Phytotaxa 408 (1): 001–029 ISSN 1179-3155 (print edition) https://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2019 Magnolia Press Article ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.408.1.1

Taxonomic revision of the (, ) in , with the first record of Ononis viscosa subsp. breviflora

ABDEL AZIZ A. FAYED1, AZZA H. EL-HADIDY2, AHMED M. FARIED1 & ASMAA O. OLWEY1* 1Botany and Microbiology Department, Faculty of Science, Assiut University, Assiut, Egypt. 2Botany and Microbiology Department, Faculty of Science, University, , Egypt. *E-mail: [email protected]

Abstract

The genus Ononis comprises 86 species worldwide distributed in temperate regions. It is represented in Egypt by ten species. During recent excursions to the mountains of southern Sinai Peninsula, a newly recorded species, namely O. viscosa subsp. breviflora, was collected. This new record was previously confused with O. sicula. These two species can be differentiated by corolla length and shape. This study provides a full taxonomic revision of the genus for the flora of Egypt, updated no- menclature for all taxa, amended descriptions, detailed hand drawings, and a diagnostic key to the species. The most valuable diagnositic characters in the of the genus in Egypt are discussed. After critical examination and literature study, we found that two names (O. serrata and O. vaginalis) needed to be lectotypified.

Key Words: Ononis, Fabaceae, Flora of Egypt, Lectotypification, Trifolieae

Introduction

Fabaceae Lindley (1836: 544) or Leguminosae Jussieu (1789: 345) is the third largest family of flowering , with approximately 19,500 species classified into approximately 770 genera (Polhill & Raven 1981, Christenhusz & Byng 2016, LPWG 2017). Recent molecular and morphological evidence supports Fabaceae as a single monophyletic family (Wojciechowski et al. 2000, Lock 2005, Bello et al. 2009, 2012, Lewis et al. 2013, LPWG 2017, Soltis et al. 2018). The genus Ononis Linnaeus (1753: 716, 1754: 321) is nested within the tribe Trifolieae Fries (1835: 100) and comprises 86 species worldwide that are distributed in temperate regions (Turini et al. 2010). Linnaeus (1753, 1754) recognized only 17 species. De Candolle (1825) classified the genus into two sections: Euononis De Candolle (1825: 158) and Lotononis De Candolle (1825: 166), based on leaf stipules (whether or not stipules adhere to the petiole). Later, Willkomm (1877) subdivided Ononis into four sections and six subsections based on habit, leaf composition, presence or absence of spines, pedicel elongation (to produce arista) and flower colour. The most comprehensive taxonomic monograph of the genus was presented by Širjaev (1932), who accepted 68 species distributed in two sections: sect. Ononis, which is characterized by reduced peduncles and few-seeded pods (2– 4); and sect. Natrix Grisebach (1843: 12), which is characterized by distinct peduncles and multi-seeded pods. More recently, Turini et al. (2010) studied the phylogeny of the genus Ononis using the plastid region trnL-F and nuclear ITS sequences and found that Ononis is a monophyletic group. However, the morphological and molecular evidence contradicted the traditional infrageneric classification; five new major lineages within the genus were suggested that are morphologically supported by peduncle length, flower colour and habit. In Egypt, Ononis has been treated only in broad floristic studies (e.g., Täckholm 1956, 1974, El Hadidy 1989, El Hadidi & Fayed 1995, Boulos 1999), which lack proper synonymy lists, full type citations, and detailed illustrations. In addition, Ononis has also been treated in studies concerning the tribe Trifolieae (e.g., Ayyad & Moore 1994, Taia 2004a, b, Lashin 2006).

Accepted by James Boatwright: 19 Mar. 2019; published: 27 Jun. 2019 1 The present study Boulos (1999: 261-262) El Hadidi & Fayed (1994/1995: 59) Täckholm (1974: 229-230 ) Täckholm (1956: 278-279) Muschler (1912: 468-475) taxa recordedin Egypt (+ = present, - absent, x a synonym was recorded). Ascherson &Schweinfurth (1889:61) Ononis ---- -++ - -- x - +-- x + +-- + + + + + - + + -- + - + + + - + + + + - + + + + + + + + + + + + + + + -- +-- x +-+ + --+ + + - x - + + + - + + - + + + + + + + Forsskål (1775:130) -+x + + - + + + + + + + + -+ + + + + + + + + -+-- x + + - + - + + - - - + -+ + + + + + +

.

Vahl L. Tenore . minor L. var Gussone (Savi) Historical review for (Boiss.) var Boissier major Linnaeus . breviflora L. subsp. L . mollis Savi Ononis serrata O. natrix Ononis Ononis cherleri . Ononis . Ononis calycina Forssk.var. Lange Syn. Ononis serrata Forsskål Ononis mitissima Linnaeus Ononis variegata Linnaeus Linnaeus subsp. natrix O. natrix Syn. Ononis diffusa stenophylla Širjaev stenophylla reclinata Moris Ononis vaginalis Syn mollis Syn. Ononis pubescens Béuinot Syn. Forsskål Ononis reclinata Syn subsp Lamarck Ononis sicula (DC.) Nyman Ononis viscosa L.subsp. TABLE 1. TABLE

2 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. The number of species of Ononis recorded in Egypt has increased over time (Table 1); Forsskål (1775) recognized four species in Egypt, two of which were validated by subsequent authors such as Greuter & Raus (1989): Ononis cherleri Forsskål (1775: 130) [= O.vaginalis Vahl (1790: 53)] and O. serrata Forsskål (1775: 130). Then, Raffeneau- Delile (1812) recorded four species in Egypt (adding two more to Forsskål’s list), namely, O. pubescens Linnaeus (1771: 267–268) and O. cherleri Linnaeus (1763: 1007). Ascherson & Schweinfurth (1889) added O. mitissima Linnaeus (1763: 1007) and O. sicula Gussone (1821: 78). Then, Muschler (1912) added one more taxon: O. natrix Linnaeus (1753: 717) subsp. stenophylla (Boissier) Širjaev (1932: 470). Täckholm (1974) listed eight species of Ononis in Egypt, while Boulos (1999, 2009) added one more species to the list for a total of nine species. Despite the fact that its natural geographic distribution is along the Mediterranean coast, Ononis viscosa Linnaeus subsp. breviflora (DC.) Nyman (1878: 161) has never been recorded in the Flora of Egypt, although Širjaev (1932: 530) suspected the presence of this taxon in Egypt. Recent excursions to the mountains of the southern Sinai Peninsula uncovered many floristic treasures (Fayed et al. 2004), one of which was the first record of O. viscosa subsp. breviflora. The aims of this study were as follows: first, to provide an updated taxonomic revision of all taxa belonging to the genus Ononis in Egypt; second, to critically evaluate previously used taxonomic characters and discuss their taxonomic validity; and third, to provide an improved diagnostic key for the taxa of Ononis in Egypt.

Materials and methods

This study was based on herbarium specimens kept in the following Egyptian herbaria: ASTU, CAI, CAIM and MAZHAR, in addition to fresh material collected from different localities representing the different habitats of Ononis in Egypt. Acronyms of herbaria are according to Thiers (continuously updated). High-quality digital photos of Egyptian material as well as the authentic type specimens kept in different herbaria worldwide were examined by the authors. The specimens examined for each species are listed according to the phytogeographical regions of Egypt, from north to south and from west to east. Within each phytogeographical region, specimens are arranged according to the date of collection. Abbreviations of Egyptian phytogeographical areas were adopted from Boulos (1999): Sinai (S), Mediterranean coastal belt (M), and Nile valley (N). Conservation assessments for each species were checked through the different Red List categories (IUCN 2018). The line drawings were made (by Olwey A.) with the help of a camera lucida attached to a Wild M8 Zewz stereomicroscope, and photographs were taken using an Olympus SC100 digital camera.

Results and discussion

The diagnositic characters for the genus Ononis in Egypt:—

Habit:—Based on habit, the genus Ononis was subdivided into sections by Wilkomm (1877), Ascherson & Graebner (1907) and Širjaev (1932). Regarding the Egyptian species of Ononis, only two species are subshrubs: O. natrix and O. vaginalis; all remaining taxa are annual herbs. Leaf composition:—All studied taxa bear two types of leaves: simple and trifoliate leaves; only Ononis variegata Linnaeus (1753: 717) is unique in having simple leaves as the dominant type, and it rarely bears trifoliate leaves. Leaf stipules:—The shape of the stipules proved to be a critical character in the discrimination of different species of Ononis in Egypt. Ononis mitissima can easily be distinguished by its large membranous stipules, while the remaining taxa have green stipules. Stipule outline is a taxonomically informative character for distinguishing Egyptian Ononis. The stipule outline is either ovate (in O. diffusa Tenore (1811: XLI), and in O. serrata), oblong (in O. mitissima and O. natrix), obovate-rhombic (in O. variegata), linear to linear-lanceolate (in O. sicula) or deltoid (in O. viscosa subsp. breviflora). The vestiture of the stipules was neglected by many previous authors; however, it appears to have merit as a diagnostic character. Two species (O. variegata and O. mitissima) can be diagnosed as having glabrous stipules instead of the glandular stipules found in all the remaining taxa under investigation. Based on the length of the stipules, Egyptian Ononis can be classified into two groups: taxa with small stipules ranging from 2.5–5.0 mm, including O. diffusa, O. serrata, O. variegata, O. reclinata subsp. mollis (Savi) Béguinot (1912: 134) and O. natrix subsp. stenophylla, and taxa with medium to large stipules of up to 18 (20) mm, including O. mitissima, O. natrix subsp. natrix, O. pubescens, O. sicula and O. viscosa subsp. breviflora.

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 3 Peduncle:—The length of the floral pedicel and whether the peduncle elongates to produce arista are paramount taxonomic characters that have regularly been used as keystone characteristics for infrageneric classification of the genus (Willkomm 1877, Ascherson & Graebner 1907, Battandier & Trabut 1890, Širjaev 1932, Turini et al. 2010). In the taxa studied, four categories can be segregated: the first category includes taxa with sessile flowers or very short pedicels (pedicel length ranges from 0.0–1.5 mm) and ex-aristate peduncles, including O. diffusa, O. serrata and O. mitissima; the second category includes one taxon with medium-length pedicels (ranging from 1.5–3.5 mm) and ex-aristate peduncles, namely, O. variegata; the third category includes taxa with well-developed pedicels (ranging from 5.0–150.0 mm) and ex-aristate peduncles, including O. pubescens and O. reclinata subsp. mollis; and finally, the last category includes taxa with well-developed pedicels (length ranging from 5.0–30.0 mm) and aristate peduncles, including O. natrix, O. vaginalis, O. sicula and O. viscosa subsp. breviflora. Length of arista:—This character was successfully employed for the infraspecific classification of O. viscosa by Huber-Morath (1970). Here, it can be used to differentiate O. viscosa subsp. breviflora and O. sicula, since the latter has an aristum of 2.0–7.0 mm, while the former has an aristum of 4.5–20.0 mm. The character can also be useful for the discrimination of the two subspecies of O. natrix, which are morphologically very similar: O. natrix subsp. natrix has long arista that ranges from 6.0 to 10.0 mm, while O. natrix subsp. stenophylla has shorter arista ranging from 1.0 to 3.0 mm. Calyx:—Širjaev (1932) discussed the taxonomic significance of calyx characters for the classification of Ononis. Using the relative length of the calyx tube and teeth (lobes), four categories can be distinguished within the studied taxa: the first category includes taxa with calyx teeth shorter than the calyx tube (i.e., O. mitissima only); the second category includes taxa with calyx teeth sub-equalling or slightly longer than the calyx tube (i.e., O. serrata and O. variegata); the third category includes taxa with calyx teeth longer than the calyx tube by 1.5-fold to 2.5-fold (i.e., O. diffusa, O. natrix and O. vaginalis), and finally, the last category includes taxa with calyx teeth longer than the calyx tube by 3-fold to 5-fold or more (i.e., O. pubescens, O. reclinata subsp. mollis, O. sicula and O. viscosa subsp. breviflora). In terms of sepal symmetry, O. diffusa can be distinguished from all the remaining taxa in having asymmetric sepals. With regard to sepal hairiness, only O. variegata has entirely glabrous sepals, which make it unique from the remaining taxa. Corolla:—The length of the corolla in relation to the length of the calyx is an important character in the taxonomy of Ononis. Egyptian species can be classified into three categories: first, taxa with a corolla obviously shorter than the calyx: O. reclinata subsp. mollis and O. sicula; second, taxa with a corolla sub-equalling the calyx or slightly longer or slightly shorter than the calyx [i.e. O. pubescens, O. mitissima and O. viscosa subsp. breviflora]; and third, taxa with a corolla longer than the calyx [i.e., O. diffusa, O. serrata, O. variegata, O. natrix and O. vaginalis]. The surface of the standard is a crucial character for diagnosis: O. diffusa and O. serrata are distinguished from all the remaining studied taxa by glandular hairs covering the standard, O. variegata has a standard covered with eglandular trichomes, and all the rest have a glabrous corolla. Wing auricles are minute projections at the base of the wing lamina, and the length of the auricle is often neglected. Three patterns of auricle size can be defined in the studied taxa: 1- very minute and hardly distinct, ranging from 0.1 mm to 0.4 mm (in O. serrata, O. mitissima, O. pubescens and O. sicula); 2- small auricles ranging from 0.5 mm to 0.75 mm (in O. variegata, O. natrix subsp. stenophylla, O. reclinata subsp. mollis and O. viscosa subsp. breviflora); and 3- large distinct auricles sized 1.0–1.5 mm (in O. vaginalis, O. natrix subsp. natrix and O. diffusa). Conjunctive teeth of wings were uniquely found in four taxa (O. diffusa, O. serrata, O. mitissima and O. pubescens) and were absent in all the remaining taxa under investigation. Legume:—The shape of the legume is a defining feature for the sectional classification of Ononis: taxa belonging to sect. Ononis have non-pendent, globose to sub-globose, ovoid to elliptic pods with a limited number of seeds (2–6), while members of sect. Natrix have pendent linear pods with many seeds (up to 25), with the exception of O. pubescens, a member of sect. Natrix subsect. Viscosae Širjaev (1932: 518) with globose pods. All Egyptian Ononis have rostrate or apiculate legumes; only O. reclinata subsp. mollis pod has a rounded apex. Fruiting calices are accrescent in O. diffusa, O. serrata and O. pubescens.

Taxonomy

Ononis Linnaeus (1753: 716, 1754: 321) ≡ Anonis Miller (1754). Type:— nom. cons. subsp. spinosa Linnaeus (1753: 716) ≡ subsp. spinosa Greuter (1986: 113).ITALY.1600– 1625, Burser J. XXI: 79, BOT:V-175559, conserved type designated by Jarvis et al. in Brummitt (1989), (UPS-image!).

4 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. =Ononis campestris Koch & Ziz (1814: 22). Type:—FRANCE. Alpes, lieux secs des pâturages aux bords du Rhône, près de Lyon, 25 July 1845, Jordan C. 364, COI00053421, (COI-image!).

Description:—Perennial or annual herbs, shrubs or rarely trees, usually glandular or villous, rarely glabrous, sometimes spiny. Branches erect, ascending, or prostrate. Leaves pinnately 1–3(–5)-foliolate, lateral leaflets often reduced or absent; margins denticulate to serrate, very rarely entire; petiolate, rarely sessile; stipules usually adnate to petiole, sometimes sheathing, cauline stipules basally connate, bifid; leaflets toothed, lateral veins extending into teeth. Flowers 1–3 in short axillary racemes; peduncle well-developed to brief and reduced, often bristle-like; floral leaves sometimes reduced to bracts when flowers crowded into a spike, pedicels often elongated and pendent back upon itself after flowering. Calyx persistent, campanulate or tubular, sometimes ±bilabiate after anthesis; sepals usually almost equal, rarely ±unequal; 3–5-nerved. Corolla pink, yellow, purple, white to versicolor; petals free from staminal tube; standard suborbicularto obovate, glabrous to pilose; wings oblique ovate to obovate tooblong-obovate, auricle ±small, conjunctive tooth sometimes present; keel ±beaked. Stamens monadelphous; filaments dilated at apex; anthers dimorphic, alternately basifixed and dorsifixed. Ovary shortly stipitate; ovules 2 to many. Style bent, glabrous to hairy on the base. Stigma terminal, small, oblique to sub-capitate. Legume linear-oblong or ovate. Seed sub-orbicular to reniform, smooth tuberculate or punctate.

Key to Ononis species in Egypt:—

1. Subshrubs or tall herbs; flowers yellow; floral pedicels up to 30.0 mm; with well-developed aristate peduncles ...... 2 - Short or medium-sized herbs; flowers white, pink or yellow; floral pedicels shorter (from 0.0 to 15.0 mm); with ex-aristate peduncles ...... 5 2. Perennial subshrubs; corolla much longer than calyx ...... 3 - Annual herbs; corolla shorter than or as long as calyx ...... 4 3. Flowers solitary; leaves mostly simple, sessile; stipules sheathing ...... O. vaginalis - Flowers arranged in racemes; leaves mostly trifoliate, petiolate; petiole 4.0–20.0 mm; stipules not sheathing ...... O. natrix 4. Leaflets narrowly elliptic (1.0–1.5 mm wide); corolla short (5.5–6.0 mm long), obviously shorter than calyx; standard elliptic with obtuse apex ...... O. sicula - Leaflets broadly elliptic (3.5–10.0 mm wide); corolla long (6.2–8.0 mm long), as long as calyx or slightly longer; standard obovate with retuse apex ...... O. viscosa subsp. breviflora 5. Stipules glabrous ...... 6 - Stipules covered with glandular hairs ...... 7 6. Leaves trifoliate; stipules large, up to 9.0 mm, membranous, conspicuous, not sheathing; calyx tube longer than teeth; style glabrous ...... O. mitissima - Leaves simple; stipules smaller, not exceeding 4.0 mm, green, inconspicuous, sheathing; calyx tube equal to teeth; style hairy at the base ...... O. variegata 7. Pod linear, many-seeded (6–10), distinctly pendent, not beaked ...... O. reclinata - Pod ovoid, few-seeded (2–5), slightly or not pendent, beaked ...... 8 8. Petiole (8.0–14.0 mm long), shorter than stipules (9.0–18.0 mm long); standard glabrous ...... O. pubescens - Petiole (2.0–7.5 mm), longer than stipules (1.5–5.0 mm long); standard glandular ...... 9 9. Leaflets obovate, with 3–6 pairs of teeth; sepals unequal, ovate, fused to 1/3 of their length or more; standard orbicular with mucronate apex, 10.5–11.5 mm; pod 3–4 mm broad ...... O. diffusa - Sepals equal, linear to linear-lanceolate, fused to 1/4 or less of their length, standard elliptic with acute apex, 5.2–7.8 mm; leaflets obovate, with 3–6 pairs of teeth ...... O. serrata

I. Section: Ononis, Syn. Bugräna Grisebach (1843: 12)

1. Ononis diffusa Tenore (1811: XLI). Type:—EUROPE. Italy, Tenore M. s.n., BM000751135, holotype, (BM-image!). Heterotypic synonyms: = Ononis dehnhardtii Tenore (1825: 11).≡ Ononis serrata Forssk. var. major Lange (1865: 156). Type:—SPAIN. Ex Tenerife: Bufadero, in saxosis camporum, 24 March 1855, Bourgeau N. 1317, COI00053519, (COI-image!) = Ononis serrata Forssk. subsp. diffusa (Ten.) Rouy (1897: 268). Type:—FRANCE. De Bastia à Biguglia, Mabille, Campbell, Ostriconi, Bonifacio, Alèria, sables maritimes de la Corse.

Description:—Annual herb, up to 40–50 cm high, much branched; branches decumbent to ascending, pubescent with glandular and eglandular hairs. Stipules green, ovate, 4.0–5.0 × 1.5 mm, acute apex, denticulate margins, with five protruding nerves, covered with glandular hairs, shorter than the petiole. Leaves mostly trifoliate, only flowering leaves simple, 17–37 mm long, petiolate; petiole 4.0–8.0 mm; leaflet suborbicular, obovate to oblong, 12.0–25.0 × 3.0–7.0

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 5 mm, acute apex, cuneate to attenuate base, dentate margins with 7–12(–20) pair of small teeth; with 1.0–4.0 mm long petiolule. Inflorescence terminal, dense raceme, 40–50 mm long, 10–15 flowered, pedunculate; peduncle 3.0–5.0 mm long, shorter than subtending leaf, not aristate. Flowers sessile or with very short pedicel, pedicel about 1.0–2.0 mm long. Calyx tubular, shorter than corolla, 10.0–11.5 × 1.3–2.0 mm, covered with glandular hairs; calyx lobes 6.5–7.0 mm, 5–nerved, ovate, with acute apex, asymmetrical; calyx tube 3.5–4.5 mm, calyx lobes longer than tube. Corolla white with purple veins on the standard; standard orbicular, covered with glandular hairs, 10.5–11.5 × 7.0–8.0 mm, mucronate apex, sinuate margins, with distinct subulate claw ca. 3.0 mm long; wings lamina 7.0–7.5 × 2.0–3.0 mm, claw ca. 2.0 mm long, auricle distinct ca. 1.0 mm long, conjunctive tooth flap-like; keel lamina 7.5 × 2.0 mm, claw ca. 3.5 mm long, wings shorter than the sub-equaling standard and keel. Stamens 7.2 mm. Ovary 2.8 × 1.2 mm, covered with glandular hairs; style straight, not deflexed, 6.5 mm long, glabrous. Pod sub-globose to ovoid, 8.0 × 4.0 mm, covered on the upper half and distal margin with glandular hairs, apex rostrate; beak 0.5–0.75 mm long; fruiting calyx accrescent; seeds few, 3–4 seeds, suborbicular, 1.7–1.75 × 1.49–1.53 mm, densely papillose, with black testa. Conservation status:—Not threatened. Phenology:—Flowering season March–June. Habitat:—Alluvial soil. Chromosome number:—2n = 30 (Palazueles et al. 1976, Dalgaard 1991). Distribution in EGYPT:—Native in two major localities (Fig. 1); N: Cairo and S: Rafah (Täckholm 1956, Boulos 1999).

FIGURE 1. Geographical distribution of the genus Ononis in Egypt, with a closer scope on Southern Sinai where Ononis viscosa subsp. breviflora is labelled with an asterisk.

Global distribution:—This species is centred along the Mediterranean coast, from which its distribution extends northwards to central and southern Europe (i.e., Corsica, Crete, Cyprus, Greece, Italy, Portugal, Sardinia, Sicily and Spain) and southwards to northwestern Asia in Palestine and the northern coasts of Africa in Libya, Algeria, Morocco, and Tunisia (Greuter & Raus 1989). Taxonomic notes:—Ononis diffusa is close ally to O. serrata. Some authors treat O. diffusa as a subspecific (Rouy 1897) or as a varietal taxon to O. serrata (Lange 1865).These species share some characteristics, such as white flowers with purple stripes on the standard, racemose inflorescences and ex-aristate, reduced peduncles. However, the two species can be discriminated by floral and pod characters. Ononis diffusa (Fig. 2) has unequal ovate sepals, fused to 1/3 or more of their length; the standard is larger (10.5–11.5 × 7.0–8.0 mm), orbicular with a mucronate apex; and the pod is obviously beaked (rostrate). In contrast, O. serrata (Fig. 2) has equal sepals, linear to linear-lanceolate, fused to 1/4 or less of their length; the standard is smaller (5.2–7.8 × 6.0 mm), elliptic with an acute apex; and the pod is rostrate with a minute beak. In earlier Egyptian Floras (Täckholm 1956, 1974), the name O. serrata Forssk. var. major Lange (1865: 156) was used to refer to this taxon. Lange (l.c.) cited the following specimen for the name [SPAIN. Ex Tenerife: Bufadero, in saxosis camporum, 24 March 1855, Bourgeau N. 1317, COI00053519, (COI!)]. The specimen was then verified by Cosson as Ononis dehnhardtii Ten., and both names were synonymized under O. diffusa Ten. by Dobignard & Chatelain (2012). We accept this opinion here. Specimens examined:—EGYPT. (N): Cairo: In verkleinerter Wald, 29 April 1885, Volkens G. L. s.n., V-815555, (UPS-image!); (S): North Sinai: Rafah, April 1921, Hefnawy M. T.s.n., (CAIM!). ALGERIA. Forêt de M’ Sila, près El-Ançor Pelouses sablonneuses, 6 May 1923, Faure A. s.n., (CAI!).

6 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. FIGURE 2. Flower morphological features of Ononis species: (1) Ononis diffusa, (2) O. serrata, (3) O. mitissima, (4) O. variegata, (5) O. natrix subsp. natrix: (a) calices, (b) standards, (c) wings, (d) keels, (e) androcia and (g) gynoecia, scale bar = 5 mm.

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 7 2. Ononis serrata Forsskål (1775: 130). Type:—EGYPT. , March 1761, Forsskål P. 1519, C10002677, (C-image!), lectotype, designated here; Forsskål P. 1520, C10002678, (C-image!); ExOriente, s.d., Forsskål P. s.n., BM000901083, (BM-image!), syntypes. Note: Hepper & Friis (1994) mentioned all three specimens as types, (Fig. 3). Heterotypic synonyms: =Ononis glaucescens Pomel (1874: 167). Type:—ALGERIA. Brezina,19 April 1862, Pomel A.N. s.n., MPU004973, (MPU-image!); Metlili, 1 April 1862, Pomel A.N. s.n., MPU004974, (MPU-image!), syntypes. =Ononis marmorata Murbeck (1922: 58). Type:—MOROCCO. Environs de Marrakech : lieux incultes à Quéliz,15 April 1921, Murbeck S. s.n, LD1213338, (LD-image!); MPU008031, (MPU-image!), syntypes.

FIGURE 3. Lectotype of Ononis serrata Forssk. , preserved in (C).

Description:—Annual herb, 4–30(–50) cm high, much branched; branches erect to ascending or decumbent, viscid with glandular and eglandular hairs. Stipules green, ovate, 2.5–3.2 ×1.0–1.5 mm, acuminate apex, margins denticulate, entire in upper leaves, with five protruding nerves, covered with glandular hairs, shorter than the petiole. Leaves mostly trifoliate, flowering leaves simple, 5–15 mm long, petiolate; petiole 2.0–6.0 mm; leaflet oblong, 4.0–13.0 × 1.5–5.0 mm, acute apex, cuneate to attenuate base, dentate margins with 3–6(–10) pair of large spreading teeth; terminal leaflet sub-equal to lateral ones, with ca. 0.2 mm long petiolule. Inflorescence terminal, ± lax raceme, 27–100 mm long, 10–30 flowered, pedunculate; peduncle 3.0–4.0 mm long, shorter than subtending leaf, not aristate. Flowers sessile or with very short pedicels. Calyx campanulate, shorter than corolla, 4.5–7.5 × 1.0–3.0 mm, covered with glandular

8 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. hairs intermixed with long eglandular setae; calyx lobes 2.0–4.5 mm, 3– or 5– nerved, linear-lanceolate to ovate, with acute apex; calyx tube 1.5–4.0 mm, calyx lobes longer than tube. Corolla white with pink veins on standard and pink notch on keel apex; standard elliptic, covered with glandular hairs, 5.2–7.8 × 6.0 mm, acute to sub-obtuse apex, entire margins, with distinct subulate claw ca. 2.3 mm long; wings narrowly oblong, lamina 4.5–5.5 × 0.8–2.0 mm, claw 2.0–3.0 mm long, auricle very small ca. 0.3 mm long, blunt conjunctive tooth; keel curved to a blunt beak, lamina 5.0–6.2 × 2.0 mm, claw 2.0–3.0 mm long, standard sub-equals keel and wings. Stamens 4.1 mm, anthers ovoid oblong, subequal, 0.2 × 0.1 mm. Ovary 3.5 × 2.0 mm, covered with glandular hairs; style straight, not deflexed, ca. 6.0 mm long, glabrous. Pod ovoid to elliptic, 5.0–6.0 × 2.0–4.0 mm, apex very shortly rostrate, beak 0.1–0.2 mm long, covered on the upper half and distal margin with glandular hairs, fruiting calyx accrescent; seeds few, 2–5 seeds, globose or sub-globose to elliptic or reniform, 1.1–1.3 × 0.8–1.4 mm, papillose, with light to dark brownish testa. Conservation status:—Not threatened. Phenology:—Flowering season March–June. Habitat:—Sandy sub-coastal soils, desert wadis and plains. Iconography:—Webb & Berthelot (1836), Townsend (1974), Jafri (1980) and Rechinger (1984). Chromosome number:—2n = 30–32 (Fernandes & Queiros 1978, Bartolo et al. 1979). Distribution in EGYPT:—Native and widely spread in northern Egypt (Fig. 1); the species occurs in three main phytogeographical regions: M, N (Nile Delta) and S (northern coasts of Sinai) (Täckholm 1956, Boulos 1999). Global distribution:—This species prefers Mediterranean littoral and maritime habitats and the Mediterranean/ Sahara transition zones. Its distribution expands from North Africa (Libya, Algeria, Morocco and Tunisia) to southern and central Europe (i.e., Greece, Crete, Cyprus, Italy, France, Corsica, Sardinia, Portugal, Sicily and Spain) and to western Asia in Turkey, Iraq, Iran, Syria, Israel, Jordan, Lebanon, Palestine, Saudi Arabia, Bahrain, Kuwait and the United Arab Emirates (Greuter & Raus 1989). Taxonomic notes:—Ononis serrata is very close to O.diffusa. However, the two species can be distinguished by flower and pod characters. O. serrata (Fig. 2) has equal sepals, linear to linear-lanceolate; the standard is small (5.2–7.8 × 6.0 mm), elliptic with an acute apex; and the pod is rostrate with a minute beak, whereas, O. diffusa (Fig. 2) has unequal ovate sepals; the standard is larger (10.5–11.5 × 7.0–8.0 mm), orbicular with a mucronate apex; and the pod is obviously beaked (rostrate). Ononis serrata exhibits a high degree of polymorphism; hence, many previous authors suggested further segregation into many infraspecific taxa (i.e., Boissier 1872, Battandier & Trabut 1888, Maire 1926); however, the variability in characters (leaf blade, sepal shape, and size and shape of pods) shows a continuum; therefore, the current belief is to reject those infraspecific taxa (i.e., Greuter & Raus 1989, Boulos 1999, 2009). Specimens examined:—EGYPT. (M): : East Sallum, Saret Retama, sandy soil, 16 April 1934, Shabetai J. R. z3385, (CAIM!); Saret Retama, 16 July 1934, Drar M.s.n. (CAI!); (M): : On the coastal road, 44 km before Mersa Matruh, 03 March 1966, Täckholm V. s.n., (CAI!); West of Mersa Matruh, s.d., Täckholm V. s.n., (CAI!); El-Garawla between Al Alamein and Mersa Matruh, 22 March 1974, Täckholm V., El-Hadidi M. N., Ayyad S. & Mahdi M. s.n., (CAI!); El-Garawla, 15 km before Mersa Matruh, 31º 15’ 34” N, 27º 22’ 12” E, Alt. 11m, 07 April 2015, Faried A., Olwey A. & Hassan M. s.n. (ASTU!); (M): Mariut: Burg El-Arab, west Alexandria, 30 March 1903, Brown T. W. s.n., (CAIM!); Abu Seer, note on sheet: common on sandy soil, 23 April 1929, (CAIM!); Mediterranean region, 02 February 1929, Anonymous s.n., (CAIM!); Mariut, sandy soil, 06 March 1930, Drar M. 649, (CAIM!); 6 km East Abu Sir, 19 March 1938, Täckholm V. s.n., (CAI!); Al Alexandria: western Mediterranean coastal region, s.d., Anonymous s.n., (CAIM!); Abu Seer-Hammam road, along the seaside 24 March 1961, Täckholm V. s.n., (CAI!); Mariut, 02 April 1971, Imam M. s.n., (CAI!); Mariut, Maritime dunes, March 1980, Fayed A. s.n. (ASTU!); Northern Mediterranean coastal region, Idku, near the town 31º 18’ 10.8” N, 30º 19’ 10.7” E,12 April 2005, El-Darir S., Bidak L., Shehata A., Rashad S., Shiha M., Marzouk R., Toto S. & Boulos L. 20460, (MAZHAR-image!); (M): : El Alamein, 16 March 1957, Sa’ad F. s.n., (CAIM!); El Alamein, 16 March 1963, El-Hadidi M. N. s.n., (ASTU!); El Alamein, 30° 49’ 25.19’’ N, 28° 57’ 5.2596’’ E, 30 March 1969, Abbas A & El Shaer A. 1464, (CAIM!); Sidi Abd-El Rahman, 31 March 1969, Abbas A. & El Shaer A. 1315, (CAIM!); (M): Burg El Arab: Burg El Arab, 18 March 1938, Täckholm V. s.n., (CAI!); Burg El Arab, Spring 1952, Botany Dept. excursion s.n., (CAI!); Burg El Arab, 08 April 1955, El-Hadidi M. N. s.n., (CAI!); Burg El Arab, the main road westwards, at the top of the ridge, alt. 29.45m, 21 March 1956, Arvidsson, I. s.n., V-066280, (UPS-image!); on the road Alexandria-Burg El Arab at Kilo 17, 22 March 1956, Imam M. s.n., (CAI!); Burg El Arab, 22 March 1956, El-Hadidi M. N. s.n., (CAI!); Burg El. Arab, 14– 17 March 1958, Täckholm V.s.n., (CAI!); Burg El Arab, 22 March 1956, Elsayed I. s.n, (CAI!); Burg El Arab, 01 April 1960, Täckholm V. s.n., (CAI!); Burg El Arab, 24 March 1961; Täckholm V. s.n., (CAI!); Burg El Arab, Spring 1961, Heiwagi E. s.n., (CAI!); Burg El Arab, 29 March 1973, Elsayed I., Mahdi M. & Sisi S. s.n., (CAI!); Burg El Arab, 29

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 9 May 1973, Mahdi M. & Sisi S. s.n., (CAI!), Burg El Arab, 24 March 1977, Gazzar A., Fayed A. & Sisi S. s.n., (CAI!); Burg El Arab, sandy soil, 03 April 1977, El Khangry S. 524, (CAIM!); Umm Sigheiu west of Alexandria, sandy soil, 02 June 1977, El Khangry S. & Mokhtar M. 363 (CAIM!); Burg El Arab, 23 March 1987, Zareh M. s.n. (ASTU!); (M): Alexandria District: , 5 April 1887, Ascherson P. et al. s.n., B-100629399 (BMBG-image!); Sidi Gaber, sables maritimes, 24 April 1908, Lwans s.n., (CAI!); Abu Kir,18 April 1913, Bollandi G. C. s.n., (CAIM!); Dans les prairies sabloneusesprès la Mamourah au nord-est d’ Alexandrie, 24 March 1911, Hartmaun E. s.n., (CAI!); Dans les prairies sabloneuses près la Montzah au nord-est d’ Alexandrie, 04 March 1911, Hartmaun E. 384, (CAIM!); Victoria, East of Alexandria, in sand, 09 February1917, Brown T. W. s.n., (CAIM!); East of Alexandria, Victoria, 13 March 1923, Simpson N. D. 1863, (CAIM!); Mandara, 17 April 1923, Simpson N. D. 2049, (CAIM!); Abu Kir, 26 March 1927, Täckholm G. s.n., (CAI!); Maamora, east of Alexandria, 14 March 1928, Hefnawy M. T.s.n., (CAI!); Mandara, 08 March 1929, Hefnawy M. T. s.n., (CAI!); Abu Kir, 25 March 1954, Boulos L. s.n., (CAI!); Abu Kir, sandy soil, 24 March 1969, Wissa R. & Nabih A. 3278 (CAIM!); 75 km from Alexandria, on the desert road to Cairo, 05 April 1976, El-Hadidi M. N. et al. s.n., (CAI!); Sidi Krir, along the west Mediterranean region, 23 March 1977, Shamass E. s.n., (CAI!); (N): Beheira Province: El Maadia on the road Alexandria-, 26 March 1974, Täckholm V., El-Hadidi M. N., Ayyad S., Mahdi M. & Fayed A. s.n., (CAI!); Rosetta, 25 April 1987, Amer A. 10281, (CAI!); Rosetta, 06 April 1988, Amer A. 16332, (CAI!); (N): Fayoum: Wadi El Natroun, 01 May 1976, Chrtek J. s.n., (CAI!); (S): North Sinai: West El Humur& El Ramla, on deep sand, 14 April 1917, Shabetai J. R. z4122 (CAIM!); Rafah, April 1921, Hefnawy T. s.n., (CAIM!); Rafah, 13 May 1921, Drar M. & Shabetai J. R. s.n., (CAIM!); in Wadi, south of the Road on sandy plain, 20 March 1922, Simpson N. D. 848, (CAIM!); Wadi Liblaba near Um Hamsa, 31 March 1922, Simpson N. D. 915, (CAIM!); Gebel Khashab at 280 m., 26 April 1922, Simpson N. D. 1192a, (CAIM!); Rafah, 01 May 1922, Anonymous s.n., (CAIM!);Wadi El (note on sheet: pods erect, prostrate),14 March 1924, Simpson N. D. 2513, (CAIM!); Wadi El Arish (note on sheet: spikes very densely flowered, plant half buried in sand), 14 March 1924, Simpson N. D. 2533, (CAIM!); Gebel Lehfen, 17 March 1928, Shabetai J. R. s.n., (CAIM!); El-Arish, 20 March 1928, Täckholm G. s.n., (CAI!); Sinai, Bir Lehfen south of El-Arish, 21 March 1928, Täckholm G. s.n., (CAI!); Rafah. Near the station, 22 March 1928, Täckholm G. s.n., (CAI!); El Arish, 16 May 1928, Shabetai J. R. s.n., (CAIM!); Gebel El Few, on sandy hill, 11 December 1929, Shabetai J. R. z641, (CAIM!); South of Arish, 05 April 1939, Mokhtar M. s.n., (CAIM!); North west Sinai. Mitla Pass, El Shatt, 10 May 1939, Shabetai J. R. 656, (CAIM!); Wadi Anquabiya, April 1950, Elsayed I. s.n., (CAI!); Wadi El-Arish, 18 March 1955, Boulos L. s.n., (CAI!); North El Busseili, sandy soil, 12 April 1955, Khattab A. 1657, (CAIM!); Rafah. Sandy soil, 31° 16’ 48.4392’’ N, 34° 14’ 24.6084’’ E, 18 April 1956, Khattab A.64, (CAIM!); El Tahrir Province, 08 November 1962, Täckholm V., Kassas M., Samy M., Girgis W.A. & Lahran M. A. s.n., (CAI!); Abu El Matamir, Gianaklis farm, 28 May 1967, Täckholm V., Elsayed I. & Mahdi M. s.n., (CAI!); Rafah, cultivated land, 28 March 2005, Amer W. s.n., (CAI!); Rafah, 29 March 2005, Amer W. s.n., (CAI!); protectorate, 29 March 2005, Amer W. s.n., (CAI!); Rafah, coastal reclaimed area, 29 March 2005, Amer W. s.n., (CAI!). PALESTINE. Om El Kilâs, sandy ground, 04 June 1934, Khattab A. z4832, (CAIM!); Khan Yunis, Goret El- Lout, 09 March 1955, Boulos, L. s.n., (ASTU!); Rafah, on the sand dunes near the sea, 31 March 1955, Boulos L. 391, (ASTU!) and (CAI!). LIBYA. Tripoli, Sidi El-Masri, Faculty of Science, 01 April 1967, Boulos L. 1759, (CAI!); 22 km west of El-Khoss on calcareous slopes, 25 April 1967, Boulos L. 1781, (CAI!); Leptis magna, 08 March 1968, Boulos L. 1860, (CAI!); Leptis magna, note on sheet: Flowers pink & white, 22 March 1970, Boulos L., Khalifa & Sherif 3855, (CAI!); Salt marsh at Al Kararima, 23 March 1970, Boulos L., Khalifa & Sherif s.n., (CAI!). ALGERIA. La Macta, prés de Mostaganem. Peloures sablonneuses, 07 May 1916, Faure A. s.n., (CAI!). MOROCCO. Remota Jahanl et Maire,1926, Lindberg H. 2071, (CAI!). IRAN. 5–8 km south-east of Bashi, south of Bushir, 50 m alt., 11 April 1974, Davis P. H. & Bokhari M. H. D56540, E00336617, (E-image!). IRAQ. Shahraban, 31 May 1957, Rechinger K.H. 9726, E00336615 (E-image!).

3. Ononis mitissima Linnaeus (1763: 1007). Type:—LUSITANIA. Barbados, Anonymous s.n., LINN896.7, lectotype, designated by Devesa (1988), then by Förther in Turland & Jarvis (1997), (LINN-image!). Homotypic synonyms: ≡ Bugranopsis mitissima (L.) Pomel (1874: 172).

Description:—Annual herb, 20–75 cm high, branched; branches erect to ascending, pubescent with glandular hairs. Stipules white, membranous, conspicuous, ovate, oblong to obovate, 3.0–9.0 × 1.5–2.0 mm, acute apex, with entire margins, lower-most stipules with denticulate margins, striate with many protruding nerves, glabrous, shorter than the petiole. Upper leaves simple, rest trifoliate, flowering leaves simple, 10–35 mm, petiolate; petiole 4.0–7.0 mm; leaflet

10 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. obovate, 6.0–20.0 × 2.0–12.0 mm, acute apex, cuneate base, dentate margins with 13–20 pairs of teeth; terminal leaflet distinctly larger than lateral ones about 1.5–2-fold, with 3.0–5.0 mm long petiolule. Terminal inflorescence, dense spike, 9–45 mm long, 10–15(20) flowered, pedunculate; peduncle 8.0–16.0 mm long, shorter than subtending leaf, not aristate. Flowers sessile or with very short pedicel. Calyx tubular, corolla equal to or slightly exceeding calyx, 9.0–12.5 × 0.5–1.5 mm, covered with glandular hairs; calyx lobes 6.0–7.0mm, 3–nerved, oblong, with acute apex; calyx tube 3.0–4.0 mm, calyx lobes longer than tube. Corolla white with pink standard; standard obovate to rhombic, glabrous, 3.5–10.0 × 2.5–3.5 mm, obtuse to acute apex, sinuate margins, with ± distinct claw 0.25–2.0 mm long; wings oblong , lamina 2.5–5.0 × 0.5–1.0 mm, claw 0.8–1.5 mm long, auricle very small 0.1–0.2 mm long, conjunctive tooth present; keel narrow, gently curved to a short subacute beak, lamina 2.0–5.0 × 1.0–1.5 mm, claw ca.2.0 mm long; standard longer than the sub-equaling keel and wings. Stamens ca. 3.0 mm, anthers oblong, 0.5 × 0.3 mm; wide filaments, strongly dilated just below anthers. Ovary 1.5–2.5 x 0.5–1.0 mm, glandular-pilose hairs; style straight, not deflexed, 2.5–4.8 mm long, glabrous. Pod elliptic, 6.0 × 3.0 mm, apex shortly rostrate, beak ca. 0.5 mm long, covered on the upper half with few glandular hairs, fruiting calyx not accrescent; seeds few, 2–5 seeds, orbicular-reniform, 1.8–1.9 × 1.13–1.7 mm, densely papillose-verruculose, with brownish to blackish testa. Conservation status:—Not threatened. Phenology:—Flowering season April–June. Habitat:—Alluvial soils and sandy desert plains. Iconography:—Dillenius (1732) and Townsend (1974). Chromosome number:—2n =30 (Fernandes & Queiros 1978). Distribution in EGYPT:—Native to Egypt; this species often occurs as a weed, especially in wheat fields, and it is centred in two regions (Fig. 1): M and N (Nile Delta), (Täckholm 1956, Boulos 1999). Global distribution:—This plant is common in the Mediterranean bushlands in Europe in Crete, Cyprus, Greece, Sicily, Italy, France, Spain, Malta, Corsica, Balearic, Portugal and Sardinia; the North African coasts (i.e., Morocco, Algeria and Tunisia) and northwestern Asia in Turkey, Iraq, Jordan, Syria, Lebanon and Palestine (Greuter & Raus 1989). Taxonomic notes:—Only Ononis mitissima and O. variegata can be differentiated from the remaining taxa by glabrous stipules instead of glandular ones. However, the stipules of O. mitissima are large and membranous, whereas the stipules of O. variegata are small and green. Ononis mitissima is also unique due to its spicate inflorescence and calyx tube that is much longer than the calyx teeth. Specimens examined:—EGYPT. In Aegypto, s.d., Godron s.n., V-815563, (UPS-image!); (N): Cairo: Tel El Kebir, 7 May 1880, Khinfũth G. s.n., (CAIM!); Shubra, on wayside of fields, 10 April 1940, Shabetai J. R. z6248 (CAIM!); El Marg, Ezbet Adès, in wheat field, 27 April 1948, Shabetai J. R. z6665 (CAIM!); (N): Bahariya: El Bawrti, 13 April 1935, Lars M. s.n., (CAIM!). ALGERIA.Oued-Imbert (Oran) Bords des champs, 20 June 1918, Faure A. s.n., (CAI!); Départ d’Alger, Maison-Carréelieux herbaux français, 30 April 1953, D’Alleizette Ch. s.n., (CAI!). MOROCCO. In pascuist ingitanis. [Tanger], s.d., Salzmann P. s.n., M0210744 (M-image!). PORTUGAL. El vas, Herdade de Ovelheira, 12 July 1956, Beliz M. & Guerra J. A. s.n., (CAI!). UNKNOWN. Sin. loc., Anonymous s.n., from Schweinfurth’s Herbarium, Berlin (now burnt), (CAI!).

4. Ononis variegata Linnaeus (1753: 717). Type:—EUROPE. in australis maritimis, Royen van, A.s.n. , Herb. A. van Royen in Naturalis Biodiversity Centre, formerly known as Leiden University, L-908.120-1904, lectotype designated by Förther in Turland & Jarvis (1997), L-0052931, syntype, (L-image!). Homotypic synonym: ≡ Bugranopsis variegata (L.) Pomel (1874: 172). ≡ Anonis variegata (L.) Lamarck (1779: 608). Heterotypic synonyms: = Ononis aphylla Lamarck (1785: 509). Type:—ITALY. Habitat in Gallo provincia. Barrelier, J. 776, in Barrelier, Pl. Galliam, Hisp., et Ital. 74 (1714), [Icon.]. = Ononis variegata L. var. erioclada De Candolle f. coronata Maire (1949: 132). Type:—ALGERIA. A Ténès, sables martitimes, May 1881, Battandier J.A. s.n., MPU004728 (MPU-image!). = Ononis variegata L. var. alleizettei Faure & Maire (1937: 350). Type:—ALGERIA. Mostaganen vers le Djebel Diss, pelouses sablonneuses, 1 May 1936, Faure A.s.n., P00465617, (P-image!).

Description:—Small annual herb, 3–15 cm high, branched; branches procumbent to ascending, pubescent with eglandular rough walled hairs. Stipules green, sheathing, obovate-rhombic, 4.0 × 1.0–1.5 mm, acute apex, with

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 11 denticulate margins, covered with eglandular hairs. Leaves mostly simple, lowermost often trifoliate, flowering leaves simple, leaves sessile, leaflet ovate, 4.0–6.0 × 1.0–2.0 mm, acute apex, cuneate base, dentate margins with 5–7 pairs of teeth. Flowers solitary; pedicel ca. 3.5 mm long; peduncle not aristate nor pendent. Calyx campanulate, shorter than corolla, 6.0–7.0 × 1.0–1.5 mm, covered with eglandular rough hairs; calyx lobes 3.0–3.5 mm, 5– nerved, ovate, with acute apex; calyx tube 3.0–3.5 mm, calyx lobes almost equal to the tube. Corolla yellow with darker veinsstandard; standard spathulate, covered with eglandular hairs, 6.5 × 3.5 mm, obtuse apex, entire margins, with distinct claw ca. 2.0 mm long; wings lamina 3.5 × 1.5 mm, claw ca. 2.5 mm long, auricle small ca. 0.5 mm long, conjunctive tooth absent; keel lamina 5.5 × 1.5 mm, with long subulate claw ca. 4.5 mm long, keel obviously longer than standard and wings, standard a little longer or equalingwings. Stamens 5.5 mm. Ovary 2.5 × 1.0 mm, covered with eglandular hairs; style straight, not deflexed, 6.5 mm long, covered with eglandular hairs near the base. Pod elliptic to linear, 4.0–5.0 × 1.5–2.0 mm, apex very shortly rostrate, covered with eglandular hairs, fruiting calyx not accrescent; seeds few, 2–6 seeds, reniform,1.25 × 0.75 mm, ±smooth, with brownish testa. Conservation status:—Not threatened. Phenology:—Flowering season March–May. Habitat:—Littoral sandy soil. Iconography:—Jafri (1980), Collenette (1999). Chromosome number:—2n = 16–30 (Sanudo et al. 1979, Runemark 2006). Distribution in EGYPT:—Rare species; collected a few times from two localities; M and S (Rafah), (Täckholm 1956, Boulos 1999). Global distribution:—A small herb growing on Mediterranean littoral and maritime sand, expanding from the North African coasts (Libya, Algeria, Morocco and Tunisia) to southern Europe (Corsica, Crete, Cyprus, Greece, Sicily, Italy, Spain, Albania, Malta, Portugal, and Sardinia) and northwestern Asia in Turkey, Palestine, Jordan, Lebanon and Syria (Greuter & Raus 1989). Taxonomic notes:—In contrast to all Egyptian Ononis, O. variegata is strictly covered with eglandular hairs. Ononis variegata and O. vaginalis are unique in possessing sheathing stipules. Both species have two types of leaves (simple and trifoliate), but only O. variegata has simple leaves as the dominant type and rarely bears trifoliate leaves. Specimens examined:—EGYPT. ExAegypto, 1823, Sieber F. W., s.n., M0232964, (M-image!). ALGERIA. La Macta: prés de Mostaganem sables maritimes, 07 May 1916, Faure A.s.n., (CAI!); Sable maritime: à Sidi-Ferruch, 18 April 1953, D’Alleizette Ch. s.n., (CAI!).

II. Section Natrix Grisebach (1843: 12)

5. Ononis natrix Linnaeus (1753: 717).

Description:—Perennial subshrub, 10–60 (–100) cm high, much branched; branches erect to ascending, densely pubescent with glandular hairs intermixed with eglandular setae. Stipules green, linear-lanceolate, 8.0–10.0 × 1.5–2.0 mm, with elongated subulate apex, and entire margins, often denticulate, striate with 5 protruding nerves, shorter than the petiole. Mostly trifoliate, upper leaves simple, some leaves compound of 5–7 leaflets, flowering leaves simple, 8–45 mm, petiolate; petiole 5.0–20.0 mm; leaflet ovate, obovate to linear-lanceolate 2.0–20.0 × 2.0–10.0 mm, acute apex, cuneate base, dentate margins with 3–20 pair of teeth; terminal leaflet distinctly larger than lateral ones about 1.5–2-fold, with 1.0–6.0 mm long petiolule. Flowers pedunculate, peduncle 6.0–20.0 mm long, as long as or longer than subtending leaf, peduncle aristate; aristum 1.0–10.0 mm long. Flowers pedicellate; pedicel 1.5–3.0 mm long. Calyx tubular, corolla much exceeding calyx, 9.0–12.5 × 0.5–1.5 mm, densely covered with glandular hairs; calyx lobes 3.5–11.0 mm, 3–nerved, linear-lanceolate, with acute apex; calyx tube 2.0–4.0 mm, calyx lobes longer than tube. Corolla yellow with red or purple veins on standard; standard obovate, glabrous, 7.5–11.0 × 5.0 mm, obtuse apex, entire margins, with distinct claw ca. 0.6 mm long; wings lamina 6.0–9.0 × 3.0–3.5mm, claw 1.5–2.0 mm long, auricle ca. 0.5 mm long, conjunctive tooth absent; keel lamina 5.0–7.0× 4.0 mm, claw ca. 2.0 mm long; standard longer than keel and wings, wings slightly longer than keel. Stamens ca. 6.3 mm. Ovary 3.0–4.0 × 1.0–1.5 mm, covered with glandular and eglandular hairs; style straight, not deflexed, 3.0–3.5 mm long, glabrous. Pod linear, 6.0 × 3.0 mm, apex rostrate, beak 1.0 mm long, densely covered with long silky glandular and eglandular hairs, fruiting calyx not accrescent; seeds many, 6–10 seeds, reniform, 1.4 × 1.6 mm, loosely papillose, with blackish testa. Conservation status:—Not threatened. Phenology:—Flowering season March–April. Chromosome number:—2n=32 (Natarajan 1978).

12 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. Global distribution:—Mediterranean region northwards to southern and central Europe: Sicily, Italy, Greece, Crete, Germany, France, Spain, Balearic, Malta, Portugal, Switzerland, Corsica and Sardinia; southwards on the North African Mediterranean coasts: Libya, Algeria, Morocco and Tunisia. Eastwards in Asia: Turkey, Jordan, Lebanon, Palestine, Syria, Iraq, Saudi Arabia and China (Greuter & Raus 1989). Taxonomic notes:—Two subspecies occur in Egypt; their morphological differences are contrasted in the following key:

- Leaves oblong to obovate (2.0–20.0 × 2.0–10.0 mm), petiole 5.0–20.0 mm long, longer than stipules; standard large, 9.0–15.0 × 9.0 mm, with subulate claw (ca. 2.0 mm) ...... O. natrix subsp. natrix - Leaves elliptic to linear-lanceolate (1.0–3.0 × 1.5–2.0 mm), petiole 4.0–4.5 mm long, shorter than stipules; standard small, 7.5– 11.0 × 5.0 mm, with short claw (ca. 0.6 mm) ...... O. natrix subsp. stenophylla

5a. Ononis natrix Linnaeus subsp. natrix Type:—NETHERLANDS. Cultivated material from the garden of George Clifford III: Holland: Hartekamp Garden, George Clifford s.n, Linn vidi Microfiche, BM000646575, holotype, (BM-image!). Note: this type was also cited in Förther & Poldlich (1991: 263). Heterotypic synonyms: = Ononis pinguis Linnaeus (1763: 1009). Type:—Ex. Herb. Magnol. (Linn. vidi. Microfiche), (MPU). = Ononis inaequifolia De Candolle (1825: 165). Type:—EUROPE. France, Corsega, St. Florent, 06 March 1851, Soleirol J. F. 1349, COI00053570, (COI-image!). = Ononis natrix L. var. media Boissier (1840: 149) ≡ Ononis natrix L. subvar. media (Boiss.) Širjaev (1932: 456). Type:—SPAIN. Altea in arenis maritimis et torrentis, s.d., Boissier P. E. s. n., G00370361, lectotype, designated by España (1958); G0037036, G00370364, syntypes (G-image!). = Ononis anomala Pomel (1874: 456) ≡ Ononis natrix L. subsp. ramosissima (Desfontaines) Briquet var. genuina (Rouy) Širjaev subvar. quinquefoliata Širjaev (1932: 463). Type:—ALGERIA. Ghar-Rouban, Pomel A.N. s.n., MPU024416, holotype, (MPU-image!). = Ononis ambigua Lange (1877: 238). Type:—EUROPE. Spain, Annonymous 833, BM000751114 (BM-image!).

Habitat:—Stony sub-coastal desert. Iconography:—Weinmann (1737), Jaume Saint-Hilaire (1825), Jafri (1980), Förther & Podlech (1991) and Collenette (1999). Distribution in EGYPT:—Very rare; this taxon was once collected from Rosetta in the Nile Delta by Muschler in 1902; then, Boulos (1999) recorded this taxon on the Mediterranean coasts. Specimens examined:—EGYPT. (N): Rosetta, October 1902, Muschler s.n., (G-image!). LIBYA. 8 km south of Al-Qubbah, 22 January 1967, Boulos L. 1326, (CAI!); 12 km East of Tripoli near seashore, 24 March 1967, Boulos, Tawil & Bouni 1615, (CAI!); Tripoli, Sidi El-Masri, barely fields facing Faculty of Science, 18 April 1967, Boulos, L. 1697, (CAI!); Wadi Derna,14 March 1968, Boulos L. 2435, (CAI!); Marawah. Gebl-Akhdar,15 March 1968, Boulos L. 2487, (CAI!); Leptis magna, April 1968, Brydolf Ch. s.n., (CAI!); Below Shahhat, note on sheet: Flowers yellow-white with brown stripes on the standard, 500–600 m alt., 26 March 1970, Boulos, Khalifa & Sherif 4034, (CAI!). PALESTINE. Gazza, 15 March 1926, Drar, M. s.n., (CAIM!). LEBANON. Sin. loc., June 1954, Täckholm, V. 85, (CAI!). SYRIA. Env. de Hammah, 1500 FT alt., 20 April 1908, Haradjian M. 1867, E00334247, (E-image!). JORDON. Medaba, 770 m alt., 24 April 1911, Meyers F. S. & Dinsmore J. E. M1648, E00334269. (E-image!). ISREAL. Yafne, Philistaean Plain, 30 December 1960, Orshan G. s.n., E00334287, (E-image!). SWITZERLAND. Collected around the alpine station “La Linnaea” Burg St. Pierre, June–August 1936, Migahid A. M. & Montasir A. H. s.n., (ASTU!). FRANCE. Marseile, May 1761, Forsskål P. 1514, C10002736, (C-image!); Card, commune: Pujaut; lieu-dit Le Planas, acciennes gravières; sable et graviers., 43º 58’ N,4º 43’ E,50 m alt., 19 June 1974, Aujard-Catot R. & Ledoux J. CA. s.n., (CAI!). ITALY. Sicily, s.d., Anonymous s.n., LINN-HS1195-23, (LINN-image!). GALLICA. Savoie, Col du Glandon, Alt. 800 m alt., 21 August 1961, Folkeson E. s.n., (CAI!).

5b. Ononis natrix L. subsp. stenophylla (Boiss.) Širjaev (1932: 470). Type:—SYRIA. Ad Joppe: Loadicream, May1817–November 1817, Sieber F.W. s.n.; lectotype, designated by Förther & Poldlich (1991: 280), hoc. loco. designates, (G-Bois).—ISRAEL. Tel Aviv District: Joppe [=Jaffa], May1817–November 1817, Sieber F.W. & Kohaut F. s.n., MEL2347544, isolectotype, (MEL-image!).—ISRAEL. Sieber F.W., s.n., MNHN-P-P00708494, isolectotype, (P- image!).—SYRIA. ad Gaza: prope Gaza, April 1846, Boissier E.s.n., MEL2347543, syntype, (MEL-image!).—ISRAEL, Sieber F.W. s.n., P00708494, syntype, (P-image!). Homotypic basionym: ≡ Ononis natrix var. stenophylla Boissier (1872: 59). ≡ Ononis stenophylla (Boiss.) Buxbaum (1926: 56).

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 13 Habitat:—Maritime coastal sand dunes. Iconography:—Förther & Podlech (1991). Distribution in EGYPT:—Native to Egypt, occurs in two main localities (Fig. 1): M and S (El Arish-Rafah region), (Täckholm 1956, Boulos 1999). Taxonomic notes:—Täckholm (1956) mentioned this taxon as a variety rather than a subspecies. Förther & Podlech (1991) provided a detailed and comprehensive study on Ononis natrix s.l. in which they raised the taxonomic level of this taxon. Specimens examined:—EGYPT. (S): North Sinai: Rafah, 19 March 1926, Shabetai J. R. s.n., (CAIM!); Rafah, near the station, 22 March 1928, Täckholm G. s.n. (CAI!); West of El Arish, sandy desert, 19 April 1930, Shabetai J. R. z 3687, (CAIM!); Rafah, desert sands of north the coast, 04 June 1934, Shabetai J. R. z 6032, (CAIM!); (M): Mediterranean region, note on sheet: common on sand and sand dunes, 09 April 1927, Drar M. 1667, (CAIM!); Mediterranean region, 23 September 1929, Drar M. 479, (CAIM!). LIBYA. Sidi Masri district 2 km east of Tripoli, 24 January 1958, Naguib M. I. s.n., (CAI!); 10–20 km East of Tripoli along the road to Khome, 22 November 1966, Boulos L. s.n., (CAI!); Guddaiem, note on sheet: Flowers yellow, standard with purple strips, 03 January 1970, Boulos L. 3433, (CAI!); Plateau at top of Escarpment, Ain Zargo, 0.7 km south-west of Jado, note on sheet: flowers yellow with brown stripes, 17 March 1970, Boulos & Khalifa 3653, (CAI!). JORDON. Along the mountain road between Jisr Damia & Salt, 14 July 1962, Täckholm V. s.n., (CAI!); Petra, 07 July–16 August 1962, Täckholm V. s.n., (CAI!); Jarash, in the mountain region, Deir El Alla. Irbid, 06 August 1962, Täckholm V. s.n., (CAI!). LEBANON. Dehour El Shouer, August 1952, Mawardi, H. A. s.n., (CAI!). PALESTINE. Gaza, April 1846, Boissier P.E. s.n., K000998649, (K-image!); Sand dunes south of Khan Yunis,14 January 1955, Boulos L. 67, (CAI!); El-Nuseirat, at the sea shore, 03 September 1955, Boulos L. 483, (CAI!).

6. Ononis vaginalis Vahl (1790: 53). Type:—EGYPT. Alexandria: Frequens in collibus peninsulae Râs et tîn, March 1762, Forsskål P. 1521, C10002679, lectotype, designated here; Forsskål P. 1522, C10002680; Forsskål P. 1523, C10002681, Forsskål P. 1524, C10002682 syntypes, (C-images!), (Fig. 4). Homotypic synonym: ≡ Ononis cherleri Forsskål (1775: 130), non Linnaeus (1763:1007). Heterotypic synonyms: = Ononis vaginalis Vahl var. rotundifolia Béguinot (1912: 132). Type:—LÝBYA . Tripolitania rocce marittime, 26 February 1912, Vaccari A.252, FI003752, holotype, (FI-image!) = Ononis vaginalis Vahl var. viviani Béguinot (1912: 132). Type:—LÝBYA. Marmarica Tobruk, 18 April 1912, Vaccari A. 61, FI003753, holotype, (FI-image!).

Description:—Perennial subshrub, 20–50 cm high, much branched; branches erect to ascending, covered with glandular hairs. Stipules green, sheathing, 2.5–8.0 × 1.0–3.0 mm, rhombic, with acute apex, anddenticulate margins, teeth on the upper half of margin, striate with multiple protruding nerves. Upper leaves simple, rest trifoliate, flowering leaves simple, 3.5–6.0 mm, sessile; leaflet ovate 1.0–3.0 × 1.5–2.5 mm, acute apex, attenuate base, dentate margins with 2–4 pairs of teeth; terminal leaflet distinctly larger than lateral ones about 1.5x–2x. Solitary flowers; flowers pedunculate, peduncle 12.0–24.0 mm long, much longer than subtending leaf, peduncle aristate, aristum 0.5–6.0 mm long. Flowers pedicellate; pedicel 2.0–3.0 mm long. Calyx tubular, corolla much exceeding calyx, 9.0–12.5 × 0.5–1.5 mm, densely covered with glandular hairs; calyx lobes 3.5–11.0 mm, 3–nerved, linear-lanceolate, with acute apex; calyx tube 2.0– 4.0 mm, calyx lobes longer than tube. Corolla yellow with red or purple veins on standard; standard obovate, glabrous, 7.5–11.0 × 5.0 mm, obtuse apex, entire margins, with distinct claw ca. 0.6 mm long; wings, lamina 6.0–9.0 × 3.0–3.5 mm, claw 1.5–2.0 mm long, auricle ca. 0.5 mm long, conjunctive tooth absent; keel lamina 5.0–7.0× 4.0 mm, claw ca. 2.0 mm long; standard longer than keel and wings, wings slightly longer than keel. Stamens ca. 6.3 mm. Ovary 3.0–4.0 × 1.0–1.5 mm, covered with glandular and eglandular hairs; style straight, not deflexed, 3.0–3.5 mm long, glabrous. Pod linear, 9.0–23.0 × 2.0–4.0 mm, apex rostrate, beak 1.0 mm long, densely covered with long silky glandular hairs, fruiting calyx not accrescent; seeds 2–6 seeds, reniform, 1.2 × 2.2 mm, very shallowly tuberculated, with brownish to blackish testa. Conservation status:—Not threatened. Phenology:—Flowering season April–August. Habitat:—Coastal sand. Chromosome number:—2n = 32 (Mohamed 1997). Distribution in EGYPT:—Near-endemic species in Egypt; very common along the eastern side of the Mediterranean coastal belt (Fig. 1). Its distribution is restricted to a single phytogeographical ecosystem (M), (Täckholm 1956, Boulos 1999).

14 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. FIGURE 4. Lectotype of Ononis vaginalis Vahl , preserved in (C).

Global distribution:—Narrowly distributed in the Mediterranean/Sahara transition zones. In North Africa: Libya and Tunisia, while in Asia: Palestine, Jordan, Lebanon and Syria (Greuter & Raus 1989). Taxonomic notes:—Perhaps one of the most common taxonomic problems in identifying Egyptian Ononis is the confusion between O. vaginalis and O. natrix. This confusion is due to the polymorphic nature of O. natrix. Both taxa are shrubby and have yellow flowers with purple or reddish stripes on the standard and flowers borne on aristate peduncles. However, the taxa can be easily distinguished by their leaves because in O. vaginalis, simple leaves are more common than trifoliolate ones, in contrast to O. natrix, where trifoliolate leaves are the dominant type of leaves. Moreover, O. vaginalis has sessile leaves and sheathing stipules, whereas O. natrix has petiolate leaves with non- sheathing stipules. Specimens examined:—EGYPT. (M): Sallum: Abbas, east of Sallum, 16 April 1934, Shabetai J. R. z3383, (CAI!); Sallum, 16 April 1934, Shabetai J. R. z3383, (CAIM!); Sallum, 26 May 1935, Anonymous 1754, (CAIM!); Wadi El Ausheily, west Matruh, rocky soil,13 April 1967, Abbas A.s.n., (CAIM!); Agiba, El Kasr, near Mersa Matruh, 31 May 1974, Helmy H. & Mokhtar M. 17, (CAIM!); Wadi Agiba, west of Mersa Matruh, 21 March 1975, Täckholm V., El Hadidi N. et al. s.n., (CAI!); Western Mediterranean coastal region; 8 km west of Agiba sand dunes,31º 26’ 16.7” N, 26º 55’ 46.6” E,-2 m alt., 21 May 2007, Hegazy A. & Sharashi O. 1745, (MAZHAR-image!); Ageba beach, Sallum to Mersa Matruh road, 31º 24’ 35” N, 27º 0’ 56”, 1 m, 07 April 2015, Faried A., Olwey A. & Hassan M. s.n., (ASTU!); (M): Mersa Matruh: Mersa Matruh, by sea coast, 16 February 1929, Melchens z681, (CAIM!); El Duba north-west coast on sand dunes, 22 May 1935, Drar M. s.n., (CAIM!); Mersa Matruh, sandy soil, 11 January 1940, Shabetai J. R.

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 15 z3978, (CAIM!); Mersa Matruh, 14 July 1959, Habbour S. s.n., (CAI!); Road between Matruh – Alexandria, in Barley field, 21 April 1971, Abbas A. & Khattab A. 75, (CAIM!); Mersa Matruh, 01 April 1972, Täckholm V. et al. s.n., (CAI!);The sea shore of Mersa Matruh, sandy soil, 27 May 1974, Helmy H. & Mokhtar M. s.n., (CAIM!); Mersa Matruh, 29 May 1974, Helmy H. & El Hediny A.s.n., (CAIM!); Hadabet Khashm El Esh,03 March 1976Täckholm V., Gazzar M. & Fayed A. s.n., (CAI!); Um Al-Rakhm road, Mersa Matruh, 31° 23’ 36.6’’ N,27° 3’ 34.488’’ E, 2 m alt., 30 May 2011, Zareh M. & Faried A. s.n., (ASTU!); Al Kasr road, Mersa Matruh, 31° 21’ 48.06’’ N, 27° 8’ 23.892’’ E, 5 m alt., 30 May 2011, Zareh M. & Faried A. s.n., (ASTU!); Al Kasr road, Mersa Matruh,31º 21’ 49” N, 27º 7’ 55” E, 3 m alt., 07 April 2015, Faried A., Olwey A. & Hassan M.s.n., (ASTU!); (M): Ras El Hekma: Ras El Hekma, 25 May 1954, Migahid & Shafey s.n., (CAI!); Ras El Hekma, 30 April 1955, Täckholm V. & The Botany department excursion s.n., (CAI); Ras El Hekma, Spring 1955, El Hadidi N. M. s.n., (CAI!); Al Iskandariyah (Alexandria) The Western Mediterranean coastal region. Ras el Hekma, 30.91667, 29.61667, 26 June 1956, Arvidsson I. s.n., V-066469, (UPS- image!); Ras El Hekma, in sandy dunes, 26 May 1963, Täckholm V., Boulos L., Girgis W., Zahran M. A. & Elsayed I. s.n., (CAI!); Ras El Hekma, 17 May 1964, Niazi & Khattab A.s.n., (CAIM!); Ras El Hekma, 03 June 1964, Täckholm V., Boulos L., Elsayed I. & Mahdi M. E. s.n., (CAI!); Ras El Hekma, 16 February 1965, Täckholm V. s.n., (CAI!); Ras El Hekma, sand plain, 25 September 1965, Osman D. & Ibrahim H. 33, (CAI!); Ras El Hekma, 02 April 1969, Abbas A. & El Shaer A. s.n., (CAIM!); Ras El Hekma. (note on sheet: very common on sandy dunes),14 April 1970, Abbas A.192, (CAIM!); Ras El Hekma, 22 April 1973, Amin A. et al. s.n., (CAI!); Ras El-Hekma, 31.45741 N, 26.79281 E,5 m alt., 01 June 2011, Zareh M. & Faried A. s.n., ( ASTU!); (M): El Alamein: On the Abu Sir to El Alamein road, in maritime sand, 01 June 1964, Täckholm V., Boulos L. Elsayed I. &. Mahdi M. E. s.n., (CAI!); Mediterranean region, between Alexandria & Alamein, 10 April 2010, Zareh M. & Aboul-Ela M. s.n., (ASTU!); Al-Hammam to Al Alameen road, 30º 51’ 17” N,29º 17’ 57” E,13 m alt.,06 April 2015, Faried A., Olwey A. & Hassan M. s.n., (ASTU!); (M): Burg El Arab: Mariut, Burg El-Arab, 15 March 1928, Täckholm G. s.n., (CAI!); El Burg, Mariut District, 24 April 1929, Drar M. 185, (CAIM!); Abu Sir, 25 March 1949, Täckholm V. s.n., (CAI!); Mariut, on the sea shore, 25 March 1949, Täckholm V. s.n., (CAI!); Mariut, 01 March 1950, Rateb, G. s.n., (CAI!); Burg El Arab, 08 March 1950, Shabetai J. R. z7636, (CAIM!); Abu Sir. Burg El Arab, 19 April 1950, Shabetai J. R. z7348, (CAIM!); Mariut, Burg El Arab,13 March 1951, Harmant B. s.n., (CAI!); On the way to Burg EL Arab, 13 March 1952, El Hadidi N. M. s.n., (CAI!); Mariut,13 March 1952, Ayyad S. s.n., (CAI!); Mariut, 13 March 1952, Kamel W. s.n., (CAI!); Burg El Arab, sandy soil, 07 June 1955, Shabetai J. R. z4533, (CAIM!); Mariut, Spring 1961, TäckholmV. s.n., (CAI!); Burg El Arab to El Alamein, along the road, 16 February1965, Täckholm V.s.n., (CAI!); On the sea coast 22 km west of Alexandria in sand dunes,15 January 1972, Zahran M. s.n., (CAI!); Burg El Arab, March 1974, Zahran & ElAraby s.n., (CAI!); Burg El Arab, 22 March 1974, Täckholm V., El Hadidi N., Ayyad S. & Mahdi M. s.n., (CAI!); Umm Sigheiu west of Alexandria, sandy soil, 07 May 1976, El Khanagry S. & Mokhtar M. 309, (CAIM!); El Gharbaniyat, 5 km from Burg El Arab on the way to Hammam, 27 January 1977, Kassas M. s.n., (CAI!); Burg El Arab at seashore, 11 March 1977, Kassas M.s.n., (CAI!); Burg El Arab, 24 March 1977, Gazzar A. Fayed A. & Sisi S. s.n., (ASTU!); Burg El Arab, 03 April 1977, El Khanagry S. 518, (CAIM!); Burg El Arab, 18 April 1977, Chrtek, Salvikova & Kosinova s.n., (CAI!); Gharbaniat, sandy dunes, 09 March 1978, Merxmüller et al. s.n., (CAI!); 60 km west of Alexandria, 01 March 1980, Fayed A.s.n., (ASTU!); Mariut, Burg El Arab, 01 March 1980, Fayed A. & El Naggar S. s.n., (ASTU!);Sidi Hatoush, 05 June 1982, Helmy H., Mokhtar M. & Ahmed K. H. 518, (CAIM!); Alexandria to Burg El Arab road, 31º 0’ 9” N, 29º 36’ 48” E,2 m alt., 06 April 2015, Faried A., Olwey A. & Hassan M. s.n., (ASTU!);(M): Abu Sir: Abu Sir, 08 October 1920, Anonymous s.n., (CAIM!); Abu Sir. Mariut District. (note on sheet: common on sandy soil), 22 April 1929, Drar M. s.n., (CAIM);Abu Sir, 19 March 1938, Täckholm V. s.n., (CAI!); Abu Sir, 27 April 1938, Shabetai J. R. z4835, (CAIM!); Abu Sir, 01 April 1952, Khattab A. E141, (CAIM!); Abu Sir, 10 March 1953, Fawzi A. M. s.n., (CAI!); Abu Sir, 20 March 1953, Kamel W. s.n., (CAI!); Abu Sir, 18 March 1954, Täckholm V. & the Botany Dept excursion s.n., (CAI!); Abu Sir, 25 April 1955, Khattab A. 174 (CAIM!); Abu Sir, sandy soil, 1 May 1961, Sharobim A. s.n., (CAIM!); Abu Sir, 25 November 1955, Khattab A. 174, (CAIM!); Abu Sir, in sand dunes, 04 April 1956, El Batanouny s.n., (CAI!); Mariut, Abu Sir, Spring 1962, Täckholm, V. & the Botany Dept excursion s.n., (CAI!); Abu Sir, in maritime sand, 10 March 1967, Täckholm V., Elsayed I. & Mahdi M. E. s.n., (CAI!); Abu Sir. Mariut, 18 April 1969, Täckholm V. s.n., (CAI!);Abu Sir, 19 October 1969, Abbas A. & El Shaer A. 2056, (CAIM!); Abu Sir,30 April 1976, Chrtek J. s.n., (CAI); Abu Sir, north Burg El Arab, sandy soil on the coast, 14 April 1981, Shabetai J. R. & Boulos, L. z7717, (CAIM!); (M): King Mariut: King Mariut, Mariut District, 17 November 1929, Drar M. s.n., (CAIM!); King Mariut, 04 May 1976, El Khanagry S. & Mokhtar M. 78, (CAIM!); King Mariut, sandy calcareous soil, 25 August 1977, El Khanagry S. 77, (CAIM!); King Mariut, 04 April 1980, Helmy H. & Mokhtar M. 8, (CAIM!); (M):Alexandria District: Alexandria, s.d., Sieber F. s.n., M0232963, (M-image!); Alexandria, 28 July 1848, Khinfũth G. s.n., (CAIM!); Prope Alexandria, 3 April 1877, Letourneux A. et al. s.n., B100629400, (BGBM-image!); Alexandria, Ramleh, 15 March 1887, Ascherson P. et al. s.n., B100629398, (BGBM-image!); Alexandrie, Chatby, sable maritime, 20 June 1907,

16 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. Lwans 281, (CAI!); sur les sables près Maamourah en Nord-est d’Alexandrie, 10 March 1908, Hartmann E. s.n., (CAI!); Alexandrie, sables maritimes,30 March 1908, Lwans 281, (ASTU!); Dans les prairies sabloneuses près la Montzah au nord-est d’ Alexandrie, 18 April 1908, Hartmann E. 181, (CAIM!);Dans les prairies sabloneuses et au les dunes près Montazah au nord d’ Alexandrie, 21 March 1911, Hartmann E. s.n., (CAI!); Ramlele, 24 May 1912, Bollandi G. B380, (CAIM!); Montazeh, 16 January 1918, Brown T. W. s.n., (CAIM!); Mandara, 17 March 1923, Simpson N. D. 2057, (CAIM!); Sidi Bishr, east of Alexandria, 17 April 1925, Simpson N. D. 3142, (CAIM!); AbuKir, 26 March 1927, Täckholm G. s.n., (CAI!); Maamora, east of Alexandria,14 March 1928, Hefnawy M.T. s.n., (CAI!); Victoria, north Alexandria, sandy soil, 09 March 1949, Shabetai J. R. s.n., (CAIM!); AbuKir, Mariut, 01 March 1954, El Hadidi N. M. s.n., (CAI!); Sidi Krir, 23 May 1974, Helmy H. & Mokhtar M. 35, (CAIM!); Sidi Krir, 23 March 1987, Fahmy A. G. 463, (CAI!); (N): Rashid, sandy soil, 01 April 1934, Drar M.s.n., (CAIM!); Fayium, sandy soil, 23 September 1936, Diwan B. H. et al. s.n., (CAIM!).

7. Ononis reclinata Linnaeus (1763: 1011). Type:—ITALY. Habitat in Delphinatu, Hispania, in Barrelier, Pl. Galliam, Hisp. et Ital. 74, t. 354 (1714), [Icon.], lectotype, designated by Förther in Turland & Jarvis (1997), (!). 7.a.Ononis reclinata L. subsp. mollis (Savi) Béguinot (1912: 134). Type:—ITALY. In collibus saxosis, in Savi, Mem. Soc. Ital. Sci. 9: 351, t. 4 (1802), [Icon.], (!). Homotypic basionym: ≡ Ononis mollis Savi (1802: 351). Heterotypic synonyms: = Ononis reclinata var. brevipila Murbeck (1897: 58). Type:—ALGERÝA. Lieux incultes à Nemours, ouest de la provience d’Oran, 8 May 1856, Bourgeau E. 141, MPU008895, (MPU-image!). = Ononis reclinata L. var. minor Moris (1937: 422). Type:—SARDINIA. Without precise locality, Moris s.n., holotype (TO).

Description:—Small annual herb, 3–20(–30) cm tall, branched; branches decumbent to ascending, densely covered with long eglandular hairs intermixed with shorter glandular hairs. Stipules green, not sheathing, 2.5–3.2 × 1.0–1.5 mm, obovate, with acute apex and denticulate margins with few teeth, striate with 5 protruding nerves. Lowermost and uppermost simple, the rest trifoliate, flowering leaves simple, 7.5–15 mm, petiolate; petiole 3.0–4.0 mm; leaflet deltoid cuneate, 4.0–10.0 × 1.0–4.0 mm, acute apex, attenuate base, dentate margins with 2–3 pairs of teeth; terminal leaflet distinctly larger than lateral ones by approximately 1.5–2x, with 0.5–1.0-mm-long petiolule. Solitary flowers; flowers pedunculate, peduncle 2.0–7.0 mm long, shorter than subtending leaf, peduncle not aristate. Flowers distinctly deflexed, pedicellate; pedicel ca. 1.0 mm long. Calyx tubular, as long as or slightly shorter than corolla, 6.0–7.0 × 0.5 mm, densely covered with short glandular hairs intermixed with long silky setae; calyx lobes 4.0–5.0 mm, 3-nerved; nerves often with red-purplish colour, linear-lanceolate, with acute apex; calyx tube 2.0–2.5 mm, calyx lobes 2x longer than tube. Corolla white with pink standard striated with purple veins and pink notch on keel apex, seldom yellowish with pink standard; standard ovate, glabrous, 4.5–5.0 × 2.2–2.5 mm, obtuse or acute apex, entire margins, with distinct claw 0.5–0.8 mm long; wing lamina 2.5–3.0 × 0.8–1.0 mm, claw ca. 1.2 mm long, auricle ca. 0.5 mm long, conjunctive tooth absent; keel lamina 2.5 × 1.5 mm, claw ca. 1.0 mm long; standard longer than keel and wings, wings as long as or slightly longer than keel. Stamens ca. 4.3 mm. Ovary 2.0 × 0.5–1.0 mm, covered with glandular and eglandular hairs; style deflexed, ca.3.7 mm long, glabrous. Pod linear, slightly longer than calyx, 5.0–5.5 × 1.5 mm, apex not rostrate, covered with glandular and eglandular hairs, fruiting calyx not accrescent; seeds 6–10, globose to sub-globose, 0.58– 0.65 × 0.65–0.7 mm, papillose, with bright brown testa. Conservation status:—Not threatened. Phenology:—Flowering season March–June. Habitat:—Alluvial soils, sandy and stony desert. Iconography:—Smith (1810), Jafri (1980) and Rechinger (1984). Chromosome number:—2n=30–60 (Montgomery et al. 1997, Runemark 2006). Distribution in EGYPT:—Native to north Egypt; distributed in three localities (Fig. 1): N, M and S (Täckholm 1956, Boulos 1999). Global distribution:—This species is centred in Mediterranean grasslands and coastal areas. It occurs from North to tropical Africa (Libya, Morocco, Tunisia, Algeria, Djibouti, Ethiopia and Kenya). It ranges from southern to central Europe (Greece, Crete, Sicily, Cyprus, France, Great Britain, Bulgaria, Albania, Balearic Islands, Corsica, Italy, Portugal, Malta, Sardinia and Spain) and occupies northern and northwestern Asia in Syria, Lebanon, Palestine, Jordan, Iran, Turkey, Iraq, Kuwait, Qatar, Saudi Arabia and Yemen (Greuter & Raus 1989).

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 17 Taxonomic notes:—Ononis reclinata subsp. mollis has often been confused with O. sicula due to their similar viscid appearance, small flowers and linear, pendent and multi-seeded pods. Although these taxa differ in their flower colour (the former has white to pink flowers, while the latter has yellow flowers), this character can easily be missed when the plants are dried and the flowers are thus discoloured. The two species can be differentiated by the following: O. reclinata has deltoid-cuneate leaflets, and its flowers are deflexed with ex-aristate peduncles, whereas O. sicula has narrowly oblong leaflets, its flowers are not deflexed, and its peduncles elongate to produce arista. Specimens examined:—EGYPT. (M): Sallum: East Sallum, Nazlt El Malfaya, sandy soil,13 April 1932, Shabetai J. R. z1786 (CAIM!); East Sallum, Saret Retama, sandy soil, 16 April 1934, Shabetai J. R. z3384, (CAIM!); 5 km east of the frontier, 24 May 1963, Täckholm V., Boulos L., Girgis W. A., Zahran M. A. & Elsayed I. s.n., (CAI!); Road between Matruh & Sallum, kilo 43, 03 April 1969, Abbas A. & El Shaer A. 2473, (CAIM!); (M): Mersa Matruh: Mersa Matruh, 22 April 1927, Drar M. s.n., (CAIM!); Wadi Habis, ca. 22 km west of Mersa Matruh, Umm El-Rakham village, sandy soil, 31º 23’ 11.7” N, 27º 02’ 29.3” E,30 April 2006, Boulos L., Hegazy A. &Sharashi O. 1103, (MAZHAR-image!); (M): El Alamein:El Hammam, Mariut, 27 February 1929, Shabetai J. R. z 822, (CAIM!); El Alamein, 30 March 1969, Abbas, A. & El Shaer A. 1483, (CAIM!); Sidi Abd-El Rahman, 31 March 1969, Abbas A. & El Shaer A. 1345, (CAIM!); (M): Burg El Arab: Mariut, March 1836, Lawerence A.s.n., (CAI!); Burg El Arab, 23 March 1961, Täckholm V. s.n., (CAI!); Burg El Arab, Spring 1961, Naguib M. I. s.n., (CAI!); Burg El Arab, 09 July 1972, Boulos L.& El Sayed I.s.n., (ASTU!); (M): Abusir: Abu Sir, 21 March 1918, Shabetai J. R.s.n., (CAIM!); (M): Alexandria: Sables maritimes, Sidi Gaber, 05 March 1908, Dumonh P. s.n., (CAI!);Aluk, short turf near the coast , 21 March 1926, Simpson N. D. 3902, (CAIM!); Abu Kir, 26 March 1927, Täckholm G. s.n., (CAI!); Abu Kir, 28 March 1957, Boulos L.s.n., (CAI!); (N):Wadi El Rokhama between Cairo & Suez, 15 April 1926, Täckholm G. s.n., (CAI!); Cairo-Suez road, near town, 14–23 April 1925, Simpson N. D. z3171, (CAIM!); (S): Northern Sinai: Wadi Gedeirat, 21 March 1918, Shabetai J. R.s.n., (CAIM!); Gebel Khashab at 280 m alt., 26 April 1922, Simpson N. D. 1192b, (CAIM!); Rafah, sandy soil, 14 April 1929, Shabetai J. R. z1008, (CAIM!); West El-Arish, near the town, 22 April 1959, Boulos L. s.n., (CAI!); Wadi El-Maghara, 23 April 1969, Boulos L.s.n., (CAI!). LIBYA. El-Gebl Al-Akhdar, 12 km west El Marj, 10 March 1968, Boulos L. 2043, (CAI!);El-Gebl Al-Akhdar, Bakur near Tokra, 10 March 1968, Boulos L. 2079, (CAI!);Between El Mersa & Ras El Hilal, 26 April 1968, Boulos L. 3092, (CAI!);Cyrenaica, 19 March 1969, Brydolf Ch. s.n., (CAI!); Below Shahhat, Cyrene,500–600 m alt., 26 March 1970, Boulos L., Khalifa & Sherif 4041, (CAI!); Umm Resum. Tamimi, sandi wadis, 30 March 1970, Boulos L. 4254, (CAI!). ALGERIA. Environs d’Orau: Pelouses sèches, 19 May 1918, Faure A. s.n., (CAI!). PALESTINE. Khan Yunis, near the railway station, 06 May 1955, Boulos L. 418, (CAI!). PORTUGAL. Baixo Alertojo maritimes, 02 March 1956, Masoto-Belia et al. 3298, (CAI!).

8. Ononis pubescens Linnaeus (1771: 267–268). Type:—EUROPE. in Europe australis, Gouan s.n., LINN 896.10, lectotype, designated by Förther in Turland & Jarvis (1997), (LINN- image!). Heterotypic synonyms: =Ononis calycina Lamarck (1785: 503). Type:—UNKNOWN. Trouvé dans son voyage en Barbarie, s.d., Desfontaines R.L. s.n., MNHN- P-P00296091, (P). = Ononis paniculata Cavanilles (1801: 69). Type:—EUROPE. Spain, May 1792, Broussonet 185, MA61630, (MA-image!). = Ononis pyramidalis Cavanilles (1801: 71). Type:—MOROCCO. Broussonet 186, MA61653, (MA-image!).

Description:—Annual herb, 10–35(–50) cm tall, branched; branches erect to ascending, viscid, covered with long patent simple eglandular hairs intermixed with unequal glandular hairs. Stipules green, not sheathing, 9.0–18.0 × 2.0–4.0 mm, ovate, with acute apex and entire margins with few teeth along margin, striate with 5 protruding nerves, as long as or longer than petiole. Lowermost and uppermost leaves simple, the rest trifoliate, flowering leaves trifoliate, 20–40 mm, petiolate; petiole 8.0–14.0 mm; leaflet elliptic to oblong, 10.0–22.0 × 2.0–9.0 mm, acute apex, cuneate base, dentate margins with 7–12 pairs of teeth; terminal leaflet subequal to lateral ones, approximately 1.0–1.25x, with ca. 5.0-mm-long petiolule. Terminal inflorescence, dense raceme, 3.5–75 mm long, ca. 5–20 flowered, pedunculate; peduncle 7.0–16.0 mm long, shorter than subtending leaf, not aristate. Flowers not pendent, pedicellate; pedicel 0.5–1.5 mm long. Calyx tubular, as long as or shorter than corolla, 10.0–18.0 × 1.5–3.0 mm, densely covered with glandular hairs intermixed with long setae; calyx lobes 7.0–14.0 mm, 5-nerved; lanceolate-oblong, with acute apex; calyx tube 3.0–4.0 mm, calyx lobes distinctly longer than tube, up to 4x. Corolla yellow with reddish-purple veins on standard and deep yellow notch on top of keel; standard orbicular to suborbicular, glabrous, (5.0–)10.5–16.0 × 4.5 mm, acute apex, entire margins, with short claw ca. 0.6 mm long; wings broadly oblong, lamina 6.5–10.0 (–12.0) × 1.5 mm, claw 0.5–0.7 mm long, auricle ca. 0.4 mm long, conjunctive tooth distinct, blunt; keel oblong, gently curved,

18 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. excavate above, lamina 6.0–9.0 × 1.5 mm, claw ca. 1.0 mm long; standard longer than the sub-equalling keel and wings. Stamens ca. 6.4 mm. Ovary 2.5 × 1.0 mm, covered with glandular hairs; style not deflexed, ca. 6.0 mm long, glandular at the base. Pod globose, slightly longer than calyx, 8.0–10 × 6.0–9.0 mm, apex rostrate, beak ca.1.0 mm, covered with short glandular hairs, fruiting calyx accrescent; seeds 2–5, globose, sub-globose to elliptic or reniform, 1.12–1.34 × 0.82–1.44 mm, loosely papillose, with brown to black testa. Conservation status:—Not threatened. Phenology:—Flowering season March. Habitat:—Inland deserts. Chromosome number:—2n=32 (Palazueles et al. 1976, Fernandes & Queiros 1978, Sanudo et al. 1979) Distribution in EGYPT:—The occurrence of this species in Egypt is doubtful; it was first recorded in Egypt by Raffeneau-Delile (1812: 69) on the border between Egypt and Syria “in arenis Aegypti et Syriaeconterminis, Savigny”. Consequently, all the subsequent floras recorded this taxon. Unfortunately, we could not find any Egyptian material for this taxon. However, in recent references, Boulos (1999) mentioned that the species existed in the Mediterranean and Sinai regions; then, Taia (2004a, b) cited one Egyptian specimen that was collected by Ayyad et al. in 1981 from Rafah (a city that lies on the border between Egypt and Gaza). Global distribution:—The plant is distributed along the Mediterranean coast in Europe (Crete, Greece, Cyprus, France, Portugal, Balearic Islands and Spain), along coastal zones in northwestern Africa (Algeria and Morocco) and in northwestern Asia (Turkey, Iraq, Jordan, Lebanon, Palestine and Syria), (Greuter & Raus 1989). Taxonomic notes:—O. pubescens is relatively similar to O. natrix, the former can be diagnosed by flowers not pendent nor aristate (vs. pendent aristate flowers in O. natrix); standard orbicular to suborbicular (vs. standard obovate); base of style hairy (vs. style glabrous) and pod globose (vs. linear). Specimens examined:—JORDON. 7 km east of Al-Hemma,13 May 1976, Täckholm V., El-Hadidi M. N., Lahham J. & Boulos L. 8907, (CAI!). ISREAL. Judaea: ad pagum Artufdit. Hierosolymae, 06 April 1909, Nàbélek F. 3013, (SAV-image!). Egyptian specimen:—(S): Northern Sinai: Rafah, 14 April 1981, Ayyad et al. s.n., (Alexandria University Herbarium). Note: the specimen was mentioned by Taia (2004a, b), unfortunately, the authors could not find this specimen.

9. Ononis sicula Gussone (1821: 78). Type:—SÝCÝLY. in rupibus volcanicis adpalagoniam, Gussone s.n. (?FI). = Ononis sicula Guss. f. microcarpa Bornmüller ex Širjaev (1932: 497). Type:—IRAN. Sinus Persicus australis, in apricis arenosis insulae Hormus, 05 February 1893, Bornmüller J. 270, HBG519341, isotype, (HBG-image!). Homotypic synonym: ≡Ononis viscosa subsp. sicula (Guss.) Huber-Morath (1970: 380). Heterotypic synonym: = Ononis leptocarpa Pau (1902: 29). Type:—SPAÝN.Valencia, Segorbe, 28 May 1910, Pau C. 1126, BM000751109, (BM-image!)

Description:—Annual herb, 5–20(–30) cm tall, branched; branches erect to ascending, viscid, covered with long patent simple eglandular hairs intermixed with unequal glandular hairs. Stipules green, not sheathing, 6.5–15.0 × 1.0– 2.0 mm, linear-lanceolate, with elongated subulate apex and entire margins, sometimes with few teeth along margin, striate with 4 protruding nerves, as long as or slightly longer than petiole. Lowermost and uppermost leaves simple, the rest trifoliate, flowering leaves simple, 10–25(–30) mm, petiolate; petiole 3.5–6.0 mm; leaflet narrowly oblong to narrowly elliptic, 6.5–16.0(–22.0) × 1.0–1.5(–6.5) mm, acute apex, cuneate base, dentate margins with 5–9 pairs of teeth; terminal leaflet distinctly larger than lateral ones, approximately 1.5–2.5x, with 1.0–3.5-mm-long petiolule. Flowers solitary, ±pendent, pedicellate; pedicel ca.1.5 mm long, pedunculate; peduncle 8.0–20.0 mm long, shorter than subtending leaf, elongated into arista; aristum 2.0–6.0 mm long. Calyx tubular, distinctly longer than corolla, 4.0–9.0 × 0.5 mm, densely covered with glandular hairs intermixed with long setae; calyx lobes 3.5–8.0 mm, 3-nerved, linear- lanceolate, with acute apex; calyx tube 0.5–1.5 mm, calyx lobes distinctly longer than tube, more than 5x. Corolla yellow; standard elliptic, glabrous, 5.5–6.0 × 2.5–3.5 mm, acute apex, entire margins with blunt claw ca. 1.0 mm long; wings oblong, lamina 3.0–4.0 × 1.5–2.0 mm, claw 1.5–2.0 mm long, auricle ca. 0.4 mm long, conjunctive tooth absent or indistinct; keel oblong, gently curved, not rostrate at apex, lamina 3.5–4.0 × 1.5–2.0 mm, claw 1.5–2.0 mm long; wings slightly shorter than the sub-equalling standard and keel. Stamens ca. 6.75 mm. Ovary 3.5 × 1.0 mm, covered with glandular hairs; style deflexed, ca. 2.2 mm long, glabrous. Pod linear, longer than calyx, 13.5–15.0 × 3.5–4.0 mm, apex shortly rostrate, beak ca.0.5 mm, covered with short glandular hairs, slightly inflated; fruiting calyx not accrescent; seeds 12–25, orbicular-reniform, 0.91 × 0.96 mm, tuberculate, with brown testa.

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 19 Conservation status:—Not threatened. Phenology:—Flowering season March–May. Habitat:—Coastal sand and stony deserts. Iconography:—Townsend (1974) and Rechinger (1984). Chromosome number:—2n = 32 (Palazueles et al. 1976, Díaz Lifante et al. 1992). Distribution in EGYPT:—Native to Egypt, distributed in two different ecosystems, namely, the Mediterranean coast and above the southern Sinai heights (Fig. 1): M and S (Boulos, 1999). Global distribution:—The species is centred in the Mediterranean and Mediterranean/Sahara regional transition zones in Europe (Sicily, Cyprus, Sardinia, Italy and Spain), in North Africa (Libya, Morocco, Algeria, Tunisia and Somalia), and in Asia (Iran, Iraq, Syria, Jordan, Lebanon, Palestine, Saudi Arabia, Bahrain and Qatar) (Greuter & Raus 1989). Taxonomic notes:—This species is diagnosed by the following characteristics: pendent flowers with peduncles elongating to produce arista; calyx densely covered with glandular hairs intermixed with long setae; corolla yellow with standard glabrous. O. sicula is very similar to O. viscosa subsp. breviflora, they can be differentiated by corolla characters. In the former, the corolla is obviously shorter than calyx and the standard elliptic with acute apex, where in the latter species, the corolla as long as calyx or slightly longer, standard obovate with retuse apex. Specimens examined:—EGYPT. (M): Alexandria: Mex, the Mediterranean Region, 17 March 1918, Brown T. W. s.n., (CAIM!); at a military camp south of Amria, 02 April 1961, Täckholm V. s.n., (CAI); (M): King Mariut: Champs Caleaur, 15 April 1908, Lwans 282, (CAI!); (S): Sinai, 1834, Bové N. s.n., V-815570, (UPS-image!); Sinai, loco Naherrek Arabiae petr. ad rupes, 09 May 1835, Schimper G.H.W. 136, HBG519343, (HBG-image!);(S): North Sinai: Mitla Pass, El Shatt, 10 May 1939, Shabetai J. R. 636, (CAIM!); (S): Southern Sinai:Rod El Ahmer, Wadi Rozneh, 12 May 1927, Kaiser A. 720, (CAIM!);Wadi Gebal region, 28° 19’ 13.8’’ N, 33° 32’ 35.52’’ E, 1895 m alt.,13 May 2004, Fayed A., El Garf I., Abdel-Khalik K. & Osman A. s.n., (ASTU!); Wadi Abu Markh, 27° 57’ 44.568’’ N, 34° 6’ 16.38’’ E,410 m alt., 12 April 2004, Fayed A., El Garf I., Abdel-Khalik K. & Osman A. s.n., (Herbarium of Loutfy Boulos-image!). JORDON.Azraq Oasis: vicinity of the hot spring, 14 May 1976, Täckholm V., El-Hadidi M. N., Lahham J. & Boulos L. 9033, (CAI!). ALGERIA. Oran, à Santa-Cruz. Pelousessèches, 03 May 1917, Faure A. s.n., (CAI!).

10. Ononis viscosa Linnaeus (1753: 718). Type:—FRANCE. Anonis lutea non viscosa lati folia minor flore pallido, in Barrellier, P1. Gall.: t. 1239, (1714), [Icon.], lectotype designated by Förther in Turland& Jarvis (1997); La Seyne, Jun 1890, Robin s.n., epitype designated by Förther in Turland & Jarvis (1997), (M); isoepitype, BM000751130, (BM-image!). Heterotypic synonym: = Ononis villosa Linnaeus (1759: 1160). Type:—SPAIN. Cadiz: in arenis et sylvaticis circa San Roque, Jun 1837, Boissier s.n., neotype designated by Förther in Turland & Jarvis (1997), (M). 10.a. Ononis viscosa L. subsp. breviflora (DC.) Nyman (1878: 161). Type:—GREECE. Crete: Monspelii et in Hispania, 08 September 1859, Sieber F.W. s.n., K000998598, K000998599, syntypes, (K- image!) Homotypic basionyms: ≡ Ononis breviflora De Candolle (1825:160). ≡ Ononis viscosavar. breviflora (DC.) Fiori (1900: 29).

Description:—Annual herb, 12–50 (–80) cm tall, branches erect to ascending, covered with long patent simple eglandular hairs intermixed with unequal glandular hairs. Stipules green, not sheathing, (2.0–) 5.0–9.0 × 0.5–2.5 mm, oblong-deltoid, with acute apex and entire margins, striate with 4 protruding nerves, shorter than petiole. Lowermost and uppermost leaves simple, the rest trifoliate, flowering leaves simple, 15–28 mm, petiolate; petiole 4.0–8.5 mm; leaflet widely elliptic, 9.0–17.5 × 3.5–10 mm, acute apex, cuneate base, dentate margins with 5–9 pairs of teeth; terminal leaflet sub-equalling lateral ones or slightly larger, with ca.2.0 mm-long petiolule. Flowers solitary, ±pendent, pedicellate; pedicel 1.5–2.0 mm long, pedunculate; peduncle 10.0–20.0 mm long, shorter than subtending leaf, elongated into arista; aristum 3.5–7.0 mm long. Calyx tubular, shorter than or as long as corolla, 5.0–7.5 × 1.3–1.5 mm, densely covered with glandular hairs intermixed with long setae; calyx lobes 4.0–6.0 mm, 3–nerved, linear-lanceolate, with acute apex; calyx tube 1.0–3.0 mm, calyx lobes longer than tube, approximately 2-fold. Corolla yellow with purple veins on standard; standard obovate, glabrous, 6.2–8.0 × 5.5–6.0 mm, retuse apex, entire margins, with blunt claw 1.2–1.5 mm long; wings oblong, lamina 4.0–5.0 × 1.5–2.0 mm, claw 1.5–2.0 mm long, auricle ca. 0.75 mm long,

20 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. conjunctive tooth indistinct or absent; keel oblong, gently curved, lamina 3.5–5.0 × 1.5 mm, claw 1.5–3.0 mm long; wings slightly shorter than the sub-equalling standard and keel. Stamens 0.5 mm. Ovary 4.0 × 1.0 mm, covered with glandular hairs; style deflexed, ca. 1.5 mm long, glabrous. Fruiting calyx not accrescent. Pod longer than calyx, linear, 11.0–13.0 × 3.5–4.0 mm, apex shortly rostrate, beak0.5 mm, covered with short glandular hairs, slightly inflated, 7–16 seeded; seeds oblong-reniform, 1.2 × 0.72 mm, papillose-tuberculate, with brown-black testa. Conservation status:—Not threatened. Phenology:–—Flowering season March–June. Habitat:—Coastal sand and stony deserts. Iconography:—Jafri (1980), Rechinger (1984) and Townsend (1974). Chromosome number:—2n = 32 (Fernandes et al. 1977, Fernandes & Queiros 1978, Colombo et al. 1980, Runemark 2006). Distribution in EGYPT:—South Sinai, co-existing with Ononis sicula. Global distribution:—The Mediterranean is the centre of the distribution of this species, from where it extends northwards to central Europe (Cyprus, Balearic Islands, Greece; Sicilia, Italy, Albania, Spain, France and Portugal) and southwards to western Asia (Turkey, Iran, Iraq Lebanon, Israel, Jordan and Syria) and the North African coasts in Libya, Morocco, Algeria and Tunisia (Greuter & Raus 1989).

FIGURE 5. Ononis viscosa subsp. breviflora, EGYPT. Southern Sinai: Wadi Gebal region, Al-Sheq: 28.3226 N, 33.5623 E, 1940 m Alt., Fayed et al. s.n., (ASTU!).

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 21 Taxonomic notes:—Ononis viscosa s.l. is a taxonomic complex that needs to be intrinsically discussed, segregated and disbanded. Álvarez & Abreu (1982) used the following diagnostic characters to discriminate six different infraspecific taxa of O. viscosa: the length of the corolla in relation to the length of the calyx, colour of the flower, size of the legume and its length in relation to calyx length, trifoliate or simple cauline leaves, leaflet apex and shape of the blade. Huber-Morath in Davis (1970: 380) treated Ononis sicula as a subspecies of O. viscosa. This opinion was based on the characters shared between the two closely related taxa (O. sicula and O. viscosa); they both have yellow solitary flowers borne on pendent aristate peduncles. Here, we discriminate between O. sicula and O. viscosa subsp. breviflora (Fig. 5 & 6) by the length of the corolla in relation to that of the calyx, shape and apex of the standard, leaf blade, leaf margin and shape of the stipules. In the former species, the corolla is much shorter than the calyx, the standard is elliptic with an obtuse apex, and the leaves are narrowly elliptic (width 1.0–1.5 mm) with ca. 5–9 pairs of teeth, while in thelatter species, the corolla equals the calyx or is slightly longer, the standard is obovate with a retuse apex (Fig. 7), and the leaves are widely elliptic (width up to 3.5–10.0 mm) with ca. 2–5 pairs of teeth on the margin of the median leaflets. Ononis sicula has linear to linear-lanceolate stipules with a long subulate apex, whereas O. viscosa subsp. breviflora has deltoid stipules with acute apices (Fig. 8).

FIGURE 6. Morphological features of Ononis viscosa subsp. breviflora. (a) habit, scale bar = 1 cm, (b) flower, (c) glandular hairs on aristum, (d) fruit (left) & leaflet (right), (e) close-up photo of the calyx, scale bar for (b–e) = 2 mm. The photographed voucher:—EGYPT: Southern Sinai. Wadi Gebal region, Al-Sheq, 28.3226 N,33.5623 E,1940 m Alt., 13 May 2004, Fayed A., El Garf I., Abdel-Khalik K. & Osman A. s.n. (ASTU!).

Specimens examined:—EGYPT. (S): Southern Sinai:Wadi Gebal region: Al-Sheq, 28.3226 N, 33.5623 E,1940 m Alt.,13 May 2004, Fayed A., El Garf I., Abdel-Khalik K. & Osman A. s.n., (ASTU!). LIBYA. Eastern Mediterranean coastal region: El-Gebel Al-Akhdar, Al-Marj to Al-Baida motorway, sand loamy to clay soil, 32º 28’ 12.7” N, 21º 03’ 36.7” E, 537m Alt., 29 April 2007, Hegazy A. & Sharashi O. 1960, (MAZHAR-image!). JORDON. Balqa Governorate: Tell Ar-Rumman, Royal Botanic Garden Sector No. 23, 10 April 2011, Shishani M. & Alkoran M.s.n., (The National Herbarium of Jordon-image!). ISRAEL. Lower Galilee: betw. Shefaram and Nazereth near Tsipori springs, 24 April 1963, Plitmann U. 224441, E00334367, (E-image!); Jerusalem: Talpioth, 20 April 1931, Amdursky I. 135, E00334385, (E-image!). IRAQ.Kurdistaniae Turcicae District Berwari: ad pagum Aradendit. Amadia, in declivitatibussiccis, 900 m alt., 14 June 1910, Nábělek F. 3021, SAV0000199, (SAV-image!).TURKEY. Kemalpasa: Karabel, near the bas-relief of Karabel (Eti-Baba), 25 May 1966, Alava, R. O. 4753, E00334379, (E-image!).

22 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. FIGURE 7. Flower morphological features of Ononis species: (6) Ononis vaginalis, (7) O. reclinata subsp. mollis, (8) O. pubescens, (9) O.sicula, (10) O. viscosa subsp. breviflora: (a) calices, (b) standards, (c) wings, (d) keels, (e) androcia and (g) gynoecia, scale bar = 5 mm.

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 23 FIGURE 8. Stipules outline: (a) Ononis diffusa, (b) O. serrata, (c) O. mitissima, (d) O. variegata, (e) O. natrix, (f) O. vaginalis, (g) O. pubescens, (h) O. sicula, (i) O. reclinata subsp. mollis, (j) O. viscosa subsp. breviflora, scale bar = 4 mm.

Acknowledgments

The authors are grateful to Dr. Mats Hjertson (UPS); Dr. Laurent Gautier (G); Dr. Robert Vogt (BGBM) and Dr. Katharina Rabe (Freie Universität Berlin) for providing high-quality digital photographs of Egyptian material kept in their herbaria. We would like to express our sincere gratitude to Dr. Olof Ryding (C) for his valuable consultation in lectotypification.

References

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28 • Phytotaxa 408 (1) © 2019 Magnolia Press FAYED ET AL. Appendix 1. Voucher specimens used for drawings. Taxon voucher caption Ononis diffusa EGYPT. North Sinai: Rafah, April 1921, &Figure 2 (1)Figure 8 (a). Hefnawy M. T. s.n., (CAIM!). Ononis serrata EGYPT. El-Garawla, 15 km before Mersa Figure 2 (2) & Figure 8 (b). Matruh, 31º 15’ 34” N, 27º 22’ 12” E, Alt. 11m, 07 April 2015, Faried A., Olwey A. & Hassan M. s.n. (ASTU!). Ononis mitissima EGYPT. Cairo: Shubra, on wayside of fields, 10 Figure 2 (3) & Figure 8 (c). April 1940, Shabetai J. R. z6248 (CAIM!). Ononis variegata ALGERIA. La Macta: pré de Mostaganem Figure 2 (4) & Figure 8 (d). sables maritimes, 07 May 1916, Faure A. s.n., (CAI!). Ononis natrix subsp. natrix LIBYA.8 km south of Al-Qubbah, 22 January Figure 2 (5) & Figure 8 (e). 1967, Boulos L. 1326, (CAI!). Ononis vaginalis EGYPT. Mersa Matruh: Al Kasr road, 31º 21’ Figure 7 (6) & Figure 8 (f). 49” N,27º 7’ 55” E,3 m alt.,07 April 2015, Faried A., Olwey A. & Hassan M. s.n., (ASTU!). Ononis reclinata subsp. mollis EGYPT. East Sallum: Saret Retama, sandy soil, Figure 7 (7) & Figure 8 (i) 16 April 1934, Shabetai J. R. z3384, (CAIM!). Ononis pubescens JORDON. 7 km east of Al-Hemma,13 May Figure 7 (8) & Figure 8 (g) 1976, Täckholm V., El-Hadidi M. N., Lahham J. & Boulos L. 8907, (CAI!). Ononis sicula EGYPT. King Mariut: Champs Caleaur, 15 April Figure 7 (9) & Figure 8 (h) 1908, Lwans 282, (CAI!). Ononis viscosa subsp. EGYPT. Southern Sinai:Wadi Gebal region: Al- Figure 7 (10) & Figure 8 (j) breviflora Sheq, 28.3226 N,33.5623 E,1940 m Alt.,13 May 2004, Fayed A., El Garf I., Abdel-Khalik K. & Osman A. s.n., (ASTU!).

TAXONOMIC REVISION OF THE GENUS ONONIS Phytotaxa 408 (1) © 2019 Magnolia Press • 29