorded in the same locality, along with two other orchid taxa—Neotinea tridentata and Ophrys scolopax Cav. subsp. cornuta (Steven) E. G. Camus. Ramets of ×Serapicamptis rousii grew on limestone, in full sunlight at 408 m asl.
Population size and flowering time—Three hybrid ramets in full bloom were recorded on 16 May 2021 on an area of 20 m2.

4. Discussion

4.1. New Species to Montenegro’s Flora

  • Gymnadenia densiflora
Gymnadenia densiflora is native to Eurasia, with a wide distribution across Europe. It can be found in the north, up to Scotland, and in southern Norway, extending southward into the Iberian, Apennine, and the Balkan Peninsulas. Its range stretches from the Atlantic coast in the west to southwestern Siberia, and the Caucasus in the east [30,37]. The closest confirmed locations for this species are in North Macedonia in the valley of the River Radika and on Korab Mt. [77], approximately 200 km away from Durmitor Mt. in Montenegro. Other notable locations include Jabučko Ravnište peatland on Stara Planina Mt. in eastern Serbia [78], which is about 285 km away, and several sites in Lika-Senj County, Croatia [34], located 320 km away. In addition to these records, this species has been recorded within the Balkan Peninsula in Slovenia [25], Romania [79], and Bulgaria [80,81], while in Bosnia and Herzegovina, there is a single unconfirmed record from 1877 [27]. Gymnadenia densiflora primarily inhabits wet habitats—alkaline to neutral marshes, fens with high content of carbonates, springs on calcites, and wet meadows, but it can also be found on dune slacks and chalk downs, from sea level to 2800 m asl [12,23,48]. In Montenegro, it was found in a small wet fragment of subalpine meadow, on limestone and dolomitic limestone, which alligns with published data so far. Besides Durmitor Mt., there are suitable habitats for this species across Montenegro, as well on nearby high mountains in Bosnia and Herzegovina (e.g., Maglić, Volujak, and Zelengora), suggesting that it might be more widespread than currently known.
The typical flowering period for G. densiflora in Europe is July and August [12,23,30], which also coincides with flowering in the territory of Montenegro. Namely, the plants from Durmitor Mt. were in the early blooming stage when they were found in the first half of July. In cases where G. densiflora lives in sympatry with related species G. conopsea, it flowers two weeks to a month later than it [30]. This was also observed on Durmitor Mt., where a population of the G. conopsea was recorded in the vicinity, which had already finished flowering and entered the fruit-bearing phase.
Gymandenia densiflora is considered Critically Endangered in the Czech Republic, Endangered in Slovakia and Ukraine, Near Threatened in the United Kingdom and Estonia, and Least Concern in Germany (Bavaria) [82]. The population of G. densiflora found in Montenegro, within the borders of the Durmitor National Park, is in good condition and does not face any immediate threats. However, there is a risk of trampling and/or picking of the plants, as many tourists and mountaineers visit this area. Additionally, the expansion of surrounding forest vegetation could lead to the overgrowth of subalpine meadows, potentially threatening the species’ habitat in the future.
  • Ophrys archipelagi
Ophrys archipelagi was initially considered endemic to the island of Korčula, located in the Adriatic Sea, south of Croatia’s Dalmatian coast [83]. However, later it was found in other locations in Croatia, from the Kvarner archipelago (Krk island), in the northwest part of the Adriatic coast, across Ivinj, the island of Čiovo, Kaštelansko Primorje, Kozjak, Drvenik to Konavle in the south of Dalmatia [34,38]. In addition to Croatia, this taxon was recorded in four regions (Abruzzo, Campania, Puglia, and Basilicata) of the southeastern part of Italy [29]. Given this disjunct range, O. archipelagi is considered an Illyrian-Apennine endemic [58]. The nearest records from our finding are in the region of Konavle [34] in the southeastern part of Croatia, 80 km away. This taxon has been recorded in thermophilic Mediterranean habitat types, in sunny or semi-shady places on Mediterranean grasslands on limestone, rare mixed evergreen forests, and maquis of holm oak and garigue, from the sea level to 680 m asl. Also, it inhabits artificial habitats like olive groves and roadsides [12,38,58,84]. Habitat characteristics of O. archipelagi in Montenegro correspond to those in other parts of its range. It was found in a highly degraded Quercus trojana woods, which is a fragment of habitat of priority importance for the European Union (Natura 2000: 9250 Quercus trojana woods) [85]. Given that suitable habitats for O. archipelagi are found along the entire Adriatic coast of Montenegro, it can be assumed that this species has a broader distribution in this country.
During both visits (2021 and 2023) to the Duga locality, most of the plants were at the peak of flowering, which aligns with literature reports, both for Croatia [38,84] and for the entire distribution area [12], where it is noted to bloom from March to April.
Ophrys archipelagi is not listed in the Red Lists of the Flora of Italy [86,87], but in Croatia, it is a strictly protected species [88] and is listed as DD in the Red Book of the Flora of Croatia [57]. At the only site recorded so far in Montenegro, the habitat itself is highly degraded, and the population of this orchid is threatened by the immediate danger of the construction works taking place in one part of the locality. For this reason, it is necessary to monitor the population and its dynamics constantly.
  • Ophrys grammica
For a long time, O. grammica was considered endemic to a small area of northwestern Greece (Kastoria, Ioannina, and Grevena prefectures). However, it has been recorded throughout the country, from Crete in the south to the Peloponnese, the Ionian islands, and Epirus. Individual findings have also been documented in all continental parts of Greece, with a greater concentration of sites in the northern mountain ranges [12,16,62]. In addition to Greece, O. grammica has recently been identified in several locations in the Svrljiške Planine Mts. in eastern Serbia [89], approximately 300 km away from our finding in Montenegro. This orchid inhabits various alkaline to slightly acidic geological substrates. It develops in full sun to shade, in eroded areas on slopes, grasslands, maquis, forest edges, and open woodlands, from sea level to 1400 m asl [12,16,62]. Ophrys grammica was recorded on the eroded slopes in the highly degraded Q. pubescens woods, therefore, the habitat preferences of this species in Montenegro correspond with those reported in the literature. Such habitat types are also present in the surrounding countries and given that the records from Serbia and Montenegro are recent, it is expected that this taxon will be recorded in new localities within these countries, as well as in neighboring ones, such as Albania and North Macedonia.
This relatively late-blooming Ophrys species typically flowers from April to June in Greece [62]. Consistent with this is the first record for O. grammica in Montenegro from the end of April, when most of the ramets were at the beginning of flowering.
Ophrys grammica is not protected in Greece and Serbia [89]. A potential threat to this newly registered species in Montenegro is tree logging by the local population, even though it is a highly degraded stand. This activity can cause mechanical damage to the orchids, particularly those growing on slopes alongside smaller trees of Q. pubescens.

4.2. New Subspecies to Montenegroˈs Flora

  • Epipactis helleborine subsp. distans
Epipactis helleborine subsp. distans is a European taxon. Its range extends from northeastern Spain through southern France and northern Italy, continuing into central Europe and southward to the Balkans and northern Greece. Isolated populations exist in northern Germany, Poland, the Baltic states, southern Sweden, and Crimea [24,25,90]. In the Balkan Peninsula, this subspecies has been published for Slovenia [25,26], Croatia [90], Bosnia and Herzegovina [27], Serbia [28], Romania [79], Bulgaria [80], and Greece [16]. The nearest known location of this orchid is on Kamena Gora Mt. in southwestern Serbia [28], approximately 15 km from Ponikvice. This proximity suggests that they are likely part of a continuous population. In addition to Serbian site, the nearest records of E. helleborine subsp. distans from Montenegro are located approximately 150 km away in Bosnia and Herzegovina [27], 240 km in Croatia [90], and 330 km in Greece [16]. Epipactis helleborine subsp. distans occurs in various habitats, including meadows, shrubby xeric slopes, forest clearings, light thermophilic forests, pine forests, and edges of different types of forests [12,16,23,25,91]. It grows in full sun to semi-shade, on moderately wet or dry soils, on limestone, from 800 to 2200 m asl [12,35]. In Montenegro, the habitat of E. helleborine subsp. distans, a mountain hay meadow, aligns with the ecological characteristics found in other regions. Suitable habitats exist in many other areas throughout Montenegro, indicating that this subspecies may be more widely distributed than currently documented. Also, considering that it was found in most of the countries of the Balkan Peninsula, its presence can probably be expected in Albania and North Macedonia as well.
Ramets of E. helleborine subsp. distans recorded in Montenegro were at the end of their flowering phase, corresponding with the typical blooming period for this orchid, which occurs in June and July. Although all ramets had already finished flowering, the identification of this subspecies is certain. Considering all known Epipactis taxa in the Balkans and their vegetative morphology, there is no doubt. The only possible confusion can be made with the typical subspecies, but E. helleborine subsp. distans has broadly elliptic leaves, which are shorter—up to twice the length of internodes, compared to up to seven times in the typical subspecies. It also has a smaller overall leaf length and a higher plant height-to-stem diameter ratio below the inflorescence. If E. helleborine subsp. distans grows in sympatry with the type subspecies, which was not the case at the Ponikvice locality, flowering time shifts two to four weeks earlier [12,16,23,25].
According to the IUCN categorization, Epipactis helleborine subsp. distans is considered Endangered in Slovakia [23], while the E. helleborine is Extinct in Cyprus, and Least Concern in 22 European countries [82]. Besides this, E. helleborine is listed in the European Red List of Vascular Plants [92] as LC. The locality in Montenegro, where E. helleborine subsp. distans was found is not a part of any protected natural areas. It was discovered at the edge of a mountain hay meadow, regularly mowed using traditional methods, which helps maintain a habitat that supports the survival of this and other orchid species. However, if traditional mowing were to cease and the meadow undergoes succession into forest vegetation, there is a risk of population extinction.
  • Epipactis leptochila subsp. neglecta
Epipactis leptochila subsp. neglecta inhabits the area of temperate and southern Europe, from southern England and Denmark, across central Europe, northeastern Italy, and the Balkans, extending to the mountains of northern Greece [12,16]. In the Balkan Peninsula, it has been recorded in Slovenia [25], Croatia [34], Serbia [28], Greece [93], and Romania [94]. The species E. leptochila was registered in Bulgaria [95], while in Bosnia and Herzegovina, there is unverified literature data regarding E. leptochila subsp. leptochila from 1891, which lacks herbarium material for confirmation [27]. The nearest known populations of E. leptochila subsp. neglecta are located in southwestern Serbia on the Jadovnik Mt. [35], approximately 90 km away from our finding in Montenegro. Also, several more localities from southwestern Serbia where this orchid was recorded are 100 km away from Savino Oko. In addition to the findings from Serbia, the closest known occurrences of E. leptochila subsp. neglecta are located in Greece [16], and in Croatia [34] about 310 km and 320 km away, respectively. This orchid grows in medium to deep shade, in beech and oak-hornbeam woodlands and other mesophilous deciduous forests [12,30]. It is also found in forests of Pinus nigra subsp. nigra and their edges in Greece [16] and in mixed forests of beech and conifers (spruce, fir) in Serbia [35]. Epipactis leptochila subsp. neglecta mainly grows on moderately moist soils, dominantly on calcareous substrates [12,30], though it can also be found on silicates [36] at altitudes up to 1500 m asl [12,30]. In Montenegro, it was found in subalpine Fagus forests, on limestone and dolomitic limestone in deep shade, which aligns with the ecological preferences of this subspecies throughout its range.
Epipactis leptochila subsp. neglecta typically blooms during June and July [25,38], less often in August, as recorded in Slovakia [23]. Similarly, ramets of this orchid found in Montenegro in July were in bloom.
Epipactis leptochila subsp. neglecta is considered Endangered in Slovakia [23], while the IUCN status of E. leptochila in European countries is: Critically Endangered in the Czech Republic, Endangered in Slovakia, Austria, Germany (Bavaria), and Greece, Vulnerable in Georgia, Denmark and Bulgaria, Near Threatened in Switzerland and Hungary, Rare in Slovenia and Luxembourg, Data Deficient in the United Kingdom, and Least Concern in Spain, France, the Netherlands, and Croatia [82]. Additionally, this species is listed in the European Red List of Vascular Plants as Least Concern [92]. The locality Savino Oko is within the borders of the Prokletije National Park, where this orchid grows in pristine habitat. Although this area attracts many tourists which poses a risk of trampling of the plants, the likelihood of such damage remains minimal because the orchids are located on a slope quite away from the main footpath, making them less accessible for visitors.
  • Pseudorchis albida subsp. tricuspis
The species P. albida sensu lato (including subsp. alba, subsp. straminea, and subsp. tricuspis) has a circumpolar distribution. Its range includes the northeastern parts of North America, Greenland, Iceland, and Europe, in the south to the mountains of the northern part of the Iberian, the Apennine, and the Balkan Peninsulas to the north of Greece, and east to Kamchatka in Asia. In southern Europe, it occurs only in mountains [12,23,66]. Apart from the mountains of the Balkan Peninsula [25,70,78], P. albida subsp. tricuspis inhabits the Swiss, Italian, and Austrian Alps, as well as Tatra Mts. and Eastern Carpathians Mts. [66]. In the area of the Balkan Peninsula, P. albida subsp. tricuspis was listed in only three countries, such as Slovenia [25], Serbia [78], and Romania [70], while P. albida was registered in the remaining countries—Croatia [58], Bosnia and Herzegovina [27], Albania [96], North Macedonia [97], Bulgaria [81], and Greece [16]. According to some authors [69], P. albida subsp. tricuspis is also present in Greece. The nearest known location of this orchid is in eastern Serbia on Stara Planina Mt. [78], approximately 270 km away from Kučke Mts.
Pseudorchis albida is a species that inhabits mountain meadows, dwarf pine meadows, montane fens, alpine grasslands, and open woodlands [12,23]. In Greece, the species is found exclusively in the forests of Pinus heldreichii [16]. It occurs predominantly in full sun, on acidic to alkaline substrates, up to 2700 m asl [12]. Pseudorchis albida subsp. tricuspis inhabits limestone habitats exclusively [12,68]. It has been recorded in various habitats in Romania, including wet meadows, grasslands, pastures, forest edges, rocky slopes, and alpine pastures [70]. In Serbia, this taxon lives in subalpine meadows [78]. Ecological data for the first finding of P. albida subsp. tricuspis in Montenegro is very limited. Based on the given site’s altitude (1764 m asl) and satellite imagery of the mountain, it is likely that the habitat of this taxon is a subalpine meadow potentially maintained by cattle grazing, as traditional katuns for cattle herding exist in the area. Given that this subspecies is often overlooked because specimens are frequently identified and published simply as P. albida, and that it is restricted to limestone habitats, which are widespread throughout Montenegro and the Balkans, its actual range may be significantly broader than currently known. A more thorough review of herbarium collections from Montenegro and neighboring countries will likely provide further confirmation of this.
In Europe, P. albida typically flowers from May to August [12,16,23], and the finding from Montenegro aligns with this timeframe.
The species P. albida is facing a general decline in population size throughout its range [71]. The primary factors contributing to its decline include habitat destruction, forestry, agricultural improvement [67], and habitat succession. Additionally, it can be assumed that climate change plays a significant role in this, as this species predominantly inhabits high mountain areas throughout its range and lowland regions of Northern Europe, which are especially vulnerable to such environmental changes [71]. The IUCN status of P. albida in European countries is as follows: Regionally Extinct in Belgium and the Netherlands, Critically Endangered in Denmark, Endangered in Ukraine, Sweden, the Czech Republic, Slovakia, and Germany (Bavaria), Vulnerable in Norway, the United Kingdom, and Bulgaria, Least Concern in Switzerland, Austria, Slovenia, and Greece, and Data Deficient in Croatia [37,82]. Evaluated as Least Concern, P. albida is included in the European Red List of Vascular Plants [92]. The presence of P. albida subsp. tricuspis in Montenegro has been recorded based on herbarium material. Therefore, the habitat condition and the potential threats it faces are still unknown.

4.3. New Variety to Montenegroˈs Flora

  • Neotinea ustulata var. aestivalis
Neotinea ustulata is an Euro-Siberian species [12,30], but var. aestivalis can be found only in Europe. Neotinea ustulata var. aestivalis has been recorded from England in the west to Denmark and the Baltic countries in the north, covering the entire central European region, as well as the Apennine and Balkan Peninsulas in the south [30,98]. Within the Balkan Peninsula, N. ustulata is present in all countries [37]. At the same time, var. aestivalis has been reported from Slovenia [25], Croatia [99], Serbia [35], Romania [100], Bulgaria [98], and Greece [16]. The closest known occurrences of N. ustulata var. aestivalis are located in southwestern Serbia [35], approximately 60 and 100 km away from our finding in Montenegro. Neotinea ustulata var. aestivalis is found in dry and semi-dry grasslands, sparse meadows, montane meadows, forest edges, and occasionally in coniferous and deciduous forests. This species primarily grows in calcareous soils at altitudes ranging from sea level to 2000 m asl [23,30]. The information on the locality and habitat of the specimen from Montenegro provided on the herbarium label is minimal. However, based on the locality’s name, we can partially reconstruct the habitat preferences of this taxon in Montenegro. Ćaf Bora is situated at an elevation of 1700 to 1800 m, with a geological base composed of sandstones, conglomerates, limestone conglomerates, limestone, siltstones, marls, sandy and marly limestone, and calcarenites [75]. According to the altitude of the locality and the information on the herbarium label that it was found in “meadows,” it is likely that those are subalpine meadows. Based on the available details, the habitat of this orchid in Montenegro appears to be consistent with its habitats found in other regions of Europe.
Given that the herbarium specimen was in bloom at the time of collection, this timing aligns with the typical flowering period across Europe, which spans from June to August [25,30,35]. The type variety and var. aestivalis exhibit different flowering periods. For instance, in Britain, it has been observed that N. ustulata var. ustulata blooms from mid-May to mid-June, while var. aestivalis flowers from the beginning of July and during August [101]. Generally, at similar latitudes and altitudes, var. aestivalis tends to bloom later than var. ustulata [30]. Based on the available material, whether the type variety also occurs at the Ćaf Bora locality is not known.
In central Europe, N. ustulata is experiencing a significant decline due to agricultural intensification and the abandonment of traditional meadow mowing cycles [15]. In terms of range reduction, it represents one of the most endangered orchids in this area [102,103]. Neotinea ustulata is Regionally Extinct in the Netherlands and Luxembourg, Critically Endangered in Lithuania, Ukraine, Belarus, the Czech Republic, and Germany (Bavaria), Endangered in Estonia, the United Kingdom, and Slovakia, Vulnerable in Liechtenstein, Georgia, Denmark, Bulgaria, Slovenia, Croatia, and Austria. Also, it is Near Threatened in Switzerland and Hungary and in the IUCN category of Least Concern in France and Greece [82,104]. This species is listed in the European Red List of Vascular Plants [92] as LC. The locality of Ćaf Bora is within the Prokletije National Park, where Montenegrin legislation ensures its protection. No known threats or endangering factors affect this taxon or its habitat at this location.

4.4. New Hybrids to Montenegroˈs Flora

  • Anacamptis ×nicodemi
Hybridization between A. morio and A. papilionacea is widespread, often resulting in hybrid swarms that can outnumber the populations of parent species in certain localities [15]. In the Balkan Peninsula, this hybrid has been recorded in several countries: Croatia [105], Serbia [78,106], Romania [107], North Macedonia [18], and Greece [69,108]. At the first registered locality in Montenegro, this taxon lived in enclosed pasture. This taxon has similar ecological preferences in other locations across the Balkan Peninsula. For example, A. ×nicodemi has been reported in Serbia within xeromesophilous grassland communities, in full light, on siliceous and carbonate sediments at elevations ranging from 332 to 639 m asl [106]. In North Macedonia, it was also registered in full sun within thermophilous grassland (ass. Astragalo-Helianthemetum marmorei Matevski, Čarni, Ċušterevska, Kostadinovski and Mucina) on dolomite limestone at 1008 m asl [18], while in Croatia, it grew in olive groves and mesophilic hay meadows [109].
  • Anacamptis ×olida
Hybrids between A. morio and A. coriophora are rare [12]. A. ×olida is one of the most morphologically variable hybrids within the genus Anacamptis due to the high variability of the parental taxa [15]. In the Balkan Peninsula, this hybrid has been recorded in several countries, including Croatia [105], Albania, Romania [110], and Bulgaria [111]. The only locality in Montenegro where this hybrid has been recorded, Kopiljsko Polje, supports cattle grazing, which may contribute to the high diversity of orchids by maintaining the ecosystem in its current state and preventing succession to less suitable shrubby and forest vegetation that could be detrimental to the growth of certain species.
  • ×Serapicamptis rousii
Besides Montenegro, on the Balkan Peninsula, ×Serapicamptis rousii was also recorded in Croatia [109] and Greece [69,108]. Parental taxa of this hybrid are of the Mediterranean range and inhabit diverse habitats, although A. laxiflora prefers moist ones [12]. Thus, the finding of this hybrid from Croatia originates from a wet Mediterranean pasture [109]. In Montenegro, ×Serapicamptis rousii was registered in a seemingly very different habitat—along the edges of roads, specifically in the cracks of the road. During the field research, they were very wet, likely due to water runoff from the surrounding higher terrain, and was inhabited by different bryophytes. So, it appears that this hybrid from Montenegro has ecological preferences similar to those from Croatia regarding substrate moisture and geological substrate (limestone).

5. Conclusions

This study presents the first records of ten orchid taxa in Montenegro, which account for approximately 10% of the country’s total orchid diversity. The findings highlight Montenegro’s ecological diversity, as newly recorded taxa were found in both relatively undisturbed ecosystems and human-altered habitats. The limited population sizes observed for several taxa further reinforce the urgency of systematic monitoring and conservation actions to mitigate threats and ensure their survival. This study demonstrates that Montenegro remains botanically underexplored and has considerable potential for future floristic discoveries and ecological research.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17050337/s1, Figure S1: Herbarium specimens of newly registered orchids in Montenegro; Figure S2: Herbarium specimens of newly registered orchids in Montenegro.

Author Contributions

Conceptualization and methodology, B.Đ.R.; investigation, B.Đ.R., B.S.B., and G.T.A.; data curation, B.Đ.R. and J.M.P.; writing—original draft preparation, B.Đ.R.; writing—review and editing, B.Đ.R., J.M.P., B.S.B., and G.T.A.; visualization, B.Đ.R. and J.M.P. All authors have read and agreed to the published version of the manuscript.

Funding

The authors gratefully acknowledge the financial support of the Ministry of Science, Technological Development and Innovation of the Republic of Serbia (Grants No. 451-03-137/2025-03/200125 and 451-03-136/2025-03/200125).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Data is contained within the article or Supplementary Material.

Acknowledgments

Authors would like to thank Miloš Ilić for creating a map used in this paper.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A

Table A1. Herbarium specimens examined.
Table A1. Herbarium specimens examined.
TaxaHerbarium Labels
Gymnadenia densifloraDurmitor Mt., Jablan Lake, N43.16753°, E19.06601°, MGRS 34TCN42, 1835 m asl, subalpine meadow, limestones, 10 July 2021, coll. et det. B. Radak (BUNS 22242)
Ophrys archipelagiMorača Gorge, Duga, near the bridge across the Morača river, N42.54113°, E19.33718°, MGRS 34TCN63, 107 m asl, highly degraded Quercus trojana woods, limestone, 11 April 2021, coll. B. Radak, det. B. Radak, J. Peškanov (BUNS 22238)
Ophrys grammicaKopitnik Mt., Velestovo, near the road to Čevo, N42.52758°, E18.94766°, MGRS 34TCN31, highly degraded Quercus pubescens woods, limestone, 30 April 2022, coll. B. Radak, det. B. Radak, J. Peškanov (BUNS 22239)
Epipactis helleborine subsp. distansStožer Mt., Ponikvice, N43.14202°, E19.51986°, MGRS 34TCN79, 1226 m asl, mountain hay meadow, limestones, 27 July 2022, coll. et det. B. Radak (BUNS 22246)
Epipactis leptochila subsp. neglectaProkletije Mts., Vusanje, Savino Oko, N42.51170°, E19.83481°, MGRS 34TDN04, 1036 m asl, subalpine Fagus sylvatica stand, limestones and dolomitic limestones, 22 July 2022, coll. et det. B. Radak (BUNS 22245)
Pseudorchis albida subsp. tricuspisKučke Mts., Katun Ljakovića, N42.58190°, E19.59019°, MGRS 34TCN84, 1764 m asl., limestones, 04 July 2021, coll. J. Lazarevic, det. B. Radak (BUNS 22244)
Neotinea ustulata var. aestivalisProkletije Mts., Ćaf Bora, MGRS 34TDN09, meadows, 23 July 2014, coll. S. Ivković, L. Horvat, det. B. Radak (BUNS 22243)
Anacamptis ×nicodemiKrute-Velje Selo, N42.02357°, E19.21719°, MGRS 34TCM52, 205 m asl, pasture, limestones, 07 April 2021, coll. B. Radak, J. Beloica, det. B. Radak, J. Peškanov (BUNS 22240)
Anacamptis ×olidaKopiljsko polje, N42.55350°, E19.24961°, MGRS 34TCN56, 601 m asl, rocky grasslands, limestones, 16 May 2021, coll. B. Bokić, M. Rat, det. B. Radak, J. Peškanov (BUNS 22241)
×Serapicamptis rousiiSkadar Lake, Gornja Seoca-Marstijepovići, N42.20085°, E19.16018°, MGRS 34TCM48, 408 m asl, road cracks, limestones, 16 May 2021, coll. B. Radak, det. B. Radak, J. Peškanov (BUNS 22247)

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Figure 1. Distribution of newly registered orchid taxa in Montenegro.
Figure 1. Distribution of newly registered orchid taxa in Montenegro.
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Figure 2. Ophrys archipelagi. Photographs by B. Radak (18 April 2023, Montenegro, Morača Gorge, Duga).
Figure 2. Ophrys archipelagi. Photographs by B. Radak (18 April 2023, Montenegro, Morača Gorge, Duga).
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Figure 3. Ophrys grammica. Photographs by B. Radak (30 April 2022, Montenegro, Kopitnik Mt., Velestovo).
Figure 3. Ophrys grammica. Photographs by B. Radak (30 April 2022, Montenegro, Kopitnik Mt., Velestovo).
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Figure 4. Epipactis helleborine subsp. distans. Photographs by B. Radak (27 July 2022, Montenegro, Stožer Mt., Ponikvice).
Figure 4. Epipactis helleborine subsp. distans. Photographs by B. Radak (27 July 2022, Montenegro, Stožer Mt., Ponikvice).
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Figure 5. Epipactis leptochila subsp. neglecta. Photograph by B. Radak (22 July 2022, Montenegro, Prokletije Mts., Vusanje, Savino Oko).
Figure 5. Epipactis leptochila subsp. neglecta. Photograph by B. Radak (22 July 2022, Montenegro, Prokletije Mts., Vusanje, Savino Oko).
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Figure 6. ×Serapicamptis rousii. (A) Habitat. (B) Inflorescence. Photographs by B. Radak (16 May 2021, Montenegro, Skadar Lake, Gornja Seoca-Marstijepovići).
Figure 6. ×Serapicamptis rousii. (A) Habitat. (B) Inflorescence. Photographs by B. Radak (16 May 2021, Montenegro, Skadar Lake, Gornja Seoca-Marstijepovići).
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Radak, B.Đ.; Peškanov, J.M.; Bokić, B.S.; Anačkov, G.T. Exploring the Orchid Flora of Montenegro: Ten Newly Identified Taxa. Diversity 2025, 17, 337. https://doi.org/10.3390/d17050337

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Radak BĐ, Peškanov JM, Bokić BS, Anačkov GT. Exploring the Orchid Flora of Montenegro: Ten Newly Identified Taxa. Diversity. 2025; 17(5):337. https://doi.org/10.3390/d17050337

Chicago/Turabian Style

Radak, Boris Đoka, Jovan Milan Peškanov, Bojana Slavenko Bokić, and Goran Timča Anačkov. 2025. "Exploring the Orchid Flora of Montenegro: Ten Newly Identified Taxa" Diversity 17, no. 5: 337. https://doi.org/10.3390/d17050337

APA Style

Radak, B. Đ., Peškanov, J. M., Bokić, B. S., & Anačkov, G. T. (2025). Exploring the Orchid Flora of Montenegro: Ten Newly Identified Taxa. Diversity, 17(5), 337. https://doi.org/10.3390/d17050337

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