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Fruit and Seed Morphology in Galium L. (Rubiaceae) and Its Importance for Taxonomic Identification

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Fruit and Seed Morphology in Galium L. (Rubiaceae) and Its Importance for Taxonomic Identification Color profile: Disabled Composite 150 lpi at 45 degrees Acta Bot. Croat. 67 (1), 1–20, 2008 CODEN: ABCRA25 ISSN 0365–0588 Fruit and seed morphology in Galium L. (Rubiaceae) and its importance for taxonomic identification KADRY ABDEL KHALIK1,MONIER ABD EL-GHANI2,AHMED EL KORDY1 ¹ Sohag University, Faculty of Science, Botany Department, Sohag 82524, Egypt ² Cairo University, Faculty of Science, The Herbarium, Giza 12613, Egypt Fruit (mericarp) and seed morphology of 11 species and two subspecies of Galium were examined with light microscope and scanning electron microscope. Macro-and micro- -morphological characters, including fruit and seed shape, colour, size, surface, epidermal cell shape, anticlinal boundaries, outer periclinal cell wall and relief of outer cell walls, are presented. Four different types of mericarp surface are described. Three types of anticlinal cell wall boundaries of seed are recognized and three different shapes of outer periclinal cell wall are described. The secondary sculpture of the cell wall varies from micro-papil- late to micro-reticulate, and smooth to fine- or coarse-folded. The fruit and seed character- istics could be used for taxonomic identification. Key words: Fruit, seed, morphology, Rubiaceae, Galium Introduction Rubiaceae form the fourth largest angiosperm family after Asteraceae, Orchidaceae and Fabaceae and comprise more than 10 000 species belonging to 640 genera. Rubiaceae are usually considered a natural monophyletic and easily circumscribed family (VERD- COURT 1958, BREMEKAMP 1966, ROBBRECHT 1988). In the flora of Egypt Rubiaceae are rep- resented by eight genera, viz. Kohautia, Oldenlandia, Galium, Valantia, Callipeltis, Cru- cianella, Pterogaillonia and Rubia (BOULOS 1995, 2000). Cruciata articulata is reported as a new record to the flora of Egypt, so the number of genera in Rubiaceae was increased to nine (ABDEL KHALIK and BAKKER 2007). Galium is one of the largest genera of Rubiaceae (WILLIS 1985, MABBERLEY 1987). In Egypt Galium is represented by 12 species (TACKHOLM 1974). BOULOS (2000) reported on- ly 10 species of Galium. In the Flora orientalis, BOISSIER (1881) reported 91 species of the genus Galium divided into three sections: section (I) Eugalium, section (II) Aparine, and section (III) Cruciata. Section (I) Eugalium is classified into four subsections: 1. Platygalia, 2. Leiogalia, 3. Chromogalia, and 4. Ceratocapa. * Corresponding author, e-mail: [email protected] ACTA BOT. CROAT. 67 (1), 2008 1 U:\ACTA BOTANICA\Acta-Botan 1-08\01 Abedl.vp 3. travanj 2008 11:18:03 Color profile: Disabled Composite 150 lpi at 45 degrees ABDEL KHALIK K., ABD EL-GHANI M., EL KORDY A. Section (II) Aparine is classified into five subsections: 1. Camptopoda, 2. Leucapari- nea, 3. Xanthaparinea, 4. Bracteata, and 5. Apera. Subsection Leiogalia includes G. mollu- go, subsection Chromogalia includes the following species: G. sinaicum and G. canum, subsection Ceratocapa includes G. ceratocarpum, subsection Camptopoda includes G. tri- corne and G. ceratopodum, subsection Leucaparinea includes G. aparine and G. spurium, subsection Xanthaparinea includes the following species: G. parisiense G. nigricans, G. decaisnei and G. setaceum,and subsection Apera includes G. murale. Most taxonomists agree that the macro- and microstructure of seeds are significant for the classification of Angiosperm taxa. During the last decades, scholars have applied SEM to morphological studies of seeds and small fruits. Micro-morphology and ultra-structural data have contributed useful infor- mation concerning the evolution and for the classification of seed plants and play an impor- tant role in the modern synthetic systems of Angiosperms (HEYWOOD 1971, DAHLGREN 1979–80). Several studies focused on intrageneric seed coat variation (CHUANG and HECKARD 1972, HEYN and HERRNSTEDT 1977, CLARK and JERNSTEDT 1978, WOFFORD 1981, JUAN et al. 2000, ABDEL KHALIK 2006) or on variation among several closely related genera (MUSSELMAN and MANN 1976; SEAVEY et al. 1977; CHANCE and BACON 1984; MATTHEWS and LEVINS 1986; FAYED and EL NAGGAR 1988, 1996; SHANMUKHA and LEELA 1993; KANAK SAHAI et al. 1997; KARAM 1997; KOUL et al. 2000). Less commonly, SEM level va- riation was used to place taxa into tribes (WHIFFIN and TOMB 1972, ABDEL KHALIK and VAN DER MAESEN 2002). More information is available concerning seed forms and internal structures (CORNER 1976, KABIL et al. 1980), seed size (ANISZEWWSKI et al. 2001), seed colours (BERGGREN 1962, DAHLGREN and CLIFFORD 1982, BARTHLOTT 1984), and epidermal cell patterns and distribution of trichomes, glands, and stomata (STACE 1965). The distribution of epidermal cells is important between species and genus level (BARTHLOTT 1981, 1984), while outer periclinal walls are also a good diagnostic for the lowest taxonomic categories (BARTHLOTT 1981, 1984, ABDEL KHALIK et al. 2002, ABDEL KHALIK 2006). The present investigations deal with the micro- and macro-morphological (LM and SEM) characters of fruits (mericarps) and seeds for 11 species and two subspecies of the genus Galium from the Rubiaceae in Egypt, to show the ranges of variability in fruit (meri- carp) and seed characters in order to establish their usefulness for future taxonomic work. Materials and methods Some of the investigated seeds were collected from mature plants in Egypt during 2004–2006. The others were taken from herbarium specimens. Only mature fruits and see- ds were taken for investigation. The dried fruits were soaked in boiling water for 2–4 minu- tes, to compensate for shrinkage and examined by light microscope (Olympus type BH-2), with 10–15 seeds for each taxon being chosen to cover the range of variation. Measuremen- ts were standardized with ocular micrometer and fruits drawn with the aid of a camera luci- da. Mature fruits and seeds were mounted on stubs with double adhesive tape. The stubs were sputter-coated with gold/palladium for 4 min. in an Apolaron E 1100 ion sputtering device. After coating, the specimens were examined with a JEOL -6300 Scanning Electron 2 ACTA BOT. CROAT. 67 (1), 2008 U:\ACTA BOTANICA\Acta-Botan 1-08\01 Abedl.vp 3. travanj 2008 11:18:03 Color profile: Disabled Composite 150 lpi at 45 degrees FRUIT AND SEED MORPHOLOGY IN GALIUM Microscope, using accelerating voltages at 20–30 KV. All photomicrographs were taken at the SEM laboratory, Sohag University, Egypt. The terminology used here follows authors such as (ABDEL KHALIK et al 2002, ABDEL KHALIK 2006) with some modification by the au- thor. List of Galium fruit and seed specimens used in light and scanning electron microscope studies. 1. Galium aparine L.: Egypt, Gebel Elba, Gebel Ekwal, J. R. Shabetai 250 (K) 2. Galium canum Req.: Palestine, Wadi Sawaanit, in rocks, P.H. Davis 5038 (K) 3. Galium ceratopodum Boiss.: Egypt, Sinai, W. Gebal region, Ain Altofaha, K. Ab- del Khalik 1956 (SHG). 4. Galium mollugo L.: Netherlands, Gelderland, NW of Wolfheze.Open low vegeta- tion on sandy soil, C.C.H. Jongkind 5226 (WAG) 5. Galium murale L.: Egypt, in cultivated land, near Maruit, Letuneux 197 (K) 6. Galium nigricanse Boiss.: Iran, Kordestan, Sanandaj, rolling limestone hills alt. 1800, M. Jacobs 6938 (BR) 7. Galium parisiense L.: France, 5 km E of Cavarllon, W. J. Reijnders 1144 (L) 8. Galium setacium Lam. subsp. setaceum: Egypt, Sinai, Gebel Serbal, alt. 1684, Fayed et al. 1946–2033 (SHG). 9. Galium setaceum subsp. decaisnei.Boiss.: Egypt, Gebel Elba, Wadi Drawina, K. AbdelKhaliks.n.(SHG). 10. Galium sinaicum (Delile ex Decne.) Boiss.: Egypt, Sinai, Gebel Serbal, alt. 1450, Fayed et al. 2034–2035 (SHG). 11. Galium spurium L. subsp. spurium: Egypt, Sinai W. Gebel region, Ain Altofaha, K. AbdelKhaliks.n.(SHG). 12. Galium spurium subsp. africanum Verdc.: Egypt, Gebel Elba, Wadi Drawina, K. Abdel Khalik 3935 K. (SHG). 13. Galium tricornutum Dandy: Egypt, Alexandria, De Bullemont s.n. (BR). Results Fruit (mericarp) characters Fruit characters are very important for interspecies distinction in the genus Galium. Fruits are of a schizocarpic nature, composed of two mericarps, each mericarp having only one seed. Mericarp shape Mericarp shape among the investigated taxa showed a large variation. Mericarps vary from cylindrical or globose, globose to sub globose or reniform (Tab. 1). Mericarp shape showed significant differences among species, but no significant differences among sub- species or varieties. However they are cylindrical in Galium murale (Figs. 2.5, 4.5); globo- se in G. aparine and G. tricornutum, (Figs.1.1, 4.1, 3.13, 5.14); sub globose in G. ceratopo- dum, G. spurium subsp. spurium and subsp. africanum (Figs. 1.3, 4.3, 3.11, 3.12, 5.12, 5.13); reniform in G. canum, G. mollugo, G. nigricans, G. parisiense, G. setacium subsp. setacium and subsp. decaisnei, G. sinaicum (Figs. 1.2, 1.4; 2.6, 2.7, 2.8, 3.10). ACTA BOT. CROAT. 67 (1), 2008 3 U:\ACTA BOTANICA\Acta-Botan 1-08\01 Abedl.vp 3. travanj 2008 11:18:03 Composite 150lpiat45degrees Color profile:Disabled 3. travanj 200811:18:03 U:\ACTA BOTANICA\Acta-Botan 1-08\01Abedl.vp A 4 ACTA BOT. CROAT. 67 (1), 2008 Tab. 1. Mericarp and seed description of Galium species. BDEL Taxon seed Seed Seed Epidermal Anticlinal Periclinal Mericarp Meri- Mericarp K shape size(mm) color cell shape boundries cell wall size (mm) carp surface HALIK color. G. aparine L. Globose 2.8–4.5 ´ 2.8–4.5 Dark 5,6 gonals, Raised-channeled, Flat to 3–5´ 3–5 Dark Hooked hairs K., A brown elongate in straight, smooth slightly con- arising from BD one direction cave, micro- tubercle-like E papillate base L -G G. canum Req. Reniform 0.5–0.8 ´ 0.3–0.5 Dark Polygonal, Slightly raised, Flat, 0.7–1 ´ 0.5–0.7 Dark Long white HANI brown elongate in straight to smooth brown simple one direction slightly sinous, straight hairs M., E smooth ´ ´ L G. ceratopodum Globose 1.6–2.2 1.3–2.0 Dark Polygonal Raised, straight Flat to slig- 1.7 –2.5 1.5 –2.5 Dark Tuberculate K Boiss.
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