Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 595 No. 3: 8 May 2023
Type: Article
Published: 2023-05-08
DOI: 10.11646/phytotaxa.595.3.1
Page range: 241-285
Abstract views: 179
PDF downloaded: 6

Pollen morphology of Scorzonera s.l. (Asteraceae, Cichorieae, Scorzonerinae) occurring in Iran

Central Herbarium of Tehran University, School of Biology, College of Science, University of Tehran, Tehran, Iran
Central Herbarium of Tehran University, School of Biology, College of Science, University of Tehran, Tehran, Iran
Department of Pharmacognosy, Faculty of Pharmacy, Tabriz medical Sciences University, Pishghadam Square, Tabriz, Iran
Department of Biology, Faculty of Sciences, Shahid Bahonar University of Kerman, 22 Bahman Blvd. Afzalipour Square, Kerman, Iran
Royal Botanic Garden Edinburgh, Inverleith Row, Edinburgh, United Kingdom
Asteraceae pollen Scorzonera Iran Eudicots

Abstract

Pollen characters of 48 species of Scorzonera s.l. distributed in Iran were investigated by using light and scanning electron microscopy. The pollen grains are isopolar, radially symmetric, generally 3-zonocolporate, rarely 2-colpororate or 4-colporate. The pollen grains are oblate-spheroidal in shape (P/E=0.89 to 0.99) with the polar axes 37.1–58.1 µm and the equatorial axes 38.8–60.6 µm. We adopted the four pollen types recognized by Blackmore based on the number and position of their lacunae. For the first time, some pollen grains of S. longipapposa and S. tortuosissima were shown to have 26 (24+2) lacunae. We compared pollen morphological characters with the findings of the most recent systematic treatment. Type A with 6 lacunae was observed in Takhtajaniantha pusilla and S. parviflora, type B with 18 lacunae in Gelasia, type C with 20 lacunae in Pseudopodospermum and type D with 24 lacunae in Ramaliella, S. renzii and the Podospermum clade (except S. laciniata) in Scorzonera. We also observed 15 or 18 lacunae in S. rupicola. The pollen in Scorzonera appears to evolve toward reduction of spines in S. parviflora, extension of spines in the Podospermum clade, enlargement of the space between ridges and increase in the number of lacunae (S. longipapposa and S. tortuosissima). 26 pollen characters of the examined species were analysed by UPGMA and PCA. Pollen size, spine size, distance between spines, number and arrangement of spines between two equatorail lacunae, exine thickness, spine length/P or E, dimensions of lacunae and polar area were shown to contribute significantly to understanding the morphology of Scorzonera.

References

<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB"><em>Askerova, R. (1969) Palynological data on the systematics and phytogeny of the genus Scorzonera </em></span><span lang="en-GB">L. </span><span lang="en-GB"><em>Isvestija AN Azerbaidzhanskoj SSR</em></span><span lang="en-GB"> 6: 11–15. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Askerova, R. (1970) Morphology of pollen of the genus </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. </span><span lang="en-GB"><em>Isvestija AN Azerbaidzhanskoj SSR </em></span><span lang="en-GB">4: 43–46. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Askerova, R. (1987) </span><span lang="en-GB"><em>Palynology of Cichorioideae, Compositae</em></span><span lang="en-GB">. Baku, 207 pp. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Blackmore, S. (1982a) Palynology of subtribe Scorzonerinae (Compositae: Lactuceae) and its taxonomic significance. </span><span lang="en-GB"><em>Grana </em></span><span lang="en-GB">21: 149–160. https://doi.org/10.1080/00173138209427691</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Blackmore, S. (1982b) A functional interpretation of Lactuceae (Compositae) pollen. </span><span lang="en-GB"><em>Plant Systematics and Evolution</em></span><span lang="en-GB"> 141: 153–168. https://doi.org/10.1007/BF00986415</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Blackmore, S., Wortley, A.H., Skvarla, J.J., Gabarayeva, N.I. &amp; Rowley, J.R. (2010) Developmental origins of structural diversity in pollen walls of Compositae. </span><span lang="en-GB"><em>Plant Systematics and Evolution</em></span><span lang="en-GB"> 284: 17–32. https://doi.org/10.1007/s00606-009-0232-2</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Boissier, E.P. (1875) </span><span lang="en-GB"><em>Flora Orientalis</em></span><span lang="en-GB">, vol. 3. H. Georg, Genevae and Basileae, 1033 pp.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Bordbar, F., Makki, P. &amp; Alifarsangi, F. (2019) Karyotype investigations in Iranian </span><span lang="en-GB"><em>Scorzonera tortuosissima</em></span><span lang="en-GB"> and </span><span lang="en-GB"><em>S. intricata</em></span><span lang="en-GB"> (Asteraceae). </span><span lang="en-GB"><em>Cytologia</em></span><span lang="en-GB"> 84: 69–72. https://doi.org/10.1508/cytologia.84.69</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Bremer, K. &amp; Anderberg, A.A. (1994) </span><span lang="en-GB"><em>Asteraceae: cladistics &amp; classification</em></span><span lang="en-GB">. Timber Press, Portland, 752 pp.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Candolle, A.P. de (1805) </span><span lang="en-GB"><em>Podospermum </em></span><span lang="en-GB">(L.) DC. </span><span lang="en-GB"><em>In</em></span><span lang="en-GB">: Lamarck, J.B. &amp; Candolle, A.P., de, </span><span lang="en-GB"><em>Flore Francaise</em></span><span lang="en-GB">, vol. 4. F. Desza, Paris, p. 61.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Candolle, A.P. de (1838) </span><span lang="en-GB"><em>Prodromus systematis naturalis regni vegetabilis</em></span><span lang="en-GB">, vol. 7. Treuttel &amp; Würtz, Paris, 801 pp.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Chamberlain, D.F. (1975) </span><span lang="en-GB"><em>Scorzonera </em></span><span lang="en-GB">L. </span><span lang="en-GB"><em>In</em></span><span lang="en-GB">: Davis, P.H. (ed.)</span><span lang="en-GB"><em> Flora of Turkey and the East Aegean Islands</em></span><span lang="en-GB">, vol. 5. Edinburgh University Press, Edinburgh, pp. 632–657.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Chater, A. (1976) </span><span lang="en-GB"><em>Scorzonera. In</em></span><span lang="en-GB">: Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M., Webb, D.A. (eds.) </span><span lang="en-GB"><em>Flora Europaea</em></span><span lang="en-GB">, vol. 4. Cambridge University Press, Cambridge, pp. 317–322.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Guardia, C.D. &amp; Blanca, G. (1987) Karyology of the </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> (Compositae) species from the Iberian Peninsula. </span><span lang="en-GB"><em>Plant Systematics and Evolution</em></span><span lang="en-GB"> 156: 29–42. https://doi.org/10.1007/BF00937199</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Dvořák, F., Dadáková, B. &amp; Růžička, I. (1979) Chromosome morphology of the Czechoslovak species of the </span><span lang="en-GB"><em>genus</em></span> <span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB">. </span><span lang="en-GB"><em>Folia Geobotanica &amp; Phytotaxonomica</em></span><span lang="en-GB"> 14: 185–199. https://doi.org/10.1007/BF00937199</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Erdtman, G. (1960) The acetolysis method – a revised description. </span><span lang="en-GB"><em>Svensk Botanisk Tidskrift</em></span><span lang="en-GB"> 54: 516–564.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Hammer, Ø., Harper, D.A. &amp; Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. </span><span lang="en-GB"><em>Palaeontologia electronica</em></span><span lang="en-GB"> 4, 9.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Hatami, E., Bordbar, F. &amp; Mirtadzadini, M. (2019) New Chromosome Numbers in </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB">, subgenera </span><span lang="en-GB"><em>Podospermum</em></span><span lang="en-GB"> and </span><span lang="en-GB"><em>Pseuodopodospermum</em></span><span lang="en-GB"> (Cichorieae, Asteraceae). </span><span lang="en-GB"><em>Cytologia</em></span><span lang="en-GB"> 84: 247–250. https://doi.org/10.1508/cytologia.84.247</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Hatami, E., Mirtadzadini, M., Bordbar, F. &amp; Jones, K.E. (2020) Delimitation of Iranian species of </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> subg. </span><span lang="en-GB"><em>Podospermum</em></span><span lang="en-GB"> and </span><span lang="en-GB"><em>S. </em></span><span lang="en-GB">subg. </span><span lang="en-GB"><em>Pseudopodospermum</em></span><span lang="en-GB"> (Asteraceae, Cichorieae) based on morphological and molecular data. </span><span lang="en-GB"><em>Willdenowia</em></span><span lang="en-GB"> 50: 39–63. https://doi.org/10.3372/wi.50.50105</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Kamelin, R. &amp; Tagaev, I. (1986) Survey of species of the genus </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. (Asteraceae). </span><span lang="en-GB"><em>Botanicheskii Zhurnal </em></span><span lang="en-GB">71: 1672–1682. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Kuthatheladze, S. (1978) </span><span lang="en-GB"><em>Kavkazskie predstaviteli podtriby Scorzonerinae Dum.</em></span><span lang="en-GB"> [The Caucasian members of the tribe Scorzonerinae Dum.]. Metsniereba, Tbilisi, 184 pp. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Lipschitz, S.J. (1935)</span> <span lang="en-GB">Fragmenta monographiae generis</span><span lang="en-GB"><em> Scorzonera</em></span><span lang="en-GB"> 1. </span><span lang="en-GB"><em>Transactions of the Rubber and Guttapercha Institute</em></span><span lang="en-GB"> 1: 1–164. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Lipschitz, S.J. (1939) </span><span lang="en-GB"><em>Fragmenta Monographiae Generis</em></span> <span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> 2. Moskovskoe Obshestvo Ispytatelei Prirody, Moscow, 165 pp. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Makbul, S., Turkmen, Z., Coskuncelebi, K. &amp; Beyazoglu, O. (2010) A morphometric study on </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. taxa (Asteraceae) from northeast Anatolia. </span><span lang="en-GB"><em>Acta Botanica Croatica</em></span><span lang="en-GB"> 69 (2): 237–247.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Makbul, S., Coşkunçelebi, K., Okur, S. &amp; Gültepe, M. (2016) Contribution to the taxonomy of Turkish </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> (Asteraceae) taxa based on vegetative anatomy. </span><span lang="en-GB"><em>Nordic Journal of Botany </em></span><span lang="en-GB">34: 670–684. https://doi.org/10.1111/njb.01159</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Makbul, S., Coskuncelebi, K., Türkmen, Z. &amp; Beyazoglu, O. (2011) Comparison of foliar anatomy of </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. (Asteraceae) taxa from northeast Anatolia. </span><span lang="en-GB"><em>Pakistan Journal of Botany</em></span><span lang="en-GB"> 43: 135–155.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Martin, E., Çetin, Ö., Duran, A., Doðan, B. &amp; Traȿ, Z. (2015) New karyotypes in some species of Asteraceae from Turkey. </span><span lang="en-GB"><em>Cytologia</em></span><span lang="en-GB"> 80: 237–248. https://doi.org/10.1508/cytologia.80.237</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Mavrodiev, E.V., Edwards, C.E., Albach, D.C., Gitzendanner, M.A., Soltis, P.S. &amp; Soltis, D.E. (2004) Phylogenetic relationships in subtribe Scorzonerinae (Asteraceae: Cichorioideae: Cichorieae) based on ITS sequence data. </span><span lang="en-GB"><em>Taxon</em></span><span lang="en-GB"> 53: 699–712. https://doi.org/10.2307/4135445</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Meo, A. &amp; Khan, M. (2004) Pollen morphology as an aid to the identification of </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> (Cichorieae-Compositae) from Pakistan. </span><span lang="en-GB"><em>Pakistan Journal of Botany</em></span><span lang="en-GB"> 36: 701–710.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Nair, P.K.K. (1985) </span><span lang="en-GB"><em>Pollen morphology of Indian Compositae</em></span><span lang="en-GB">. Today &amp; Tomorrow’s Print, Delhi, 182 pp.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Nazarova, E.A. (1990) </span><span lang="en-GB"><em>Takhtajaniantha</em></span><span lang="en-GB"> Nazarova and </span><span lang="en-GB"><em>Lactucella</em></span><span lang="en-GB"> Nazarova, two new genera of the tribe Lactuceae (family Asteraceae). </span><span lang="en-GB"><em>Biologicheskiĭ Zhurnal Armenii</em></span><span lang="en-GB"> 43: 179–183. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Nazarova, E.A. (1997) Karyosystematic investigation of the genus </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. s.l. (Lactuceae, Asteraceae). </span><span lang="en-GB"><em>Caryologia </em></span><span lang="en-GB">50: 239–261. https://doi.org/10.1080/00087114.1997.10797399</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Norouzi, M., Ghahremaninejad, F., Maassoumi, A.A. &amp; Safavi, S.R. (2016) Anatomical studies on </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> (Asteraceae) species, subgenera </span><span lang="en-GB"><em>Podospermum</em></span><span lang="en-GB"> and </span><span lang="en-GB"><em>Pseudopodospermum</em></span><span lang="en-GB"> in Iran. </span><span lang="en-GB"><em>Modern Phytomorphology </em></span><span lang="en-GB">9: 51–68.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Osman, A.K. (2006) Pollen types of the Egyptian species of tribe Lactuceae (subfamily Cichorioideae-Compositae). </span><span lang="en-GB"><em>Acta Botanica Croatica</em></span><span lang="en-GB"> 65: 161–180.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Owen,W.M., D’Amato, G., De Dominicis, R.I., Salimbeni, P. &amp; Tucci, G.F. (2006) A cytological and molecular study of the genera </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. and </span><span lang="en-GB"><em>Podospermum</em></span><span lang="en-GB"> (L.) DC. (Asteraceae). </span><span lang="en-GB"><em>Caryologia </em></span><span lang="en-GB">59: 153–163. https://doi.org/10.1080/00087114.2006.10797911</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Pinar, N.M., Coþkunçelebý, K., Þimsek, D., Okur, S., Makbul, S. &amp; Acar, A. (2016) Pollen morphology of </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> (Asteraceae) in Turkey. </span><span lang="en-GB"><em>Phytotaxa</em></span><span lang="en-GB"> 252: 233–262. https://doi.org/10.11646/phytotaxa.252.4.1</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Podani, J. (2000) Simulation of random dendrograms and comparison tests: some comments. </span><span lang="en-GB"><em>Journal of Classification</em></span><span lang="en-GB"> 17: 123–142. https://doi.org/10.1007/s003570000007</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Qureshi, S.J., Khan, M.A. &amp; Rashid, A. (2008) Diameter, exine thickness, and sculpturing in genera </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L., </span><span lang="en-GB"><em>Sonchus</em></span><span lang="en-GB"> L. and </span><span lang="en-GB"><em>Tragopogon</em></span><span lang="en-GB"> L. of Asteraceae in Pakistan. </span><span lang="en-GB"><em>International Journal of Science &amp; Technology</em></span><span lang="en-GB"> 3: 139–149.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Rechinger, K.H. (1977) </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB">. </span><span lang="en-GB"><em>In</em></span><span lang="en-GB">: Rechinger, K.H. (ed.) </span><span lang="en-GB"><em>Flora Iranica</em></span><span lang="en-GB">, vol. 122. Akademische Druck- u. Verlagsanstalt, Graz, pp. 16–83.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Safavi, S.R. (2013) </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB">.</span><span lang="en-GB"><em> In</em></span><span lang="en-GB">: Safavi, S.R., Naseh, Y., Jafari, E, Tavakoli, Z. &amp; Heidarnia, N., Tribe Cichorieae, Asteraceae. </span><span lang="en-GB"><em>Flora of Iran</em></span><span lang="en-GB">, vol. 77. Research Institute of Forest and Rangelands, Tehran, pp. 16–83. [In Persian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Safavi, S.R. (2016) A new species of </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> (Asteraceae) from Natanz, Iran. </span><span lang="en-GB"><em>Iranian Journal of Botany</em></span><span lang="en-GB"> 22: 1–5.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Safavi, S.R. (2019) </span><span lang="en-GB"><em>Scorzonera tomentosa</em></span><span lang="en-GB"> L. (Asteraceae), a new record for the Flora of Iran. </span><span lang="en-GB"><em>Iranian Journal of Botany</em></span><span lang="en-GB"> 25 (1): 40–43.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Tomb, A.S. (1975) Pollen morphology in tribe Lactuceae (Compositae). </span><span lang="en-GB"><em>Grana</em></span><span lang="en-GB"> 15: 79–89. https://doi.org/10.1080/00173134.1975.11864623</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Tomb, A.S., Larson, D.A. &amp; Skvarla, J.J. (1974) Pollen morphology and detailed structure of family Compositae, tribe Cichorieae. I. Subtribe Stephanomeriinae. </span><span lang="en-GB"><em>American Journal of Botany</em></span><span lang="en-GB"> 61: 486–498. https://doi.org/10.1002/j.1537-2197.1974.tb10788.x</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Türkmen, Z., Makbul, S., Çoşkunçelebi, K. &amp; Beyazoðlu, O. (2010) Palynological observations on the genus </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> L. (Asteraceae) from north-east Anatolia (Turkey). </span><span lang="en-GB"><em>Turkish Journal of Botany</em></span><span lang="en-GB"> 34: 495–512. https://doi.org/10.3906/bot-0909-17</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Tzvelev, N. (1988) About some plant species from European part of USSR and from the Far East. </span><span lang="en-GB"><em>Novosti Sistematiki Vysshikh Rastenii</em></span><span lang="en-GB"> 25: 183–191. [In Russian]</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Vega, A.J. &amp; Dematteis, M. (2011) Pollen morphology of some species of </span><span lang="en-GB"><em>Vernonanthura</em></span><span lang="en-GB"> (Asteraceae, Vernonieae) from southern South America. </span><span lang="en-GB"><em>Palynology</em></span><span lang="en-GB"> 35: 94–102. https://doi.org/10.1080/01916122.2010.549290</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Wagenitz, G. (1976) Systematics and phylogeny of the Compositae (Asteraceae). </span><span lang="en-GB"><em>Plant Systematics and Evolution </em></span><span lang="en-GB">125: 29–46. https://doi.org/10.1007/BF00986129</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Wodehouse, R.P. (1935) </span><span lang="en-GB"><em>Pollen grains. Their structure, identification and significance in science and medicine</em></span><span lang="en-GB">. McGraw–Hill Book Company, Inc., New York.</span></span></span></span></p>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span lang="en-GB">Zaika, M.A., Kilian, N., Jones, K., Krinitsina, A.A., Nilova, M.V., Speranskaya, A.S. &amp; Sukhorukov, A.P. (2020) </span><span lang="en-GB"><em>Scorzonera</em></span><span lang="en-GB"> sensu lato (Asteraceae, Cichorieae)–taxonomic reassessment in the light of new molecular phylogenetic and carpological analyses. </span><span lang="en-GB"><em>PhytoKeys</em></span><span lang="en-GB"> 137: 1–85. https://doi.org/10.3897/phytokeys.137.46544</span></span></span></span></p>