Rhamnus spathulifolia Fischer & Meyer (1835: 46)

Alijanpoor, Behnaz, Khodayari, Hamed, Assadi, Mostafa, Rahiminejad, Mohammadreza & Mehregan, Iraj, 2021, A taxonomic revision of Rhamnus L. and Atadinus Raf. (Rhamnaceae) in Iran, Phytotaxa 521 (3), pp. 127-158 : 133-134

2. Rhamnus spathulifolia Fischer & Meyer (1835: 46) .

Typus — AZERBAIJAN. Elisabethpol : In rupestribus ad rivulum Gandscha circa coloniam Helenendorf ( LE 01042935, image!) .

Description— Trees or shrubs, 1.5–5 m high, bark grey or rarely brown. Two types of branches: long (spiny) branches ending in spines or without spines, mostly with alternate leaves at the top; and short (scaly) branches bearing flowers or fruits and fasciculate leaves at the end. Leaves 10–35 × 4–15 mm, coariceous, thick, sometime pubescent, polymorphic; base of leaves cuneate or narrowly cuneate; apex of leaves obtuse or rarely acute; margin with 5–12 pairs of teeth; primary lateral nerves almost erect, inconspicuous, (2–)3–6 pairs with 20–25 pairs of secondary lateral nerves; petioles 3–15 mm long, pubescent. Inflorescence fasciculate. Flowers polygamous on leaf axis, 2.5–3 mm long; calyx 3 mm long with acute apex; petals four or rarely two; anthers basifixed; disk very small; ovary mostly with 2(–3) locular and mostly with 2(–3) styls; peduncles glabrous. Fruits freshy, drupe, glabrous, in mature stage black, in immature stage green-red, with 2(–3) seeds, with 2(–3) deep (opened) furrows.

Discussion— R. spathulifolia was reported as a hybrid species R. cathartica var. caucasica Kusn. × R. pallasii Fisch. & C. A. Mey. ( Browicz & Zielinski (1977: 25) but is here described as a species R. spathulifolia Fisch. & C. A. Mey. , because plants are not depended on their parent, stable in their habitat and distributed in low altitude (while the putative parents are distributed at high altitude). They are fertile and have mature fruit. Their distribution altitudes is ranged from 20 m to 1500 m, which is a possible reason for polymorphic leaves, including narrow elliptic, lanceolate, ramoide and so on. Plants of R. spathulifolia also have a different habit in different altitudes: erect with a few spines at low altitudes and shrubby with some spins at high altitudes. Sometimes their fruits are differed in size and color and two sizes of leaves could be observed. There are also some specimens with many thick and long spins. Of course, there is a few differences between R. pallasii and R. spathulata genetically (Hauenschild et al., 2016).

Habitat and distribution —Endemic to N IRAN and ARMENIA.

R. spathulifolia is an Euro-Siberian element of Gilan, Golestan and Mazandaran Provinces of N Iran (rarely in Tehran). It grows at an altitude of 20–1500 m, between 49° and 56° E and between 36° and 37° N on sandy coastal and forest habitats in different habit with polymorphic leaves. It is related to R. pallasii and R. cathartica . It is also reported from TURKEY, KENYA and ARMENIA under this name.

Phenology —Flowering is in May, fruits and seeds maturie in August.

Examined specimens— IRAN. Gilan: From Rasht towards Rudbar, 2 Km towards Rostamabad , pasturage– forest habitat, 49°25ʹ58ʺ E, 36°44ʹ57ʺ N, 160–170 m, 03 May 2017, Alijanpoor & Alijanpoor s.n. ( TARI 103864 !); Golestan: Around Aliabad-e-Katul , beside the Negarestan dam, forest habitat, 54°52ʹ48ʺ E, 36°52ʹ25ʺ E, 180 m, 30 April 2017, Alijanpoor & Alijanpoor s.n. ( TARI 103862 !); 10 Km from Kalaleh towards Maraveh Tapeh , Eastern slope, forest and sandy habitat, 55°37ʹ15ʺ E, 37°34ʹ20ʺ N, 210 m, 30 April 2017, Alijanpoor & Taheri s.n. ( TARI 103891 !); Northeastern border of the Golestan National Park , 9 km of Mirza-Baylu towards Soolegerd , Jakhti- Kalan pass, 56° 11ʹ E, 37 °24ʹ N, 1850 m, 21–22 June 1995, H . Akhani s.n. ( W 02537 !); Northeastern, ca. 6 km S Soolegerd , Degarmanli river bed, 56°08ʹ E, 37°24ʹ N, 1500 m, 10 July 1995, Akhani s.n. ( W 02539!); Mazandaran: From Babolsar towards Bahnamir, near to Mirod village , sandy beach habitat, 52°44ʹ31ʺ E, 36°43ʹ13ʺ N, 45 m, 02 May 2017, Alijanpoor & Hosseini s.n. ( TARI 103844 !); in delivibus boreal, jugi Gaduk, 1900 m, Gauba & Mirdamadi s.n. ( W 08336!); Kelardash t, 1200 m, 29 May 1998, Gauba & Sabeti s.n. ( W 08327!); From Karaj towards Siah bisheh, ca. 10 km after Siah Bisheh dam, Nothtern slope, forest habitat, 51°19ʹ12ʺ E, 36°12ʹ48ʺ N, 1500 m, 04 May 2017, Alijanpoor & Azizi s.n. ( TARI 103885 , 103892 !); From Sari towards Neka, before Neka industrial city, on the hills, Western slope, forest and sandy habitat, 53°20ʹ08ʺ E, 36°39ʹ52ʺ N, 75 m, 02 May 2017, Alijanpoor & Hosseini s.n. ( TARI 103863 !); In valle fluvi Talar inter jugi Gaduk et Abbasabad ca. 1800 m, 04 October 1937, s.n. ( W 6353 !); Persian borealis: 2000 m., 20 September 1937, D. E . Gauba, 1460 Plantae Gaubaeanae Iranica; In vally Lur montium Elburs occid ad pagum Gotachenal, 09 June 1902, 2200 m, J . Bornmüller 65034 ( W!) ; Persia borealis: In valle Lur montium Elbur s Occid , 2200 m, 06 July 1902, J . et A. Borrnmϋller, J. Bornmüller 6558 ( W!) ; ARMENIA. Abovyan, ad meridiem versus a vico Gami in valle fluminis Azat. , 1200–1500 m, 03 October 1974, V . Vasak s.n. ( W 00785!); Aboryan, ad vicum Arzni , in declivibus vallis fluminis, 1500–1600 m, 07 October 1974, V . Vasak s.n. ( G 335973 !); Ashtarak, in declivibus montis, Aragac , in valle fluminis Amberd , situ austro–occidental a vico Byurakan., 1500–1600 m, 04 October 1974, V . Vasak s.n. ( W 11495!) .

The specimens from the exsiccate series Inter transcaspica–persicum 648 (in TURKEY) were reported R. spathulifolia , but in this review is recognized as R. Pallasii subsp sintenisii because of the larger number of spine, shape and size of leaves, and the specimens of (W 10219!) (in KENYA), (W 0007562!) (in GEORGIA) and (W 088536, 0007563 0007564!) (in ARMENIA) were reported as R. spathulifolia , but in this rewiew are identified as R.cathartica because of the shape of the leaves and the number of lateral nerves.