Rubus dolichocarpus Juz.

Salahi, Razieh, Afsharzadeh, Saeed & Sochor, Michal, 2025, Taxonomic and nomenclatural revision of Rubus L. (Rosaceae) in Iran, Phytotaxa 700 (1), pp. 1-17 : 6-8

Rubus dolichocarpus Juz. in Trudy Prikl. Bot. Selekts. 14(3): 159 (1925). Type: Georgia, prope opp. Mtzkhet, in silva frondosa, 20 August 1924, Juzepczuk 274 (LE, holotype —image! in Juzepczuk 1925).

= R. charadzae Sanadze in Trudy Tbilissk. Univ. 42: 84 (1951), syn. nov. Type: Kachethia, in viciniis p. Lagodekhi, in faucibus Koczla, 27 July 1947, Sanadze s.n. (TBI, holotype!).

= R. gaubae Rech.f. in Ann. Naturhist. Mus. Wien 53: 345 (1942, publ. 1943), syn. nov. Type: Gilan: Waldlichtungen zwischen Resht und Lahidjan, 14 May 1937, Rechinger 69 (W, holotype —image!).

= R. juzepczukii Sanadze in Trudy Tbilissk. Univ. 42: 89 (1951), syn. nov. Type: Kachethia, in faucibus Batzara, 15 July 1947, Sanadze s.n. (TBI, holotype!).

= R. kacheticus Sanadze in Trudy Tbilissk. Univ. 42: 92 (1951), syn. nov. Type: Kachethia, in viciniis p. Lagodekhi, in faucibus Koczla, 26 July 1947, Sanadze s.n. (TBI, holotype!).

= R. ketzkhovelii Sanadze in Trudy Tbilissk. Univ. 42: 80 (1951), syn. nov. Type: Kachethia, in viciniis p. Lagodekhi, in faucibus Koczla, 30 July 1947, Sanadze s.n. (TBI, holotype!).

= R. kudagorensis Sanadze in Trudy Tbilissk. Univ. 42: 81 (1951), syn. nov. Type: Kachethia, in viciniis p. Lagodekhi, in monte Kudagora, 28 July 1947, Sanadze s.n. (TBI, holotype!).

= R. lahidjanensis Rech.f. in Ann. Naturhist. Mus. Wien 53: 345 (1942, publ. 1943), syn. nov. Type: Gilan: Waldlichtungen zwischen Resht und Lahidjan, 14 May 1937, Rechinger 59 (W; lectotype —image!, inflorescence only, designated here). Note: The type specimen cited in the protologue (holotype) consists of an inflorescence belonging to R. dolichocarpus and a primocane section from another, unidentified bramble. Because the protologue is based solely on this mixed specimen, none of its parts can be considered as “corresponding more nearly with the original description” (Art. 9.14, Turland et al. 2018). Since the identity of the primocane is unclear and its description in the protologue, for the bigger part, is not in serious conflict with the variation of R. dolichocarpus observed by us, we designate here the inflorescence as a lectotype in conformity with Art. 9.11 ( Turland et al. 2018) and recommendations of Sennikov (2016).

= R. longipetiolatus Sanadze in Trudy Tbilissk. Univ. 42: 85 (1951), syn. nov. Type: Kachethia, in viciniis p. Lagodekhi, in faucibus Koczla, 27 July 1947, Sanadze s.n. (TBI, holotype!).

Shrub, usually up to 2 m tall. Primocanes erect or arching; stems angular and slightly furrowed, ± 4–8 mm in diameter, green to vinaceous, sparsely to moderately hairy with short stellate and simple hairs, sometimes with scattered stalked glands, with 8 to 15 prickles per 50 mm of stem length arranged along stem angles; prickles straight or slightly curved, erect or slightly declining, ± 1–4 mm long and ± 1–4 mm wide at base, usually red or green-reddish with yellowish tip. Stipules linear, ± 10–22 × ca. 1 mm, hairy with short stellate and long simple hairs, sometimes with stalked glands. Leaves on primocanes palmately 5-foliolate, very sparsely hairy above with long simple hairs, white/greyish tomentose beneath with short stellate hairs; petiole ± as long as basal leaflets, sparsely hairy with short stellate hairs and long simple hairs and 10 to 20(–35) small curved prickles; terminal leaflet usually narrowly to broadly obovate, sometimes ovate, base distinctly cordate, apex acuminate, with ± 10–15(–20) mm long tip, margins almost flat, periodically dentate, incisions 1–4 mm deep, petiolule 38–41(–60)% of the length of leaflet lamina; lateral leaflets obovate or elliptic-obovate, with petiolules 12–30 mm long; basal leaflets elliptic or subobovate, with petiolules 4–7(–10) mm long (± 5.5–8.5% of lamina length). Inflorescence usually a loose and rich pyramidal panicle; inflorescence leaves ternate; inflorescence axis angular, usually hairy with dense stellate and simple longer hairs; prickles 5 to 15 per 50 mm of axis length, mostly slightly curved, 1–4 mm long, 1–5 mm wide at base, sometimes with dense stalked glands (20 to 30 per 10 mm), ± 1 mm long, sometimes without stalked glands; pedicels (2–) 5–25 mm long, tomentose, with 0–10 thin, straight to somewhat curved, 1–2 mm long prickles and 0–15 ± 1 mm long stalked glands per 10 mm. Flowers 25–30 mm in diameter; sepals greyish tomentose with short stellate and long simple hairs, with stalked glands and prickles, distinctly reflexed during and after anthesis, 5–12 mm long; petals white, much longer than sepals, 15 mm long, obovate to elliptic. Stamens much longer than styles; filaments white; anthers glabrous. Carpels glabrous. Flower receptacle densely hairy. Aggregate fruit usually large, long cylindrical, composed of many small drupelets, sometimes small and semiglobose, green turning red and finally black at maturity, shiny, juicy and sweet ( Fig. 3 ).

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.

Distribution: Hyrcanian mixed forests ecoregion in northern Iran and Azerbaijan ( Fig. 2B ); 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).

Figures

FIGURE 2. Distribution of Rubus species in Iran. A: R. caesius; B: R. dolichocarpus; C: R. glandulosus; D: R. ulmifolius.

FIGURE 3. Rubus dolichocarpus.A: habit;B: primocane leaf;C: primocane stem; D:inflorescence;E:inflorescence axis; F: infructescence with unripe fruits; G: ripe and unripe fruits; H: abaxial surface of terminal leaflet of primocane leaf; I: adaxial surface of terminal leaflet of primocane leaf.