Abstract
This study aims to revise the taxonomic status of Rhamnus L. and Atadinus Raf. (Rhamnaceae) for the Flora of Iran. Nine species, including 15 taxa are recognized, described and compared with their closely related taxa. These include one new species, R. rahiminejadii Alijanpoor & Assadi, one new subspecies, R. pallasii subsp. mazandaranica Alijanpoor & Assadi, and one hybrid species, R. ×mehreghanii Alijanpoor & Khodayari (R. persica Boiss. × R. kurdica Boiss. & Hohen.). In addition, a hybrid species i.e, ×R. spathulifolia Fisch. & C. A. Mey., is accepted as a distinct species, R. spathulifolia Fisch. & C. A. Mey. and R. cornifolia and R. cornifolia var. denudata, from sect. Rhamnustrum Rouy. are transfered to genus Atadinus Raf. as A. cornifolius (Boiss. & Hohen.) Alijanpoor and A. cornifolius var. denudatus (Bornm.) Alijanpoor.
References
Bentham, G. & Hooker, J.D. (1862) Genera Plantarum 1. London, pp. 371–386.
Boissier, E. (1872) Flora Orientalis 2. H. Georg, Basel & Genève, pp. 14–22.
Boissier, E. (1859) Diagnoses plantarum novarum praesertim Orientalium nonnulis Europaeis boreali-africanisque additis. Ser. 2, 3 (6). B. Herrmann, Lipsiae [Leipzig]; J.-B. Baillière, Parisiis [Paris] (Typis Ramboz et Schuchardt, Genève), 148 pp.
Bolmgren, K. & Oxelman, B. (2004) Generic limits in Rhamnus L. s. l. (Rhamnaceae) inferred from nuclear and chloroplast DNA sequence phylogenies. Taxon 53 (2): 383–390. https://doi.org/10.2307/4135616
Brongniart, A. (1826) Memoire sur la Famille des Rhamnées. Didot, Paris, pp. 14–76.
Browicz, K. & Zielinski, J. (1977) Rhamnaceae In: Rechinger, K.H. (Ed.) Flora Iranica 157. Akademische Druck- u. Verlagsanstalt, Graz, pp. 14–25.
Canale, A., Benvenuti, S., Raspi, A. & Benelli, G. (2016) Insect pollinators of the late winter flowering Rhamnus alaternus L., a candidate for honeybee-friendly scrubland spots in intensively managed agricultural areas. Plant Biosystems 150 (4): 611–615. https://doi.org/10.1080/11263504.2014.993742
Chalabi-Ka<sup>,</sup>bi, Z. (1980) Rhamnus L. In: Townsend , C.C. & Guest, E. (eds.) Flora of Iraq, vol. 4. Ministry of Agriculture and Agrarian Reform Republic of Iraq , Baghdad, pp. 438–442.
Candolle, A.P. de. (1825) Rhamneae. In: Candolle, A.P. de (ed.) Prodromus Systematis Naturalis Regni Vegetabilis 2.Treuttel et Würtz, Paris, pp. 19–42.
Davis, P.H. & Yaltriv, F. (1967) Rhamnus L. In: Davis, P.H. (Ed.) Flora of Turkey and the East Aegean Islands 2. [Edinburgh] University Press, Edinburgh , pp. 526–541.
Dodoens, R. (1583) Stirpium Historiae Pemptades Sex Sive Libri XXX. Christophori Plantini, Antwerpen, pp. 741–743. https://doi.org/10.5962/bhl.title.149774
Doweld, A.B. (2017) New names of fossil Atadinus, Endotropis and Frangula (Rhamnaceae). Phytotaxa 321 (2): 194–200. https://doi.org/10.11646/phytotaxa.321.2.4
Endlicher, S. (1836-1840) Genera Plantarum Secundum Ordines Naturales disposite. Vindobona, Beck Universitatis Biblioplam, pp. 1094–1103. https://doi.org/10.5962/bhl.title.728
Fisch, F.E.L. & Meyer, C.A. (1837) Index seminum, quae Hortus Botanicus Imperialis Petropolitanus pro mutua commutatione offert :accedunt Animadversiones botanicae nonnullae 4. Hortus, St. Petersburg, pp. 46–47.
Grubov, V.I. (1949) Monography of Rhamnus L. s. l. In: Schischkin, B.K. (ed.) Flora et Systematica Plantae Vasculares 6. Academiae Scientiarum USSR, Leningrad, pp. 14–425.
Gulýas, J. & Traveset, A. (2012) Altitudinal variation in the reproductive performance of the Mediterranean shrub Rhamnus lycioides L.. Plant Ecology. 5 (3): 330–336. https://doi.org/10.1093/jpe/rtr047
Hauenschild, F., Favre, A., Salazar, G.A. & Muellner-Riehl, A.N. (2016c) Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s. l. supports the description of a new genus Venetia. Taxon 65 (1): 65–78. https://doi.org/10.12705/651.5
Hauenschild, F., Favre, A., Michalak, I. & Muellner-Riehl, A.N. (2018a) The influence of the Gondwanan breakup on the biogeographic history of the ziziphoids (Rhamnaceae). Journal of Biogeography 45: 1–9. https://doi.org/10.1111/jbi.13448
Hauenschild, F., Favre, A., Salazar, G.A. & Muellner-Riehl, A.N. (2016) Erratum: Corrigendum to Hauenschild, F. & al., Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s. l. supports the description of a new genus Venetia. Taxon 65 (4): 926–927. https://doi.org/10.12705/651.5
Hauenschild, F., Matuszak, S., Muellner-Riehl, A.N. & Favre1, A. (2016b) Phylogenetic relationships within the cosmopolitan buckthorn family (Rhamnaceae) support the resurrection of Sarcomphalus and the description of Pseudoziziphus gen. nov. Taxon 65 (1): 47–64. https://doi.org/10.12705/651.4
Johnston, M.C. & Johnston, L.V. (1978) Rhamnus. In: Flora Neotropica Monograph 20. New York Botanical Garden, New York, pp. 1–98.
Jahandiez, E. & Maire, R. (1932) Catalog of Morocco Plants 2. Imprimerie Minerva, Paris, pp. 475–477.
Linnaeus, C. (1753) Species plantarum 1. Salvius, Holmiae [Stockholm], 560 pp.
Linnaeus, C. (1762) Species plantarum 1. Salvius, Holmiae [Stockholm], 784 pp.
Medan, D. Montaldo, N.H. (2005) Ornithophily in the Rhamnaceae: The pollination of the Chilean endemic Colletia ulicina, Flora - Morphology Distribution Functional Ecology of Plants 200 (4): 339–344. https://doi.org/10.1016/j.flora.2004.12.003
Medan, D. & Schirarend, C. (2004) Rhamnaceae in Kubitzki, K. (Ed.) The Families and Genera of Vascular Plants 6. Springer-Verlag Berlin Heidelberg, pp. 332–338. https://doi.org/10.1007/978-3-662-07257-8_37
Miller, P. (1754) The gardeners dictionary 3. Printed for the author and sold by John and James Rivington, London, 590 pp. https://doi.org/10.5962/bhl.title.79061
Onstein, R.E., Carter, R.J., Xing, Y.W., Richardson, J.E. & Linder, H.P. (2015) Do Mediterranean-type ecosystems have a common history—Insights from the buckthorn family (Rhamnaceae). Evolution 69: 756–771. https://doi.org/10.1111/evo.12605
Pallas, P.S. (1776) Reise aus Sibirien zur?ck an die Wolga im 1773 sten Jahr. Reise durch verschie Provinzen des russischen, Kayserl Akademi der Wissenschaften, Saint Petersburg 3: 458-760.
Pool, A. (2013) New species, combinations, and lectotypifications in Neotropical and northern Mexican Frangula (Rhamnaceae). Novon 22: 447–467. https://doi.org/10.3417/2013009
Qaiser, M. & Nazimuddin, S. (1981) Rhamnaceae In: Nasir, E. & Ali, S.I. (Eds.) Flora of Pakistan, vol. 140. Department of Botany, University of Karachi, pp. 17–23.
Rafinesque, C.S. (1838) New genera and species of tree and shrubes of North America and other regions on the earth, Philadelphia pp. 26–34. https://doi.org/10.5962/bhl.title.23764
Richardson, J.E., Fay, M.F., Cronk, Q.C.B., Bowman, D. & Chase, M.W. (2000a) A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences. American Journal of Botany 87: 1309–1324. https://doi.org/10.2307/2656724
Richardson, J.E., Fay, M.F., Cronk, Q.C.B. & Chas, M.W. (2000) A Revision of the Tribal Classification Rhamnaceae. Kew Bulletin, Springer 55 (2): 311–340. https://doi.org/10.2307/4115645
Richardson, J.E., Weitz, F.M., Fay, M.F., Cronk, Q.C.B., Linder, H.P., Reeves, G. & Chase, M.W. (2001) Phylogenetic analysis of Phylica L. (Rhamnaceae) with an emphasis on island species: evidence from plastid trnL-F and nuclear internal transcribed spacer ibosomal) DNA sequences. Taxon 50: 405–427. https://doi.org/10.2307/1223889
Shishkin, BK. & Bobrov, EG. (1974) Flora of the USSR 14. Academy of Sciences of the USSR, pp. 494–516.
Sofian, Kh. & Dinarvand (2007) Rhamnus L. In: Assadi, M. (ed.) Flora Iran 55. Researches Institute of Forests and Rangelands, Tehran, pp. 22–44.
Suessenguth, K. (1953) Pflanzenfam In: Engler, A. & Prantl, K. (eds.) Angiosperm Reibe Rhamnales. Dunker & Humblot Berlin, Band 20d, pp. 65–65.
Takhtajan, A.L. (1973) (Ed.) Flora of Armenia Ericaceae to Elaegnaceae 6. Yerevan: Armenian Academy of Sciences Press, 485 pp.
The Angiosperm Phylogeny Group [APC] (2016) An update of the Angiosperpm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical journal of the Linnean Society 181 (1): 1–20. https://doi.org/10.1111/boj.12385
Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M. & Webb, D.A. (1968) Flora Europaea, vol. 2, Cambridge University, Cambridge, pp. 244–245.
Vent, W. (1962) Monographie der Gattung Oreoherzogia W. Vent gen. nov. Feddes Repertorium. Specierum. Novarum. Regni Vegetabilis 65: 3–132.
Weberbauer, A. (1895) Rhamnaceae In: Engler, A. & Prantl, K. (eds.) Die natürlichen Pflanzenfamilien, vol. 5. Wilhelm Engelmann, Leipzig, pp. 393–427.
Boissier, E. (1872) Flora Orientalis 2. H. Georg, Basel & Genève, pp. 14–22.
Boissier, E. (1859) Diagnoses plantarum novarum praesertim Orientalium nonnulis Europaeis boreali-africanisque additis. Ser. 2, 3 (6). B. Herrmann, Lipsiae [Leipzig]; J.-B. Baillière, Parisiis [Paris] (Typis Ramboz et Schuchardt, Genève), 148 pp.
Bolmgren, K. & Oxelman, B. (2004) Generic limits in Rhamnus L. s. l. (Rhamnaceae) inferred from nuclear and chloroplast DNA sequence phylogenies. Taxon 53 (2): 383–390. https://doi.org/10.2307/4135616
Brongniart, A. (1826) Memoire sur la Famille des Rhamnées. Didot, Paris, pp. 14–76.
Browicz, K. & Zielinski, J. (1977) Rhamnaceae In: Rechinger, K.H. (Ed.) Flora Iranica 157. Akademische Druck- u. Verlagsanstalt, Graz, pp. 14–25.
Canale, A., Benvenuti, S., Raspi, A. & Benelli, G. (2016) Insect pollinators of the late winter flowering Rhamnus alaternus L., a candidate for honeybee-friendly scrubland spots in intensively managed agricultural areas. Plant Biosystems 150 (4): 611–615. https://doi.org/10.1080/11263504.2014.993742
Chalabi-Ka<sup>,</sup>bi, Z. (1980) Rhamnus L. In: Townsend , C.C. & Guest, E. (eds.) Flora of Iraq, vol. 4. Ministry of Agriculture and Agrarian Reform Republic of Iraq , Baghdad, pp. 438–442.
Candolle, A.P. de. (1825) Rhamneae. In: Candolle, A.P. de (ed.) Prodromus Systematis Naturalis Regni Vegetabilis 2.Treuttel et Würtz, Paris, pp. 19–42.
Davis, P.H. & Yaltriv, F. (1967) Rhamnus L. In: Davis, P.H. (Ed.) Flora of Turkey and the East Aegean Islands 2. [Edinburgh] University Press, Edinburgh , pp. 526–541.
Dodoens, R. (1583) Stirpium Historiae Pemptades Sex Sive Libri XXX. Christophori Plantini, Antwerpen, pp. 741–743. https://doi.org/10.5962/bhl.title.149774
Doweld, A.B. (2017) New names of fossil Atadinus, Endotropis and Frangula (Rhamnaceae). Phytotaxa 321 (2): 194–200. https://doi.org/10.11646/phytotaxa.321.2.4
Endlicher, S. (1836-1840) Genera Plantarum Secundum Ordines Naturales disposite. Vindobona, Beck Universitatis Biblioplam, pp. 1094–1103. https://doi.org/10.5962/bhl.title.728
Fisch, F.E.L. & Meyer, C.A. (1837) Index seminum, quae Hortus Botanicus Imperialis Petropolitanus pro mutua commutatione offert :accedunt Animadversiones botanicae nonnullae 4. Hortus, St. Petersburg, pp. 46–47.
Grubov, V.I. (1949) Monography of Rhamnus L. s. l. In: Schischkin, B.K. (ed.) Flora et Systematica Plantae Vasculares 6. Academiae Scientiarum USSR, Leningrad, pp. 14–425.
Gulýas, J. & Traveset, A. (2012) Altitudinal variation in the reproductive performance of the Mediterranean shrub Rhamnus lycioides L.. Plant Ecology. 5 (3): 330–336. https://doi.org/10.1093/jpe/rtr047
Hauenschild, F., Favre, A., Salazar, G.A. & Muellner-Riehl, A.N. (2016c) Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s. l. supports the description of a new genus Venetia. Taxon 65 (1): 65–78. https://doi.org/10.12705/651.5
Hauenschild, F., Favre, A., Michalak, I. & Muellner-Riehl, A.N. (2018a) The influence of the Gondwanan breakup on the biogeographic history of the ziziphoids (Rhamnaceae). Journal of Biogeography 45: 1–9. https://doi.org/10.1111/jbi.13448
Hauenschild, F., Favre, A., Salazar, G.A. & Muellner-Riehl, A.N. (2016) Erratum: Corrigendum to Hauenschild, F. & al., Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s. l. supports the description of a new genus Venetia. Taxon 65 (4): 926–927. https://doi.org/10.12705/651.5
Hauenschild, F., Matuszak, S., Muellner-Riehl, A.N. & Favre1, A. (2016b) Phylogenetic relationships within the cosmopolitan buckthorn family (Rhamnaceae) support the resurrection of Sarcomphalus and the description of Pseudoziziphus gen. nov. Taxon 65 (1): 47–64. https://doi.org/10.12705/651.4
Johnston, M.C. & Johnston, L.V. (1978) Rhamnus. In: Flora Neotropica Monograph 20. New York Botanical Garden, New York, pp. 1–98.
Jahandiez, E. & Maire, R. (1932) Catalog of Morocco Plants 2. Imprimerie Minerva, Paris, pp. 475–477.
Linnaeus, C. (1753) Species plantarum 1. Salvius, Holmiae [Stockholm], 560 pp.
Linnaeus, C. (1762) Species plantarum 1. Salvius, Holmiae [Stockholm], 784 pp.
Medan, D. Montaldo, N.H. (2005) Ornithophily in the Rhamnaceae: The pollination of the Chilean endemic Colletia ulicina, Flora - Morphology Distribution Functional Ecology of Plants 200 (4): 339–344. https://doi.org/10.1016/j.flora.2004.12.003
Medan, D. & Schirarend, C. (2004) Rhamnaceae in Kubitzki, K. (Ed.) The Families and Genera of Vascular Plants 6. Springer-Verlag Berlin Heidelberg, pp. 332–338. https://doi.org/10.1007/978-3-662-07257-8_37
Miller, P. (1754) The gardeners dictionary 3. Printed for the author and sold by John and James Rivington, London, 590 pp. https://doi.org/10.5962/bhl.title.79061
Onstein, R.E., Carter, R.J., Xing, Y.W., Richardson, J.E. & Linder, H.P. (2015) Do Mediterranean-type ecosystems have a common history—Insights from the buckthorn family (Rhamnaceae). Evolution 69: 756–771. https://doi.org/10.1111/evo.12605
Pallas, P.S. (1776) Reise aus Sibirien zur?ck an die Wolga im 1773 sten Jahr. Reise durch verschie Provinzen des russischen, Kayserl Akademi der Wissenschaften, Saint Petersburg 3: 458-760.
Pool, A. (2013) New species, combinations, and lectotypifications in Neotropical and northern Mexican Frangula (Rhamnaceae). Novon 22: 447–467. https://doi.org/10.3417/2013009
Qaiser, M. & Nazimuddin, S. (1981) Rhamnaceae In: Nasir, E. & Ali, S.I. (Eds.) Flora of Pakistan, vol. 140. Department of Botany, University of Karachi, pp. 17–23.
Rafinesque, C.S. (1838) New genera and species of tree and shrubes of North America and other regions on the earth, Philadelphia pp. 26–34. https://doi.org/10.5962/bhl.title.23764
Richardson, J.E., Fay, M.F., Cronk, Q.C.B., Bowman, D. & Chase, M.W. (2000a) A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences. American Journal of Botany 87: 1309–1324. https://doi.org/10.2307/2656724
Richardson, J.E., Fay, M.F., Cronk, Q.C.B. & Chas, M.W. (2000) A Revision of the Tribal Classification Rhamnaceae. Kew Bulletin, Springer 55 (2): 311–340. https://doi.org/10.2307/4115645
Richardson, J.E., Weitz, F.M., Fay, M.F., Cronk, Q.C.B., Linder, H.P., Reeves, G. & Chase, M.W. (2001) Phylogenetic analysis of Phylica L. (Rhamnaceae) with an emphasis on island species: evidence from plastid trnL-F and nuclear internal transcribed spacer ibosomal) DNA sequences. Taxon 50: 405–427. https://doi.org/10.2307/1223889
Shishkin, BK. & Bobrov, EG. (1974) Flora of the USSR 14. Academy of Sciences of the USSR, pp. 494–516.
Sofian, Kh. & Dinarvand (2007) Rhamnus L. In: Assadi, M. (ed.) Flora Iran 55. Researches Institute of Forests and Rangelands, Tehran, pp. 22–44.
Suessenguth, K. (1953) Pflanzenfam In: Engler, A. & Prantl, K. (eds.) Angiosperm Reibe Rhamnales. Dunker & Humblot Berlin, Band 20d, pp. 65–65.
Takhtajan, A.L. (1973) (Ed.) Flora of Armenia Ericaceae to Elaegnaceae 6. Yerevan: Armenian Academy of Sciences Press, 485 pp.
The Angiosperm Phylogeny Group [APC] (2016) An update of the Angiosperpm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical journal of the Linnean Society 181 (1): 1–20. https://doi.org/10.1111/boj.12385
Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M. & Webb, D.A. (1968) Flora Europaea, vol. 2, Cambridge University, Cambridge, pp. 244–245.
Vent, W. (1962) Monographie der Gattung Oreoherzogia W. Vent gen. nov. Feddes Repertorium. Specierum. Novarum. Regni Vegetabilis 65: 3–132.
Weberbauer, A. (1895) Rhamnaceae In: Engler, A. & Prantl, K. (eds.) Die natürlichen Pflanzenfamilien, vol. 5. Wilhelm Engelmann, Leipzig, pp. 393–427.