taxonID	type	description	language	source
FB60206E7B01FFA8B7B8AAC281A3FF15.taxon	biology_ecology	Ecology: mainly along margins of forests or woodlands, mostly in partly shady situations, rarely in full sun. Rubus dolichocarpus is the most common forest Rubus species in northern Iran. Diagnostic characters: terminal leaflets of primocane leaves obovate with thin lamina, white-felted underneath with short stellate hairs only (longer hairs absent) and dull above; inflorescences usually rich, loose, (almost) leafless; petals white; aggregate fruits long cylindrical with numerous small drupelets.	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B01FFA8B7B8AAC281A3FF15.taxon	distribution	Distribution: Hyrcanian mixed forests ecoregion in northern Iran and Azerbaijan (Fig. 2 B); the distribution range further extends along the Greater Caucasus to central Georgia (Juzepczuk 1925, Zieliński 1978, Kasalkheh et al. 2024). Notes: Rubus dolichocarpus is characterized by its long, cylindrical fruits, tomentose leaves abaxially, and the presence of stalked glands (but see below). Therefore, Juzepczuk (1925) classified it under the series Radula in the Caucasus. However, the analysis of collections reveals that this species exhibits extraordinary variability in several characteristics. For instance, the fruit shape is not consistently long and cylindrical, and the presence of pubescence in the inflorescences varies considerably among specimens. Additionally, the density of stalked glands on stem and inflorescence axis ranges from absent to scattered on stem and absent to very dense in inflorescence, with many transitional forms between these extremes. Importantly, there is no significant geographical distinction (Zieliński 1978) nor genetic differentiation (Kasalkheh et al. 2024) between the glandular and eglandular forms. It has recently been established that R. dolichocarpus is diploid and reproduces sexually (Kasalkheh et al. 2024). Therefore, it should be treated taxonomically as such, that is as a morphologically and genetically variable species under biological species concept (Mayr 1942). As an ancestral species, it contributed to the evolution of many polyploids (Kasalkheh et al. 2024). Published records from Iran: This species was reported from Iran by Parsa (1948; as R. lahidjanensis and R. gaubae), Zieliński (1978), and Khatamsaz (1992).	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B03FFA7B7B8AB568189FF15.taxon	description	Shrub or shrublet, usually up to 50 cm tall. Primocanes low-arching or creeping; stems terete or slightly angular, ± (3 –) 6 – 8 mm in diameter, green, hairy with long simple and tufted hairs, distinctly glandular with stalked glands ˂ 1 – 2 mm long and (sub) sessile glands, with 15 to 40 prickles per 50 mm of stem length and many pricklets forming gradient from stalked glands through glandular and non-glandular acicles / pricklets to prickles; prickles unequal in size, slender, thin, straight, erect to slightly declining, ± 1 – 6 mm long and 1 – 5 mm wide at base, yellow or sometimes green with yellowish tip. Stipules linear, ± 12 – 15 × ca. 1 mm, hairy with long simple hairs, with stalked and sessile glands. Leaves on primocanes 3 – 5 - foliolate, distinctly hairy to touch, usually light green, moderately to densely hairy above with long simple hairs, green beneath with sparse short stellate and dense long simple hairs; petiole ± as long as lateral leaflets, sparsely hairy with short stellate hairs and long simple hairs and 15 to 40 small, thin, mostly straight, erect to declining prickles, with numerous stalked glands; terminal leaflet usually elliptic to broadly obovate, base distinctly cordate, apex acuminate or caudate, with ± 15 – 20 mm long tip, margins almost flat, periodically to almost regularly dentate, incisions ca. 1 – 3 mm deep, petiolule ± 23 – 37 % length of leaflet lamina; lateral leaflets elliptic or elliptic-ovate, with petiolules ± 5 – 10 mm long; basal leaflets obovate to elliptic, with petiolules 1 – 2 mm long (± 1.5 – 3.5 % of lamina length). Inflorescence usually poorly branched, racemose; inflorescence leaves ternate; bracts simple to trifid; inflorescence axis terete or somewhat angular, densely hairy with stellate and long simple hairs and dense stalked glands (20 to 130 per 10 mm), 1 – 1.5 mm long; prickles (5 –) 15 – 35 per 50 mm of axis length, mostly slender, straight, erect or slightly curved, ˂ 1 – 3 (– 6) mm long, ca. 1 – 1.5 mm wide at base; pedicels 5 – 25 mm long, with stellate hairs and dense long patent simple hairs, with 1 – 15 (– 26) thin, straight to somewhat curved, ca. 1 – 1.5 mm long prickles, mostly densely glandular with stalked glands. Flowers 25 – 30 mm in diameter; sepals greyish, woolly, with many stalked glands and prickles, distinctly erect during and after anthesis, 8 – 12 mm long; petals white, much longer than sepals, 10 – 15 mm long, narrowly obovate to elliptic. Stamens much longer than styles; filaments white; anthers glabrous. Carpels glabrous. Flower receptacle densely hairy. Aggregate fruit medium to large, with ± 15 – 30 drupelets, usually longer than wide, green turning red and finally black at maturity, shiny and juicy (Fig. 4).	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B03FFA7B7B8AB568189FF15.taxon	biology_ecology	Ecology: Rubus glandulosus is an exclusively forest species; it occurs mainly along the forest margins or in gaps / clearings, but also in the forest interior, from lower elevations (370 m a. s. l.) almost to the tree line in western Hyrcania (Gilan), but progressively getting restricted to high-mountain forests and avoiding alkaline / calcareous bedrocks to the east (Mazandaran, Golestan). Diagnostic characters: primocane stems rather thin, low-arching to creeping; stems, petioles and inflorescence axis and branches with dense long-stalked glands; primocanes with many transitions between stalked glands, glandular and eglandular acicles, pricklets, and prickles; prickles on primocane stems thin, slender, acicular; petals white; sepals erect.	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B03FFA7B7B8AB568189FF15.taxon	distribution	Distribution: the ecoregion of Caspian Hyrcanian mixed forests, mostly in the west (Gilan, Mazandaran), scattered to rare in the east (Golestan; Fig. 2 C). Iran represents the eastern extremity of the species distribution, which covers the Caucasus and most of southern, central, and western Europe (Sochor et al. 2024 a, b). Published records from Iran: This species was reported from Iran by Parsa (1948), Zieliński (1978; as R. hirtus agg.) and Khatamsaz (1992; as R. hirtus agg.).	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B0CFFA6B7B8ACC083D1F82B.taxon	description	= R. dalmatinus (Tratt.) Guss. in Fl. Sicul. Syn. 1: 567 (1843). Type: Dalmatia ad sepes, s. dat., Portenschlag s. n. (holotype, not found). = R. inermis Pourr. in Hist. & Mém. Acad. Roy. Sci. Toulouse 3: 326. (1788), nom. rej. Type: Spain, Barcelona, s. dat., Pourret 3168 (MAF, lectotype — image!). designated by Van de Beek in Gorteria 9: 206 (1979). = R. sanctus Schreb. in Icon. Descr. Pl. [Schreber] 15, t. 8 (1766), nom. illeg.? Type: Crete, s. dat., Schreber 43 (M, holotype — image!). = R. sanguineus Friv. in Flora (Regensb.) 18 (21): 334 (1835). Type: Macedonia, s. dat., Frivaldsky von Frivald s. n. (KIEL, lectotype — not seen), designated by Monasterio-Huelin and Weber (1996). = R. turcomanicus Freny. in Bull. Herb. Boissier 6: 209 (1906). Type: Turkmenistan / Balkan, Kisil Arwat, Karakala, ad radices montis Sundsodagh, in valle humid, 31 May 1901, Sintenis 1846 a (W, JE 00014920, syntype — image!).	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B0CFFA6B7B8ACC083D1F82B.taxon	description	Shrub, usually up to 2 m tall and often forming large thickets. Primocanes arching, sometimes climbing; stems distinctly angular with flat to slightly furrowed sides, ± 5 – 10 mm in diameter, (blueish-) green to violet-vinaceous on sides exposed to sun, moderately to densely hairy to tomentose, without stalked glands, with 3 to 8 prickles per 50 mm of stem length arranged along stem angles; prickles almost uniform, medium to large, straight to curved, sometimes slightly declining, strong and broad-based, 5 – 8 (– 10) mm long and 4 – 8 mm wide at base, ± concolourous with stem (usually greyish) with yellowish tip. Stipules linear / filiform, ± 10 – 12 × ca. 1 mm, hairy with stellate hairs and long simple hairs, without stalked glands. Leaves on primocanes palmate, 5 - foliolate, distinctly softly hairy to touch, sparsely hairy above with short stellate hairs and long simple hairs, white / greyish tomentose beneath with many short stellate and sparse long simple hairs; petiole ± as long as basal leaflets, pruinose, densely hairy to tomentose and with 4 to 10 small curved prickles; terminal leaflet usually narrowly to broadly obovate or elliptic to suborbicular, truncate and usually shortly cuspidate, base rounded or subcordate, apex shortly acuminate or submucronate or sometimes rounded, with ± 0 – 15 mm long tip, margins almost flat, periodically dentate, incisions 1 – 3 mm deep, petiolule ± 26 – 32 (– 51) % of the length of leaflet lamina; lateral leaflets obovate, sometimes suborbicular, with petiolules 10 – 15 mm long; basal leaflets obovate to elliptic, with petiolules 0 – 5 mm long (± 0 – 10 % of the lamina length). Inflorescence usually richly branched, narrowly pyramidal, racemose; inflorescence axis angular to furrowed, villous and tomentose, pruinose, hairy with dense subadpressed stellate hairs and longer patent hairs; prickles 2 to 15 per 50 mm of axis length, strong, curved, ± 1 – 4 mm long, ca. 1 – 4 mm wide at base; pedicels 3 – 25 mm long, villous, tomentose with adpressed stellate hairs and dense long patent simple hairs, with 0 – 6 (– 9) very small, straight to curved, ca. 1 – 2 mm long prickles. Flowers 20 – 25 mm in diameter; sepals greyish tomentose with short stellate and long simple hairs, distinctly reflexed during and after anthesis, 4 – 7 mm long; petals pale pink to deep magenta, rarely white, much longer than sepals, 10 – 12 mm long, broadly elliptic to suborbicular. Stamens equal or much longer than styles; filaments usually pink; anthers hairy. Carpels densely hairy. Flower receptacle glabrous. Aggregate fruit large with ± 10 – 40 drupelets, semiglobose or globose, green turning red and finally black at maturity, shiny, juicy and sweet (Fig. 5).	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B0CFFA6B7B8ACC083D1F82B.taxon	biology_ecology	Ecology: Rubus ulmifolius is the most common non-forest bramble species in Iran. It occurs mainly on roadsides, in gardens, tree plantations, pastures, ruderal and other secondary habitats, along rivers or water lines, predominantly in sunny or semi-shaded situations, rarely in forests under semi-closed canopy. Diagnostic characters: stems thick, greyish / blueish pruinose and tomentose; prickles strong, broad-based, mostly straight and erect to declining; petals usually pink to deep magenta; filaments pink; anthers hairy.	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B0CFFA6B7B8ACC083D1F82B.taxon	distribution	Distribution: widely distributed species, mostly in Hyrcanian regions in Gilan, Mazandaran, Golestan, and also in East and West Azerbaijan. Further south, it becomes progressively rarer but remains relatively common in central Iran (provinces of Tehran, Qazvin, Kermanshah, Kurdistan, Lorestan, Isfahan, Chaharmahal & Bakhtiari, Kohgiluyeh & Boyer-Ahmad, Khuzestan) and rarely occurs also in the Kerman, Fars, and Bushehr provinces (Fig. 2 D). The overall distribution of R. ulmifolius extends further east to Afghanistan and west through the Mediterranean regions of Europe and Africa to Western and Northwestern Europe and Macaronesia (Kurtto et al. 2010). Notes: The name Rubus ulmifolius has been recently conserved against R. vulgaris and R. inermis (Wilson 2024). The conservation against the so-called eastern taxon (R. sanctus sensu Monasterio-Huelin and Weber 1996) has not been approved yet. Although the nomenclatural committee in principle supported this conservation, it was not clear whether R. creticus L. (the oldest name known for the species) is a validly published name and whether R. sanctus Schreb. (the commonly used name for the eastern taxon; Kurtto et al. 2010) is legitimate. Considering the nomenclatural inclarity, as well as the available evidence for the conspecificity of the two taxa (western and eastern; Sochor et al. 2017, 2024 a), we use here the most commonly used name for the diploid species in question, that is R. ulmifolius. Published records from Iran: This species was reported from Iran by Parsa (1948; as R. anatolicus and R. dalmaticus), Gilli (1969; as R. anatolicus), Zieliński (1978; as R. anatolicus), and Khatamsaz (1992; as R. sanctus agg.).	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B0EFFA3B7B8AC3A818EFD8C.taxon	biology_ecology	Ecology: The polyploids of Rubus ser. Discolores prefer disturbed habitats in sunny situations, typically along roads, less commonly also in forest margins and along rivers or water lines. Diagnostic characters: primocane stem glabrous to sparsely hairy, not pruinose; primocane leaves leathery, dark green and glossy above, white-felted beneath; prickles strong, large, erect or slightly declining; stem and inflorescence without stalked glands; petals mostly pink.	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
FB60206E7B0EFFA3B7B8AC3A818EFD8C.taxon	distribution	Distribution: In Iran, polyploids of R. ser. Discolores occur mostly in northern regions (Fig. 7). General distribution includes most of the temperate and submediterranean regions of Europe (Kurtto et al. 2010) and through the Caucasus and Iran extends up to Central Asia (Kyrgyzstan, Tajikistan; Kasalkheh et al. 2024). Published records from Iran: Three species of this series were described from Iran, but none of them can be confirmed now as an acceptable apomictic species. Zieliński (1978) also reported on the occurrence of R. procerus P. J. Müll. ex Boulay, a species described from eastern France, but more data is needed for identification and characterization of that species at this moment.	en	Salahi, Razieh, Afsharzadeh, Saeed, Sochor, Michal (2025): Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran. Phytotaxa 700 (1): 1-17, DOI: 10.11646/phytotaxa.700.1.1, URL: https://doi.org/10.11646/phytotaxa.700.1.1
