Bull 435 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Bull. Iraq nat. Hist. Mus. (2023) 17 (3): 435-458. https://doi.org/10.26842/binhm.7.2023.17.3.0435 ORIGINAL ARTICLE GENUS RETAMA RAF., 1838 (FABALES, FABACEAE): TAXONOMIC REVISION IN EGYPT SUPPORTED BY MOLECULAR FINGERPRINTING Reham A. Youssef, Wafaa M. Amer* and Azza B. Hamed Department of Botany and Microbiology, Faculty of Science, Cairo University, Giza 12613, Egypt. ⃰ Corresponding author: wamer@sci.cu.edu.eg Recived Date: 04 Febraury 2023, Accepted Date 18 March 2023, Published Date:20 June 2023 This work is licensed under a Creative Commons Attribution 4.0 International License ABSTRACT In Egypt, the taxonomic identity of the taxa under genus Retama Raf., 1838 (Fabaceae) is still unclear and its morphological resemblance precludes its identification. The current study aims to resolve the taxonomic identity of the species belonging to genus Retama in Egypt and clarify the morphological, molecular, and geographic distribution characterised each species. To achieve these goals, the fresh and herbarium Retama specimens were used for morphological investigations using 94 macro-morphological characters while the Inter Simple Sequence Repeats (ISSR) markers were used for the molecular identity. This revision revealed the identification of two distinct species namely: Retama raetam (Forsskk.) Webb. and Retama monosperma (L.) Boiss with five forms under R. raetam (Forms 2, 4, 6, 7, and 8). In addition to Form 5 from under R. monosperma; the morphological and molecular identities of forms showed variations in the meanwhile it's clustering were congruent. The geographic distribution of the Retama taxa in Egypt was elucidated. This study highlights the urgent need for Retama species conservation, due its vulnerability to climatic change. Keywords: Flora of Egypt, Retama monosperma, R. raetam, Forms, ISSR, Vulnerable species. INTRODUCTION Globally, Fabaceae is among the most prominent families of Angiospermae, it includes more than 19500 species grouped into approximately 770 genera (Bruneau et al., 2013; Abusaief and Boasoul, 2021). Economically, Fabaceae is in the second position to Poaceae (Mabberley, 1997; Yahara et al., 2013). The Egyptian flora comprises a unique mixture of native African and Asiatic species with over 2079 species (Amer, 2008), where Fabaceae is represented by 212 species (Boulos, 1999). Genus Retama Raf.m 1838 (Fabaceae), previously named Lygos Adanson (GBIF Secretariat, 2022); Spartium L. and Retama Boiss. (Lopez et al., 1998; Benmiloud- Mahieddine et al., 2011), distributed in the western Mediterranean region (Chiapella et al., BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Iraq Natural History Research Center & Museum, University of Baghdad https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home Copyright © Bulletin of the Iraq Natural History Museum Online ISSN: 2311-9799-Print ISSN: 1017-8678 https://doi.org/10.26842/binhm.7.2023.17.3.0435 https://orcid.org/0000-0002-0083-1619 https://orcid.org/0000-0003-0126-6719 https://orcid.org/0000-0001-7971-1048 mailto:wamer@sci.cu.edu.eg https://creativecommons.org/licenses/by/4.0/ https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home 436 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 2009). It comprises four closely related species included in the tribe Genisteae, namely R. monosperma (L.) Boiss., R. sphaerocarpa Boiss., R. raetam Webb and Berthel., and R. dasycarpa Coss. It also contains several subspecies and varieties distributed in North Africa and their diversification center is in the western Mediterranean basin, Canary Islands, and Southern Europe (Benmiloud-Mahieddine et al., 2011). The morphological resemblance between these four species in this genus obstructs their taxonomical identification (Käss and Wink, 1997; Pardo et al., 2004; León-González et al., 2018). Retama species can tolerate severe drought conditions in various ecosystems, including deserts, coastal dunes, and maquis (León-González et al., 2018). In Egypt, Retama grows in the desert wadis, sandy and maritime plains (Boulos, 1999, 2009; Amer et al., 2021). Economically, Retama species are important fodder for livestock and considered as alternative to the conventional forage species in the arid Mediterranean ecosystems (Barakat et al., 2013). They are used for protection and stabilization against wind or water erosions, irradiation, and overheating (Maghrani et al., 2005; Morsy et al., 2015). Also, they are widely used for construction and ornamental purposes (Barakat et al., 2013); and have several medicinal benefits for humans (Maghrani et al., 2005; Awen et al., 2011; Djeddi et al., 2013; Muñoz Vallés et al., 2013; Chouaibi et al., 2019). The taxonomic identity of the taxa grouped under genus Retama Raf. is still unresolved. Several reports have shown divergence such as the following: in Algeria, this genus is represented by R. raetam Webb, R. spherocarpa (L.) Boiss. and R. monosperma (L.) Boiss. subsp. bovei (Spach.) Maire (Quezel and Santa, 1962); in Europe, it represent by three species, according to Tutin et al. (1968) are Lygos raetam (Forssk.) Heywood (syn. Retama raetam (Forssk.) Webb & Berth.) subsp. gussoniei (Webb) Heywood, L. spherocarpa (L.) Heywood. (syn. Retama spherocarpa (L.) Boiss.) and L. monosperma (L.) Heywood. (syn. Retama monosperma (L.) Boiss.) or only L. monosperma (L.) Heywood. (syn. R. monosperma (L.) Boiss. (Polunin, 1969). In Palestine, Zohary (1972) reported the presence of R. raetam (Forssk.) Webb with two varieties, var. raetam (Forssk.) Webb and var. sarcocarpa Zoh. While, in Lebanon and Syria, this genus is represented by two species R. raetam (Forssk.) Webb. and Retama duriaei (Spach) Willd. (Mouterde, 1966), and only one species, R. raetam (Forssk.) Webb was seen in Libya (Jafri, 1980). The literature data of the previous taxonomic treatment of genus Retama in Egypt increased its complexity. Where Täckholm (1974), reported it as Lygos Adans which represented by Lygos raetam (Forssk.) Heywood (syn. Retama raetam (Forssk.) Webb with two varieties, var. sarcocarpa (Zoh.) Täckh. et Boulos and var. bovei (Spach) Täckh. et Boulos, in addition to three unidentified anonymous varieties in Sinai. Later, Boulos (2009), reported the genus Retama Raf. inEgypt by two species, R. raetam (Forssk.) Webb & Berthel (syn. Lygos raetam (Forssk.) Heywood) and Retama monosperma (L.) Boiss. subsp. bovei (Spach) Maire (syn. Spartium bovei Spach. and Retama bovei (Spach) Webb). The complexity of this genus in Egypt also includes some taxa, such as var. bovei, which was treated as a variety under Lygos raetam (Täckholm, 1974) and a subspecies of R. monosperma by Boulos (2009). 437 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Although the genus Retama is important in the Mediterranean region, several species have recently become vulnerable to climate change, mainly rainfall shortage and temperature increase resulting in the poleward shift of the xeric species. Among them, R. raetam, which recently showed distribution, density, and genetic decline; accordingly, it needs urgent conservation (Amer et al., 2020; Elshayeb, 2020). It is crucial to understand the genetic profile of Retama taxa (species/populations/taxa) in Egypt to promote future conservation strategies. Molecular markers are powerful tools for detecting polymorphism in related species as they are not influenced by developmental stages or environmental factors (Helm et al., 2009; Karimi et al., 2009). ISSR molecular markers are among the molecular tools that can distinguish closely related species, even at the interspecies level (Zietkiewicz et al., 1994; Abdelhameed et al., 2020; Amer et al., 2021), and identify the diversity within and among populations (Mosaferi et al., 2015; Minaeifar et al., 2016; Abdelhameed et al., 2020; Muraseva and Guseva, 2021). The current study aims to resolve the following points concerning genus Retama in Egypt, naming the number of taxa representing this genus; clarifying the morphological, genetic and geographic features of these taxa. MATERIALS AND METHODS Plant materials: The fresh Retama specimens (63 from 21 populations) and their seeds were collected and examined from the current geographical range in Egypt between 2020–2022 (Tab. 1) and a distribution map was conducted using the geographic position system. The dry specimens (220 specimens) were kept in the Egyptian herbaria at Cairo University (CAI), National Research Center (CAIRC), Desert Research Institute (CAIH), and Agricultural Museum (CAIM) and the online virtual herbaria (Royal Botanic Garden Herbarium at Kew (K)), JSTOR Global Plants database and the Global Biodiversity Information Facility database were additionally studied. The acronyms for herbaria follow Thiers (2019). Fresh samples were preserved in formalin-aceto-alcohol (FAA: 50 mL ethyl alcohol, 10 mL formaldehyde, 5 mL glacial acetic acid, and 35 mL distilled water (Amer et al., 2021). Morphological investigation: Morphological investigations were conducted based on 94 macro-morphological characters including stem, leaves, flowers, inflorescences, fruits, and seeds of the fresh and herbarium Retama specimens (see Appendix 1). The identified Retama taxa were based on previous taxonomic treatments (Quezel and Santa, 1962; Tutin et al., 1968; Polunin, 1969; Zohary, 1972; Täckholm, 1974; Jafri, 1980; Wickens, 1998; Boulos, 1999, 2009). Molecular investigation: Genomic DNA was extracted from juvenile leaf samples according to Abdelhameed et al. (2020) and purified them using the Plant DNeasy Mini Kit (Qiagen, Santa Clarita, CA). Twelve ISSR primers were used and sequenced as listed in Table (3). The PCR mixture (25 μL) included 30 ng template DNA ( this was applied to triple samples/taxa), 2.5 μL of 10X PCR buffer, 1.5 μL of 25-mM MgCl2, 2.5 μL of the dNTPs mix, 30 pm of ISSR primer, and 1 U Taq DNA polymerase (Promega, WI, USA). DNA amplification was 438 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 performed according to Amer et al. (2021) using a thermal cycler (Applied Biosystems, USA). The PCR cycle consisted of initial denaturation at 94 °C for 5 mins, 40 cycles of denaturation at 94 °C for 2 mins, annealing at 50 °C for 45 secs, extension at 72 °C for 2 mins, and final elongation for 7 mins at 72 °C. The PCR products were separated using 1.5 % agarose gel in 1x TBE buffer and visualized using ethidium bromide. The gels (3 per each studied taxa) were visualized under a UV-transilluminator, documented in Gel-Doc XR (Bio- Rad), and photographed. The amplicon sizes were determined using a 1 Kplus DNA ladder RTU (Genedirex). Statistical analysis: The morphological data (expressed as mean ± standard error) were analyzed and plotted using Graph Pad Prism version 8.0.1. One-way analysis of variance was used for all statistical comparisons followed by Tukey’s procedure for posthoc analysis. Values with P < 0.05 were considered significant. The radar plot was developed using the Excel program (Microsoft 365). The Jaccard’s coefficient was applied using the SPSS program (version 20 for Windows) to estimate the similarity between the studied Retama taxa based on the macro-morphological characters. A dendrogram was generated by cluster analysis using the unweighted pair group method of the arithmetic averages (UPGMA) using Past software (version 3.26 for Windows). For the molecular part the generated/amplified bands were scored based on the presence (1) and absence (0) of bands using NTSYS-PC 2.21 software (Rohlf, 2009) and indirect gradient analysis was performed by Detrended Correspondence Analysis (DCA) using Past software (version 3.26 for Windows). RESULTS The current revision of genus Retama Raf. in Egypt revealed the presence of two distinct species, Retama raetam (Forssk.) Webb & Brantel and Retama monosperma (L.) Boiss. along with five forms of Retama raetam (Forssk.) Webb (Form 2, Form 4, Form 6, Form 7, and Form 8). In addition to Form 5 of Retama monosperma (L.) Boiss. The morphology of each form was interdisciplinary and shared the features of one of the two identified species (Retama raetam and R. monosperma) with distinct dissimilar characters rendering it consistent with the typical species; accordingly they were treated as forms. The differential morphological traits were tabulated in Appendix 1, details description will be followed: Morphological features of Retama raetam (Forssk.) Webb & Berthel. Hist. Nat. Iles Canaries 3(2; 2): 56 (1842) Synonyms: Genista raetam Forssk. in Fl. Aegypt.-Arab.: 214 (1775) Lygos raetam (Forssk.) Heywood in Feddes Repert. 79: 53 (1968) Spartium raetam (Forssk.) Spach in Ann. Sci. Nat., Bot., sér. 2, 19: 288 (1843) Perennial, woody spartoid (branched from the base), “nebka-forming shrub = sand-mound forming shrub” up to 3 m in height. Stem green-grooved, branches ascending or spreading, young twigs richly branched, sparsely-densely covered with silky hair. Leaves 2–8 (-23) × 0.5–3.0 mm, simple, sericeous, deciduous, linear-oblong, entire margin and acute-obtuse apex, petiole 0.25 mm. Inflorescence: lax–dense raceme, with 1–7 flowers (up to 15 flowers in the traced Egyptian forms). Flower papilionate, 8–12 mm, pedicle 1–3 mm. Calyx 2–4 × 1.5–3 439 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. mm, green-purple, occasionally with purple spots, up to ⅓ flower length, calyx tube with 2- lipped: upper lip with 2 broader triangular teeth and the lower lip with 3 shorter teeth almost equal in length, caducous after anthesis. Corolla white occasionally with purplish tips and/or purple spots. Standard 8–12 × 5–7 mm, as long as the wings, elliptical with purple main vein, apex retuse, hairy on tips and main vein. Wings 8–12 × 1.5–2.5 mm, oblong with obtuse apex, hairy on tips and base, occasionally with spots. Keel 8–12 × 3–5 mm, oblong, apex acute, hairy on base, tips, and main vein. Stamens 10, monadelphous, connate all below into a closed tube, 6–10 mm, filaments one long and the rest random in length. Anthers 5 heart-shaped, apiculate, dorsal fixed, 0.25–0.5 mm, and 5 oval-shaped, terminal fixed, 0.5–1.0 mm. Ovary 3–6 × 1–2 mm, elliptical, attenuate on both sides. Style 5–7 mm. Stigma 0.25 × 0.25 mm, capitate. Fruit pod 10–15 × 3–7 mm green indehiscent, glabrous, elliptical attenuates gradually to a short erect peak 0.25–2.0 mm, 1-seeded. Fruit pericarp is fleshy, leathery smooth when dry. Seeds 4–6 × 3–4 mm, reniform to round, smooth, olive-green to brown (Pl. 1a-e). Flowering from January to April. Distribution mainly in the desert wadis, sandy and gravel plains in Egypt. Morphological features of Retama monosperma (L.) Boiss Voy, Bot. ESpagne 2: 144 (1840) Synonyms: Genista monosperma (L.) Lam. in Encycl. 2: 616 (1788) Lygos monosperma (L.) Heywood in Feddes Repert. 79: 53 (1968) Spartium gracile Salisb. in Prodr. Stirp. Chap. Allerton: 329 (1796), nom. superfl. Spartium monospermum L. in Sp. Pl.: 708 (1753) Perennial, woody spartoid, up to 3 m in height. Stems erect, ascending or spreading, green- grooved, young twigs sparsely-densely with silky hairs. Leaves simple, 2–7 × 0.5–1.25 mm, simple, sericeous, deciduous, linear to lyrate, entire margin and acute-obtuse apex, petiole 0.25–0.5 mm. Inflorescence lax raceme, with 1–7 flowers (up to 15 flowers in the traced Egyptian form). Flower papilionate, 7–10 mm, pedicle 0.25–1.5 mm. Calyx 2–4 × 1.5–2 mm, purple, up to ⅓ of the flower, 2-lipped: upper lip with 2 broader triangular teeth and the lower lip with 3 shorter teeth equal in length, caducous after anthesis. White corolla, standard 4–6 × 6–8 mm, shorter than the wings, orbicular with purple mid-vein, apex retuse, hairy on tips, and mid-vein. Wings 7–10 × 2–3 mm, oblong with obtuse apex, hairy along the main vein, tips, and base. Keel 7–10 × 4 mm, oblong, apex obtuse, hairy along mid-vein, tips, and base. Stamens 10, monadelphous, connate all below into a closed tube, 7–8 mm, filaments nearly equal in length. Anthers 5 heart-shaped, apiculate, dorsal fixed, 0.25–0.5 mm, and 5 oval- shaped, terminal fixed, 0.5–1 mm. Ovary 2–3 × 0.5–1 mm, ovate. Styles 4–5 mm. Stigma 0.25 × 0.25 mm, capitate. Fruit pod 10–15 × 7–10 mm, green indehiscent, glabrous, globoid abruptly short-apiculate to erect peak, 1–2 mm, 1 to 2 seeds. The pericarp is leathery smooth and wrinkled when dry. Seeds 5–8 × 4–6 mm, reniform to round, smooth, olive-green to brown (Pl. 1f-j). Flowering from January to June. Distributed along the Mediterranean coast from Sallum in the west to Rafah at the Palestinian authority borders. 440 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Identification key to the traced Egyptian Retama taxa: 1. Standard shorter than wings...………………………………………………….….….…… 2 – Standard as long as wings……….……………..………………..…………….…….......... 3 2. Fruit elliptical, peak gradually apiculate, seed yellow to orange ……....... R. raetam Form 2 – Fruit globose, peak abruptly apiculate, seed olive-green to brown…….... R. monosperma 3. Fruit length to 1.5 cm, with one seed…………………………………..………………..… 4 – Fruit length exceeds 1.5 cm, with 1–2 seeds………………..………......…….………...…5 4. Standard elliptical, calyx green-purple……………………………….………..… R. raetam – Standard rhombic, calyx purple………………………………..……..…. R. raetam Form 4 5. Flower length not exceeding 1 cm, seed olive-green to brown …..……...…………..…… 6 – Flower length up to 1.5 cm, seed yellow to orange ……………...………………………..7 6. Calyx up to ½ flower, standard elliptical……………………….……...... R. raetam Form 8 – Calyx up to ⅓ flower, standard orbicular………………………….…… R. raetam Form 7 7. Fruit globose, with inflorescence from 1 to 15 flowers…... ….…... R. monosperma Form 5 – Fruit elliptical, with inflorescence from 1 to 7 flowers.……….…...…… R. raetam Form 6 Morphological traits distinguish the Retama Forms from the identified species R. raetam Form 2: This form represents the three typical forms (1, 2 and 3) with standard shorter than the wing and the keel; fruit elliptical, with a gradually apiculate peak. Yellow to orange seeds (Pl. 2a). Diagram 1 (a-f) shows that this form is significantly different from R. raetam in both leaf length and width, standard length and keel length (P = 0.0001), wing length (P = 0.0335). R. raetam Form 4: standard as long as the wing and the keel. Standard shape rhombic (Pl. 2b) keel mid-vein white. Fruit length to 1.5 cm with one seed. Diagram 1 (b & c), shows that this form is significantly different from R. raetam in leaf width (P = 0.0019) and flower length (P = 0.0001). R.monosperma Form 5: standard is as long as the wing and the keel. Flower length up to 1.5 cm, with inflorescence from 1–15 flowers (Pl. 2c). Fruit globose exceeds 1.5 cm in length with 1–2 yellow to orange seeds (Pl. 2a). Diagram 1 (a–f) shows the significant differences from R. monosperma in leaf length and width, flower, standard, wing and keel lengths (P < 0.0001). R. raetam Form 6: standard as long as the wing and the keel. Flower up to 1.5 cm (Pl. 2d). Inflorescence 1–7 flowers. Fruit elliptical exceeds 1.5 cm with 1–2 yellow to orange seeds. Diagram (1 a, b, d and f) shows that this form is significantly different from R. raetam in leaf length and width (P < 0.0001), as well in standard length and keel length (P = 0.0053, P = 0.0015; respectively). R. raetam Form 7: standard as long as the wing and the keel. Flower not exceeding 1 cm. Calyx up to ⅓ flower (Pl. 2e). Standard orbicular (Pl. 2f). Fruit exceeds 1.5 cm with 1–2 olive-green to brown seeds. Diagram 1 (a-e) shows that form 7 is significantly different when 441 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. compared with R. raetam in leaf length and width, standard length (P < 0.0001), and wing length (P = 0.0020). R. raetam Form 8: standard as long as the wing and keel. Flower not exceeding 1 cm. Calyx up to ½ flower (Pl. 2g). Standard elliptical (Pl, 2h). Fruit exceeds 1.5 cm with 1–2 olive-green to brown seeds. Diagram 1 (a, b and f) shows that this form is significantly different from R. raetam in leaf length & width and keel length (P < 0.0001), as well as the standard length (P = 0.0006) and wing length (P = 0.0001) as shown in Diagram 1 (d and e). Morphological similarity of the identified Retama species and forms: Diagram (2) shows the UPGMA dendrogram based on 94 macro-morphological characters. It demonstrates the grouping of the studied Retama taxa into two main clusters (I and II). Cluster I included R. monosperma and R. monosperma Form 5, and cluster II included R. raetam and five forms of R. raetam (Form 2, Form 4, Form 6, Form 7, and Form 8). While the Jaccard’s coefficient (Tab, 2) was used to estimate the similarity between the studied Retama species and the forms based on the 94 macro-morphological characteristics. The similarity value between R. raetam and R. monosperma was 60.4 %. Evaluation of the similarity between Retama species and the identified forms denoted that the highest similarity (91.6 %) was between R. raetam and R.raetam Form 4. Also the lowest similarity was between R. monosperma and R. raetam Form 6 (49.4 %). Diversity in morphological features of the identified Retama species and forms: There were notable differences among and between the morphological features of the studied Retama taxa such as leaves, flowers, standard, wing, and keel lengths (Diag. 1a-f). The comparable features of the standard (Diag. 3), verify that R. monosperma had the shortest standard length compared to R. raetam and the identified forms. While R. monosperma Form 5 had the widest standard among the rest of the forms, followed by R. raetam Form 4 and R. monosperma were nearly equal in standard width. While R. raetam Form 8 had the narrowest standard. The wings also showed diversity among the studied taxa. Diagram (4) shows that R. monosperma Form 5 had the longest wings, while R. raetam Form 8 had the shortest wing compared with R. raetam, R. monosperma, and other forms. Also, R. monosperma had the widest wing and R. raetam Form 7 had the narrowest wing, while R. raetam, R. raetam Form 2, R. raetam Form 6 and R. raetam Form 8 were equal in wing width. The molecular identity of the identified Retama species and forms: The ISSR marker analysis using 12 primers produced a total of 152 bands, of which 85 bands are polymorphic and 37 are monomorphic (Tab. 3). The band sizes varied between 150 and 2500 bp. The highest matching primers were primers 10 and 14 with 20 bands/each, followed by primers 13 and 19. Further, the lowest number of bands (8 bands/each primer) was obtained with primers 6, 9, and 20. Primer 13 produced bands (13 bands) showing the highest polymorphism (81.3 %). Primer 19 produced more number of unique bands (7 bands) with R. monosperma and Forms (4, 6, and 8). 442 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Retama monosperma was distinguished by five unique bands at 900, 1400, 500, 1250 & 1400 bp with primers 9, 10, 14, and 19; respectively. While, R. raetam could be distinguished from R. monosperma by 2 unique bands at 450 bp (primer 9) and at 360 bp (primer 10). R. raetam Form 2 was distinguished by two unique bands with primers 10 and 20 at 180 and 220 bp, respectively, and an absent band with primer 5 at 200 bp. The two closely related forms, R. raetam Form 7 and R. raetam Form 8, could be distinguished using primer 14 (unique bands at 2500 and 490 bp, respectively). Primer 20 could distinguish between the R. monosperma Form 5 and R. raetam Form 6 (two bands at 400 and 450 bp were absent and one band at 150 bp was unique, respectively). Similarly, primers 5, 6, 10, 13, and 19 produced eight unique bands for R. raetam Form 4 (Tab. 3). Molecular grouping of the identified Retama taxa based on ISSR data: The DCA plot (Diag. 5) based on the ISSR markers data revealed that the identified Retama species and forms could be grouped into two main clusters (I and II). Cluster I included R. monosperma and R. monosperma Form 5; while cluster II included R. raetam and its Forms 2,4,6,7 and 8. This grouping is congruent with the grouping derived from the macro-morphological characters (Diagram 2). Geographic distribution of the identified taxa in Egypt: This revision revealed that Retama raetam was distributed in all the phytogeographic regions in Egypt, mainly in the desert wadis, sandy and gravel plains. Contrastingly, R. monosperma was distributed along the Mediterranean coast from Sallum in the west to Rafah at the Palestinian authority border & the Red Sea coast and close to the salt-affected littoral plains. The identified forms were basically from South Sinai with a westward extension to the northern part of the Eastern Desert (Map 1). DISCUSSION The confusion regarding the taxonomy of the Retama Raf. species is not only based on its generic characters (Lopez et al., 1998) but also closely related to phenology (León-González et al., 2018) and the presence of several varieties (Benmiloud-Mahieddine et al., 2011). This confusion extends to the species description reported previously and the authors noticed misalignments in several points, which will be clarified in this part. Retama raetam (Forssk.) Webb & Branthl: The current revision of the Retama taxa in Egypt revealed the identification of two species and five forms. The first species was R. raetam (Forssk.) Webb, which was characterized by linear-oblong leaves, is consistent with previous reports (Zohary, 1972; Jafri, 1980). However, these reports mentioned larger leaf dimensions (5–20 × 3–8 mm) while the studied specimens showed smaller dimensions (2–8 (- 23) × 0.5–3 mm). This reduction in leaf size might be due to the arid environment of the Egyptian desert (Barakat et al., 2013; Amer et al., 2021). While the specimens we examined exhibited lax–dense inflorescence with 1–7 flowers (Pl. 1e), previous studies only reported 1– 5 (Zohary, 1972; Jafri, 1980; Boulos, 1999). These authors also reported longer flower length (15 mm) which did not exceed 12 mm in the current study (Pl. 1a). The calyx color has also been debatable among authors. For instance, reports have shown purple (Zohary, 1972; Jafri, 443 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. 1980) and reddish calyx (Mouterde, 1966) while this work reported a greenish-purple calyx (Pl. 1a). The standard elliptic, as long as wings (Pl. 1a) with standard length as reported previously (Mouterde, 1966; Zohary, 1972; Täckholm, 1974; Jafri, 1980; Boulos, 1999). While the shape varies, it was obovate-orbicular (Zohary, 1972; Jafri, 1980); rhombic-ovate (Webb, 1853); ovate-oblong (Tutin et al., 1968). The fruit length was 10–15 mm (Pl. 1c), while previous reports showed 20 mm (Tutin et al., 1968; Zohary, 1972; Jafri, 1980; Boulos, 1999). Authors have attributed the smaller fruit size to the aridity in Egypt (Amer et al., 2021). The observed one-seeded, indehiscent pods were consistent with Boulos (1999) and Täckholm (1974). Fruits were elliptical (Pl. 1b, c) consistent with Webb (1853) while relevant shapes were also mentioned as ovoid-oblong (Zohary, 1972; Jafri, 1980;), and ovoid (Mouterde, 1966); oblong- ovate -Webb and Berthelot, 1836) and obovate-ellipsoid (Tutin et al., 1968). The fruit attenuated gradually to an erect peak (Pl.1b, c), allied with Zohary (1972) and Jafri (1980). In contrast, the oblique-peak shape described by Mouterde (1966) did not observe. Fruit pericarp is fleshy, leathery smooth when dry (Pl.1b, c), similar to Jafri (1980) and Zohary (1972). The same authors supported the observed seed color (olive-green to brown; Pl. 1d), as well as Boulos (1999). The seed was black (Webb, 1853; Täckholm, 1974). Chiapella et al. (2009) reported that Retama raetam (Forssk.) Webb subsp. raetam is a Saharo-Arabian taxon, distributed in North Africa, Syria, Israel, Lebanon, and the Arabian Peninsula. Retama monosperma (L.) Boiss.: The second species, was R. monosperma (L.) Boiss., the shrub height was up to 3 m, consistent with Tutin et al. (1968); Polunin (1969); Boulos (1999); it reached 4 m (Quezel and Santa, 1962) or up to 4.5 m (Muñoz Vallés et al., 2013). The inflorescence was lax raceme with 1–7 flower/inflorescence (Pl. 1j), and it is consistent with Tutin et al. (1968) and Muñoz Vallés et al. (2013). While non-parallel data (10–26 flowers/inflorescence) was reported by the later. The flower length was 7–10 mm (Pl. 1f), similar to Tutin et al. (1968) and Muñoz Vallés et al. (2013). While previous studies showed larger flower, it reached 15 mm (Polunin, 1969) and 15–17 mm (Täckholm, 1974; Boulos, 1999). The flowers had purple-colored calyx (Pl. 1f), it appears similar to the reddish calyx reported by Muñoz Vallés et al. (2013). The notable characteristics of Retama monosperma were: (1) The shorter standard compared to wings (Pl. 1f), which has been previously verified (Quezel and Santa, 1962; Täckholm, 1974; Boulos, 1999). The detected standard was orbicular while Tutin et al., (1968) described it as rhombic-ovate. (2) The globoid fruit (Pl. 1g, h) was consistent with previous reports (Polunin, 1969; Täckholm, 1974; Muñoz Vallés, 2013). (3) The fruit was 1–2 seeded, abruptly attenuated to a short-apiculate peak (Pl. 1g) as shown by Muñoz Vallés et al. (2013) while the fruit pericarp was wrinkled (Pl. 1h), consistent with that shown by Barker- Webb and Berthelot (1836) in both R. raetam and R. monosperma fruits. 444 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 The identified species and Forms vs. the taxa reported by Täckholm (1974): Täckholm (1974) mentioned the presence of Lygos raetam (Forssk.) Heywood var. sarcocarpa (Zoh.) Täckh.et Boulos (syn. R. raetam subsp. gussonei (Webb) Greuter) in Egypt. Some of the identified forms (R. raetam Form 2, R. raetam Form 4, R. raetam Form 8) have characters similar to subsp. gussonei, such as diffusely deflexed-branching with silky-silvery indumentum (Zohary, 1972) and the fleshy fruit pericarp (Täckholm, 1974). The dehiscent ellipsoidal-ovoid pod distinguishes subsp. gussonei (Zohary, 1972), this inconsistent trait did not fit into any of the identified forms, where all forms possess indehiscent elliptical-globoid pods. R. monosperma Form 5 that have yellow seeds (Pl. 2a), and wrinkled fruit pericarp was similar in color to subsp. gussonei (Zohary, 1972; Täckholm, 1974). While, the presence of an indehiscent pod and the inflorescence with 15 flowers in R. monosperma Form 5 hampered the fitting it with subsp. gussonei. In addition, Chiapella et al. (2009) restricted the presence of subsp. gussonei to Italy on the Ionian coast of Calabria and Southern Sicily. Täckholm (1974) was expected the presence of at least three nameless varieties in south Sinai. This revision identified four Forms (Form 2, Form 6, Form 7, and Form 8) from South Sinai, its seed length was 7–8 mm and the fruits were green, unlike three nameless varieties given by Täckholm (1974). She reported that the seeds ~ 6 mm and black to dirty-yellow fruit or seeds: ~ 3 mm with white hilum. Therefore, all the varieties expected by Täckholm (1974) were not consistent with the observations retrieved in the current work. The identified Forms vs. Retama monosperma subsp. bovei recorded in Egypt by Boulos (1999): Retama monosperma (L.) Boiss. subsp. bovei (Spach) Maire (syn. R. bovei (Spach) Webb), found in Egypt, were characterized by flowers ranging from 15–17 mm, standard shorter than wings, pod obovoid and seeds reddish-brown or olive-green. The identified R. monosperma Form 5 has a globular pod exceeding 15 mm and a yellow-orange seed, unlike that of subsp. bovei (Boulos, 1999). Additionally, R. monosperma Form 5 lacking the pulvinus-branches base characterized the subsp. bovei (Webb, 1853). Moreover, subsp. bovei is endemic to Algeria and Morocco (Benmiloud-Mahieddine et al., 2011). The morphological and molecular similarity of the identified Retama taxa: The UPGMA dendrogram based on the macro-morphological characters grouped the Retama taxa into two main clusters (I & II). Cluster I included R. monosperma and R. monosperma Form 5, and cluster II included R. raetam and R. raetam Forms (2, 4, 6, 7, and 8; Diagram 2). The similarity value between R. raetam and R. monosperma was 60.4 % while the similarity values between the R. raetam & forms were between 64.8 % to 91.6 % (Tab. 2). The morphological similarities among the Retama taxa (species and forms) might be attributed to similarities in chromosome numbers. This postulation, based on the karyotype analysis of the R. monosperma (4 populations) and R. raetam (17 populations) from Algeria, showed that the chromosome number was the same (2n = 48) (Benmiloud-Mahieddine et al., 2011). 445 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. The DCA plot based on ISSR markers (Diag. 5), showed congruent clustering whereas R. monosperma and its Form 5 and R. raetam as well as its forms were grouped in different clusters. The molecular grouping was consistent with the morphological grouping (Diag. 2) and might be attributed to the difference in the B chromosome, as reported by Chiapella et al. (2009) who revealed that the chromosome number in Retama taxa was 2n = 48 with up to 6 copies of the B chromosomes. For instance, R. monosperma from Portugal (2n = 48) and Spain (2n = 48 + 4B). R. monosperma var. webbii from different regions in Morocco were 2n = 48 + 4B and 48 + 6B; while R. raetam subsp. raetam from Algeria, Libya, Israel, and Algeria was 2n = 24, 48, 48 + 3B, and 48 + 3B, respectively. The subsp. gussonei populations from different regions in Italy had different numbers (2n = 48, 48 + 4B & 48 + 6B). Benmiloud-Mahieddine et al. (2011), clarify the significance of infraspecific variation in chromosome numbers when correlated with the geographic distribution of the studied Retama populations, and the identified forms might be newly adapted populations, consistent with Amer et al. (2021) who reported that a recent increase in aridity induced molecular variations in R. raetam populations and the pressure of grazing increased the use of these shrubs as fodder in these areas (Barakat et al., 2013). Also, Stebbins (1971) claimed that polyploids are better adapted to extreme conditions than their diploid progenitors. Map (1): Distribution pattern of Retama species and Forms in Egypt. 446 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Plate (1): Characteristic morphological features of Retama raetam and R. monosperma; (A–E.) Represent R. raetam, (A) Flower showing standard as long as wings, (B) Elliptic fruit and gradually apiculate peak, (C) Dry one-seeded fruit with leathery smooth pericarp, D: Olive green and brown seeds, and E: Dense inflorescence; (F-J) Represent R. monosperma, (F) Flower showing standard shorter than wings, (G) Globose fruit and abruptly apiculate peak, (H) Dry two seeded fruit with wrinkled pericarp, (I) Olive green and brown seeds, and (J) Lax inflorescence. 447 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Plate (2): Characteristic morphological features of the identified forms; (A) Yellow and orange seed in R. raetam Form 2 and R. monosperma Form 5, (B) Rhombic standard in R. raetam Form 4, (C) Lax inflorescence with 15 flowered in R. monosperma Form 5, (D) Flower length up to 1.5 cm in R. raetam Form 6, (E) Calyx ⅓ flower length in R. raetam Form 7, (F) Orbicular standard in R. raetam Form 7, (G) Calyx ½ flower length in R. raetam Form 8, and (H) Elliptical standard R. raetam Form 8. 448 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Diagram (1): Histograms of different morphological characters between Retama raetam (Rr), Retama monosperma (Rm), and the identified forms (Rr Forms 2, 4,6,7,8 & Rm Form 5). *: a Significant difference. (a) *: P = 0.0183; (b) **: P < 0.0019; (c) **: P = 0.0023 & ***: P < 0.0006; (d) **: P = 0.0053, ***: P = 0.0006; (e) *: P < 0.0335, **: P < 0.0020, ***: P < 0.0001; (f) **: P = 0.0015; ****: P < 0.0001 & ns: non-significant. 449 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Diagram (2): UPGMA dendrogram of the identified R. raetam (Rr), Retama monosperma (Rm), and the identified forms (Rr Forms 2, 4,6,7,8 & Rm Form 5), based on macro-morphological characters using the Jaccard similarity matrix. Diagram (3): Diversity in standard length and width of R. raetam (Rr), Retama monosperma (Rm), and the identified forms (Rr Forms 2, 4,6,7,8 & Rm Form 5), by Rader plot. 450 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Diagram (4): Diversity in wing length and width of R. raetam (Rr), Retama monosperma (Rm), and the identified forms (Rr Forms 2, 4,6,7,8 & Rm Form 5), by Rader plot. Diagram (5): Detrended correspondence analysis (DCA) based on the ISSR markers data of the studied taxa. 451 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Table (1): Localities of the collected fresh populations (used for taxonomic and molecular investigations) and their GPS coordinates arranged from South to North. Locality Location Latitude (N) Longitude (E) 1. Wadi Nasb, South Sinai 28° 27’ 30.6” 34° 4’ 26.292” 2. Wadi Nasb, South Sinai 28° 28’ 4.008” 34° 5’ 58.308” 3. Wadi Nasb, South Sinai 28° 28’ 53.868” 34° 6’ 30.708” 4. Wadi Nasb, South Sinai 28° 29’ 13.632” 34° 6’ 55.98” 5. Wadi Nasb, South Sinai 28° 29’ 42.432” 34° 0’ 5.688” 6. Saint Catherine, South Sinai 28° 33’ 18.144” 33° 56’ 48.3” 7. Saint Catherine, South Sinai 28° 33’ 34.164” 33° 55’ 2.28” 8. Saint Catherine, South Sinai 28° 33’ 34.632” 33° 56’ 51.395” 9. Abo Sela, South Sinai 28° 35’ 21.264” 33° 55’ 44.939” 10. Abo Sela, South Sinai 28° 35’ 36.384” 33° 55’ 30.251” 11. El Sheikh Awad, South Sinai 28° 38’ 23.208” 33° 53’ 23.064” 12. El Sheikh Awad, South Sinai 28° 38’ 55.464” 33° 53’ 22.2” 13. El Sheikh Awad, South Sinai 28° 39’ 4.392” 33° 53’ 54.347” 14. Eltarfa, South Sinai 28° 41’ 40.164” 33° 56’ 35.16” 15. Eltarfa, South Sinai 28° 41’ 49.164” 33° 57’ 39.636” 16. Wadi Hagoul, Cairo-Suez Road 29° 51’ 04.4” 32° 16’ 06.8” 17. Wadi Hagoul, Cairo-Suez Road 29° 53’ 11.3” 32° 14’ 12” 18. Wadi Hagoul, Cairo-Suez Road 29° 58’ 08.3’ 32° 08’ 13.1” 19. Western Mediterranean coast, Omyed 30° 49’ 41.7” 29° 12’ 15.4” 20. Western Mediterranean coast 24 km to Matrouh 30° 49’ 41.7” 29° 12’ 15.5” 22.Western Mediterranean coast, Marakia 30° 55’ 54” 29° 28’ 40.3” 452 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Table (2): Jaccard’s similarity coefficient between the studied Retama species and forms based on macro-morphological characters. Taxa Jaccard’s similarity coefficient R. monosperma R. raetam Form 2 R. raetam R. raetam Form 4 R. monosper ma Form 5 R. raetam Form 6 R. raetam Form 7 R. raetam Form 8 R. monosperma 100 R. raetam Form 2 67.1 100 R. raetam 60.4 75.3 100 R. raetam Form 4 65.2 72.4 91.6 100 R. monosperma Form 5 67.1 55.2 67.4 68.6 100 R. raetam Form 6 49.4 65.9 71.8 76.8 67.9 100 R. raetam Form 7 68.3 74.1 64.8 65.9 65.9 67.1 100 R. raetam Form 8 70.2 71.8 71.4 72.2 64.0 71.1 86.1 100 Table (3): Polymorphism percentages and characteristic bands detected using ISSR marker analysis for the Retama species and forms; test was applied to triple for species and forms. ISSR Primers Number of bands Number of total amplified bands % P o ly m o r p h ism Unique bands between bracts M o n o m o r p h ic P o ly m o r p h ic T o ta l & (U n iq u e ) b a n d s R . m o n o sp e rm a R . ra e ta m F o r m 2 R . ra e ta m R . ra e ta m F o r m 4 R . m o n o sp e rm a F o r m 5 R . ra e ta m F o r m 6 R . ra e ta m F o r m 7 R . ra e ta m F o r m 8 Primer 3 5`-ACACACACACACACACYT-3` 3 9 12 (0) 7 (0) 8 (0) 8 (0) 6 (0) 9 (0) 7 (0) 8 (0) 8 (0) 75 % Primer 5 5`-GTGTGTGTGTGTGTGTYG-3` 1 7 11 (3) 4 (0) 3 (0) 4 (0) 8 (3) 7 (0) 8 (0) 8 (0) 7 (0) 63.64 % Primer 6 5`-CGCGATAGATAGATAGATA-3` 2 4 8 (2) 3 (0) 3 (0) 4 (0) 8 (2) 5 (0) 5 (0) 5 (0) 5 (0) 50 % 453 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. CONCLUSIONS The current revision was conducted to resolve the morphological diversity and addressed the nomenclature debates regarding the genus Retama Raf. in Egypt. We identified two distinct species, R. raetam (Forssk.) Webb and R. monosperma (L.) Boiss. along with five forms of Retama raetam (Forssk.) Webb & Berthel. (Form 2, Form 4, Form 6, Form 7, and Form 8). In addition to Form 5 of Retama monosperma (L.) Boiss. These forms showed significant differences in both morphological characteristics and genetic identity. Each of the identified forms showed inconsistent characteristics with its related species (raetam or monosperma). The results revealed that most of the forms were located in South Sinai, this may highlights the influence of geographic and climatic factors on taxa features and taxonomic identity. Further molecular investigations were highly recommended to resolve the taxonomic identity of the identified forms in genus Retama. CONFLICT OF INTEREST STATEMENT "The authors have no conflicts of interest to declare". Primer 9 5`-GATAGATAGATAGATAGC-3` 2 4 8 (2) 3 (1) 6 (0) 6 (1) 6 (0) 5 (0) 5 (0) 5 (0) 5 (0) 50 % Primer 10 5`-GACAGACAGACAGACAAT-3` 2 12 20 (6) 9 (1) 8 (1) 9 (1) 11 (1) 6 (1) 10 (0) 11 (1) 3 (0) 60 % Primer 11 5`-ACACACACACACACACYA-3` 3 8 13 (2) 9 (0) 9 (0) 6 (0) 4 (0) 7 (0) 5 (2) 10 (0) 10 (0) 61.5 % Primer 13 5`-AGAGAGAGAGAGAGAGYT-3` 2 13 16 (1) 8 (0) 7 (0) 6 (0) 12 (1) 11 (0) 11 (0) 3 (0) 2 (0) 81.3 % Primer 14 5`-CTCCTCCTCCTCCTCTT-3` 8 9 20 (3) 10 (1) 16 (0) 14 (0) 14 (0) 14 (0) 15 (0) 16 (1) 17 (1) 45 % Primer 16 5`-TCTCTCTCTCTCTCTCA-3` 8 1 10 (1) 8 (0) 9 (0) 9 (0) 9 (0) 9 (0) 9 (0) 10 (1) 9 (0) 10 % Primer 18 5`-HVHCACACACACACACAT-3` 3 8 12 (1) 5 (0) 4 (0) 3 (0) 5 (0) 10 (0) 11 (0) 10 (0) 11 (1) 66.7 % Primer 19 5`-HVHTCCTCCTCCTCCTCC-3` 1 6 14 (7) 8 (2) 6 (0) 4 (0) 6 (1) 1 (0) 4 (1) 1 (0) 8 (3) 42.9 % Primer 20 5`-HVHTGTGTGTGTGTGTGT-3` 2 4 8 (2) 5 (0) 5 (1) 4 (0) 4 (0) 2 (0) 5 (1) 6 (0) 6 (0) 50 % Total 37 85 152 (30) 79 (5) 84 (2) 77 (2) 93 (8) 86 (1) 95 (4) 93 (3) 91 (5) 454 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 LITERATURE CITED Abdelhameed, A., Amer, M. W., Hassan, W. and Aboellil, A. 2020. Auto-taxonomy of Brassica tournefortii Gouan. (Brassicaceae) in Egypt. Bangladesh Journal Plant Taxon, 27 (2): 233-250. [CrossRef] Abusaief, H. M. A. A. and Boasoul, S. H. 2021. A taxonomic study of twelve wild forage species of Fabaceae. Heliyon, 7(2): 1-12. [CrossRef] Amer, W. M. 2008. Egyptian flora: status and future prospective. Täeckholmia, 28: 29- 47.[Click here] Amer, W. M., Elshayeb, N. F., Hegazy, A. K., Abbas, M. S., Soliman, A. S. and Abdel Wahab, M. M. 2020. Species diversity and climate an intimate relationship over the last decades in the Mediterranean region: the case study of Sallum Sector, Egypt. Flora Mediterranea, 30: 65-79.[Click here] Amer, W. M., Elshayeb, N. F., Hegazy, A. K., Abbas, M. S. and Soliman, A. S. 2021. Long- term species diversity and climate change: An intimate relationship over the last ten decades: case study in Egypt. In: Leal Filho, W. et al. (Ed.). Handbook of Climate Change Management, p. 1-24. Springer Nature, Switzerland. [CrossRef] Awen, B. Z. S., Unnithan, C. R., Ravi, S., Kermagy, A., Sasikumar, J. M., Khrbash, A. S. and Ekreem, W. L. 2011. Essential oils of Retama raetam from Libya: chemical composition and antimicrobial activity. Natural Product Research, 25(9): 927-933. [CrossRef] Barakat, N. A. M., Laudadio, V., Cazzato, E. and Tufarelli, V. 2013. Potential contribution of Retama raetam (Forssk.) Webb & Berthel as a forage Shrub in Sinai, Egypt. Arid Land Research and Management, 27(3): 257-271. [ResearchGate] Barker-Webb, M. M. P. and Berthelot, S. 1836. Histoire naturelle des Iles Canaries. Saint- Andre-Des-Art, Paris, 56 pp. Benmiloud-Mahieddine, R., Abirached-Darmency, M., Brown, S. C., Kaid-Harche, M. and Siljak-Yakovlev, S. 2011. Genome size and cytogenetic characterization of three Algerian Retama species. Tree Genetics and Genomes, 7 (issue number): 987-998. [CrossRef] Boulos, L. 1999. Flora of Egypt (Azollaceae-Oxalidaceae). Al Hadara Publishing, Cairo, Egypt, 419 pp. Boulos, L. 2009. Flora of Egypt checklist revised annotated edition. Al-Hadara Publishing, Cairo, Egypt, 410 pp. http://dx.doi.org/10.3329/bjpt.v27i2.50664 https://doi.org/10.1016%2Fj.heliyon.2021.e06077 https://www.academia.edu/35431756/Flora_of_Egypt_status_and_future_prospectives.pdf https://www.herbmedit.org/flora/FL30_065-080.pdf https://doi.org/10.1007/978-3-030-22759-3_103-2 https://doi.org/10.1080/14786419.2010.503612 https://www.researchgate.net/profile/Eugenio-Cazzato/publication/235760000_Potential_Contribution_of_Retama_raetam_Forssk_Webb_Berthel_as_a_Forage_Shrub_in_Sinai_Egypt/links/561cf10008aea803672665ff/Potential-Contribution-of-Retama-raetam-Forssk-Webb-Berthel-as-a-Forage-Shrub-in-Sinai-Egypt.pdf https://doi.org/10.1007/s11295-011-0389-z 455 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Bruneau, A., Doyle, J. J., Herendeen, P., Hughes, C., Kenicer, G., Lewis, G., Mackinder, B., Pennington, R. T., Sanderson, M. J. and Wojciechowski, M. F. 2013. Legume phylogeny and classification in the 21 st century: progress, prospects and lessons for other species-rich clades. Taxon, 62 (2): 217-248. [CrossRef] Chiapella, L. F., Velari, T. C., Kosovel, V. and Pellizzari, L. 2009. Karyological notes on some genera of Genisteae from the Mediterranean region. Bocconea, 23: 187-202. Chouaibi, M., Rezig, L., Lakoud, A., Boussaid, A., Hamdi, S., Hassouna, M. and Ferrari, G. 2019. Exploring potential new galactomannan source of Retama raetam seeds for food, cosmetic and pharmaceuticals: Characterization and physical, emulsifying and antidiabetic properties. International Journal of Biological Macromolocules, 124: 1167-1176. [CrossRef] Djeddi, S., Karioti, A., Yannakopoulou, E., Papadopoulos, K., Chatter, R. and Skaltsa, H. 2013. Analgesic and antioxidant activities of Algerian Retama raetam (Forssk.) Webb & Berthel extracts. Records of Natural Prodroducts, 73: 169-176.[ResearchGate] Elshayeb, N. F. M. 2020. Vegetation ecology in the western Mediterranean stripe under the climatic changes during the past one hundred years. Ph.D. thesis, Cairo University, Faculty of African Postgraduates studies, University of Cairo (unpublished thesis), 174 PP. GBIF Secretariat. 2022. GBIF Backbone Taxonomy. Checklist dataset. [Click here] Helm, A., Oja, T., Saar, L., Takkis, K., Talve, T. and Pärtel, M. 2009. Human influence lowers plant genetic diversity in communities with extinction debt. Journal of Ecology, 97(6): 1329-1336. [CrossRef] Jafri, S. M. H. 1980. Fabaceae. In: Jafri, S.M.H. and El-Gadi A. (Ed.). Flora of Libya, p. 31- 33. Tripoli University, Libya. Karimi, H. R., Kafkas, S., Zamani, Z., Ebadi, A. and Fatahi, M. R. 2009. Genetic relationships among Pistacia species using AFLP markers. Plant Systematics and Evolution, 279: 21-28.[Click here] Käss, E. and Wink, M. 1997. Phylogenetic relationships in the Papilionoideae (Family Leguminosae) based on nucleotide sequences of cpDNA (rbcL) and ncDNA (ITS 1 and 2). Molecular Phylogenetics and Evolution, 8(1): 65-88. [CrossRef] León-González, A. J., Navarro, I., Acero, N., Muñoz Mingarro, D. and Martín-Cordero, C. 2018. Genus Retama: A review on traditional uses, phytochemistry, and pharmacological activities. Phytochemistry Reviews, 17 (4): 701-731. [CrossRef] https://doi.org/10.12705/622.8 http://dx.doi.org/10.1016/j.ijbiomac.2018.12.007 https://www.researchgate.net/publication/237078139_Analgesic_and_Antioxidant_Activities_of_Algerian_Retama_raetam_Forssk_Webb_Berthel_Extracts https://www.gbif.org/dataset/search?type=CHECKLIST https://doi.org/10.1111/j.1365-2745.2009.01572.x https://www.academia.edu/51912675/Genetic_Relationships_Among_Pistacia_Species_Using_AFLP_Markers#:~:text=DOI%2010.1007/s00606%2D008%2D0117%2D9 https://doi.org/10.1006/mpev.1997.0410 https://doi.org/10.1007/s11101-018-9555-3 456 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Lopez, J., Devesa, J. A., Ruiz, T. and Ortega-Olivencia, A. 1998. Seedling morphology in Genisteae (Fabaceae) from south-west Spain. Botanical Journal of Linnean Society, 128 (3): 229-250. [Click here] Mabberley, D. J. 1997. The Plant-Book. Cambridge Univ. Press, Cambridge, 680 pp. Maghrani, M., Zeggwagh, N. A., Haloui, M. and Eddouks, M. 2005. Acute diuretic effect of aqueous extract of Retama raetam in normal rats. Journal of Ethnopharmacology, 99 (1): 31-35. [CrossRef] Minaeifar, A. A., Sheidai, M., Attar, F., Noormohammadi, Z. and Ghasemzadeh-Baraki, S. 2016. Biosystematic study in the genus Cousinia Cass. (Asteraceae), section Cousinia. Biochemical Systematic and Ecology, 69: 252-260. [CrossRef] Morsy, A. A., Khatab, H. K., El Sherbiny, E. A. and Eldemirdash, J. E. 2015. Floristic diversity and vegetation analysis of Wadi Sudr, South-West Sinai Peninsula. Taeckholmia, 35: 99-119. [CrossRef] Mosaferi, S., Sheidai, M., Keshavarzi, M. and Noormohammadi, Z. 2015. Genetic diversity and morphological variability in Polygonum aviculare s.I. (Polygonaceae) of Iran. Phytotaxa, 233 (2): 166-178. [CrossRef] Mouterde, P. 1966. Nouvelle Flore du Liban et de la Syrie. Dar EL-Machreq Editeurs, Liban. p. 227-228. Muñoz Vallés, S., Gallego Fernández, J. B. and Cambrollé, J. 2013. The biological flora of coastal dunes and wetlands: Retama monosperma (L.) Boiss. Journal of Coastal Research, 29(5): 1101-1110. [CrossRef] Muraseva, D. S. and Guseva, A. A. 2021. ISSR primer screening for analysis of genetic diversity among Scutellaria tuvensis (Lamiaceae) populations. In: Bio Web of Conferences. (Vol. 38, P. 00082). EDP Sciences. [CrossRef] Pardo, C., Cubas, P. and Tahiri, H. 2004. Molecular phylogeny and systematics of Genista (Leguminosae) and related genera based on nucleotide sequences of nrDNA (ITS region) and cpDNA (trnL-trnF intergenic spacer). Plant Systematics and Evolution, 244: 93-119. [CrossRef] Polunin, O. 1969. Flowers of Europe. Oxford University press, New York, P. 186. Quezel, P. and Santa, S. 1962. Nouvelle flore de l'Algérie et des régions désertiques méridionales. Editions du centre national de la recherche scientifique, France, 475 pp. https://www.zobodat.at/pdf/PHY_39_1_0107-0129.pdf https://doi.org/10.1016/j.jep.2005.01.045 https://doi.org/10.1016/j.bse.2016.10.008 https://doi.org/10.21608/taec.2015.12223 https://doi.org/10.11646/phytotaxa.233.2.4 https://doi.org/10.2112/JCOASTRES-D-12-00013.1 https://doi.org/10.1051/bioconf/20213800082 https://doi.org/10.1007/s00606-003-0091-1 457 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Youssef et al. Rohlf, F. J. 2009. NTSYS–PC: numerical taxonomy system. version 2.21. Setauket, NY, USA. Stebbins, G. L. 1971. Chromosomal evolution in higher plants. Arnold, London, 216 pp. Täckholm, V. 1974. Student’s flora of Egypt. Cairo University Press, Cairo, 666 pp. Thiers, B. 2019. The World’s Herbaria 2018: A summary report based on data from index herbariorum. New York Botanical Garden’s Virtual Herbarium. [Click here] Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. and Webb, D. A. 1968. Flora Europaea (Rosaceae-Umbelliferae). Cambridge University Press, Cambridge, 101pp. Webb, P. B. 1853. Plantarum Rabiorum. Visctor Masson, Bibliopola, Parisis, p. 22-25. Wickens, G. E. 1998. Ecophysiology of economic plants in arid and semi-arid lands. Springer, United Kingdom, 232 pp. Yahara, T., Javadi, F., Onoda, Y., de Queiroz, L. P., Faith, D. P., Prado, D. E., Akasaka, M., Kadoya, T., Ishihama, F., Davies, S. and Slik, J. F. 2013. Global legume diversity assessment: concepts, key indicators, and strategies. Taxon, 62(2): 249-266. [CrossRef] Zietkiewicz, E., Rafalski, A. and Labuda, D. 1994. Genome fingerprinting by simple sequence repeat (SSR)-anchored polymerase chain reaction amplification. Genomics, 20 (2): 176-183. [CrossRef] Zohary, M. 1972. Flora Palaestina (Platanaceae-Umbelliferae). The Israel academy of sciences and humanities, Jerusalem, p. 47-48. https://sweetgum.nybg.org/science/ih/ https://doi.org/10.12705/622.12 https://doi.org/10.1006/geno.1994.1151 458 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Genus Retama Raf., 1838 Bull. Iraq nat. Hist. Mus. (2023) 17 (3): 435-458. في مصر مدعوما بالبصمة الجزيئية Retama Raf., 1838مراجعة تصنيفية لجنس ريهام يوسف، وفاء محروس عامر و عزة حامد ، 12613قسم علوم النبات و االحياء املجهرية، كلية العلوم، جامعة القاهرة، الجيزة مصر. 20/6/2023 ، تأريخ النشر:18/3/2023القبول: ، تأريخ 4/2/2023تأريخ االستالم: الخالصة العائلة من Retama Raf., 1838تحت جنس وفاتفي مصر، الهوية التصنيفية لألصن تحديد هويته الخارجي يجعلال يزال غير واضح وتشابهه الشكل ((Fabaceaeالبقولية بمصر وذلك فيما يتعلق هذا الجنسالدراسة الحالية إلى مراجعة تهدف. أمرا صعبا هذا ىية ووراثية وجغرافية. وللوصول الملوجودة وما يميزها من صفات شكلباألنواع ا لقد أجرينا املراجعة . واملعشبية ريةالط Retama جنس عينات ال تاستخدم ،الهدف سمة شكلية ودرسنا الهوية الجزيئية باستخدام عالمات تكرار 94املظهرية باستخدام Retama ديد نوعين متميزين هماالتسلسل البسيط و كشفت هذه املراجعة عن تح raetam (Forsskk.) Webb و Retama monosperma (L.) Boiss إلى جانب خمسة ، النموذج 6، النموذج 4، النموذج 2النموذج ) Retama raetam اصنوفة أشكال تحت وقد أظهرت . R. monospermaمن تحت 5باإلضافة إلى النموذج (. 8، والنموذج 7 الهوية املظهرية والجزيئية لهذه األشكال اختالفات كبيرة في نفس الوقت كونت وتسلط . تم توضيح التوزيع الجغرافي لألصناف في مصر. مجموعات إحصائية متطابقة هو من األنواع املهمة، معرضة للتأثر سلبا Retamaهذه الدراسة الضوء أيًضا على أن . طلب برامج صون عاجلة بالتغير املناخي وتت