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