255 Taxonomic Revision of Astragalus L. sect. Hymenostegis Bunge (Leguminosae) S. ZARREM. &D. PODLECH Abstract: Zarre M., S. & PODLECH, D.: Taxonomic revision of Astragalus L. sect. Hymenostegis Bunge (Leguminosae). - Sendtnera 3: 255-312. 1996. - ISSN 0944-0178. The present work comprises the revision ofAstragalus L. sect. Hymenostegis Bunge. 22 species are recognized. Detailed descriptions, complete lists ofsyn- onyms and distribution maps are given for all taxa. One species and one sub- species are newly described. The sect. Hymenocoleus Bunge is reduced to the rank of subsection. Differences between the section and the closely related sections are discussed. Zusammenfassung: Eine Revision vonAstragalus L. sect. Hymenostegis wird vorgelegt. Ausführ- liche Beschreibungen, vollständige Synonymic und Verbreitungskarten aller Sippen werden angegeben, eine Art und eine Unterart neu beschrieben. Die Sektion Hymenocoleus wird als Subsektion von Hymenostegis geführt, die Unterschiede zuverwandten Sektionendiskutiert. Introduction Astragalus sect. Hymenostegis was placed by BUNGE (1868/69) in the artificial subgen. Calycophysa, together with its closest relatives, i.e. sect. Megalocystis Bunge, which was reviewed recently, see TIETZ & ZARRE (1994) and sect. Acidodes Bunge, for which no modem revision is available. The entire section was revised by RECHINGER et al. (1958) and later for certain areas e.g. the flora of Turkey (CHAMBERLAIN & MATTHEWS 1970), the flora of URSS (GONTSCHAROV 1946) and flora of Iraq (TOWNSEND & GUEST 1974). However, because of the great variability in many species of the section, none of the available keys are useful for naming the species. The extensive variability present caused the taxonomists to describe a lot ofnew species in this relatively small section. The goal of this work was to provide an exact taxonomic definition for each taxon in the section, to recognize the limits of morphological diversity for each taxon and to prepare an useful diagnostic key for all taxa. The present work is mainly based on the study of herbarium material kindly provided by the following herbaria (abbreviated according HOLMGREN et al. 1990): B, BG, BRNM, C, E, G, G-BOIS,JE, K, L, M, MSB, P, PR,TARI,TUH, W, WUandZT. 256 Some additional field studies were carried out during an excursion to Iran (by S. ZARRE). We want to thank the directors of the above mentioned herbaria for sending the material. We are grateful to Mr. CHEHREGANI from Bou-Ali University of Hamadan (Iran) for his support, especially in organizing the excursion in Iran. This work is partly supported by the Ministry of Culture and Higher Education of Islamic Republic Iran. Position ofthe sect. Hymenostegis within the genus Within the subgen. Astragalus sect. Hymenostegis belongs to the group of thorny Astragali (with exception of A. vaginans, whose leaves are imparipinnate) with a calyx inflated in fruit and unilocular fruits, which remain enclosed by the calyx up to maturity. The most conspicuous characters of the section are the ± large bracts and panduriform standards. Since some species of the closely related and very hetero- geneous sect. Megalocystis have similarly ± large bracts and panduriform standard, there are some difficulties in separating the sections. However, the flowers in sect. Megalocystis are pedicellate in opposite to sect. Hymenostegis with sessile flowers. Moreover most of the species with such a standard in the former section have black hairs beside the white ones on the calyx, but in sect. Hymenostegis black hairs are always absent. For an exact view on the limits between these two sections see the diagnostic key. Diagnostic key to the closely related sections 1 Short lateral shoots usually present at the base of each leaf on the main stem; inflorescences born on the short lateral shoots, only in A. pachyrhachis short lateral shoots sometimes not clearly developed sect. Poterion - Short lateral shoots absent; inflorescences arising from the main stem 2 mm 2 Standard at least 15 long, if shorter, then the limb hastate or auriculate at base 3 - Standard up to 13(-14) mm long, limb rounded or somewhat angulately passing into the claw 5 3 Limb of standard rounded at the base sect. Megalocystis - Limb ofstandard hastate-auriculate at base 4 4 Flowers pedicellate; black hairs on inflorescence present, ifabsent (inA. ebenoi- des and A. szovitsii), then fruit laterally compressed and two bracteoles on the base ofeach calyx present sect. Megalocystis - Flowers sessile; black hairs absent; fruit dorsi-ventrally compressed; bracteoles absent orrarely present at the base ofsome calyces and mostly not in pairs sect. Hymenostegis 5 Fruiting calyx only slightly larger than flowering one, ± campanulate to globose, mm mm not rupturing, 3-4,5 wide, the tube 3-5 long, only inA. argyrostachyus mm clearly enlarging, 6-9 wide, campanulate; calyx teeth as long as tube to dis- mm tinctly longer than it; longerhairs on the calyx 2-A sect. Campylanthus - Fruiting calyx bladdery inflated, contracted at the teeth, only in A. diopogon mm rarely rupturing, (4)5-10 wide, the tube 5-15 rmn long; calyx teeth distinctly mm shorter than the tube; calyx hairs up to 2 long 6 6 Inflorescence longer than the leaves, if shorter, then densely flowered; fruit laterally compressed sect. Microphysa 257 - Inflorescence shorter than or as long as the leaves, remotely few-flowered (with 2-6, rarely up to 13 flowers); fruit dorsi-ventrally compressed sect. Megalocystis (A. diopogon andA. eriostomus) Valuation of taxonomic characters Many characters, which are normally important for the delimitation ofneighboring sections, are not significant in sect. Hymenostegis, either because of homogeneity of the section, or because they are influenced strongly by ecological conditions. Rela- tively short flowering and fruiting period is another problem in naming the species. Our key is primarily useful when the flowers are still adherent to the inflorescence. Importance, variability and applicability ofthe characters are subsequently discussed: Life forms, branching mode and habit Except forA. vaginans, all species ofthe sec- : tion are thorny cushions-forming subshrubs of alpine habitats. All are arising from a woody underground caudex. The stems are mostly branched from the base. In response to different ecological conditions the branching pattern can vary from remote to dense, even in the same species. Therefore none of these characters are of taxonomical importance. Indument The hairs are basifix (stable character of the subgen. Astragalus) and : mostly pure white. A. recognitus and A. hymenocystis subsp. hymenocystis are the only taxa of the section with brownish-yellow hairs. Length of the hairs (especially on the calyx), their density and form are often very important characters. Stem Only A. vaginans has a stem with relatively long internodes, which is almost : glabrous. Other species have stems with long internodes, which are hairy but become glabrescent later on. Stipules They have mostly the same texture as the bracts and can be useful for sep- : arating the species. Stipules are in many species long adnate to the petioles and otherwise connate. They become mostly glabrescent. The shape shows httle variabil- ity and is therefore of no taxonomic value. The size of the stipules is very important and makes some species easily separable. In some species such as A. chrysostachys venation of the stipules shows only one main nerve at free portion, but the number of the nerves may increase under different ecological conditions. Because of this variability in some species, the stipule venation is ofno taxonomic importance. Leaves As mentioned above with the exception of A. vaginans all species of the : section possess paripinnate leaves. In some species the length of the leaves is very variable, but in some cases it can be used as a distinctive character. This is also true for the orientation of the rachides: In some species such as A. persicus both forms of rachides, straight and recurved, can be seen. The relative length petiole to the whole rachid is more or less equal in all of the species. End-thorn in subsect. Hymenostegis is mostly shorter than the uppermost leaflets, with some exceptions in A. hirticalyx. The number of leaflet-pairs is not very variable, and is ofno taxonomical importance. A. laguriformis with 1-4 pairs of leaflets is the only species which can be charac- terized by this feature. Leaflet size, shape and indumentum are sometimes very good characters. Inflorescence Although some species show variability, the relative size ofthe inflor- : escence to the leaves is mostly of taxonomic significance. Its size and shape is im- portant too. But in some withered specimens it is not easy to recognize, because the flowers fall offquickly. In this case the distance between flower traces can be used to determine whether the inflorescence is lax, which is very important for distinguishing some closely related species (for example A. uraniolimneus from A. lagopodioides). 258 The indumentum on peduncles is another important character. It can be villose or composed ofstraight, ± appressed hairs. Bracts: Texture, size, tip, indumentum and colour of the bracts are the most decisive characters in the section. Very thin and hyahne bracts characterize for example A. chrysostachys. Acutely tipped bracts ofA. kohrudicus make it easily distinguishable from the related A. glumaceus. The bracts in most of the species are glabrous inside, but in some others such as A. hymenostegis, A. lagopoides, A. persicus (not always) and A. tahrizianus they are hairy inside, especially at tip. Moreover the venation of the bracts ofA. hymenostegis differs in one aspect from that of other species: The ends of the nerves are connected reticulately instead of ending parallely. However, this charachter is not easy to observe. Bracteoles They are rarely developed in some species and of no taxonomic im- : portance. Pedicel All the species have nearly sessile flowers. : Calyx: In spite of its very thin texture the calyx remains unruptured during its matu- ration in all species. With exceptions ofA. paralurges and A. sciureus the calyx is bladdery inflated immediately after anthesis. The shape ofcalyx is at first tubular and after inflation globose or elliptic. Calyx is parallely nerved, and the number of the nerves changes in a narrow range. The calyx indumentum and form of the teeth are almost constant in all species. However the size ofthe calyx, colour ofthe nerves and proportion ofthe tooth length to the tube are sometimes distinguishing characters. Corolla: Almost all species have standards with yellowish white claws. But the limb colour is a good character to distinguish for example A. chrysostachys andA. recog- nitus from other closely related species. They have always yellowish white limbs, whereas other species may have pink, red, purple or violet limbs. As the colour ofthe corolla after collecting changes determination of the colour is sometimes very diffi- cult. In most species the claws ofthe wings and keel may be adnate up to 1 mm to the staminal tube, but in A. sciureus they are nearly free. This character is difficult to measure and the differences are not large enough to be ofuse in the key. Standard: the standard is homogeneously panduriform in the whole section. But its size and the proportion oflimb to claw is sometimes taxonomically useful. Wings: The size ofthe limbs is their most important character. Some species, such as A. hymenocystis, A. nervistipulus, A. strausii andA. uraniolimneus have wings with a conspicuously large limb, although there are usually no sharp limits between species with respect to this character. In the isolated A.glumaceous group the auricles of the wings are larger than in other species ofthe section. Keel: It is always distinctly shorter than the wings. The limb size is a very good char- acter to separate the A.glumaceous group from other species of the section. Further- more the limb outline is because of the variability in some taxa of no taxonomical importance. The auricle is tiny in all species. Stamens and ovary Except for the length of the segment, where the stamens are free : from each other, none ofthe characters ofthese two organs are taxonomically impor- tant. Fruit Their characters are oflimited use in the key: Most ofthe herbarium specimens : are without ripe fruits, because as soon as they are ripe, flowers with fruits fall offin many species, and the specimens in this state have nearly no value for collecting. The fruits are always dorsi-ventrally compressed. Their form and indumentum do not pro- vide distinctive characters, as they are very homogenous in the whole section. However the size of them may be used for separating two closely related species A. chrysostachys andA. recognitus. Seeds: None oftheir characters are used here, firstly because they are mostly not ripe on herbarium sheets, and secondly the section is very homogeneous in their charac- ters. 259 Grouping in the section The section is divided in two subsections in this work: subsect. Hymenocoleus and subsect. Hymenostegis. The former differs from the latter mainly in having impari- pinnate leaves. But the structure of inflorescence and flowers in both subsections is identical. Therefore we have preferred to reduce the monotypic sect. Hymenocoleus to the rank of a subsection. Subsect. Hymenostegis is very homogeneous and can be scarcely grouped into natural units. In respect to similarity in some important characters we have recognized the following groups in the subsection. However there are no sharp morphological limits between the groups and the characters overlap in many cases. Therefore no formal rank is considered for the groups. Group 1: A. ghimaceous and A. kohrudiciis: It is a somewhat isolated and easily de- terminable group within the subsection. It is characterized by very broad inflor- escence, large bracts and flowers. Group 2 A. chehreganii and A. strausii: Relatively broad inflorescence and ± large : standards are the features, which connect this group with the preceding, but long peduncle and short hairs on the calyx make it easily separable from group 1. Group 3 A. chrysostachys, A. hirticalyx, A. hymenostegis, A. lagopoides, A. lagiiri- : formis, A.nerxistipulus, A. pediculariformis, A. persicus, A. recognitus, A. tabrizi- anus andA. velenovskyi: This is the central group in the subsection. Most species are very variable, and some forms are so different from the typical form, that it is difficult to believe they belong to a same species, but mostly there is a continuous range of variability. Separating the species can be difficult because some oftheir more extreme morphs can approach neighboring species. For example short-pedunculated forms of A. persicus might be mistaken forA. hirticalyx, large bracteate forms ofthe former for A. lagopoides and so on. Because of this problem it was very difficult to prepare a diagnostic key for the group and for example A. persicus is cited many times in the key. Group 4 A. lagopodioides, A. paralurges, A. rubrostriatus and A. sciureus: Lax in- : florescences with remote flowers are the conspicuous characters of the group. More- over, A. paralurges and A. sciureus have a calyx which becomes not distinctly inflated with age. A. sciureus possesses the longest inflorescence in the subsection. The group can not be differentiated exactly from group 3, because A. sciureus some- times has a dense inflorescence like some forms ofA. persicus with long cylindrical inflorescences. However, inA. persicus the calyx inflates itselfsoon after anthesis. Group 5: A. hymenocystis andA. uraniolimneus: It is very near to the formergroup in nearly all respects but has dense inflorescences. Geographical distribution and ecology Almost all members ofthe sect. Hymenostegis are Irano-Turanian elements and are common in mountain regions ofthe Iranian highland (see map 1). Iran is the center of diversity ofthe section,with 17 species ofwhich 11 are endemic in the region.Turkey with 7 species and 3 endemics, Azerbeidzhan and Armenia each with 3 species and Iraq with 2 species are other countries in which the section is native. The widest ranging species in the section are A. chrysostachys, A. glumaceous andA. persicus. Local endemism plays a very important role in the section. Some of the species with such a distribution pattern are: A. chehreganii (from Guoushchi mountains N of NW Oroumieh lake, Iran), A. hymenocystis and A. hymenosteWgis (same area as the former), A. pediculariformis (near Sultanieh in Prov. Zanjan, Iran) and A. laguri- . 260 formis (from Iraqish-Turkish border, Prov. Kordestan). It is ofinterest, that three ofthe local endemics can be found in the montain region next to the northern part of the Uroumieh lake. Apart from the species named above there are 7 more taxa of this W section in the Prov. Azarbaijan: A. chrysostachys, A. glumaceous, A. hirticalyx, A. lagopoides, A. persicus, A. tahrizianus and A. uranioUmneus The species ofthe sect. Hymenostegis prefer to live in higher areas between (400-) 800-3000 m. Most of them are cushion-forming plants of alpine habitats, which can be found in steppes with Astracantha, Artemisia, Cousinia, Thymus and others. Similar to other thorny Astragali, the species of the sect. Hymenostegis are adapted to dry and windy condition. Flowering and fruiting of the section occur in the months (May-)June-August. Taxonomic enumeration Astragalus L. sect. Hymenostegis Bunge, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 57. 1868. Lectotype (Podlech 1990): A. hymenostegis Bunge. Plants perennial, mostly cushion forming dwarf shrublets, 10-60 cm in diameter, 10-70 cm high, densely or loosely branched from the base. Hairs basifix, white or mm yellow, 0.1-6 long, with a sharp tip, the longest ones mostly on peduncle and on the calyx, the longer ones mostly somewhat thicker as the remainder. Caudex light brown to grey or black, prostrate, 0.5-3 cm thick. Stems prostrate or ascending, up to 20 cm long, 1-5 mm in diameter, hairy or glabrous, below the stipules always densely hairy, glabrescent. Stipules membranaceous orchartaceous, yellow, with 1-13 parallel mm nerves in free portion arising from a loose reticulate net at the base, 8-30 long, up to lower half adnate to petiole, otherwise connate, free portions triangular-lanceolate, acute or acuminate, younger ones appressedly hairy, later on glabrescent, sometimes ciliate. Leaves 1-20 cm long; rachides remote or dense, thin or thick, mostly rigid, with appressed or spreading hairs; petiole 1/7-1/2 the length of the rachid; end-thorn mostly present l/10-l/l(-3 times) the length ofthe uppermost leaflets; terminal leaflet mm mm only present inA. vaginans. Leaflets in 1-13 pairs, 3-30 long and 1-12 wide, flattened or complicate, narrowly oblong-elliptic to elliptic or oblong, rarely obovate, obtuse or acute, mostly mucronate, both sides densely or sparsely appressed hairy, rarely spreadingly hairy, sometimes glabrescent, rarely glabrous. Inflorescence shorter than or overtopping the leaves; flowering part 2-25 cm long and 2-4.5 cm wide, globose to long cylindrical, sometimes lax and spicate; peduncle 0.5-30 cm long, shorter or longer than the leaves, densely appressed hairy to villose. Bracts thinly mm membranaceous to glumaceous, yellow, sometimes purple at tip, 8-27 long and mm 2-12 wide, broadly ovate at the base ofinflorescence to elliptic-lanceolate further up, acute or acuminate at tip, densely or sparsely appressed hairy on midrib and apex, becoming glabrous, sometimes glabrous from the beginning, ciliate, inside of the bracts mostmlmy glabrous, sometimemsmhairy especially at tip. Bracteoles rarely present, 1 or 2, 4-7 long and 0.5-1.5 wide, linear or subulate, mostly villose. Flowers mm mm sessile. Calyx yellow, sometimes purple at tip, 10-28 long and 3-16 wide, at first tubular, mostly inflating after anthesis, with 15-30 parallel nerves, which ana- stomose towards the teeth, densely appressed pilose, hairs initially straight, then mm crispate, irregularly villose or tomentose; teeth4-12 long, from a shortly triangular base subulate to filiform. Corollawith white oryellow claws, limbs the same colour as the claw, orpink to purple, sometimes blue to violet towards margins. Standard 13-31 mm long; limb panduriform, hastate at base; claw cuneate. Wings slightly shorter than the standard; limb oblong, obtuse; claw somewhat longer than limb. Keel distinctly shorter than wings; limbs 5-6 mm long and 2-3 mm deep, obovate-elliptic, triangular 261 or rarely oblong, obtuse or minutely mucronulate, minutely auriculate at base; claw mm longer than the limb. Stamens as long as the keel, at upper 2.5-5 free from each other. Ovary 2-8 ovulate, densely appressed hairy; style hairy up to lower half. Fruit mm dorsi-ventrally compressed, 4-11 long, narrowly elliptic to elliptic orrarely ovate, ventral side straight, dorsal side curved, valves slightly expanded, densely appressed pilose; beak 0.6-2.5 mm long, straight, mostly hooked at tip. Seeds always single, 3- mm mm 4.5 long and 2-3 wide, elUptic-reniform. Key to the subsections ofAstragalus sect. Hymenostegis 1 Stems elongated, intemodes 1-1.5 cm long; leaves remote, imparipinnate; leaflets glabrous on adaxial surface A. subsect. Hymenocoleus - Stems short, intemodes up to 0.7 cm long; leaves dense, paripinnate; leaflets hairy on adaxial surface A. subsect. Hymenostegis Astragalus L. subsect. Hymenocoleus (Bunge) Podlech & Zarre, comb. nov. = Astragalus L. sect. Hymenocoleus Bunge, Mem. Acad. Imp. Sei. Saint Peters- bourg 11(16): 57. 1868. Type (monotypic): Astragalus vaginans DC. Leaves imparipiimate. Otherwise see description ofthe species. \. Astragalus vaginans DC, Astragal.: 210, t. 37. 1802 = Tragacantha vaginans (DC.) Kuntze, Revis. Gen. 2: 949. 1891. Lectotype (here designated): 'A. orientahs flore ochroleuco' Toumefort{P: Hb.VAILLANT!). , Figures: DE CANDOLLE, Astragalogia: tab. 37. 1802. Plants c. 30 cm high, up to 40 cm including inflorescence. Hairs 0.1-2 mm, except those on calyx mostly straight, thick. Stems ascending, up to 23 cm long, growing 2- 20 cm per year, in first year 2-3.5 mm in diameter, below stipules glabrous or sparsely pilose. Stipules thinly membranaceous, greenish to yellowish white, with 8-13 parallel nerves at free portion, 10-30 mm long, at a length of4-15 mm adnate to the petiole, mm otherwise 2-14 connate, triangular-lanceolate, acuminate, glabrous or ciliate. Leaves imparipinnate, with 5-13 pairs of leaflets, 3-11 cm long; rachides remote, ± thin, flexible, straight or curved, obliquely erect or rarely deflexed, sparsely appressed hairy; petiole 1/7-1/5 the length ofthe rachid; terminal leaflet ± as long as the leaflets mm of the next pair; leaflets light green, remote or rarely dense, 10-27 long and 2-6 mm mm wide, flattened, narrowly elliptic to elliptic, acute, with a mucro up to 1 long, upper surface glabrous, lower surface sparsely appressed hairy. Inflorescence dis- tinctly overtopping the leaves, dense, 3.5-5 cm long and c. 4 cm in diameter, globose to ovate; peduncle 10-15 cm long, longer than the leaves, densely covered with short and long appressed hairs, longer hairs thicker than the reaminder. Bracts chartaceous mm sometime hyaline at margins, yellowish white, younger ones purplish at tip, 12-17 mm long and 4-6 wide, ovate-elliptic, long acuminate, glabrous, ciliate, rarely sparsely mm appressed pilose on midrib. Calyx yellowish white with purple teeth, 13-17 long and 4-6 mm wide, at first tubular later on slightly elliptically inflating, with 22-30 parallel nerves, densely long appressed hairy becoming ±densely tomentose; teeth 4- mm 6 long. Corolla limb purple; claws of wings and keel at the base adnate to the mm mm mm staminal tube. Standard 18-22 long; limb 12-14 long and 6-8.5 wide, elliptic or oblong-panduriform, rounded or shallowly retuse at the apex, obtusely 262 hastate-auriculate at base; claw 5-8 mm long. Wings 17.5-21 mm long; limbs 8.5-9.2 mm long and 2.5-3 mm wide, oblong, obtuse or rarely minutely mucronulate at tip; auricle 0.2-0.4 mm long; claw 9-12 mm long. Keel 15.5-17 mm long; limbs ca. 6 mm long and 3 mm deep, obovate-elliptic, with broadly curved lower edge and ± straight mm upper edge, obtuse, rarely minutely mucronulate; claw 9.5-1 1 long. Stamens at mm the upper 3 free from each other. Fruit immature. Seeds immature. Floweringand fruiting time: VI-VII(-VIII). Occurence: Loose forests (for example of Piniis hrutia), mostly in somewhat humid climat. Distribution: Turkey (AnatoUa). Map 10. Specimens seen: Turkey. Prov. Adana Amanus-Gebirge (Nurdaglari), ca. 3 km S oberhalb Yarpuz (29 km : E Osmaniye), 1200 m, 13.7.1978, Ehrendofer et al. 787-38 (MSB) - Cappadocia, Hadjin, Kala Sekisi, 27.6.1893, Förster 912 (M). - Prov. Maras: 78 km N Maras, zwischen Maras und Göksun, 1280 m, 7.7.1981, Nydegger 16713 (M, MSB) - Maras-Göksun, 60 km S Göksun, 540 m, 1.6.1987, Nydegger 42588 (MSB) - Bulghar Maaden, Al Chodscha, 1300 m, 1896, Siehe 594 (P). A. vaginans is the only species of the subsect. Hymenocoleus. Except for impari- pinnate leaves and long internodes there is no difference between the two subsec- tions. Here is the same situation as between sect. Anthylloidei DC. and sect. Megalocystis Bunge. As it was mentioned (TIETZ & ZARRE 1994), the end thorn can be absent in some forms of the species of sect. Megalocyctis, namely A. ebenoides andA. raddei, thus this character is bad to use for separating two sections. Within the sect. Hymenostegis A. vaginans is comparable with A.chehregani and A. strausii, because of the very dense and globose to shortly cylindrical inflorescence and the short hairs on the calyx. In hb. DECANDOLLE (G-DC) there is no original material ofA. vaginans, therefore the specimen of hb. VAILLANT in Paris, which surely has been seen by DE CANDOLLE, is selected as lectotype. Astragalus L. sect. Hymenostegis Bunge subsect. Hymenostegis Leaves always paripinnate. Otherwise see description ofthe section. Key to the species and subspecies ofsubsect. Hymenostegis 1 Inflorescence lax, the axis visible, ifnotclearly visible (some formsofA. sciureus), then 13-27 cm long 2 - Inflorescence dense, globose to long cylindrical, the axis not visible, shorter 5 2 Stipules 17-27 mm long; median leaves 10-16 cm long A. sciureus - Stipules 6-15 nam long; median leaves up to 8(-10) cm long 3 3 Rachides spreadingly hairy; peduncle villose A. lagopodioides - Rachides appressedly hairy; peduncle appressedly hairy or longer hairs rarely subappressed to ± patent 4 4 Median leaves 3-8(-10) cm long; rachides obliquely erect to patent; calyx at mm anthesis campanulate-tubular, immediately after anthesis inflated, 5-8 wide, elliptic-cylindrical A. rubrostriatus 263 - Median leaves 2-3.5 cm long; rachides mostly deflexed; calyx at anthesis tubular, mm after anthesis notor slightly inflated, 3-4 wide, tubularto narrowly elliptic A. paralurges 5 Keel 19-26mm long, the limbs 7-10mm long, 3.5-4 mm wide 6 mm mm mm 6 LKeeaeflle1t1s-02.05-3 mlmongw,idteh,e lliinmebasr4t-o6n.a5rrowllyonobglaonngd-e2l.l5i-p3t.i5c, ± fowliddeed; bracts acut7e or shortly acuminate A. kohrudicus - Leaflets 3-12 mm wide, elliptic to broadly elliptic, flattened; bracts longly acu- minate A. glumaceus I Bracts thinly membranaceous, hyaline, neverpurple at tip, glabrous or only ciliate at the margins; calyx teeth and nerves of same colour as tube; corolla limb sul- phureous or pale yellow A. chrysostachys - Bracts thickly membranaceous to glumaceous, sometimes purple at tip, if thinly membranaceous and hyaline at margins then calyx teeth and nerves red to purple, bracts densely pubescent or hairy only on the midrib; corolla limb pink, purple, violet, yellow to white 8 8 Inflorescence shorter than or as long as the leaves i) 9 Inflorescence distinctly overtopping the leaves 14 9 Leaflets in l-3(^) pairs A. laguriformis - Leaflets in (3-)4-10 pairs 10 10 Corolla limb pale yellow A. velenovskyi - Corolla limb pink to purple 11 mm II Bracts longer than or as long as the calyx, lowerones 13-20 long 12 mm Bracts distinctly shorter than the calyx, lower ones 9-11(13) long 13 12 Bracts densely pubescent or hairy at least on midrib and apex; stipules charta- ceous, not hyaline; leaflets densely appressed hairy, silvery green A. tabrizianus - Bracts glabrous; stipules membranaceous, hyaline; leaflets sparsely appressed hairy, becoming glabrous, green A. pediculariformis mm 13 Peduncle covered with short subappressed hairs, long hairs up to 2 long A.persicus - Peduncle villose, long hairs 2-3.5 mm long A. hirticalyx 14 Bracts longer than or as long as the calyx, persistent in fruit 15 - Bracts shorter than the calyx, caducous, rarely persistent 17 15 Leaflets densely to sparsely spreadingly hairy; young rachides patent hairy A. persicus - Leaflets densely appressed hairy; young rachides ± appressedly hairy 16 mm 16 Bracts very broad, lower ones 7-12 wide, inside hairy at apex, shortly or rarely long acuminate, acumen shorter than halfofthe limb, if longer, then purple tipped A. hymenostegis - Bracts narrower, lower ones 5-8 mm wide, inside glabrous at the apex, long aristate, arista more than halfas long as limb, the apex mostly the same colour as the limb A. lagopoides mm mm mm 17 Calyx 16-23 long, the hairs up to 6 long; wings 18-22 long, their mm hmbs 8-12 long; inflorescence at least 2-times longeras wide A. nervistipulus - Calyx 12-18 mm long, the hairs up to 4.5 mm long; wings 14-18 mm long, if wings longer, then inflorescence globose or up to 1.5-times longer as wide 18 1) In youngerforms ofA. uraniolimneus inflorescences sometimes are shorterthan the leaves. This specimens can be determined from A. hirticalyx by their thin, hyaline and flexible stipules. 264 18 Corolla limb white oryellow 2) 19 - Corolla limb pink to purple or violet 21 19 Calyx nerves distinctly red to purple A. persicus - Calyx nerves whitish yellow, without any contrast with the rest of tube 20 20 Hairs mostly brownish yellow; leaflets with double indumentum, i.e. densely hairy with long, subappressed straight hairs and under them shortly tomentose; pods mm 8-10 long A. recognitus - Hairs pure white; leaflets simply sericeous or spreadingly hairy; pods 6-7 mm long A.persicus 21 Median leaves up to 5 cm long 22 - Median leaves 6-16 cm long 24 22 Standard limb ± as long as the claw or slightly longer A. persicus Standard limb 1.7-3 times longer than the claw 23 23 Rachides straight, thick, rigid, obliquely erect to subhorizontal, older ones not broken A. uraniolimneus Rachides recurved, thin, flexible, olderones mostly broken A. hymenocystis a Leaflets covered with yellow hairs, thick, elliptic to broadly elliptic or obovate, obtuse at tip A. hymenocystis subsp. hymenocystis b Leaflets white hairy, thin, narrowly oblong-elliptic, rarely narrowly obovate, mostly acute at tip A. hymenocystis subsp. confiniorum 24 Inflorescence 2-3 cm wide, ovate to long cylindrical; standard 13-22 mm long, the limb up to 1.5 times longer than the claw; bracts pubescent or glabrous A. persicus - Inflorescence 3^.5 cm wide, globose to ovate; standard 18-27 mm long, the hmb 2-3 times as long as the claw; bracts glabrous or only ciliate at margins 25 mm mm 25 Peduncle patent hairy; standard 20-27 long; wing limb 8-11 long; mm leaflets 7-30 long, linear to narrowly oblong A. sti-aussii - Peduncle appressedly hairy; standard 18-21 mm long; wing limb 7-8(-9) mm mm long; leaflets 6-18 long, narrowly elliptic A. chehreganii & 2.Astragalus chehreganii WZarre Podlech sp. nov. Holotype: Persia, Prov. Azarbaijan, injugo Qushchi inter Shahpur et Rezaiyeh, 1600-1850 m, 13.6A911, Rechinger41877(V^Ulso: MSB!). Fig. 2a Differt abA. straussii foliolis brevioribus et latioribus, pedunculo appresse (nee pa- mm tenter) piloso, vexillo 18-21 (nee 20-27) longo et lamina alarum 7-8 (nee 8-11) mm longo. Plantae suffruticosae, ad 40 cm altae. Caules saepe ascendentes, ad 22 cm longi, parte homotino 1-1,5 cm longo. Stipulae firme membranaceae, flavidae, 12-17 mm mm longis, per 6-10 petiolo adnatae, acuminatae, glabrae vel margine ciliatae. FoUa 2,3-11,5 cm longae, rachidibus primo flexilibus demum rigidis validis, oblique erectis ad patentibus, dense serieeis, demum glabrescentis, petiolo laminae attingente, spina terminali 1/4-1/2 longitudinis foliolorum apicalium attingente. Foliola (3-)5-ll juga, remota, 7-20 mmmmlonga et 1,5-4 mm lata, linearia ad anguste elliptica, acuta, mucrone terminali ad 2 longo, plana, utrimque dense vel sparse sericea. Inflorescntia dense 2)A. chehreganii has acorolla which sometimes seems yellow, but withexact observation it will be clear that they are tinged with red or pink, and through drying the colour was changed. Theglobose inflorescenceofthelattercanbeusedasanotherdiagnosticcharacter fordistinguishing itfromyellow-floweredspeciesofthe section.