Fruit, seed morphology and seed coat sculpturing were recorded for 10 species belongs to 9 genera and five tribes of Brassicaceae in eastern region of Saudi Arabia by using light microscope (LM) and scanning electron microscope (SEM). Four seed patterns were recognized based on surface sculpturing pattern; Reticulate, with four subtypes, undulate, verrucate and ocellate. The results of numerical analysis showed that species were grouped into two major clusters and each cluster divided into two groups. The cluster analysis indicates that the characters of fruit and seed are very important characters for identification and taxonomical classification of the tribes. The results indicate that the most tribes of Brassicaceae are polyphyletic, and some are related to each other’s as Lepidieae, Alyssieae and Sisymbrieae.
KeywordsFruitSeed CoatMorphologyScanning Electron Microscope
Warwick, S.I., Francis, A. and Al-Shehbaz, I.A. (2006) Brassicaceae: Species Checklist. Plant Systematics and Evolution, 259, 249-258. https://doi.org/10.1007/s00606-006-0422-0
Chaudhary, S.A. (1999) Flora of the Kingdom of Saudi Arabia. Ministry of Agriculture and Water, Riyad, Kingdom of Saudi Arabia, Volume 1, 462-523.
Mandaville, J.P. (1990) Flora of Eastern Saudi Arabia. Kegan Paul International, London&N.Y. jointly with the National Commission for Wildlife Conservation and Development, Riyadh, 130-155.
Bentham, G. and Hooker, J.D. (1862) Genera Plantarum. Reeve, London, Vol. 1, 57-102.
Prantl, K. (1891) Cruciferae. In: Engler, A. and Prantl, K., Eds., Die Natürlichen Pflanzenfamilien, Engelmann, Leipzig, 145-208.
Hayek, A.V. (1911) Entwurf eines Cruciferae-Systems auf Pflogenetischer Grundlage. Beihefte zum Botanischen Centralblatt, 27, 127-335.
Post, G. (1932) Flora of Syria, Palestine and Sinai. American Press, Beirut, Vol. 1, 500-574.
Schulz, O.E. (1936) Cruciferae. In: Engler, A. and Prantl, K., Eds., Die natürlichen Pflanzenfamilien, Vol.17B, Engelmann, Leipzig, 227-658.
Janchen, E. (1942) Das system der Cruciferen. Oesterreichische botanische Zeitschrift, 91, 1-18. https://doi.org/10.1007/BF01257342
Zohary, M. (1966) Clypeola L. In: Zohary, M., Ed., Flora Palaestina, Vol. 1, Israel Academy of Sciences and Humanities, Jerusalem, 289-294.
Davis, P.H. (1975) Flora of Turkey Edinburgh. Edinburgh University Press, Edinburgh, 158-174.
Avetisian, V.E. (1976) Some Modifications of the System of the Family Brassicaceae. Botaniceskij Zhurnal, 61, 1198-1203.
Avetisian, V.E. (1983) The System of the Family Brassicaceae. Botaniceskij Zhurnal, 68, 1297-1305.
Al-Shehbaz, I.A. (1984) The Tribes of Cruciferae (Brassicaceae) in the Southeastern United States. Journal of the Arnold Arboretum, 65, 343-373.
Al-Shehbaz, I.A., Beilstein, M.A. and Kellogg, E.A. (2006) Systematics and Phylogeny of the Brassicaceae: An Overview. Plant Systematics and Evolution, 259, 89-120. https://doi.org/10.1007/s00606-006-0415-z
Zunk, K., Mummenhoff, K. and Hurka, H. (1999) Phylogenetic Relationships in Tribe Lepidieae (Brassicaceae) Based on Chloroplast DNA Restriction Site Variation. Canadian Journal of Botany, 77, 1504-1512.
Bailey, C.D., Price, R.A. and Doyle, J.J. (2002) Systematics of the Halimolobine Brassicaceae: Evidence from Three Loci and Morphology. Systematic Botany, 27, 318-332.
Koch, M., Al-Shehbaz, I.A. and Mummenhoff, K. (2003) Molecular Systematics, Evolution, and Population Biology in the Mustard Family (Brassicaceae). Annals of the Missouri Botanical Garden, 90, 151-171. https://doi.org/10.2307/3298580
Warwick, S.I. and Sauder, C. (2005) Phylogeny of Tribe Brassiceae (Brassicaceae) Based on Chloroplast Restriction Site Polymorphisms and Nuclear Ribosomal Internal Transcribed Spacer and Chloroplast trnL Intron Sequences. Canadian Journal of Botany, 83, 467-483. https://doi.org/10.1139/b05-021
Beilstein, M.A., Al-Shehbaz, I.A. and Kellogg, E.A. (2006) Brassicaceae Phylogeny and Trichome Evolution. American Journal of Botany, 93, 607-619. https://doi.org/10.3732/ajb.93.4.607
Warwick, S.I., Sauder, C.A., Al-Shehbaz, I.A. and Jacquemoud, F. (2007) Phylogenetic Relationships in the Tribes Anchonieae, Chorisporeae, Euclidieae, and Hesperideae (Brassicaceae) Based on Nuclear Ribosomal Its DNA Sequences. Annals of the Missouri Botanical Garden, 94, 56-78. https://doi.org/10.3417/0026-6493(2007)94[56:PRITTA]2.0.CO;2
German, D.A. and Al-Shehbaz, I.A. (2008) Five Additional Tribes (Aphragmeae, Biscutelleae, Calepineae, Conringieae and Erysimeae) in the Brassicaceae (Cruciferae). Harvard Papers in Botany, 13, 165-170. https://doi.org/10.3100/1043-4534(2008)13[165:FATABC]2.0.CO;2
Warwick, S.I., Sauder, C.A., Mayer, M.S. and Al-Shehbaz, I.A. (2009) Phylogenetic Relationships in the Tribes Schizopetaleae and Thelypodieae (Brassicaceae) Based on Nuclear Ribosomal ITS Region and Plastid ndhF DNA Sequences. Botony, 87, 961-985. https://doi.org/10.1139/B09-051
Franzke, A., German, D., Al-Shehbaz, I.A. and Mummenhoff, K. (2009) Arabidopsis Family Ties: Molecular Phylogeny and Age Estimates in the Brassicaceae. Taxon, 58, 425-437.
Mohamed, A.H. (2009) Molecular Systematic of Some Brassicaceae Taxa in Egypt Based on Electrophoretic Isoenzymes Pattern and RAPD Markers. Australian Journal of Basic and Applied Sciences, 3, 1499-1511.
Couvreur, T.L.P., Franzke, A., Al-Shehbaz, I.A., Bakker, F., Koch, M.A. and Mummenhoff, K. (2010) Molecular Phylogenetics, Temporal Diversification and Principles of Evolution in the Mustard Family (Brassicaceae). Molecular Biology and Evolution, 27, 55-71. https://doi.org/10.1093/molbev/msp202
Liu, L., Zhao, B., Tan, D. and Wang, J. (2012) Phylogenetic Relationships of Brassicaceae Species Based on Matk Sequences. Pakistan Journal of Botany, 44, 619-626.
Özüdogru, B., Akaydin, G., Erik, S., Al-Shehbaz, I. and Mummenhoff, K. (2015) Phylogeny, Diversification and Biogeographic Implications of the Eastern Mediterranean Endemic Genus Ricotia (Brassicaceae). Taxon, 64, 727-740. https://doi.org/10.12705/644.5
Li, Y., Feng, Y., Lv, G.H., Liu, B. and Qi, A. (2015) The Phylogeny of Alyssum (Brassicaceae) Inferred from Molecular Data. Nordic Journal of Botany, 33, 715-721. https://doi.org/10.1111/njb.00588
Heywood, V.H. (1971) Scanning Electron Microscopy. Systematic and Evolutionary Applications. Journal of the Arnold Arboretum, 65, 343-373.
Cole, G.T. and Behnke, H.D. (1975) Electron Microscopy and Plant Systematics. Taxon, 24, 3-15. https://doi.org/10.2307/1218989
Barthlott, W. (1981) Epidermal and Seed Surface Applicability and Some Evolutionary Aspects. Nordic Journal of Botany, 1, 345-355. https://doi.org/10.1111/j.1756-1051.1981.tb00704.x
Barthlott, W. (1984) Microstructural Features of Seed Surface. In: Heywood, V.H. and Moore, D.C., Eds., Current Concepts in Plant Taxonomy, Academic Press, London, 95-105.
Gunn, C.R. (1981) Seed Topography in the Fabaceae. Seed Science and Technology, 9, 737-757.
Zeng, C.L., Wang, J.B., Liu, A.H. and Wu, X.M. (2004) Seed Coat Microsculpturing Changes during Seed Development in Diploid and Amphiploid Brassica Species. nnals of Botany, 93, 555-566. https://doi.org/10.1093/aob/mch080
Musil, A.F. (1948) Distinguishing Species of Brassica by Their Seeds. U.S. Dept. Agr. Washington, D.C. Misc. Pub. No. 643, 35.
Murley, M.R. (1951) Seeds of the Cruciferae of North Eastern America. American Middle Naturalichen, 46, 1-81. https://doi.org/10.2307/2421948
Berggren, G. (1962) Reviews on the Taxonomy of Some Species of the Genus Brassica, Based on their Seeds. Svensk Botanisk Tidskrift, 56, 65-135.
Vaughan, J.G. and Whitehouse, J.M. (1971) Seed Structure and the Taxonomy of Cruciferae. Botanical Journal of the Linnean Society, 64, 383-409. https://doi.org/10.1111/j.1095-8339.1971.tb02153.x
Stork, A.L. (1971) Seed Characters in European Taxa of Malcolmia R. Br. Svensk Botanisk Tidskrift, 65, 283-292.
Stork, A.L. (1972) Seeds and Seedlings of Malcolmia sens. lat. Svensk Botanisk Tidskrift, 66, 417-433.
Stork, A.L. and Wüest, J. (1978) SEM Studies of Seed Coats in Malcolmia (Cruciferae). Archives des Sciences Physiques et Naturelles, 31, 229-237.
Jonsell, B. (1975) Lepidium L. (Cruciferae) in Tropical Africa. A Morphological, Taxonomical and Phytogeographical Study. Botaniska Notiser, 128, 151-163.
Prasad, K. (1976) Seed Coat Structure and Development in Certain Species of Cruciferae. New Botanist, 3, 95-103.
Mulligan, G.A. and Bailey, L.G. (1976) Seed Coat of Some Brassica and Sinapis Weeds and Cultivated in Canada. Economic Botany, 30, 143-148. https://doi.org/10.1007/BF02862959
Stork, A.L., Songerup, S. and Wüest, J. (1980) Seed Characters in Brassica Section Brassica and Some Related Groups. Candollea, 35, 421-450.
Jonsell, B. (1979) New Taxa of Cruciferae from East Tropical Africa and Madagascar. Botaniska Notiser, 132, 521-535.
Stork, A.L. and Wüest, J. (1980) Morettia DC. (Cruciferae): A Morphological and Taxonomical Study. Boletim da Sociedade Broteriana, 53, 241-273.
Jonsell, B. (1986) A Monograph of Farsetia (Cruciferae). Systematic Botany, 25, 1-106.
Fayed, A.A. and El Naggar, S.M. (1988) Taxonomic Studies on Cruciferae in Egypt 2. Taxonomic Significance of the Seed Coat Sculpture in Species of Tribe Brassiceae. Taeckholmia, 11, 87-95.
Fayed, A.A. and El Naggar, S.M. (1996) Taxonomic Studies on Cruciferae in Egypt 4. Seed Morphology and Taxonomy of the Egyptian Species of Lepidieae. Bulletin of the Faculty of Science. Assiut University, 25, 43-50.
Koul, K., Ranjna, N. and Raina, S.N. (2000) Seed Coat Microsculpturing in Brassica and Allied Genera Subtribes Brassicinae, Raphaninae, Moricandiinae). Annals of Botany, 86, 85-97. https://doi.org/10.1006/anbo.2000.1197
Abdel Khalik, K. and Maesn, V. (2002) Seed Morphology of Some Tribes of Brassicaceae Implication for Taxonomy and Species Identification for the Flora of Egypt. Blumea, 47, 363-383.
Tantawy, M.E., Khalifa, S.F., Hassan, S.A. and Al-Rabiai, G.T. (2004) Seed Exomorphic Characters of Some Brassicaceae (LM and SEM Study). International Journal of Agriculture & Biology, 6, 821-830.
El Naggar, S.M. (2005) Seed Coat Micro-Sculpturing and the Systematic of the Egyptian Brassicaceae (Magnoliopsida). Flora Mediteranea, 15, 581-598.
Kasem, W.T., Ghareeb, A. and Marwa, E. (2011). Seed Morphology and Seed Coat Sculpturing of 32 Taxa of Family Brassicaceae. Journal of American Studies, 2, 166-178.
EL-Habashy, I.E., Abdel-Hameed1, U.K., Abu Ziada, M.E.A. and Abd El-Moteleb, M. (2013) Inter-Specific Relationships among Some Taxa of Brassicaceae (L) Based on Macromorphology, Lamina Architecture, Stomatography and Palynological Criteria. Vegetos—An International Journal of Plant Research, 26, 1-14. https://doi.org/10.5958/j.2229-4473.26.2.047
Bona, M., (2013) Seed-Coat Microsculpturing of Turkish Lepidium (Brassicaceae) and Its Systematic Application. Turkish Journal of Botany, 37, 662-668.
Özüdogru, B., Akaydinb, G., Erika, S. and Mummenhoff, K. (2016) Seed Morphology of Ricotia (Brassicaceae) and Its Phylogenetic and Systematic Implication. Flora —Morphology, Distribution, Functional Ecology of Plants, 222, 60-67. https://doi.org/10.1016/j.flora.2016.03.013
Hani, M., Lebazda, R. and Fenni1, M. (2017) Studies of Morphological Characteristics and Production of Seeds Weeds of Species of Family Brassicaceae (Cruciferous) in Setifian High Plateau, Algeria. Annual Research & Review in Biology, 12, 1-9. https://doi.org/10.9734/ARRB/2017/33473
Barthlott, W. (1990) Scanning Electron Microscopy of the Epidermal Surface in Plants. In: Claugher, D., Ed., Scanning Electron Microscopy in Taxonomy and Functional Morphology, Clarendon Press, Oxford, 69-94.
Stearn, W.T. (1992) Botanical Latin. David & Charles Pub, London.
Bailey, C.D., Koch, M.A., Mayer, M., Mummenhoff, K., O’Kane, S.L., Warwick, S.I., Windham, M.D. and Al-Shehbaz, I.A. (2006) Toward a Global Phylogeny of the Brassicaceae. Molecular Biology and Evolution, 23, 2142-2160. https://doi.org/10.1093/molbev/msl087
Dudley, T.R. and Cullen, J. (1965) Studies in the Old World Alysseae Hayek. Feddes Repertorium, 71, 218-228. https://doi.org/10.1002/fedr.19650710105
Warwick, S.I., Sauder, C.A. and Al-Shehbaz, I.A. (2008) Phylogenetic Relationships in the Tribe Alysseae (Brassicaceae) Based on Nuclear Ribosomal ITS DNA Sequences. Botany-Botanique, 86, 315-336. https://doi.org/10.1139/B08-013
German, D.A. (2009) A Check-List and the System of the Cruciferae of Altai. Komarovia, 6, 80-88.
Koch, M.A., Dobes, C., Schmickl, R., Klimes, L. and Lysak, M.A. (2007) Supernetwork Identifies Multiple Events of Plastid trnF(GAA) Pseudogene Evolution in the Brassicaceae. Molecular Biology and Evolution, 24, 63-73. https://doi.org/10.1093/molbev/msl130
Khosravi, A.R., Mohsenzadeh, S. and Mummenhoff, K. (2009) Phylogenetic Relationships of Old World Brassicaceae from Iran Based on Nuclear Ribosomal DNA Sequences. Biochemical Systematics and Ecology, 37, 106-115. https://doi.org/10.1016/j.bse.2009.01.010
Beilstein, M.A., Al-Shehbaz, I.A., Mathews, S. and Kellogg, E.A. (2008) Brassicaceae Phylogeny Inferred from Phytochrome A and ndhF Sequence Data: Tribes and Trichomes Revisited. American Journal of Botany, 95, 1307-1327. https://doi.org/10.3732/ajb.0800065
Mitchell, A.D. and Heenan, P.B. (2000) Systematic Relationships of New Zealand Endemic Brassicaceae Inferred from nrDNA ITS Sequence Data. Systematic Botany, 25, 98-105. https://doi.org/10.2307/2666676
Galloway, G.L., Malmberg, R.L. and Price, R.A. (1998) Phylogenetic Utility of the Nuclear Gene Arginine Decarboxylase: An Example from Brassicaceae. Molecular Biology and Evolution, 15, 1312-1320. https://doi.org/10.1093/oxfordjournals.molbev.a025859
Koch, M. and Hurka, H. (1999) Isozyme Analysis in the Polyploid Complex Microthlaspi perfoliatum (L.) F. K. Meyer: Morphology, Biogeography and Evolutionary History. Flora, 194, 33-48. https://doi.org/10.1016/S0367-2530(17)30877-0
Koch, M.A., Haubold, B. and Mitchell-Olds, T. (2001) Molecular Systematics of the Brassicaceae: Evidence from Coding Plastidic matK and Nuclear Chs Sequences. American Journal of Botany, 88, 534-544. https://doi.org/10.2307/2657117
Al-Shehbaz, I.A. (1988) The Genera of Sisymbrieae (Cruciferae; Brassicaceae), the Southeastern United States. Journal of the Arnold Arboretum, 69, 213-237. https://doi.org/10.5962/bhl.part.2393
Warwick, S.I., Al-Shehbaz, I.A., Price, R.A. and Sauder, C. (2002) Phylogeny of Sisymbrium (Brassicaceae) Based on ITS Sequences of Nuclear Ribosomal DNA. Canadian Journal of Botany, 80, 1002-1017. https://doi.org/10.1139/b02-089
Warwick, S.I., Al-Shehbaz, I.A., Sauder, C., Harris, J.G. and Koch, M. (2004) Phylogeny of Braya and Neotorularia (Brassicaceae) Based on Nuclear Ribosomal Internal Transcribed Spacer and Chloroplast trnL Intron Sequences. Canadian Journal of Botany, 82, 376-392. https://doi.org/10.1139/b04-012
Warwick, S.I., Al-Shehbaz, I.A., Sauder, C., Murray, D.F. and Mummenhoff, K. (2004) Phylogeny of Smelowskia and Related Genera (Brassicaceae) Based on Nuclear ITS DNA and Chloroplast trnL Intron Sequences. Annals of the Missouri Botanical Garden, 91, 99-123.
Zunk, K., Mummenhoff, K. and Hurka, H. (1993) Chloroplast Restriction Site Variation in the Brassicaceae, Tribe Lepidieae. Plant Molecular Evolution Newsl, 3, 40-44.
Price, R.A., Palmer, J.D. and Al-Shehbaz, I.A. (1994) Systematic Relationships of Arabidopsis: A Molecular and Morphological Approach. In: Meyerowitz, E. and Somerville, C., Eds., Arabidopsis, Cold Spring Harbor Press, Cold Spring Harbor, NY, 719.
Les, D.H. (1994) Molecular Systematic and Taxonomy of Lake Cress (Neobeckia aquatica; Brassicaceae), an Imperilled Aquatic Mustard. Aquatic Botany, 49, 149-165. https://doi.org/10.1016/0304-3770(94)90035-3
Franzke, A., Pollman, K., Bleeker, W., Kohrt, R. and Hurka, H. (1998) Molecular Systematics of Cardamine and Allied Genera (Brassicaceae): ITS and Non-Coding Chloroplast DNA. Folia Geobotanica, 33, 225-240. https://doi.org/10.1007/BF03216204
Sweeney, P.W. and Price, R.A. (2000) Polyphyly of the Genus Dentaria L. (Brassicaceae): Evidence from trnL Intron and ndhF Sequence Data. Systematic Botany, 25, 468-478. https://doi.org/10.2307/2666690
Bleeker, W., Franzke, A., Pollmann, K., Brown, A.H.D. and Hurka, H. (2002a) Phylogeny and Biogeography of Southern Hemisphere High-Mountain Cardamine Species (Brassicaceae). Australian Systematic Botany, 15, 575-581. https://doi.org/10.1071/SB01026
Bleeker, W., Weber-Sparenberg, C. and Hurka, H. (2002b) Chloroplast DNA Variation and Biogeography in the Genus Rorippa Scop. (Brassicaceae). Plant Biology, 4, 104-111. https://doi.org/10.1055/s-2002-20442
Warwick, S.I. and Black L.D. (1997) Molecular Phylogenies from Theory to Application in Brassica and Allies (Tribe Brassiceae, Brassicaceae). Opera Botanica, 132, 159-168.
Warwick, S.I. and Black, L.D. (1997b) Phylogenetic Implications of Chloroplast DNA Restriction Site Variation in Subtribes Raphaninae and Cakilinae (Brassicaceae, Tribe Brassiceae). Canadian Journal of Botany, 75, 960-973. https://doi.org/10.1139/b97-107
Lysak, M.A., Koch, M.A., Pecinka, A. and Schubert, I. (2005) Chromosome Triplication Found across the Tribe. Genome Research, 15, 516-525. https://doi.org/10.1101/gr.3531105
Mandáková, T. and Lysak, M.A. (2008) Chromosomal Phylogeny and Karyotype Evolution in x = 7 Crucifer Species (Brassicaceae). Plant Cell, 20, 2559-2570. https://doi.org/10.1105/tpc.108.062166