Nordic Journal of Botany 27: 328
335, 2009 doi: 10.1111/j.1756-1051.2009.00187.x, # The Author. Journal compilation # Nordic Journal of Botany 2009 Subject Editor: Arne Strid. Accepted 17 February 2009 Astragalus sect. Trachycercis (Fabaceae) in Iran Massoud Ranjbar M. Ranjbar (ranjbar@basu.ac.ir), Dept of Biology, Herbarium division, Bu-Ali Sina Univ., PO Box 65175/4161, Hamedan, Iran. A diagnostic key, and new data are provided for seven species of Astragalus sect. Trachycercis, which occur within the area covered by the Flora of Iran. Astragalus brevipedunculatus Ranjbar is described as new, and A. armeniacus Boiss. is reported as new for the flora of Iran. The micromorphology of the seed coat surface of ten Astragalus taxa has been studied by SEM. Considerably different types of sculpturing at species level and similarities between related taxa were observed. In addition, differences between A. sect. Trachycercis and the closely related sections A. sect. Erioceras and A. sect. Wettsteiniana are discussed. Astragalus durandianus, A. pseudoshebarensis and A. shebarensis, which all have strongly inflated pods, are transferred from A. sect. Erioceras to A. sect. Wettsteiniana. Astragalus brevipedunculatus sp. nov., described from a small region near Aras River in Iran, is the only geographically isolated species of the section. Members of the section prefer habitats from coastal dune climates of the Azarbaijan Sharqi Province to the Armenian and Azarbaijan frontiers, although possibly also growing in the adjacent southern parts of Armenia and Azarbaijan. Astragalus brevipedunculatus sp. nov. is the only species of the section in Iran and neighboring countries that has few pairs of leaflets and is completely glabrous. The new species should be considered as a ‘Critically Endangered’ (CR) following the IUCN criteria. A distribution map for A. brevipedunculatus and A. armeniacus is presented. Astragalus, with nearly 3000 species, is one of the largest genera of flowering plants (Podlech 1986, Lock and Simpson 1991, Mabberley 1997, Maassoumi 1998, Ranjbar and Karamian 2002). Iran alone has more than 840 species and is one of the main centers of diversity of the genus (Lock and Simpson 1991, Mabberley 1997, Maassoumi 1998, Ranjbar and Karamian 2002). Astragalus has been divided into approximately 150 sections of which A. sect. Trachycercis is one of the most diverse and variable. It belongs to A. subgen. Cercidothrix Bunge, which is characterized by perennial growth and the presence of bifurcate hairs (Bunge 1868
1869). This sub-genus with more than 800 species has its main center of diversity in central (Yakovlev et al. 1996) and southwest Asia (Ranjbar 2007a, 2007b). In the course of the work on Astragalus, the majority of bifurcate-haired species known from Iran have been (Rechinger et al. 1961, 1969, Ghahreman et al. 1996, Maassoumi and Ranjbar 1998, Ranjbar and Maassoumi 1998, Maassoumi et al. 1999, 2000, Podlech 2001, Podlech and Sytin 2002, Podlech and Maassoumi 2003, Podlech and Zarre 2003, Ranjbar and Karamian 2003, 2004, Zarre et al. 2005, Karamian and Ranjbar 2005b, Ranjbar et al. 2005, 2007a, 2007b, Ranjbar 2007a, 2007b). During field investigations to the mountains of Iran, some peculiar populations of A. sect. Trachycercis were found. They were distributed above the sub-mountains around Maku, Khoi (Seid Hajian), and across a dune coastal habitat on the Aras River. Investigations on living 328 material and herbarium specimens suggest that populations growing along dune coasts of Aras river and Seid Hajiadin in Khoy are closely related to A. theodori Grossh., but several differences in their morphology allow us to treat them as new distinctive species. The applications of Scanning Electron Microscopy (SEM) to vegetative and reproductive organs have amply demonstrated the importance and impact of this technique in systematic studies. Brisson and Peterson (1977) listed 273 references to microanalysis of seed surfaces in a bibliography. The most detailed information about seed surfaces in Astragalus has been obtained for 9 species belonging to Astragalus L. sect. Astragalus L., Ammodendron Bunge and Caraganella Bunge (Karamian and Ranjbar 2005a, Ranjbar et al. 2007b) from Iran and forty-eight species of Astragalus L. sect. Onobrychoidei DC., Uliginosi Gray and Ornithopodium Bunge (Fabaceae) from Turkey (Vural et al. 2008) using SEM observations. The correlation between surface patterns and habitats has been discussed and these characters have been declared to be of minor taxonomic significance (Chakrabarty and Mukherjee 1986). Classification and terminology of plant epicuticular waxes for comparative descriptions have been obtained from an analysis of a number of species including Astragalus spp. The result stressed the taxonomic and diagnostic significance of these characteristics. The main aim of this study is to assess the value and utility of micromorphological characters of the seeds as an additional source of information for the taxonomy of A. sect. Trachycercis and A. sect. Erioceras. Material and methods This study is mainly based on herbarium material and field observations during several excursions in Iran, supported by the Bu-Ali-Sina Univ. The collected materials were in vegetative or fruiting phase. Several herbarium sheets of each species have been examined from the following herbaria: PR, W, WU, TARI, FUMH and BASU. All vouchers are deposited at BASU. Seed surface sculpturing of the species belonging to A. sect. Trachycercis and A. sect. Erioceras were studied by scanning electron microscopy to evaluate the diagnostic value of this character. Seeds were taken from herbarium specimens or were directly collected in the field, and documented by voucher specimens (Table 1). They were soaked in water to remove the adherent outermost layer before gold coating, and was then studied by a Cambridge 360 scanning electron microscope. Notes on Astragalus sect. Trachycercis Astragalus L. sect. Trachycercis Bunge (1868, p. 114). Lectotype (Podlech 1990): A. humilis M. Bieb. Bunge (1868
1869) introduced A. sect. Trachycercis as a new section with 16 species. It belongs to the bifurcatehaired Astragali group from the sub-genus Cercidothrix. Today, about 60 species are referred to this relatively large section (Lock and Simpson 1991, Yakovlev et al. 1996, Maassoumi 1998). The majority of the species that belong to this section are distributed in central Asia (Yakovlev et al. 1996). The section has been revised for the flora of the former Soviet Union (Gončarov et al. 1946), the flora of Turkey (Chamberlain and Matthews 1970), the flora of Iraq (Townsend and Guest 1974) and the flora of Iran (Maassoumi 2005). Members of A. sect. Trachycercis are characterized by small, often stemless, habit and small pods, without or with reduced peduncles. It seems that this section belongs to the bifurcate-haired astragali group together with A. sect. Cystodes, A. sect. Dissitiflori, A. sect. Cremoceras, A. sect. Cystium, A. sect. Erioceras, A. sect. Trachycercis and A. sect. Incani. In part, this hypothesis is supported by molecular data (Kazempour Osaloo et al. 2003). Astragalus sect. Trachycercis is not sharply separated from A. sect. Incani and A. sect. Erioceras. It differs from A. sect. Erioceras mainly by stemless habit (or with reduced stem), and by peduncles and inflorescences with few flowers (Ranjbar and Karamian 2004). It can be distinguished from A. sect. Incani by small habit, pods densely covered with sub-appressed long hairs (or rarely glabrous) and the absence of dots on both surfaces of leaflets. Key to the species of Astragalus sect. Trachycercis in Iran 1. Plant glabrous, pod unilocular, glabrous, ca 9.5 ca 2.5 mm ..........................................A. brevipedunculatus
Plant densely covered with sub-appressed long hairs, pod bilocular, hairy ..................................................... 2 2. Corolla white, calyx exclusively covered by dense white hairs ............................................................................. 3
Corolla violet to pale violet, calyx densely covered by black and white hairs................................................... 5 3. Plant densely covered with spreading long hairs, standard 24
30 mm long ........................................ A. barnassari
Plant densely covered with appressed hairs, standard 15
25 mm long .......................................................... 4 4. Pod sessile, 12
15 ca 2.5 mm, leaflets in 5
8 pairs, standard 15
16 mm long ...................... A. poliothrichus
Pod stipitate, 10
175
10 mm, leaflets in 7
14 pairs, standard 17
25 mm long............... A. armeniacus 5. Plant densely covered with spreading long hairs .............................................................. A. durandianus
Plant densely covered with appressed to sub-appressed hairs ............................................................................. 6 6. Leaflets in 3
5 pairs, flowers 2
5, pods 15
18  7
8 mm ................................................... A. shebarensis
Leaflets in 3
4 pairs, flowers 1
2, pods 15
25  8
10 mm.........................................A. pseudoshebarensis N.B. A. durandianus, A. shebarensis and A. pseudoshebarensis doubtfully belong here. I prefer to include them in A. sect. Wettsteiniana. Table 1. Sections, vouchers and status of the Astragalus species studied in Iran. Taxa A. brevipedunculatus Ranjbar A. armeniacus Boiss. A. sympileicarpus Rech. f. A. pentanthus Boiss. A. neo-sytinii Ranjbar A. keredjensis Podlech A. khajehensis F. Ghahremani A. catacamptus Bunge A. anacamptus Bunge A. pakravaniae Podlech & Maassoumi Section Trachycercis Trachycercis Erioceras Erioceras Erioceras Erioceras Erioceras Erioceras Erioceras Erioceras Collector Locality Date Rahiminejad and Ranjbar Rahiminejad and Ranjbar Maassoumi et al. Ranjbar Namati Maassoumi and Jalili Assadi and Ranjbar Ranjbar Ranjbar Assadi and Ranjbar Jolfa Maku Birjand Hamadan Yazd Karaj Haris Isfahan Semnan Gorgan 12 Jul 2003 11 Jul 2003 3 May 2003 10 May 2004 25 May 1998 20 May 2003 1 Jun 2004 8 May 2003 5 May 2002 8 May 2002 Voucher number Status BASU 5494 BASU 5465 TARI 83362 BASU 5057 BASU 1250 TARI 82404 BASU 5465 BASU 5701 BASU 8278 BASU 5465 endemic native native endemic endemic endemic endemic endemic endemic endemic 329 Astragalus brevipedunculatus Ranjbar sp. nov. (Fig. 1) Ab A. theodori Grossh. planta glabra (nec dense appresse pilosa), foliis 3
4-jugis (nec 7
8-jugis), foliolis 5
12 nec (2
3) mm longis, leguminis obovatis (nec oblongo
ellipticis) glabris (nec appresse pilosis) differt. Type: Iran. North Azarbaijan Sharqi: Bazargan toward Jolfa, margin Aras River, 900 m a.s.l., 12 Jul 2003, Ranjbar 5494 (holotype: BASU!, isotype: TARI!). Plant (7
)10
20 cm tall. Caudex up to 1
2 cm in diameter, divided, with few short branches, densely covered with remnants of old leaves. Stipules papery or membranous pale brown, 5
12 2
3 mm, linear to lanceolate
triangular, the longest adnate to the petiole for 2
3 mm, glabrous or sparsely to loosely sub-appressed white hairy, with 6
11 parallel nerves in the free portion, margins ciliate with spreading 0.5
1.0 mm long hairs. Leaves 5
15 cm long; petiole 2
6 cm long, as the rachis finely striate, 1.5
2.0 mm thick, glabrous. Leaflets in 3
4 pairs, ovate
orbicular to elliptic, 5
128
15 mm, cuneate to rounded at the base, truncate
rounded or rarely obtuse at the apex, often minutely mucronulate, sometimes only at margins ciliate with appressed white 0.2
3.0 mm long hairs, otherwise glabrous. Peduncle 8
10 mm long, erect, filiform. Raceme 1- or 2-flowered, strongly contracted in fruit, up to 20 mm long, densely covered with remnants of old leaves. Pedicels ca 2 mm long. Corolla unknown. Pods erect to ascending, obovate, rarely spherical, sessile, slightly oblique, 7
8 mm long and ca 5 mm high and wide, at the apex abruptly narrowed into an acuminate 1
3 mm long beak, unilocular, opening at both sides beginning at the apex; valves straw colored to pale brown, glabrous. Seeds ca 2 in each locule, Figure 1. Astragalus brevipedunculatus sp. nov. (A) habit, (B) pod, (C) leaflets. Scale: (A) 1 cm, B3 cm. 330 brownish, irregularly rectangularly reniform, ca 3 ca 3 mm (Fig. 1). Additional specimens examined (paratypes) Iran. North Azarbaijan Garbi: 35 km Khoy to Salmas, Seid Hajiadin, 1400
1500 m a.s.l., 7 Jun 2004, Assadi and Ranjbar 6088 (BASU!). Taxonomic remarks Astragalus brevipedunculatus is closely related to A. theodori Grossh., especially because of similar shape and size of the pod, the length of the peduncle and the number of leaflets. I have not seen any material belonging to this species from Azarbaijan in herbarium W, WU or PR., but it was recorded from there by Gončarov et al. (1946). Astragalus theodori have probably evolved from Iranian species with fewer pairs of leaflets, e.g. A. brevipedunculatus. In turn, A. brevipedunculatus is related to perennial legume species, especially A. shelkovnikovii Grossh. Astragalus brevipedunculatus and A. armeniacus are two rare allopatric species. They both occur as a small number of individuals. This study provides more information about their distribution, ecology, and taxonomy. Their distribution is across a dune coastal habitat in the northern Azarbaijan Sharqi Province. To investigate their taxonomy, I used morphology as a predictor of their relationships. A. brevipedunculatus with an estimate of 40 plants per hectare was very abundant and well distributed at two localities. Astragalus armeniacus was less abundant and occurring at only one locality. I often found both of them growing side-by-side in regions exposed to sunlight and wind. The highest abundance of A. brevipedunculatus was on the river edges. These astragali species were found in Figure 2. Distribution map of A. brevipedunculatus (star), A. armeniacus (black circle), A. barnassari (white circle), A. poliothrichus (white square), A. pseudoshebarensis (black square), A. durandianus (black triangle) and A. shebarensis (white triangle) in Iran. Garbi Provinces. It was collected from the dry-steppe and sand zones on the southern side of Aras River. Astragalus brevipedunculatus is threatened by grazing. Therefore, it should be regarded as ‘Critically Endangered’ (CR) following the IUCN criteria (IUCN 1994). Etymology The specific epithet is ‘brevipedunculatus’, which means ‘with short peduncle’. Astragalus armeniacus Boiss. (1843, p. 86) new record for Iran
Holotype: in Armenia, P. M. R. Aucher-Eloy 4450 (G-BOIS; isotype: BM, G, K, P: foto MSB), Fig. 3. The presence of this species in Iran has been substantiated by the following specimens: Iran. Azarbayejan Garbi: Keshmeshtapeh toward Maku, before Ghezelbolagh, Badoly village, 2000 m a.s.l., 29 Jun 2003 Assadi and Ranjbar 5379 (BASU!); Chaldoran toward Keshmeshtapeh, 50 km before Keshmeshtapeh, 1833 m a.s.l., 11 Jul 2003, Rahiminejad and Ranjbar 5465 (BASU!). Taxonomic and distributional remarks Figure 3. Astragalus armeniacus. (A)
(B) habit, (C) calyx, (D) standard, (E) keel, (F) androecium, (G) pods. Scale: (A), (B), (G) 1.5 cm, (C)
(F)3.5 cm. different habitats. Each species was distinct and they did not occur in combination. Distribution and suggested conservation status Astragalus brevipedunculatus has been seen in the field by the author and is known from seven specimens only from two localities deposited at BASU (Fig. 2). It is endemic to northwest Iran and occurs in the Azarbaijan Sharqi and The new record was seen in the field by the author but is known only from nine specimens from two localities. It occurs in the Azarbaijan Garbi Province (Fig. 2). It has been collected from northwest Iran close to eastern Turkey and the southern Armenian frontier in sub-mountainous slopes around Maku. It grows on limestone at altitudes 1800
2000 m a.s.l. I did not find any material from Armenia belonging to this species in herbarium W. It has been recorded by Chamberlain and Matthews (1970) from the surroundings of Erzurum in Turkey. The texture of the fruits is unknown and lacking in both the type specimens and the original description, but the new Iranian specimens fit in most respects with the original description of A. armeniacus. However, it is a further example of the close relationship between the bifurcate-haired astragali group in Iran, Turkey and Armenia. Some differences between these closely related species are presented in Table 2. Table 2. Diagnostic morphological characters of A. theodori, A. armeniacus and A. brevipedunculatus. Taxa characters Indumentum Leaf length (cm) Leaflet number (pairs) Leaflet length (mm) Flower number in inflorescence Pod length (mm) Pod shape Pod indumentum A. theodori A. brevipedunculatus A. armeniacus densely appressed, hairy 2
5 7
8 2
3 (5) few-flowered 8
10 oblong
ovoid appressed white hairs glabrous 5
15 3
4 5
12 1
2-flowered 7
8 obovate to rarely sperical glabrous sparsely appressed, hairy 3
10 7
14 6
10 3
4-flowered 10
17 orbicular
ovoid often appressed white hairs 331 An overview of the position of the Astragalus durandianus group Here the positions of A. durandianus, A. shebarensis and A. pseudoshebarensis are reviewed. They can be placed in an isolated group. The two former species have previously been referred to A. sect. Caraganella (Lock and Simpson 1991), but were transferred to the new A. sect. Wettsteiniana by Širjaev and Rechinger (Rechinger 1955). Moreover, the extremely asymmetrically bifurcate hairs on leaflets and strongly inflated pods are diagnostic characters for all species belonging to this section. However, Astragalus sect. Wettsteiniana was not accepted by Podlech (pers. comm., Lock and Simpson 1991, ref. 1143). These species were excluded from the new section and assigned to A. sect. Erioceras and then to A. sect. Trachycercis by Podlech (Lock and Simpson 1991, Podlech 2004, Maassoumi 2005). Astragalus durandianus, A. pseudoshebarensis and A. shebarensis occupy an intermediate position between A. sect. Trachycercis and other sections of bifurcate-haired Astragalus, especially A. sect. Erioceras, and have been treated differently in important taxonomic works. Based on the type of indumentum and the shape of leaflets, it seems that the A. durandianus group is more closely related to A. sect. Erioceras than to A. sect. Trachycercis (Ranjbar and Karamian 2004). On the other hand, this group can be placed in A. sect. Wettsteiniana since it has strongly inflated pods. I agree with this treatment and prefer to maintain A. sect. Wettsteiniana as a separate section with three species (A. durandianus, A. pseudoshebarensis, A. shebarensis) rather than as a group within A. sect. Trachycercis. However, more material, field examination and molecular data are needed for clarifying the exact taxonomic position of the A. durandianus group. Geographical distribution and ecology Almost all members of A. sect. Trachycercis are Irano
Turanian elements. The species of this section are distributed in central Asia, south Asia, the Middle East, east Asia, north Asia, west Asia, the Caucasus and Europe. China and Mongolia with about 20 species are the centers of diversity of the section. Kazakhstan with 11 species, Russia with 8 species, Iran with 7 species, Turkey with 7 species, Kirgizistan with 7 species, Azarbaijan with 3 species, Turkmenistan with 2 species, Ukraine with 2 species, Uzbekistan with 2 species and Tadzhikistan with 1 species are other countries in which the section is native. Nearly 12% of the species of the section exist in Iran, with 7 species distributed in mountainous regions (Fig. 2). The widest ranging species are A. barnassari, A. durandianus and A. pseudoshebarensis. The Iranian species are distributed from an altitude of about 900 m a.s.l. (A. brevipedunculatus near Aras River in the Azarbaijan Sharqi Province) to 2350
3500 m a.s.l. (A. pseudoshebarensis in Fars, Kerman and Yazd Provinces). In all collections from throughout the distribution, the altitudes were often between 900
3500 m a.s.l. Flowering and fruiting in species of the section occur during (Apr
) May
Jun (
Jul). In general, it seems that local endemism plays an important role in the Iranian species of the section. Some of the species with such narrow 332 distribution pattern are A. pseudoshebarensis (Sare-Sefid Mountain, around Neyriz in Fars Province), A. brevipendunculatus (from Aras River in Azarbaijan Sharqi Province), A. poliothrichus (Kohrud Mountain in Esfahan Province, C Zagros) and A. durandianus (Bizg Mountain in Khorassan Province). Taxonomic importance of seed surface micromorphology Seed morphology is of considerable significance for species delimitation. According to our observations, seed features such as seed shape, size, and seed coat sculpturing vary between species. These features can be used to classify and circumscribe the species. The shape of the Astragalus seed is reniform, ovoid, orbicular or rectangular, and the colour is light or dull brown. In A. sect. Trachycercis and A. sect. Erioceras, the outline of the seed is usually rectangular. The seed surface is usually rough, which is only easily observable with the SEM. SEM studies have shown micromorphological diversity, indicating the potential taxonomic value of this feature (Karamian and Ranjbar 2005a, Ranjbar et al. 2007b). The seed-coat surface of Astragalus brevipendunculatus consists of a wavy ridge around the cells that bears small invaginations, merging loops and protuberaces (Fig. 4). In contrast, the A. armeniacus seed-coat surface is a monolayer reticulate formed by polygonal cells with corrugated radial walls of medium depth uniformly thickened, and tangential walls irregularly compressed (Fig. 5). The separate specific status of A. brevipendunculatus is thus confirmed by details of the seed surface micromorphology. In addition, A. brevipendunculatus is readily distinguishable from A. armeniacus by having small and unilocular pod. The seed surface is reticulate with an obvious rugose layer in A. sympileicarpus (Fig. 6). The seed coat surfaces of A. pentanthus and A. neo-sytinii have a double reticulate pattern formed by polygonal cells with radial walls of relatively large depth, evenly thickened, and tangential walls, each with papilla of the colliculate type, i.e. small rounded and hill-like elevations covered with granules or micropapilla-like projections seen at high magnifications (Fig. 7, 8). In A. keredjensis the seed coat sculpturing has a single reticulate pattern and resembles the secondary sculpturing on the seed surface of A. pentanthus and A. neo-sytinii (Fig. 9). The close relationship between these species is also confirmed by macromorphological characters. However, A. keredjensis is readily distinguishable from A. pentanthus and A. neo-sytinii by having triangular and completely glabrous pods. The seed surface of A. khajehensis is typical for the section. Topographically, it consists of an uneven surface with depressed central parts and thickened, wrinkled, anticlinal walls (Fig. 10). In A. catacamptus and A. anacamptus the seed coat sculpturing has a dense irregular
reticulate pattern with wavy ridges (Fig. 11, 12) and the reticulate patterns appear similar to each other. In addition, the close relationship between these species is supported by macromorphological data. In A. pakravaniae, the seed-coat sculpturing consists of a monolayer and has a rugose stellate
reticulate pattern with obvious pits (Fig. 13). This special pattern in A. pakravaniae is different from those in the other species. This result is consistent with Figure 4
13. Seed SEM micrographs of Astragalus sect. Trachycercis and A. sect. Erioceras. (4) A. brevipedunculatus, 3000. (5) A. armeniacus, 3000. (6) A. sympileicarpus, 2500. (7) A. pentanthus, 2500. (8) A. neo-sytinii, 2500. (9) A. keredjensis, 2500. (10) A. khajehensis, 2500. (11) A. catacamptus, 2500. (12) A. anacamptus, 2500. (13) A. pakravaniae, 2500. 333 morphological data. However, the similar seed surface patterns confirm the close morphological relationship between A. sect. Trachycercis and A. sect. Erioceras. They are not geographically well separated from each other. Considering that the surface characteristics of mature seeds are stable features, which are not effected by environmental conditions and are genetically controlled, hyperdiversity (which is typical for each species in Astragalus) may be evaluated for diagnostic purposes, but it is necessary to understand how plastic these interspecific surface structures are for species that grow in different habitats, in order to utilize this parameter as a fingerprint of a species. In a study on the seed surface of Epilobium, it was concluded that real genotypic variability is very small (Skvortsov and Rusanovitch 1974). The genetically fixed epidermal patterns may not change with large representative samples (Chakrabarty and Mukherjee 1986). Extensive scale to test phenotypic variability of the seed surface characteristics are worth more numerous investigations in individuals of a widely distributed taxon, rather than for local endemics. Two general types of surface patterns were determined among the species examined here. Reticulate and rugose sculpturing in seed surfaces may indicate the phylogeny of A. sect. Trachycercis and A. sect. Erioceras. The species having seeds with similar surface ornamentations may have common phylogenetic relations, but the relationship inferred from these micro-characteristics need to be evaluated by molecular data. Furthermore, it may be possible to identify Astragalus species using a seed sample only by comparing with a data base where the information on the surface patterns obtained in a large sampling of Astragalus genus is collected. Acknowledgements
The great help of Dr Vitek, Dr Wallnofer and Dr Till during our visit to herbarium W and WU in Vienna and the help of Dr Otakar Šı́da during the visit to herbarium PR in Prague is much appreciated. I would like to thank the director of the herbarium Res. Inst. of For. and Rangelands (TARI) and the herbarium of Ferdowsi, Univ. of Mashhad (FUMH) for making the herbarium facilities available and for lending us herbarium specimens. This project has received financial support from the Bu-Ali Sina University (project no: 32-723 ‘Pollen study of Astragalus in Iran’). In addition, I wish to thank Mr Moradi for preparing the illustration. References Brisson, J. D. and Peterson, R. L. 1977. The scanning electron microscope and x-ray micro-analysis in the study of seeds: a bibliography covering the period of 1967
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