Sendtnera Mitteilungen der Botanischen Staatssammlung und des Instituts für Systematische Botanik der Universität München Herausgeber: Jürke Grau - Franz Schuhwerk Band 7 München 2001 Sendtnera Mitteilungen der Botanischen Staatssammlung und des Instituts für Systematische Botanik der Universität München Herausgeber: Jürke Grau - Franz Schuhwerk Band 7_ München 2001 Sendtnera (Mitteilungen der Botanischen Staatssammlung und des Instituts für Systematische Botanik der Ludwig-Maximilians-Universität München) Band 7 Erscheinungsdatum: 31.8.2001 Herausgeber: JÜRKE GRAU und FRANZ SCHUHWERK Redaktion: CHRISTINE EHRHART Anschrift: Botanische Staatssammlung München - Institut für Systematische Botanik der Ludwig-Maximilians-Universität München Menzinger Straße 67, D-80638 München, Deutschland. ISSN 0944-0178 Inhalt BOENDER, R. & ULMER, T.: Passiflora tina spec. nov., a new species of Passiflora, subgenus Astrophea (Passifloraceae) from Ecuador..................020000- Davies, A.M.R. & FACHER, E.: Achene hairs and their diversity in the genus Chaetanthera Ruiz & Pav. (Mutisieae, Asteraceae) ................0 cece cece DOBBELER, P.: Polytrichadelphus magellanicus sensu lato (Musci) and its asco- Bugs ie different tunpi On ditierent NOSIS : ......... 2... won e See os ee EHRHART, C.: Zephyra compacta (Tecophilaeaceae) — eine neue Art aus Chile... . ERBEN, M.: Bemerkungen zur Taxonomie der Gattung Limonium VII ........... FÖRTHER, H. & PODLECH, D.: Contributions to the Flora of Northern Africa I, New BEROIEWOTNYATAXA a! es ee a ee ee HERTEL, H.: Floristic and taxonomic notes on saxicolous lecideoid lichens........ KÖBELE, C.P. & TILLICH, H.-J.: Die Infloreszenzen der Juncaceae.............. PODLECH, D.: Contributions to the knowledge of the genus Astragalus L. (Legumi- BOSSEHNT-X 22... 0 errr le a meee ee Rene PODLECH, D. & MAASSOUMI, A.A.: New species of Astragalus L. sect. Hymeno- Biepis omalranı.. 22.200 ode BS eee ie oe ee ee TRIEBEL, D. & SCHOLZ, P.: Lichenicolous fungi from Bavaria as represented in the Botanische Staatssammlung München : . 0.25.3). 2235.22. 8 ee EN ZARRE, SH. & PODLECH, D.: Taxonomic Revision of Astragalus sect. Acantho- PRES (EP ADSCCAL) nn. ol 55 &6-8 ac) sin ee Hehe A IE eee New taxa and new combinations published in Sendtnera 7 (31.8.2001).......... We thank the reviewers for their helpful comments and contributions to this volume 163 203 211 233 253 veraggelim - ui raten apc er. ae Me Em m ut ie reihen yma, = soit (ihe sgh eter « poate fe fi, De, re N I \e ba i Bam u Anbdarra eu KR rg N ellis ” u ei! yA : ri “i i. en Passiflora tina spec. nov., a new species of Passiflora, subgenus Astrophea (Passifloraceae) from Ecuador R. BOENDER & T. ULMER Abstract: BOENDER, R. & ULMER, T.: Passiflora tina spec. nov., a new species of Passiflora, subgenus Astrophea (Passifloraceae) from Ecuador. — Sendtnera 7: 5-12. 2001. ISSN 0944-0178. A new species of Passiflora is described from northwestern Ecuador. Passiflora tina R.Boender & T.Ulmer is characterized by its flowers grouped in dense fascicles, the wavy outer corona filaments surrounding the androgynophore in a campanulate fashion, the deeply cleft operculum, and the structure and length of the inner corona series. The new species is somewhat intermediate between different sections of subgenus Astrophea and thus questions the current classification. It most closely resembles P. callistemma, P. maguirei and P. pittieri. The differences between the four species are discussed. Zusammenfassung: Mit Passiflora tina wird eine neue Art der Untergattung Astrophea aus dem nord- westlichen Ecuador vorgestellt. Damit hat sich die Anzahl der in Ecuador beheima- teten Vertreter der Untergattung Astrophea seit 1988 mehr als verdoppelt. Charak- teristisch fiir Passiflora tina sind im wesentlichen die vielblumigen Bliitenstande und die Struktur der äußeren Strahlenkranzreihe. Diese neue Art ist besonders eng mit P. pittieri verwandt und wurde zeitweise auch für diese gehalten. Eine Unterscheidung beider ist jedoch leicht möglich. Während P. tina einen aufrechten, glockenförmigen Strahlenkranz aufweist, liegt dieser bei P. pittieri hernieder. Darüber hinaus bieten die Anzahl der Elemente der äußeren Strahlenkranzreihe sowie die Menge der Blüten per Nodium eine gute Unterscheidungsmöglichkeit. Weitere verwandte Arten sind P. calli- stemma und P. maguirei. Von beiden kann P. tina mit Hilfe ihres 5-reihigen Strah- lenkranzes, des breiteren Blütenkelches, sowie der im oberen Drittel stark verbreiterten Strahlenkranzelemente unterschieden werden. Introduction During various trips to Tinalandia, a pre-montane rain forest site on the western slope of the Andes in northwestern Ecuador, the senior author worked on the life history of Heliconius sapho candidus Brown (in press). This spectacular butterfly was previously very common. Its host plant and that of a closely related butterfly, Heliconius eleuchia primularis Butler, was in Ecuador listed as Passiflora macrophylla Spruce ex Mast. by BENSEN et al. (1976). Both species of butterflies mate while the female is still in the pupal stage. These closely related species would be in competition for the same host and therefore a second host passion vine had to exist in the area, probably P. pittieri Mast. or something similar (Gilbert, pers. comm.). The widespread neotropical H. sapho feeds on P. pittieri in Central America. During the 15th trip to Tinalandia in 1996, Heliconius sapho candidus was observed leaving a nectaring flower and circle a small area a number of times as if there were a host plant and then fly away. A search of the location revealed 3 small juvenile treelets of P. tina. They were growing in pre-montane rain forest on a north facing slope of about 35 to 40 degrees. The site was remarkably rich in litter of 10-15 cm deep, and the soil is typical laterite. In culture, the treelets turned into climbing vines with well developed tendrils, which bloom about 2-3 times per year from old wood on shady parts of the plant. Remarkable is the extreme amount of nectar and the very nice odor of the flowers. A careful examination of its characteristics and a comparison with other Astrophea species revealed P. tina differs from its close relatives, P. callistemma Escobar, P. maguirei Killip and P. pittieri Mast., in many characters and should be regarded as an independent species. Passiflora tina R.Boender & T.Ulmer, spec. nov. Type: Ecuador. Prov. Pichincha, Tinalandia, southeast of Santo Domingo de las Colorados on road to Alluriquin, 700 m, 6 May 1996, R. Boender 744 (holotype QCA; isotype M). Species haec in subg. Astrophea pertinens; liana; ubique ovario excepto glaberrima; petioli 0,9-1,9 cm longi; laminae foliorum integrae, oblongae, oblongae-ovatae, 7,5-30 cm longae, 3,5-10 cm latae; flores fasciculati, 6-7,5 cm diametro; coronae filamenta 5-seriata, filamentis exterioribus 2—2,8 cm longis; operculum tubulosum, erectum, supra medium irreguliter laciniatum. Passiflora callistemma, P. maguirei et P. pittieri affinis sunt. Treelet turning into woody vine up to 9 m tall, with well developed tendrils on younger stems, glabrous throughout with exception of ovary and styles; stem terete, brown, younger parts green; stipules setaceous, 0.1-0.2 cm long, soon deciduous; petioles 0.9-1.5(-1.9) cm long, 2-glandular at junction to midrib of blade, with sessile oval nectaries ca. 0.2 cm long and 0.1 cm wide; leaves entire, oblong to oblong-ovate, (7.5-)12.5-25(-35) cm long, (3.5-)5-9(-13.5) cm wide, acute or abruptly acuminate at apex, cuneate or rounded at base, with 9-12 pairs of major lateral veins, subcoriaceous, drying olive-green; peduncles grouped in dense fascicles on old stems, 0.5-0.7 cm long; bracts subulate, 0.2-0.3 cm long; flowers 3.5-5.5 cm long, 6-7.5 cm in diameter, white with rose to purplish-brown and yellow corona; floral stipe 0.4-0.6 cm long, up to 0.3 cm in diameter; floral tube (hypanthium) cylindrical to campanulate-cylindrical, 0.8-1.2 cm long, 1-1.5 cm wide at apex, tapering towards base; sepals oblong, 2.9-4.1 cm long, 0.8-1.3 cm wide, outside green with fine darker lines, inside white tinged greenish on the edges, obtuse, fleshy; petals oblong, nearly as long as the sepals, 1-1.5 cm wide, white, membranaceous; corona in 5 series, outer series curved, forming a campanulate structure around the androgynophore, filaments of outer series (44-)47-53, laterally compressed, wavy, creme at base, purplish-orange to purplish-brown above middle and yellow at the apex, (2-)2.3-2.8 cm long, 0.2 cm wide at base, abruptly dilated in upper third, 0.4 cm wide, tapering to apex, ca. 0.1 cm wide, filaments of second series laterally com- pressed, 0.9-1.3 cm long, dilated up to 0.2 cm near apex, creme to yellow at base, tinged more and more purple above middle, those of third series laterally compressed, 0.6-0.8 cm long, about 0.1 cm wide, purplish-orange to purplish-brown, those of fourth series filiform, about Icm long, inserted inside the hypanthium, curled in lower third, erect in upper half, those of inner series narrowly linear, about 0.5 cm long, placed 0.3 cm above base of oper- culum; operculum inserted slightly above middle of hypanthium, tubular, erect, exserted above apex of hypanthium, irregulary cleft in upper half, 1-1.5 cm long, reddish; andro- gynophore 3-3.5 cm long, reddish; ovary ovoid, ca. 0.5 cm long, ca. 0.3 cm wide, densely pubescent; styles ca. 1 cm long and 0.2 cm wide, densely pubescent; fruit unknown. Additional collection examined: Ecuador. Prov. Esmeraldas: Bilsa Reserve, Green Trail, Station 35, 559 m, 6 Aug 1998, D.D. Kapan & S. Delgados 125 (TEX). Taxonomic remarks Passiflora tina belongs to the subgenus Astrophea, but it does not fit well into any of the six sections given by KILLIP (1938) and taken on by ESCOBAR (1994). The new species is intermediate between the sections Dolichostemma Killip (exserted, tubular operculum), Pseudoastrophea (Harms) Killip (morphology of the flowers, especially the corona) and Botryastrophea (Harms) Killip (flowers in dense fascicles, cauliflorous) and thus questions the current classification. KILLIP (1948) mentioned the same problems with P. maguirei, which he placed next to P. deficiens Mast. of section Pseudoastrophea. ESCOBAR (1994) described P. callistemma a species characterized by its white flowers in dense fascicles, a short hypanthium and an exserted operculum. She indicated that P. callistemma closely resembles P. ovata Martin ex DC of section Pseudoastrophea, which also has an tubular operculum. There is at least one more species of subgenus Astrophea, P. haughtii Killip, which has a tubular operculum around the androgynophore with its margin above apex of hypanthium. On the basis of these findings, it seems to be necessary to question whether the long, tubular operculum should be regarded as a distinctive mark of section Dolichostemma. How- ever, further studies are required to establish whether the current circumscription the sections are monophyletic entities. We are anxiously looking forward to the revision of this subgenus begun by Escobar and to be completed by Hansen in the near future. Passiflora tina is closely related to P. callistemma from northwestern Colombia, P. ma- guirei from British Guiana, Brazil and probably Venezuela, and the widespread P. pittieri which is known from Belize, Costa Rica, Panama, Colombia and Venezuela. JORGENSEN & MACDOUGAL (1997) mentioned that P. pittieri occurs also in Ecuador. Using ESCOBAR’s key to subgenus Astrophea (1994) it key out as P. callistemma. In the Flora of Ecuador HOLM-NIELSEN et. al. (1988) it does not fit in the key of subgenus Astrophea, because only four species not resembling P. tina were known. P. tina differs from P. callistemma and P. maguirei by having 5 corona series, a wider hypanthium and strongly dilated corona filaments of outer, second and third series. From P. pittieri, the new species can be distinguished by its flowers borne in dense fascicles, shorter peduncles, the wavy outer corona series which is forming a campanulate structure around the androgynophore, and the higher number of the strongly dilated outer corona filaments. The differences between the four species are summarized in Table 1. Etymology The epithet honors Tina Garzon who was born in Odessa. She fled the Russian Revolution in the early part of the 20th century and built the Tinalandia hotel located next to a small forest and an adjoining tract which are about the only forest left in this part of Ecuador. Tina recently died at about 88. We thank Dr Thomas Emmel at the University of Florida for introducing the senior author to the country of Ecuador and his letters of support. We thank Dr Lawrence Gilbert at the University of Texas for the inspiration to study Passifloras and Heliconius butterflies. We thank John MacDougal at the Missouri Botanical Gardens for the constant encouragement to keep going in our search and promotion of this plant family. We thank Dr Alberto Padilla and Dr William Patricio Ponce at the Catholic University in Quito, Ecuador for their invaluable help in sponsoring and obtaining permits to bring this material back for more detailed study. We thank the people at INEFAN for granting the permits that allowed us to study their natural resources. We thank Dr Durrell Kapan of the University of Texas for his helpful information and photographs of P. tina from Bilsa Reserve location. Finally we are grateful to Dr Peter Moller Jorgensen for his constructive criticism on this manuscript. References BENSEN, W.W., BROWN, K.S.Jr. & GILBERT, L.E. 1976: Coevolution of plants and herbi- vores: passion flower butterflies. — Evolution 29: 659-680. BOENDER, R. & EMMEL, T. (in press): The Heliconius butterflies of Western Ecuador and their associated Passifloraceae foodplants in Pichincha Province. Association of tropical Lepidoptera. ESCOBAR, L.K. 1994: Two new species and a key to Passiflora subg. Astrophea. — Systematic Botany 19(2): 203-210. HOLM-NIELSEN, L.B., JORGENSEN, P.M. & LAWESSON, J.E. 1988: Passifloraceae. — In: HARLING, G: & ANDERSON, L. (eds.): Flora of Ecuador 31. Kopenhagen. JORGENSEN, P.M. & MACDOUGAL, J.M. 1997: Three new species of Passiflora (Passi- floraceae) from Ecuador and Notes on Passiflora viridescens. — Novon 7: 379-386. KILLIP, E.P. 1938: The American species of Passifloraceae. — Publ. Field. Mus. Nat. Hist., Bot. Ser. 19: 1-612. — 1948: Passiflora maguirei. — Bull. Torr. Bot. Club 75: 415. — 1960: Supplemental notes on the American species of Passifloraceae with descriptions of new species. — Contr. U.S. Natl. Herb. 35: 1-23. Table 1. Comparison of Passiflora tina and its closest relatives. reits VB callsene Be Number of flowers/node Length of pe- duncles including floral stipe Hypanthium paired or grouped in fascicles, 2-8 flowers/node cylindrical, ca. 0.9 cm long, ca. 0.6 cm wide grouped in dense fascicles, about 5 flowers/node funnel shaped, 1.5-2 cm long, ca. 0.8 cm wide peduncles soli- tary, often bifur- cate, usually 1-2 flowers/node cylindric-campa- nulate, 0.8—1 cm long, 0.6—0.7 cm wide grouped in dense fascicles, 4-10 flowers/node 0.9-1.3 cm cylindrical to cam- panulate-cylindri- cal, 0.8-1.2 cm long, 1-1.5 cm wide at apex Number 4 4 4-5 5 of corona series Outer corona se- ries Number, length and form of outer corona filaments Filaments of second series Operculum wavy, surrounding the androgyno- phore in a campa- nulate fashion Ca) 69,.ca.3icm long, very slender ca. 0.5 cm long, dilated at apex ca. | cm long probably not wavy, probably more or less flat number unknown, 2-2,2 cm long, linear-dolabri- form, 0.2 cm at widest point ca. 1 cm long, narrowly linear ca. 1.2 cm long, deeply cleft in upper half not wavy, more or less flat 28-37(-42), 1.5—2 cm long, sub-dolabriform, 0.2 cm at widest point 0.5-0.8 cm long, dilated, 0.1 cm wide at apex ca. 1 cm long, minutely denticu- late wavy, SUITOUN- ding the andro- gynophore in a campanulate fashion (44-)47-53, (2-)2.3-2.8 cm long, abruptly dilated in upper third, up to 0.4 cm at widest point 0.9-1.3 cm long, dilated up to 0.2 cm near apex 1-1.5 cm long, deeply cleft in upper half Ronald BOENDER, Butterfly World, 3600 West Sample Road, Coconut Creek, FL 33073, U.S.A. Torsten ULMER, Universität Essen, Fachbereich 9/Botanik, 45117 Essen, Germany. 10 Figure 1: a: Flowers of Passiflora tina R.Boender & T.Ulmer grouped in dense fascicles (Boender 744, cultivated); b: Flower, close up showing the corona series; c: Heliconius sapho candidus Brown. Figure 2: Habit of Passiflora tina. \ 2cm Figure 3: Schematic detail of longitudinal section through the flower of Passiflora tina. 13 Achene hairs and their diversity in the genus Chaetanthera Ruiz & Pav. (Mutisieae, Asteraceae) A.M.R. DAVIES & E. FACHER Summary: DAVIES, A.M.R. & FACHER, E.: Achene hairs and their diversity in the genus Chaet- anthera Ruiz & Pav. (Mutisieae, Asteraceae). — Sendtnera 7: 13-33. 2001. ISSN 0944-0178. A detailed and comprehensive survey of the achenes belonging to 35 of the 42 accepted species of the South American genus Chaetanthera Ruiz & Pav. revealed new micro-morphological characters. In particular the achene hairs yielded important and taxonomically significant characters. Three achene hair types were distinguished: “Zwillingshaare” (in various forms), filiform hairs, and single-celled papillae. Glabrous achenes were also recorded; the lack of achene hairs was species specific and, not unexpectedly, was not a useful discriminator for bigger groups. Within a species the hair types were consistent. The diversity of hair types and their distribution throughout the genus was compared to the traditional sub-generic structure proposed by CABRERA (1937). The achenes of those taxa belonging to the current subgenera Proselia, Eu- chaetanthera, Tylloma and Glandulosa are never glabrous and have very similar “Zwillingshaare” (90-120 um long, obovate to elliptic or lanceolate in outline). Achenes of the subgenus Oriastrum show consistent variations of spherical or conical- deltoid “Zwillingshaare” (10-50 um in length) while achenes of taxa in the subgenus Carmelita are glabrous or have tiny (7-15 um in length) spherical “Zwillingshaare”. Achenes of the subgenus Egania are mostly glabrous, with a couple of taxa showing single-celled papillae thereon, and two taxa long filamentous (but not “Zwillings- haare”) hairs (approx. 450 um in length). Based on the hair types, clear divisions within the genus can be seen. However these need to be supported by correlating characters. A revision of the genus Chaetanthera is currently in progress. Zusammenfassung: Eine detaillierte und umfassende Untersuchung der Achänen von insgesamt 35 der 42 akzeptierten Arten der südamerikanischen Gattung Chaetanthera Ruiz & Pav. zeigt neue mikromorphologische Merkmale auf. Insbesondere die Achänenhaare liefern wichtige und für die Taxonomie verwertbare Merkmale. Drei grundlegende Achänen- haartypen können unterschieden werden: Zwillingshaare (in verschiedenen Formen), filiforme Haare und einzellige Papillen. Relativ häufig treten kahle Achänen auf; das Fehlen von Haaren ist artspezifisch, faßt jedoch, wie zu erwarten, keine größeren Gruppen zusammen. Innerhalb einer Art sind die auftretenden Haartypen einheitlich. Die Verteilung der Haartypen innerhalb der Gattung wird mit der von CABRERA (1937) vorgeschlagenen Unterteilung verglichen. Dabei ergibt sich folgendes Bild: Bei den in den Untergattungen Proselia, Euchaetanthera, Tylloma und Glandulosa zusammen- gefaßten Arten sind die Achänen nie kahl und weisen große Ähnlichkeit in den 14 Zwillingshaaren auf: Elliptisch oder lanzettförmig mit einer Länge von 90-120 um. Die Arten der Untergattung Oriastrum tragen auf den Achänen kugel- oder kegel- bis deltaförmige Zwillingshaare mit einer Länge von 10-50 um. Die Achänen der Arten der Untergattung Carmelita sind kahl oder besitzen kleine, kugelförmige Zwillings- haare mit einer Länge von 7-15 pm. Die Arten der Untergattung Egania haben über- wiegend kahle Achänen mit Ausnahme von zwei Arten mit einzelligen Papillen auf den Achänen und zwei weiteren Arten mit filiformen, bis zu 450 um langen Haaren (keine Zwillingshaare). Aus diesen Ergebnissen kann der Schluß gezogen werden, daß aufgrund der auf den Achänen gefundenen Haartypen eine deutlichere Gliederung der Gattung vorgenommen werden kann. Es ist jedoch notwendig, diese durch zusätzliche Merkmale zu stützen. Eine komplette Revision der Gattung Chaetanthera ist in Bear- beitung. Introduction Achene characters are often used to taxonomic advantage in many genera of the Compositae, (OBERPRIELER 1998, VINCENT 1996, ROMMEL 1979). Of particular interest is the orna- mentation of the pericarp epidermal cells — that is the surface characters of the fruit bodies. It has been said that while using ultrastructure features can be useful in establishing concepts, they cannot be used as key characters (KING & ROBINSON, 1970). Nevertheless micro- morphological characters are often studied as generic markers in the Asteraceae. In some cases they are found to obscure generic relationships (Astereae, SUNDBERG, 1985), but equally so, micro-morphological characters can be both useful and consistent markers for generic delimitation (LANE 1985, SCOTT 1985). |. Infra-generic taxonomy The predominantly Chilean genus Chaetanthera, first described in 1794 by Ruiz and PAVON, now circumscribes approximately 40 species. The most recent revision by CABRERA (1937) divided the genus into seven subgenera. The defining generic character linking these species is said to be the possession of a winged membrane on the margins of the phyllaries, the outer series of which are foliaceous. The subgenera are distinguished by characters including capitulum size, ornamentation of the apices of the inner series of phyllaries, style arm dimensions, leaf shape, habit, life cycle and achene pubescence (after CABRERA, 1937). Table 1 below summarises the subgenera structure of Chaetanthera proposed by Cabrera and the species included in this study. Since the 1937 revision was published five new taxa have been described: Chaetanthera chiquianensis Ferreyra, C. boliviensis J.Kost., C. aymarae Martic. & Quezada, C. lepto- cephala Cabrera, and C. perpusilla (Wedd.) Anderb. & Freire (the last solely distinguished from Chaetanthera by the presence of papillose hairs on the achenes). With the exception of C. boliviensis, these are broadly included in the subgenus “Euchaetanthera”. None of these were analysed here, and neither were examples of C. australis Cabrera, C. brachylepis Phil. or C. flabellifolia Cabrera. 2. Inter-generic taxonomy Chaetanthera has two putative sister genera — Pachylaena D.Don (distinguished by having a plumose pappus) and Brachyclados Hook. & Arn. (distinguished by having non-foliaceous external phyllaries, pappus arrayed in 3-4 series and shrubby with + pedunculate capitulae). These two very small genera have not been revised since the 1830s. Examples of Pachylaena atriplicifolia Hook. & Arn., Brachyclados lycioides D.Don and B. megalanthus Speg. were observed with a view to their potential for outgroup comparison. 15 Table 1: Summary of the traditional infra-generic structure sensu CABRERA (1937) of Chaetanthera including only the taxa that were surveyed. (Author names after BRUMMITT & POWELL, 1992) Glandulosa C. glandulosa Remy C. acerosa (Remy) Benth. & Hook.f. C. pulvinata (Phil.) Hauman C. apiculata (Remy) F.Meigen C. revoluta (Phil.) Cabrera Egania C. cochlearifolia (A.Gray) B.L.Rob. C. sphaeroidalis (Reiche) Hicken C. dioica (Remy) B.L.Rob. C. stuebelii Hieron. C. pentacaenoides (Phil.)Hauman C. lycopodioides (Remy) Cabrera C. planiseta Cabrera Oriastrum C. gnaphalioides (Remy) I.M.Johnst. C. pusilla (D.Don) Hook. & Arn. C. minuta (Phil.) Cabrera : C. villosa D.Don C. spathulifolia Cabrera C. lanata (Phil.) I.M.Johnst. C. glabrata (DC.) F.Meigen C. renifolia (Remy) Cabrera C. limbata (D.Don) Less. C. splendens (Remy) B.L.Rob. C. elegans Phil. C. serrata Ruiz & Pav. Euchaetanthera C. chilensis (Willd.) DC. C. valdiviana Phil. C. ciliata Ruiz & Pav. C. microphylla (Cass.) Hook. & Arn. C. euphrasioides (DC.) F.Meigen C. moenchioides Less. C. flabellata D.Don C. peruviana A.Gray C. incana Poepp. C. tenella Less. C. linearis Poepp. 3. Achene hairs in the Compositae — morphology and importance The description and publication concerning the hairs seen on the fruits of the Compositae has been accepted for many years. Although typified by HANAUSEK (1910), HEss (1938) wrote the seminal paper on the typical trichomes unique to the Compositae. They are named “Zwillingshaare” (twin hairs). Achenes and often their hairs have frequently proven to be fundamentally useful indicators of generic and subgeneric relationships (Felicia, GRAU 1973; Amellus, ROMMEL 1979; Grangea, Grauanthus and Dichrocephala, FAYED 1979; Gerbera-complex, JEFFREY 1967, HANSEN 1989; Anthemis L., OBERPRIELER 1998), or even tribal distinctions (DITTRICH, 1985, 1996) in the Astereae. VELEZ (1981) demonstrated that some genera (of the American Asteraceae) have unique hairs, while in other cases several genera can share one hair type. One advantage of using achene characters, and in particular the hairs, is that they are considered to be independent of environment (Grau, pers. comm.). Therefore the variation expressed in these characters forms a source of reliable taxonomic indicators. The presence or absence of achene hairs has been pivotal to the keying out of the seven subgenera of Chaetanthera in the past. CABRERA (1937) recognised two achene hair types - papillose and seriaceous-villous, as well as glabrous achenes. HANSEN (1991) noted the presence of both hairy and glabrous achenes in the genus, the former carrying either of two hair types: inflated and tapering, or orbicular. The aim of this study was to establish on a firm basis the existing variation in the achenes and whether this variation might have taxonomic implications. Methods Over 650 specimens of the genus Chaetanthera were received from the herbaria of GH, K, M, NY, P, W (abbreviated after HOLMGREN et al., 1990) and the private collections of Prof. J. Grau and Dr C. Ehrhart. Nearly 200 collections of 35 Chaetanthera species (covering all seven subgenera), 1 species of Pachylaena, and 2 of Brachyclados were sampled. Citations for typical material observed can be found in Appendix 1. Achenes from both ray and disc flowers were carefully extracted from the herbarium material and placed directly, without further treatment on carbon-coated stubs. These were then sputtered with platinum using a Bal-tec sputterer. Digital images were taken under a high vacuum using a LEO 483VP Scanning Electron Microscope (SEM) at both low and high magnifications. Using these images the achenes were assessed for 51 characters covering 6 quantitative measures and 45 features including achene shape, achene surface, achene hair shape and construction, and 5 pappus characters. Results 1. SEM observations and images During this study three factors indirectly related to the research became apparent. These were the effect of collection age on sample quality, the consistency of achene surface structures relative to maturity and, perhaps most importantly, the SEM focus error. 1.1. Specimen age The age of the herbarium specimens, with only few exceptions, did not contribute to, or detract from, the quality of achene hair preservation. One of the more distressing features of some specimens was the abundance of hyphal growth obscuring the characters of interest. The disintegration of some features has been attributed to environmental conditions at the time of collection. Only one taxon showed a tendency to have fused achenes — C. villosa. 1.2. Mature and immature or sterile achenes The maturity of the achenes is a difficult parameter to quantify without precise glasshouse surveys. However, casual observation shows that the maturity and fecundity of the achenes particularly affects the size and shape of the achenes, but does not seem to alter the presence, or type, of hairs. As seen in one collection, an achene can increase in length by 30% and in width by 60% as it matures, while the changes in hair dimensions are negligible. 1.3. Focus error Unusually for SEM observations, the total size of the material samples was sometimes quite large (> 4.00 mm). As a result, it was not always possible to have the complete object entirely in focus, affecting the apparent length of the object. The mean error for large achenes over 2.35 mm in length was calculated at 2.4% of the length. The greatest error was measured for the largest achenes, but rapidly tails off as the general size of the achene shrinks, and the error becomes almost negligible for achene lengths below 2.35 mm (0.4%). 2. Within the genus Chaetanthera — quantitative characters Six quantitative characters were observed. These were achene length and width, achene apex width, achene base width, and achene hair length and width. The measurements of the various dimensions were taken using the guidelines shown in Plate 1 (Figs. 1-2). The Logo values of each character were plotted in rank order, with the graphs drawn to the same scale so that each character could be easily compared (see Graphs 1-4). Fig. 1: Location from which the quantitative measurements on the whole achene were taken. Achene Length: The length of the achene taken from the longest axis from the pappus insertion Achene Apex Width (apex) to (but not including) the carpopodium (base). Pappus Achene Width Achene Width: taken from the widest point (perpendicular to the long axis) of the fruit. Achene Apex Width: taken from below the pappus insertion. Achene Base Width: taken from the where the carpopodium and fruit meet. When no carpopodium is present, the base width is taken from the homologous region. Ss = © 4 o = oO a oS < Achene Base Width Carpopodium Fig. 2: Location from which the quantitative measurements on the achene hairs were taken. Achene Hair Length: the length is taken parallel to the central cell wall of the 'twin hairs', always from the point where the hair is inserted in the Achene pericarp epidermis. Hair Width Achene Hair Width: taken from the widest point across the hair perpendicular to the long axis. N.B. This figure is adapted from an image of a ‘twin hair' of the species C. linearis. Achene Hair Length Plate 1: Images showing measuring planes used to collect the qualitative information from the achenes and their hairs (when present). 18 2.1. Achene length and width: Achene length varies from 0.5 mm (C. pusilla) to 4.8 mm (C. lanata). The achene widths vary from 0.04 mm (C. linearis, immature) to 1.51 mm (C. villosa). Achene lengths and widths are continuously distributed in this sample (Graph 1). 2.2. Achene apex and base widths: The apex widths vary from 0.13 mm (C. linearis) to 1.17 mm (C. spathulata). Achene base widths vary from 0.07 mm (C. gnaphalioides) to 0.79 mm (C. villosa). The measurements are continuously distributed in this sample (Graph 2). Although there are no discrete groupings, there is still a significant variation by over a factor of ten in achene length and width as well as achene apex and base widths across the data set. In all four cases the linear distribution (see Graphs 1-2) is approximately y = 0.002x, indi- cating that each quantitative character exhibits similar variation. Thus we deduce that all of these four characters (achene length and width, and achene apex and base width) are equally useful for analysing the data, even though they do not show clear evidence of clustering. D Achene Length (upper) — | | © Achene Apex Width (upper) | + Achene Width (lower) | 1% _* Achene Base Width (lower) | 0,5 oo ob = S Se = ——— 0 100 200 300 400 0 100 200 300 400 Ranked Order of Values Ranked Order of Values Graph 1: Ranked Order of Log (Achene Graph 2: Ranked Order of Log (Achene Length) & Log (Achene Width) Apex Width & Achene Base Width) 2.3. Achene hair length and width: When hairs are present on the achenes they can vary from 10 um to over 450 um in length. The hairs vary in width from 5 um to 75 um. Graphs 3 and 4 have comparatively less smooth, flatter curves, with much bigger tails, particularly at the lower end. They both vary in the same way, the flat part indicating that for about half the achenes, the size of achene hairs is not a good discriminator. 2.901 2 Log (Hair Length) Log (Hair Width) Big I 0 100 200 300 400 0 100 200 300 400 Ranked Order of Achenes Ranked Order of Achenes Graph 3: Ranked Order of Log (Achene Graph 4: Ranked Order of Log (Achene Hair Length) Hair Width) 19 However, the dramatic tails on the graphs indicate that for large and small extremes hair size, particularly length, becomes an increasingly valuable discriminator. Plotting a linear regression on these curves would not be useful, as it would only apply to about half the sample. Quantitative characters like these that indicate some clustering, together with those which show less clear patterns of segregation, can be combined to be more informative. We have plotted the Logo values of achene lengths and achene hair lengths against each other (Graph 5). A third qualitative character, achene hair shape, is also represented on the plot. Glabrous achenes are included for illustrative purposes. The various hair types (see qualitative characters) are represented by different symbols. We can see that the 4 hair types have broad hair length distributions. Three obvious bands of hair lengths are seen which correlate with the following hair shapes; filiform, globular and obovate-lanceolate. The glabrous achenes show a similar broad distribution along the x-axis. Graph 5 indicates that for nearly the same wide range of achene lengths it is possible to have hairs or no hairs. When present the length of these hairs is really tightly associated with their shape. Thus globular hairs are always, in this data set, much shorter in length than any of the obovate-lanceolate hairs. These are in turn always shorter than the filiform hairs. Graph 5 clearly demonstrates that achene hair length and shape are, to some extent, correlated. Furthermore, the hair shape allows the data set to be divided into discrete clusters that were not apparent from the hair length distribution alone. 3. Within the genus Chaetanthera — qualitative characters Qualitative characters of the achenes included shape, cross-section, apical morphology, and surface ornamentation and, for the achene hairs, types, orientation and features. Achene shape varied from pyriform, turbinate, fusiform to oblong, and in cross-section from compressed, terete, semi-terete or triangular. The angles in cross-section can be rounded and shallow, or prominent and sharply angular. The pericarp epidermal cells were classed as parenchymatous, subisodiametric, papillate, striate, centrally depressed or having no surface sculpture. The achene hairs when present were 1-, or 2-celled (parallel or stacked) and displayed a range of shapes (after STEARN, 1995) lanceolate or elliptic, to very small globular, conical or ampulliform hairs, long filiform hairs (RAMAYYA, 1962), or papillae (METCALFE & CHALK, 1979). The tips varied from obtuse or acute and mucronate or apiculate, sometimes shortly bi- furcated, rarely depressed. They were variously found to be erect to appressed, and either deflated or inflated (symmetrically or asymmetrically) about the horizontal axis. They covered the achenes sparsely to densely. Achene hairs were recorded in 30 different species. Within these, three basic hair types were identified: “Zwillingshaare”, filiform hairs and single-celled papillae. The variation is outlined below and the species placements are given in Table 3. The achene hairs were found to be mostly consistent in both shape and size within species (i.e. between collections). 3.1. “Zwillingshaare” 3.1.1. Obovate elliptic to lanceolate or rarely naviculate These hairs are large, varying in length from (80)-90-120-(130) um. This hair type is associated with a relatively featureless epidermal surface. The 17 species with these sorts of hairs (C. chilensis, C. ciliata, C. elegans, C. glabrata, C. glandulosa, C. incana, C. lanata, C. limbata, C. linearis, C. microphylla, C. moenchioides, C. peruviana, C. renifolia, C. ser- rata, C. splendens, C. tenella, and C. valdiviana) generally have hairs on both the disc and ray achenes, and are very rarely nearly glabrous. This is the most common hair type in Chaet- anthera and it shows so much variation that it is seldom possible to distinguish collections of species without practice. The extremes of variation in this group of hairs are illustrated with images that can be seen in Plate 2 (Figs. 3-10). 20 ‘Kay ut sjoquiAs 0} FupI090e poyouap sadAq Irey Jusyde JusIsyyIp YIM ‘(yI3U9T Are Susyoy) 307] ysuredse (Y3U97] susyoy) 307 :¢ ydeip (y3u97 ausyay) 307 8°0 9°0 r0 To 0 CO v0 00K «3K — 000 0K «25 KEITH EEK X —3¢ x ! ro ® ee = 3 2 2 ° oo 2 0 2 > 2 co. > o ° ° ° ° ° ° o ° ° = x ° E 2 oFee o ° ° © ° ° ‘ 3 oo =: ° ° o AS || ° ; = 5 ° : o° : One ° °° o ° fr S00 = v va a ee. x 4 ea = % v Ki s = v viv «wy v N ww wy v x v og Ey, v7” y, "Cav te, Vor, „v wo“ & RL ee eae E Vv v v v v snolge]3 x PO]]99-9]3UIS @ + re [Se n ER eet eh oyeJooue[-2JeAOgO v JengoJ3 o LO + 21 3.1.2. Spherical (to rarely conical) Seven species fall into this category. The hairs are smaller but consistently spherical, vary in length from 8-50 um, and can be subdivided into 3 sections. In C. planiseta and C. pusilla (section 3.1.2.a.) the hairs are found on both disc and ray achenes and, while often deflated to some extent, have a distinctly thickened cell wall. The length of these hairs varies from 35-50 um (Plate 3, Fig. 15). Section 3.1.2.b. includes three taxa (C. euphrasioides, C. flabellata and C. lycopodioides). The hairs have an indistinct dividing cell wall, in some cases tending towards conical rather than spherical in outline, and in one species the ray achenes are often glabrous The hair length measures between 20-40 um (Plate 3, Figs. 11-14). Lastly C. lanata and C. spathulifolia, distinguished as section 3.1.2.c., possess spherical hairs that vary from 7-15 um and are sparsely distributed over the surface. They are however frequently poorly preserved. In these cases the achenes can appear to be glabrous (Plate 3, Fig. 16). 3.1.3. Ampulliform to nearly deltoid Only two species belong to this group - C. gnaphalioides and C. minuta. These hairs can be found with an occasional recurved, rostrate apical tip and may or may not appear to be collapsed. The hairs vary in length from 7-20 um. These images can be seen in Plate 3, Figs. 17-18. 3.2. Filiform hairs Only two species (C. apiculata and C. acerosa) possessed these hairs. The long, filamentous, seriaceous-villous hairs are constructed of two vertically stacked cells; one short basal cell, and the second, elongated apical cell. They have an approximate length of 450 um. Some collections of the two species can be completely glabrous. No correlation to disc and ray distribution was observed. Filiform hairs confined to a sparse ring near the pappus insertion were only rarely recorded (two samples, not C. apiculata and C. acerosa). Their cell con- struction remains unclear, but it seems most likely that they are an artefact of the pappus morphology rather than an achene epidermal character. Plate 4, Fig. 23. illustrates the filiform hairs. 3.3. Single-celled papillae Single-celled papillae were only found in a couple of species (C. pulvinata and C. revoluta), the oblate flattened hairs densely covering the achenes, and with a mean length between 25-45 um. These images can be seen in Plate 4, Figs. 21-22. 3.4. Glabrous achenes within the genus Chaetanthera Twelve Chaetanthera species had glabrous achenes. Half of these were consistently glabrous both within and between capitula of the same and different collections. These were C. cochle- arifolia, C. dioica, C. pentacaenoides, C. pulvinata, C. sphaeroidalis, and C. stuebelii. These achenes had the most interesting epidermal surfaces; for example the image shown on Plate 4, Fig. 19. Further, these taxa are arranged in the same sub-genus “Zgania”. In six taxa (C. ace- rosa, C. apiculata, C. lanata, C. minuta, C. revoluta, and C. villosa) combinations of both glabrous and pubescent achenes in collections of the same species were observed. These did not follow any set pattern, nor do the species concerned come from the same group of taxa. The epidermal surfaces in these cases were relatively featureless (Plate 4, Fig. 20.). The varying combinations of glabrous and pubescent achenes within and between collections of the same taxa are presented in Table 2. 22 Table 2: Table summarising combinations of glabrous (-) and pubescent (+) achenes in collections of the same species. 3 (C. acerosa, C. lanata, C. minuta) + + 2 (C. villosa, C. revoluta) [Notobserved - | Not observed + (rare) + SSE ae 4. Between the genera — Pachylaena and Brachyclados Pachylaena and Brachyclados were also studied. Pachylaena atriplicifolia was represented by 2 collections. The first had only glabrous achenes, while the second possessed small (20-35 um L.) lozenge-shaped achene hairs — again, twin-celled (see Table 3, section 3.1.4.). They were however, sparsely distributed, and rather poorly preserved. Brachyclados lycioides material possessed both long (120-175-(200) um in length) oblate- lanceolate “Zwillingshaare” (section 3.1.5.), and shorter (25—45 um in length) globular hairs (Table 3, section 3.1.2.a.) B. megalanthus only had the former hair type, but marginally smaller in length (80-130 um). The results are pictured in Plate 4, Figs. 24-26. + 1 (€. apiculata) 5. Summary of results The information in Table 3 presents a summarised view of the achene hair types, which species carry these hair types, and to which subgenus these species are traditionally ascribed. The section numbers given in Table 3 correspond mostly to the section numbers in the text. Table 3: Summary table showing achene hair type and the species (and subgenera) which carry these types. (*indicates which species are illustrated in this paper) Hair Type ie ee Taxon Names - Subgenus [Species] 3.1. “Zwillingshaare” Proselia [C. chilensis*, C. elegans, C. serrata, C. valdiviana] Euchaetanthera |C. ciliata*, C. incana*, C. linearis, C. micro- phylla*, C. moenchioides*, C. peruviana, C. tenella} Tylloma [C. glabrata, C. limbata*, C. renifolia*, C. splendens] Glandulosa [C. glandulosa*] Carmelita [C. lanata (?)] 2. Oriastrum [C. planiseta, C. pusilla*] Brachyclados lycioides Cu Oriastrum [C. lycopodioides* | Euchaetanthera [C. euphrasioides*, C. flabellata* | 3.2. Filiform 3.3. Papillose 3.4. Glabrous Egania [C. acerosa*, C. apiculata] Egania [C. pulvinata*, C. revoluta*] Egania [C. acerosa*, C. apiculata, C. cochlearifolia, C. dioica, C. pentacaenoides, C. pulvinata, C. revoluta, C. sphaeroidalis*, C. stuebelii] Carmelita [C. lanata, C. villosa] Oriastrum | C. minuta] P. atriplicifolia 23 Fig. 3: bar 20um Fig. 4: bar 10um 7: bar lOum Fig. 9: bar 20um Be Fig. 10: bar 10um Plate 2: 3: C. moenchioides Less., Pirion 492 (GH); 4: C. ciliata Ruiz & Pav., Rosas 1890 (M); 5: C. microphylla (Cass.) Hook. & Arn., Wagenknecht 18489 (GH); 6: C. chilensis (Willd.) DC., Elliott 292 (K); 7: C. glandulosa Remy, Rosas 1592 (M); 8: C. limbata (D.Don) Less., Morrison 16740 (GH); 9: C. incana Poepp., Jiles 5001 (M) ; 10: C. renifolia (Remy) Cabrera, Seibold 2850 (W). 24 11: bar 10um Fig. 13: bar 20um Fig. 14: bar 10um Fig. 16: bar 3um Fig. 17: bar 2um Fig. 18: bar 10um Plate 3: 11: C. euphrasioides (DC.) F.Meigen, Morrison 17071 (GH); 12: C. flabellata D.Don, Philippi s.n. (W); 13: C. flabellata D.Don, Grau 2440 (M); 14: C. lycopodioides (Remy) Cabrera, Werdermann 632 (M); 15: C. pusilla (D.Don) Hook. & Am., Hastings 432 (NY); 16: C. spathulifolia Cabrera, Kiesling, Ulibarri & Krapovikao 7467 (NY); 17: C. gnaphalioides (Remy) I.M.Johnst., Wagenknecht 18590 (GH); 18: C. minuta (Phil.) Cabrera, Castellanos 7 3 Fig. 20: bar 10m Nees Fig. 22: bar 10 um Fig. 24: bar 100 um Fig. 26: bar 30 um Plate 4: 19: C sphaeroidalis (Reiche) Hicken, Werdermann 253 (K); 20: C. acerosa (Remy) Benth. & Hook., Werdermann 627 (M); 21: C. pulvinata (Phil.) Hauman, Vervoorst 3239 (W); 22: C. revoluta (Phil.) Cabrera, Werdermann 1020 (M); 23: C. acerosa (Remy) Benth. & Hook., Ehrhart & Grau 95/818 (M); 24: Brachyclados lycioides D.Don, Jiles 4870 (M); 25: Brachyclados megalanthus Speg., King & Heinz 9390 (M); 26: Pachylaena atriplicifolia Hook. & Arn., Schajovsky s.n. (M). 26 Discussion l. The taxonomic value of achene hairs above other characters Historically achene hairs have been rather tentatively used in Chaetanthera classifications, even though often used to taxonomic advantage in other Compositae taxa. Now that modern technology has the wherewithal to reveal micro-characters, the true value of these achene variations can be utilised. Many characters, both qualitative and quantitative were considered. The quantitative characters, such as achene length, did not show useful, discrete discontinuities. Given the large amount of variation they could, however, form a useful contribution to a multivariate analysis of a larger data set. Qualitative characters such as achene shape were found to be unreliable because of fecundity, maturity etc. The achene hairs appeared to be independent of these factors. The shape and size, especially length, was found to be statistically and visually a good discriminator for taxa in this genus. The absence of hairs, or the glabrous state, is not a reliable taxonomic indicator above the species level. 2. Achene hair diversity in the context of subgeneric divisions of Chaetanthera The information regarding achene hair type distribution across the genus is summarised in Table 3. The table shows that clusters of species, traditionally placed in the same subgenera possess similar achene hairs. The patterns of hair type and subgenus fall into one of two categories: e one hair type groups several subgenera, and e individual subgenera group several hair types. This is also illustrated in Plate 5. The hairs are schematically represented (to scale) and accompanied by transverse sections through the hairs, showing hair cell-wall variations, hair width ranges, and hair cell-number. 2.1. One hair type — several subgenera. This grouping represents those species and even subgenera whose achene characters group them together within Chaetanthera. The taxa in the subgenera Proselia, Euchaetanthera, Tylloma, and Glandulosa are virtually inseparable on the basis of their achene features. With only two exceptions (C. euphrasioides and C. flabellata) the species traditionally placed within these four subgenera share the same type of achene hairs (elliptic or obovate to lanceolate “Zwillingshaare”) on both disc and ray achenes and, within a broader sense, have similar sized achenes. There is also little variation in the achene characters among the subgenera themselves. This lack of diversity itself is distinctive. They are quite separate from the remaining taxa in the study. The taxa in these four subgenera, in particular within Proselia and Euchaetanthera, are very closely related and often express a wide range of morphological variation. This is reflected in the taxonomy where examples of taxa with several varieties are described (e.g. C. chilensis), as well as suites of species that are hard to distinguish from each other (e.g. C. microphylla and C. linearis). 2.2. One subgenus — several hair types. The taxa in this grouping pose more of a challenge to interpretation, and in effect represent the taxa that are least well-collected and tricky to sample. Three subgenera fall into this group. The first two, Egania and Oriastrum show 2-3 hair types, each type found in only 2 or 3 species. The third, Carmelita, forms somewhat of an outlier as it, for the most part, exhibits only one hair type. However, there were a couple of oddities that require further investigation. 127 BETT RIAI[Og ee eR CE ) 2 [oıy v7] un a er (y) esseureyey ME 4.) Lemon]. EL (v) Byes BEE (\) Antone a ©) Bote SOT ee (J) elueonely eI XI — (D) of OLE 19P TITA | | | (9) arme [op TIA === Se (9) SUISSIH.O IA _ EE 5) 5103 0 BD er me] (dD) osresedjeA A | | | | nn 1027" | gen) (J) eureseyy III | Det | = (9) esedese] | va Ad) > un un | mn Ss o © a 9 un un | u oo _ || Io ©|| O0 2 & 8 ae) RE om | | 4 oS co \© x u01331 y9e9 ur sa19adg Jo JOqUINN a Regions of Chile (C) & Argentina (A), Peru & Bolivia Graph 6: Distribution of Chaetanthera species with Lanceolate ("L") and Glabrous or Globular ("G") achene hairs over South America 28 2.2.1. Nine species are traditionally grouped into the subgenus Egania. United mostly by the possession of interior phyllaries with coloured apical scales and having buds on the lower stem regions, these taxa represent the most under-collected species in the genus, and are consequently the most poorly sampled taxa in this study. Two hair types were revealed: long filamentous hairs and papillae. This was the only subgenus in which some species had con- sistently glabrous achenes. These hair types are not duplicated elsewhere in the genus Chaetanthera. Five of the species had consistently glabrous surfaces on both disc and ray achenes. This is not a very exciting taxonomic feature, and certainly not a good discriminator considering the propensity of glabrous achenes to appear in amongst otherwise pubescent achene collections. As an aspect of variation this is, however, worth mentioning. The reversal or reduction to the glabrous state is frequent within species of the tribe Mutisieae s.str. (HANSEN, 1991). C. apiculata and C. acerosa were glabrous or had achenes with long filamentous hairs. The distribution of hairy and glabrous achenes in these two species appears to be largely a feature of population variation. A variety of C. acerosa (C. acerosa var. dasycarpa Cabrera) is de- scribed as having long seriaceous-villous hairs on the achenes, as opposed to the glabrous achenes of var. typica. These two varieties are reputed to co-exist in mixed populations. This is a good example where a detailed population survey could determine whether this is really population variation or a true taxonomic difference. The repeated occurrence of filiform hairs on the fruits of the Compositae is documented. These are considered to be plesiomorphic in the Nassauviinae at least (HANSEN, 1989). Uniquely in this study, achenes from collections identified as C. revoluta and C. pulvinata were densely clothed in single-celled deflated papillae. Glabrous achenes were seen in collections of C. pulvinata. 2.2.2. The subgenus Oriastrum includes 5 species (after CABRERA, 1937). Three sorts of “Zwillingshaare” were identified in this group, and are not found in any other taxa of Chaetanthera. C. pusilla and C. planiseta had nearly identical achenes and hairs. The species are morphologically close, separated from each other by phyllary apex colour, and the pappus bristle form. The small spreading floccose species of C. minuta and C. gnaphalioides share distinct achene hairs. There is a slightly different third hair shape in the collections that may indicate an hitherto undescribed taxon. Only C. minuta has both glabrous and pubescent achenes. C. lycopodioides is grouped here with 2 taxa normally included in Euchaetanthera — C. flabellata, and C. euphrasioides. All three species share achene hairs on both disc and ray achenes that are strictly spherical to softly conical (on same achene). The achene is also often a distinct pyriform shape. Currently there is little else to support the grouping of these three taxa. 2.2.3. The three taxa in the subgenus Carmelita are generally very distinct, on account of the individual nature of the leaves and phyllary morphology. C. /anata and C. spathulifolia have very similar but sparse hairs, all occurring in some state of deterioration. C. Janata however also had examples of completely glabrous achenes, like those of C. villosa, and, very rarely, large hairs, as seen in the taxa belonging to Tylloma. These collections of C. Janata are currently under more intensive review. They have not been illustrated in Plate 5 because of the puzzling nature of the result. From this brief discussion regarding the distribution of the achene hairs across the genus Chaetanthera the taxonomic significance of the achene hairs can be seen to vary in different subgenera. The variation in these achene characters raises the question “Is the rank of subgenus the correct rank for the separation of these groups of species?” The informative potential of the geographical distribution of the hair types is considered next. 29 Brachyclados - never glabrous. Oriastrum (incl. C.euphrasioides & C.flabellata) - can be glabrous. () @: =e oO va) Ya on C) dD: = oO =a =) [iS Egania - can be glabrous. il (\ @ dO Carmelita - can be glabrous. = Yoyılaoyıl Proselia, Euchaetanthera, Glandulosa, + Tylloma - never glabrous. Pachylaena - poorly preserved hairs and sparse. Normally glabrous. Plate 5: Different achene hairs and their distribution among the subgenera of Chaetanthera (Euchaetanthera, Proselia, Glandulosa, Tylloma, Oriastrum, Egania and Carmelita) and the two sister genera Brachyclados and Pachylaena. The hair form is schematically represented and additional diagrams of the transverse cross-sections and scale bar of the width are included. 30 3. Achene hairs and their geographical distribution The species in Chaetanthera can be arbitrarily divided into two large groups based on their achene hair type. The species numbers in both groups also include those described taxa interpreted as belonging to the group from a morphological point of view, although no achenes were seen. The distribution of each taxon was collected using primarily the herbarium collection information, but supplemented by literature citations too. The first group comprises all 22 species, in the subgenera Proselia, Euchaetanthera, Tylloma, and Glandulosa, possessing the obovate-lanceolate hair type on both achenes. It is identified by the epithet “L” (this is effectively Table 3, section 3.1.1.). The second group “G” contains 21 species mostly from subgenera Carmelita, Egania and Oriastrum (Table 3, sections 3.1.2., 3.1.3., 3.2., 3.3., and 3.4.). It includes all taxa described as being glabrous or having globular (or filiform or papillose) hairs. A histogram showing the geographical distribution over South America of numbers of species in each group (“L”) and (“G”) is plotted in Graph 6.). This shows us the hotspots of species diversity in each group; “L” is centred in Chilean Regiön Metropolitana de Santiago (RMS), and “G” is centred over the high Andean border between RMS (Chile), and the neighbouring regions of San Juan and Mendoza in Argentina. Furthermore, the “L” group is confined (with the exception of 2 records in San Juan and Neuquen, Argentina) to the west of the Andean mountain range with 2 species exclusive to Peru. The “G” group is confined to the Andes, but spread both to the east and west of the main mountain chain, with over 70% of the species in this group found in both Chile and Argentina. Thus we see two different patterns of distribution for the two artificial species groupings based solely on achene hair type. If we consider the altitude information as well, as a general rule the “L” species are distributed at altitudes ranging from sea level to just over 2000 m. The “G” species are usually to be found above this altitude (2000 m) up to 5000 m. The summary of distribution, altitude and geography is in Table 4. Table 4: Summary of information supporting the 2 groups of achene hairs. Source “L” (= Proselia, Euchaetanthera, “G” (= Egania, Oriastrum and Tylloma, Glandulosa) Carmelita) Distribution Chile & Peru. Concentrated in RMS Chile, Argentina, Bolivia & Peru. (Chile). Concentrated over RMS (Chile) & Geography & Altitude San Juan & Mendoza (Argentina). Andean regions. Mostly 0-2000 m. 2000-5000 m. It seems that several sources of information (achene characters, and distribution) point towards a split within the genus Chaetanthera that is not really reflected by the assignment of the rank “subgenus”. Nevertheless it is well recognised that a single character — here strongly weighted towards achene hairs — is not sufficient to bind or separate ranks in a taxonomic hierarchy. Correlating suites of morphological characters drawn from other sources should support the achene characters. Further correlating character progressions are currently under investigation. These include parallel progressions seen in the two groups in capitulum size and floret ratios, life cycles, and habit, among others. 4. Achene hair diversity between the genera The significance of the results from Pachylaena and Brachyclados cannot be extensively interpreted. Pachylaena achenes show preliminary similarities to the achenes and hairs seen on species included in the Chaetanthera subgenus Tylloma, but much smaller. The two Brachyclados species both had “Zwillingshaare” (Table 3, section 3.1.5.) of a similar nature 31 to the obovate-lanceolate achene hairs but distinctly longer. However, B. /ycioides also had a second hair type — the spherical “Zwillingshaare” (Table 3, section 3.1.2.b). The presence of two hair types on the same achenes, which in Chaetanthera are so clearly segregated, is interesting. The achene hair information from these two genera Pachylaena and Brachyclados indicates that they are indeed outliers to Chaetanthera, and do not form part of this carpo- logically diverse genus as it stands. KARIS et al. (1992) sampled collections of six Chaetanthera species in their phylogenetic analysis of the Cichorioideae. They found that Chaetanthera formed part of a monophyletic cluster with, among others, Gerbera, Mutisia and Trichocline. Unfortunately they did not consider Pachylaena or Brachyclados. BREMER (1994) postulated that Pachylaena could have its sister group within a paraphyletic Chaetanthera. Were Chaetanthera to be monophyletic, then this could be resolved with a cladistic analysis of Chaetanthera and Pachylaena with Brachyclados as an outgroup. Other studies (GRAU, 1980; BREMER, 1994), conducted at the generic level, cite Chaet- anthera results that are often based on material pertaining only to taxa belonging to the “L” group. In the light of the achene hair results, these studies have limited usefulness for a discussion of generic limits of Chaetanthera. Conclusion Many studies indicate that where divergent achene hair types are found, they are frequently indicative of natural generic boundaries. The question then remains: do the divergent achene hairs types found in this study represent a fundamental split in the “genus” Chaetanthera different to that hitherto reflected in the current classification? Several points can be highlighted: e There is more variation among the achenes of the genus Chaetanthera, in particular regarding the achene hairs, than hitherto described. The variation in the achene hairs shows a dramatic division of the species found in the genus Chaetanthera Ruiz & Pav. e Although achene hair length is closely correlated to shape, achene hair type was found to be a clear descriptor of species clusters above the specific level, at the subgeneric level. The absence of achene hairs (the glabrous state) is not, as a rule, a taxonomically useful feature at the sub-generic level. e 4 subgenera Proselia, Euchaetanthera, Tylloma, and Glandulosa form one group with the same achene hair type (obovate-lanceolate “Zwillingshaare”). 3 subgenera, Oriastrum, Egania and Carmelita, express 6 different achene hair types and include the majority of glabrous achenes. Geographical distribution data and altitude information show correlating trends with species groupings based on achene hair characters. e The two genera Brachyclados and Pachylaena were shown to be distinct from all groupings within the genus Chaetanthera based on achene hairs. This study implies that achene diversity could be representative of significant divisions within the genus Chaetanthera as it stands. In particular the separation of the subgenera Proselia, Euchaetanthera, Tylloma, and Glandulosa (as one group) from Oriastrum, Egania and Carmelita seems a possibility. The achene characters of the outlying genera Pachylaena and Brachyclados while interesting, do not help clarify these divisions. The variation we have found has highlighted those areas of the genus which urgently need reviewing. The search for more characters to resolve the question of the generic limits of Chaetanthera continues and a full revision is currently in progress. 32 Grateful thanks are extended to Prof. J. Grau, the Curators and Directors of the herbaria who generously loaned the material used in this research and to Dr R. Davies. References BREMER, K. 1994: Asteraceae: Cladistics and Classification. Portland, Oregon. BRUMMITT, R.K. & POWELL, C.E. 1992: Authors of Plant Names. Kew. CABRERA, A.L. 1937: Revision del genero Chaetanthera (Compositae). — Revista Mus. La Plata Secc. Bot. 1(3): 87-215. DITTRICH, M. 1985: Morphologische und anatomische Untersuchungen an den Blüten und Früchten der Gattung Carlina (Compositae). — Bot. Jahrb. Syst. 107: 591-609. — 1996: Die Bedeutung morphologischer und anatomischer Achänen-merkmale für die Systematik der Tribus Echinopeae Cass. und Carlineae Cass. — Boissiera 51: 7-102. FAYED, A. 1979: Revision der Grangeinae (Asteraceae — Astereae). — Mitt. Bot. Staatssamml. München 15: 425-576. GRAU, J. 1971: On the generic delimitation of some South African Astereae. — Mitt. Bot. Staatssamml. München 10: 275-279. — 1973: Revision der Gattung Felicia (Asteraceae). — Mitt. Bot. Staatssamml. München 9: 195-705. — 1980: Die Testa der Mutisieae und ihre systematische Bedeutung. — Mitt. Bot. Staats- samml. München 16: 269-332. HANAUSEK, T.F. 1910: Beiträge zur Kenntnis der Trichombildungen am Perikarp der Kompositen. — Oesterr. Bot. Z. 60: 132-136, 184-187. HANSEN, H.V. 1989: Taxonomy and phylogeny of the genera in the scapose group of Compositae, tribe Mutisieae, subtribe Mutisiinae. — Nordic J. Bot. 9(5): 469-485. — 1991: Phylogenetic studies in Compositae, Tribe Mutisieae. — Opera Bot. 109: 1-50. HEss, R. 1938: Vergleichende Untersuchungen über die Zwillingshaare der Compositen. — Bot. Jahrb. Syst. 68: 435-496. HOLMGREN P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990: Index Herbariorum Part I: The Herbaria of the World. 8" Ed. New York. JEFFREY, C. 1967: Notes on Compositae Il. The Mutisieae in East Tropical Africa. — Kew Bull. 21: 177-224. KARIS, P.O., KALLERSJO, M. & BREMER, K. 1992: Phylogenetic anaysis of the Cichorioideae (Asteraceae), with emphasis on the Mutisieae. — Ann. Missouri Bot. Gard. 79(2): 416-427. KING, R.M. & ROBINSON, H. 1970: The new Synantherology. — Taxon 19: 6-11. LANE, M.A. 1985: Features observed by electron microscopy as generic criteria. — Taxon 34(1): 38-43. METCALF, C.R. & CHALK, L. 1979: Anatomy of the Dicotyledons. 2™ Ed. Oxford. OBERPRIELER, C. 1998: The Systematics of Anthemis L. (Compositae, Anthemideae) in W and C North Africa. — Bocconea 9: 1-328. RAMAYYA, N. 1962: Studies on the trichomes of some Compositae I. General Structure. — Bull. Bot. Surv. India 4(1-4): 177-188. ROMMEL, A. 1979 Die Gattung Amellus L. (Asteraceae — Astereae) systematischer Teil. — Mitt. Bot. Staatssamml. München 13: 579-728. Ruiz, H.L. & PAVON, J.A. (1794): Flora peruvianae, et chilensis prodromus. Madrid. SCOTT, R.W. 1985: Microcharacters in the Eupatorieae. — Taxon 34(1): 26-30. STEARN, W.T. 1995: Botanical Latin. 4" Ed. Devon. SUNDBERG, S. 1985: Micromorphological characters as generic markers in the Astereae. — Taxon 34: 31-37. VELEZ M.C. 1981: Karpologische Untersuchungen an Amerikanischen Astereae (Compo- sitae). — Mitt. Bot. Staatssamml. München 17: 1-170. 33 VINCENT, P.L.D. 1996: Progress on clarifying the generic concept of Senecio based on an extensive world-wide sample of taxa. — In: HIND, D.H.J & BEENTJE, H.J. (eds.): Com- positae: Systematics. Proceedings of the Inernational Compositae Conference, Kew, 1994 vol. 1: 597-611. Kew. Alison M.R. DAVIEs, Institut fiir Systematische Botanik der Ludwig-Maximilians-Universitat München, Menzinger Straße 67, D-80638 München, Germany. Appendix The collections cited here represent specimens (limited to a maximum of two) that had typical achene hairs for the species. The collections that contributed to the wider understanding of this project are not cited. The collections in bold are photographically illustrated in this paper. Typical Collections Surveyed Buchtien 84 (M); Johnston 5925 (GH) Ehrhart & Grau 95/818 (M); Werdermann 627 (M) Elliott 292 (K); Grau 2495 (JG) C. ciliata Ruiz & Pav. Grau 2400 (M); Rosas 1890 (M) C. cochlearifolia (A. Gray) B. L. Rob MacBride & Featherstone 845 (GH) Sleumer 1875 (W) Rosas 1875 (M); Werdermann 1256 (M Morrison 17071 (GH); Philippi s.n. (W Grau 2440 (M); Philippi s.n. (W Rosas 1076 (M); K. & W. Rechinger 63504 (M Rosas 1592 (M); Morrison & Wagenknecht 17104 (K Jiles 5001 (M); K. & W. Rechinger 63743 (W Werdermann 934 (M); Jiles 2487 (M Rosas 1226 (M); Morrison 16740 (GH K. & W. Rechinger 63321 (M); Werderman 890 (M C. gnaphalioides (Remy) |. M. Johnst. Jiles 4158 (M); Wagenknecht 18590 (GH Morrison 17300 (K); Werdermann 632 (M). Grau 2542 (M); Wagenknecht 18489 (G Castellanos 71644 (W); Johnston 6094 (GH Grau 2360 (JG); Pirion 492 (GH Sparre 1572 (K Weberbauer 6876 (GH Germain #1246 (W); Hastings 432 Sleumer 374 (W); Vervoorst 3239 (W Grau 3293 (M); Seibold 2990 (W C. Grandjot s.n. (M); Seibold 2850 (W Werdermann 1020 (M Grau s.n. (M); Pennell 12836 (GH Kiesling, Ulibarri & Krapovickao 7467 (K); Malme 2940 (GH Castellanos 71647 (W); Werdermann 253 (M Jiles 4816 (M Venturi 9266 (GH Grau 2407 (M); Werdermann 815 (GH Philippi 359a (P Hollermayer 415 (W); Schajovskoy s.n. (M Jiles 4870 (M); Schajovskoy 150 (M). King & Heinz 9390 Schajovskoy 32] (M); Schajovskoy s.n. ve ! | N 2 a a - By ; . yo hee rn, y Lined 7 ‘ ne yyne saa iF Abi ui ue iim uty Aa Bas I ie, iV Raerth a ‘ he phe, os vet = rar : — ra T % ‘ At ‘ Nah Di P . ehh wae 21 gi & 5 e f - ‚Ale 2, ’ Eu " : am 7 Ast aloe Pi 1 wath oh wok eae . e Ba. ne ie ae a eee 2 224 m, tt testis nro Fi Ivy es ube 43 us. RIC ic, reel ug net 3 IH it en tis wr ita ad A HT 35 Polytrichadelphus magellanicus sensu lato (Musci) and its ascomycetes — different fungi on different hosts P. DÖBBELER Abstract: DÖBBELER, P.: Polytrichadelphus magellanicus sensu lato (Musci) and its ascomycetes — different fungi on different hosts. — Sendtnera 7: 35-45. 2001. ISSN 0944-0178. Forty-six herbarium specimens of the moss Polytrichadelphus magellanicus s.l. (Poly- trichaceae) from all parts of its (sub)antarctic distribution area were analysed for associated fungi. On thirty-eight specimens fruit-bodies representing ten species of ascomycetes were found. P. magellanicus s.str. from Patagonia and Tierra del Fuego was found to be an excellent host, infected by nine fungal species belonging to seven genera. Each of the fifteen infected collections was associated, on average, with four different fungal species. However, on the twenty-three infected specimens of P. inno- vans from New Zealand, the Auckland Islands, Australia and Tasmania, only Epibryon pogonati-urnigeri was present with regularity. Except for the unspecific Bryochiton perpusillus both host mosses have their own parasites. This underlines the distinctness of P. magellanicus s.str. from southern South America and P. innovans from Austral- asia. These taxa have probably developed independently from each other over a long time period. Relations between P. innovans and Pogonatum urnigerum are discussed. Zusammenfassung: Sechsundvierzig Herbarbelege des Laubmooses Polytrichadelphus magellanicus s.l. (Polytrichaceae) aus allen Teilen des (sub)antarktischen Verbreitungsgebietes wurden auf Pilzbefall hin analysiert. Auf achtunddreißig Proben konnten Fruchtkörper nach- gewiesen werden, die zehn Arten von Ascomyceten repräsentieren. P. magellanicus s.str. aus Patagonien und Feuerland ist mit neun vorkommenden Pilzarten, die zu sie- ben Gattungen gehören, ein ausgezeichneter Wirt. Jede der fünfzehn befallenen Auf- sammlungen weist im Durchschnitt vier verschiedene Pilzarten auf. Auf den dreiund- zwanzig Proben von P. innovans aus Neuseeland, den Auckland Inseln, Australien und Tasmanien ist hingegen fast nur Epibryon pogonati-urnigeri vorhanden, allerdings mit erstaunlicher Regelmäßigkeit. Abgesehen von dem unspezifischen Bryochiton perpusillus haben beide Wirtsmoose ihre eigenen Parasiten. Das unterstreicht die Eigenständigkeit von P. magellanicus s.str. aus dem südlichen Südamerika und P. in- novans aus Australasien. Vermutlich haben sich beide Sippen über lange Zeiträume getrennt voneinander entwickelt. Diskutiert werden BeziehungenzwischenP. innovans und Pogonatum urnigerum. Introduction Two randomly collected specimens of Polytrichadelphus magellanicus (Hedw.) Mitt. (Polytri- chales, Musci) from Tierra del Fuego proved to be extremely good hosts for parasitic asco- 36 mycetes. Six and seven species, respectively, were recorded, representing no less than five genera and four orders. Apparently, hundreds of ascomata may be formed on a single host plant without causing any symptoms visible to the naked eye (DOBBELER 1999b). The present study analyses the fungal diversity within different populations of P. magellanicus in southern South America and P. innovans (Müll.Hal.) A.Jaeger in Australasia. The latter taxon often was not separated from P. magellanicus (e.g. BEEVER et al. 1992, FIFE 1995, RAMSAY 1997, SCOTT& STONE 1976, van ZANTEN 1978: 460). Material and methods Specimens of Polytrichadelphus magellanicus s.\. from the bryological herbaria B, F, G, GZU, JE, M and W were screened for the presence of fungi. Before studying the plants it was necessary to wet them thoroughly. This clearly demonstrates their endohydric organization as well as highlightingthe water repellentlamellae margins. Small or sparsely distributed ascomata were only detectable by microscopic screening of individual host leaves. Lugol’s solution (Merck 9261) was used to test iodine reactions. Infected plants, or parts of them, were separated and deposited under the fungal name in the corresponding host herbarium. Specimens examined Polytrichadelphus magellanicus s.str. as host (specimens 12 and 13, reported by DOBBELER at are included) Chile. Prov. de Cautin, Volcano Villarrica, ca. 1000 m, Hollermayer 56 (W). Recorded species: Crocicreas magellanicum, Dawsophila polycarpa, Potriphila neurogena, Rogellia nectrioidea. 2 Chile. Prov. de Cautin, Andes of Villarrica, (probably 1897), Neger (JE). Recorded species: Cro- cicreas magellanicum, Dawsophila polycarpa, Potriphila navicularis, P. neurogena. 3 Chile. Prov. de Valdivia, Corral, near Quitaluto, ca. 400 m, 2.2.1935, Hosseus 698 (JE). Recorded species: Bryochiton perpusillus, Crocicreas magellanicum. 4 Chile. Prov. de Valdivia, Cordillera Pelada, road from La Union to El Mirador, 800 m, 12.12. 1985, Hellwig 8560 (M). Recorded species: Dawsophila polycarpa, Rogellia nectrioidea. 5 Chile. Prov. de Valdivia, West slope of Cordillera Pelada, 8.8 km by road W of El Mirador on road between La Union and Punta Hueicolla, 580 m, 40°07'S, 73°16'W, 18.2.1976, Crosby 12758 (B, W). Recorded species: Crocicreas magellanicum, Dawsophila polycarpa. 6 Chile. Prov. de Chiloé, Trumao, path to “Madrina”, 200 m, 28.5.1932, Junge 424 (F). Recorded species: Bryochtion perpusillus, Crocicreas magellanicum, Dawsophila polycarpa. 7 Chile. “Waldhagen”, [Nothofagus] pumilio-tree line, 1200 m, 1.1.1938, Schwabe 34 (JE). Recorded species: Dawsophila polycarpa, Epibryon eremita, Potridiscus polymorphus, Potriphila navicularis, P. neurogena, Rogellia nectrioidea. 8 Chile. Prov. de Coihaique, Checkpoint Cisnes Medio of the Carretera Austral, IV.1985, Hellwig 811 (M). Recorded species: Bryochiton perpusillus, Crocicreas magellanicum, Dawsophila poly- carpa. 9 Chile. Prov. de Magallanes, Punta Arenas, 1917, Hicken (B). Recorded species: Crocicreas magel- lanicum, Dawsophila polycarpa, Potriphila navicularis, P. neurogena. 10 Chile. Prov. de Antartica Chilena, Canal Beagle, II.1922, Gusinde 339 (W). Recorded species: Crocicreas magellanicum, Dawsophila polycarpa, Potriphila navicularis, P. neurogena, Ro- gellia nectrioidea. 11 Argentina. Tierra del Fuego, Dep. Ushuaia, Rio Olivia, 27.9.1979, Matteri NC2153 (B). Recorded species: Bryochiton perpusillus, Crocicreas magellanicum, Dawsophila polycarpa. 12 Argentina. Tierra del Fuego, Dep. Ushuaia, about 30 km E of Ushuaia, road from Ushuaia to Paso Garibaldi, ca. 190 m, 22.3.1997, Mues (M). Recorded species: Crocicreas magellanicum, Daw- sophila polycarpa, Epibryon eremita, E. interlamellare, Potriphila navicularis, P. neurogena (Type). ‘3 14 15 37 Argentina. Tierra del Fuego, Dep. Ushuaia, near Paso Garibaldi, ca. 430 m, 20.3.1997, Mues (M). Recorded species: Crocicreas magellanicum (Type), Dawsophila polycarpa (Type), Epibryon eremita (Type), E. interlamellare, Potriphila navicularis, P. neurogena, Rogellia nectrioidea (Type). Argentina. Tierra del Fuego australis, Dep. Ushuaia, Montes Martiales, 320 m, 54°46'S, 68°18'W, 30.12.1969, Roivainen (Plantae Argentinae 1480, sub Polytrichadelphus magellanicus), (G). Recorded species: Dawsophila polycarpa. Argentina. Tierra del Fuego, Dep. Ushuaia, Cerro Martial, 350 m, 5.1.1989, Schäfer-Verwimp & Verwimp 10849 (G). Recorded species: Crocicreas magellanicum, Dawsophila polycarpa, Epi- bryon eremita, E. interlamellare, Potriphila navicularis, P. neurogena. Polytrichadelphus innovans as host 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 New Zealand. (without further notes), com. Dr. Schwarz (W). Recorded species: Epibryon pogo- nati-urnigeri. New Zealand. North Island, North Auckland, Waipoua forest, 50 km NW of Dargaville, ca. 300 m, 9.1.1974, van Zanten 7401214 (B). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Paradise Valley, Fish hatchery, near Rotorua, 7.1.1969, O. & I. Degener 31830 (GZU). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Waiotapu, IX.1929, Meebold (M). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Bay of Plenty, Kaingaroa plantation forest, SE of Rotorua, ca. 600 m, 25.10.1959, van Zanten 1251 (B). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Tauhara, Taupohill, about 2500', IX.1929, Meebold (M). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Waitakerei Range, 1206', IX.1929, Meebold (M). Recorded species: Bryochiton perpusillus. New Zealand. North Island, Gisborn district, Lake Waikaremoana, along road from Waiotukupuna-bridgeto Cascade Falls, ca. 600 m, 29.8.1968, van Zanten 681886 (JE). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Mt. Egmont, 3000', VIII.1929, Meebold (M). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Mt. Egmont, Dawson Falls, 6.3.1959, H. & E. Walter (M). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Hawke’s Bay, near Whakarara, Gardiner’s Mill, 31.12.1950, Sainsbury (W). Recorded species: Epibryon pogonati-urnigeri. New Zealand. North Island, Wellington, York Bay, V.1929, Meebold (M). Recorded species: Epibryon pogonati-urnigeri. New Zealand. South Island, Westland, Teramakau Valley, 600 ft, 6.2.1903, Naylor Beckett (JE). Recorded species: Epibryon pogonati-urnigeri. New Zealand. South Island, West Coast Area, N of Whataroa, 7.2.1970, Vitt 4505 (B). Recorded species: Bryochiton perpusillus, Epibryon pogonati-urnigeri. New Zealand. South Island, South Otago, The Chaslands, along highway 93, 14.1.1973, Visch (B). Recorded species: Epibryon pogonati-urnigeri. New Zealand. Stewart Island, 1.1889, Bell (M). Recorded species: Epibryon pogonati-urnigeri. Auckland Islands. Auckland Island, South end of North Harbour, 12.1.1973, Vitt 10223 (M). Recorded species: Epibryon pogonati-urnigeri. Australia. Victoria, Baw Baw National Park, 6 km SE of Mt. Erica, road cutting on Mt. Erica, 700-800 m, 37°35'S, 146°23’E, 10.1.1986, Rambold 3276 (M). Recorded species: Epibryon pogonati-urnigeri. Australia. Victoria, Baw Baw National Park, 6 km SE of Mt. Erica, road cutting on Mt. Erica, 900-1000 m, 37°35'S, 146°23'E, 10.1.1986, Rambold 3280 (M). Recorded species: Epibryon pogonati-urnigeri. Australia. Victoria, Mt. Donna Bvang, 5 km N of Warburton, 1200 m, 12.4.1974, Streimann 991 (B). Recorded species: Epibryon pogonati-urnigeri. Tasmania. (without further notes), Lucas (M). Recorded species: Epibryon pogonati-urnigeri. Tasmania. NW of Mt. Field, Florentine Valley, 21.1.1959, H. & E. Walter (M). Recorded species: Epibryon pogonati-urnigeri. Tasmania. Hobart, Mt. Wellington, 14.4.1903, Fleischer B2074 (B). Recorded species: Epibryon pogonati-urnigeri. 38 The ascomycetes recorded Bryochiton perpusillus Döbbeler (DÖBBELER, Mitt. Bot. Staatssamml. München 14: 226, 1978) Hosts: Polytrichadelphus magellanicus, P. innovans (and other bryophytes, see below) Distribution on Polytrichadelphus: Chile, Argentina, New Zealand Specimens examined: 3, 6, 8, 11, 22, 29 Bryochiton perpusillus is a very widespread and variable species recorded from distantly related bryophytes, often with a withered appearance, indicating that taxonomic uniformity should be proved. Partially damaged hepatics, like Prilidium occur within the spectrum of hosts besides the Polytrichaceae, including Dawsonia (DÖBBELER 1978, 1981). B. perpusillus most regularly colonises Polytrichum piliferum Hedw. where it develops subcuticularly from a conspicuous network of thick brown hyphae, especially in the apical, abaxial region of normal looking or hardly altered leaves. On Polytrichadelphus, Bryochiton perpusillus occurs in lower concentrations. Ascomata are sparse on the lower, withering leaves of a plant, suggesting the saprophytic tendency of the fungus. Generally, apical regions of individual leaves are preferred and often both sides of a leaf have fruit-bodies as well as the interlamellar spaces being infected. Here the ascomata may look very similar to those of Epibryon pogonati-urnigeri. However, the latter species prefers upper healthy leaves where the fungus is completely surrounded by the leaf lamellae. A positive iodine reaction within the ostiolar region immediately reveals E. pogonati- urnigeri. Crocicreas magellanicum Döbbeler (DÖBBELER, Haussknechtia Beih. 9: 81, 1999b) Host: Polytrichadelphus magellanicus Distribution: Chile, Argentina Specimens examined: 1-3, 5, 6, 8-13, 15 Due to a lack of contrast with its substrate, the white to nearly hyaline apothecia of Crocicreas magellanicum are often difficult to detect, especially when hidden between the leaf lamellae and sparse in numbers. Microscopically screenedleaves repeatedly proved to be infected, though apothecia could not be seen by low stereo-microscopic magnification. Dawsophila polycarpa Döbbeler (DÖBBELER, Haussknechtia Beih. 9: 83, 1999b) Host: Polytrichadelphus magellanicus Distribution: Chile, Argentina Specimens examined: 1, 2, 4-15 The two species Dawsophila callichroma Döbbeler and D. pygmaea Döbbeler frequently colonise different members of the genus Dawsonia and are present within its whole Australasian distribution range (DÖBBELER 1981). Though special attention was paid to seeking Dawsophila on Polytrichadelphus innovans, which shares its distribution for the most part with Dawsonia, ascomata could not be detected. Epibryon eremita Döbbeler (DÖBBELER, Haussknechtia Beih. 9: 85, 1999b) Host: Polytrichadelphus magellanicus Distribution: Chile, Argentina Specimens examined: 7, 12, 13, 15 Several ascomata of collection 15 had approximately thirty-two spores per ascus unlike the eight spores per ascus found in the other three specimens, including the type specimen. Spore size among specimens was similar. The larger asci (up to 60 xm long) found in the 32-spored asci 39 simply accommodate a larger spore number than the smaller asci (35-40 um long). As there are no other differences to the type material, spore number and ascus size do not seem to be of taxonomic significance. Relations to Epibryon elegantissimum Döbbeler on Dawsonia superba Grev. in New Zealand (DÖBBELER 1981) may exist, judging by fruit-body morphology, hymenial features and the peculiar microhabitat preference. Epibryon interlamellare (Racov.) Döbbeler (DÖBBELER, Mitt. Bot. Staatssamml. München 14: 279, 1978) Hosts: Polytrichadelphus magellanicus (and other Polytrichaceae) Distribution on Polytrichadelphus: Argentina Specimens examined: 12, 13, 15 This frequent and widespread species infects different Polytrichaceae (see DÖBBELER 1999b). Epibryon pogonati-urnigeri Döbbeler (DOBBELER, Mitt. Bot. Staatssamml. München 14: 296, 1978) Hosts: Polytrichadelphus innovans (and other Polytrichaceae, see below) Distribution on Polytrichadelphus: New Zealand, Auckland Islands, Australia, Tasmania Specimens examined: 16-21, 23-38 The young, upper leaves of an infected plant normally contain only a few ascomata which are restricted to the apical leaf region. Due to a lack of space, they are very laterally compressed. Often the ascomatal walls adjacent to the lamellae are reduced or completely missing. In older leaves ascomata are more evenly distributed and spherical. Heavily infected, 6 mm long leaves of different collections commonly yielded about hundred ascomata. In one case an individual leaf of 8 mm length was colonised by approximately 180 ascomata. The species is quite uniform on Polytrichadelphus innovans, although slight differences in spore size and other features between collections on different Polytrichaceae or from different geographical regions apparently exist. An essential feature of Epibryon pogonati-urnigeri is its iodine reaction, which is not known in other bryophilous ascomycetes. The ostiolar region turns reddish in Lugol’s solution and KOH pretreatment gives a more distinct bluish colour (hemiamyloid reaction, BARAL 1987). Four polytrichaceous hosts are known for Epibryon pogonati-urnigeri in addition to Polytrichadelphus innovans, namely Pogonatum urnigerum (Hedw.) P.Beauv., P. dentatum (Brid.) Brid. (DOBBELER 1978, 1985), Dawsonia longiseta Hampe and D. polytrichoides R.Br. (DOBBELER 1981). With the exception of P. dentatum, for which only a few data are available, these mosses are very regularly or (in certain regions) even indiscriminately infected by E. pogonati-urnigeri. Both species of Pogonatum represent the subgenus Dendroidea (HYVONEN 1989). It is rather well documented that E. pogonati-urnigeri occupies wide areas of the geographical range of P. urnigerum (DOBBELER 1978, 1985) which is distributed in northern Eurasia, North America and in scattered high mountain localities in Africa, southeast Asia and Papua New Guinea (HYVONEN 1989). After studying hundreds of European Polytrichaceae it is obvious that species like Pogonatum aloides (Hedw.) P.Beauv. (P. subgen. Pogonatum) or Polytrichastrum alpinum (Hedw.) G.L.Sm. (syn. Pogonatum alpinum (Hedw.) Röhl.) do not host E. pogonati-urnigeri. Dawsonia longiseta and D. polytrichoides make up the more primitive section of the genus Dawsonia (D. sect. Dawsonia) and are restricted to eastern Australia (van ZANTEN 1973). On studying representative material DOBBELER (1981) observed that ascomata of Epibryon pogonati-urnigeri were always present on both D. longiseta and D. polytrichoides whereas for D. superba (D. sect. Superba) the situation is quite different. Though occurring within the range of the parasite in New Zealand, Australia and Tasmania, parasitism by 40 E. pogonati-urnigeri occurred only exceptionally. The mosses in question are not ecologically separate as they (as most Polytrichaceae) prefer open, exposed soil as on river banks and road cuttings. How should one interpret the range of hosts for Epibryon pogonati-urnigeri? It exhibits specificity between sections within Dawsonia and Pogonatum and between Pogonatum urnigerum and Polytrichastrum alpinum, but it does not discriminate among genera as different as Pogonatum, Polytrichadelphus and Dawsonia. The differences between these genera may not be as great as one might initially suppose because they are distinguished by sporophytic characters, i.e. capsule morphology. The gametophytic plants are well-developed and share several common features. The leaves are characterised by a distinct sheath and swelling tissue. They bear densely formed adaxial lamellae with specialised marginal cells which are covered in Polytrichadelphus and Dawsonia with angular flakes of epicuticular wax. In this respect they differ from other Polytrichaceae (PROCTOR 1992). Though species of Pogonatum vary greatly regarding waxy deposits, P. urnigerum appears “to be highly distinctive, with epicuticular wax on the papillose lamella margins in the form of flakes or platelets not unlike those of Dawsonia and Polytrichadelphus” (CLAYTON-GREENE et al. 1985: 552; see also Proctor 1979). On a macroscopic level, the upper leaf surface of these mosses displays a glaucous bloom caused by the waxy coverings. Another leaf character common to Pogonatum urnigerum and Polytrichadelphus innovans are similar lamellae hairs (DOBBELER 1999a, Figs. 4, 5). Nevertheless, recent molecular results on the phylogeny of the Polytrichales by HYVONEN et al. (1998) do not support a very close relationship between the genera in question. Potridiscus polymorphus Döbbeler & Triebel (DOBBELER & TRIEBEL, Hoppea 61: 72, 2001) Hosts: Polytrichadelphus magellanicus (and other Polytrichaceae, especially Polytrichum Juniperinum Hedw., DÖBBELER & TRIEBEL 2001) Distribution on Polytrichadelphus: Chile Specimen examined: 7 This distinctive species with a palaeoaustral distribution is more frequent than previously thought. Polytrichadelphus seems only to be an occasional host. Within a rather rich collection of P. innovans from New Zealand (collection 20) a few intermingled plants of Polytrichum Juniperinum were heavily infected by Potridiscus polymorphus, whereas no ascomata could be detected on the predominant Polytrichadelphus. Potriphila navicularis Döbbeler (DOBBELER, Nova Hedwigia 62: 62, 1996) Host: Polytrichadelphus magellanicus (and other Polytrichaceae) Distribution on P. magellanicus: Chile, Argentina Specimens examined: 2, 7, 9, 10, 12, 13, 15 Potriphila navicularis has a bipolar distribution type and is very frequent on Polytrichastrum alpinum (DOBBELER 1996). On Polytrichadelphus magellanicus, as in nearly all southern hemisphere records, only the anamorphicstate is present. In addition to normal leaves, bracts and scale leaves should also be screened. Usually only a few pycnidia are detectable on dozens of potential host plants. The host of Potriphila navicularis from the Auckland Islands is not Polytrichadelphus magellanicus, as recorded by DOBBELER (1996), but Polytrichastrum longi- setum (Brid.) G.L.Sm. 41 Potriphila neurogena Döbbeler (DÖBBELER, Haussknechtia Beih. 9: 88, 1999b) Host: Polytrichadelphus magellanicus Distribution: Chile, Argentina Specimens examined: 1,2, 7, 9, 10, 12, 13, 15 Ascomata also develop within male bracts surrounding the antheridia. Rogellia nectrioidea Débbeler (DOBBELER, Haussknechtia Beih. 9: 91, 1999b) Host: Polytrichadelphus magellanicus Distribution: Chile, Argentina Specimens examined: 1, 4, 7, 10, 13 In several cases the ascomata of both Rogellia nectrioidea and Potriphila neurogena were observed on the same host leaf. Geographical range of Polytrichadelphus magellanicus and P. innovans Austral populations of Polytrichadelphus are often regarded as a single species, P. magellanicus. All of the herbarium material cited (specimens 1-38) was determined as P. magellanicus (specimen 7 as P. horridus Mitt.). However, SMITH (1971) considers Polytrichadelphus of New Zealand and Australia as a distinct taxon, P. innovans. “Compared with the American P. magellanicus, the plants are smaller and more slender, with shorter, subsecund leaves. The leaf sheath is broader, with more pronounced shoulders, and the teeth on the margin of the blade less well developed” (SMITH 1971: 41). He additionally noted differences in capsule morphology. Moreover, the presence of two distinct species is clearly indicated by completely different lamellae hair types as documented by DOBBELER (1999a) and repeatedly observed in this study. If one accepts P. innovans as a distinct taxon, then P. magellanicus s.str. is confined to Chile and Argentina south of about 40 latitude, the Fuegian region, Juan Fernandez Islands, the Falkland Islands, Tristan da Cunha and Gough Island (GREENE 1986, HE 1998, PROCTOR 1992, ROBINSON 1975, SCHIAVONE 1993, SCHIAVONE & HyYVONEN 1992, SMITH 1971). The assumed disjunct occurrence in the northern Andes of Peru (SCHULTZE- MOTEL & MENZEL 1987, GRADSTEIN& FRAHM 1987, MENZEL 1992) is based on a BRYOTROP-specimen [Peru. Dep. Amazonas, Prov. Chachapoyas, road Chachapoyas-Cajamarca km 413, mountain rain forest, 3000 m, 5.9.1982, Frahm, Geissler, Gradstein, Philippi & Schultze-Motel 916 (Herbarium Frahm!). Female plants with only one capsule and several successive, apparently lateral setae typical for the genus Polytrichadelphus.]. This species clearly differs by habit and leaf lamellae characters from the southern South American Polytrichadelphus magellanicus. The Australasian P. innovans is known from New Zealand, southeastern Australia, Tasmania,the Auckland Islands and Campbell Island (SMITH 1971, Vitt 1974, 1979). Comparison of the ascomycetes on Polytrichadelphus magellanicus and P. innovans Careful analysis of forty-six collections of both species of Polytrichadelphus revealed the presence of ten different ascomycetes, eight only present on the American, one only on the Australasiatic host and one common to both. Of the seventeen collections of Polytrichadelphus magellanicus from Chile and Argentina, fifteen are infected by one to seven species of ascomycetes. Altogether fifty-eight specimens of ascomycetes are found, on average nearly four 42 on each host collection. Thus P. magellanicus represents one of the best hosts, not only within Polytrichaceae but also within bryophytes in general, with regard to the number of associated fungi and the frequency of infection. Dawsophilapolycarpa, the most common species, is present on fourteen host specimens; Crocicreas magellanicum, on twelve; Potriphilaneurogena, on eight; P. navicularis, on seven; Rogellia nectrioidea, on five; Bryochiton perpusillus and Epibryon eremita, on four and Potridiscus polymorphus, on one host collection. Twenty-eight specimens of Polytrichadelphus innovans from New Zealand, Australia and Tasmania and one collection from the Auckland Islands were examined. Twenty-three proved to be colonised. Epibryon pogonati-urnigeri was present on twenty-two specimens, whereas Bryochiton perpusillus was encountered only twice. This means that on average only one associated fungus per host collection was found. The present inventory should not imply that the fungal diversity of the two austral species of Polytrichadelphus is completely covered. Only the more frequent, previously described parasites are discussed. On several occasions ascomata of an undescribed species were detected but are not discussed in this paper. In most cases the potential hosts consisted of only a few plants gathered for bryological purposes. More extensive collections are required to obtain a sufficient number of ascomata for a proper investigation of the other sparsely distributed parasites. Field studies are also required to seek out the relatively conspicuous, necrotrophic fungi that are unlikely to be representedin bryophyte herbarium material. An exampleis Lizonia, which occupies the antheridial cups of various polytrichaceous mosses. One should also search for orange coloured apothecia formed on the ground between individual plants, as they may belong to rhizoidparasitic species of Octospora or Leucoscypha. Whether Polytrichadelphus belongs to the host range of the latter genera and also Lizonia is not currently clear. Interpretation of bryomycological evidence The most surprising result of this bryomycological analysis is the diversity of the fungal associates of the two austral species of Polytrichadelphus studied. Only one species out of ten was encountered several times on both hosts, namely Bryochiton perpusillus, a very common and critical taxon which is also recorded from hepatics. Five ascomycete species are known exclusively on P. magellanicus s.str., among them Potriphila neurogena and Rogellia nectrioidea. The latter two species develop within the leaf nerves, a distinct niche rarely occupied by the multitude of polytrichophilous ascomycete species. The marked differences in suitability of the two moss species as hosts was also unexpected. Polytrichadelphus magellanicus is an excellent substrate, whereas P. innovans is less attractive. The absence of common fungal parasites supports the distinctness of both moss taxa as has already been shown by morphological features. Another question concerns the phylogenetic relationship of the Polytrichadelphus species. Geographical isolation even for long geological periods does not automatically lead to genetic diversification as demonstrated by Frey et al. (1999) using molecular methods. In the ancient Lopidium concinnum (Musci) which has exactly the same distribution range as P. magellanicus s.l. little genetic divergence has occurred within the isolated populations since their separation by geotectonic events about 80-60 million years ago. The vicariant taxa P. magellanicusand P. innovans (SMITH 1972: 46) are “undoubtedly closely related” (PROCTOR 1992: 317). This inferred relationship is not mirroredby an overlap of the more significant fungal parasites. Furthermore, Dawsophila frequently colonises various species of Dawsonia (DOBBELER 1981) and P. magellanicus, but avoids P. innovans. Epibryon elegan- 43 tissimum occurring on Dawsonia and E. eremita infecting P. magellanicus seem to represent a further link between Australasianand South American polytrichaceous hosts, leaving P. innovans apart. The bryomycological facts documented herein indicate that P. magellanicus and P. in- novans have had a very long, separate evolution. It would be interesting to know more about the ascomycetes on the relatives of Polytrichadelphus, both in South America and Australasia. I am indebted to the curators of the herbaria B, F, G, GZU, JE, M and W for loans of specimens. Prof. Dr. F. Hellwig (Jena) provided several specimens collected in Chile many years ago. Dr. H. No- wak-Krawietz (Berlin) helped with the literature. Prof. Dr. W. Frey (Berlin) and Prof. Dr. H. Hertel (München) made comments on an early draft of the manuscript. I am gratefulto Prof. Dr. T. H. Nash III (Tempe) and M. Cullen (Dublin) for linguistic correction of this paper. Literature BaRAL, H.O. 1987: Lugol’s solution/IKI versus Melzer’s reagent: hemiamyloidity, a universal feature of the ascus wall. — Mycotaxon 29: 399-450. BEEVER, J., ALLISON, K.W. & CHILD, J. 1992: The mosses of New Zealand, sec. ed. Dunedin. CLAYTON-GREENE, K.A., COLLINS, N.J., GREEN, T.G.A. & Proctor, M.C.F. 1985: Surface wax, structure and function in leaves of Polytrichaceae. — J. Bryol. 13: 549-562. DÖBBELER, P. 1978: Moosbewohnende Ascomyceten I. Die pyrenocarpen, den Gametophyten besiedelnden Arten. — Mitt. Bot. Staatssamml. München 14: 1-360. — 1981: Moosbewohnende Ascomyceten V. Die auf Dawsonia vorkommenden Arten der Botanischen Staatssammlung Miinchen. — Mitt. Bot. Staatssamml. Miinchen 17: 393-473. — 1985: Moosbewohnende Ascomyceten VII. Neufunde einiger Arten der Gattung Epibryon. — Mitt. Bot. Staatssamml. München 21: 757-773. — 1996: Potriphila navicularis gen. et sp. nov. (Ostropales, Ascomycetes), ein bipolar ver- breiteter Parasit von Polytrichum alpinum. — Nova Hedwigia 62: 61-77. — 1999a: Lamellen- und Epidermishaare als neues Merkmal der Polytrichaceen (Musci). — Nova Hedwigia 69: 357-380. — 1999b: Polytrichadelphus magellanicus - a mycological Eldorado: five new ascomycetes on a single collection from Tierra del Fuego. — Haussknechtia Beth. 9: 79-96. — & TRIEBEL, D. 2001: Potridiscus polymorphus (Leotiales) — a new ascomycete on Poly- trichaceae (Musci) with palaeoaustral distribution. — Hoppea 61 (2000): 71-83. FIFE, A.J. 1995: Checklist of the mosses of New Zealand. — Bryologist 98: 313-337. Frey, W., STECH, M. & MEISSNER, K. 1999: Chloroplast DNA-relationship in palaeoaustral Lopidium concinnum (Hypopterygiaceae, Musci). An example of stenoevolution in mosses. Studies in austral temperate rain forest bryophytes 2. — Plant Syst. Evol. 218: 67-75. GRADSTEIN, S.R. & FRAHM, J.-P. 1987: Die floristische Höhengliederung der Moose entlang des BRYOTROP-Transektes in NO-Peru. — Beih. Nova Hedwigia 88: 105-113. GREENE, D.M. 1986: A conspectus of the mosses of Antarctica, South Georgia, the Falkland Islands and southern South America. — British Antarctic Survey: Cambridge. HE, S. 1998: A checklist of the mosses of Chile. — J. Hattori Bot. Lab. 85: 105-189. HyvONneEN, J. 1989: A synopsis of genus Pogonatum (Polytrichaceae, Musci). — Acta Bot. Fenn. 138: 1-87. — & HEDDERSON, T.A., SMITHMERRILL, G.L., GIBBINGS, J.G. & KOSKINEN, S. 1998: On phylogeny of the Polytrichales. — Bryologist 101: 489-504. MENZEL, M. 1992: Preliminary checklist of the mosses of Peru (Studies on Peruvian Bryophytes IV). — J. Hattori Bot. Lab. 71: 175-254. 44 Proctor, M.C.F. 1979: Surface wax on the leaves of some mosses. — J. Bryol. 10: 531-538. — 1992: Scanning electron microscopy of lamella-margin characters and the phytogeography of the genus Polytrichadelphus. — J. Bryol. 17: 317-333. RAMSAY, H.P. 1997: Cytotaxonomic studies on some Polytrichales from Australia, New Zealand, Papua New Guinea and Vanuatu. — J. Hattori Bot. Lab. 82: 213-226. RoBINSON, H. 1975: The mosses of Juan Fernandez Islands. — Smithsonian Contr. Bot. 27: I-IV, 1-88. SCHIAVONE, M.M. 1993: Bryophyta, Musci: Polytrich[in]ales.— Flora criptogämica de Tierra del Fuego 14 (12): 1-61. — & Hyvönen, J. 1992: Distribution maps of mosses from Tierra del Fuego, Argentina. 3. — Lindbergia 17: 47-49. SCHULTZE-MOTEL, W. & MENZEL, M. 1987: Die Laubmoosflora im BRYOTROP-Transekt von Peru. — Beih. Nova Hedwigia 88: 9-59. SCOTT, G.A.M. & STONE, I.G. 1976: The mosses of southern Australia. — London. SMITH, G.L. 1971: Conspectus of the genera of Polytrichaceae. - Mem. New York Bot. Gard. 21(3): 1-83. — 1972: Continental drift and the distribution of Polytrichaceae.— J. Hattori Bot. Lab. 35: 41-49. Vitt, D.H. 1974: A key and synopsis of the mosses of Campbell Island, New Zealand. — N. Zealand J. Bot. 12: 185-210. — 1979: The moss flora of the Auckland Islands, New Zealand, with a consideration of habitats, origins, and adaptations. — Canad. J. Bot. 57: 2226-2263. ZANTEN, B.O. van 1973: A taxonomic revision of the genus Dawsonia R. Brown. — Lindbergia 2: 1-48, pl. I-X1. — 1978: Experimental studies on trans-oceanic long-range dispersal of moss spores in the southern hemisphere. — J. Hattori Bot. Lab. 44: 455-482. ; Dr. Peter DOBBELER, Institut fiir Systematische Botanik der Ludwig-Maximilians-Universitat München, Menzinger Straße 67, D-80638 München, Deutschland. Table 1: The hosts, the associated ascomycetes and their frequency (by number of specimens recorded). Fifteen specimens of Polytrichadelphus magellanicus s.str. (southern South America) are infected by: Bryochiton perpusillus (4) Crocicreas magellanicum (12) Twenty-three specimens of Polytrichadelphus innovans (Australasia) are infected by: Bryochiton perpusillus (2) Epibryon pogonati-urnigeri (22) 45 co an an E an loa) | te] oe] se] se] ez] se] sz] ve] ee] ze] ve] 02] si] er] er] st e| /e/le/eo UNIIUD]JABDUL SD3491904) | | el | el el SE (er ern nnn inner EIGEN sna1upjjadpu Snydjappy214d]04 ISOH "sojo0AWOOse Imıseled Moy) pue supaouuı ‘g pue snorupjJedpw snydjappy>14Adj0g JO susunsads pajooJUt OY] :T oIgeL 47 Zephyra compacta (Tecophilaeaceae) — eine neue Art aus Chile C. EHRHART Zusammenfassung: EHRHART, C.: Zephyra compacta (Tecophilaeaceae) — eine neue Art aus Chile. — Sendtnera 7: 47-52. 2001. ISSN 0944-0178. Aus dem Küstenbereich des nördlichen Chile (IV Region de Coquimbo, Prov. de Elqui, 29°17'S — 71°18'W) konnte im Nifo-Jahr 1997 eine neue Tecophilaeacee der Gattung Zephyra gesammelt werden. Sie wird hier als Zephyra compacta spec. nov. beschrieben. Sie unterscheidet sich von der bislang einzigen Art der Gattung, Zephyra elegans D.Don durch den viel kompakteren Wuchs, durch die reichverzweigte, aus bis zu 15 Partialfloreszenzen zusammengesetzte Infloreszenz, die breiteren, fleischigeren Blätter/Tragblätter und die deutlich kleineren, reinweißen Blüten mit Tepalen, die im Gegensatz zu den Tepalen von Z. elegans nicht radförmig ausgebreitet, sondern zum Blütenstiel eingerollt sind. Resumen: Durante el afio del Nifio 1997 se colectö en la zona costera del norte de Chile (IV Region de Coquimbo, Prov. de Elqui, 29°17'S — 71°18'W) una especie nueva del género Zephyra (Tecophilaeaceae), que se describe como Zephyra compacta spec. nov. Este nuevo taxon se distingue de la ünica especie del género hasta ahora conocida, Zephyra elegans D.Don, por su habito compacto, su inflorescencia muy ramificada, por sus hojas mas anchas y mas suculentas, por sus flores blancas con tépalos, que son a diferencia de los tépalos de Z. elegans, mucho mas pequefios y no planos pero curvados hacia el envés. Einleitung Von der als monotypisch geltenden Gattung Zephyra kannte man bislang nur die im Küstenbereich des nördlichsten Chile (von der I. Region Tarapaca bis etwa zur III. Region Atacama) weitverbreitete Zephyra elegans D.Don. Im Nifo-Jahr 1997 konnten während einer Sammelreise im Küstengebiet zunächst wenig nördlich von La Serena aus der IV. Region, später auch noch etwa 100 km weiter nördlich, Populationen von Zephyra gesammelt werden, die sich in mehreren Merkmalen von Z. elegans unterscheiden. Diese Pflanzen sind wesentlich gedrungener gebaut, die Infloreszenz ist viel reicher verzweigt, und auch im Blütenbereich finden sich deutliche Unterschiede zu Z. elegans. Es handelt sich augen- scheinlich um eine gut differenzierte Art, die hier als Zephyra compacta neu beschrieben wird. Vom Wildstandort konnten sowohl Samen, als auch Lebendmaterial in Form von Knollen gesammelt und in München erfolgreich kultiviert werden. Dabei hat sich gezeigt, daß die 48 Merkmale der Art weitgehend konstant sind. Dies ist umso wichtiger in der Beurteilung der Abgrenzung der beiden Arten, da Z. elegans, wie sich bei Populationen verschiedener Wildstandorte zeigte, in einigen Merkmalen relativ variabel zu sein scheint und es damit erst zu klären galt, ob Z. compacta nicht nur als eine weitere, in das Variabilitätsspektrum von Z. elegans fallende Form anzusehen ist. Die Kultur von Wildmaterial mehrerer Aufsammlungen beider Arten zeigte jedoch keine kontinuierliche Verbindung zwischen den beiden Arten, so daß eine Etablierung dieser südlichsten Populationen als eigene Art gerechtfertigt ist. Zur Taxonomie von Zephyra Die monotypische Gattung Zephyra mit Zephyra elegans wurden 1832, basierend auf einer Aufsammlung von Boellaert, von Don beschrieben. 1864 glaubte Miers Zephyra-Belege, die ebenfalls von Boellaert gesammelt worden waren, als eine neue Art ansprechen zu können und benannte sie Zephyra amoena. Nach Überprüfung der Typusbelege beider Taxa steht fest, daß es sich bei Z. amoena um ein Synonym von Z. elegans handelt und die Beschreibung von Z. amoena höchstwahrscheinlich sogar auf der gleichen Aufsammlung von Boellaert beruht. Miers hat vermutlich nur Teilstücke einer Pflanze gesehen, zumindest zeigen die beiden Typusbelege von Z. amoena in BM ganz eindeutig einen Teilbereich der Infloreszenz, wie er für Z. elegans charakteristisch ist. Die von Miers beschriebenen Merkmalsunterschiede der beiden Arten liegen durchaus im Variabilitätsbereich der Merkmale von Z. elegans. Offensichtlich in Unkenntnis der Existenz von Zephyra, beschrieb Philippi 1873 eine neue Gattung Dicolus mit der einzigen Art Dicolus caerulescens Phil., die nach Typus und Diagnose ebenfalls völlig mit Z. elegans übereinstimmt und in die Synonymie zu dieser zu stellen ist. Die Typusaufsammlung von Philippi stammt von Carrizal Bajo (III. Region), einem Gebiet südlich von Copiapö, in dem Z. elegans auch in jüngster Zeit häufig gesammelt werden konnte. Einer Zusammenfassung der Gattungen Tecophilaea und Zephyra unter Zephyra, wie sie 1988 RAVENNA vorschlug, kann nicht zugestimmt werden. Wie er selbst erkannte, trennt die beiden Gattungen ein ganz wesentlicher Unterschied im generativen Bereich: Tecophilaea besitzt drei Staminodien, Zephyra dagegen nur zwei. Die relative Merkmalsarmut vieler großblütiger Monokotylen, so auch der Vertreter der Tecophilaeaceae, geben dem genannten Unterschied eine große Bedeutung. Zephyra compacta C.Ehrhart, spec. nov. Holotypus: Chile, TV Region, Prov. de Elqui: Quebrada Los Choros, Sandflächen westlich Choros Bajos, 20.10.1997, Ehrhart & Grau 97/1147 (M; Iso SGO, CONC). Planta perennis, bulbosa, plerumque pluricaulis, caulibus erectis, valde ramosis, ad 12-20 cm altis; inflorescentia sessilis, paniculata, compressa, in inflorescentiis partialibus ad 15 divisa, etiam in axillis foliorum basalium inflorescentiae partiales surgentes; internodia ad 2 cm longa; folia radicalia parte basali amplexicaulia et carnosa, ad 15 cm longa et 14 mm lata; folia media ad 12 mm lata et carnosa, folia ultima distincte minora et lineari-lanceolata. Flores numerosi albi; tepali ad 7 mm longa apice mucronata reflexa, 3 inferioribus paulum latioribus, basaliter canaliculatis et marginibus ciliatis. Capsula ad 12 mm longa et 8 mm lata. Semina acuti-elliptica ad 2,2 mm longa, atrofusca, grosse rugosa. Numerus chromosomatum On —2x'= 26. Weitere bekannte Aufsammlungen: Chile. III Regiön. Prov. de Huasco: ca. 10 km westlich Canto del Agua, Quebrada, 25.10.1997, Ehrhart & Grau 97/1281 (M). — IV Region. Prov. de Elqui: Carretera Panamericana a Choros Bajos km 5, 250 m, 21.10.1971, Marticorena, Rodriguez & Weldt 1673 (CONC 35536). 49 Zephyra compacta überdauert mit bis zu 12 cm tief im Boden verborgenen Knollen, die — ähnlich wie bei anderen Tecophilaeaceen — außen von den trockenen Resten der Leitbündel als feines Netz umhüllt sind. Von einer Knolle können mehrere Sprosse gleichzeitig ausgehen. Die oberirdischen Sprosse messen selten mehr als 15 cm, sind reich verzweigt, die Internodien der Hauptachse erreichen meist nur 2 cm Länge. Die leicht sukkulenten, flachen Blätter sind in sich rinnig gebogen und schmiegen sich jeweils etwa im unteren Drittel dem Sproß eng an; sie sind bis zu 15 cm lang und 14 mm breit. Bereits aus den Achseln der ersten grundständigen Blätter entstehen meist schon blühende Seitensprosse, so daß der gesamte oberirdische Sproß der Pflanze im Prinzip eine Infloreszenz darstellt. Diese kann sich aus bis zu 15 Teilinfloreszenzen zusammensetzen. Zumindest die basalen Teilinfloreszenzen tragen bis zu 25 Blüten. Die Tragblätter der Teilinfloreszenzen nehmen zu den distalen Knoten hin nur allmählich an Länge und Breite ab und sind hier noch bis zu 2 cm lang und 5 mm breit. Die Blüten sind immer rein weiß und messen nicht mehr als 10 mm im Durchmesser; sie sind wie die Blüten von Z. elegans leicht asymmetrisch gebaut. Von den äußeren, mit einer grünlich-fleischigen Spitze versehenen, bis zu 2,5 mm breiten und etwa 5 mm langen Tepalen ist das abaxiale Tepalum immer etwas schmäler; die leicht breiteren inneren Tepalen sind basal rinnig zusammengedrückt, der Rand ist in diesem Bereich leicht gekräuselt und mit fadigen Auswüchsen besetzt. Von den sechs Staubblättern sind im äußeren Kreis die beiden adaxialen steril, die drei des inneren Kreises sind fertil; die fertilen Staubblätter sind bis zu 2 mm lang, die Filamente sind grün und leicht fleischig; die gelb gefärbten Antheren öffnen sich apikal mit je einer großen Öffnung pro Theka; die sterilen Staubblätter bestehen nur aus den grünlichen, fleischigen Filamenten. Der monocolpate Pollen mißt bis zu 23 um in der Länge und 14 um in der Breite. Die Frucht ist eine trockenhäutige, dreikantige, vielsamige Kapsel mit kleinen Flügelleisten an den Verwachsungsnähten, die sich apikal öffnet. Der eiförmige, leicht zugespitzte, dunkelbraune Same mißt ca. 1,2 mm im Durchmesser und besitzt eine leicht runzelige Oberfläche. Die Chromosomenzahl ist 2n = 2x = 26. Vergleich mit Z. elegans Z. elegans besitzt, wie schon eingangs erwähnt, in einigen Merkmalen eine gewisse Plastizität, die durch den Vergleich von einer breiteren Besammlung und Kultivierung noch untersucht werden sollte. Vor allem die Tepalenlänge ist nicht besonders festgelegt und schwankt zwischen 12 und 20 mm, und auch die Blütenfarbe kann von überwiegend weiß bis fast vollständig blau variieren. Die geringe Tepalenlänge war auch der Hauptunterschied, der Miers zur Beschreibung von Z. amoena veranlasste. Wie sich aber gezeigt hat, variiert die Länge an ein und demselben Individuum in diesen Grenzen, so daß dieses Merkmal nicht zur Unterscheidung einer Art ausschlaggebend sein kann. Die Tepalen bei Z. compacta hingegen sind innerhalb der Population ganz einheitlich und mit Abstand kleiner, als die von Z. elegans. Die „eleganten“ Blüten von Z. elegans sind außerdem immer + flach radförmig ausgebreitet, während die Tepalen von Z. compacta an der Spitze nach hinten eingerollt sind. Die auffälligsten Unterschiede zwischen den beiden Arten zeigen sich allerdings im Wuchs: Im Gegensatz zu Z. elegans, bei der sich pro Knolle jeweils nur ein Sproß ausbildet, entwickeln sich bei Z. compacta sehr häufig mehrere Sprosse gleichzeitig. Auffällig ist außerdem der hohe, schlanke Wuchs bei Z. elegans, während die gedrungenen, kompakten Sprosse von Z. compacta sich kaum über den Boden erheben. Die prinzipiell gleichen Standortsbedingungen, die für beide Arten gelten, geben diesen Unterschieden einiges Gewicht. Bei Z. elegans hebt sich die Infloreszenz durch ein lang gestrecktes Internodium weit vom Boden ab, während dieses bei Z. compacta gestaucht ist und die ersten Teilinfloreszenzen schon aus den Achseln der untersten Laubblätter entstehen. Die schmalen, vom Sproß abspreizenden Blätter von Z. elegans, die als Tragblätter der Teilinfloreszenzen distal schnell zu sehr schmalen und nicht einmal 1 cm langen Blättchen reduziert sind, heben 50 sich deutlich gegenüber den wesentlich breiteren, dem Sproß stärker angeschmiegten Blättern von Z. compacta ab, die auch als Tragblätter der oberen Teilinfloreszenzen noch ihre breite, fleischige Gestalt behalten. Tabelle 1 gibt einen Überblick über die Merkmale der beiden Arten im Vergleich. In den ökologischen Ansprüchen unterscheiden sich die beiden Arten von Zephyra kaum. Beide kommen auf sandigem bis sandig-steinigem Boden im Bereich der Küste bis wenige km ins Landesinnere hin vor. Das Areal der neuen Art schließt sich südlich an das Areal von Z. elegans an. In einem überraschenden zweiten Fundort von Z. compacta etwa 100 km weiter nördlich des Erstfundes konnten beide Arten sympatrisch wachsend gesammelt werden — was auch für die Eigenständigkeit von Z. compacta spricht: Z. elegans direkt an der Küste in ihrer typischen schlanken, hohen Gestalt und Z. compacta etwa 15 km weiter im Landesinneren bei Canto del Agua; erstere in voller Blüte, letztere bereits fruchtend, unverkennbar allerdings in ihrer typischen gedrungenen Gestalt. Dieser zweite Fundort zeigt, daß es sich bei Z. compacta nicht um eine lokale Entwicklung, d.h. eine relativ junge Abspaltung von Z. elegans handelt, sondern daß die Art doch eine weitere Verbreitung besitzt, wobei das heutige Areal relativ zerrissen erscheint (oder schlecht besammelt ist?). Z. compacta könnte daher als eine ältere, durchaus etablierte Art betrachtet werden, die nicht eine junge, durch relativ rezente klimatische Veränderungen entstandene Entwicklung darstellt. Leider konnte die zweite, nördlichere Aufsammlung von Z. compacta noch nicht kultiviert werden; interessant wäre es jedoch zu sehen, ob sich zwischen den beiden Aufsammlungen bereits Unterschiede zeigen. Tabelle 1: Merkmalsvergleich von Z. compacta mit Z. elegans bes mean Lacoste iw LCL. ‚un. Sn Wuchshöhe 12-20 cm 25-40 cm mit mehreren Sprossen pro Knolle mit nur einem Sproß pro Knolle bis 15 cm lang, max. 14 mm breit bis 22 cm lang, max 8 mm breit im basalen Drittel den Sproß + Tragblätter der umhüllend Teilinfloreszenzen vom Sproß abspreizend immer auffällig laubblattartig entwickelt; die unteren bis zu 14 mm breit und auch die oberen an den distalen Knoten bis zu 2 cm lang und 5 mm breit unauffällig, nur das der untersten 1-2 Teilinfloreszenzen laubblattähnlich und bis zu 4 mm breit, die Tragblätter der oberen Teilinfloreszenzen nur noch fädig dünn gering: nach ein bis zwei lang- gestreckten Internodien, d.h. erst aus der zweiten oder dritten Blattachsel ein Seitentrieb der Infloreszenz auszweigend meist mit nicht mehr als 7 Teilinfloreszenzen zwischen den Teilinfloreszenzen oder rein blau Breite der Tepalen 1,5-3,5 mm bis 8 mm hoch: bereits aus den Achseln der untersten Blatter der erste Seitentrieb der Infloreszenz auszweigend Verzweigungsgrad der Infloreszenz mit bis zu 15 Teilinfloreszenzen Abb. 1: Zephyra elegans bei Carrizal Bajo (Ehrhart & Grau 97/1260 (M)) 51 Abb. 2: Zephyra compacta bei Choros Bajos (Ehrhart & Grau 97/1147 (M)) Abb. 3: Fundorte von Z. compacta: @ = Typusaufsammlung (Ehrhart & Grau 97/1147 (M)) O = Fundort bei Canto del Agua (Ehrhart & Grau 97/1281 (M)) 32 Den Konservatoren von BM möchte ich für die rasche Ausleihe der Typen von Zephyra danken. Herrn Craig Brough (Kew) danke ich für seine Hilfe in der Literaturbeschaffung. Literatur Don, D. 1832: On the characters and affinities of certain genera, chiefly belonging to the Flora Peruviana. — Edinburgh New Philos. J. 13: 233-244. PHILIPPI, R.A. 1873: Descripciön de algunas plantas nuevas incorporadas ultimamente en el herbario chileno por el Dr. Don Rodolfo A. Philippi (1). — Anales Univ. Chile 43: 479-583. Miers, J. 1864: On the Conanthereae. — Trans. Linn. Soc. London 24: 501-510. RAVENNA, P. 1988: New or noteworthy Tecophilaeaceae. — Phytologia 64(4): 288-289. Dr. Christine EHRHART, Institut fiir Systematische Botanik der Ludwig-Maximilians- Universitat Miinchen, Menzinger Str. 67, D-80638 Miinchen, Deutschland. 53 Bemerkungen zur Taxonomie der Gattung Limonium VII M. ERBEN Zusammenfassung: ERBEN, M.: Bemerkungen zur Taxonomie der Gattung Limonium VII. — Sendtnera 7: 53-84. 2001. ISSN 0944-0178. Acht neue Arten werden für die Gattung Limonium beschrieben. L. portovecchiense wächst auf Korsika. L. caralitanum, L. multifurcatum, L. capitis-eliae, L. malfatanicum und L. carisae kommen auf Sardinien vor. L. fesianum und L. comosum haben ihr Areal in Marokko bzw. in Tunesien. Da sich L. coralliforme nicht von L. ursanum un- terscheiden läßt, wird es nun als Synonym der zuletzt genannten Art geführt. Statice bollei wurde zu Limonium bollei umkombiniert. L. dubium und L. tunetanum wurden durch die Auswahl eines Lectotypus typifiziert. Abstract: Eight new species are described of the genus Limonium. L. portovecchiense grows in Corsica. L. caralitanum, L. multifurcatum, L. capitis-eliae, L. malfatanicum and L. ca- risae are from Sardinia. L. fesianum and L. comosum are locally distributed within Marocco and Tunisia. L. coralliforme does not differ from L. ursanum, therefore has been reduced to a synonym of the latter name. Statice bollei has been recombined as Limonium bollei. The typification of L. dubium and L. tunetanum is completed by the selection of lectotypes. I. Neue Taxa 1. Limonium portovecchiense Erben, spec. nov. Holotypus: Korsika, Dépt. Corse-du-Sud: Nordéstlich von Porto-Vecchio, Golfo di Sogno, ca. 2 km östlich von Sta.-Trinité an der Straße D 468, auf feucht-sandigen Flächen am Meer, 5.9.1990, Erben E 872 (MSB 85555; Isotypen: Herb. Erben). Abb.: 1-2 Planta perennis, glabra, pluricaulis. Caudiculi 10-80 mm longi, in parte superiore laxe ad dense ramosi. Folia florendi tempore persistentia, dense rosulate disposita, 15-80 mm longa et 4-12 mm lata, oblanceolata ad spatulata, acuta ad obtusa, sublaevia, 1—3-nervia, in pe- tiolum 1/3 longitudinis laminae attingentem 1-3 mm latum sensim attenuata. Caules 15-40 cm longi, erecti, flexuosi, in partibus 3/4 ad 2/3 superioribus ramosi. Rami laxe distiche di- chotomiforme dispositi, oblique sursum spectantes, sub angulis 40°-50° abeuntes, inferiores 1-3 steriles, non ramosi, 2-6 cm longi, superiores fertiles, 5-20 cm longi, recti ad sub- flexuosi, laxe secunde ad distiche ramosi. Inflorescentia paniculata, plana, forma obtrullata ad 54 cuneata. Spicae in parte 1/4 superiore inflorescentiae dispositae, 5-50 mm longae, subrectae; axis spicae subtiliter flexuosus. Spiculae 5,5—6,5 mm longae, laxe ad 2-5 in | centimetro dispositae, non vel partim mutue attingentes. Bractea inferior 1,9-2,3 mm longa et 1,9-2,0 mm lata, triangulari-ovata, acuta, margine anguste membranaceo, parte centrali subcarnosula, acuminata, acumine tenui marginem fere contingente. Bractea media membranacea, 1,9-2,2 mm longa et 1,7-2,0 mm lata, late elliptica, rotundata. Bractea superior 5,0-5,3 mm longa et 3,6-4,0 mm lata, elliptica, obtusa, dorso longitudinaliter subarcuata, margine late membrana- ceo, parte centrali subcarnosula, subdura, 3,6-4,1 mm longa et 1,9-2,2 mm lata, oblongo- elliptica, acuminata, acumine 1,0-1,2 mm longo marginem fere contingente. Calyx fere infun- dibuliformis, 5,0-5,7 mm longus, ex bractea superiore 1,0-1,5 mm exsertus, tubo imprimis in parte inferiore unilateraliter sparsim breviter piloso; dentes calycis ca. 0,4 x 1,0 mm, trans- verse late semi-elliptici; costae tubi ad vel breviter supra basim dentium calycis desinentes. Flores caeruleo-violacei, diametro 7,5-8,0 mm. Chromosomatum numerus: 2n = 27. Pflanze ausdauernd, kahl, mehrstengelig. Stimmchen 10-80 mm lang, am oberen Ende locker bis dicht verzweigt. Blätter dicht rosettig angeordnet, bei älteren Pflanzen die unteren braun und verwelkt, die oberen grün und lebend, ganzrandig, 15-80 mm lang und 4-12 mm breit, oblanzeolat bis spatelförmig, am oberen Ende spitz bis stumpf, an den Rändern sehr schmal weißhäutig, nahezu glatt, kleinere Blätter 1-, größere 3-nervig, allmählich in einen 1-3 mm breiten, ca. 1/3 der Spreitenlänge erreichenden Stiel übergehend. Stengel 15-40 cm lang, aufrecht, mehr oder weniger zickzackförmig gebogen, ab dem unteren Viertel bis Drittel ver- zweigt. Äste locker dichotomartig abzweigend, meist zweiseitswendig angeordnet, schräg nach oben gerichtet (Verzweigungswinkel 40°-50°), die unteren 1-3 steril, unverzweigt, 2-6 cm lang, die oberen fertil, 5-20 cm lang, gerade bis schwach zickzackförmig gebogen, locker ein- oder zweiseitswendig verzweigt. Infloreszenz flach, rispenförmig, im Umriss obtrullat bis dreieckig-eiförmig. Ähren im oberen Viertel der Infloreszenz sitzend, 5-50 mm lang, nahezu gerade; Ährenachse fein zickzackförmig gebogen. Ährchen 5,5-6,5 mm lang, 2-4-blütig, locker zu 2-5 pro Zentimeter angeordnet, sich gegenseitig nicht oder nur teilweise berührend. Äußere Braktee 1,9-2,3 mm lang und 1,9-2,0 mm breit, dreieckig-eiförmig, am oberen Ende spitz, häufig leicht gekielt; Rand schmal weißhäutig; zentraler Bereich etwas fleischig, mit einer feinen, fast bis zum Rand reichenden Spitze. Mittlere Braktee häutig, 1,9-2,2 mm lang und 1,7-2,0 mm breit, breit elliptisch, am oberen Ende abgerundet. Innere Braktee 5,0-5,3 mm lang und 3,6-4,0 mm breit, elliptisch, am oberen Ende stumpf, am Rücken in Längs- richtung schwach bogenförmig; Rand breit weißhäutig; zentraler Bereich etwas fleischig und ziemlich hart, 3,6-4,1 mm lang und 1,9-2,2 mm breit, länglich-elliptisch, mit einer 1,0-1,2 mm langen, fast bis zum Rand reichenden Spitze. Kelch annähernd trichterförmig, 5,0-5,7 mm lang, die innere Braktee um 1,0—1,5 mm überragend; Kelchröhre nur in der unteren Hälf- te auf einer Seite spärlich kurz behaart; Kelchzähne sehr zart, ca. 0,4 x 1,0 mm, transvers breit halbelliptisch; Rippen der Kelchröhre an oder kurz unterhalb der Kelchzahnbasis endend. Blüten blau-violett, mit einem Durchmesser von 7,5-8,0 mm. Chromosomenzahl: 2n = 27; untersucht wurden Exemplare der Aufsammlungen Li-574, Li-1258, Li-1274, Li-1315 und Li-1519. Limonium portovecchiense ist sicher näher mit L. glomeratum (Tausch) Erben verwandt. Da beide Arten in mehreren Merkmalen übereinstimmen, die sie von anderen Sippen stärker trennen, erschien mir zunächst eine Zusammenfassung beider Sippen zu einer Art die sinn- vollste Lösung zu sein. Nun haben aber Kulturversuche mit L. portovecchiense und L. glome- ratum gezeigt, dass sich beide Arten in Kultur noch viel deutlicher von einander unterschei- den als an ihren Wildstandorten. Da diese Eigenschaften also genetisch fixiert sein dürften, erscheint mir auch die Behandlung dieser Sippen als zwei selbständige Arten gerechtfertigt. 35 Betas theories BEM Az, Bo MC REA EO 86 HERnARICM MM. ERDEN Eimanium parwvecchiense Erben, spec uva km aztlich vor Tiotlo di Soma, za Tremolecia atrata Figure 11: Subantarctic distribution of Tremolecia atrata (based on specimens seen). Tremolecia atrata is tiny but usually easily recognizeable species. It is not yet known from continental Antarctica, Bouvet Island, Macquarie Island, Campbell and Auckland Islands and from the North Island of New Zealand. Tremolecia atrata, an alpine element, seems to be con- fined to areas with a humid climate. I am grateful to Dr. Thomas H. Nash III (Tempe, AZ, U.S.A) and Dr. David Galloway (Rox- burgh, Central Otago, N.Z.) for helpful comments and Dr. Christian Leuckert (Berlin) for the chemical analysis of various lichen specimens. Spezial thanks are due to Dr. Alan Prather and Dr. Alan Fryday for their invitation to visit the Michigan State Herbarium, East Lansing, (courtesy of U.S. National Science Foundation award DBI-9808735), where Dr. Henry A. Imshaug’s very rich and valuable lichen collections from Subantarctic and alpine North Ameri- can areas are housed. 131 Literature AHTI, T., SCOTTER, W. & VANSKA, H. 1973: Lichens of the Reindeer Preserve, Northwest Ter- ritories, Canada. — Bryologist 76: 48-76. ANDERSON, R.A. 1962: The lichen flora of the Dakota sandstone in North-Central Colorado. — Bryologist 65: 242-261. — 1964: The genus Lecidea (Lichenized Fungi) in Rocky Mountain National Park, Colorado. — Univ. 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Es wird der Infloreszenzbau von Vertretern aller Gattungen der Juncaceae s.l. (Juncus, Luzula, Prionium, Marsippospermum, Rostkovia, Distichia, Oxychloe und Patosia) analysiert. In der Literatur wurden die Blütenstände der Familie bisher mit Begriffen wie z.B. Rispe (Panicula), Trichterrispe (Spirre, Anthela) oder Drepanium (Sichel) beschrieben. Da diese Termini monotelen Synfloreszenzbau suggerieren, wird der Frage nachgegangen, ob bei den Juncaceen tatsächlich monotele Synfloreszenzen vor- kommen. Bei Prionium und bei Juncus subgen. Juncus ist polyteler Bau offenkundig, da die Baueinheiten (Floreszenzen) des Blütenstandes mehrblütige, offene Ähren bzw. Köpfchen darstellen. In Juncus subgen. Poiophylli sowie bei allen anderen Gattungen finden sich distal an jeder Achse Einzelblüten, die an ihrer Basis von zwei Brakteen begleitet werden. Diese Blüten wurden bisher als terminal, die sie begleitenden Brak- teen als Vorblätter bezeichnet. Es werden jetzt Befunde vorgelegt, die diese Ein- zelblüten mitsamt ihren Brakteen eindeutig als einblütige, terminal offene Ährchen ausweisen. Damit ist für die gesamte Familie polyteler Synfloreszenzbau nachgewie- sen. Reduktionstendenzen führen einerseits zur Beschränkung der Blütenbildung auf die Hauptfloreszenz (mehrblütig z.B. bei Juncus triglumis, einblütig bei Marsipposper- mum und Rostkovia), andererseits zur Prolifikation und Blütenbildung nur noch an Paracladien (Oxychloe, Patosia und Distichia). Bei Luzula sect. Gymnodes sind die Paracladien differenziert in distale Kurzparacladien, die zu einem Ährchen 2. Ordnung zusammentreten, und proximale Langparacladien, die das Verhalten der distalen Re- gion der Hauptachse wiederholen. Das Achsensystem des Gesamtblütenstandes ist bei Prionium dem einer monotelen Kegelrispe analog, die Infloreszenz muss korrekt als Paniculodium bezeichnet werden. Bei reicher verzweigten Juncus-Arten sowie bei den Sektionen Anthelaea und Pterodes von Luzula entspricht das Achsensystem der Infloreszenz dem einer monotelen Trichterrispe, der Blütenstand ist ein Anthelodium. Das Drepanium ist ein Spezialfall des Verzweigungsmusters eines Paracladiums. Es entsteht durch die monochasiale Verkettung von Sympodialgliedern, die jeweils außer einem adossierten Vorblatt nur ein weiteres Blatt und distal ein einblütiges Ährchen tragen. Das Drepanium wurde nur in den Sektionen Caespitosi und Tenageia von Juncus gefunden. Infloreszenzen nach dem Muster des Paniculodiums und des Anthelodiums, die starke Tendenz zur Bildung einblütiger Ährchen und deren Zusammentreten zu Ährchen 2. Ordnung stimmen bis in viele Details mit den entsprechenden Mustern der Cyperaceae überein. 138 Abstract: A detailed analysis of the inflorescence structure of members of all genera of Jun- caceae s.]. is presented (Prionium, Juncus, Luzula, Marsippospermum, Rostkovia, Oxychloe, Patosia, Distichia). In the literature the inflorescences of the family are mostly interpreted as panicles or anthelas, details are described using vaguely defined terms aS cymose, racemose, or drepanium. Most of these terms suggest a monotelic structure of the inflorescence, which would be very unusual among derived Monocots. This problem stimulated the present study. The basic elements of the inflorescence in Prionium and Juncus subgen. Juncus are many-flowered, terminally open spikelets or small heads (fig. 13a). In these cases the complete system is undoubtedly a polytelic synflorescence. In Juncus subgen. Poiophylli and in all the other genera the main axis, as well as all branches, bear a distal, solitary flower embraced with two bracts. This structure represents an one-flowered spikelet. The only flower develops in the axil of the lower bract and takes a pseudoterminal position. The axil of the upper bract is empty. These one-flowered spikelets are arranged in Juncus subgen. Poiophylli and in the sections Anthelaea and Pterodes of Luzula in the same manner as the multi- flowered spikelets in Prionium and Juncus subgen. Juncus (fig. 13b). Thus all mem- bers of the family have polytelic synflorescences. At family level several tendencies of reduction of the branching system are found. All paracladia may be suppressed, flowers are developed only in the main florescence, either multi-flowered, e.g. in Jun- cus triglumis (fig. 13d), or one-flowered, as in Marsippospermum and Rostkovia (fig. 13e). In other cases the main florescence fails to develop, the vegetative main axis grows permanently, and flowers are developed only at paracladia (Distichia, Rostko- via, Patosia; fig. 13f). Luzula sect. Gymnodes has two kinds of paracladia. The distal ones are short paracladia, forming a spike of second order, the proximal ones are long paracladia, which repeat the behaviour of the distal area of the main axis (fig. 13c). The branching system of the complete inflorescence of Prionium is analogous to that of a monotelic panicle, the inflorescence is correctly termed a paniculodium, i.e. the polytelic analogon to the monotelic panicula. The complete inflorescences of most Juncus species and of the sections Anthelaea and Pterodes of Luzula are analogous to the monotelic anthela and have to be termed an anthelodium. The drepanium is a specialized case of a branching system of a paracladium. The sympodial elements of it consist of an axis with an adaxial prophyll, one further bract and a distal, obviously always one-flowered, spikelet. In our material we have found a drepanium only in the sections Caespitosi and Tenageia of Juncus. Among Cyperaceae inflorescences similar to those of the Juncaceae are found. These include typical paniculodia, anthelodia, and one-flowered spikelets, which are often crowded forming a spikelet of second order. 1. Einleitung In der traditionellen Umgrenzung gehören zu den Juncaceae die beiden weltweit verbreiteten Gattungen Juncus und Luzula sowie die südhemisphärischen Gattungen Distichia, Marsippo- spermum, Oxychloe, Patosia, Rostkovia und Prionium. Das in der Kapprovinz endemische Prionium wurde kürzlich von MUNRO & LINDER (1998) als monotypische Familie Prioniaceae von den Juncaceae abgetrennt. Die reichbliitigen Infloreszenzen von Juncus, Luzula und Prionium werden allgemein als Rispe (Panicula) bzw. Spirre (Anthela, Trichterrispe) bezeichnet. Rispe und Spirre sind eindeutig definiert als monotele Synfloreszenzen (vgl. TROLL 1964b, S. 152). Eine erhebliche Verwirrung der Terminologie entsteht durch die Benutzung des Begriffspaares cymos/ racemos für die Juncaceen-Blütenstände. NOVARA (1976) beschreibt die Infloreszenzen von Juncus Subgenus Poiophylli als cymos, die des Subgenus Juncus als racemos. Diese Terminologie übernehmen auch SNOGERUP (1993) und BALSLEV (1996). Bezüglich der prazisen Definition und Verwendung der Begriffe cymos bzw. Cyme sei auf die Arbeit von MULLER-DOBLIES et al. (1992) verwiesen. BALSLEV (1998) verstarkt die begriffliche Ver- 139 wirrung noch, wenn er meint, einige Juncus-Arten hätten „determinate sympodial panicles“ (!). Monotelie ist auch zu vermuten, wenn er angibt: „Rostkovia and Marsippospermum have a solitary, perfect, terminal flower“. Ein weiteres Problem liegt in dem für einige Juncus-Arten so typischen Drepanium (Sichel). MULLER-DOBLIES et.al. (1992) haben zwar geklärt, dass dieses sympodiale System nicht zu den Cymen gehört, da die Verzweigung nicht aus der Achsel eines Vorblattes erfolgt. Andererseits bezweifeln sie nicht, dass die distale Bltite jedes Sympodialgliedes eine termi- nale Position hat und vermuten, dass das Drepanium von einem reicher verzweigten System herzuleiten ist: „the drepanium of Juncus has in any case to be derived from richer branched, i.e. paniculate structures with a racemose branching. Die Frage nach der möglichen Mono- telie bleibt damit wiederum offen. Zweifel am Vorkommen von Monotelie bei den Juncaceen ergeben sich zunächst aus der Tatsache, dass die Teilblütenstände einiger Juncus-Arten einfache, offene Ähren bzw. Köpf- chen (Floreszenzen) darstellen. Diese Arten weisen einen ganz klar polytelen Synfloreszenz- bau auf. Es müssten also innerhalb einer Familie und sogar innerhalb einer Gattung monotele und polytele Synfloreszenzen nebeneinander vorkommen. BUCHENAU (1890) hat die beiden Verzweigungstypen bereits genau erkannt und die Arten der Gattung Juncus daraufhin in die informellen Gruppen Flores prophyllati und Flores eprophyllati untergliedert. VIERHAPPER (1930) blieb bei dieser Gliederung. Erst KIRSCHNER et al. (1999) haben den Sachverhalt nomenklatorisch bereinigt und die Subgenera Juncus und Poiophylli mit insgesamt zehn Sektionen beschrieben (s. Tab. 1). Tabelle 1: Die taxonomische Gliederung der Gattung Juncus L. (nach KIRSCHNER et al. 1999). Table I: The taxonomic subdivision of the genus Juncus L. (after KIRSCHNER et al. 1999). Juncus L. subgenus Juncus Juncus L. subgenus Poiophylli Buchenau sect. Juncus 7. sect. Tenageia Dumort. sect. Graminei (Engelm.) Engelm. 8. sect. Steirochloa Griseb. sect. Caespitosi Cout. 9. sect. Juncotypus Dumort. sect. Stygiopsis Kuntze 10. sect. Forskalina Kuntze sect. Ozophyllum Dumort. sect. /ridifolii Snogerup & Kirschner GN) ON) =a Beim Subgenus Poiophylli trägt jeder blattachselständige, einbliitige Ast ein grundstän- diges, adossiertes Vorblatt sowie, direkt unter der distalen Blüte, zwei weitere Brakteen, die in der Bestimmungsliteratur fälschlich ebenfalls als Vorblätter bezeichnet werden. Beim Sub- genus Juncus besitzen die Einzelblüten des mehrblütigen Ährchens keine Blattorgane an ihrem Blütenstiel, sie stehen direkt in der Achsel eines Tragblattes. BUCHENAU (1890) hatte sich schon Gedanken über die phylogenetischen Beziehungen zwischen den beiden Inflores- zenztypen gemacht. Er hält zunächst die flachblättrigen Juncaceae für die ursprünglichen Sippen. Die Blüten vieler Arten dieser Gruppe sind durch je zwei Brakteen, die auch BUCHENAU als Vorblätter bezeichnet, eingehüllt. Als spezielle Modifikation findet man in der Gattung Luzula z.T. das Phänomen der Trunkation der Hauptachse und, bei manchen Blüten, gleichzeitig den Ausfall der oberen Braktee. BUCHENAU sieht darin ein Indiz für den Über- gang vom Infloreszenztyp des Subgenus Poiophylli zu dem des Subgenus Juncus: „In dieser Richtung ist es nun bedeutungsvoll, dass bei den Luzu/a-Arten, deren Blüten in Ähren oder Köpfchen zusammengedrängt sind, sich eine Neigung zum Schwinden der Vorblätter zeigt. Wenn dieses Schwinden fortschritte, so würden zuletzt vorblattlose Blüten in den Achseln von Hochblättern vorhanden sein. Aus diesem Grund glaube ich annehmen zu dürfen, dass die Juncus-Arten mit vorblättrigen Blüten die ältere Gruppe bilden. Ich nehme daher auch an, 140 dass der Urtypus der Juncaceen vorblättrige Blüten hatte und dass er also den Juncis poio- phyllis entsprach“ (S. 53f.). Übertragen in die heutige Terminologie bedeutet dies: Der Grundtypus der Infloreszenz der Juncaceen besitzt aus den Achseln von Tragblättern entspringende Paracladien, die an ihrer Basis je ein adossiertes Vorblatt tragen und in einer Terminalblüte enden. Direkt unterhalb dieser Blüte stehen zwei Brakteen. Durch den Ausfall der beiden Brakteen und des Vorblattes, zusätzliche Trunkation der Hauptachse und Verkürzung der Internodien der Seitenäste sind die Ähren bzw. Köpfchen der Untergattung Juncus entstanden (Abb. la, b). Sie entsprächen damit einer monotelen Rumpfsynfloreszenz. Das vermeintliche Vorkommen der phylogenetisch ursprünglichen Monotelie inmitten der ganz überwiegend polytelen Monocotylen (Ausnahmen wohl nur bei wenigen Alismatidae und den Burmanniaceae, s. TROLL 1964a, 1969) hat eine gründlichere Untersuchung angeregt. Mit der vorliegenden Arbeit wird versucht, Antwort auf zwei Fragen zu geben: l. Kommen bei den Juncaceae tatsächlich monotele Synfloreszenzen vor? 2. Welche phylogenetische Beziehung besteht zwischen den verschiedenen Infloreszenzen der Familie? 2. Material und Methoden Die Untersuchungen erfolgten zum größten Teil an frischen Pflanzen oder an konserviertem Material (70% Ethanol), das von C. P. Köbele 1999-2000 gesammelt worden ist. Von diesen Herkünften wurden Belegexemplare im Herbarium des Instituts für Systematische Botanik der LMU-München (MSB) hinterlegt. Zusätzlich wurde Herbarmaterial aus der Botanischen Staatssammlung München (M) untersucht. Das für die vorliegende Arbeit herangezogene Ma- terial ist in Tab. 2 zusammengestellt. Es umfasst alle Gattungen der Familie, aus der Gattung Juncus konnten Beispiele aus 8 der insgesamt 10 Sektionen berücksichtigt werden. Für die morphologischen Analysen und zum Zeichnen wurde ein Stereomikroskop Leica Wild MZ 8 mit Zeichenspiegel verwendet. Die Aufriss-Diagramme der Infloreszenzen wur- den mit dem Programm CorelDraw am Computer erstellt. Um die Übersichtlichkeit zu ge- währleisten, werden alle Achsen mit gestreckten Internodien dargestellt, Achsenstauchungen und Übergipfelungen bleiben in den Schemata weitgehend unberücksichtigt. Über die ver- wendeten Symbole gibt Abb. Ic Auskunft. Um die Orientierung in den z.T. sehr komplexen Infloreszenzen zu erleichtern, sind alle Blätter und Blüten der Hauptachse jeweils schwarz ausgefüllt. Rudimentäre Blüten sind nur als Blütenstiel dargestellt. Tabelle 2: Das Untersuchungsmaterial. Table 2: The investigated material. Herkunft und Beleg Distichia muscoides Nees & Meyen | Bolivien, Depto. La Paz, T. Feuerer & H. Preiss Nr. 4760a, 10.8.1980. Juncus acutiflorus Ehrh. ex Hoffm. Deutschland, Bayern, Landkr. FFB, Köbele Nr.23. Juncus articulatus L. em. Richter Deutschland, Bayern, Landkr. FFB, Köbele Nr.24 u. 40. Marokko, Hoher Atlas, Oukaimeden, Köbele Nr. 31. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 38. Südafrika, Cape Province, W. Griess Nr. 1214, 10.11.1946. Deutschland, Bayern, Mittelfranken, O. Prechtelsbauer s. n., Juli 1886. Deutschland, Bayern, Unterfranken, Aug. Vill s. n., 8.8.1896. Österreich, Kärnten, Joh. Drobny s. n., 21.7.1922. Österreich, Tirol, Merxmiiller & Wiedmann s. n., 21.7.1957. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 11, 18, 19 u. 20. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 6, 10 u. 14. Juncus articulatus L. em. Richter Juncus bufonius L. Juncus capensis C. P. Thunberg Juncus capitatus Weigel Juncus capitatus Weigel Juncus castaneus SM. Juncus castaneus SM. Juncus conglomeratus L. em. Leers Juncus inflexus L. 141 Herkunft und Beleg Juncus inflexus L. Marokko, Hoher Atlas, Köbele Nr. 32 u. 34. Juncus inflexus L. Marokko, Mittlerer Atlas, Köbele Nr. 35. Juncus maritimus L. Marokko, Antiatlas, Köbele Nr. 28. Juncus maritimus L. Marokko, Hoher Atlas, Köbele Nr. 33. Marokko, Mittlerer Atlas, Köbele Nr. 36. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 21 u. 39. Deutschland, Bayern, Allgäuer Alpen, E. Dörr s. n., 21.8.1970. Deutschland, Bayern, Blauberge, W. Lippert & J. Sellmair s. n., 8.9.1976 Deutschland, Bayern, Landkr. STA, Köbele Nr. 15. Deutschland, Sachsen-Anhalt, Harz, Köbele Nr. 25. Italien, Trentino, Köbele Nr. 27. kultiviert im Bot. Garten München, Köbele Nr. 41 Deutschland, Bayern, Landkr. BGL, Köbele Nr. 26. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 1, 2, 9 u. 12. Juncus punctorius L. fil. Juncus tenuis Willd. Juncus triglumis L. Juncus triglumis L. Luzula campestris (L.) DC. Luzula campestris (L.) DC. Luzula campestris (L.) DC. Luzula forsteri (SM.)DC. Luzula glabrata (Hoppe) Desv. Luzula luzuloides (Lamk.) Dandy & Wilmott Luzula multiflora (Ehrh.)Ley. Luzula pilosa (L.) Willd. Luzula pilosa (L.) Willd. Marsippospermum grandiflorum J. D. Hooker Oxychloe andina Phil. Patosia clandestina (Phil.) Buch. Prionium serratum L. fil. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 8. Deutschland, Bayern, Landkr. FFB, Köbele Nr. 3, 4, 7 u. 17. Deutschland, Bayern, Landkr. STA, Köbele Nr. 16. Chile, Prov. Nuble, E. Werdermann Nr. 302, Marz 1927. Chile, Prov. Atacama, E. Werdermann Nr. 937, Jan. 1926. Chile, Prov. Malleco, H. Merxmiiller s. n., 27.12.1968. Siidafrika, Cape Province, P. Goldblatt Nr. 3277, 3.11.1974. South Georgia Islands, Vill Nr. 26, 8.1.1983. Rostkovia magellanica (Lamk.) J. D. Hooker 3. Ergebnisse 3.1. Allgemeines zum Bau der Infloreszenzen Bei den reich verzweigten Arten von Juncus und Luzula nimmt, ebenso wie bei Prionium, der Verzweigungsgrad der Paracladien von proximal nach distal kontinuierlich ab. Die Trag- blatter der unteren Paracladien sind noch laubblattartig (frondos) gestaltet, zur Infloreszenz- spitze hin werden sie rasch kleiner und erreichen schlieBlich brakteose Gestalt. Jedes Para- cladium trägt basal ein meist deutlich zweikieliges, adossiertes Vorblatt. Das Epipodium ist bei Juncus und Luzula meist deutlich gestreckt, wodurch die Abstammungsachse übergipfelt wird (Abb. Ic). In der Gattung Prionium fehlt die Übergipfelung. Bei stärkeren Paracladien sind zwischen das Vorblatt und die unter den Blüten stehenden Brakteen noch weitere Blätter eingefügt. Wie an der Hauptachse sind diese Blätter proximal noch frondos und werden distal brakteos. Aus den Achseln dieser Blätter erfolgt weitere Verzweigung. Nur bei wenigen Ar- ten entspringt auch aus der Vorblattachsel ein Bereicherungstrieb. Die Ährchen des Subgenus Juncus sind nach einem einheitlichen Prinzip gebaut. Jede Blüte steht in der Achsel eines Tragblattes (Braktee). Auf das letzte fertile Tragblatt folgen entweder mehrere Brakteen, in deren Achseln nur verkümmerte und distal schließlich über- haupt keine Blüten stehen, oder nur eine einzige sterile Braktee ohne Spur einer Blüte. Die Zahl der Blüten pro Ährchen nimmt an Paracladien steigender Ordnung kontinuierlich ab. Die letzten Verzweigungen reichblütiger Infloreszenzen entwickeln oft nur noch Floreszenzen mit drei oder zwei Blüten. Immer folgt aber der obersten Blüte noch mindestens eine sterile Brak- tee. Im Extremfall wird die Floreszenz auf eine einzige Blüte reduziert. Dies wird am Beispiel von Juncus maritimus in Abb. 2 demonstriert. Das einblütige Ährchen ist nicht von den mit zwei Brakteen umhüllten Blüten des Subgenus Poiophylli oder der Gattung Luzula zu unter- 142 scheiden. Schon diese Beobachtung führte uns zu der Vermutung, dass in der Familie die mit zwei Brakteen umgebenen Einzelblüten als einblütige Ährchen zu interpretieren sind. In den Verzweigungsschemata ist diese Interpretation bereits berücksichtigt (s. Abb. Ic). Zusätzliche Argumente werden weiter unten in der Diskussion vorgestellt. Unter Zugrundelegung dieser Arbeitshypothese können die Gattungen mit reichblütigen Infloreszenzen (Juncus, Luzula, Prionium) in zwei Hauptgruppen zusammengefasst werden (Abb. 3). Infloreszenzen der Hauptgruppe 1 besitzen mehrblütige Ährchen, diejenigen der Hauptgruppe 2 nur einblütige Ährchen. Subtypen sind in beiden Fällen danach zu unter- scheiden, ob zumindest kräftige Exemplare eine Verzweigung aus der Achsel des adossierten Vorblattes hervorbringen (Gruppe la bzw. 2a), oder ob die Vorblattachsel immer steril bleibt (Gruppe 1b bzw. 2b). Wenn ein Paracladium aus der Achsel eines Vorblattes gebildet wird, bleibt es stets deutlich schwächer als die aufwärts folgenden. In der Hauptgruppe 2 tritt zu- sätzlich noch eine dritte Variante auf, bei der einblütige Ährchen zu dichten Ähren 2. Ord- nung zusammentreten (Gruppe 2c, s. Tab. 3). Tabelle 3: Die Zuordnung der im Text behandelten Juncaceen-Taxa zu den beschriebenen Inflores- zenzformen. Table 3:Assignment of the taxa dealt with in the text to the inflorescence types. Hauptgruppe | vielblütige Ährchen Gruppe la Gruppe 1b mit Verzweigung aus ohne Verzweigung aus der Vorblatt-Achsel der Vorblatt-Achsel Juncus subgen. Juncus sect. Juncus sect. Graminei J. maritimus J. capensis sect. Caespitosi sect. Stygiopsis J. capitatus J. castaneus J. triglumis sect. Ozophyllum J. acutiflorus é articulatus J. punctorius Gruppe 2a Gruppe 2b Gruppe 2c mit Verzweigung aus er Verzweigung aus Floreszenzen 2. Ordnung der Vorblatt-Achsel der Vorblatt-Achsel Juncus subgen. Poiophylli sect. Tenageia sect. Steirochloa J. bufonius J. tenuis sect. Juncotypus J. conglomeratus Luzula sect. Anthelaea Luzula sect. Gymnodes L. glabrata L. campestris L. luzuloides L. multiflora Luzula sect. Pterodes L. forsteri L. pilosa 143 3.2. Infloreszenzen mit mehrblütigen Ährchen (Hauptgruppe 1) Durch Blütenstände dieses Typs sind die Gattung Prionium sowie Juncus subgenus Juncus charakterisiert. Gruppe la. Die Blütenstände sind dadurch gekennzeichnet, dass die Paracladien auch aus der Achsel von Vorblättern Seitenäste entwickeln können. Dieses Verhalten findet sich be- sonders an den proximalen Paracladien bei Arten mit reich verzweigten Infloreszenzen. Als Beispiel ist der Blütenstand von Juncus maritimus (Sektion Juncus) wiedergegeben (Abb. 4). Bei Arten mit nur wenig verzweigten Blütenständen der Sektion Caespitosi wurden Vorblatt- Achselsprosse nur vereinzelt festgestellt. Sie treten fakultativ bei besonders kräftigen Indi- viduen an einzelnen Paracladien auf. Der annuelle Juncus capitatus besitzt stets nur ein ein- ziges Paracladium dicht unterhalb der Hauptfloreszenz (Abb. 5a, b). An kräftigen Individuen kann aus dessen Vorblatt-Achsel ein Paracladium 2. Ordnung hervorgehen (Abb. Sc, d). Dies wurde auch schon von BUCHENAU (1865) und von v. LAMPE (1996) beschrieben. Ein insge- samt vermutlich ursprüngliches Verhalten zeigt die Gattung Prionium. Die Hauptachse der Infloreszenz und die stärkeren Paracladien haben gestreckte Internodien und bleiben domi- nant gegenüber allen ihren Seitenachsen. Die für Juncus- und Luzula-Arten so typische Über- gipfelung fehlt hier noch vollkommen. Die Hemmung der Seitenachsen höherer Ordnung führt dazu, dass deren Floreszenzen zu dichten Knäueln zusammengedrängt werden (Abb. Se, Gruppe 1b. Bei den hierher gehörenden Blütenständen wurden selbst bei reichlicher Ver- zweigung keine Seitenachsen in der Achsel von Vorblättern gefunden. Dieses Verhalten trifft in der Gattung Juncus auf die untersuchten Arten der Sektionen Graminei, Ozophyllum und Stygiopsis zu. Nicht nur im Bau der Blütenstände, sondern auch bei den vegetativen Merk- malen sind die Grenzen zwischen diesen drei Sektionen fließend. Die Abgrenzung gegen- einander ist schwierig (vgl. KIRSCHNER et al. 1999). In Abb. 6a ist als Beispiel ein für die ganze Gruppe typischer Blütenstand von Juncus articulatus (sect. Ozophyllum) im sche- matischen Aufriss wiedergegeben. Durch eine oft nur geringe Neigung zur Verzweigung zeichnen sich die in Mitteleuropa heimischen Vertreter der Sektion Stygiopsis aus (Abb. 6b). Im Extremfall unterbleibt die Bildung von Paracladien vollkommen, so dass der gesamte Blütenstand auf die Hauptfloreszenz beschränkt ist. Ein bekanntes Beispiel hierfür ist der alpine Juncus triglumis (sect. Stygiopsis). Bei dieser Art fällt auf, dass hier, im Gegensatz zu den meisten Arten der Gattung, die Blüten häufig deutlich gestielt sind (Abb. 6c). 3.3. Infloreszenzen mit einblütigen Ährchen (Hauptgruppe 2) Einblütige Ährchen sind kennzeichnend für die Gattung Luzula sowie für Juncus subgenus Poiophylli. Außerdem gehören hierher alle fünf südhemisphärischen Gattungen der Juncaceae s.str. Bei diesen tritt allerdings die Besonderheit auf, dass die gesamte Infloreszenz auf ein einziges einblütiges Ährchen reduziert ist. Deshalb sollen diese Gattungen gesondert bespro- chen werden (s. u.). Gruppe 2a. Typische Beispiele für diesen Infloreszenztyp finden sich in der Sektion Junco- typus, zu der die weit verbreiteten Arten Juncus effusus, J. inflexus und J. conglomeratus gehören. Sie sind durch reichblütige Infloreszenzen gekennzeichnet, bei denen Seitenzweige auch aus der Achsel der Vorblätter proximaler Paracladien gebildet werden (Abb. 6d, 7). Erst an distalen Paracladien bzw. an Paracladien höherer Verzweigungsordnung bleiben die Vor- blätter steril. Außer in der Sektion Juncotypus wurde dieses Verzweigungsverhalten nur noch bei der aus annuellen Arten bestehenden Sektion Tenageia beobachtet. So sind z.B. bei Jun- cus bufonius an nicht zu schwachen Exemplaren regelmäßig Vorblatt-Achselsprosse zu beobachten (Abb. 8). Dies ist eine interessante Parallele zu der ebenfalls aus annuellen Arten bestehenden Sektion Caespitosi (Subgenus Juncus), bei der auf der Basis der Infloreszenz- Gruppe la ebenfalls die Vorblattachseln fertil sein können (vgl. Abb. 5c, d). Hier bietet sich die Spekulation an, dass Annuelle den Nachteil ihrer kurzen Lebensdauer durch Aus- 144 schöpfung aller Verzweigungsmöglichkeiten und damit maximaler Blüten- und Samenpro- duktion ausgleichen. Gruppe 2b. Unter den Juncus-Arten mit einblütigen Ährchen wurden Infloreszenzen mit fehlender Verzweigung aus den Vorblattachseln in unserem Material nur bei J. tenuis (Sect. Steirochloa) gefunden (Abb. 9a). Für die Gattung Luzula scheint dieses Merkmal dagegen konstitutiv zu sein. Selbst sehr reich verzweigte Infloreszenzen, wie die von Luzula glabrata (Sektion Anthelaea) zeigten niemals eine Verzweigung aus der Achsel von Vorblättern (Abb. 9b). In der Sektion Prerodes ist der Verzweigungsgrad geringer. Da die Epipodien der Para- cladien sich erheblich strecken, wirken die Blütenstände sehr locker (Abb. 10a). Die Vor- blattachseln bleiben aber auch hier stets steril. Gruppe 2c. In der Zuzula-Sektion Gymnodes weisen die Blütenstände einen von den übri- gen Vertretern der Gattung abweichenden, offenbar stark abgeleiteten Bau auf. Die Paracla- dien sind in zwei scharf voneinander getrennte Typen geschieden: Im distalen Bereich der Synfloreszenz stehen Kurzparacladien, die jeweils nur ein Vorblatt und das einblütige Ähr- chen besitzen. Insbesondere an den distalen Gliedern fällt häufiger die obere, sterile Braktee des Ährchens aus. Die Kurzparacladien stehen dicht gedrängt und bilden dadurch ein Ährchen bzw. ein Köpfchen 2. Ordnung. Proximal folgen Langparacladien, die das Verhalten des distalen Abschnitts der Hauptachse wiederholen. Ihr Vorblatt ist jeweils nahe der Basis in- seriert. Auf das verlängerte Epipodium folgen zahlreiche brakteose Tragblätter, in deren Ach- seln Kurzparacladien nach dem Muster der Hauptachse stehen (vgl. Abb. 3, 10d). Als weitere Besonderheit ist festzuhalten, dass das terminale Ährchen der Hauptachse und der Langpara- cladien häufig nicht mehr ausgebildet wird. Stattdessen findet man im Distalbereich der Achse nur noch einige sterile Brakteen (Abb. 10c, d). Es handelt sich hier um eine fakultative Trunkation der Synfloreszenz, die oft auch auf die Langparacladien übergreift. 3.4. Die Infloreszenzen der südhemisphärischen Gattungen Rostkovia und Marsippospermum. Die beiden circum-antarktisch verbreiteten Gattungen sind einander habituell sehr ähnlich. Sie tragen eine einblütige Infloreszenz, die durch ein schaft- artiges Internodium weit emporgehoben wird (Abb. 11a). Die Blüte wird von zwei Brakteen begleitet. Bei Rostkovia ist die untere Braktee laubblattartig gestaltet und überragt die Blüte deutlich (Abb. 11b). Marsippospermum fällt durch den Besitz der größten Blüten der Familie auf, trägt unter der Blüte aber nur eine oder zwei winzige Brakteen (Abb. 11c). Die Inflores- zenz ist in beiden Fällen als einblütiges Ährchen zu interpretieren. Die gesamte Synfloreszenz ist auf die einblütige Hauptfloreszenz reduziert, Paracladien werden nicht mehr angelegt. Distichia, Oxychloe und Patosia. Die drei Gattungen sind als Polsterpflanzen auf hoch- andine Lebensräume beschränkt. Bis auf eine Oxychloe-Art sind sie alle diözisch. Die einblü- tigen Infloreszenzen aller drei Gattungen entspringen jeweils der Achsel eines Laubblattes im distalen Bereich des vegetativen Sprosses. Bei Distichia und Patosia wird die männliche In- floreszenz durch ein lang gestrecktes Epipodium deutlich über das Polster emporgehoben (Abb. 11f). Die weibliche Infloreszenz bleibt dagegen in der Blattachsel verborgen, es werden nur die Narben im freien Luftraum exponiert. Bei Oxychloe sind die Blüten beiderlei Ge- schlechts etwa gleich lang gestielt (Abb. 11d). Direkt unterhalb der Blüte stehen in der Regel zwei Brakteen, Patosia besitzt aber nur noch eine Braktee. Für Distichia gibt VIERHAPPER (1930) 1-2 Brakteen an, BALSLEV (1996) spricht von 2-4 Brakteen. In dem von uns unter- suchten Material fanden sich stets zwei Brakteen. Das Gesamtbild deutet auch in diesen drei Gattungen wieder darauf hin, dass es sich bei den Infloreszenzen um einblütige Ährchen handelt. Die Infloreszenzen aller drei Gattungen sind jeweils als Paracladium mit einer einblütigen Cofloreszenz aufzufassen. Die gesamte Synfloreszenz ist auf ein einziges, ein- blütiges Paracladium reduziert, die Hauptachse proliferiert (Abb. 1 1e, g). 145 4. Diskussion 4.1. Das einblütige Ährchen Entscheidend für das Verständnis der Juncaceen-Infloreszenzen ist die Deutung der von zwei Brakteen begleiteten Einzelblüte. Die vergleichende Analyse hat uns zu der in sich wider- spruchsfreien Auffassung geführt, dass diese Blüte, zusammen mit den zwei Brakteen, ein einblütiges Ährchen repräsentiert. Obwohl der entwicklungsgeschichtliche Nachweis für die axilläre Anlegung der pseudoterminalen Blüte noch aussteht, sprechen die morphologischen Beobachtungen deutlich für diese Interpretation. Ein erstes Argument gegen die Terminalität der Einzelblüte liefern Ährchen von Luzula Sektion Gymnodes, bei denen die obere Braktee ausgefallen ist. Der Anschluss der Blattspira- le des äußeren Perigonkreises einer Terminalblüte müsste sich stets nach dem zuletzt unter der Blüte angelegten Blatt richten. Tatsächlich zeigt das Perigon aber eine festliegende Po- sition zur unteren Braktee, egal, ob die obere angelegt wird oder nicht. Abb. 12a zeigt dia- grammatisch den Normalfall mit zwei Brakteen. Es fällt auf, dass die obere Braktee etwas aus der Mediane verschoben ist. Von den Organen des äußeren Perigonkreises liegt das innerste direkt vor der unteren Braktee. Das ist die gleiche Position wie bei den Achselblüten von Juncus subgen. Juncus. Wenn die obere Braktee ausfällt, ändert sich an dieser Stellung nichts (Abb. 12b). Die Blüte ist mit der unteren Braktee in genau der Weise korreliert, wie es für eine Achselblüte zu fordern ist. In diesem Zusammenhang ist noch einmal an den in Abb. 2c dargestellten Fall zu erinnern, bei dem ein Ährchen von Juncus maritimus (Juncus subgen. Juncus ) bis auf eine Blüte ver- armt ist. Es kann kein Zweifel daran bestehen, dass es sich hierbei um ein einblütiges Ähr- chen handelt. Die einzige verbliebene Blüte ist in eine pseudoterminale Position gedrängt. Aus dem Vergleich mit den zunehmend verarmenden Ährchen des Gesamtblütenstandes ist kein anderer Schluss zu ziehen, als dass die Blüte der Achsel der unteren Braktee entspringt, und dass ihr aufwärts, als letztes Organ der terminal offenen Achse, eine sterile Braktee folgt. Dass die Einzelblüten der übrigen Juncaceen einschließlich der Arten von Juncus subgen. Poiophylli tatsächlich einem Ährchen homolog sind, wird durch einen weiteren Befund gestützt: Bei der Analyse eines Blütenstandes von Juncus conglomeratus fand sich ein Ähr- chen, das statt der üblichen Einzelblüte zwei Blüten entwickelt hatte (Abb. 7, 12c). Auf die zweite (distale) Blüte folgte auch hier wieder eine sterile Braktee. Dieses Ährchen unter- scheidet sich nicht von einem auf zwei Blüten verarmten Ährchen des Juncus maritimus (vgl. Abb. 12c und d). 4.2. Zur Typologie und Phylogenie der Juncaceen-Synfloreszenz Wenn die von zwei Brakteen umfassten Einzelbliiten der Juncaceen je einem Ahrchen ent- sprechen, gehören alle Blütenstände der Familie dem Typus der polytelen Synfloreszenz an. Die Floreszenzen sind Ährchen bzw. Köpfchen, die bis zur Einblütigkeit reduziert sein kön- nen. Die Anordnung der ein- oder mehrblütigen Floreszenzen im Gesamtverzweigungssystem erfolgt analog der Anordnung von terminalen Einzelblüten in monotelen Systemen. Dadurch ergibt sich in den reicher verzweigten Blütenständen eine täuschende Ähnlichkeit zu be- stimmten monotelen Synfloreszenzen. Der an eine Kegelrispe erinnernde Blütenstand von Prionium hat tatsächlich ein der Rispe (Panicula) entsprechendes Achsensystem, trägt aber am Ende jeder Achse statt der für die Rispe zu fordernden Endblüte eine offene Floreszenz. Bei den reicher verzweigten Juncus-Arten sowie bei Luzula sect. Anthelaea und sect. Ptero- des sind die Hauptachse und die Achsen der großen Paracladien so stark gestaucht, dass sie von ihren jeweiligen Seitenachsen deutlich übergipfelt werden. Auf diese Weise entsteht ein Achsensystem, das mit dem einer Trichterrispe bzw. Spirre vollkommen identisch ist. Wiede- rum unterscheidet sich der Blütenstand aber von der Trichterrispe (Anthela) dadurch, dass er am Ende jeder Achse statt der Terminalblüte eine offene Floreszenz trägt. TROLL (1968, S. 94) hat für diese polytelen Blütenstände, die ein Analogon zu entsprechenden Monotelen 146 darstellen, die Begriffe Paniculodium bzw. Anthelodium vorgeschlagen. Mit diesen Begriffen sind die Blütenstände von Prionium bzw. von Juncus und Luzula treffend zu bezeichnen. Erst auf der Basis des Anthelodiums ist auch ein spezieller Paracladientyp, das Drepanium (die sogenannte Sichel), zu verstehen. Es handelt sich hierbei um äußerst spezialisierte, sym- podial verzweigte Paracladien mit einer größeren Anzahl wiederholter Übergipfelungen. Voraussetzung für die Entstehung eines Drepaniums ist die fortlaufende Bildung besonders gestalteter, stets gleichbleibender Sympodialglieder. Jedes dieser Glieder muss ein adossiertes Vorblatt, ein mit einem Divergenzwinkel von 180° folgendes weiteres Blatt und das distale Ährchen besitzen. Das zwischen dem Vorblatt und dem Ährchen stehende Blatt fällt somit immer abaxial in die Medianebene und steht genau über dem Tragblatt des jeweiligen Sym- podialgliedes. Nur aus der Achsel des zwischen dem Vorblatt und dem Ährchen stehenden Blattes erfolgt die weitere Verzweigung (s. z.B. Pc2 in Abb. 8). Diese fixierte Blatt- und Ver- zweigungsfolge konnte in unserem Material bisher nur bei Vertretern mit einblütigen Ähr- chen, und zwar in den Sektionen Tenageia und Steirochloa gefunden werden (vgl. Abb. 8 und 9a). Das Drepanium kennzeichnet keinesfalls generell die Gattung Juncus und findet sich offenbar überhaupt nicht in der Gattung Luzula. Überlegungen zu den Beziehungen zwischen den Blütenstandstypen der Juncaceae s.l. sind schematisch in Abb. 13 dargestellt. Am ursprünglichsten dürften die Infloreszenzen von Prio- nium mit mehrblütigen Ährchen, reicherer Verzweigung und gestreckten Internodien sein (Abb. 13a). Den prinzipiell gleichen Bau haben auch noch die Infloreszenzen von Juncus subgen. Juncus, nur wird das äußere Erscheinungsbild durch die Achsenstauchung und die sich daraus ergebenden Übergipfelungen modifiziert. Von hier aus sind mehrere Wege der Verarmung und/oder der Spezialisierung zu erkennen. Durch Wegfall der Paracladien resul- tiert eine auf die Hauptfloreszenz reduzierte Synfloreszenz, wie sie etwa von J. triglumis repräsentiert wird (Abb. 13d). Bei Juncus subgen. Poiophylli und allen übrigen Gattungen sind die Floreszenzen einblütig geworden. Mehr oder weniger, zum Teil aber sehr reich verzweigt sind die Infloreszenzen von Juncus subgen. Poiophylli sowie von Luzula sect. Anthelaea und Pterodes (Abb. 13b). Bei Luzula sect. Gymnodes sind die blühenden Seitenäste in distale Kurzparacladien und proximale Langparacladien differenziert (Abb. 13c). Dazu kommt eine deutliche Tendenz zur Verrumpfung (Trunkation). Dieses Muster findet sich in vielen Familien mit polytelem Synfloreszenzbau (vgl. WEBERLING 1981). Durch Verarmung bis zur Einblütigkeit der gesamten Synfloreszenz sind die in kühlen Klimaten der Südhemi- sphäre beheimateten Gattungen gekennzeichnet. Marsippospermum und Rostkovia besitzen nur noch die einblütige Hauptfloreszenz, während die Paracladienbildung völlig unterdrückt wird (Abb. 13e). Durch das Merkmal der Prolifikation sind die Gattungen Distichia, Oxychloe und Patosia gekennzeichnet. Die Blütenbildung ist dadurch nur noch an Paracladien möglich. Jeder blühende Spross dieser Gattungen besitzt nur noch ein einziges, einblütiges Kurzpara- cladium (Abb. 13f). Mit diesen Vorstellungen werden die Annahmen von BUCHENAU (1890) vom Kopf auf die Füße gestellt (vgl. Abb. la, b). Bei der Gegenüberstellung von Juncus subgen. Poiophylli (Abb. 1a) und Juncus subgen. Juncus (Abb. 1b) muß der Pfeil von rechts nach links weisen. Außerdem ist nicht das einblütige Paracladium im Subgenus Poiophylli einer axillären Einzelblüte des Ährchens von Subgenus Juncus homolog. Vielmehr entspricht das dreiblütige Ährchen in Abb. 1b dem distalen, einblütigen Ährchen in Abb. la. Um ganz genau zu sein: Die Blüte des terminalen Ährchens in Abb. 1a entspricht der untersten Blüte des Ährchens in Abb. 1b. 4.3. Vergleich mit dem Infloreszenzbau der Cyperaceae Als sehr eng verwandt mit den Juncaceae s.l. gelten die Cyperaceae und die Thurniaceae. Zu diesem Ergebnis führen sowohl intensive Analysen aller klassischen Merkmale (DAHLGREN & RASMUSSEN 1983, DAHLGREN et al. 1985, SIMPSON 1995) als auch jüngste molekulare 147 Analysen (DUVALL et al. 1993, LINDER & KELLOGG 1995, CHASE et al. 2000, MUASYA et al. 2000). Eine Analyse der kugelförmig-kompakten Blütenstände von Thurnia steht noch aus. Infloreszenzen der Cyperaceae sind dagegen schon häufig untersucht worden. Aus neuerer Zeit sind die Arbeiten von BROWNING & GORDON-GRAY (1999), EITEN (1976), KUKKONEN (1984, 1986, 1994), TIMONEN (1998), VEGETTI (1992, 1994) sowie VEGETTI & TIVANO (1991) zu nennen. Nachdem vielfach Ährchen mit echten Terminalblüten beschrieben worden sind (z.B. KOYAMA 1971), ist inzwischen für die meisten derartigen Fälle Pseudoterminalität zumindest sehr wahrscheinlich gemacht worden. Für die ganze Familie darf wohl polyteler Bau der Infloreszenzen angenommen werden. Es bleiben nur ganz wenige unklare Fälle, wie z. B. Scirpodendron und die sehr aberrante Gattung Chrysitrix, die eine scheinbar terminale Blüte besitzen, bei denen aber aus vergleichenden Gründen ebenfalls Pseudoterminalität anzunehmen ist (GOETGHEBEUR 1998). Die elementare Baueinheit der Synfloreszenz der Cyperaceen ist das Ährchen. Wie bei den Juncaceen ist es im ursprünglichen Fall mehrblütig. Es trägt manchmal basal, häufig aber distal sterile Spelzen. Weit verbreitet ist die Tendenz zur Reduktion auf die Einblütigkeit (EITEN 1976, GOETGHEBEUR 1998). Besonders in diesen Fällen nimmt die einzig verbleiben- de Blüte eine pseudoterminale Position ein, distal der einzigen fertilen Braktee steht auch hier noch eine sterile Braktee (EITEN 1976, vgl. Abb. 12e, f). Ähnlich wie bei Luzula sect. Gym- nodes treten die einbliitigen Ahrchen oft zu Ahrchen 2. Ordnung zusammen (z.B. die weib- lichen „Ähren“ von Carex). Eine weitere Parallele zu den Juncaceae besteht in der Organisa- tion des Gesamtbliitenstandes. Er ist im urspriinglichen Fall nach dem Muster einer elongaten Rispe gebaut und stellt ein Paniculodium dar (z. B. Rhynchospora, s. RAYNAL 1971). Mehr- fach ist daraus das polytele Analogon der Trichterrispe hervorgegangen. Die Infloreszenzen von Scirpus, Cyperus oder Bolboschoenus liefern dafür anschauliche Beispiele. Auf diese Infloreszenzen wenden z.B. RAYNAL (1971) und GOETGHEBEUR (1998) noch den Terminus „anthele“ bzw. „anthela“ an, während KUKKONEN (1994) und BROWNING & GORDON-GRAY (1999) sie zutreffend als Anthelodium bezeichnen. Diese kurze Übersicht macht deutlich, dass die Infloreszenzen der Juncaceae und der Cyperaceae in vielen Merkmalen übereinstimmen. Die nahe Verwandtschaft beider Familien kann jetzt auch mit dem Merkmalsbereich der Blütenstände gut belegt werden. Literatur BALSLEV, H. 1996: Juncaceae. — Flora Neotropica 68. New York. — 1998: Juncaceae. — In: KUBITZKI, K. (ed.): The Families and Genera of Vascular Plants. Vol. IV: Flowering Plants. Monocotyledons: Alismatanae and Commelinanae (except Gramineae), 252-260. Berlin. BROWNING, J. & GORDON-GRAY, K.D. 1999: The inflorescence in southern African species of Bolboschoenus (Cyperaceae). — Ann. Bot. Fenn. 36: 81—97. BUCHENAU, F. 1865: Der Blüthenstand der Juncaceen. Jahrb. Wiss. Bot. 4: 385-435. — 1890: Monographia Juncacearum. Leipzig. CHASE, M.W. et al. 2000: Higher-level systematics of the monocotyledons: an assessment of current knowledge and a new classification. — In: WILSON, K.L., MORRISON, A. (eds.): Monocots. Systematics and Evolution, 3-16. Melboune. DAHLGREN, R., CLIFFORD, H.T. & YEO, P.F. 1985: The families of the Monocotyledons. Berlin. — & RASMUSSEN, F.N. 1983: Monocotyledon evolution: Characters and phylogenetic estimation. — In: HECHT, M.K., WALLACE, B. & PRANCE, G.T. (eds.): Evolutionary Biology 16: 255-395. New York. DUVALL. MLR. et al. 1993: Phylogenetic hypothesis for the monocotyledons constructed from rbcL sequence data. — Ann. Missouri Bot. Gard. 80: 607-619. 148 EITEN, L.T. 1976: Inflorescence units in the Cyperaceae. — Ann. Missouri Bot. Gard. 63: 81-112. GOETGHEBEUR, P. 1998: Cyperaceae. — In: KUBITZKI, K. (ed.): The Families and Genera of Vascular Plants. Vol. IV: Flowering Plants. Monocotyledons: Alismatanae and Commeli- nanae (exept Gramineae), 141-190. Berlin. KIRSCHNER, J., NOVARA, L.J., NOVIKOV, V.S., SNOGERUP, S. & KAPLAN, Z. 1999: Supra- specific division of the genus Juncus (Juncaceae). — Folia Geobot. Phytotax. 34: 377-390. KOYAMA, T. 1971: Systematic interrelationships among Sclerieae, Lagenocarpeae and Ma- panieae (Cyperaceae). — Mitt. Bot. Staatssamml. München 10: 604-617. KUKKONEN, I. 1984: On the inflorescence structure in the family Cyperaceae. — Ann. Bot. Fenn. 21: 257-264. — 1986: Special features of the inflorescence in the family Cyperaceae. — Ann. Bot. Fenn. 23: 107-120. — 1994: Definition of descriptive terms for the Cyperaceae. — Ann. Bot. Fenn. 31: 37-43. LAMPE, M. v. 1996: Wuchsform, Wuchsrhythmus und Verbreitung der Arten der Zwergbin- sengesellschaften. Dissertationes Botanicae 266. LINDER, H.P. & KELLOGG, E.A. 1995: Phylogenetic patterns in the commelinid clade. — In: RUDALL, P.J., CRIBB, P.J., CUTLER, D.F. & HUMPHRIES, C.J. (eds.): Monocotyledons. Systematics and Evolution, 473-496. Kew. MUASYA, A.M., SIMPSON, D.A., CHASE, M.W. & CULHAM, A. 2000: Phylogenetic relation- ships within the heterogeneous Scirpus s. lat. (Cyperaceae) inferred from rbcL and trnL-F sequences data. — In: WILSON, K. L. & Morrison, D. A (eds.): Monocots. Systematics and Evolution, 610-614. Melbourne. MÜLLER-DOBLIES, D., STÜTZEL, T. & WEBERLING, F. 1992: A drepanium is not a cyme. — Flora 187: 61-65. MUNRO, S.L. & LINDER, H.P. 1998: The phylogenetic position of Prionium (Juncaceae) within the order Juncales based on morphological and rbcL sequence data. Systematic Botany 23: 43-55. NOVARA, L.J. 1976: Contribucion al conocimiento de las inflorescencias de Juncus y su significacion taxonomica. — Kurtziana 9: 41-61. RAYNAL, J. 1971: Quelques notes morphologiques sur les Cyperacées. — Mitt. Bot. Staats- samml. München 10: 589-603. SIMPSON, D. 1995: Relationships within Cyperales. — In: RUDALL, P.J., CRIBB, P.J., CUTLER, D.F. & HUMPHRIES, C.J. (eds.): Monocotyledons. Systematics and Evolution, 497-509. Kew. SNOGERUP, S. 1993: A revision of Juncus subgen. Juncus (Juncaceae). — Willdenowia 23: 23-73. TIMONEN, T. 1998: Inflorescence structure in the sedge tribe Cariceae (Cyperaceae). — Publications in Botany from the University of Helsinki 26: 5-35. TROLL, W. 1964a: Bericht der Kommission fiir Biologische Forschung. Botanischer Teil. — Akad. Wiss. Jahrb. 1963: 119-129. — 1964b: Die Infloreszenzen. Bd. 1. Jena. — 1968: Bericht der Kommission für Biologische Forschung. Botanischer Teil. — Akad. Wiss. Jahrb. 1967: 92-103. — 1969: Bericht der Kommission für Biologische Forschung. Botanischer Teil. — Akad. Wiss. Jahrb. 1968: 95-106. VEGETTI, A.C. 1992: Typology of the inflorescence in species of Schoenoplectus (Cypera- ceae) of Austral America. — Beitr. Biol. Pflanzen 67: 241-249. — 1994: Typology of the inflorescence in species of /solepis. — Beitr. Biol. Pflanzen 68: 21-26. 149 — & Tivano, J.C. 1991: Inflorescence typology in Schoenoplectus californicus. (Cypera- ceae). — Beitr. Biol. Pflanzen 66: 323-345. VIERHAPPER, F. 1930: Juncaceae. — In: ENGLER, A. & PRANTL, K. (eds.): Die Natiirlichen Pflanzenfamilien, 2. Aufl., 15a: 192-224. Leipzig. WEBERLING, F. 1981: Morphologie der Blüten und der Blütenstände. Stuttgart. Christian P. KOBELE & Prof. Dr. Hans-Jiirgen TILLICH, Institut fiir Systematische Botanik der Ludwig-Maximilians-Universität München, Menzinger Strasse 67, D-80638 München. Deutschland. Abb. 1: a, b: Hypothetischer Übergang des Infloreszenzbaues von Juncus subgen. Poiophylli (a) zum subgen. Juncus (b) nach BUCHENAU (1890). e: Juncus subgen. Poiophylli. Verzweigungsschema mit Angabe der im folgendenText verwendeten Symbole. B: sterile Braktee; Br: fertile Braktee; Cf*: Cofloreszenz eines Paracladiums aus der Vorblattachsel; Cf 1 bzw. 1.1: Cofloreszenz eines Paracladiums 1. bzw. 2. Ordnung; Ha: Hauptachse; Hf: Hauptfloreszenz; Pcl bzw. Pc 1.1: Para- cladium 1. bzw. 2. Ordnung; TB: Tragblatt; VB: Vorblatt. Fig. 1: a, b: Hypothetical transformation of the inflorescence type of Juncus subgenus Poiophylli (a) to that of subgenus Juncus (b) after BUCHENAU (1890). e: Juncus subgenus Poiophylli. Branching pattern with indication of symbols used in the following text. B: sterile bract; Br: fertile bract; Cf*: coflorescence of a paracladium originating from the axil of a prophyll; Cf 1. and 1.1: coflorescence of a paracladium of first and second order, respectively; Ha: main axis; Hf: main florescence; Pc 1 and 1.1: paracladium of first and second order, respectively; TB: pherophyll; VB: prophyll. 150 0 2,5 5 mm Blüte 1 Abb. 2: Juncus maritimus. Armblütige Ährchen. a: dreiblütig; b: zweiblütig; c: einblütig. Beachte die sterile Braktee (B), die jeweils distal der zuletzt gebildeten Blüte steht! Fig. 2: Juncus maritimus. Few-flowered spikelets. a: three-flowered; b: two-flowered; c: one-flowered. Note the sterile bract distal to the uppermost flower of each spikelet! 151 SAN: 5 ae Y dh SR an "in NV Y WE NR , Gs N N = oN) IA th (<= 17 N Ge „Nas g ASS y (/ SEZ Gruppe 1a. Gruppe 1b. Hauptgruppe 1 O Hauptgruppe 2 Q) ; 55 (DO Q) Yo LD, © WO LZ N D Ne TI 2 Z x = N 7 ( > > IZ Gruppe 2b. Gruppe 2c. Abb. 3: Schema der Grundtypen des Infloreszenzbaues der Juncaceae s.l. Oben: Infloreszenzen mit mehrbliitigen Ahrchen (Hauptgruppe 1); Unten: Infloreszenzen mit einblütigen Ahrchen (Hauptgruppe 2). Nähere Erläuterungen im Text. Fig. 3: Basic types of inflorescence structure in Juncaceae s.l. Above: Inflorescences with many-flowered spikelets (main group 1); Below: Inflorescences with one-flowered spikelets (main group 2). Further explanation in the text. 152 Abb. 4: Juncus maritimus. Aufriss einer Infloreszenz. Fig. 4: Juncus maritimus. Schematic elevation of an inflorescence. 153 Abb. 5: Infloreszenzen der Gruppe la. a-d: Juncus capitatus. a: Typische Infloreszenz; b: Aufriss von Fig. a; c: Infloreszenz mit Verzwei- gung aus der Achsel des Vorblattes; d: Aufriss von Fig. c. e, f: Prionium serratum. e: Teilinfloreszenz; f: Aufriss von F ig. e. Fig. 5: Inflorescences of the group la. a-d: Juncus capitatus. a: typical inflorescence; c: inflorescence with a branch from the axil of a prophyll; b, d: schematic elevations of figs. a and c. e, f: Prionium serratum. e: partial inflorescence; f: schematic elevation of fig. e. 154 Abb. 6: a-c: Infloreszenzen von Juncus sectiones Ozophyllum und Stygiopsis. a, b: schematischer Auf- riss der Infloreszenzen, a: J. articulatus; b: J. castaneus; c: J. triglumis, eine vollständige Inflores- zenz, auf der Rückseite der dritten Blüte befindet sich eine sterile Braktee; d: J. inflexus (sect. Juncotypus), Paracladium mit Verzweigung aus der Vorblattachsel. Dicht unterhalb jedes Ährchens steht ein Tragblatt (Tb°) mit einer rudimentären Blütenanlage. N: Resektionsfläche des Tragblatts von Pc 1. Fig. 6: a-c: Inflorescences of Juncus sectiones Ozophyllum and Stygiopsis. a, b: schematic elevation of inflorescences, a: J. articulatus; b: J. castaneus; c: J. triglumis, a complete inflorescence, on the back-side of the third flower there is a sterile bract; d: J. inflexus (sect. Juncotypus), paracladium with a branch from the axil of the prophyll. Closely below each spikelet there is a bract (Tb°) with a rudimentary flower bud in its axil. N: scar of the dissected pherophyll of Pc 1. ON Q Ss QQ) A N OND 9\Q \ \ \ Q Ss : SA) Y) OQ SS NV S)\« AN) AN N Q N OQ) SI N I = o\ TN IE © > a I) oO Q Q) >I NSS Q) 2 Ss N Q \ O An Qe A) _V > SV) Qy WY’ Y =, Y)s as Q) DIR I. N \) Sa Q JO N Q } Ov Dye Ny © Ö) ASS UNS Q N 3 er N on N oa I Yay N Q ANAND PO 5 Nv) Q Q 0 EN y) WY) 2 SAS), 9% > ZN \' Nv SAY | a —I OQ =y y) Q IS N O S % Say N Abb. 7: Juncus conglomeratus. Aufriss einer In- floreszenz; TB: das pseudoterminale, frondose Tragblatt, welches die gesamte Infloreszenz in eine pseudolaterale Position abdrängt. Der Pfeil markiert ein anomales, zweiblütiges Ährchen. Fig. 7: Juncus conglomeratus. Schematic elevation of an inflorescence; TB: the pseudoterminal leaf, which shifts the complete inflorescence into a pseudolateral position. The arrow marks an abnormal, two-flowered spikelet. NY u \g — Sg ip SSeS N¢ NS = N= ©) Zi NS III ~ Ne TB OW Zen > I. = = AN SOL Wan LO N 156 Peta Abb. 8: Juncus bufonius. a: distaler Bereich einer Infloreszenz; b: das dazugehörige Aufrissdia- gramm. Fig. 8: Juncus bufonius. a: distal region of an inflorescence; b: schematic elevation of the same object. Abb. 9: Aufrisse von Infloreszenzen der Gruppe 2b. a: Juncus tenuis: b: Luzu- la glabrata. Fig. 9: Schematic elevations of inflorescences representing group 2b. a: Juncus tenuis: b: Lu- zula glabrata. 158 Q N y \ (CD 2 N (Ye 3 O = Y ——T i CO WS \\ Y/ NN yy Vy AQ N TG Q WV) » N b C) (CO L> E229 A GFZ F224 Abb. 10: Infloreszenzbau bei Luzula. a, b: sect. Pterodes. a: L. pilosa, Detail der Verzweigung; b: L. forsteri, Aufriss einer Infloreszenz. c, d: sect. Gymnodes. c: L. campestris, schematischer Aufriss einer Infloreszenz; d: Pc2 im Detail. Fig. 10: Inflorescence structure in Luzula. a, b: sect. Pterodes. a: L. pilosa, detail of branching; b: L. forsteri, schematic elevation of an inflorescence. c, d: sect. Gymnodes. c: L. campestris, schematic elevation of an inflorescence, d: Pc2 in detail. 159 Abb 11: Infloreszenzbau südhemisphärischer Juncaceae. a, b: Rostkovia magellanica; c: Marsippo- spermum grandiflorum; d, e: Oxychloe andina; f, g: Distichia muscoides. Nähere Erläuterung im Text. Fig. 11: Inflorescence structure in south hemispherical Juncaceae. a, b: Rostkovia magellanica; c: Marsippospermum gran - diflorum; d, e: Oxychloe andina; f, g: Distichia muscoides. Further explanation in the text. 160 VB 4 ‘ i\ \ 1 i} i} N / Gynoeceum Br TB ee ee neo ert 5mm 2mm 2,5 A Abb. 12: Ahrchenbau bei Juncaceae und Cyperaceae. a, b: Luzula campestris, Ährchen-Diagramme. a: Ahrchen mit zwei Brakteen; b: obere Braktee ausgefallen. c, d: armblütige Ahrchen von Juncus. c: ausnahmsweise zweibliitiges Ahrchen von J. conglomeratus. Die sterile obere Braktee befindet sich auf der vom Betrachter abgewandten Seite. d: Auf zwei Bliiten verarmtes Ahrchen von J. ma- ritimus. e, f: Ährchen von Cyperaceen, schematisch. e: Eleocharis; f: Cyperus (nach EITEN 1976, S. 83, geändert). Fig. 12: Spikelets in Juncaceae and Cyperaceae. a, b: Luzula campestris, diagrams of spikelets. a: spikelet with two bracts; b: the upper bract is missing; c, d: few-flowered spikelets in Juncus. c: unusually two-flowered spikelet of J. con- glomeratus, the sterile bract is at the back of the spikelet; d: impoverished, two-flowered spikelet of J. maritimus. e, f: spikelets of Cyperaceae, schematically. e: Eleocharis; f: Cyperus (after EITEN 1976, p. 83, modified). 161 Oe = SON 3. = N X )) Wy, =, —- N X )) =V, © ZEN) 2 X =) - jae r BR EI Ey, = L =) ven TEN k= In) O N SY) RRA SU KYU — Z = => > = 35 > > > = Oo ---->> = — <= © — == = S N = x ro O Ka O [57 = De] 7 A 1 A 1 1 1 1 1 1 1 1 [a7 ' ve : WO _ OVS 1 OO \A i U IC) - OF JO) - © OF v Bm Ss ——E——EEe = En N) TE @s N OSINO > VW © IR Op) O Abb. 13: Vermutete Beziehungen zwischen den Infloreszenz-Typen der Juncaceae. Nähere Erläu- terung im Text. Fig. 13: Supposed relationships between inflorescence types of Juncaceae. Further explanation in the text. 4 ) | 163 Contributions to the knowledge of the genus Astragalus L. (Leguminosae) VII-X' D. PODLECH Abstract: PODLECH, D.: Contributions to the knowledge of the genus Astragalus L. (Legumi- nosae) VII-X. — Sendtnera 7: 163-201. 2001. ISSN 0944-0178. VI. A survey of Astragalus L. sect. Leucocercis. The section, endemic in Iran with six species, is revised. Synonymy, descriptions, the investigated specimens and a key for the species are given. VIII. New typifications and changes of typification in Astragalus-species. 12 wrongly typified species are re-typified. 19 taxa are typified here. IX. Some new species in genus Astragalus: 27 new species, one subspecies and one section are described here. They belong to the following sections: Sect. Caprini: A. behbehanensis, A. bozakmanii, A. spitzenbergeri. Sect. Cenantrum: A. tecti-mundi subsp. orientalis. Sect. Chlorostachys: A. poluninii, A. rhododendrophila. Sect. Cystium: A. owirensis. Sect. Dissitiflori: A. argentocalyx, A. bingoellensis, A. doabensis, A. fruticulosus, A. lanzhouensis, A. montis-karkasii, A. pravitzii, A. recurvatus, A. saadatabadensis, A. sata-kandaoensis, A. wakha- nicus. Sect. Hemiphaca: A. sherriffii, A. tsangpoensis. Sect. Incani: A. olurensis, A. zaraensis. Sect. Komaroviella: A. damxungensis. Sect. Onobrychoidei: A. ras- montii. Sect. Polycladus: A. austrotibetanus, A. cobresiiphila, A. conaensis. Sect. Pseudotapinodes, sect. nov.: A. dickorei. X. New names and combinations are given. Four illegitimate names are changed, two taxa have been raised in rank. Zusammenfassung: VII. A survey of Astragalus L. sect. Leucocercis. Eine Revision von Astragalus L. sect. Leucocercis wird vorgestellt. Die Sektion ist endemisch im Iran und umfasst sechs Arten. Ausführliche Synonymie, Beschreibung der Arten, Aufzählung der untersuchten Belege und ein Bestimmungsschlüssel werden gegeben. VIII. New typi- fications and changes of typification in Astragalus-species. Neue Typifizierungen und Änderungen früherer Typifizierungen werden vorgestellt. 12 Arten waren bis- her falsch typifiziert, 19 Taxa werden hier typifiziert. IX. Some new species in genus Astragalus: 27 neue Arten, eine neue Unterart und eine neue Sektion werden beschrieben. Sie gehören zu folgenden Sektionen: Sect. Caprini: A. behbehanensis, A. bozakmanii, A. spitzenbergeri. Sect. Cenantrum: A. tecti-mundi subsp. orientalis. Sect. Chlorostachys: A. poluninii, A. rhododendrophila. Sect. Cystium: A. owiren- sis. Sect. Dissitiflori: A. argentocalyx, A. bingoellensis, A. doabensis, A. fruticu- losus, A. lanzhouensis, A. montis-karkasii, A. pravitzii, A. recurvatus, A. saada- ' PoDLECH, D.: Beiträge zur Kenntnis der Gattung Astragalus I. Neue und bemerkenswerte Arten aus Afghanistan. Bot. Jahrb. Syst. 107: 55-73. 1985. — Beiträge zur Kenntnis der Gattung Astragalus L. (Leguminosae) II. Astragalus renzianus species nova aus dem Iran. Mitt. Bot. Staatss. München 22: 1-3. 1986. — Beiträge zur Kenntnis der Gattung Astragalus L. (Leguminosae) III-VI. Sendtnera 1: 267-272. 1993. 164 tabadensis, A. sata-kandaoensis, A. wakhanicus. Sect. Hemiphaca: A. sherriffii, A. tsangpoensis. Sect. Incani: A. olurensis, A. zaraensis. Sect. Komaroviella: A. damxungensis. Sect. Onobrychoidei: A. rasmontii. Sect. Polycladus: A. austro- tibetanus, A. cobresiiphila, A. conaensis. Sect. Pseudotapinodes, sect. nov.: A. dickorei. X. New names and combinations are given. Vier illegitime Namen wer- den geändert, zwei Sippen werden im Rang verändert. VIL A survey of Astragalus L. sect. Leucocercis Bunge. The species of the small section Leucocercis are all endemic in Iran. They have all a characteristic habit and are all close together. It is a section with no closer allies within the genus. The section has been revised twice in recent years, but in both cases important characters especially in the indument structure have not been taken into consideration and several types have not been investigated. Therefore several changes of names and new circumscription of the species are offered here. Astragalus sect. Leucocercis Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 132. 1868. Lectotype (PODLECH 1990): A. mucronifolius Boiss. Plants low, mostly densely cushion-like spiny shrublets with short growth in one year. Hairs merely white, mostly appressed, symmetrically to strongly asymmetrically medifixed, in part even basifixed. Stipules adnate to the petiole, free from each other. Leaves paripinnate, with thickly leathery, acuminate-cuspidate leaves with prominent nerves. Rhachis rigid, pungent, persistent, of variable length. Racemes with a mostly short peduncle, loosely 3-15- flowered. Bracts short. Bracteoles present, in upper part of the pedicel or at the base of the calyx. Calyx campanulate to shortly tubular, hairy, with slightly to distinctly pungent teeth. Petals glabrous, pink (always ?), deciduous. Standard blade elliptic to suborbicular, slightly retuse at the apex. Wings and keel as long or only slightly shorter than the standard. Staminal tube truncate at the mouth. Legumes sessile, short, unilocular; valves coriaceous, hairy. Seeds 18. Key to the species Remark: The form of legumes is very characteristic in some species. If they are not available, it is necessary to compare carefully all the characters in the descriptions. l. Bracts densely hairy; calyx teeth 1.5-2 mm long, often recurved; standard 9-10 mm long; legumes 5—8 mm long, with only one seed A. curviflorus Boiss. _ Bracts only ciliate at the margins; calyx teeth longer, not recurved; standard distinctly longer; legumes distinctly longer, with several seeds 2 2. Stems of the current year nearly glabrous; rhachis of the leaves covered with strongly asymmetrically medifixed hairs up to 1.5 mm long; legumes ovoid, 9-15 mm long, 7-11 mm high and wide, rounded ventrally and dorsally; leaflets with a cusp 1.5-3 mm long; peduncles 2-8 cm long A. ovoideus Sirj. & Rech.f. — Stems of the current year densely hairy (hairs concealed by stipules); rhachis of the leaves covered with shorter and at least partly + symmetrically medifixed hairs; 165 peduncles short, at most up to 2.5 cm long, rarely up to 5 cm long (in A. mucronifolius); legumes elliptic or obliquely elliptic, carinate ventrally, if rounded (in A. kentrophyllus) than leaflets shortly acuminate with a cusp up to 0.5(-1) mm long 3 3. Calyx covered with basifixed to strongly asymmetrically medifixed hairs; standard blade elliptic to widely elliptic, gradually narrowed at the base into the claw 4 = Calyx covered with + symmetrically medifixed hairs; standard blade suborbicular, subabruptly narrowed at the base into the claw 5 4. Stems of the current year with hairs protruding from the stipules; leaflets long acuminate-aristate, with a cusp up to 6 mm long; legumes with up to 8 seeds A. fischeri Fisch. — Stems of the current year with hairs nearly completely concealed by the stipules; leaflets shortly acuminate-aristate with a cusp up to 3 mm long; legumes with 2-3 seeds A. mucronifolius Boiss. 5. Leaflets in 4-5 pairs, elliptic to obovate, 2-3 mm wide; legumes slightly compressed from dorsal side, ca. 6 mm high and 8 mm wide, rounded ventrally and dorsally A. kentrophyllus Podlech — _ Leaflets in 6-9 pairs; narrowly elliptic, 1-2 mm wide; legumes strongly compressed laterally, 6-6.5 mm high and 3 mm wide, subcarinate ventrally and dorsally A. talimansurensis Sir}. & Rech.f. Astragalus curviflorus Boiss., Diagn. pl. orient., ser. 1, 2: 67. 1843 = Tragacantha curviflora (Boiss.) Kuntze, Revis. Gen. 2: 944. 1891. Syntypes: Persia australis, P.M.R. Aucher-Eloy 4388 (G!, G-BOIS!, K!, OXF!, W!); Persia australis, inter Jaroun [Jahrom] et Darab, P.M.R. Aucher-Eloy 4391. Lectotype (PODLECH 1998): P.M.R. Aucher-Eloy 4391 (G-BOIS!; iso: G!, K!, P!: foto MSB!). Plants horribly spiny, densely cushion-like shrublets 12-5(-30) cm tall. Main stem up to 6 cm long and up to 5 mm thick, branched, densely covered with remnants of old spines, densely covered with appressed to subappressed short white hairs. Stipules yellowish, 5-6 mm long, from wide base subabruptly narrowly triangular-acuminate, acute but not pungent, adnate to the petiole for 2-3 mm, densely covered at the base and at the margins with subappressed, extremely asymmetrically medifixed hairs, the top often glabrescent or glabrous. Leaves 2—4 cm long; petiole and rhachis stout, up to 1.5 mm thick, ngid, pungent, densely covered with appressed to subappressed, asymmetrically to extremely asym- metrically medifixed hairs up to 0.5 mm long and with much fewer, more ascending hairs up to 0.8 mm long. Leaflets in 4-6 pairs, thickish-leathery, elliptic to obovate, 4-5 x 1.5-2 mm, at the apex abruptly acutish or with a yellowish spine up to 1 mm long, densely covered on both sides with appressed, asymmetrically medifixed hairs up to 0.6 mm long. Peduncles 0.5-1.5 cm long, slender, hairy like the rhachis. Racemes loosely 2-5-flowered. Pedicels 1 mm long. Bracts narrowly triangular 1-2 mm long, densely hairy. Bracteoles ca. 0.5 mm long, in upper part of the pedicel or at the base of calyx. Calyx 5—6 mm long, tubular, attenuate at the base, densely hairy; teeth subulate, 1.5-2 mm long, often somewhat curved, not pungent. Petals pink? Standard 9-10 mm long; blade upcurved above the claw, widely elliptic to nearly circular, 5.5-6 mm wide, slightly retuse at the apex, gradually narrowed into the cuneate short 166 claw. Wings ca. 9 mm long; blades slightly curved, oblong, obtuse at the apex, 6 x 2 mm; auricle ca. 0.5 mm long, claw curved, ca. 3.5 mm long. Keel 9 mm long; blades elliptic- triangular, with widely curved lower edge and nearly straight upper edge, long acuminate at the apex, 5 x 2.5 mm; auricle short, claw 4.5 mm long. Ovary sessile, pilose, with 4 ovules. Pods sessile, widely elliptic seen from the side, laterally strongly compressed, 5-8 mm long, 4-5 mm high and 2-3.5 mm wide, carinate ventrally, rounded dorsally but with prominent midnerve, at the apex minutely acuminate, unilocular; valves pale brownish, thin but tough, loosely covered with medifixed to subbasifixed, tangled, subappressed hairs 0.1-1 mm long, soon glabrescent. Seed only one. Distribution: in southwest Iran (Prov. Fars). Specimens seen: Iran. Fars: 2 km after Sivand on the road to Abadeh, 1350-1400 m, 14.7.1994, Chehregani & Zarre 17904 (MSB); circa Kaserun, Stapf 1071 (WU) — Kuh-i-Bamu, NE of Shiraz, 1830 m, 28.7. 1966, Archibald 2939 (K); 32 km W Shiraz, 1820 m, Zohary 477 (HUJ) — prope urbem Shiras, 1842, Kotschy 58a (W) — dto.bei Schiras, 16.6.1885, Stapf (K, WU) - dto. 23.8.1985, Stapf (K) — dto., 27.8.1960, Zohary 453 (HUJ) — Persepolis, 12.7.1985, Stapf (K) — Hadchiabad [Hajiabad], 12.7.1885, Stapf 1070 (WU) — near Fasa, ca. 150 km SE Shiraz, 7.6.1965, Ledingham & Assefi 4132 (W) - inter Jaroun [Jahrom] et Darab, Aucher-Eloy 4391 (G, G-BOIS, K, P) — Lake Maharlu, 35 km SE of Shiraz, 1500 m, 18.11.1963, Grant 15082 (W) — Takht-e Jamshid, above the ruins, 22.5.1959, Wendelbo 786 (BG) — in montibus 12 km NW Hoseynabad, 37 km ab opp. Darab, 30.5.1973, Sojak 5043, 5047 (PR) — Kotal Komared, 4.5.1885, Stapf 685 (K). Astragalus fischeri Fisch., Bull. Soc. Imp. Naturalistes Moscou 26(2): 429. 1853. Type: [bei Gäsnabad, 12.5.1849], F.A. Buhse 1413 (H!). = A. phyllokentrus Hausskn. & Bornm., Mitth. Thiiring. Bot. Vereins, n.s. 23: 11. 1908. Syntypes: Persia occ., in monte Latetar, 10.6.1895, Th. Strauss (B!: foto K!, JE!); et ad Gulpaigan, vi.1899, Th. Strauss; Tschchar-Khatun, ditionis montis Raswend, vi.1902, Th. Strauss (B!, G!, K!, W!). Lectotype (PODLECH, designated here): In dit. urbis Sultanabad, Gulpaigan, vi.1899, Th. Strauss (JE!; iso: B!, K!, W!, Z!). = A. cornu-caprae Sirj. & Rech.f., Anz. Österr. Akad. Wiss., Math.-Naturwiss. Kl. 1954 (91): 171. 1954. Holotype: [Iran] entre Gomsche et Yezd-i Khast, route d'Ispahan a Chiraz, a 200 km SE de Ispahan, 1800 m, 11.6.1930, J. Gautier 14 (P!: foto MSB!). Plants spiny, cushion-like shrublets up to 30 cm tall, with only white hairs. Caudex up to 10 mm in diameter, with short, dark brown branches, in upper part covered with remnants of old leaves. Stems of the year 1-2 cm long, densely covered with ascending, fine, subbasifixed white hairs up to 2.5 mm long, protruding well from the upper stipules. Stipules membraneous, 8-12 mm long, triangular-acuminate, not spiny, distinctly reticulately nerved, adnate to the petiole for 2-4 mm, sometimes only densely ciliate at the margins with basifixed hairs up to 1 mm long, otherwise glabrous or hairy also on the outer surface. Leaves 4-25 cm long, in the lower 2.5-6 cm without leaflets; petiole and rhachis up to 2.5 mm thick, rigid, pungent, sparsely to rather densely covered with appressed short, + medifixed hairs 0.2-0.3 mm long and with asymmetrically medifixed to subbasifixed hairs up to 1 mm long, often soon glabrescent or glabrous. Leaflets in 3-6 pairs, thickish-leathery, elliptic to obovate, at the apex abruptly acuminate into a spine 1.5-6 mm long, 8-15 x 5-9 mm, prominently nerved, on 167 upperside densely, on underside sparsely to loosely covered with appressed + medifixed hairs 0.4-0.8 mm long. Peduncles 1-2 cm long, sparsely hairy or glabrous. Racemes loosely 2-15- flowered, later on often elongated and up to 9 cm long. Bracts membraneous, narrowly triangular to linear, 1-3 mm long, ciliate. Pedicels 1 mm long. Bracteoles ca. | mm long, in upper part ofthe pedicel or at the base of calyx. Calyx campanulate, 6-7 mm long, glabrous or very sparsely hairy; teeth narrowly triangular to subulate, 2-4 mm long, acuminate-pungent. Petals pink. Standard 12-14 mm long; blade upcurved above the claw, widely elliptic, 8 mm wide, slightly retuse at the apex, gradually narrowed into the cuneate, short claw. Wings 10-14 mm long, slightly curved; blades oblong, rounded at the apex, 8.5 x 3 mm; auricle ca. 0.5 mm long, claw curved, ca. 5.5 mm long. Keel 9-12 mm long; blades elliptic-triangular, with widely curved lower edge and straight upper edge, acute at the apex, 5.5-6 x 3.5 mm; auricle short, claw 6.5 mm long. Ovary sessile, hairy. Pods sessile, obliquely elliptic, straight to slightly curved, 10-20 mm long, 3.5—6.5 mm high and 3-4 mm wide, obtusely carinate by the thick median nerve ventrally, carinate or rounded dorsally but with prominent midnerve, at the apex shortly acuminate, unilocular; valves brownish, thick, with obliquely transverse- reticulate nerves, loosely covered with extremely asymmetrically medifixed to subbasifixed ascending hairs 0.8—1(—2) mm long, becoming glabrescent. Seeds up to 8, irregularly triangular to rectangular, 3.5 x 3 mm, brown, slightly pitted. Distribution: In southwest and central Iran. Specimens seen: Iran. Markasi: in ditione urb. Sultanabad [Arak], Gulpaigan, vi.1899, Strauss (B, JE, K, W, Z) — near Arak, 1520 m, 2.4.1963, Bowles sholarship Bot. Exp. 610 (K) - 60 km from Delijan to Qum, 1140 m, 23.5.1972, Amin & Mousavi 16119 (W) — in monte Tschchar Khatun (dit. Raswend), VI.1902, Strauss (W) — inter Teheran et Dilijan, 500-1000 m, 23.4.1956, Schmid 5158 (M) - 10 miles W Qom, 27.4.1961, Stutz 747 (W) - 15 km SQom, 1130 m,, 22.4.1961, Pabot 12436-E (W) & 5669 (G). — Esfahan: Julfa, Armenia cemetry, 29.5.1959, Wendelbo 909 (BG) — dto., 20.4.1948, Rechinger, Aellen & Esfandiari 2688 (K, M, W) — 60-70 km ab Ardestan boreo-occidentem versus inter Ab-i Azan (Abyazan) et Rahmatabad, 1150 m, 26.5.1974, Rechinger 46504 (M, W) — Iahabad, 20 km NW Najafabad, 45 km W Esfahan, 1900 m, 12.6.1965, Ledingham & Assefi 4140 (W) - 2 km N Najafabad, 12.6.1965, Ledingham & Assefi 4134 (W) — 14 km SE Natanz, 1480 m, 12.5.1961, Pabot 7175 (G, W) - road from Natanz to Ardestan, near Emamzade Abyazan, 1380 m, 15.5.1974, Wendelbo & Foroughi 11462 (W) — 24 km NE Murcheh Khvort, 1750 m, 27.5.1974, Rechinger 46638 (M, W) — 70 km NW Nain versus Ardestan, 1700 m, 18.5.1975, Rechinger 52022 (M, W) - Ardestan to Nain, 29.11.1956, Sabeti 12085-E (W) — Isfahan district, 1500-1800 m, Trott (K) — Kuh-i-Shir, 7 km SW Isfahan, 1650 m, 27.4.1953, Soder 216 (W) — Ispahan, Aucher-Eloy 4400 (K) — 10 km W Esfahan, 1600 m, 28.5.1960, Pabot 3465 (G) — 27 km S Esfahan, 1700 m, 24.4.1961, Pabot 5753 (G) - 65 km SW of Esfahan, 1400 m, 22.4.1929, Cowan & Darlington 1166 (K) — 10 km SE Nain, 1530 m, 11.5.1961, Pabot 7104 (G, W) - 70 km NE Nain, 1700 m, 18.5.1975, Iranshahr 41057-E (W) - 43 km NW Nain, 1970 m, 11.5.1961, Pabot 7112 (G). — Yazd: ad pagum Seinudin, 1600-1700 m, 11.4.1892, Bornmüller 3710 (K, W) — 54 km S Mahallat to Golpaiegan, 1750 m, 16.5.1973, Babakhanlu & Amin 15271(W). — Fars: entre Gomsche et Yezd-i Khast, route d'Ispahan a Chiraz, a 200 km SE de Ispahan, 1800 m, 11.6.1930, J. Gautier 14 (P) — zwischen Jezdikhast und Aminabad, 11.9.1885, Stapf 1008 (WU) — bei Jezdikhast, 11.9.1985, Stapf 1008 (WU) - 16 km W of Abarqu, 1600 m, 3.4.1964, Grant 15410 (W) — N of Abadeh, 2100 m, 22.4.1966, Archi- bald 1473 (K) — Abadeh, 1800 m, 12.5.1915, Pravitz 727 (W) - entre Abadeh et Daulatabad, 1500-2000 m, 26.4.1956, Schmid 5235 (G) - 30 km SE Abadeh, 1810 m, 26.4.1961, Pabot 5814 (G). 168 Astragalus kentrophyllus Podlech, Sendtnera 6: 156, 1999. Holotype: Iran, Prov. Khuzestan: 25 km N of Haftgel towards Izeh, 550 m, 25.5.1973, /ranshahr & Mousavi 15593-E (W!). Plants spiny, cushion-like shrublet up to 15 cm tall, in most parts densely covered with appressed, + symmetrically medifixed, white hairs. Caudex up to 10 mm in diameter, with short, dark brown branches, in upper part covered with remnants of old leaves. Stems of the year 1-3 cm long, the dense indumentum of fine, basifixed white hairs up to | mm long is not visible because of the very dense cover of leaves with their stipules. Stipules pale yellowish, 5-8 mm long, triangular-acuminate, shortly spiny, distinctly reticulately nerved, adnate to the petiole for 3-5 mm, sparsely ciliate at the margins with basifixed hairs up to 0.3 mm long, otherwise glabrous. Leaves 5-12 cm long, in the lower half to two third without leaflets; petiole and rhachis up to | mm thick, rigid, pungent, densely covered with appressed hairs 0.2-0.6 mm long. Leaflets in 4-5 pairs, thickish-leathery, elliptic to obovate, 4-7 x 2-3 mm, at the apex abruptly acuminate into a minute spine up to 0.5(—1) mm long, on upperside densely, on underside loosely to rather densely covered with appressed, symmetrically medifixed hairs 0.3-0.5 mm long. Peduncles 1-1.5 cm long, curved, hairy like the rhachis. Racemes loosely 3-flowered. Bracts whitish-hyaline, narrowly triangular, 2 mm long, ciliate. Pedicels at fruiting time thickish, 3-4 mm long, rather densely hairy. Bracteoles 0.5-1.5 mm long, linear, in the upper part of the pedicel. Calyx campanulate, ca. 7 mm long, obliquely cut at the mouth, rather densely covered with short, + symmetrically medifixed hairs, at the upper margins and teeth with basifixed cilia; teeth narrowly triangular, 3-3.5 mm long, acuminate but not pungent, with elevated midnerve. Standard 13 mm long; blade upcurved above the claw, suborbicular, 7 mm wide, emarginate at the apex, subabruptly narrowed into the wide claw. Wings 11-12 mm long, slightly curved; blades oblong-obovate, rounded at the apex, 6 x 3 mm; auricle ca. | mm long, claw curved, 5-6 mm long. Keel 11-12 mm long; blades strongly obliquely elliptic-curved, with very widely rectangular-curved lower edge and concave upper edge, acute at the apex, 5-6 x 3-3.5 mm; auricle 0.5 mm long, claw 6 mm long. Pods sessile, elliptic, straight, 12-15 mm long, 6 mm high and 8 mm wide, flat ventrally, widely rounded dorsally, at the apex very shortly acuminate, unilocular; valves pale brownish, thick, slightly rugulose, covered with appressed, very short, + symmetrically medifixed hairs and with distinctly longer, strongly asymmetrically medifixed hairs up to 0.8 mm long, soon glabrescent and nearly glabrous with age. Seeds 3, nearly globular, 5 x 4 x 3 mm, pale brownish, smooth. Distribution: S.W. Iran, endemic. Occurence: on gypsaceous soil Specimens seen: Iran. Ilam: 10 km S of Zarrin-Abad, Anaran Mountain, 1200-1300 m, 6.8.1989, Akhani 5540 (MSB). — Khuzestan: 11 km W Haft-Kel, 300 m, 13.3.1959, Pabot 315 (G) - 25 km N of Haftgel towards Izeh, 550 m, 25.5.1973, I/ranshahr & Mousavi 15593-E (W) -— 8 km W Behbehan, 15.4.1959, Pabot 12435-E (W) & 621 (G) — Zeydoun, 65 km SE Soveyreh, road from Mahshahr to Genaveh, 350 m, 29.2.1972, Iranshahr & Termeh 14957-E (W) - Anhal, 47 km from Masdjed-Suleiman to Lali, 16.3.1978, Jranshahr & Termeh 14964-E (W). — Boyer Ahmadi: 45 km from Dogonbadan to Behbehan, 500 m, 3.3.1972, Iranshahr & Termeh 40843-E (W). — Ears: 4 km S Fahlian, 900 m, 10.5.1959, Pabot s.n. (G) — bei Kaserun, 8.5.1885, Stapf 1068 (K, WU) - Kazerun to Rudak, 800 m, 10.4.1974, Davis & Bokhari D.56516 (K) — Kotal Komarech, 4.5. 1885, Stapf 1143 (WU). 169 Astragalus mucronifolius Boiss., Diagn. pl. orient., ser. 1, 2: 68. 1843 = Tragacantha mucro- nifolia (Boiss.) Kuntze, Revis. Gen. 2: 946. 1891. Holotype: [Iran] Persia prope Ispahan, P.M.R. Aucher-Eloy 4400 (G-BOIS!; iso: BM!, G!, LE!, OXF!, P!, W!). = A. mucronifolius var. brevidentatus Sir). & Rech.f., Ann. Naturhist. Mus. Wien 58: 72. 1951 = A. mucronifolius f. brevidentatus (Sirj. & Rech.f.) Parsa, Fl. Iran 9: 132. 1966. Holotype: [Iran] Inter Jezd et Kermanshahan, 22.-23.4.1948, K.H. Rechinger, P. Aellen & E. Esfandiari 2842 (W!; iso: B!, E!, G!, IRAN, K!). = A. taftanicus Parsa, Kew Bull. 1948: 195. 1948. Holotype: [Iran] S. Persia, Taftan (South), 2200 m, 9.5.1939, A. Parsa 658 (K!). A. taftanicus var. multiflorus Parsa, Kew Bull. 1948: 196. 1948. Holotype: [Iran] S. Persia, Shiraz, Khafr, 12.4.1939, A. Parsa (K!). Plants spiny, cushion-like shrublet up to 12 cm tall, with appressed to subappressed, symmetrically to strongly or extremely asymmetrically medifixed or subbasifixed, merely white hairs. Caudex up to 10 mm in diameter, with short, dark brown branches, in upper part covered with remnants of old leaves . Stems of the year 1-2 cm long, the dense indumentum of fine, subbasifixed white hairs up to 2 mm long is not visible because of the very dense cover of leaves with their stipules. Stipules pale yellowish, 5-11 mm long, triangular-acuminate, shortly spiny, distinctly reticulately nerved, adnate to the petiole for 3-5 mm, densely ciliate at the margins with basifixed hairs up to 0.5(—1) mm long, otherwise mostly glabrous. Leaves 6-15 cm long, in the lower half without leaflets; petiole and rhachis up to 2 mm thick, rigid, pungent, sparsely to loosely covered with appressed, irregularly medifixed hairs 0.4-0.7 mm long, often soon glabrescent to glabrous. Leaflets in 3-6 pairs, thickish-leathery, narrowly elliptic or elliptic to obovate, 6-10 x 2-7 mm, at the apex abruptly to subabruptly acuminate into a spine 1.5-3 mm long, prominently nerved, on upper side rather densely, on underside sparsely to loosely covered with subappressed, symmetrically to strongly asymmetrically medifixed hairs up to 1 mm long. Peduncles 1-2.5 cm long, sparsely to rather densely hairy. Racemes loosely 3-10-flowered. Bracts whitish-hyaline, narrowly triangular, 1-2 mm long, ciliate. Pedicels 1-1.5 mm long, sparsely hairy. Bracteoles 0.5-1.5 mm long, linear, at the base of calyx. Calyx campanulate, 6-7 mm long, sparsely covered with extremely asymmetrically medifixed and basifixed hairs 0.2-0.5 mm long; teeth narrowly triangular, ca. 3 mm long, acuminate-pungent, with elevated midnerve. Petals pink, the keel whitish. Standard 12-14 mm long; blade upcurved above the claw, elliptic, 7 mm wide, slightly retuse at the apex, gradually narrowed into the cuneate short claw. Wings 11-13 mm long, slightly curved; blades oblong, rounded at the apex, 7 x 3 mm; auricle ca. 1 mm long, claw curved, 5-6 mm long. Keel 10-12 mm long; blades obliquely elliptic-triangular, with widely curved lower edge and straight upper edge, acute at the apex, 5 x 3 mm; auricle 0.5 mm long, claw 6-7 mm long. Ovary sessile, villous; style glabrous. Pods sessile, obliquely elliptic to elliptic, straight, 10-12(-15) mm long, 4-6(-9) mm high and 3.5-5(-6) mm wide, obtusely carinate by the thick median nerve ventrally, carinate or rounded dorsally but with prominent midnerve, at the apex shortly acuminate, unilocular; valves brownish, thick, slightly transversely rugulose-nerved, covered with very fine subappressed to ascending hairs up to 1 mm long, soon glabrescent and often glabrous when ripe. Seeds 2-3, rectangular, 3 x 2 mm, brown, irregularly pitted. Distribution: In southwest, southeast and central Iran. 170 Specimens seen: Iran. Esfahan: inter Ab-ı Azan (Abyazan) et Rahmatabad, 60-70 km NW Ardestan, 1150 m, 26.5.1974, Rechinger 46504 (M, W) — 15 km SW of Isfahan, 4.6.1974, Alava 13536 (TUR) — Kouhhaye Kolah-Ghazi, 40 km from Esfahan to Shiraz, 1700-2000 m, 6.6.1984, Termeh & Tehrani 41408-E (W) — Natanz, 1500 m, 19.5.1975, Rechinger 52058 (M, W) - 20° SE of Esfahan, 1730 m, 11.4.1962, Furse 1361 (K, W) — prope Ispahan, Aucher-Eloy 4400 (BM, G, G- BOIS, LE, OXF, P, W) — between Aran and Kashan, 850 m, 28.4.1974, Dinie & Bazargan 7972 (W). — Fars: pr. Schiras, 10.5.1842, Kotschy 354 (MSB, WU) — pr. Gere inter Abuschir et Schiras, 18.3.1842, Kotschy 58 (MSB, WU) - Shiraz, Khafr, 12.4.1939, Parsa (K)! Jahrom, Koelz 14606 (W) — Sarvestan, Gauba 525 (W). — Kerman: entre Bam et Jiroft, 1450 m, 19.4.1972, Leonard 5617 (K, M, MSB, W) — 9 km N Kerman, 1790-1900 m, 8.5.1961, Pabot 6952 (G) — 3 km NE Sirjan, 1720 m, 7.5.1961, Pabot 6852 (G) — Bashagard, Ghorichi, 1000 m, 20.2.1973, Jranshah & Mousavi 15466-E (W) — 150 km SE Kerman versus Bam, 26.3.1965, Rechinger 27152 (W) - Mahan, entre Kerman et Bam, 13.6.1960, Dadashzadeh DK430 (G) — 24 km S Mahan, 26.3.1965, Rechinger 27147 (K, M, W) — inter Mashiz et jugum Khan-e Sorck (inter Kerman et Sirdjan), 2000-2580 m, 27.4.1948, Rechinger, Aellen & Esfandiari 3031 (W) — 6 km from Darzin on the road to Jiroft, 1450 m, 14.4.1975, Foroughi 15908 (W) — prope Kerman, IV.1859, Bunge (K) - S. of Kerman, 26.3.1965, Lamond 75 (M) - hills S of Deh Bakri, 1150-1700 m, 1.5.1975, Parris 74.264 (K) — 4 km SW of Baft, 2000 m, 2.5.1975, Parris 75.319 (K). — Markasi: 42 km from Qum to Arak, 1360 m, 21.5.1972, Amin & Mousavi 15961 (W). — Semnan: 30 km SW of Damghan, 1200 m, 5.5.1974, Wendelbo & al. 11217 (H, W) - 20 km E of Damghan, 1100 m, 5.5.1974, Wendelbo & al. 11191 (W). — Sistan: Khash, Karavandar Gebirge, 1.4.1949, Mirzajan 583-E (W) — Gebirge bei Khash, 26.3.1949, Mirzajan 581-E (W) — Sangan, Ladis road, 1550 m, 15.3.1974, Foroughi 10819 (W) — Sangan, between Khash and Ladis, 1500 m, 15.3.1974, Jranshahr & Ershad 16332-E (W) — 76 km NE from Zarbaz versus Qale Zaboli, 1200 m, 11.3.1974, /ranshahr & Ershad 16357-E (W) — prope pag. Deh Pabid, 1420 m, 28.3.1973, Sojak 307 (PR) — prope opp. Khash, 28.3.1973, Sojak 461 (PR) - in declivibus australibus vulcani Taftan prope Torshab, 1900-2300 m, 26.4.1977, Rechinger 54782 (W) - dto., 2500-2700 m, Assadi in Hb. Rechinger 54838 (M, W), 54839 (M, W). — Yazd: inter Yezd et Kermanshahan, 22.-23.4. 1948, K.H. Rechinger, P. Aellen & E. Esfandiari 2842 (B, E, G, IRAN, K, W). Astragalus ovoideus Sir}. & Rech.f., Ann. Naturhist. Mus. Wien 58: 72. 1951 = A. mucronifolius Boiss. var. ovoideus (Sirj. & Rech.f.) Parsa, Fl. Iran 9: 131. 1966. Holotype: [Iran] Prov. Kerman, Montes Djamal Bariz inter Bam et Djiroft, infra Deh Bakri, ca. 1800 m, 8.-10.5.1948, K.H. Rechinger, P. Aellen & E. Esfandiari 3865 (W!; iso: E!, G!). = A. mucronifolius Boiss. subsp. robustus Sirj. & Rech.f., Ann. Naturhist. Mus. Wien 58: 71. 1951 = A. mucronifolius var. robustus (Sirj. & Rech.f.) Parsa, Fl. Iran 9: 131. 1966. Holotype: [Iran] Prov. Kerman, inter Saidabad et jugum Chah Choghuk, 1700-1900 m, 28.4.1948, K.H. Rechinger, P. Aellen & E. Esfandiari 3172 (W!; iso: B!, E!, G!, K!, M!). Plants nearly acaulescent, caespitose-spiny, up to 25 cm tall, sparsely covered with appressed to subappressed, partly asymmetrically to extremely asymmetrically or even subbasifixed white hairs 0.3-1(-1.5) mm long. Caudex branched with short to elongated branches up to 6 cm long, densely covered with remnants of old stipules and rhachides. The very short stems densely hairy. Stipules thickish, whitish to straw-coloured, 5-10 mm long, narrowly triangular, with distinct longitudinal nerves, adnate to the petiole for 1-3 mm, ciliate at the margins, at the base covered with long basifixed white hairs, otherwise glabrous, more rarely appressed hairy. Rhachis strong, at the base up to 2 mm thick, pale greenish, (4—)6—18 cm long, pungent, finely striate-sulcate, sparsely hairy to glabrescent, with fine, extremely asymmetrically medifixed hairs up to 1.5 mm long, the lower half or 2/3 without leaflets, the 171 free apex 1-3 cm long. Leaflets in 3-5 pairs, remote, rigidly thickish, narrowly to widely elliptic or obovate, 5-10 x 2-6 mm, shortly acuminate and with a yellowish cusp 1.5-3 mm long, the midvein on underside prominent, yellowish, both sides loosely to rather densely covered with appressed, asymmetrically to more rarely symmetrically medifixed hairs up to | mm long. Peduncles 2-8 cm long, curved, hairy like the rhachis. Racemes 3-12-flowered, at fruiting time elongated and up to 8 cm long. Bracts whitish-hyaline, 1-2 mm long, narrowly triangular, ciliate with basifixed hairs. Pedicels ca. | mm long, hairy. Calyx 5-7 mm long, campanulate-tubular, sparsely covered with subbasifixed hairs up to 1 mm long; teeth from the narrowly triangular base long acuminate, somewhat pungent, 2-4 mm long. Petals pink, the standard with darker longitudinal stripes. Standard 10-13 mm long; blade obovate to nearly orbicular, 6-8 mm wide, slightly retuse at the apex, gradually narrowed into the short claw. Wings 10-12 mm long; blades narrowly obovate, slightly curved, rounded at the apex, 8 x 3 mm; auricle ca. 1 mm long, claw 4.5 mm long. Keel 9-10 mm long; blades triangular, with widely rectangular-curved lower edge and + straight upper edge, acutish at the apex, 5 x 3 mm; auricle short, claw 5 mm long. Pods sessile, ovoid, 9-15 mm long, 7-11 mm high and 7-10 mm wide, ventrally and dorsally with prominent flat vein, at the apex with a minute beaklet, unilocular; valves hard, brownish, finely and densely transversely rugulose-veined, rather densely covered with fine, ascending, basifixed hairs up to 2 mm long and with few very short, medifixed, appressed hairs, soon glabrescent and mostly completely glabrous when fully ripe. Seeds 4-5, nearly quadrangular, 3 x 3 mm, greyish, densely irregularly rugulose-pitted. Distribution: In southwest, south and central Iran. Specimens seen: Iran. Fars: 25 km W of Shiraz, 28.8.1960, Zohary 583 (HUJ) — Dudehak, 25 km NW of Shiraz, 1900 m, 8.6.1965, Grant 17668 (W) — 20 km N Shiraz, 1760 m, 28.4.1961, Pabot 6013 (G) — 27 km N Shiraz, 1630 m, 28.4.1961, Pabot s.n. (G) — 10 km SW of Shiraz, 1600 m, 12.4.1964, Grant 15498 (W) — entre Shiraz et Persepolis, 800-1000 m, 2.5.1956, Schmid 5513 (W) — 53 km NW Jahrom vers Shiraz, ca. 1100 m, 30.4.1961, Pabot 6292 (G) — 75 km SE Shiraz, 1450 m, 30.4. 1961, Pabot 6240 (G, W) — 80 km SE Shiraz, 1430 m, 30.4.1961, Pabot 6249 (G) — 12 km N Shiraz, 30.5.1965, Ledingham & Assefi 4075 (W) — 25 km E Lar, 660 m, 2.5.1961, Pabot 6442 (G).— Hormozgan: Hajiabad, 134 km S Sirjan, 1290 m, 6.5.1961, Pabot 40815-E (W) & 6795 (G). — Kerman: 3 km NE Sirjan, 1720 m, 7.5.1961, Pabot 6851 (G) — 20 km NE Sirjan, 1820 m, 7.5.1961, Pabot 6861 (G) — inter Saidabad et jugum Chah Choghuk, 1700-1900 m, 28.4.1948, Rechinger, Aellen & Esfandiari 3172 (B, E, G, K, M, W) — zwischen Sirjan und Hajjiabad, 5.6.1976, Manutshehri 508 (MSB) — Djiroft, 2.4.1965, Soltan 6390-E (W) — Montes Djamal Bariz inter Bam et Djiroft, infra Deh Bakri, ca. 1800 m, 8.-10.5.1948, Rechinger, Aellen & Esfandiari 3865 (E, G, W) — 40 km S Kerman, 1500 m, 1.4.1964, Grant 15393 (W) — prope Mahan, 1800 m, 6.5.1977, Rechinger 55242 (M, W) — prope opp. Mahan, 1700 m, 24.3.1973, Sojak 177 (PR) — inter Chabbis [Shahdab] et Kerman, IV.1859, Bunge (K). — Khuzestan: N of Dezful, above Damsite, 600 m, 28.3.1959, Wendelbo 55 (BG) — Near the big Dez dam [N Dezful], 600 m, 28.3.1959, Wendelbo 55 (BG). — Sistan: 12 km SE a vico Bazman, 13.4.1973, Sojak 1910 (PR) — inter Khash et Iranshahr, 1600 m, 17.5.1948, Rechinger, Aellen & Esfandiari 4041 (W) — Iranshahr, 25.2.1949, Sharif 592-E (W) — Iranshahr to Bam, Kuh-e Pansareh, 25 km N Bazman, 900-1300 m, 27.4.1983, Termeh, Moussavi & Tehrani 41101-E (W) — Kahnouk, btween Zahedan and Khash, 1700 m, 22.4.1983, Termeh, Moussavi & Tehrani 41108-E (W). 172 Astragalus talimansurensis Sirj. & Rech.f., Anz. Österr. Akad. Wiss., Math.-Naturwiss. Kl. 1953 (90): 180. 1953 = A. mucronifolius Boiss. var. talimansurensis (Sirj. & Rech.f.) Parsa, Fl. Iran 9: 131. 1966. Holotype: [Iran] Bakhtiari, Talimansur, 28.4.1940, W. Koelz 15090 (W!: foto K!). Plant spiny, cushion-like shrub up to 15 cm tall (up to 45 cm tall according to the collector), in most parts densely covered with appressed to subappressed, + symmetrically to more rarely asymmetrically medifixed white hairs 0.3-1 mm long. Old stems thick, blackish, covered with remnants of old leaves, the stems of the year 1-2 cm long, the dense indumentum of fine, subbasifixed white hairs up to 1 mm long is not visible because of the very dense cover of leaves with their stipules. Stipules pale yellowish, 6-8 mm long, triangular-acuminate, not spiny at the tip, adnate to the petiole for ca. 3 mm, densely ciliate at the margins with basifixed hairs up to 0.5 mm long, otherwise mostly glabrous. Leaves 3—7 cm long, in the lower half without leaflets; petiole and rhachis up to 1 mm thick, pungent, densely covered with appressed to subappressed, irregularly medifixed hairs 0.2-0.6 mm long. Leaflets in 6-9 pairs, narrowly elliptic to elliptic, 4-6(-7.5) x 1-2 mm, at the apex subabruptly acuminate into a spine 0.5-1 mm long, on both sides densely covered with appressed to subappressed, mostly + symmetrically medifixed hairs up to 0.5(-0.8) mm long. Peduncles 1-2 cm long, densely hairy. Racemes loosely 2—4-flowered. Bracts whitish, narrowly triangular, ca. 2 mm long, ciliate. Pedicels at fruiting time thickish, ca. 3 mm long, hairy. Bracteoles 1.5 mm long, linear, in upper part of the pedicel. Calyx campanulate, ca. 7 mm long, loosely covered with + symmetrically medifixed, short hairs; teeth narrowly triangular, ca. 3 mm long, acuminate, scarcely pungent, with elevated midnerve. Petals pink. Standard ca. 12 mm long; blade suborbicular, ca, 8 mm wide, subabruptly narrowed into the short claw. Wings ca. 11 mm long, blades narrowly oblong, rounded at the apex, 5 x 1.5 mm; auricle ca. 1 mm long, claw 6 mm long. Keel 10 mm long; blades obliquely elliptic-triangular, with widely curved lower edge and nearly straight upper edge, acute at the apex, 5 x 2 mm; auricle 0.5 mm long, claw 5 mm long. Pods sessile, elliptic, straight, 15-17 mm long, 6-6.5 mm high and 3 mm wide, carinate ventrally and dorsally, at the apex shortly acuminate, unilocular; valves pale brownish, thick, slightly transversely rugulose, rather densely covered with very short subappressed hairs. Distribution: Iran, endemic; known only from the type. Specimen seen: Iran. Chahar Mahal Bakhtiari: Talimansur, 26.4.1940, Koelz 15090 (W). Excluded species: The following species, which are included into the section by RECHINGER, DULFER & PATZAK do not belong to Leucocercis: Astragalus semnanensis Bornm. & Rech.f. has only basifixed hairs, persistent petals in fruit and hairy standard. It is the representant of the monotypic sect. Semnanenses Podlech & Zarre. Astragalus crassispinus Bunge was described by the author based on a plant without flowers and fruits. Because of superficial similarities it was erroneously assigned to section Leucocercis. It has only basifixed hairs and clearly belongs to the plant which was later on described as A. noziensis Sirj. & Rech.f. of sect. Anthylloidei (RECHINGER 1958). 173 VI. New typifications and changes of typification in Astragalus-species. A. angarensis Bunge 1868, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 126 in clave [et lc. 15(1): 222. 1869]. — Lectotype (Flora USSR 1946): [Russia] Sibiria baicalensis ad Angaram prope Balagansk, [1830], P.K.N.S. Turczanninov (LE!; iso: BR!, C!, G!, H!, HAL!, K!, LE!, M!, OXF!, P!, W!, WAG!). The lectotypification by PODLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. A. coronilloides Ulbr. 1913, Bot. Jahrb. Syst. 1, Beibl. 110: 14. — Lectotype (designated here): Zentral-China, West-Hupeh, Fang, vii.1901, E.H. Wilson 2386 (K!; iso: E!, P!, W!). A. crassispinus Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 132(1868) in clave [et l.c. 15(1): 230. 1869]. Holotype: Afghanistan, W. Griffith distr. no. 1592 (K). Although the type of A. crassispinus has no flowers and fruits there can be no doubt about the identity with A. noziensis Sirj. & Rech.f., Dansk Biol. Skr. 9(3): 124. 1958. An epitype has to be choosen to show the important characters of the flowers and fruits. — Epitype (designated here): Afghanistan, Prov. Ghazni, Okak, NE altoplanitiei Dasht-e Nawar, ca. 3000 m, ca. 33°50’ N, 67°55’ E, 4.7.1962, K.H. Rechinger 17760 (as A. noziensis) (M; iso: W). A. cuscutae Bunge var. pulcher Beck 1886, Denkschr. Kaiserl.. Akad. Wiss., Math.- Naturwiss. KI. 51: 339. Syntypes: [Iran] prope Kaebuterchan, 1882, Th. Pichler, ac inter Hamadan et Tschitschian. — Lectotype (designated here): near Käbuterchan [Kabutara- hang], 2130 m, 14.5.1982, Pichler (W!; iso: K!). A. cytisoides Bunge 1868, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 128 in clave [et l.c. 15(1): 224. 1869]. — Lectotype (Flora USSR 1946): [Kazakhstan] Karatau occ. Turkestaniae, 5000', 11.5.1866, N.A. Sewerzow (LE!; iso: G-BOIS: fragment!, P!. The lectotypification by PODLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. A. dilutus Bunge, Del. Sem. Dorpat. 1840: 7. 1840. — Lectotype (Fl. URSS 1946): ad rivulum Tuktugem in deserto Tschujae, 10.7.1839, A. Politov (LE!; iso: K!, P!, TUB!). The lectotypification by PoDLEcH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. A. feddei Sirj. 1943, Repert. Spec. Nov. Regni Veg. 52: 229. — Lectotype (designated here): Armenia turcica, Egin, Kota, 6.6.1890, P.E.E. Sintenis 2568 (W!; iso: B!, BR!, K!, LE!, M!, MSB!, WU!). A. gypsaceus Beck var. angustifolius Bornm. 1910, Beih. Bot. Centralbl. 27(2): 332. — Lectotype (designated here): [Iran] in monte Elwend, Th. Strauss (JE!; iso: B!). The lectotype has been named holotype in PODLECH 1988. Because there is a specimen seen by Bornmiiller also in B, a lectotypification is necessary. A. kadschorensis Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 103 in clave [et l.c. 15(1): 182. 1869]. — Lectotype (Flora USSR 1946): Iberia caucasica prope Tiflis, Kadshory, supra ruinas Ker-ogly, 5.6.1866, 4.P. Owerin LE!; iso: G-BOIS!, P!). The lectotypification by PoDLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. A. leansanicus Ulbr., Bot. Jahrb. Syst. 36, Beibl. 82: 62. 1905. Type: [China] Shaanxi merid., in monte Lean shan, Giraldi 4241 (B: destroyed: at K a good drawing and fragment). — Lectotype (designated here): Giraldi 4241: the drawing in connection with the fragment at K!; iso: W!: fragment. 174 . lepsensis Bunge 1868, Mém. Acad. Imp. Sci. Saint Petersbourg 11(16): 25 in clave [et Le. 15(1): 29. 1869]. — Lectotype (Flora USSR 1946): [Kazakhstan] jugo Alatau ad fl. Lepsa et Sarchan, G.S. Karelin & IP. Kirilov 1362 (LE!; iso: BM!, K!, LE!, M!, P!, W!). The lectotypification by PODLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. . leptothalamus Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 135. 1868 [et Lc. 15(1): 232. 1869]. — Holotype: [Turkey] ad ripas Sari prope Karakoi ad montem Argaeum Ciliciae, 4500', 26.5.1859 Th. Kotschy suppl. 198 (G-BOIS!; iso: HUJ!, P!: foto MSB!, W!). The lectotypification by PODLECH 1998 with the type in P is superfluous, because BUNGE quotes the specimen in hb. Boissier only. . leucophanus Bornm. 1906, Beih. Bot. Centralbl. 19(2): 242. — Lectotype (designated here): [Iran] Hamadan, in monte Elwend, v.1897, Th. Strauss (B!; iso: JE!). . macropus Bunge 1847, Arbeiten Naturf. Vereins Riga 1: 238. Syntypes: im Gouvern. Orenburg bei Spaskoje, 4.6.1839 (K!); am Flusse Ilek, 22.5.1841, A. Lehmann. — Lectotype (designated here): am Flusse Ilek, 22.5.1841, A. Lehmann (LE!; iso: K!). No specimen at P. Syntypes have been distributed as Rel. A. Lehmann. no. 351 (K!). . macrotropis Bunge 1868, Mém. Acad. Imp. Sci. Saint Petersbourg 11(16): 127 in clave [et l.c. 15(1): 223. 1869]. — Lectotype (Flora USSR 1946): [Kazakhstan] Songaria, inter Arganaty et Keyssyk-auss, 3.5.1857, P.P. Semenov (LE!; iso: P!: foto MSB!). The lectotypification by PODLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. . macrourus Hohen. subsp. pseudotauricola (‘pseudotauricolus') Ponert, Feddes Repert. 83: 620. 1973. — Lectotype (designated here): [Turkey] Prov. Antalya, Elmali, 9.6.1860, Bourgeau (sub A. tauricolus) (E!: foto MSB; iso: W!). The author cites only the data of the type collection but gives no reference, where the type is housed. A. lectotypification is therefore necessary. . monozyx Bornm. 1908, Mitth. Thüring. Bot. Vereins, n.s. 23: 18. — Lectotype (designated here): [Iran] Hamadan in montibus Wafs, 10.6.1905, Th. Strauss (B!; iso: B!, LE!). . oreades C.A.Mey. 1831, Verz. Pfl. Cauc.: 141. — Lectotype (Podlech & Sytin, designated here): in regione alpina mt. Kasbek, 7200-8600', 27.9.1829, C.A. Meyer (LE!; iso: LE!). . pullus N.D.Simpson 1913, Notes Roy. Bot. Gard. Edinburgh 8: 125. — Lectotype (designated here): [China] Lichiang Range, Yunnan, 10000-10500', vi.1906, Forrest 2379 (&!; iso; El, LEN): . salatavicus Bunge 1868, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 117 [et l.c. 15(1): 201. 1869]. — Lectotype (Flora USSR 1946): [Russia] in alpe Chanakoi-tau Salataviae, i.e. Daghestaniae borealis, 8800', 24.7.1861, A.P. Owerin (LE!; iso: G-BOIS!, P!). The lectotypification by PoDLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. . Semenovii Bunge 1866, Bull. Soc. Imp. Naturalistes Moscou 39(2): 22. — Lectotype (Flora USSR 1946): in der Nähe von Mai-tübe [in valle Ili], 11.5.1857, P.P. Semenov (LE!; iso: P!). The lectotypification by PODLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. . semibilocularis Bunge 1868, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 103 in clave [et lc. 15(1): 183. 1869]. Syntypes: Altai, Ledebour, Meyer, Bunge et alii. [Riddersk, Bunge (LE!)]. — Lectotype: Riddersk (LE!). The lectotypification by PODLECH 1998 with the type in P (hb. Bunge: Fl. orient. Altaica 1839, A. von Bunge: with flower dissection) is 175 superfluous because the type is indicated in Fl. USSR as to come from Ridders and to be in LE. A. sesbanoides Benth. 1835, in Royle, Illustr. Bot. Himal. Mount.: 199. — Lectotype (designated here): Mussooree, J.F. Royle (LIV!; iso: K!, LIV!). A. sungpanensis E.Peter 1938, Acta Horti Gothob. 12: 34. — Lectotype (designated here): China, Szechuan, Sungpan, 6.7.1922, H. Smith 2238 (W!; iso: LD!, PE!). A. thionanthus Bornm. 1906, Beih. Bot. Centralbl. 19(2): 243. Syntypes: Luristan, in monte Schuturunkuh, vi.1899, Th. Strauss; inter Sultanabad et Kum in districtu Chaladschistan, v.1899, Th. Strauss (B!, JE!). — Lectotype (designated here): Luristan, in monte Schuturun- kuh, vi.1899, Strauss (JE!; iso: B!). A. tschujensis Bunge 1868, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 22 in clave [et l.c. 15(1): 24. 1869]. — Lectotype (Flora USSR 1946): in lapidosis alpinis ad Tschujam, A. Politov (LE!; iso: G!, LE!, MSB!, P!). The lectotypification by PopLEcH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. A. tumbatsicus C.Marquand & Airy Shaw 1929, J. Linn. Soc., Bot. 48: 171. — Lectotype (here designated): SE. Tibet, Tumbatse, 3400 m, 5.8.1924, F.F. Kingdon Ward 6057 (K!; iso: E!). A. umbellatus Bunge 1868, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 24 in clave [et l.c. 15(1): 29. 1869]. — Lectotype (Flora USSR 1946): [Russia] insula Nowaja-Semlia ad sinum Kostin-schar, 11.8.1837, A. Lehmann (LE!; iso: P!). The lectotypification by PODLECH 1998 with the type in P is superfluous because the type is indicated in Fl. USSR 1946 as to be in LE. A. variegatus Franch. var. koksuensis Popov 1928, Trudy Sredne-Aziatsk. Gosud. Univ., ser. 8b, Bot. 3: 38. — Lectotype (designated here): Tianschan occid., ad cacumina montis Tschimgan minoris, 4.7.1926, M. Popov & A.I. Vvedensky in Herb. Fl. As. Med. 373 (LE!; iso: B!, BR!, BRNU!, E!, K!; LE!, MA!, MSB!, MW). A. xanthomeloides Korovin & Popov 1928, Trudy Sredne-Aziatsk. Gosud. Univ., Ser. 8b, Bot. 3: 57. — Lectotype (designated here): ad cacumina montium Samarkandicorum prope Sazagan, 13.5.1925, M. Popov in Herb. Fl. As. Med. 400 (LE!; iso: BRNU!, LE!, MA!, MW, PR!, TASH). A. xipholobus Popov 1928, Trudy Sredne-Aziatsk. Gosud Univ., Ser. 8b, Bot. 3: 39. — Lectotype (designated here): Tian-schan occ., prope stat. Sary-agatsch, 28.5.1928, Granitov in Herb. Fl. As. Med. 374 (LE!; iso: B!, BP!, BRNU!, E!, K!, LE!, MA!, MW, P!, PR!, W!). IX. Some new species in genus Astragalus: Astragalus behbehanensis Podlech, spec. nov. (Sect. Caprini). Holotypus: Iran, Kuhzestan, Behbehan, 400 m, 3.3.1972, H. Foroughi 3278 (MSB; iso: HUJ). Differt ab A. akhanii Podlech rhachide pilis ad 3 mm longis (nec 0,3—1 mm longis) obtecto, foliis 11-15-jugis (nec 7—9-jugis), foliolis 6-8 x 4-6 (nec 10-22 x 8-17) mm, apice late rotundatis (nec acutiusculis vel breviter acuminati-mucronulatis), supra glabris (nex laxe pilosis), subtus densiuscule pilis oblique patentibus ad 2,5 mm longis obtectis, racemis 7-10 (nec 2-3)-floris, pedicellis 3-4 (nec 5-8) mm longis, calyce 13-14 (nec ca. 17) mm longo, 176 pilis longioribus obtecto, dentibus 2,5-3 (nec 4-6) mm longis. Lamina alarum auricula 2,5—3 (nec ca. 1) mm longa provisa. Plants acaulescent, ca. 20 cm tall, with white hairs up to 3 mm long. Caudex up to 8 mm thick, at the apex shortly branched. Stipules whitish-membranous, oblong, 10-15 mm long, the lower ones obtuse, the upper ones acute, adnate to the petiole for 2-5 mm, ciliate at the margins with spreading hairs 0.8-1.5 mm long and very sparsely furnished with minute subsessile glands, otherwise glabrous. Leaves 12-20 cm long; petiole 2-6 cm long, like the rhachis striate-sulcate, loosely covered with + spreading hairs |.5—2.5(—3) mm long. Leaflets in 11-15 pairs, elliptic to widely elliptic, 6-8 x 4-6 mm, at the apex widely rounded to slightly emarginate, on upper side glabrous, on underside and at the margins loosely to rather densely covered with ascending hairs 1.8—2.5 mm long. Peduncles 5-8 cm long, hairy like the rhachis. Racemes 7-10-flowered. Bracts whitish-hyalinee, 6-9 mm longe, very narrowly triangular, long hairy. Pedicels 3-4 mm long, spreadingly hairy. Calyx 13-14 mm long, tubular, at the base slightly gibbous, slighty obliquely truncate at the mouth, straw-colored, rather densely covered with very thin, tangled, spreading hairs 1.5—2 mm long; teeth unequal, the upper two from triangular base attenuate, ca. 2.5 mm long, the lower three narrowly triangular, ca. 3 mm long, all densely hairy on innerside. Petals yellow. Standard 22-23 mm long; blade slightly recurved, obovate, 9-11 mm wide, at the apex emarginate, at the base subangularly narrowed into the rather short claw. Wings 21-23 mm long; blade narrowly oblong, rounded at the apex, 9-10 x 3 mm; auricle 2.5-3 mm long, claw 12-14 mm long. Keel 20-22 mm long; blade obliquely elliptic-curved, with widely and nearly rectangular-curved lower edge and distinctly concave upper edge, at the apex obtuse, 8 x 4 mm: auricle ca. | mm long, claw 11-13 mm long. Staminal tube very oblique at the mouth. Ovary with a stipe 1-2 mm long, narrowly elliptic, white-hairy; style glabrous. Legumens unknown. Remark: This new species clearly belongs to the A. caprinus-group (see PODLECH 1988) because of the long spreading hairs and the long auricles of wing blades. It is somewhat isolated here by the leaflets glabrous on upper side and by the rather short calyx teeth. Astragalus bozakmanii Podlech, spec. nov. (Sect. Caprini). Holotypus: Türkei, Prov. Ankara, zwischen Kizilcahamam und Celtikgi Yolu, 6.6.1970, 7. Bozakman & K. Fitz 906 (W). Differt ab A. multijugo DC. indumento e pilis ad 3 mm (nec 0,2-0,5(-1) mm) longis consistente, stipulis a petiolo fere liberis (nec per 4-5 mm adnatis), foliis ca. 15-jugis (nec 20-30-jugis), foliolis utrinque subappresse (nec oblique patenter ad patenter) pilosis, pedun- culis 2-4 cm (nec 7-17 cm) longis, bracteis 5-9 (nec 2-4) mm longis. Plants ca. 15 cm tall, subacaulescent, covered with spreading, white to reddish hairs up to 3 mm long. Caudex slender. Stems up to 3 cm long, sparsely hairy. Stipules whitish-mem- branous, 10-12 mm long, narrowly triangular, nearly free from the petiole, sparsely to loosely hairy. Leaves 15-20 cm long; petiole 4-6 cm long, like the rhachis striate-sulcate, rather 177 densely covered with fine, spreading hairs 1-3 mm long. Leaflets in ca. 15 pairs (in first leaves in ca. 10 pairs), elliptic, 10-13(-18) x 5-6(-8) mm, at the apex minutely acuminate-mucro- nulate, on both sides loosely to rather densely covered with subappressed hairs up to 2 mm long. Peduncles 2-4 cm long, hairy like the rhachis. Racemes loosely 15-20-flowered. Bracts whitish, narrowly ovate to narrowly triangular, 5-9 mm long, loosely hairy. Pedicels 3-5 mm long, spreadingly hairy. Calyx 12-13 mm long, tubular, slightly obliquely gibbous at the base, slightly obliquely cut at the mouth, loosely to rather densely covered with fine spreading hairs up to 3 mm long; teeth 5—6 mm, unequal, the upper two from triangular base attenuate, the lower three narrowly triangular, all densely hairy on innerside. Petals glabrous, yellow, the standard slightly reddish-suffused. Standard 18-20 mm long; blade ca. 9 mm wide, obovate, slightly emarginate at the apex, at the base gradually narrowed into the cuneate claw. Wings 17-19 mm long; blades narrowly oblong, rounded at the apex, 9-10 x 3 mm; auricle ca. 1 mm long, claw 7-9 mm long. Keel ca. 15 mm long; blades obliquely triangular, with widely, nearly rectangular-curved lower edge and slightly curved upper edge, acutish at the apex, 6 x 4 mm; auricle very short, claw 9 mm long. Staminal tube oblique at the mouth. Ovary wit a stipe ca. 2 mm long, densely hairy; style hairy up to the middle, below the stigma with a dorsal line of short spreading hairlets. Legumes unknown. Astragalus spitzenbergeri Podlech, spec. nov. (Sect. Caprini). Holotypus: Türkei, Prov. Antalya, Kuhu Dag S von Elmali, Gebiet von Ciglikara, 16.6. 1969, K. Fitz & F. Spitzenberger 875 (W). Differt ab A. antalyensis Podlech planta sparse pilosa (nec glaberrima), stipulis per 1-2 (nec 3-7) mm petiolo adnatis, foliis 18-22 (nec 7-15)-jugis, calyce 12-13 (nec 10) mm longo dentibus 5-7 (nec 2-3) mm longis, leguminibus semibilocularibus (nec fere complete bilocularibus, pilis tenuibus 2-3 mm long obtectis (nec glabris). Plants 20-30 cm tall, subacaulescent, sparsely covered with white spreading hairs. Caudex strongly divided, with many short, subterranean branches. Stems up to 6 cm long, angular- sulcate, glabrous. Stipules whitish-membranous, 9-12 mm long, narrowly triangular, adnate to the petiole for 1-2 mm, ciliate at the margins with hairs 0.5—1 mm long, otherwise glabrous. Leaves 12-30 cm long; petiole 4-10 cm long, like the rhachis sparsely to loosely furnished with spreading hairs 1-1.5 mm long. Leaflets in 18-22 pairs, narrowly elliptic to elliptic, obtuse or more rarely acutish at the apex, 6-15 x 3-6 mm, glabrous on upper side or with few scattered hairs only, at the margins and on underside especially at the midvein sparsely to loosely furnished with subappressed hairs up to | mm long. Peduncles 0.5-1.5 cm long, glabrous. Racemes loosely 5-9-flowered, several at the base of leaves, sometimes forming a dense , globular head up to 6 cm in diameter in fruit. Bracts whitish, narrowly triangular to linear-acute, 5-7 mm long, sparsely hairy. Pedicels 6-10 mm long, often flexuose. Calyx shortly tubular, 12-13 mm long, sparsely hairy, especially toward the teeth; teeth subulate, 5-7 mm long. Petals yellow. Standard 20 mm long; blades slightly recurved, elliptic to obovate, ca. 10 mm wide, slightly retuse at the apex, at the base gradually narrowed into the cuneate claw. Wings 18 mm long; blades narrowly oblong, rounded at the apex, 9-10 x 3 mm; 178 auricle ca. 1 mm long, claw 9 mm long. Keel 15 mm long; blades obliquely elliptic, with widely rectangular-curved lower edge and slightly curved upper edge, acute at the apex, 5.5 x 4 mm; auricle short, claw 9 mm long. Staminal tube slightly oblique at the mouth. Ovary subsessile, elliptic, densely hairy; style below the stigma with a line of short spreading hairlets. Pods subsessile, elliptic to obliquely elliptic seen from the side, 10-15 mm long, 5-6 mm high and 2.5-3 mm wide, strongly carinate ventrally, slightly grooved dorsally, subabrurptly attenuate at the apex into a beak 2-3 mm long, semibilocular; valves straw-colored, thin, rather densely covered with very fine, ascending to spreading hairs 2-3 mm long. Seeds ca. 5 in each locule, 3.5—4 x 2-2.5 mm, brown. Astragalus tecti-mundi Freyn subsp. orientalis Podlech, subsp. nov. (sect. Cenantrum). Holotype: Pakistan, Hunza- und Nagar-Gebiet, Daintar, 3300-3900 m, 1959, F. Lob- bichler 115 (M; iso: MSB). Differt a subsp. tecti-mundi foliis 5-7-jugis (nec 4-5-jugis), foliolis minoribus, subtus sparse ad laxe appresse ad subappresse pilosis (nec glabris vel subglabris). Other specimens seen: Pakistan. Gilgit: Tal NE von Chalt, 3270 m, 27.7.1989, Bosshard, Klötzli & Schaffner 053.10 (MSB) — Gilgit, 1880, Tanner 148 (K) — Karakorum, Shinghai Gah to Pahot Gali, 35°48-55 N, 74°10-17’ E, 3840 m, 31.7.1990, G. & S.Miehe 1302 (MSB) — Hunza- und Nagar- Gebiet, Baltar, 4200 m, Lobbichler 575 (M) — Gharesa Glacier 12 miles E of Nagar, 3890 m, 14.8.1960, Polunin 6239 (E). — Baltistan: Tibet, Balti, 17.7.1856, Schlagintweit cat. 6077 (M) — Shagarthang Valley, 3360 m, 18.7.1892, Duthie 12109 (E) — Ahane Rama, Astor Distr., 3360 m, 5.7.1946, R.R. Stewart 22906 (K). India. Kashmir: Ladak, Tsukse, 3960 m, 26.7.1931, Koelz 2453 (E, W) — Rungdum, 4 km NW of Zuildo, 4130 m, 1980, Southampton University 145 (K) — Khardang La above Leh, Heyde (K) — Zanskar, NW of the Pensi-la, 3650-4270 m, Stolitzka (K) — Suru-Siriwang, Ladak side, really Zans-kar, 1.9.1912, R.R. Stewart 290 (K). — Himachal Pradesh: Lahul, Rangcha Galli, 4420 m, 8.7. 1941, Bor 14018 (E, K) — Kulu-Lahaul, near Pasparag, 6.7.1888, Drummond 23484 (E, K). China. Xinjiang: Distr. Taxkorgan, 2.7.1978, Xinjian Exped. 1160 (WUK). — Xizang: SE. Ti- bet, Gyamda Chu, NE tributary, Pasum Tso NE, 4700-4800 m, 30°03’ N, 93°59’ E, 31.8.1994, Miehe & Wiindisch 94-250-2 (MSB) — Tibetan Himalaya, Everest E, Kama Chu, N bank of upper Kangchung Gl., 5130 m, 28°59’ N, 87°03’ E, 14.10.1989, Dickoré 6383 (Hb. Dickore). Astragalus poluninii Podlech, spec. nov. (sect. Chlorostachys). Holotypus: Nepal, Gum, near Rara, 7.000 ft, 27.8.1952, O. Polunin, W.R. Sykes & L.H.J. Williams 3049 (E!). Differt ab A. chlorostachyde Lindl. cui similis sparsissime ad sparse pilosus (nec partim laxe ad densiuscule vel ipse dense pilosus), bracteis 1,5—3 (nec 3-8) mm longis, caducissimis, vexillo 15-16 mm (nec 10-13 mm) longo. Plants more than 40 cm tall (basal parts missing), very sparsely to sparsely furnished with + appressed hairs 0.3-0.5 mm long. Stem more than 40 cm long, angular-sulcate, in middle part 3-4 mm in diameter, erect, glabrous. Stipules green, 3-6 mm long, narrowly triangular, free, sparsely black and white ciliate, at the base and the base of petiole with minute, sessile 179 glands. Leaves 6-20 cm long; petiole 1-2.5 cm long, like the rhachis dinstinctly grooved on upper side, sparsely hairy or with scattered hairs only. Leaflets in 9-13 pairs, narrowly elliptic, 8-20 x 3-7 mm, acute to shortly acuminate at the apex, on upper side glabrous or with few scattered hairs only, on underside sparsely white hairy. Peduncles several in one stem, in the axils of the middle and upper leaves, 6-10 cm long, striate, glabrous or subglabrous. Racemes 4-11 cm long, loosely many-flowered, later on elongating and up to 25 cm long. Bracts falling at the beginning of anthesis, greenish to whitish-membranous, narrowly triangular, 1.5-3 mm long, black and white ciliate with hairs up to 1 mm long, near the base with few minute sessile glands. Pedicels at the beginning of anthesis 1-2 mm long, sparsely hairy, in fruit up to 5 mm long. Bracteoles absent. Calyx campanulate, 6-7 mm long, at the base slighly oblique, slightly obliquely cut at the mouth, subglabrous or sparsely black hairy, at the upper margin and teeth sparsely ciliate; teeth unequal, the lower three subulate, 1-1.5 mm long, the upper two remote, triangular, 0.4-0.5 mm long. Petals pale yellow, with age or dry sometimes pink-suffused. Standard 15-16 mm long; blade elliptic, 6-7 mm wide, emarginate at the apex, gradually narrowed at the base into the rather long claw. Wings ca. 15 mm long; blades narrowly oblong, rounded at the apex, ca. 7 x 2 mm; auricle ca. 2 mm long, claw ca. 8 mm long. Keel ca. 14 mm long; blades obliquely elliptic-triangular, with in the middle widely curved lower edge and = straight upper edge, obtuse at the apex, ca. 6 x 3 mm; auricle 0.5-1 mm long, claw ca. 8 mm long. Staminal tube oblique at the mouth. Ovary on a stipe 5-7 mm long, glabrous; stigma glabrous. Legumes (immature) on a stipe 8-10 mm long, narrowly elliptic to oblong, 15-17 mm long, ca. 6-7 mm high and wide, at the apex subabruptly narrowed into the straight beak 3-4 mm long; valves thin, pale yellowish, glabrous. Astragalus rhododendrophila Podlech & L.R.Xu, spec. nov. (sect. Chlorostachys). Holotypus: China, Xizang, S. Tibet, N of Lhiinze, Subansiri tributary, Xezar W of Sangngaggoiling, 4070 m, 28°36’ N, 92°32’ E, Betula-Rhododendron-forest, 27.7.1994, G. Miehe & U. Wündisch 94-107-7 (MSB!). Differt ab A. tumbatsico C.Marquand & Airy Shaw caulibus in parte basali densiuscule pilis albis confuse patentibus ad 1,5 mm longis instructis (nec glabris vel pilis nigris brevibus appressis tantum instructis), foliolis 10-14 (nec 7-9)-jugis, bracteis albi-membranaceis, anguste triangularibus, 3 mm longis (nec viridibus late ovatis, 4-7 mm longis), lamina carinae distincte majore, 7 x 3,5 (nec 4 x 2,5) mm, leguminibus dense praecipue nigri-pilosis (nec glabris). Plants 50-60 cm tall. Stem 45-52 cm long, 1-3 mm in diameter, hollow, striate, in lower parts rather densely covered with tangled, spreading, white hairs 0.5-1 mm long and with few appressed black hairs 0.3-0.4 mm long, more rarely partly glabrescent and with age also completely glabrous, in upper parts sparsely to loosely covered with short appressed black hairs 0.3-0.5 mm long. Stipules greenish, narrowly triangular, 5-9 mm long, free from the petiole and from each other, sparsely black and white hairy. Leaves 5-8 cm long, subsessile; rhachis slender, loosely furnished with subappressed to ascending, predominantly black hairs 0.1-0.3 mm long. Leaflets in 10-14 pairs, of thin texture, narrowly elliptic, rounded at the apex, 5-13 x 1-3.5 mm, on upper side glabrous, on underside loosely, in the youth more 180 densely covered with appressed white hairs 0.3-0.5 mm long. Peduncles 6-8 cm long, angular- sulcate, hairy like the upper part of stem. Racemes 4-6 cm long, loosely many-flowerd; axis hairy like the peduncle, flowers + spreading. Bracts whitish-membranous, narrowly triangular, ca. 3 mm long, black hairy, at the margins with minute, sessile glands. Pedicels 3-5 mm long, densely black hairy. Calyx campanulate, 4-5 mm long, strongly obliquely gibbous at the base, obliquely cut at the mouth, loosely to rather densely covered with subappressed black hairs 0.3-0.6 mm long and sometimes also with a few somewhat longer white hairs; teeth slightly unequal, the three abaxial near together, narrowly triangular, ca. 1.5 mm long, the two adaxial remote, triangular, ca. 1 mm long, all densely black hairy on inner side. Petals cream (?), sometimes flushed with violet, all of nearly the same length. Standard ca. 13 mm long; blade ca. 5 mm wide, elliptic, with slightly distinct short tongue-like, widely emarginate apex, below the middle slightly constricted and narrowed into the widely cuneate claw. Wings ca. 12 mm long; blades narrowly oblong, obtuse at the apex, 7 x 2 mm; auricle ca. 2 mm long, claw ca. 5 mm long. Keel ca. 12 mm long; blades obliquely obovate, with widely curved lower edge and slightly concave upper edge, obtuse at the apex, 7 x 3.5 mm; auricle ca. 1 mm long, claw ca. 6 mm long. Staminal tube + truncate at the mouth. Ovary on a slender stipe ca. 5 mm long, spindle like, densely white hairy; style and stigma glabrous. Legumes (immature) with a slender stipe ca. 5 mm long, densely covered with ascending white and predominantly black hairs. Astragalus owirensis [Ali ex] Podlech, spec. nov. (sect. Cystium). Holotype: Pakistan, Chitral, Owir (Nichagh), 11.000 ft, 36°7’ N, 71°55’ E, 11.6.1958, S.A. Bowes Lyon 874 (BM!; iso: L!). Differt ab A. skorniakovii B.Fedtsch. foliis brevibus, 3-4 cm longis (nec 5-20 cm longis) 4-6-jugis (nec 10—20-jugis), pedunculis brevibus, calyce saltem partim pilis arcte asymmetrice medifixis oblique patentibus ad patentibus nigris vel griseis obtecto (nec pilis appressis nigris albisque, + symmetrice medifixis) obtecto, petalis dilute flavis violacei-suffusis (nec violaceis) vexillo obovato apice breviter lingulatim angustato. Plants 10-30 cm tall, acaulescent or nearly so, covered in vegetative parts with + appressed to subappressed, mostly + symmetrically medifixed, white hairs. Caudex up to 10 mm in diameter, slightly to strongly divided, with short branches, covered with remnants of old leaves. Stems, if present, up to 1 cm long, hairy, hidden by the dense stipules. Stipules whitish-hyaline to membranous, 5-8 mm long, adnate to the petiole for 3-4 mm, connate behind the stem up to 1/3 to 1/2 of the length, the free tips often greenish, triangular in the lower stipules, narrowly triangular to subulate-acuminate in the upper ones, glabrous or with few asymmetrically medifixed to basifixed hairs at the margins, especially toward the tips. Leaves 3-4 cm long; petiole 1-2 cm long, like the rhachis rather densely covered with appressed to subappressed, often somewhat flexuose hairs 0.8-1 mm long. Leaflets in 4-6 pairs, narrowly ovate to narrowly obovate, acute to shortly acuminate at the apex, 5-9 x 1.5-4 mm, on both sides loosely to rather densely covered with appressed hairs 0.8-1.5 mm long. Peduncles 2.54.5 cm long, angular-sulcate, sparsely to loosely hairy like the rhachis but in upper part often with mixed black and white hairs. Raceme ovoid, 2.5—4 cm long, rather densely few to many-flowered; axis with black hairs. Bracts whitish-membranous, ovate to narrowly triangular, 4-5 mm long, sparsely to loosely black hairy, at the margins also with 181 basifixed hairs. Pedicels 3-4 mm long, densely covered with tangled, ascending black hairs. Calyx 11-13 mm long, tubular to slightly inflated, scarcely oblique at the base, strongly obliquely cut at the mouth, rather densely covered with strongly asymmetrically medifixed, appressed to ascending, partly even spreading black or black and greyish hairs 0.5-1 mm long; teeth narrowly triangular to linear-acute, 2.5—4 mm long, on innerside black hairy. Petals pale yellow, tinged with violet. Standard 21 mm long; blade 9 mm wide, obovate, at the apex shortly but distinctly tongue-like narrowed, widely retuse, at the base gradually cuneately narrowed. Wings 18 mm long; blades narrowly oblong, shortly narrowed toward the obtuse apex, 7.5 x 2.5 mm; auricle ca. | mm long, claw 10-11 mm long. Keel 16 mm long; blades obliquely elliptic with widely rounded lower edge and + straight upper edge, obtusish at the apex, 7 x 3.5 mm. Staminal tube truncate at the mouth. Ovary with a stipe 1.5-2 mm long, linear, glabrous. Legumes unknown. The type collection was first named A. owirensis by Ali but later on he thought it to be identical with A. skorniakovii B.Fedtsch. The description of the latter species in Fl. Pakistan 100 (Papilionaceae) fits well the description of that species in Fl. USSR but not that of the cited specimen. This is in fact a new species described here. Astragalus argentocalyx [Ali ex] Podlech, spec. nov. (sect. Dissitiflori). Holotype: Pakistan, Chitral, Yarkhun, bezween 36°30’ N, 72°40’ E and 36°48’ N, 73°05’ E, 3050 m, 19.6.1958, S.A. Bowes Lyon 965 (BM). Differt ab A. urgutinus Lipsky foliis 4-5 (nec 6-10) cm longis; foliolis ca. 5-jugis (nec 5-9- jugis), linearibus 10-17 x 1.5-2 (nec 7-10 x 1.5-3) mm, complicatis (nec planis); racemis tantum 5—7-floribus (nec multi-floribus), axi eorum pilis + symmetrice medifixis subappressis albis 0.4-0.5 mm longis et pilis basifixis, patentibus albis et praecipue nigris ca. 1(—1.5) mm longis obtecto (nec pilis symmetrice ad asymmetrice bifurcatis ascendentibus nigris 0.3-0.4 mm longis et pilis basifixis albis tantum ad 1 mm longis obtecto); bracteis ovatis (nec anguste triangularibus), densissime villosis pilis basifixis praecipue nigris obtectis (nec densiuscule pilis partim asymmetrice medifixis albis nigrisque obtectis); calycis tubo late cylindraceo ad leviter ventricoso (nec cylindraceo), densiuscule pilis asymmetrice medifixis ad basifixis subappressis ad fere patentibus nigris 0.3-0.5 mm longis et densiuscule ad dense pilis basifixis, ascendentibus albis 1.5-2 mm longis obtecto (nec pilis nigis subappressis et albis tantum ad | mm longis obtecto); petalis flavi-brunnescentibus (nec albi-violaceis); petalis longioribus; ovario dense pilis albis oblique patentibus ad 1.5 mm longis obtecto. Plants suffrutescent, ca. 40 cm tall. Caudex with a pluricipital root-crown. Stems several, branched, parts of the current year 9-12 cm long, densely to very densely covered with + symmetrically medifixed, appressed white hairs 0.4-0.7 mm long, with some black hairs mixed in. Stipules narrowly triangular, 3-4 mm long, shortly adnate to the petiole, with appressed white and black hairs. Leaves 4-5 cm long, those of the lateral branches mostly distinctly shorter, all subsessile or with petiole up to 0.8 cm long; rhachis slender, hairy like the stem. Leaflets in ca. 5 pairs (on leaves of side-branches often only in 3 pairs), linear, 10-17 x 1.5-2 mm, at the apex acute, on upper side loosely, on underside densely covered 182 with appressed white hairs 1-1.2 mm long, folded. Peduncles 9-12 cm long, rather densely to densely appressed hairy like the stem, below the raceme with increasing black hairs. Racemes 3-4 cm long, loosely 5-7 -flowered, scarcely elongating in fruit; axis covered with + symmetrically medifixed, subappressed white hairs 0.4-0.5 mm long and with basifixed, spreading white and predominantly black hairs up to 1(-1.5) mm long. Bracts 3-4 mm long, ovate, very densely villous, with basifixed, ascending to spreading, predominantly black hairs ca. | mm long. Pedicels 1-2 mm long, white and black hairy; flowers subhorizontally spreading. Calyx widely tubular to slightly ventricous, 13-15 mm long, gibbous at the base, obliquely cut at the mouth, rather densely covered with asymmetrically to strongly asymmetrically medifixed or basifixed subappressed to + spreading black hairs 0.3-0.7 mm long and rather densely to densely with basifixed, tanglex ascending white hairs 1.5-2 mm long; teeth narrowly triangular, 2-3 mm long, hairy on inner side, at the base densely white, at the apex predominantly black hairy. Petals buff yellow. Standard ca. 24 mm long; blade obovate, ca. 8 mm long, slightly emarginate at the apex, at the base slightly angularly passing into the rather long, narrowly cuneate claw. Wings ca. 24 mm long, as long as the standard; blades narrowly obovate, contricted above the auricle, rounded at the apex, 9-9.5 x 3.5 mm; auricle ca. 0.5 mm long, claw 14-15 mm long. Keel 20-21 mm long; blades obliquely elliptic, with widely curved lower edge and nearly straight upper edge, subacute at the apex, 7.5 x 4 mm; auricle minute, claw ca. 13 mm long. Staminal tube truncate at the mouth. Ovary sessile, fusiform, densely covered with ascending white hairs up to 1.5 mm long; style glabrous. Legumes unknown. Remark: The type of this species was recognized by Ali as a new species but later on he has changed his mind according to the determination by A. Borissova who named it A. urgutinus Lipsky. In spite of some superficial resemblance it is a species distinct from the latter which is endemic in the western Zeravshan and Turkestan Ranges. Astragalus bingoellensis Podlech, spec. nov. (§ Dissitiflori). Holotype: Turkey, Prov. Mus, [B8] Buglan Gegidi (pass) E Solhan, between Bingöl and Mus, 1640 m, 17.7.1983, M. Nydegger 18265 (MSB; isotype: BASBG). Plantae 35 cm altae, dense pilosae. Stipulae membranaceae, 4-8 mm longae, petiolo adnatae. Folia 5-10 juga, foliolis ellipticis 7-18 x 2,5-7 mm, laxe ad densiuscule pilosis. Pedunculus ca. 13 cm longus, pilosus. Racemus ca. 20 cm longus, laxe multiflorus. Bracteae membranaceae, 2,5-3 mm longae, praecipue nigri-pilosae. Pedicelli 2-2,5 mm longi. Calyx 7-8 mm longus, campanulati-tubulosus, pilis asymmetrice medifixis appressis obtectus, dentibus 2—2,5 mm longis. Vexillum 15-17 mm longum et ca. 6 mm latum. Alae ca. 10 mm longae. Legumina sessiles, oblique elliptica, leviter curvata, dense pilis ascendentibus arcte asymmetrice medifixis obtecta. Plants 35 cm tall, caulescent, densely hairy. Stem ca. 10 cm long, branched, angular-sulcate, very densely covered with symmetrically to asymmetrically, often somewhat flexuose, appressed to subapressed white hairs ca. | mm long. Stipules membranous, 4-8 mm long, adnate to the petiole for 1-4 mm, narrowly triangular to nearly subulate, rather densely 183 covered with symmetrically to strongly asymmetrically medifixed white, near the apex also with some black hairs. Leaves 9-13 cm long; petiole 3-4 cm long, like the rhachis grooved on upper side, loosely to densely hairy like the stem. Leaflets in 5-10 pairs, + elliptic, 7-18 x 2.5-7 mm, at the apex shortly acuminate, loosely to rather densely covered with + symmetrically medifixed, appressed white hairs 0.8-1 mm long. Peduncle 13 cm long, angular- sulcate, loosely white hairy like the stem, towards the raceme with increasing, distinctly shorter black hairs. Raceme ca. 20 cm long, loosely many-flowered; axis hairy like upper part of peduncle. Bracts membranous, narrowly triangular, 2.5-3 mm long, with predominantly black hairs. Pedicels 2-2.5 mm long, white and black hairy, flowers erect. Bracteoles none. Calyx 7-8 mm long, campanulate-tubular, slightly obliquely cut at the mouth, rather densely covered with asymmetrically medifixed, often flexuose, appressed to subappressed white hairs 1-1.2 mm long and with sometimes shorter black hairs; teeth subulate, 2-2.5 mm long, densely white hairy on innerside. Corolla violet fading to yellowish-brown or yellowish fading to violet. Standard 15-17 mm long; blade ca. 6 mm wide, elliptic, rounded at the apex, subabruptly narrowed into the short, cuneate claw. Wings ca. 10 mm long; blades narrowly oblong, obtuse at the apex, 6 x 2 mm; auricle 0.8 mm long, claw ca. 5 mm long. Keel ca. 9 mm long, blades obliquely oblong, with in the middle slightly curved lower edge and slightly sigmoid-concave upper edge, acutish at the apex, 4.5 x 2 mm; auricle ca. 0.3 mm long, claw ca. 4 mm long. Staminal tube truncate at the mouth. Ovary sessile, elliptic, white hairy; style glabrous. Legumes (unripe) sessile, obliquely elliptic, slightly curved, with strongly curved dorsal side and slightly concave ventral side, ca. 10 mm long, 3 mm high, attenuate at the apex into a beak ca. 3 mm long; valves densely covered with strongly asymmetrically medifixed, ascending white hairs 1—1.2 mm long. Astragalus doabensis Podlech, spec. nov. (sect. Dissitiflori). Holotype: North Afghanistan, Doab [Doabe Mekhe Zarin], 5700’, 30.4.1937, R. Meinertz- hagen (BM!). Plantae ca. 30 cm altae, pilis symmetrice medifixis appressis obtecta. Stipulae 4-6 mm longae. Folia 5-12 cm longa. Foliola 5-6-juga, remota, elliptica. 9-17 x 3-8 mm, supra laxe subtus densiore pilosa. Racemus pedunculo ca. 18 cm longo albi-piloso suffultus, laxe 5-6- florus. Bracteae 2,5-3 mm longae. Pedicelli 2 mm longi. Calyx 18-20 mm longus, tubulosus, pilis albis nigrisque 0,4-1 mm longis obtectus, dentibus 3-4 mm longis. Vexillum 25 mm longum, 8 mm latum, longe unguiculatum. Alae 25 mm longae, lamina 6 x 2 mm. Carina 21 mm longa. Ovarium breviter stipitatum lineare, albi-pilosum. Legumina ignota. Plant 30 cm tall, covered with + appressed, mostly symmetrically medifixed hairs. Caudex branched. Stems few, erect, 10-12 cm long, densely covered with white hairs 0.9-1 mm long. Stipules greenish, narrowly triangular, 4-6 mm long, shortly adnate to the petiole, loosely to rather densely white and black hairy. Leaves 5-12 cm long; petiole 2-4 cm long, like the rhachis densely hairy like the stem. Leaflets in 5-6 pairs, remote, elliptic, 9-17 x 3-8 mm, acuminate at both ends, on upper side loosely, on underside loosely to rather densely covered with white hairs 0.6-1.2 mm long. Peduncle ca. 18 cm long, erect, loosely to rather densely white hairy. Raceme short, loosely 5-6-flowered. Bracts whitish, narrowly triangular or from ovate base acuminate, 2.5-3 mm long, white and black or predominantly black hairy. Pedicels 184 2 mm long, white and black hairy. Calyx 18-20 mm long, tubular, obliquely gibbous at the base, obliquely cut at the mouth, densely covered with appressed to subappressed, + symmetrically medifixed white and black hairs 0.4-1 mm long; teeth subulate, 3-4 mm long, on innerside densely covered with very thin white hairs. Petals color unknown. Standard ca. 25 mm long; blade widely elliptic, ca. 8 mm wide, emarginate at the apex, at the base gradually narrowed into the long claw. Wings 25 mm long; blades narrowly oblong, rounded at the apex, 6 x 2 mm; auricle short, claw 19 mm long. Keel 21 mm long; blades obliquely elliptic, with widely curved lower edge and nearly straight upper edge, obtuse at the apex, 6 x 3 mm; auricle short, claw 15 mm long. Staminal tube truncate at the mouth. Ovary with a stipe ca. 1.5 mm long, linear, white hairy; style glabrous. Legumes unknown. Astragalus fruticulosus Podlech, spec. nov. (sect. Dissitiflori). Holotype: Afghanistan, Prov. Badghis, 8 km E Qades, 1380 m, Lössboden, 63°30’ E, 34°48’ N, 4.5.1977, D. Podlech & Kh. Jarmal 29825 (MSB!). Affinis A. xyloclado Rech.f. & Gilli, sed differt stipulis 1,5-2,5 mm tantum (nec 4-5 mm) longis, foliolis in caule principali 4-5 (nec 7—8)-jugis, supra laxe pilosis (nec marginem versus tantum pilosis), lamina alarum late elliptica, 5 x 4,5 mm (nec anguste oblonga, 6—7 x 2 mm). Plants fruticulose, 8-18 cm tall, in vegetative parts covered with appressed to sub- appressed white hairs, the stipules also with black hairs. Caudex diffusely branched. Stems several, prostrate to erect, branched, with numerous non-flowering, short brachyblasts, the older parts with greyish-brown bark, stems of the current year 2-15 cm long, very densely covered with + symmetrically medifixed, appressed white hairs 0.3-0.6 mm long. Stipules greenish, triangular, 1.5—2.5 mm long, shortly adnate to the petiole, rather densely white and black or sometimes predominantly black hairy. Leaves of the main stems 2-3.5 cm long; petiole 0.5—0.8 cm long, like the rhachis narrowly grooved on upper side, rather densely to densely white hairy like the stem; leaves of the brachyblasts shorter, nearly sesssile. Leaflets in 4-5 pairs, narrowly elliptic, 3-8 x 1-2.2 mm, those of brachyblast leaves smaller, all rounded or obtuse at the apex, on upper side loosely, on underside densely covered with appressed white hairs 0.6-1 mm long. Peduncle singular in lower part of the stem, 1.5-5 cm long, at fruiting time up to 10 cm long, rather densely hairy like the stem, but toward the raceme also with black hairs. Raceme at anthesis short, densely 4-10-flowered, at fruiting time strongly elongated and 4-6 cm long; axis densely covered with spreading, mostly basifixed hairs white hairs 0.3-1 mm long and with few medifixed to basifixed, + appressed to ascending, shorter black hairs. Bracts whitish-membranous, 1.5-2 mm long, ovate, covered with + spreading, mostly basifixed white and black or predominantly black hairs. Pedicels 0.5-1 mm long, white and black hairy. Calyx 11-14 mm long, tubular, slightly gibbous at the base, nearly straightly cut at the mouth, densely villous, with thin, tousled, spreading, basifixed white hairs 1-1.5 mm long and with ascending to + spreading, basifixed to asymmetrically bifurcate, distinctly shorter black hairs; teeth linear-acute to narrowly triangular, 1-2 mm long, densely white hairy on innerside. Petals yellowish.. Standard ca. 20 mm long; blade obovate, ca. 7 mm wide, rounded to scarcely retuse at the apex, at the base long cuneately narrowed. Wings ca. 20 mm long; blades widely elliptic, rounded at the apex, 6 x 4.5 mm; auricle ca. | mm long, claw 13-14 mm long. Keel ca. 17 mm long; blades obliquely 185 elliptic with in the middle widely and nearly rectangular-curved lower edge and straight to slightly concave upper edge, acute at the apex, 6 x 3 mm; auricle minute, claw ca. 13 mm long. Staminal tube truncate at the mouth. Ovary sessile, linear, hairy; style glabrous. Legumes (unripe) sesssile, linear, strongly falcate, ca. 40 mm long, | mm in diameter; valves densely covered with appressed, symmetrically to strongly asymmetrically medifixed white and a few black hairs mixed in up to 1 mm long. Astragalus lanzhouensis Podlech & L.-R.Xu, spec. nov. (sect. Dissitiflori). Holotypus: China, Prov. Gansu, Lanzhou, 1600 m, 12.6.1996, Lang-Ran Xu 96-005 (MSB; iso: WUK). Differt ab A. stenoceras stipulis 3-4 (nec 1,5-2) mm longis, alte connati-vaginantibus (nec inter se liberis), foliis 8—12-jugis (nec 4-7-jugis), bracteis 3-4 (nec 1-1,5) mm longis, calyce 11-12 (nec 8-10) mm longo, dentibus subulatis 4-5 mm longis (nec anguste triangularibus, 1-1,5 mm longis), vexillo ca. 16 mm (nec 20-22 mm) longo, leguminibus curvatis, 17-20 mm longis, mere albi-pilosis (nec rectis, 20-35 mm longis, albi-nigri-pilosis). Plants 10-25 cm tall, with + symmetrically medifixed, appressed to subappressed hairs. Stems branched near to the base, 7— 20 cm long, angular-sulcate, rather densely to densely covered with appressed white hairs, at the nodes sometimes also with a few black hairs, 0.6-1 mm long. Stipules whitish-membranous, triangular, 3-4 mm long, free from the petiole, behind the stem high up connate, with white and sometimes a few black hairs. Leaves 2-5 cm long; petiole 0.5—1 cm long, like the rhachis slender, loosely to rather densely white hairy as stem. Leaflets in 8-12 pairs, narrowly elliptic, acute at the apex, 7-15 x 1-3 mm, covered on both sides loosely or more rarely rather densely with white hairs 0.8-1 mm long. Peduncles 2.5-8.5 cm long, angular-sulcate, loosely hairy like as stem, below the raceme sometimes with a few black hairs. Raceme at fruiting time 2-4 cm long, loosely 4-8-fruited. Bracts whitish- membranous, ovate, long acuminate, 3-4 mm long, sparsely white and black hairy, at the margins also with subbasifixed to basifixed hairs. Pedicels ca. 2 mm long, white and black hairy. Calyx 11-12 mm long, tubular, slightly obliquely cut at the mouth, loosely covered with subappressed white and black, but mostly predominantly white hairs ca. 1 mm long; teeth subulate, (3-)4-5 mm long, white hairy on innerside. Standard ca. 16 mm long; blade narrowly elliptic-oblong, ca. 6.5 mm wide, widely emarginate at the apex, at the base shortly and indistinctly narrowed. Wings ca. 14 mm long, blades narrowly oblong, slightly obliquely emarginate at the otherwise rounded apex, ca. 7 x 2 mm; auricle ca. 0.8 mm long, claw ca. 7 mm long. Keel ca. 11 mm long; blades obliquely obovate-triangular, with widely corved lower edge and straight upper edge, acute at the apex, 5 x 2.5 mm; auricle ca. .8 mm long, claw ca. 6 mm long. Staminal tube truncate at the mouth. Legumes obliquely erect or spreading, shortly stipitate, linear, slightly curved, terete, 17-20 mm long, 2-2.4 mm high, ca. 2 mm wide, carinate ventrally, grooved dorsally, at the apex shortly attenuate into a straight slender beak ca. | mm long, nearly fully bilocular; valves thin but tough, loosely to rather densely covered with appressed to subappressed, somewhat flexuose white hairs 0.6—0.8 mm long. Other specimen seen: China. Prov. Gansu: mountains Beitaschan, N of Lanzhou, 24.6.1957, M.P. Petrov (LE). 186 Astragalus montis-karkasii Podlech, spec. nov. (sect. Dissitiflori). Holotype: Iran, Prov. Esfahan, Kuh-e Karkas (Kuh-i-Kargiz), 33°27’ N, 51°48’ E, in declivibus supra Tar, 2300-2500 m, 27.5.1974, K.H. Rechinger 46555 (M; iso: W). Differt ab A. sitiens Bunge stipulis vaginati-amplectentibus, truncatis, sine apiculis distinc- tis, foliis brevioribus, brevissime petiolatis, foliolis pilis 0,5-1 (nec 1-1,5) mm longis obtectis, calyce 6-7 (nec 7-10) mm longo, dentibus triangularibus, ca. | mm longis (nec subulatis, 1,5—3 mm longis). Plants 13-25 cm tall, covered with appressed, + symmetrically medifixed, white hairs. Caudex divided. Stems several, branched in lower part, the older parts ligneous, short, covered with greyish-brown bark, parts of the current year 3-5 cm long, angular-sulcate, very densely covered with hairs 0.6-1 mm long. Stipules whitish-hyaline, 1.5-3 mm long, adnate to the petiole for ca. 1 mm, behind the stem vaginate-connate, truncate, without distinct free tips, sparsely to loosely hairy. Leaves 2-3 cm long; petiole 0.1-0.6 cm long, together with the short rhachis hairy like the stem. Leaflets in 1-2 pairs, 12-23 x 1.5-2 mm, densely covered on both sides with hairs 0.5-1 mm long. Peduncle 5-7.5 cm long, erect, hairy like the stem. Raceme 5-8 cm long, remotely 5-10-flowered; axis hairy like the peduncle. Bracts membranous, ovate, | mm long, sparsely black hairy. Pedicels ca. 3 mm long, erect, densely predominantly shortly black hairy. Calyx 6-7 mm long, turbinate-tubular, not gibbous at the base, obliquely cut at the mouth, rather densely covered with appressed to subappressed black hairs 0.3-0.5 mm long and with mostly fewer, slightly longer white hairs; teeth triangular, ca. | mm long, on innerside densely covered with very thin, white, basifixed hairs. Petals pale bluish to pale violet. Standard ca. 16 mm long; blade ca. 6 mm wide, elliptic, emarginate at the apex, with a rather long, narrowly cuneate claw. Wings ca. 14 mm long; blades narrowly oblong, obtuse to slightly obliquely retuse the apex, 7.5 x 2 mm; auricle ca. | mm long. claw 7 mm long. Keel ca. 12 mm long; blades obliquely elliptic-curved, with slightly widely curved lower edge and slightly concave upper edge, acutish at the apex, 5 x 2.5 mm; auricle ca. 0.5 mm long, claw 7 mm long. Ovary subsessile, narrowly elliptic, long white hairy; style glabrous. Legumes unknown. Astragalus pravitzii Podlech, spec. nov. (sect. Dissitiflori). Holotypus: Iran, Prov. Fars: Didegan, in alpe, 2000 m, 16.5.1915, A. Pravitz 696 (S!). A. saadatabadensi Podlech ob calycem petalaque brevia similis sed differt stipulis non difformibus, foliolis (1—)2—3(-4) paria (nec 1-3 foliolis tantum), distincte majoribus. Plants 12-15 cm tall, with + symmetrically medifixed, appressed, white hairs, at the stipules and in the inflorescense also with black hairs. Caudex divided with short branches, covered with greyish bark. Stems several, ascending to erect, 5-8 cm long, in basal parts densely, in upper parts loosely to rather densely covered with hairs 0.4-0.8(-1) mm long. Stipules slightly greenish, narrowly triangular, 2-2.5 mm long, shortly adnate to the petiole, sparsely to loosely white hairy, sometimes with a few black hairs at the base. Leaves 1.5-3 cm long; petiole 0.5—1.5 cm long, hairy like the stem. Leaflets in (1—)2—3(-4) pairs, elliptic to obovate, 4-9 x 1.5-4 mm, acute to rounded and minutely acuminate at the apex, on both sides rather densely covered with hairs 0.6-1 mm long. Peduncles 4-5 cm long, loosely white hairy 187 like the stem. Racemes loosely 8-12-flowered; axis hairy like the peduncle. Bracts whitish- membranous, narrowly triangular, ca. 2 mm long, sparsely to loosely covered with partly strongly asymmetrically medifixed, predominantly black hairs. Pedicels 0.5-1 mm long, white hairy. Calyx ca. 7 mm long, tubular, scarcely obliquely gibbous at the base, obliquely cut at the mouth, rather densely to densely covered with appressed white, or white and black hairs 0.6-1 mm long; teeth narrowly triangular, ca. 1 mm long, on innerside nearly glabrous. Petals in dry state yellowish. Standard 14 mm long; blade elliptic, ca. 6 mm wide, slightly emarginate at the apex, at the base cuneately narrowed. Wings 12 mm long; blades narrowly oblong, rounded at the apex, 5 x 2 mm; auricle ca. 1 mm long, claw 7 mm long. Keel 10-11 mm long; blades obliquely obovate, with in upper part widely, nearly rectangular-curved lower edge and slightly concave upper edge, acutish at the apex, 3.5 x 2.2 mm; auricle short, claw 7-8 mm long. Staminal tube truncate at the mouth. Ovary with a stipe ca. 2 mm long, linear, shortly hairy; style glabrous. Legumes (immature) linear, densely covered with appressed white hairs. Other specimen seen: Iran. Fars: Abadeh, Nadjaf-abad, 16.5.1962, Boroumand 12412-E (IRAN). Astragalus recurvatus Podlech, spec. nov. (sect. Dissitiflori). Holotype: Persia, Kashan, Mooteh protected region, ad bifurcationem viae publicae versus Muteh (Mooteh) ducentis, c. 140 km NW Esfahan, 1950 m, 30.5.1974, K.H. Rechinger 46775 (M; iso: W). Differt ab A. sitiens Bunge statura altiore, caulibus pilis brevioribus, 0,2-0,5 mm (nec 0,7-1,1 mm) longis, stipulis longioribus, 6-8 mm (nec 3-4 mm) longis, distincte vaginati- amplectentibus (nec liberis), foliis 4-6 (nec 1-3)-jugis, sparse ad laxe pilosis pilis brevioribus, leguminibus recurvatis ad laterum dorsalem versus, laxe et breviter pilosis. Plant 50 cm tall, covered with appressed hairs. Caudex slender, with a pluricipital root- crown. Stems few, 22-34 cm long, erect, slightly angular-striate, loosely to densely covered with symmetrically medifixed white hairs 0.2-0.5 mm long. Stipules hyaline-membranous, 6-8 mm long, nearly free from the petiole, behind the stem distinctly to up to the middle vaginate-connate, the free tips narrowly triangular to subulate, sparsely to loosely white hairy, sometimes nearly glabrous. Leaves 4-9 cm long; petiole 1.5—2 cm long, like the rhachis loosely hairy. Leaflets in (3-)4-6 pairs, narrowly linear, 12-20 x 1-1.2 mm, loosely covered on both sides with hairs 0.5-1 mm long. Peduncle 4-11 cm long, erect, angular-striate, loosely hairy like the stem. Raceme remotely 10-15-flowered, at fruiting time elongated and up to 17 cm long; axis white hairy like the peduncle but also with a few black hairs mixed in. Bracts whitish-membranous, narrowly triangular, ca. 3 mm long, sparsely to loosely white and black hairy. Pedicels (at fruiting time) 2-3 mm long, white and black hairy. Calyx whitish- membranous, 7-8.5 mm long, tubular, scarcely obliquely gibbous at the base, slightly obliquely cut at the mouth, loosely to rather densely covered with + appressed white hairs up to 1.2 mm long and with black hairs 0.3-1 mm long; teeth subulate, 2-3 mm long, mostly curved, on innerside covered with very thin hairs. Petals color unknown. Standard ca. 20 mm long; blade ca. 7 mm wide, elliptic, emarginate at the apex, with a rather long, narrowly cuneate claw. Wings and keel unknown. Staminal tube truncate at the mouth. Legumes sessile, ascending to spreading or even slightly pendulous, narrowly linear, subterete, slightly to 188 strongly recurved to the dorsal side, 50-60 mm long, ca. 2 mm high and 1.5 mm wide, carinate ventrally, flattish dorsally, at the apex attenuate into a beak 6-10 mm long, incompletely bilocular; valves straw-colored, loosely covered with appressed, + symmetrically medifixed white hairs 0.2-0.4 mm long, later on partially glabrescent. Seeds ca. 8 in each locule, rectangular, 3 x | mm, blackish. Astragalus saadatabadensis Podlech, spec. nov. (sect. Dissitiflori). Holotypus: Iran, Prov. Fars, Pasagarde, 15 km N of Saadatabad, semidesert, 2000 m, 21.5.1964, M.L. Grant 15784 (W). Differt ab A. argyroide Beck stipulis difformibus, basalibus quam superiores distincte longioribus et petiolo longe adnatis (nec omnes conformibus), foliis cum 1-3 foliolis tantum (nec (3—)4—6(—7)-jugis, calyce 7-8 (nec 11-13) mm longo, petalis brevioribus, vexillo ca. 14 (nec 18-22) mm longo. Plants ca. 12 cm tall, with + symmetrically medifixed, appressed, white, only in the inflorescence also with few black hairs. Caudex with a pluricipital root-crown. Stems several, 3-5 cm long, erect, densely covered with hairs 0.8-1 mm long. Stipules different, the lower ones membranous, 5—7 mm long, at the base wide, adnate to the petiole for 2-3.5 mm, densely covered with somewhat flexuose, partly asymmetrically medifixed hairs 1-1.5 mm long, the free tips up to 4 mm long, narrowly triangular, sparsely to loosely hairy, the upper ones greenish, distinctly shorter, narrowly triangular, adnate to the petiole for 1-2.5 mm, sparsely hairy. Leaves 1-4 cm long; petiole 0.5—2.5 cm long, hairy like the stem. Leaflets 1-3, narrowly elliptic to obovate, 7-10 x 2-4, the end-leaflet mosly larger, 12-16 x 4-6 mm, all acute to shortly acuminate at the apex, on both sides loosely to rather densely covered with hairs 0.8-1.2 mm long. Peduncles 5-7 cm long, loosely hairy like the stem. Racemes loosely 6-8- flowered; axis hairy like the peduncle. Bracts whitish-membranous, narrowly triangular, ca. 2 mm long, sparsely to loosely covered with partly strongly asymmetrically medifixed white or white and black hairs, at the base also with few sesssile, minute glands. Pedicels 1 mm long, white hairy. Calyx 7-8 mm long, tubular, slightly obliquely gibbous at the base, obliquely cut at the mouth, rather densely to densely covered with appressed, white hairs 0.8—1 mm long, at the teeth sometimes also with very few black hairs; teeth narrowly triangular, 1-1.5 mm long, on innerside nearly glabrous. Petals in dry state yellowish, slightly flushed with pink. Standard 14 mm long; blade obovate, ca. 7 mm wide, slightly emarginate at the apex, at the base cuneately narrowed. Wings 11 mm long; blades narrowly obovate, rounded at the apex, 5 x 2 mm; auricle ca. 0.5 mm long, claw 6 mm long. Keel 10 mm long; blades obliquely obovate, with in upper part widely, nearly rectangular-curved lower edge and slightly concave upper edge, obtuse to acutish at the apex, 4 x 2 mm; auricle short, claw 6 mm long. Staminal tube truncate at the mouth. Ovary with a stipe ca. 1 mm long, linear, shortly hairy; style glabrous. Legumes unknown. 189 Astragalus sata-kandaoensis Podlech, spec. nov. (sect. Dissitiflori). Holotype: Afghanistan, Prov. Paktia, 9-20 km SE Gardez versus jugum Sata Kandao, 2300 m, 3.6.1967, K.H. Rechinger 35430 (MSB!; iso: W!). Differt ab A. xyloclado Rech.f. & Gilli stipulis 1,5-2 mm (nec 4-5 mm) longis, foliis 2—4- jugis (nec 7-8-jugis), racemis elongatis, remote 3—8 (nec brevibus, dense 7-15-floris), axi racemorum pilis medifixis (nec basifixis) obtecto, calyce pilis + medifixis, subappressis (nec basifixis, patulis) obtecto, ovario 3-4 mm longe stipitato (nec sessile). Plants fruticose, 10-20 cm tall, in vegetative parts with + symmetrically medifixed, appressed to partly subappressed hairs. Caudex up to 10 mm in diameter, strongly branched, covered with grey-brownish bark. Stems several, ascending to erect, branched, with numerous non-flowering, short brachyblasts, the older parts with dark brownish bark, stems of the current year 4-5 cm long, very densely covered with white hairs, near the nodes also with black hairs 0.4-0.3 mm long. Stipules membranous to greenish, triangular, 1.5-2 mm long, very shortly adnate to the petiole, densely white and black or predominantly black hairy. Leaves 1-2 cm long, nearly sesssile; rhachis densely white hairy like the stem, sometimes also with some black hairs. Leaflets in 2-4 pairs, linear or narrowly elliptic, often somewhat recurved, 3-10 x 0.5-1.5(-2) mm, subobtuse to rounded at the apex, often folded, on both sides densely covered with appressed white and sometimes a few black hairs mixed in, 0.8-1.2 mm long. Peduncle 3-6 cm long, white hairy like the stem, in upper part some black hairs. Raceme 2-7 cm long, remotely 3-8-flowered; axis covered with + medifixed, predominantly white hairs. Bracts whitish-membranous, ca. 2 mm long, ovate-triangular, predominantly black hairy, at the margins with subbasifixed to basifixed hairs. Pedicels ca. 2 mm long, densely predominantly black hairy. Calyx 9-11 mm long, tubular, strongly obliquely gibbous at the base, obliquely cut at the mouth, densely covered with somewhat tousled, subappressed to slightly ascending, + medifixed white and black hairs up to 1.5 mm long; teeth narrowly triangular-acuminate, ca. 2 mm long, on inner side densely covered with very thin, white hairs. Petals purplish-violet, fading to pale yellowish. Standard 20 mm long; blade obovate, 9-10 mm wide, slightly to distinctly emarginate at the apex, at the base cuneately narrowed. Wings 18-19 mm long; blades narrowly oblong, rounded at the apex, 9-10 x 3-4 mm; auricle 1-1.2 mm long, claw 10 mm long. Keel 16-17 mm long; blades obliquely obovate, with widely curved lower edge and straight to slightly concave upper edge, obtuse at the apex, 8-9 x 3.54 mm; auricle ca. 1 mm long, claw 9 mm long. Staminal tube truncate at the mouth. Ovary on a stipe 3-4 mm long, linear, hairy; style glabrous. Other specimens seen: Afghanistan. Bamian: hill on west side of Shibar pass, 3000 m, 14.6.1962, Hedge & Wendelbo W.4198 (BG). — Parwan: Shibar-Pass, nahe der Passhöhe, 2900 m, 27.6.1967, Freitag 1201 (MSB) - east side of Shibar pass, c. 2800 m, 14.6.1962, Hedge & Wendelbo W.4247 (BG). Astragalus wakhanicus Podlech, spec. nov. (Dissitiflori). Holotype: Afghanistan, Prov. Badakhshan, Wakhan, oberes Baroghil Tal und Baroghil Pass, 73°22’ E, 36°53’ N, 3300-3800 m, 30.7.1971, O. Anders 7905 (MSB!; iso: S!). Differt ab A. viridis Bunge stipulis brevibus, 1,5-3 (nec 5-7) mm longis, foliis 4-5 (nec 5-9)-jugis, foliolis apice acuto, rubri-brunneo, breviter cartilaginei-cucullato, axi racemi pilis 190 brevibus appressis albis nigrisque obtecto (nec pilis oblique patentibus ad patentibus albis, 0,3-1,2 mm longis obtecto), bracteis 2-3 (nec 4-6) mm longis, leguminibus pilis longissimis ad 1,5 mm (nec ad 3) mm longis obtectis. Plants suffruticose, 30-35 cm tall, vegetative parts covered with + symmetrically medi- fixed, appressed hairs. Caudex branched. Stems at the base ligneous, covered with dark greyish bark, branched, stems of the current year 6-17 cm long, very densely covered with white, near the nodes also with a few black hairs 0.5—0.7 mm long. Stipules narrowly triangular, 1.5-3 mm long, adnate to the petiole for 0.5-1.5 mm, white hairy, those of upper leaves sometimes also with black hairs. Leaves 4-7 cm long; petiole 0.3-0.5 cm long, like the rhachis slender, densely hairy like the stem. Leaflets in 4-5 pairs, linear to filiform, 10-20 x 0.8-1 mm, at the apex acute, with a short, reddish-brown, cartilaginous, cucullate tip, on both sides densely covered with white hairs 0.8-1 mm long. Peduncles 11-16 cm long, loosely to rather densely white hairy like the stem, toward the raceme also with some black hairs, sometimes partly glabrescent. Racemes (in fruiting state) loosely 4-12-flowered; axis subappressed hairy like the peduncle but with mixed black and white hairs. Bracts narrowly triangular to ovate- acuminate, 2-3 mm long, white and black hairy, at the margins also with basifixed hairs. Pedicels (in fruit) ca. 3 mm long, with ascending white and black hairs. Calyx ca. 15 mm long, tubular, rather densely covered with subappressed, + symmetrically to asymmetrically medi- fixed black hairs 0.3-0.7 mm long and with ascending, strongly asymmetrically medifixed to subbasifixed white hairs 1-1.5 mm long; teeth subulate, 3-3.5 mm long, densely white hairy on innerside. Petals unknown. Staminal tube truncate at the mouth. Legumes sessile, erect, oblong, ca. 20 mm long, 4-5 mm high and wide, carinate ventrally, slightly grooved dorsally, at the apex subabruptly attenuate into a beak 2-3 mm long, fully bilocular; valves thinly coriaceous, brownish to blackish when ripe, densely villous with ascending to spreading, basifixed white hairs 1-1.5 mm long and with short, asymmetrically medifixed, subappressed black hairs, at the dorsal side also with ascending, subbasifixed black hairs up to 1 mm long. Astragalus sherriffii Podlech, spec. nov. (Sect. Hemiphaca). Holotype: India, Kashmir, Gya. Ladah, 13.500 ft, 14.7.1941, F. Ludlow & G. Sherriff 8491 (E). Differt ab A. /horongensis P.C.Li & C.C.Ni stipulis inferioribus post tergum caulis distincte vaginanti-connatis nec omnibus inter se liberis), foliis 3-4 (nec 4-10) cm longis, 4-5 (nec 5—7)-jugis, lamina alarum apice retusa ad emarginata (nec rotundata). Plants ca. 25 cm tall, covered with appressed to subappressed hairs 0.2-0.3 mm long. Caudex up to 10 mm thick, with a pluricipital root-crown. Stems several, ascending to erect, very sparsely to sparsely furnished with white and some black hairs especially below the nodes. Stipules greenish, the lower ones ca. 3 mm long, ovate, free from petiole, distinctly vaginate-connate behind the stem, the upper ones narrowly triangular, up to 4 mm long, free from each other, all loosely white or white and black hairy. Leaves 3-4 cm long; petiole 0.8—2 cm long, like the rhachis slender, sparsely to loosely hairy like the stem. Leaflets in 4-5 pairs, linear, 7-12 x 1.5—3 mm, obtuse at the apex, on upper side glabrous, on underside sparsely to loosely appressed white hairy. Peduncles 3—7 cm long, sparsely to loosely, toward the raceme more densely hairy like the stem. Racemes 2-3 cm long, loosely many-flowered. Bracts 191 whitish-membranous, narrowly triangular, 1-1.5 mm long, predominantly black hairy. Pedi- cels ca. 0.5 mm long, predominantly black hairy; flowers + spreading. Calyx campanulate, ca. 3 mm long, obliquely cut at the mouth, rather densely covered with predominantly black hairs; teeth unequal, the upper two triangular, 0.5—1 mm long, the lower three narrowly triangular, 1—1.5 mm long, hairy on innerside. Petals flushed with pink. Standard ca. 7 mm long; blade elliptic to widely elliptic, 4-5 mm wide, incised at the apex, narrowed at the base without a distinct claw. Wings 5.5 mm long; blades narrowly oblong, retuse to emarginate at the apex, 4 x 1.5 mm; auricle 0.5 mm long, claw 2 mm long. Keel 4 mm long; blades obliquely obovate- triangular, with rectangular-curved lower edge and straight to slightly concave upper edge, subobtuse at the apex, 2.5 x 1.5 mm; auricle indistinct, claw 1.5 mm long. Staminal tube trun- cate at the mouth. Ovary shortly stipitate, glabrous; style thick, short. Legumes unknown. Astragalus tsangpoensis Podlech & L.R.Xu, spec. nov. (sect. Hemiphaca). Holotypus: China, Xizang, SE. Tibet, Tsangpo valley, c. 5 km W of Gonggar (Lhasa Airport), 29°17" N, 90°49’ E, 3620 m, 30.7.1989, B. Dickoré 3064 (MSB). Plantae ultra 30 cm altae, pilis albis nigrisque subappressis ad adscendentibus 0,3-0,7 mm longis instructae. Caules ramosi. Stipulae triangulares, 3-5 mm longae, liberae. Folia 3-7 cm longa. Foliola 8-11 juga, 4-9 x 1-2,5 mm, acuta, complicata, supra glabra, subtus laxe ad densiuscule appresse pilosa. Pedunculi 5-6 cm longi, sparse ad laxe pilosi. Racemi 1,5-2,5 cm longi, laxe 6-10-flori. Bracteae 1,5-2 mm logae. Calyx campanulatus, c. 3 mm longus, albi- vel albi-nigri-pilosus, dentibus 1 mm longis. Vexillum c. 7 mm longum, orbiculare, 6 mm latum, brevissime unguiculatum. Alae 6,5 mm longae, lamina apice rotundata, 4,5 x 2,5 mm. Carina c. 5 mm longa. Ovarium sesssile, albi-pilosum. Legumina ignota. Plants at least 30 cm tall (basal parts missing), covered with subappressed to tangled- ascending hairs 0.3-0.7 mm long. Stem ascending to erect, branched, 2-3 mm in diameter, glabrous in basal parts, sparsely to loosely white and black hairy in upper part. Stipules greenish, 3-5 mm long, triangular, sparsely white and black hairy, free from petiole and from each other. Leaves 3-7 cm long; petiole 0.5— 1.5 cm long, like the rhachis loosely to rather densely covered with white and sometimes with fewer black hairs. Leaflets in 8-11 pairs, narrowly ovate to narrowly elliptic, acute at the apex, 4-9 x 1-2.5 mm, glabrous on upper side, loosely to rather densely subappressed white hairy on underside, mostly folded. Peduncles 5-6 cm long, hairy like the stem. Racemes 1.5-2.5 cm long, loosely 6-10-flowered. Bracts whitish-membranous to slightly greenish, 1.5-2 mm long, narrowly triangular, sparsely white and black hairy, at the margins with few sessile, minute glands. Pedicels 0.5 mm long, black and white hairy. Calyx campanulate, slightly obliquely cut at the mouth, ca. 3 mm long, loosely to rather densely covered with white or white and black hairs; teeth linear, ca. 1 mm long, hairy on inner side. Petals (in dry state) whitish yellow or very slightly flushed pale bluish. Standard ca. 7 mm long; blade orbicular, ca. 6 mm long, widely emarginate at the apex, at the base abruptly constricted into the very short claw. Wings ca. 6.5 mm long; blades elliptic, rounded at the apex, 4.5 x 2.5 mm; auricle ca. 0.5 mm long, claw ca. 2 mm long. Keel ca. 5 mm long; blades obliquely elliptic-triangular, with in the middle widely curved lower edge and slightly convex upper edge, obtusish to acute at the apex, 3.5 x 2 mm; auricle minute, claw ca. 2 mm long. Staminal tube slightly oblique at the mouth. Ovary sessile, ellipsoid, white hairy; style glabrous. Legumes unknown. 192 Astragalus olurensis Podlech, spec. nov. (sect. Incani). Holotype: Turkey, Prov. Coruh (Artvin), between Olur and Yusufeli, towards Ishan, 870 m, 25.5.1990, M. Nydegger 45536 (MSB; isotype: BASBG). Differt ab A. /atifolio Lam. statura humiliore, calyce breviore, tubo ejus intus glabro, petalis violaceis distincte brevioribus, forma leguminis diversissima, leguminibus 18-20 mm (nec 40-60--80) mm longis, ca. 4 mm (nec 2,5 mm) altis. Plants 5-15 cm tall, covered with appressed, mostly symmetrically medifixed, merely white hairs 0.3-0.5 mm long. Caudex up to 10 mm in diameter, unbranched or slightly and shortly branched, covered in upper part with remnants of old leaves. Stipules narrowly tri- angular, ca. 5 mm long, adnate to the petiole for ca. 2 mm, densely hairy, sometimes glabrescent. Leaves 4-7 cm long; petiole 1.5-4.5 cm long, densely hairy. Leaflets only 1 or 1-6 paired, often with quite different number of leaflets in one plant, ovate to widely elliptic, at the base widely cuneate to truncate, at the apex acute to acuminate or widely rounded, mostly minutely mucronulate, if only one leaflet, this 10-25 x 6-21 mm, if more leaflets, than those 3-7 x 2-5 mm but end-leaflet mostly larger, on both sides loosely to rather densely hairy and mostly spotted with minute reddish dots. Peduncle erect or ascending, 4-9 cm long, loosely to rather densely hairy. Raceme loosely 6-8-flowered. Bracts membranous, narrowly triangular, 1-2.5 mm long, sparsely hairy, at the margins near the base with minute, globular, sesssile glands. Pedicels erect, 1 mm long, white hairy. Bracteoles linear, 0.5—1 mm long, at the base of the calyx. Calyx 11-13 mm long, tubular, obliquely cut at the mouth, loosely to rather densely hairy, inner side of tube glabrous; teeth subulate, 5-6 mm long, on inner side densely hairy. Corolla violett. Standard 17-18 mm long; blade slightly upcurved, 7 mm wide, elliptic, at the apex emarginate, at the base gradually narrowed into the short, cuneate claw. Wings 13-14 mm long; blades narrowly oblong, gibbous above the auricle, at the apex rounded, laterally obliquely slightly emarginate, 7 x 2 mm; auricle | mm long, claw 7 mm long. Keel ca. 11 mm long; blades obliquely obovate, with widely curved lower edge and slightly concave upper edge, at the apex subacute, 5 x 2.5 mm; auricle ca. 0.5 mm long, claw 7 mm long. Stami- nal tube truncate at the mouth. Ovary linear, sessile, white hairy; style glabrous. Legumes sessile, erect linear, slightly to distinctly upcurved, 18-20 mm long, ca. 4 mm high, 3 mm wide, carinate ventrally, slightly grooved dorsally, tapering into a straight beak ca. 3 mm long, bilocular; valves coriaceous, dirty pale brownish, loosely to rather densely hairy. Astragalus zaraensis Podlech, spec. nov. (sect. Incani). Holotype: Turkey, Prov. Sivas, [B7] N of Karabel, at road from Divrigi to Zara, 1560 m, 18.6.1992, M. Nydegger 46325 (MSB; isotype: BASBG). Differt ab A. ancistrocarpo Boiss. & Hausskn. calyce breviore, 7-8 (nec 11-14) mm longo, dentibus ejus 0,5-1,5 (nec 2-4 mm longis, petalis distincte brevioribus, vexillo apicem versus non lingulato-angustato, laminis alarum anguste oblongis, ca. 2 mm latis (nec apicem versus distincte dilatatis, 3,8-4 mm latis), leguminibus erectis 16-20 mm longis (nec saepissime pendulis vel patentibus, 20-35-40 mm longis). Plants acaulescent, ca. 12-16 cm tall, covered in vegetative parts with appressed, symmetrically medifixed white hairs 0.4-0.5 mm long. Caudex simple or shortly branched, 193 covered with remnants of old leaves. Stipules narrowly triangular to narrowly oblong, 6-8 mm long, adnate to the petiole for 1.5-2 mm, rather densely hairy. Leaves 5-11 cm long; petiole 2-4 cm long, like the rhachis loosely to rather densely hairy. Leaflets in 5-11 pairs, elliptic to obovate, 3-8 x 2-5 mm, at the apex obtuse to retuse, on both sides densely hairy when young, loosely hairy with age, on upper side sometimes spotted with minute pellucid to reddish dots. Peduncle 7-10 cm long, loosely white hairy. Raceme at anthesis rather densely 10-20-flowered, strongly elongating in fruit; axis hairy like the peduncle. Bracts whitish- to brownish-membranous, narrowly triangular, 1.5—3 mm long, sparsely white and black hairy, at the margins with minute, sessile, globular glands. Pedicels ca. 2 mm long, black and white hairy; flowers erect. Bracteoles minute, 0.3— 1 mm long, whitish, at the base of calyx or just below the calyx at the pedicel. Calyx 8-9 mm long, tubular, gibbous at the base, obliquely cut at the mouth, loosely covered with appressed white and black hairs 0.2-0.4 mm long and with somewhat longer white hairs; teeth narrowly triangular to linear, 0.5-1.5 mm long, hairy on inner side. Petals violet. Standard ca. 20-21 mm long; blade distinctly upcurved above the middle, ca. 6-6.5 mm wide, narrowly elliptic, not attenuate towards the slightly retuse apex, subabruptly narrowed at the base into the rather long claw. Wings ca. 15 mm long; blades narrowly oblong, obliquely laterally emarginate at the apex, 7-8 x 2 mm; auricle ca. | mm long, claw ca. 8 mm long. Keel 13 mm long; blades obliquely obovate, with widely curved lower edge and sigmoid-concave upper edge, at the apex obtuse to subacute, 5 x 2.5 mm; auricle 0.5 mm long, obtuse, claw ca. 7 mm long. Staminal tube truncate at the mouth. Ovary subsessile, linear, sparsely appressed hairy; style glabrous. Legume sessile, erect, linear, distinctly upcurved to sickle-shaped, 16-20 mm long, ca. 2 mm high and 2.5 mm wide, slightly carinate ventrally, narrowly grooved dorsally, attenuate at the apex into a slender, + straight beak 3-4 mm long, fully bilocular; valves thinly coriaceous, pale brownish, distinctly purplish mottled, loosely covered with appressed white hairs 0.5-0.6 mm long, sometimes glabrescent. Other specimen seen: Turkey. Sivas: [B6] Bestepeler Gegidi at road from Kayseri to Sivas, 1400 m, 12.6.1992, Nydegger 46236 (MSB). Astragalus damxungensis Podlech & L.R.Xu, spec. nov. (sect. Komaroviella). Holotype: China, Prov. Xizang, Nyaingentanglha Shan, between Yangbajain and Damxung, valley SE of Nyaingentanglha Feng, 5060 m, 30°19’ N, 90°35’ E, 12.8.1989, B. Dickoré 3887 (MSB). Plantae fere acaules, humiles, 3-4 cm altae, subglabrae, pilis perpaucis nigris 0,2-0,3 mm longis instructae. Stipula 3-4 mm longae, a petiolo liberae, inter se distincte vaginanti- connatae. Folia 4-6 juga, foliolis 3-5 x 1-2,8 mm. Racemi pedunculo 0,5—1 cm longo suffulti, 1-2-flori. Calyx campanulatus, ca. 6 mm longus, dentibus 2,5-3 mm longis. Petala purpurei- violacea. Vexillum 12-13 mm longum, lamina ca. 10 mm latum, suborbiculare. Alae 10 mm longae. Carina 11 mm longa. Legumina ignota. Plants 3-4 cm tall, nearly acaulescent, in vegetative parts subglabrous, with few sub- appressed, black hairs 0.2-0.3 mm long. Caudex with short stolones. Stems inconspicuous, glabrous. Stipules greenish to membranous, 3-4 mm long, narrowly ovate to oblong, with 194 obtusish tips, nearly free from the petiole, behind the stem distinctly to highly up vaginate- connate, glabrous. Leaves 1.5-2.5 cm long; petiole 0.4-0.7 cm long, like the rhachis glabrous or with scattered hairs. Leaflets in 4-6 pairs, elliptic, 3-5 x 1-2.8 mm, at the apex obtuse to acutish, glabrous or more rarely with scattered hairs on underside, glabrous on upper side, conspicously and densely rugulose when dry on underside. Peduncle 0.5—1 cm long, glabrous or below the raceme with scattered black hairs. Raceme 1-2-flowered. Bracts whitish- membranous, ca. 3 mm long, narrowly ovate, glabrous. Pedicels ca. 2 mm long, rather densely hairy. Bracteoles present, linear-subulate, 1-2 mm long, whitish, sparsely hairy, at the base of calyx. Calyx campanulate, ca. 6 mm long, rather densely hairy; teeth linear, slightly unequal, 2.5—3 mm long, hairy on innerside. Petals purple-violet. Standard 12-13 mm long; blade ca. 10 mm wide, widely obovate-orbicular, at the apex emarginate, at the base gradually narrowed into a short, cuneate claw. Wings ca. 10 mm long; blades narrowly obovate, rounded at the apex, 7x 3 mm; auricle ca. 1 mm long, claw 3 mm long. Keel 11 mm long; blades obliquely obovate, with in upper part widely curved lower edge and + straight upper edge, at the apex obtuse, 8x 4 mm; auricle ca. 0.5 mm long, claw 3.5 mm long. Staminal tube truncate at the mouth. Ovary with a stipe ca. 1.5 mm long, glabrous. Pods unknown. Astragalus rasmontii Podlech, spec. nov. (sect. Onobrychoidei). Holotype: Turkey, Prov. Sivas, Yazyurdu, 1760 m, 38°48’14’’ N, 36°56’07’’ E, 11.7. 1995, P. Rasmont PR105 (MSB!; iso: LG). Plantae ca. 50 cm altae, pilis bifurcatis albis nigrisque 0,1-0,3 mm longis obtectae. Stipulae 2-4 mm longae, inferiores vaginanti-connatae, superiores inter se liberae. Folia 7-9 cm longa, 10-13 juga, foliolis anguste ellipticis, 6-13 x 1-2,5 mm, supra glabris, subtus sparse ad laxius- cule albi-pilosis. Racemi pedunculo 6-9 cm longo laxiuscule piloso suffulti, 2,5-4,5 cm longi, dense multiflori. Bracteae ovatae, 1,5-2 mm longae. Calyx ca. 4 mm longus, campanulatus, albi-nigri-pilosus, dentibus 1—1,5 mm longis. Petala violacea. Vexillum 14-15 mm longum, 5-6 mm latum, ovatum, apicem versus gradatim angustatum. Alae 9-10 mm longae. Carina 7-7,5 mm longa. Legumina ca. 5 mm longa, bilocularia, pilis albis arcte asymmetrice medifixis ca. 0,5 mm longis obtecta. Plants ca. 50 cm tall, covered with appressed bifurcate hairs 0.1-0.3 mm long. Stem up to 45 cm long, branched, sparsely white and black hairy, partly glabrescent. Stipules 2-4 mm long, the lower ones membranous, very shortly adnate to the petiole, behind the stem distinctly vaginate-connate, subglabrous, the upper ones often greenish, free from the petiole, mostly clasping the stem partly with a narrow rim but not connate, sparsely hairy at the margins. Leaves 7-9 cm long, subsessile, those of side-branches shorter; rhachis sparsely to loosely white and black hairy. Leaflets in 10-13 pairs, narrowly elliptic, 6-13 x 1-2.5 mm, obtuse at the apex, on upper side glabrous, on underside sparsely to rather loosely white hairy. Peduncles 6-9 cm long, rather loosely white and black hairy. Racemes ovoid to shortly cylindric, 2.5-4.5 cm long, densely many-flowered. Bracts membranous, 1.5—2 mm long, ovate, sparsely hairy. Pedicels ca. 0.5 mm long, predominantly black hairy. Calyx campa- nulate, ca. 4 mm long, loosely to rather densely covered with black hairs and with somewhat longer white hairs up to 0.5 mm long; teeth narrowly triangular, 1-1.5 mm long, hairy on innerside. Petals deeply violet. Standard 14-15 mm long; blade 5-6 mm wide, ovate, gradually 195 narrowed toward the minutely emarginate apex, at the base subabruptly narrowed into the claw ca. 3 mm long. Wings 9-10 mm long; blades narrowly oblong, slightly dilated toward the slightly and obliquely retuse apex, ca. 6 x 2.5 mm; auricle ca. 0.6 mm long, claw 3.54 mm long. Keel 7-7.5 mm long; blades obliquely elliptic, with in the middle widely curved lower edge and slightly curved upper edge, obtuse at the apex, 3.5 x 2.5 mm; auricle ca. 0.5 mm long, claw 3.54 mm long. Staminal tube oblique at the mouth. Ovary subsesssile, ellipsoidic, white hairy; style glabrous. Legumes (immature) subsessile, elliptic, ca. 5 mm long, 2 mm high, compressed laterally, carinate ventrally, grooved dorsally, at the apex with a short oblique beak, nearly completely bilocular; valves thin, rather densely covered with strongly asymmetrically medifixed white hairs ca. 0.5 mm long. Seeds two in each locule, brown. Astragalus austrotibetanus Podlech & L.R.Xu, spec. nov. (sect. Polycladus). Holotypus: China, Xizang, SE. Tibet, Lhasa valley, c. 30 km SW of Lhasa, 29°26’ N, 90°55’ E, 3650 m, 30.7.1989, B. Dickoré 3093 (MSB; iso: hb. Dickore). A. leucocephalo Benth. similis sed differt pilis albis nigrisque (nec mere albis) instructus, caulibus pilis subappressis ad leviter ascendentibus (nec patentibus) obtectis, racemis oblongis, demum elongatis et laxiusculis (nec globosis ad ovoideis, densis), bracteis 3-4 (nec 5-6) mm longis, petalis caerulei-violaceis (nec pallide flavis), vexillo latiore, leguminibus curvatis (nec rectis). Plants up to 30 cm tall, in vegetative parts covered with subappressed to ascending, often flexuous, tangled white hairs 0.5-1.5 mm long. Caudex up to 8 mm thick, branched. Stems several, erect or at the base slightly ascending, sometimes branched, 10-20 cm long, angular- sulcate, loosely to rather densely hairy. Stipules green, narrowly triangular, 6-9 mm long, free from petiole, the lower ones behind the stem distinctly connate, the upper ones free from each other, loosely hairy. Leaves 4-8 cm long, nearly sessile; petiole up to 1 cm long, like the rhachis grooved on upper side, loosely to rather densely hairy. Leaflets in 12-15 pairs, elliptic, acute to obtuse, 5-14 x 1.5-7 mm, on upper side glabrous or subglabrous to sparsely hairy, on underside loosely to rather densely hairy. Peduncles 1-3 in the axils of upper leaves, 2-7 cm long, hairy like the stem but mostly also with black hairs toward the raceme. Racemes shortly cylindric, 2-5 cm long, rather loosely many-flowered, later on slightly elongated. Bracts membraneous, narrowly triangular, 3-4 mm long, predominantly white but also black hairy, at the margins with few minute glands. Pedicels ca. 1 mm long, predominantly white hairy, recurved, flowers therefore pendant. Calyx campanulate, 4-5 mm long, slightly obliquely cut at the mouth, rather densely covered with subsppressed to ascending white hairs 0.5-]1 mm long and with mostly somewhat shorter black hairs; teeth slightly unequal, narrowly triangular, 1.5-3(-3.5) mm long, hairy also on inner side. Petals blue-violet or more rarely white. Standard 7-8(-9) mm long; blade widely elliptic to suborbicular, 5-6(-7) mm wide, at the apex slightly to deeply emarginate, at the base shortly gradually narrowed without distinct claw. Wings 5.5-7(-9) mm long; blades narrowly elliptic, obtuse at the apex, 4-4.5(-6) x 2-2.5 mm; auricle ca. 0.5 mm long, claw 1.5-2.5(-3.5) mm long. Keel 5-6(-7) mm long; blades obliquely elliptic-triangular, with in the middle + rectangular-curved lower edge and slightly curved upper edge, obtuse at the apex, 3-3.5(-4) x 2-2.5 mm; auricle minute, claw 2-3 mm. Staminal tube oblique at the mouth. Ovary subsessile, elliptic, hairy; style 196 short, glabrous. Pods subsessile, obliquely elliptic, curved, ca. 6-7 mm long, 2-2.5 mm high and wide, carinate ventrally, flattened to shallowly grooved dorsally, narrowed toward the acute apex, often with persistent, hooked style, unilocular; valves rather densely covered with ascending white or white and black hairs up to 0.8 mm long. Seeds two, 3 x 1.5 mm, blackish. Other specimens seen: China. Qinghai: NE. Tibet, along Golmud - Lhasa highway, NE of Tanggulashanqu, 4580 m, 34°18’ N, 92°34’ E, 4.8.1993, G. & S. Miehe 9413/15 (MSB, Hb. Miehe). — Xizang: Dochen Lake, c. 4600 m, 24.6.1939, Gould 2192 (K) - Tingri, 4580-4890 m, 4.7.1974, Hingston 201, 213 (K) — Rama, 4870 m, 29.6.1939, Gould 2209 (K) — Samoda, 4280 m, 9.8.1936, Chapman 507 (K) — Nalu La to Champitang, 3660-4280 m, 1.8.1936, Chapman 1037 (K) — Gooring valley, 90°25’ E, 30°12’ N, 5050 m, vii.-vili.1895, Littledale (K) — Gyantse, 4020 m, 19.8.1935, Cutting & Vernay 43 (K) - SE. Tibet, Lhasa valley, Lhasa, W. City, 3660 m, 1.8.1989, B.Dickore 3134 (MSB, hb. Dickoré) — Lhasa, 21.5.1943, Ludlow & Sherriff 9514 (E) — dto., 6.7.1943, Lud- low & Sherriff 9739 (E) — dto., ix.1904, Waddell (E) — Tsangpo Valley, Gonggar (Lhasa Airport), 3620 m, 29°17’ N, 90°50’ E, 30.7.1989, Dickore 3025 (MSB, Hb. Dickoré) — ca. 10 km NNE of Lhasa, 4350 m, 2.8.1989, Dickoré 3284, 3299 (MSB, hb. Dickoré) — Lhasa valley, c. 30 km SW of Lhasa, 29°26’ N, 90°55’ E, 3650 m, 30.7.1989, Dickoré 3093 (MSB) — Lhasa, above Chup- sang Monastery, 4300-4400 m, 29°41” N, 91°08’ E, 9.8.1997, Miehe, Huang, Otsu & Tunsu 97- 008-15 (MSB, Hb. Miehe) — dto., 4050 m, 13.8.1997, Miehe, Huang, Otsu & Tunsu 97-014-02 (MSB) — Upper Kyi Chu, above Reting Monastery, 4300 m, 30°18’ N, 91°31’ E, 22.8.1997, Mie- he, Huang, Otsu & Tunsu 97-027-01 (MSB) — SE Tibet, upper Kyi Chu basin, S of Damxung, 4550 m, 30°33’ N, 91°27’ E, 7.9.1995, G. & S. Miehe 95-37-05 (MSB) — Mekong - Salween divide, Salween tributary, Bamda - Nujiang, NE of pass, 4450 m, 30°10’ N, 97°17’ E, 5.7.1994, Dickoré 8896 (MSB, Hb. Dickoré) — Mekong - Salween divide, Salween tributary, right flanc of Yü Qu N of Bamda, 4130 m, 30°20’ N, 97°15’ E, 7.7.1994, Miehe & Wiindisch 94-52-1 (MSB, Hb. Miehe) — Mekong - Salween divide, Mekong tributary, Bamda - Qamdo, Zi Qu tributary SW of Gyitang, 3450 m, 30°42’ N, 97°19’ E, 7.7.1994, Dickoré 9012 (MSB, hb. Dickoré) — Upper Gyamda Chu (Nyang Chu), W of Gongbogyamda, 4700-4850 m, 29°52’ N, 92°17’ E, 2.9.1994, Miehe & Wiindisch 94-256-1 (MSB, Hb. Miehe) — EC. Tibet, Yamco Yumco (Yamdrok Yutsho), above Dzamtschili, 4570 m, 28°59’ N, 90°27’ E, 5.9.1997, Miehe, Huang, Otsu & Tunsu 97- 057-25 (MSB, Hb. Miehe) — dto., 4450 m, 12.9.1995, G. & S. Miehe 95-52-05 (MSB) — Yamco Yumco, above Shangding Gompa, 4570 m, 28°58’ N, 90°29’ E, 4.9.1997, Miehe, Huang, Otsu & Tunsu 97-054-17 (MSB, Hb. Miehe) — Yamco Yumco, (Yamdrok Yutsho), above Baidi, 4500 m, 29°07’ N, 90°26’ E, Miehe, Huang, Otsu & Tunsu 97-063-21 (MSB, hb. Miehe) — Lhasa - Nagarze, Yamdruk Yumco, Zamu E of Nagarze, 4520 m, 28°59’ N, 90°32’ E, 24.7.1994, Dickore 9909 (MSB) — upper Subansiri, Lhünze - Qayü, 15 km NW of Qayü, 3800 m, 28°23’ N, 92°42’ E, 31.7.1994, Dickoré 10268 (MSB) — Chayul Dzong, Loro Chu, Ludlow & Sherriff 1326 (W). Astragalus cobresiiphila Podlech & L.R.Xu, spec. nov. (sect. Polycladus). Holotypus: China, Xizang, valley ca. 8 km SSE of Lhasa, 3950 m, B. Dickoré 3384 (MSB; iso: hb. Dickoré). Differt ab A. stricto Benth. foliolis 5-8 (nec 8-12)-jugis, 14 mm (nec 4-10 mm) longis, vexillo 6-7 mm (nec 7-9 mm) longo, leguminibus pilis ad 0,2 mm longis (nec 0,3-0,7 mm longis) obtectis. Plants up to 12 cm tall, caespitose, furnished with appressed to subappressed hairs. Caudex with a pluricipital root-crown. Stems numerous, diffusely prostrate to ascending, up to 5 cm long, slender, angular-sulcate, loosely furnished with appressed, somewhat flattened hairs 0.1-0.2 mm long, partly glabrescent to glabrous, especially in lower part. Stipules 197 greenish or membranous, 2-3.5 mm long, ovate-triangular, free from the petiole, distinctly connate behind the stem, sparsely hairy to sometimes glabrous. Leaves 1-3.5 cm long; petiole 0.3-1.2 cm long, like the rhachis slender, somewhat flattened, sparsely to loosely hairy like the stem. Leaflets in 5-8(-9) pairs, elliptic, 14 x 0.6-2 mm, at the apex rounded to emarginate, on upper side glabrous, on underside especially at the midvein sparsely to loosely covered with appressed hairs up to 0.3 mm long. Peduncles several in upper leaf axils, 1.5—S cm long, slender, sparsely to loosely hairy like the stem but with increasing black hairs toward the raceme. Racemes short, densely 3—10-flowered. Bracts whitish-membranous, ovate to narrowly triangular, 1—1.5 mm long, white and black hairy at the margins often with short stalked minute glands. Pedicels 0.5—1 mm long, black hairy. Calyx campanulate, (2.5—)3—4 mm long, slightly obliquely cut at the mouth, loosely covered with appressed black hairs 0.1-0.3 mm long and more rarely also with some white hairs; teeth unequal, the lower three narrowly triangular, 0.5—1.5 mm long, the other two triangular, 0.3—1 mm long. Petals blue to purple. Standard 6—7 mm long; blade widely elliptic to orbicular, ca. 5 mm wide, deeply emarginate at the apex, at the base gradually narrowed without distinct claw. Wings ca. 5.5 mm long; blades narrowly elliptic, obtuse at the apex, 3.5 x 1.8-2 mm; auricle 0.5 mm long, claw 2 mm long. Keel 5 mm long; blades obliquely elliptic, with widely curved lower edge and sigmoid upper edge, subacute at the apex, 3 x 1.8-2 mm; auricle minute, claw 2 mm long. Staminal tube slightly oblique at the mouth. Ovary with a stipe ca. 1 mm long, white hairy; style glabrous. Legumes subsessile or with a stipe ca. | mm long, pendulous, narrowly elliptic, curved, ca. 6 mm long, 2 mm high and wide, carinate ventrally, grooved dorsally, acute at the apex with the hooked persistent style, unilocular; valves thin, pale brownish, loosely covered with appressed black hairs 0.1-0.2 mm long, soon glabrescent. Seeds 3-4, 2 x 1.2 mm, blackish. Other specimens seen: China. Qinghai: Qinghai Nan Shan, S of Qinghai Hu, E of Jiangxigou, 3460 m, 36°33’ N, 100°28’ E, 26.7.1993, G. & S. Miehe 9316/28 (MSB, Hb. Miehe) — Qilian Shan, Riyue Shan, Lyewa Pass, SE of Qinghai Hu, 3350 m, 36°25’ N, 101°05’ E, 26.7.1993, G. & S. Miehe 9313/10 (MSB, Hb. Miehe) — Tangula Shan N, upper Yangtse basin, Bi Qu, Tangula Pass - Yanshiping, Tangula Station, 4890 m, 33°15’ N, 91°51’ E, 3.9.1989, Dickoré 4637 (MSB, Hb. Dickoré) - Tangula Shan N, upper Yangtse basin, Gar Qu, Mt. Geladadong, NE glacier tongue, 5000 m, 33°34’ N, 91°27’ E, 30.8.1989, Dickore 4545 (MSB, Hb. Dickore). — Xinjiang: Karakorum, Aghil Shan, northern declivity, Aghil valley, ca. 19 km SW of Ylik, 4160 m, 36°15’ N, 76°33’ E, 30.8.1986, Dickoré 479 (MSB, Hb. Dickoré) — Aghila Shan, ca. 7 km NW of Aghil Pass, 4200 m, 36°14’ N, 76°34’ E, 30.8.1986, Dickoré 486 (MSB, Hb. Dickoré). — Xizang: Upper Tsangpo basin, Ngamring — Sangsang, 4620 m, 29°19’ N, 86,53’ E, 20.8.1993, G. & S. Miehe 9518/07 (MSB, Hb. Miehe) — EC. Tibet, Central Plateau, Nagqu - Siling Co, 4620 m, 31°24’ N, 90°59’ E, 12.8.1993, G. & S. Miehe 9469/02 (MSB, Hb. Miehe) — Central Plateau, E of Siling Co, 4600 m, 31°42’ N, 89°36’ E, 13.8.1993, G. & S. Miehe 9478/06 (MSB, Hb. Miehe) — Lhasa, 3670 m, 21.5.1943, Ludlow & Sherriff 9515 (E) - dto., 3970 m, 25.6.1943, Ludlow & Sherriff 9715 (E) - Lhasa, 5200 m, 6.9.1904, Waddell (K) — Nyaingentanghla Shan, Yangbajain - Damxung NW. of Lhasa, SE. of Nyainquent, 4900 m, 30°17’ N, 90°37’ E, 9.8.1989, Dickore 3538, 3569 (MSB, hb. Dickoré) — dto., 5060 m, Dickore 3786 (MSB, Hb. Dickoré) — 10 km NNE of Lhasa, 4280 m, 29°45’ N, 91°09’ E, 2.8.1989, Dickoré 3249 (MSB, Hb. Dickoré) — Nagarze - Lhozak, Pomo Co (C7), 5010 m, 28°30’ N, 90°33’ E, 18.7.1994, Miehe & Wündisch 94-76-10 (MSB, Hb. Miehe) — SE Tibet, upper Yi’ong Zangbo, above Lhari, 4530 m, 31°39’ N, 93°12’ E, 2.9.1995, G. & S. Miehe 95-07-03 (MSB, Hb. Miehe). 198 Astragalus conaensis Podlech & L.R.Xu, spec. nov. (Sect. Polycladus) Holotype: China, Xizang, Tibetan Himalaya NE of Bhutan, N of Cona, 28°07’ N, 91°54’ E, 4720 m, 30.7.1994, B. Dickoré 10235 (MSB; iso Hb. Dickoré). Differt ab A. conferto cui habitu similis foliolis 3-7 (nec 5-9)-jugis, pedunculis ad 1,5 (nec ad 4) cm long, calyce pilis nigris 0,1-0,3 mm longis et pilis albis 0,6-1 mm longis (nec imprimis pilis nigris 0,1-0,3 mm tantum longis) obtecto, vexillo ca. 9 (nec 10-12) mm longo, lamina carinae 4,5 x 2,5 mm (nec 6-7 x 3 mm). Plants acaulescent to subacaulescent, densely caespitose, 2-5 cm tall, furnished with sub- appressed, in vegetative parts white, in inflorescence also black hairs 0.4-0.5 mm long. Caudex up to 10 mm in diameter, repeatedly divided with short thick, subterranean branches. Stems or basal leaf-rosettes numerous, stems if present up to 0.5 cm long, angular-sulcate, loosely hairy. Stipules whitish-membranous, 3-4 mm long, ovate-triangular, free from the petiole or shortly adnate, but distinctly connate before the petiole, behind the stem connate up to the middle, rather densely hairy. Leaves 1-2 cm long; petiole 0.5-0.8 cm long, like the rhachis slender, rather densely hairy. Leaflets in 3-7 pairs, narrowly elliptic to elliptic, 1-3 x 0.5-1.5 mm, obtuse at the apex, on upper side sparsely to loosely, on underside rather densely to densely hairy, often somewhat folded. Peduncles 0.3-1.5 cm long, white and black hairy. Racemes short, 2-5-flowered. Bracts whitish-membranous, narrowly triangular, 1.5—2.5 mm long, predominantly black hairy, at the base often with minute, sessile glands. Pedicels ca. 1 mm long, densely black hairy. Calyx narrowly campanulate, 4-5 mm long, slightly obliquely cut at the mouth, rather densely to densely covered with appressed, black hairs 0.1-0.3 mm long and with subappressed white hairs 0.6-1 mm long; teeth subulate, 2-2.5 mm long, densely hairy on innerside. Petals deeply violet. Standard ca. 9 mm long; blade widely elliptic to suborbicular, ca. 6 mm wide, retuse at the apex, at the base gradually narrowed, without distinct claw. Wings ca. 8 mm long; blades narrowly oblong, apex obtuse to rounded, ca. 5.5 x 1.8-2 mm; auricle ca. 1 mm long, claw 3.5 mm long, slightly curved. Keel ca. 8 mm long; blades obliquely elliptic, with in the middle widely to rectangular-curved lower edge and + straight to slightly concave upper edge, obtuse at the apex, ca. 4.5 x 2.5 mm; auricle ca. 0.8 mm long, claw ca. 3.5 mm long. Staminal tube slightly oblique at the mouth. Ovary with a slender stipe ca. 1 mm long, narrowly elliptic, white hairy; style glabrous. Legumes unknown. Other specimen seen: China. Xizang: Tibetan Himalaya NE of Bhutan, N of Cona, 28°07’ N, 91°54’ E, 4610 m, 30.7.1994, G. Miehe & Wiindisch 94-123-12 (MSB) — N slope Shisha Pangma [Xixabangma], S. Tibet, 5100-5200 m, 10.9.1984, Miehe 1515 (BM) — dto., 5530 m, 13.9.1984, Miehe 1533 (BM) - up from Nyalam, 4300 m, 3.9.1984, Miehe 1403 (BM). Astragalus tibeticola Podlech, spec. nov. (sect. Polycladus). Holotype: China, Prov. Xizang: S. Tibet, Chayul Dzong, Loro Chu, 23.4.1936, F. Ludlow & G. Sherriff 1326 (BM). Plants 15-25 cm tall. Stems 10-17 cm long, branched at the base, ascending to erect, striate-sulcate, sparsely to loosely covered with appressed to subappressed white, at the nodes also black hairs 0.4-0.7 mm long. Stipules green, 6-9 mm long, ovate-acuminate to 199 narrowly triangular, free from the petiole, behind the stem high up vaginate-connate, sparsely to loosely covered with appressed white and sometimes also with some black hairs. Leaves 6-12 cm long; petiole 1-3 cm long, like the rhachis loosely covered with appressed white hairs 0.2-0.5 mm long. Leaflets in 9-11 pairs, narrowly elliptic to elliptic, 7-17 x 3-6 mm, rounded to slightly emarginate at the apex, glabrous on upper side, sparsely and mainly at the midrib furnished with appressed white hairs 0.5-1 mm long. Peduncles 5-6 cm long, hairy like the stem but in upper part with predominantly black hairs. Racemes ovoid, rather loosely 7-15-flowered, Bracts whitish, linear-acute, 2.5-3 mm long, predominantly black hairs. Pedicels ca. 1 mm long, black hairy. Calyx campanulate, 4-5 mm long, slightly oblique at the base, slightly obliquely cut at the mouth, rather densely covered with appressed black hairs 0.2-0.3 mm long and with fewer, slightly longer white hairs; teeth subulate, 1.5-2 mm long. Petals mauve with whitish center, keel purplish-mauve. Standard ca. 6 mm long; blade 5 mm wide, transverse-elliptic, deeply incised at the apex, at the base subabruptly contracted into the very short claw. Wings ca. 6 mm long; blades oblong, rounded at the apex, 3.5 x 2 mm; auricle wide, ca. 0.5 mm long, claw 2.5 mm long. Keel 5 mm long; blades oblique-elliptic, rounded at the apex, 3 x 2 mm; claw 2 mm long. Staminal tube truncate et the mouth. Ovary sessile, white-hairy; style glabrous. Legumes unknown. Astragalus L. sect. Pseudotapinodes Podlech & L.R.Xu, sect. nov. Differt ab A. sect. Polyclado Y.C.Ho stipulis petiolo adnatis et post tergum caulis distincte connatis (nec a petiolo liberis), leguminibus quoad cognatis semibilocularibus (nec unilocularibus). Typus sectionis: A. orbicularifolius P.C.Li & C.C.Ni, Acta Phytotax. Sin. 17: 112. 1979. Holotypus: [China] Chongba, 5500 m, 9.8.1975, Chinghai-Tibetan Complex Exp. 6724 (PE). The following species belongs to this section up to now: A. dickorei Podlech & L.R.Xu, A. longiscapus C.C.Ni & P.C.Li, A. orbicularifolius P.C.Li & C.C.Ni. Astragalus dickorei Podlech & L.R.Xu, spec. nov. (Sect. Pseudotapinodes). Holotypus: China, Sichuan, Litang to Batang, Jinsha (Jangtse) - Yalong divide, Pass to Litang Plateau E of Yidun, 4800 m, 30°14’ N, 99°33’ E, 28.6.1994, B. Dickoré 8532 (MSB). Differt ab A. longiscapo foliis 0,8—1,5 (nec 1,5—9) cm longis, glabris vel subglabris (nec laxe ad densiuscule pilosis), bracteis ellipticis 3-4 mm longis (nec anguste triangularibus 2 mm longis, vexillo ca. 10 (nec 7) mm longo, carina alis multo breviore (nec ca. | mm tantum breviore). Plants acaulescent, 1.5-2 cm tall, subglabrous in vegetative parts. Caudex slender, some- times branched. Stems absent. Stipules whitish-membraneous, ovate, 3-4 mm long, widely rounded at the apex, adnate to the petiole for 1.5-2 mm, vaginate-connate behind the stem for 1-2 mm, toward the apex and at the margins with tousled, white and black hairs up to 0.3 mm long. Leaves 1-1.5 cm long, glabrous or leaflets with few white hairs up to 0.4 mm long; 200 petiole 0.4-0.6 mm long. Leaflets in 3-4 pairs, narrowly elliptic, 3-5 x 1-1.5 mm, obtuse at the apex. Peduncles inconspicuous, up to 0.3 cm long, densely covered with subappressed to slightly ascending, tousled black hairs ca. 0.3 mm long. Racemes short, densely 4-5-flowered. Bracts whitish-membranous, 3-4 mm long, elliptic, rather densely hairy like the peduncle. Pedicels 0.5—1 mm long, black hairy. Calyx campanulate, 3-4 mm long, rather densely covered with flexuous, tangled black hairs up to 0.3 mm long; teeth narrowly triangular to oblong, 1.5-2 mm long, black hairy on innerside. Petals blue-violet. Standard ca. 10 mm long, elliptic, ca. 6 mm wide, shortly incised at the apex, at the base subabruptly narrowed into the cuneate claw ca. 2 mm long. Wings ca. 9 mm long; blades narrowly obovate, rounded at the apex, 6-6.5 x 2.5 mm; auricle ca. 0.5 mm long, claw 2-2.5 mm long. Keel ca. 5 mm long; blades obliquely elliptic, with slightly but widely curved lower and upper edge, acutish at the apex, 3.5 x 2.2 mm; auricle short, claw 2 mm long. Staminal tube slightly oblique at the mouth. Ovary sessile, glabrous; style short, thick. Legumes unknown. X. New names and combinations in Astragalus Astragalus angorensis Podlech, nom. nov. — Typonym: A. brevidentatus Podlech, Sendtnera 6: 151. 1999, illeg. [non C.H.Wright 1906] Astragalus keredjensis Podlech, nom. nov. — Typonym: Astragalus triqueter Bornm. & Gauba, Repert. Spec. Nov. Regni Veg. 39: 106. 1935, illeg. [non A.Gray 1878]. Astragalus maowensis Podlech, & L.R.Xu, nom. nov. — Typonym: Astragalus aridicola P.C.Li, Acta Bot. Yunnanica 11(3): 295. 1989, illeg. [non Sosn. 1948]. Astragalus obtusifoliolus (S.B.Ho) Podlech & L.-R.Xu, comb. nov. Basion.: A. nobilis B.Fedtsch. var. obtusifoliolus S.B.Ho, Bull.Bot. Res. North-East. Forest. Inst. 3(4): 58. 1983. Holotype: [China] Xinjiang, Wugia Xian, Jigen, Ximuhana, 3000 m, 29.7.1978, Exped. Xinjiang Inst. Bot. Bor.-Occ. 2128 (WUK: foto MSB!; iso: WUK!) Figures: Ho, S.B. loc. cit. 67, tab. 6, fig. 1-11; Flora Reipublicae Popularis Sinicae 42(1): 346, tab. 92, fig. 14-23. 1993 This plant has nothing to do with A. nobilis because of the spreading hairs at the calyx and the bilocular fruit. It is nearest to A. sphaerocystis Bunge. Because the taxon is based on a very short diagnosis only, a complete description is given here. Plants 7-8 cm cm tall, acaulescent, in vegetative parts covered with + symmetrically medifixed, appressed white hairs. Caudex divided with very short branches, covered with remnants of old leaves. Stipules whitish-membranous, ca. 5 mm long, adnate to the petiole for 2 3-3.5 mm, densely covered with hairs up to 1 mm long, at the margins also with basifixed hairs, the free tips triangular, glabrous on the innerside. Leaves 2-6 cm long; petiole 1-3 cm long, like the rhachis grooved on upper side, densely covered with hairs 0.5-1 mm long. Leaflets in 2-4 pairs, elliptic to obovate, 4-9 x 2-5 mm, at the apex rounded to very shortly acuminate, on both sides densely hairy like the rhachis. Peduncles 4-5 cm long, rather densely to densely white hairy like the rhachis, in upper part also with few black hairs, often 201 glabrescent. Racemes shortly ovoid, densely many-flowered. Bracts whitish-membranous, 4-5 mm long, narrowly triangular, with white and black hairs, at the margins with basifixed hairs. Pedicels ca. 1 mm long, white and black hairy. Calyx soon ovoid-inflated, 9-11 mm long, 7-9 mm in diameter, loosely covered with + spreading, basifixed to subbasifixed white hairs 1-1.5 mm long and with fewer and slightly shorter, + medifixed, subappressed black hairs, the whole surface finely rugulose-warty; teeth subulate, 1-2 mm long. Petals purple-red (according to the diagnosis). Standard ca. 20 mm long; blade ovate, ca. 7 mm wide, emarginate at the apex, subabruptly constricted in lower third and then gradually narrowed. Wings ca. 18 mm long; blades narrowly oblong, obliquely emarginate at the apex, ca. 9 x 2 mm; auricle 0.5 mm long, claw 9-10 mm long. Keel 13-15 mm long; blades obliquely elliptic, with widely curved lower edge and slightly convex upper edge, acutish at the apex, ca. 5 x 3 mm; auricle minute, claw 8-10 mm long. Staminal tube truncate at the mouth. Ovary sessile, densely appressed hairy; style hairy only at the base. Legumes subsessile, oblong, 6-7 mm long, 3-3.5 mm high and 2 mm wide, carinate ventrally, slightly grooved, at the apex abruptly terminating into a slender, hooked beak ca. 2 mm long with persistent style, bilocular; valves thin, straw-colored, densely covered with subappressed to partly ascending, subbasifixed white hairs 0.5-0.8 mm long. Astragalus pseudotauricola (Ponert) Podlech, comb. et stat. nov. — Basion.: A. macrourus Hohen. subsp. pseudotauricola (‘pseudotauricolus') Ponert, Feddes Repert. 83: 620. 1973. Astragalus parrisii Podlech, nom. nov. — Typonym: Astragalus glaberrimus Podlech, Sendt- nera 6: 146. 1999, illeg. [non Sirj. & Rech.f.]. References ALI, S.I. 1977: Flora of West Pakistan 100 (Leguminosae). — Department of Botany, Uni- versity of Karachi, Karachi. MAASSOUMI, A.A. & RANIBAR, M. 1998: Revision of the genus Astragalus L. sect. Leuco- cercis Bunge (Leguminosae) from Iran. — Iran. J. Bot. 7: 239-248 (1997). PODLECH, D. 1988: Revision von Astragalus L. sect. Caprini DC. (Leguminosae). — Mitt. Bot. Staatss. München 25: 1-924. — 1990: Die Typifizierung der altweltlichen Sektionen der Gattung Astragalus L. (Legumi- nosae). — Mitt. Bot. Staatss. Miinchen 29. 461-494. — 1998: Typification of Astragalus species II. Species mainly of the herbaria of Paris (P) and Geneva (G). — Sendtnera 5: 247-263. RECHINGER, K.H. 1958: Leguminosae in KoEIE, M. & RECHINGER, K.H., Symbolae Afghanicae II. — Biol. Skr. 9(3): 1-208. RECHINGER, K.H., DULFER, H. & PATZAK, A. 1959: Sirjaevii fragmenta astrogologica. IX—XI. Chronopus, Microphysa, Leucocercis. — Sitzungsber. Österr. Akad. Wiss., math.-nat. K1., Abt. I. 168: 693-718. Prof. Dr. Dietrich PoDLEcH, Institut für Systematische Botanik der Ludwig-Maximilians- Universität München, Menzinger Straße 67, D-80638 München, Germany. cn) ta y \ergvagl) = u Pu} eg. rn Zu Ber) zu data i > Du _ neared D 2 x) as ‘ Ma 0 P : , Aa! WIG ra! | al > © i¢yy 4 owt Ath a 3 sip ’ 4 ” 2 If wy ¢ ¥ ’ ; GA, 4 bes, En i ee ee hive gro ale TE + nani me | dark OM 7 60-12 ean Lp pantie? wills, 4 en wire ren! er ann ur Br ea rer rear vel An hei mt Ha N. Pe re Be ee a EL ie Fans ay +> ia -_ ' an |) 203 New species of Astragalus L. sect. Hymenostegis (Fabaceae) from Iran D. PODLECH & A.A. MAASSOUMI Abstract: PODLECH, D. & MAASSOUMI, A.A.: New species of Astragalus L. sect. Hymeno- stegis from Iran. — Sendtnera 7: 203-209. 2001. ISSN 0944-0178. During the work on “Flora Iranica” seven new species of Astragalus sect. Hyme- nostegis could be discovered among the rich material collected in recent times. They are described here: A. anguranensis, A bashmaghensis, A. bradosticus, A. doghrunensis, A. marivanensis, A. pseudopersicus, A. goturensis. Zusammenfassung: Im Verlauf der Bearbeitung der Gattung Astragalus für die „Flora Iranica“ konnten sieben neue Arten der Sect. Hymenostegis aufgefunden werden, die hier beschrieben werden: A. anguranensis, A bashmaghensis, A. bradosticus, A. doghrunensis, A. marivanensis, A. pseudopersicus, A. goturensis. Astragalus anguranensis Podlech & Maassoumi, spec. nov. Holotype: Iran, prov. Zanjan, distr. Mahneshan, Anguran, Belgheis mts., 2800 m, 26.7.1998, R.R. Maassoumi & V. Mozaffarian 78520 (TARI; iso: MSB). Differt praecipue et conspicue ab A. pediculariforme Maassoumi pedunculis pilis rectis appressis ad subappressis vel leviter ascendentibus 0,3-1 mm longis (nec dense pilis confuse patentibus ad 0,5 mm longis et paucioribus oblique patentibus ad patentibus 1,5-2,5 mm longis) obtectis. Plants suffruticose, caespitose, 15-22 cm tall, spiny. Stems 6-15 cm long, the lower parts ligneous, branched, densely covered with blackish remnants of old leaves, parts of the current yearl-3 cm long. Stipules membranous-hyaline, 15-20 mm long, adnate to the petiole for ca. 8-10 mm, connate behind the stem for 1.5-4 mm, the free tips narrowly triangular, 8-10 mm long, glabrous or mostly ciliate at the margins with hairs up to 1.5 mm long, in basal parts with somewhat netlike nerves, slightly netlike dissoluting with age. Leaves 3-8 cm long; petiole 1.5-3 cm long, like the rhachis rather densely covered with appressed hairs 0.3-0.7 mm long, sometimes glabrescent. Leaflets in 5-7 pairs, remote, narrowly elliptic, 10-20 x 24 mm, at the apex acuminate, pungent, with a cusp 0.5-3 mm long, on both sides in the youth rather densely to densely appressed hairy like the rhachis, with age mostly only sparsely to loosely hairy. Peduncles 2-4 cm long, shorter than the leaves, erect, densely covered with 204 appressed to slightly ascending straight hairs 0.3-1 mm long. Racemes ovoid to shortly cylindrical, 3.5-7 cm long and 2.5-3 cm in diameter, densely many-flowered. Bracts + persistent, chartaceous-glumaceous, scarcely hyaline toward the margins, 10-12 mm long and 3-5 mm wide, ovate-acuminate, shortly aristate, glabrous or sparsely ciliate at the margins. Calyx tubular-inflated, 13-15 mm long and 4-5 mm wide, whitish or creamy, in upper part purplish, with purple parallel nerves and teeth, sparsely to rather densely covered with short hairs up to 0.5 mm long and rather densely with mostly ascending or sometimes partly spreading hairs 1—2.5 mm long; teeth ca. 5 mm long. Petals pink to purplish. Standard 17-18 mm long; blade ca. 8 mm wide, obovate-panduriform, rounded at the apex, slightly to distinctly constricted in lower part, at the base acutely hastate-auriculate, with widely cuneate claw. Wings ca.16 mm long; blades narrowly obovate, at the apex rounded, ca. 7 x 3 mm; auricle ca. 0.3 mm long, claw 9-10 mm long. Keel ca. 14 mm long; blades obliquely obovate, with in upper part up to rectangularly curved lower edge and nearly straight upper edge, acutish at the apex, 4—x 2.5 mm; auricle short, claw 9-10 mm long. Ovary ca. 4-5 mm long, narrowly ellipsoidic, densely appressed hairy; style hairy at the base. Legumes unknown. Other specimens seen: Iran. Zanjan: ca. 77 km from Mahneshan to Anguran, 2300 m, 24.5.1987, Maassoumi 64809 (TARI) - on the road to Mahneshan, meck mountain [SE of Mahnestan], 1950 m, 25.7.1998, Maassoumi & Mozaffarian 78505 (MSB). Astragalus bashmaghensis Maassoumi & Podlech, spec. nov. Holotype: Iran, Kordestan, Saqqez, between Sonnate and Divandarreh, Kuh-e Ghaleh, neck mt. between Bashmagh and Jafarabad, 2300 m, 14.7.1991, V. Mozaffarian 70063 (TARI; iso: MSB!). Differt ab A. doghrunense Maassoumi & Podlech foliis 10-17 (nec 2-6) cm longis, foliolis 10-22 x 3-4 (nec 5-9 x 1,5-3) mm, pedunculis 12-18 (nec 5—8) cm longis, racemis elongatis, 6-10 (nec 3-4) cm longis, bracteis 10-15 x 5-7 (nec 9-10 x 3-5) mm, calyce pilis longioribus 2—3(—4) (nec 1-1,5) mm longis obtecto. Plants 22-30 cm tall, subacaulescent. Stems up to 7 cm long, the lower parts ligneous, branched, densely covered with blackish remnants of old leaves, part of the current year 1-2 cm long. Stipules hyaline-membranous, whitish, 15-20 mm long, narrowly triangular, adnate to the petiole for 5-8 mm, shortly connate behind the stem, the free tips many-nerved, glabrous, ciliate at the margins with ascending hairs up to 1 mm long. Leaves 2—6 cm long; petiole 3-5 cm long, like the rhachis loosely to rather densely covered with appressed to partly slightly ascending hairs 0.2-0.8 mm long. Leaflets in 6-9 pairs, narrowly elliptic, 10-22 x 3-4 mm, acuminate with a cusp 1-2 mm long, on both sides sparsely to loosely covered with appressed to subappressed hairs 0.3-0.6 mm long. Peduncles 12-18 cm long, up to 3 mm thick, + as long as the leaves, loosely to densely covered with appressed to subappressed hairs 0.3-1(-1.5) mm long. Racemes cylindric, 6-10 cm long and ca. 2 cm in diameter, densely many-flowered. Bracts chartaceous, not hyaline, yellowish, partly falling, 10-15 mm long, 5-7 mm wide, ovate-acuminate, glabrous, rarely very sparsely shortly ciliate. Calyx pale yellowish, at first tubular-ventricous, soon becoming elliptically inflated, 13-15 mm long, 205 rather densely covered with short hairs 0.2-0.5 mm long and with ascending hairs 2-3(-4) mm long; teeth subulate, ca. 5 mm long. Petals pale yellow. Standard ca. 17 mm long; blade obovate-panduriform to ovate, ca. 7 mm wide, rounded but minutely mucronulate at the apex, slightly constricted below the middle or not, angular-hastate at the base, with widely cuneate claw. Wings ca. 15 mm long; blades narrowly oblong, rounded at the apex, 6 x 2 mm; auricle ca. 0.3 mm long, claw ca. 10 mm long. Keel ca. 15 mm long; blades obliquely obovate, with in upper part rectangularly curved lower edge and slightly convex upper edge, at the apex subacute, ca. 5 x 2.8 mm; auricle minute, claw 10 mm long. Ovary narrowly ellipsoid, ca.5 mm long, densely appressed hairy; style hairy at the base. Legumes unknown. Astragalus bradosticus Maassoumi & Podlech, spec. nov. Holotype: Iran, Azerbayjan, Oroumieh, Mavana, Hakki mt. W of the village Hakki, 2100-2700 m, 31.7.1995, V. Mozaffarian, R.R. Maassoumi & Safavi 75442 (TARI; iso: MSB). Differt ab A. demonstrato Maassoumi stipulis 8-10 (nec 15-18) mm longis, foliolis minoribus, 6-10 x 1,5-3 (nec 10-23 x 2-3) mm, calyce 12-15 (nec 16-19) mm longo, vexillo ca. 16 (nec 22-26) mm longo, lamina alarum 6.5-7 x 2-2,5 (nec 11-12 x 2-3) mm. Plants suffruticose, caespitose, spiny, 17-20 cm tall, subacaulescent. Stems up to 10 cm long, branched at the base, in older parts densely covered with greyish-brown remnants of old leaves, parts of the current year ca. 1 cm long. Stipules thinly hyaline-membranous, pale yellowish, 8-10 mm long, adnate to the petiole for 4-5 mm, shortly connate behind the stem, the free tips narrowly triangular-acuminate, glabrous or sparsely ciliate at the margins. Leaves 4-7 cm long; petiole 1-1.5 cm long, like the rhachis rather densely to densely covered with subappressed to partly slightly ascending hairs 0.2-0.5 mm long and with few ascending hairs up to 1 mm long, spiny. Leaflets in 6-7 pairs, narrowly elliptic, 6-10 x 1.5-3 mm, at the apex acuminate-aristate, pungent, with a cusp 0.5-1 mm long, on both sides rather densely to densely covered with subappressed to partly slightly ascending hairs 0.3-0.5 mm long. Peduncles very short to nearly absent, with spreading hairs. Racemes nearly globular, 2.5—3 cm long and ca. 2.5 cm in diameter, rather densely many-flowered. Bracts partly falling, thinly membranous, 8-15 mm long and 4-5 mm wide, ovate to narrowly ovate, acuminate, glabrous or sparsely hairy, especially near the base and along the mid-line, ciliate at the margins. Calyx during the anthesis tubular, later on slightly inflated, 12-15 mm long and ca. 4 mm wide, whitish-yellow or sometimes towards the apex with purplish nerves, rather densely covered with ascending to spreading hairs 0.3-0.5 mm long and with ascending hairs 1.5-3 mm long; teeth subulate, purple, 6-7 mm long. Petals purplish-violet, soon fading to yellowish. Standard ca. 16 mm long; blade ca. 7 mm wide, obovate, slightly emarginate at the apex, at the base not hastate-auriculate but subangularly narrowed into the short cuneate claw. Wings 15-16 mm long; blades narrowly elliptic, at the apex obtuse, 6.5-7 x 2-2.5 mm; 11-12 x 2-3 mm; auricle ca. 0.3 mm long, claw 10-11 mm long. Keel ca. 13 mm long; blades obliquely obovate, acutish at the apex, ca. 4.5 x 2 mm; auricle short, claw 10-11 mm long. Ovary subsessile, ca. 4 mm long, densely subappressed hairy; style glabrous. Legumes unknown. 206 Other specimens seen: Iran. Azarbaijan Garbi: Oroumieh, Silvana, mts. W of the village, 1700-2100 m, 31.7.1995, Mozaffarian, Maassoumi & Safavi 75424 (MSB, TARI). Astragalus doghrunensis Maassoumi & Podlech, spec. nov. Holotype: Iran, Azarbayejan Sarki: Arasbaran protected area, Doghrun mountain, 2500-2800 m, 13.7.1977, M. Assadi & H. Sardabi 23974 (TARI). Differt ab A. recognito Fisch. stipulis hyalino-membranaceis, albidis (nec chartaceis, flavidis), foliolis utrinque pilis appressis vel subappressis laxe vel interdum densiuscule obtectis (nec dense ad densissime pilosis), pedunculis densiuscule pilis appressis, + rectis brevibus (nec pilis confusis crispis subappressis) et paucioribus 1—1,5 mm longis obtectis, bracteis glabris (nec saltem partim pilosis), calyce 11-13 (nec 13-18) mm longo, dentibus 4-5 (nec 5-8) mm longis. Plants 18-20 cm tall, caespitose. Stems 6— 10 cm long, the lower parts ligneous, branched, densely covered with blackish remnants of old leaves, part of the current year 1-2 cm long. Stipules hyaline-membranous, whitish, 12-18 mm long, adnate to the petiole for 6-10 mm, connate for 1-2 mm, the free tips narrowly triangular-acuminate, many-nerved, glabrous or more rarely the lower ones hairy, ciliate at the margins with ascending hairs up to 1 mm long. Leaves 2-6 cm long; petiole 1—2 cm long, like the rhachis densely covered with appressed to partly slightly ascending hairs 0.2-0.5 mm long. Leaflets in 5-7 pairs, narrowly elliptic, 5-9 x 1.5-3 mm, acuminate with a cusp 0.5—I mm long, on both sides loosely to more rarely rather densely covered with appressed to subappressed hairs 0.4-0.8 mm long. Peduncles 5-8 cm long, + as long as the leaves, rather densely covered with + appressed, short hairs 0.3-0.5 mm long and with much fewer hairs 1-1.5 mm long. Racemes 3-4 cm long and 2-2.5 cm in diameter, rather densely many-flowered. Bracts chartaceous, not hyaline, yellowish, lower ones or all mostly easily falling, 9-10 mm long, 3-5 mm wide, ovate, long acuminate, glabrous, ciliate. Calyx pale yellowish, at first tubular, soon becoming elliptic- to globose-inflated, 11-13 mm long, 5-7 mm wide, loosely to rather densely covered with short hairs 0.2-0.5 mm long and with ascending hairs 1—1.5 mm long; teeth subulate, 4-5 mm long. Petals pale yellow. Standard ca. 19 mm long; blade obovate-panduriform, ca. 7 mm wide, rounded but minutely mucronulate at the apex, slightly constricted below the middle, angular-hastate at the base, with widely cuneate claw. Wings ca. 16 mm long; blades narrowly obovate, rounded at the apex, 6-6.5 x 2.8-3 mm; auricle ca. 0.3 mm long, claw ca. 10 mm long. Keel ca. 15 mm long; blades obliquely obovate, with in upper part rectangularly curved lower edge and slightly convex upper edge, at the apex subacute, ca. 5 x 2.8 mm; auricle minute, claw 10 mm long. Ovary narrowly elliptic, ca.4 mm long, densely appressed hairy; style hairy at the base. Legumes unknown. Other specimen seen: Iran. Azarbaijan Sarki: Arasban protected area, Doghrun mountain, 2500 m, 15.7.1977, Assadi & Sardabi 23921 (TARI). 207 Astragalus marivanensis Podlech & Maassoumi, spec. nov. Holotype: Iran, prov. Kordestan, 50 km from Baneh to Marivan, 1650 m, 30.5.1978, H. Runemark & V. Mozaffarian 29326 (TARI; iso: MSB). Differt ab A. chrysostachyde Boiss. petiolo rhachideque pilis laxe ad dense patentibus (nec appressis vel subappressis obtecto, foliis 6-9-jugis (nec 4-7-jugis), utrinque pilis oblique patentibus (nec appressis ad subappressis) obtectis, pedunculo pilis patentibus (nec appressis) obtecto, bracteis dorso in linea mediana pilosis (nec glabris), calyce 17-20 (nec 13-16) mm longo, dentibus 8-10 mm (nec 4-6 mm) longis. Plants suffruticose, caespitose, spiny, 20-30 cm tall. Caudex very thick, up to 2 cm in diameter, branched. Stems 5—10 cm long, the lower parts ligneous, covered with dark brownish bark and remnants of old leaves, branched, parts of the current year 1.5—3 cm long, completely covered by leave stipules. Stipules hyaline-membranous, 10-18 mm long, adnate to the petiole for 5-9 mm, connate behind the stem for 1-3 mm, the free tips narrowly triangular-acuminate, glabrous. Leaves 4-12 cm long; petiole 1.5-3.5 cm long, like the rhachis loosely to densely covered with spreading hairs 0.2-0.5 mm long, spiny, at the insertion of leaflets with minute ovoid, subsessile glands. Leaflets in 6-9 pairs, narrowly elliptic, 6-20 x (1.5-)2-5 mm, acuminate, pungent, with a cusp 0.5-2 mm long, on both sides loosely to densely covered with ascending to sometimes + spreading hairs (0.3—)0.6—1.5 mm long. Peduncles 8-15 cm long, erect or ascending, rather densely to densely covered with ascending to spreading, tangled, curly hairs 0.2-0.5 mm long and with + straight, spreading hairs 1-2 mm long, sometimes glabrescent in upper part. Racemes ovoid to shortly cylindric, 4-10 cm long and 3-3.5 cm in diameter, densely many-flowered. Bracts partly falling, membranous, hyaline towards the margins, pale yellowish, 10-15 mm long and 4-6 mm wide, narrowly ovate- acuminate, shortly aristate, hairy along the midline. Calyx 17-20 mm long, at the beginning of anthesis tubular, slightly ventricous, later on ovoid-inflated and 5-6 mm wide, pale yellowish, rather densely covered with ascending to + spreading hairs 2.5-3.5 mm long; teeth subulate, 8-10 mm long. Petals pale yellow. Standard ca. 20 mm long; blade ca. 6 mm wide, obovate- panduriform, obtuse to minutely mucronulate at the apex, distinctly constricted in lower part, at the base + sharply angular-auriculate, abruptly narrowed into the narrowly cuneate claw. Wings ca. 18 mm long; blades elliptic, at the apex obtuse, ca. 7 x 2.5 mm; auricle minute, ca. 0.3 mm long, claw ca. 11 mm long. Keel ca. 15 mm long; blades obliquely obovate; with widely, sometimes nearly rectangular-curved lower edge and nearly straight upper edge, acutish at the apex, ca. 5 x 2.5 mm; auricle minute, claw ca. 10 mm long. Ovary ca. 5 mm long, elliptic, very densely ascending hairy; style glabrous. Legumes unknown. Other specimens seen: Iraq. Rowanduz distr. (MRO): in montis Kuh-Sefin, 1100-1200 m, 2.5.1893, Bornmüller 1177, 1177b (B) — in monte Handarin [Handren], 1300 m, 28.7.1893, Bornmiiller 1178 (B). Iran. Kordestan: ca. 35 km from Baneh on road to Mahabad, after Kavalan, Ganeh-Dar village, 1650 m, 13.6.2000, Maassoumi & Nikchehreh 80225 (MSB, TARI) — ca. 46 km from Sanandaj on the road to Saghez, 5 km after Iran-shah, 2000 m, 13.6.2000, Maassoumi & Nikchehreh 80213, 80214 (MSB, TARI). 208 Astragalus pseudopersicus Podlech & Maassoumi, spec. nov. Holotype: Iran, prov. Azarbaijan Sarki, ca. 5 km S of Tabriz to Sperkhan, 1600 m, 3.7.1991, V. Mozaffarian 69853 (TARI; iso: MSB). Differt ab A. persicus pedunculis crassis, ca. 2.5 mm diametro (nec sat tenuibus, ca. 1 mm diametro), bracteis 14-17 (nec 5-8) mm longis, 6-7 (nec 4-5) mm latis, sat longe acuminati- aristatis (nec breviter acuminatis), calyce 14-16 (nec 11-13) mm longo petalis + aequilongo (nec eis distincte breviore), petalis pallide lilacina nec obscure purpurei-violaceis. Plants suffruticose, caespitose, spiny, 25-35 cm tall. Stems 10-15 cm long, the older parts ligneous, branched, covered with blackish remnants of old leaves, parts of the current year 1.5-3 cm long, densely covered with nearly spreading hairs up to 2 mm long, but mostly completely covered by the stipules. Stipules chartaceous, pale yellowish, 15—18 mm long, with in basal part reticulate nerves, adnate to the petiole for 8-10 mm, connate behind the stem for 2-3 mm, the free tips triangular to long acuminate, ciliate at the margins with hairs up to 1 mm long. Leaves 7-12 cm long; petiole 1.5-3 cm long, like the rhachis rather densely to densely covered with + spreading, tangled, often flexuose hairs 0.3-0.8 mm long. Leaflets in 5-6 pairs, narrowly elliptic, 10-20 x 2-3 mm, at the apex acuminate-aristate, pungent, with a cusp 1—2 mm long, on both sides densely covered with subappressed to ascending hairs 0.5—1 mm long. Peduncles 7-13 cm long, thick, up to 2.5 mm in diameter, as long as to longer than the leaves, densely covered with tousled, + spreading hairs 0.2-2 mm long. Racemes ovoid to shortly cylindric, 4-6 cm long, 2.5-3 cm wide, densely many-flowered. Bracts partly falling, membranous, but scarcely hyaline at the margins, whitish-yellowish, towards the tip mostly purple, (12-)14-17 mm long, 6-7 mm wide, ovate to widely ovate, + long acuminate-aristate, rather densely to densely subappressed hairy, with hairs 0.3-1 mm long. Calyx at first tubular, later on slightly inflated, 14-16 mm long, 3-4 mm wide, whitish with pale purplish teeth, rather densely covered with ascending to spreading hairs 2.5—-4 mm long; teeth subulate, 4-6 mm long. Petals in dry state pale lilac to violet. Standard 15-17 mm long, + as long as calyx teeth; blade 5-5.5 mm wide, obovate-panduriform, rounded to slightly retuse at the apex, slightly constricted below the middle, at the base shortly angular-hastate, gradually narrowed into the cuneate claw. Wings 14-16 mm long; blades narrowly elliptic, obtuse at the apex, 5-7 x 1.8-2.4 mm; auricle short, claw 9-10 mm long. Keel 13-14 mm long; blades obliquely obovate, with in upper part curved lower edge and + straight upper edge, subacute at the apex, 5 x 2.5-3 mm; auricle indistinct, claw 8-9 mm long. Ovary sesssile, ca. 5 mm long, ellipsoidic, densely hairy; style hairy in lower part. Legumes unknown. Other specimen seen: Iran. Azarbaijan Sarki: Sahand, S. of Sperkhan, 2500-2850 m, 10.9.1981, Mozaffarian & Mohammadi 37535 (TARI). Astragalus goturensis Podlech, spec. nov. Holotype: Iran, Azarbaijan Garbi, Kuh Kani Ziarat N Habashi Bala prope Qotur, 2300-3000 m, 18.7.1974, W.Rechinger & J. Renz in Rechinger 49644 (M; iso: TARI, W). Differt ab omnibus speciebus sectionis Hymenostegis vexillo dorso sparse subappresse piloso. Plants suffruticose, caespitose, spiny, 18-22 cm tall. Stems 6-10 cm long, the older parts ligneous, divided, in upper parts covered with dark grey-brownish remnants of old leaves, parts of the current year 1.5-2 cm long. Stipules hyaline-membranous, whitish, 12-15 mm long, with many, at the base netlike nerves, therefore netlike dissoluting with age, adnate to the petiole for 6-8 mm, connate behind the stem for 2-3 mm, the free tips narrowly triangular, ciliate at the margins with spreading hairs up to | mm long. Leaves (2-)3-9 cm long; petiole 0.8-2.5 cm long, like the rhachis densely covered with tangled, appressed to subappressed hairs 0.05—0.4 and with fewer hairs up to | mm long, spiny. Leaflets in (2-)6-8 pairs, narrowly elliptic, 7-12 x 1.5-3 mm, at the apex acuminate-aristate, pungent, with a cusp ca. | mm long, on both sides densely to very densely covered with subappressed hairs 0.3-0.8(-1) mm long. Peduncles 5-8 cm long, as long as or slightly longer than the leaves, densely covered with short, appressed to subappressed hairs 0.3—0.8 mm long and with fewer, longer, subappressed to slightly ascending hairs up to 2 mm long. Racemes globular to ovoid, 2.5—-4 cm long, 2-2.5 cm wide, densely many-flowered. Bracts partly falling, chartaceous, yellowish, 7-12 mm long, 34.5 mm wide, ovate-acuminate, partly with an awn up to 5 mm long, loosely to rather densely covered with subappressed to slightly ascending hairs. Calyx at first tubular, slightly inflated, later on ovoid-inflated, 12-14 mm long, 4-5 mm wide, whitish with purplish teeth, rather densely covered with short, flexuose hairs 0.3-0.5 mm long and with ascending to nearly spreading hairs I-2 mm long; teeth subulate, 4-5 mm long. Petals violet, with standard sparsely hairy on the back. Standard ca. 17 mm long; blade ca. 7 mm wide, oblong, retuse at the apex, not or scarcely constricted below the middle, at the base indistinctly obtusely angular, gradually narrowed into the cuneate claw. Wings ca. 16 mm long; blades narrowly elliptic, obtuse at the apex, ca. 7 x 2.5 mm; auricle ca. 0.4 mm long, claw ca. 9 mm long. Keel ca. 13 mm long; blades obliquely obovate, with widely rectangularly curved lower edge and slightly convex upper edge, acutish at the apex, ca. 5 x 3 mm; auricle indistinct, claw ca. 9 mm long. Ovary sessile, ellipsoidic, ca, 5 mm long, densely hairy; style hairy in lower part. Legumes unknown. Prof. Dr. Dietrich PoDLECH, Institut für Systematische Botanik der Ludwig-Maximilians- Universität München, Menzinger Straße 67, D-80638 München, Germany. Dr. A.A. MAASSOUMI, Research Institute of Forests & Rangelands, P.O.Box 13185-116, Tehran, Islamic Republic of Iran. . Bee | : VRR TR DV) ne wu Sth clint er Ahad er nik i a | ae | hl ; Ly eat 0; f pray aa A M Kin en AI i "BIST Per Wo 0 " gh fo ” 1. EEE dia Sa ar ei 6 ame (1 f av ee we RN IR. airy Nie ek A Peer Are ii rnp Aw BR SEE Boar) KETTE N a) beg hee ini: i ee Wieser Far shies a Ved Th: | OY ee MAR: we \ ee as a DE MN N a I ee ote nae iy Wie 4 i sad einer > yee, sai De a iti Bu Dre Wear ie Aa aunt ee Le nee: ee yh Se ee ne Klh ZU] Lichenicolous fungi from Bavaria as represented in the Botanische Staatssammlung München D. TRIEBEL & P. SCHOLZ Abstract: TRIEBEL, D. & SCHOLZ, P.: Lichenicolous fungi from Bavaria as represented in the Botanische Staatssammlung München. — Sendtnera 7: 211-231. 2001. ISSN 0944-0178. The alphabetical list below includes the lichenicolous fungi from Bavaria as represented in the herbarium of lichenicolous fungi at the Botanische Staats- sammlung München (M). It compiles 136 taxa from Bavaria, 20 of these are for the first time reported from Germany and 9 of these are for the first time reported from Bavaria. The new combination Stigmidium lichenum (Arnold) Triebel & P. Scholz comb. nov. is proposed. Lectotype and isolectotype specimens for this taxon are designated. Zusammenfassung: Es werden die lichenicolen Pilze aus Bayern, die im Herbarium lichenicoler Pilze der Botanischen Staatssammlung München (M) aufbewahrt werden, alphabetisch ge- listet. Die Liste umfaßt 136 Taxa aus Bayern, 20 davon werden erstmals für Deutschland, 9 erstmals für Bayern angegeben. Die Neukombination Stigmidium lichenum (Arnold) Triebel & P.Scholz wird vorgeschlagen. Für diese Art werden Lecto- und Isolectotypus festgelegt. Introduction When preparing a checklist of the lichens and lichenicolous fungi of Germany (ScHoLz 2000) it became once more evident that the knowledge about the occurence and distribution of lichenicolous fungi in Germany is rather poor. Many names of lichenicolous fungi with records from Germany published by KEISSLER (1930) and some other authors are now ambiguous. The reason is that major changes of genus and species concepts took place and that within certain groups it is more or less impossible today to use some names without checking the original herbarium material. The largest collection of lichenicolous fungi in Germany is housed at the Botanische Staatssammlung München (M). The majority of the specimens there was collected by F. Arnold, A. von Krempelhuber and H. Rehm in the second half of the 19" century. These authors had an excellent knowledge of this group of fungi, described a number of new species and distributed them in several exsiccatae. Therefore, still today the knowledge of 212 lichenicolous fungi in Bavaria is mostly based on the collections made by them. Apart from specimens where the lichenicolous fungi are discovered by the collector, a large number of specimens are collected incidentally as part of lichen samples. In these cases the collector given on the label is the person who collected the host lichen and not the person who realised the presence of the lichenicolous fungus. Its presence was discovered later by someone studying the lichen more closely or by someone who was explicitly screening herbarium material of potential host lichens for lichenicolous fungi. In certain cases the later specific search was done by the first author (“det. D. T.” in the list below). The list includes all species of lichenicolous fungi from Bavaria represented in the herbarium of lichenicolous fungi at the Botanische Staatssammlung Miinchen until the end of 2000. For each species with up to five specimens in M, full information on the governmental district “Regierungsbezirk” (underlined), locality (as given on the label), collector(s), date of collecting (month, year) and host lichen is given. Host lichens [in square brackets] are named according to SCHOLZ (2000). In certain cases notes are added. Species with more than five Bavarian collections are cited by only four representative specimens and notes on the number of the additional material. References to recently published records from Bavaria based on material from M are given instead of repeating these records. Records of lichenicolous fungi in Bavaria were published especially by KREMPELHUBER (1861) and ARNOLD (various publications, see references). The material cited in these publications can generally be found in M (see references after the citation of the specimens). The list includes 136 taxa from Bavaria. More than 40 of them are not mentioned in the first checklist of lichenicolous fungi of Germany (WIRTH 1994) for various reasons, some of course because they were not yet described at that time. The recent online-checklists of FEUERER (2001a, b) give only about 30 species of lichenicolous fungi from Bavaria. Species which are — as far as known — newly published for Germany are marked with two asterisks (20 species), those newly recorded for Bavaria with one asterisk (9 species). In this context has to be mentioned that some of this information is already included in ScHoLz (2000) referring to TRIEBEL & SCHOLZ (in prep.). Lichenicolous fungi of Bavaria were distributed in the following exsiccatae: Arnold, Lich. Exs. Lich. Jur.; Arnold, Lich. Exs.; Arnold, Lich. Monac. Exs.; Hepp, Flechten Europ.; Hertel, Lich. Alp.; Poelt, Lich. Alp.; Poelt & Steiner, Lich. Alp.; Rabenhorst, Fungi Europ. Exs.; Rabenhorst, Lich. Europ. Exs.; Rehm, Ascomyc.; Thiimen, Mycoth. Univ.; Triebel, Microf. Exs. The abbreviations follow Triebel (2001). These series are effective publications and the material is widely distributed in public herbaria. List of species Abrothallus bertianus De Not. Schwaben: Landkreis Füssen, Bleckenau, Schréppel & Doppelbaur, June 1963, sub Karschia bayrhofferi, [on Melanelia glabratula]. — Oberbayern: Hirschpark bei Eichstätt, Arnold, June 1865, sub Abrothallus smithii, Arnold, Lich. Exs. Lich. Jur. 319, [on Melanelia glabratula], rev. J. Hafellner, (ARNOLD 1885: 222) — Forstenrieder Park westlich von Baierbrunn, München, Arnold, July 1896, sub Abrothallus parmeliarum, Arnold, Lich. Monac. Exs. 453, [on Melanelia glabratula], (two additional collections: Forstenrieder Park, Arnold) — Wallgauer Berge, Landkreis Tölz, Oberes Isartal zwischen Wallgau und Vorderriß unweit “Schröfeln”, Hertel 19757, Mar. 1979, sub Abrothallus parmeliarum, [on Melanelia glabratula]. — Several additional specimens from Bavaria in M. 213 * Abrothallus caerulescens C.Kotte Oberpfalz: Vorderer Bayerischer Wald, am Seidelberg östlich Brennberg, Schuhwerk 1437, Dec. 1979, [on Xanthoparmelia somloensis], det. C. Hauger. Abrothallus cetrariae C.Kotte Oberbayern: Im Walde vor der vorderen Oesteralpe, Partenkirchen, Arnold, Aug. 1875, [on Platismatia glauca], (ARNOLD 1877a: 281). Abrothallus microspermus Tul. Schwaben: Schwalbenwald bei Wemding, Arnold, Aug. 1865, [on Flavoparmelia caperata], (ARNOLD 1865: 598). — Oberbayern: Beim Buchhof, Starnberg bei München, Arnold, Apr. 1900, [on Flavoparmelia caperata], (ARNOLD 1891: 13) — Wald bei der Menterschwaige, München, Kummer, Mar. 1854, [on Flavoparmelia caperata], with Vouauxiomyces truncatus — Föhren bei der kalten Herberge, München, Arnold, Aug. 1855, [on Flavoparmelia caperata], (KREMPELHUBER 1861: 275). — Two additional specimens from Bavaria in M. Abrothallus parmeliarum (Sommerf.) Arnold Schwaben: Höhe zwischen Breitenberg und Aggenstein bei Pfronten, Allgäuer Alpen, Poelt, Sept. 1956, [on Parmelia saxatilis], with Phacopsis oxyspora var. oxyspora. — Oberbayern: Heim- garten, Schauer, July 1964, [on Parmelia saxatilis]) — Im Schweinsparke unterhalb der Eustachius Kapelle bei Eichstätt, Arnold, Nov. 1861, sub Abrothallus smithii, [on Parmelia sulcata] — Berchtesgaden, Krempelhuber, 1855, sub Parmelia saxatilis var. parasitica, [on Parmelia sul- cata], (KREMPELHUBER 1861: 275). — A number of additional specimens from various regions of Bavaria in M, see also ARNOLD (1897: 38). Abrothallus peyritschii (Stein) C.Kotte Schwaben: Rappenalpenthal im Allgäu, Rehm, 1860, [on Vulpicida pinastri] — Auf Bretter- wänden in Oberstdorf im Allgäu, Rehm [?], [no date], [on Vulpicida pinastri] — Allgäuer Alpen, Rehm, [no date], [on Vulpicida pinastri], (REHM 1890: 360). — Oberbayern: Bretterplanken bei Bad Kreuth, Bausch, 1858, [on Vulpicida pinastri]. Abrothallus prodiens (Harm.) Diederich & Hafellner Mittelfranken: Windsbach, Mittelfranken, Vill, Mar. 1926, sub Abrothallus parmeliarum, [on Hypogymnia physodes]. — Schwaben: Landkreis Augsburg, nordöstlich Rommelsried, Doppelbaur, Sept. 1949, sub Abrothallus parmeliarum, [on Hypogymnia physodes]. — Oberbayern: Bei Groß- hesselohe, München, Schnabl, Apr. 1896, sub Abrothallus parmeliarum, Arnold, Lich. Monac. Exs. 452, [on Hypogymnia physodes], two specimens of this exsiccata — Pietenfelder Höhe bei Eichstätt, Arnold, 1857, sub Abrothallus smithii, [on Hypogymnia physodes], (ARNOLD 1885: 222). — Eight additional specimens from Bavaria especially from the surroundings of München in Adelococcus alpestris (Zopf) Theiss. & Syd. Oberbayern: Dolomit oberhalb Schambach zwischen Eichstätt und Kipfenberg, Arnold, 1859, sub Polyblastia discrepans, [on Acarospora glaucocarpa], rev. D. T., conf. M. Matzer & J. Ha- fellner. Adelococcus interlatens (Arnold) Matzer & Hafellner Oberbayern: [Fränkischer Jura] Kalkfelsen der Schluchten bei Obereichstätt, Arnold, Febr. 1876, sub Verrucaria interlatens, conf. M. Matzer & J. Hafellner. Arthonia apotheciorum (A.Massal.) Almqu. Mittelfranken: Auf Keupersandstein bei Sugenheim in Franken, Rehm, 1868, sub Conida cle- mens, Arnold, Lich. Exs. Lich. Jur. Exs. 396b, [on Lecanora albescens], (ARNOLD 1885: 220) — An Süßwasserdolomitblöcken des kahlen Abhangs oberhalb Bubenheim bei Weissenburg, Arnold, 214 May 1868, sub Conida clemens, Arnold, Lich. Exs. Lich. Jur. 396a, [on Lecanora albescens], (ARNOLD 1868: 523, 1885: 220). — Oberbayern: Ziegel der Kirchhofmauer in Dornach bei Mün- chen, Arnold, Sept. 1891, [on Lecanora albescens], (ARNOLD 1892: 26). Arthonia destruens Rehm Mittelfranken: Bei Obernesselbach in Franken, Rehm, 1867, sub Conida destruens, Arnold, Lich. Exs. Lich. Jur. 377, [on Physcia stellaris (P. aipolia, P. tenella)| — Bei Obernesselbach (Sugenheim) in Franken, Rehm, 1867, sub Arthonia destruens, Rabenhorst, Lich. Europ. Exs. 816, [on Physcia stellaris], isolectotype — Bei Sugenheim in Franken, Rehm, 1867, [on Physcia stellaris], (KEISSLER 1930: 83, REHM 1891: 423). — Schwaben: Pfronten-Berg, Mayr, July 1924, [on Physcia stellaris}. Arthonia epiphyscia Nyl. Mittelfranken: Bei Sugenheim in Franken, Rehm, 1868, sub Arthonia destruens var. maculans, Arnold, Lich. Exs. Lich. Jur. 397, [on Xanthoria parietina], (KEISSLER 1930: 83). Arthonia glaucomaria Nyl. Syn. ? Lichen varians Davies (to be typified on the host lichen) Oberfranken: Bayreuth, Walther & Thümen, 1874, sub Celidium grumosum, Thümen, Mycoth. Univ. 572, [on Lecanora swartzii] — Quarzblöcke zwischen Neudorf und Pegnitz in Oberfranken, Arnold, May 1861, sub Celidium varians, Arnold, Lich. Exs. Lich. Jur. 210, [on Lecanora rupi- cola], (ARNOLD 1862: 313, 1885: 221). — Oberpfalz: Auf Basaltblöcken der rauhen Culm bei Neu- stadt in der Oberpfalz, Arnold, Sept. 1861, sub Celidium varians, Arnold, Lich. Exs. Lich. Jur. 211, [on Lecanora rupicola]. — Schwaben: Allgäuer Alpen (Gottesackerwände), Gümbel, [on Lecanora rupicola]. — Ten additional specimens from Bavaria in M. Arthonia intexta Almqu. Oberbayern: Berchtesgadener Land, Hocheck-Gipfel des Watzmanns, Wunder 1058, Aug. 1969, [on Lecidella patavina] — Berchtesgadener Land, Hochkalter, Gipfelbereich, Türk 24445, July 1997, [on Lecidella spec.] — Bayerische Alpen, Breitenstein über Fischbachau, Hertel 17767, July 1977, [on Lecidella stigmatea]. — For further specimens from Bavaria in M see HERTEL (1969), TRIEBEL (1989: 62). Arthonia molendoi (Heufl. ex Frauenf.) R.Sant. Oberbayern: Kalkwand ober der Weitalm am Hochgern bei Wessen, [Chiemgauer Berge], Arnold, Sept. 1868, [on Caloplaca spec.], two specimens (ARNOLD 1869: 254) — Mutmaßlich bei Eichstätt (scr. Lahm), Fuisting, [no date], [on Caloplaca spec.]. ** Arthonia peltigerea Th.Fr. Schwaben: Allgäu, [without indication of collector and date], [on Solorina bispora var. bispora], det. D. T. Arthonia phaeophysciae Grube & Matzer Schwaben: Landkreis Füssen, in Pfronten-Ried, Klement, 1954, [on Phaeophyscia orbicularis], det. D. T. — Erroneously omitted by SCHOLZ 2000 (see GRUBE & MATZER 1997). Arthonia punctella Nyl. Oberfranken: Kalkfelsen des Görauer Angers östlich von Weismain in Oberfranken, Arnold, Sept. 1887, [on Buellia epipolia], (ZOPF 1897: 143). Arthonia subfuscicola (Linds.) Triebel Mittelfranken: Obernesselbach unweit Sugenheim in Mittelfranken, Rehm, 1867, sub Celidium varians var. pallidae (C. grumosum), Arnold, Lich. Exs. Lich. Jur. 376, [on Lecanora albella], 215 lectotype and isolectotype of Arthonia glaucomaria var. pallidae, (ARNOLD 1885: 221) — Bei Obernesselbach in Franken, Rehm, 1871, sub Celidium varians f. pallidae, Rehm, Ascomyc. 576, two specimens, [on Lecanora albella] (ARNOLD 1885: 221). — Oberbayern: Südlich bei Schleiss- heim, München, Arnold, April 1893, sub Conida pallidae, [on Lecanora cf. albella], (ARNOLD 1897: 38, sub Conida apotheciorum), rev. D. T. Arthonia subvarians Nyl. Syn. Arthonia vagans var. lecanorina Almqu. Oberfranken: Kalkfelsen des Görauer Angers östlich von Weismain in Oberfranken, Arnold, Sept. 1887, [on Lecanora muralis]. — Oberbayern: Vor dem Hirschparke bei Eichstätt, Boll, 1887, [on Lecanora carpinea] — Hornsteine bei Nassenfels, Eichstätt, Arnold, 1859, [on Lecanora hagenii] (ARNOLD 1868: 523, 1885: 220), det. D. T. — Basispyramide östlich bei Föhring, München, Arnold, May 1888, [on Lecanora hagenii], (ARNOLD 1891: 130) — Steinhaufen zwi- schen Dettenhausen und Egling, München, Arnold, Sept. 1888, [on Lecanora polytropa] — Same locality and host data, Arnold, Aug. 1890, (ARNOLD 1891: 130) — Bei Ebenhausen, München, Arnold, May 1894 sub Conida pallidae [on Lecanora cf. carpinea], (ARNOLD 1897: 38 sub Conida apotheciorum), rev. M. Grube, growing together with Vouauxiella lichenicola. Arthopyrenia microspila Körb. Syn. Stigmidium microspilum (Körb.) D.Hawksw. Schwaben: An Buchen bei Oberstdorf, Krempelhuber (?), [no date], [on Graphis scripta]. — Oberbayern: An Buchen im Walde des Frühauf bei Eichstätt, Arnold, July 1862, [on Graphis scripta] — Im Wald zwischen Percha und Neufahrn bei Starnberg, München, Arnold, Sept. 1889, [on Graphis scripta], (ARNOLD 1891: 120). — Niederbayern: Im Donauthal zwischen Kelheim und Weltenburg, Arnold, Aug. 1860, [on Graphis scripta]. — Three additional specimens collected by Arnold at different localities in the vicinity of München in M. Arthopyrenia punctillum Arnold Syn. Stigmidium punctillum (Arnold) D. Hawksw. Oberbayern: Kampenwand bei Niederaschau, Bayerische Alpen, Arnold, Sept. 1873, [probably not lichenicolous], (ARNOLD 1874b: 380). Athelia arachnoidea (Berk.) Jülich Mittelfranken: [?]region of Ansbach, Kaiser, 1862, sub /llosporium roseum, [on Xanthoria parietina, Physcia adscendens], det. P. Diederich. — Oberpfalz: Lichtenberg, Umgebung Regens- burg, Killermann, Nov. 1918, sub Corticium centrifugum?, [on or beside lichens], det. W. Jülich. — Schwaben: Pfronten, Friedhof, Doppelbaur, 1952, sub Illosporium roseum, [on Physcia spec.], det. P. Diederich. Bachmanniomyces uncialicola (Zopf) D.Hawksw. Oberfranken: Glashütten, Arnold, Sept. 1884, sub Cladonia uncialis f. leprosa, [on Cladonia uncialis] (ARNOLD 1885: 228) — Muthmannsreuth, Bayreuth, Arnold, Sept. 1884, sub Cladonia uncialis f. leprosa, [on Cladonia uncialis]. — Mittelfranken: Nürnberg, Rehm, 1853, sub Cladonia uncialis f. leprosa, [on Cladonia uncialis]. — Oberpfalz: Auerbach, Oberpfalz, Arnold, Sept. 1884, sub Cladonia uncialis f. leprosa, [on Cladonia uncialis], (ARNOLD 1885: 228). — Schwaben: Allu- vialsand bei den Schwalbmühlen, Wemding, Arnold, Aug. 1865, sub Cladonia uncialis f. leprosa, [on Cladonia uncialis], (ARNOLD 1884: 77). Buellia pulverulenta (Anzi) Jatta Oberbayern: An der Kalkhornsteinwand auf dem Hochgern in den bayerischen Alpen, Arnold, Aug. 1869, sub Celidium muscigenae, [on Physconia muscigena], rev. J. Hafellner 1979, (AR- NOLD 1870b: 236). 216 Carbonea supersparsa (Nyl.) Hertel Schwaben: Obermädli-Alpe, Allgäuer Alpen, Rehm, [no date], [on Lecanora polytropa agg.], det. G. Rambold & D. T. Carbonea vitellinaria (Nyl.) Hertel Mittelfranken: Sandsteinfelsen des braunen Jura zwischen Wülzburg und Weißenburg, Arnold, Aug. 1861, sub Lecidea vitellinaria, Arnold, Lich. Exs. Lich. Jur. 193a, [on Candelariella vitelli- na], (ARNOLD 1862: 389). — Schwaben: Sandsteinblöcke bei Pfronten und Zell bei Pfronten, Schröppel & Poelt, 1957, sub Lecidea vitellinaria, Poelt, Lich. Alp. 31, [on Candelariella vitellina]. — Oberbayern: Hirnschnitt eines Holzpfostens bei der Station Gauting, München, Ar- nold, Aug. 1890, sub Lecidea vitellinaria, [on Candelariella vitellina], (ARNOLD 1892: 26) — Steinhaufen zwischen Dettenhausen und Egling, München, Arnold, [no date], sub Lecidea vitellinaria, Arnold, Lich. Monac. Exs. 139, [on Candelariella vitellina], (ARNOLD 1891: 129). — Additional specimens from various regions of Bavaria; see also under Polycoccum microsticticum. Cercidospora epipolytropa (Mudd) Arnold Oberbayern: Gneisblock im Walde zwischen Irschenhausen und Wadlhausen, München, Arnold, June 1891, sub Arthopyrenia lichenum, [on Lecanora polytropa], rev. D. T. Further specimens collected by F. Arnold were sent on loan and therefore their label data are actually not citable. Cercidospora macrospora (Uloth) Hafellner & Nav.-Ros. Syn. Cercidospora ulothii Körb. Schwaben: Allgäu, Lkrs. Füssen, Zell, Moor am Sulzbach, Schröppel, Aug. 1951, [on Lecanora muralis], det. D. T. — Two specimens from quite the same locality; occuring together with Stigmidium squamariae. Further specimens collected by F. Arnold were sent on loan and therefore their label data are actually not citable. ** Cercidospora verrucosaria (Linds.) Arnold Schwaben: Allgäu, Krs. Füssen, am Aggenstein, Klement, 1950, [on Megaspora verrucosa], det. D. T. — Oberbayern: Berchtesgadener Land, Nationalpark, Funtensee Tauern, Gipfelbereich, Türk 23764, July 1995, [on Megaspora verrucosa}, det. D. T. Clypeococcum hypocenomycis D.Hawksw. Oberbayern: Forstenrieder Park, siidlich Miinchen, Schwaiger 670, Sept. 1985, [on Hypo- cenomyce scalaris] — Forstenrieder Park, südlich München, Schwaiger 718, Sept. 1985, [on Hy- pocenomyce scalaris] — Lkrs. Fürstenfeldbruck, Nannhofer Wald, c. 2.5 km NW of Mammendorf, Triebel & Rambold, Sept./Oct. 1993, Triebel, Microf. Exs. 78, [on Hypocenomyce scalaris]. ** Cornutispora lichenicola D.Hawksw. & B.Sutton Schwaben: Landkreis Oberallgäu (Allgäuer Alpen), Hindelang, Iseler Berg südlich von Oberjoch, Triebel, Sept. 1983, [on Lecanora pulicaris]. Corticifraga fuckelii (Rehm) D.Hawksw. & R.Sant. Oberbayern: Kiesgrube am Anfang des Gleißenthals bei Deisenhofen, München, Arnold, Sept. 1893, [on Peltigera canina agg.], (ARNOLD 1897: 39). — Niederbayern: Kalkboden bei Gundels- hausen an der Donau (Regensburg), Arnold, Oct. 1889, sub Phragmonaevia fuckelii, Rehm, Ascomyc. 1011, [on Peltigera canina agg.]. Corticifraga peltigerae (Nyl.) D.Hawksw. & R.Sant. Mittelfranken: Waldgräben bei Ezelheim in Franken, Rehm, Sept. 1869, sub Melaspilea pelti- gerae, Rehm, Ascomyc. 19, [on Peltigera spec.], two specimens. — Oberbayern: Am Buchen- waldsaum des Gehänges zwischen Schöngeising und dem Jexhof, München, Arnold, Sept. 1894, [on Peltigera praetextata], (ARNOLD 1897: 39). 217. Cyphelium sessile (Pers.) Trevis. Mittelfranken: Sugenheim an alter Eiche (gegen Markt Bibart zu), Rehm, [no date], [on Pertusaria coccodes], det. A. Schmidt 1962, L. Tibell 1969, (KREMPELHUBER 1861: 274). — Ober- bayern: Grünwalder Park, alte Eiche, Lederer, May 1895, sub Acolium inquinans, [on Pertusaria spec.], det. A. Schmidt, (ARNOLD 1897: 30, SCHMIDT 1962: 117) — Eichstätt, Arnold, 1859, sub Acolium stigonellum, [on Pertusaria spec.], det. A. Schmidt 1962, L. Tibell 1969 — An einer alten Eiche hinter Sappenfeld bei Eichstätt, Arnold, 1859, [on Pertusaria coccodes], det. A. Schmidt 1962, L. Tibell 1969, two specimens, (ARNOLD 1885: 49, SCHMIDT 1962: 117). Dacampia engeliana (Saut.) A.Massal. Schwaben: Obermädli-Alpe im Allgäu, Rehm, [no date], [on Solorina saccata]. — Oberbayern: Kalkboden der Frauenalpe, Partenkirchen, Bayer. Alpen, Arnold, Aug. 1874, [on Solorina saccata] — Berchtesgaden, Krempelhuber, [no date], sub Sagedia engeliana, [on Solorina saccata] — Kalkwand an der Teufelskirche auf dem Hochgern in den Bayer. Alpen, Arnold, Sept. 1871, [on Solorina saccata]. — Two additional specimens from alpine regions of Schwaben and Oberbayern. Dacampia hookeri (Borrer) A.Massal. Schwaben: Am Kratzer und auf der Obermädli-Alpe bei Oberstdorf im Allgäu, Rehm, 1859, Arnold, Lich. Exs. Lich. Jur. 126, (KREMPELHUBER 1861: 232) — Allgäu, Kemtenkopf, Sendtner, [no date]. — Oberbayern: Watzmann, Krempelhuber, 1855 (KREMPELHUBER 1861: 232) — Bene- diktenwand, Gattinger, 1847. — A number of additional specimens from alpine regions of Schwa- ben and Oberbayern in M. Dactylospora athallina (Müll.Arg.) Hafellner Oberfranken: An Sandsteinfelsen des braunen Jura in schattigen Hohlwege zu Burglesau bei Scheßliz in Oberfranken, Arnold, July 1860, sub Buellia saxatilis, Arnold, Lich. Exs. Lich. Jur. 166a, [on Baeomyces rufus}, det. J. Hafellner. — Mittelfranken: Im Keupersandsteinbruche bei Deutenheim in Franken, Rehm, 1860, sub Buellia saxatilis, Arnold, Lich. Exs. Lich. Jur. 166b, [on Baeomyces rufus], det. J. Hafellner — Auf Sandstein bei Dietenhofen in Franken, Rehm, [no date], sub Buellia saxatilis, Rabenhorst, Lich. Europ. Exs. 800, [on Baeomyces rufus], det. J. Hafellner. — Oberbayern: Blomberg bei Tölz in Oberbayern, Arnold, Sept. 1880, sub Buellia athallina, [on Baeomyces spec.], conf. J. Hafellner. — A number of additional specimens from various regions of Bavaria in M, see also HAFELLNER (1979: 102). ** Dactylospora deminuta (Th.Fr.) Triebel Oberfranken: “Oberpfalz”, Neuhaus, Veldensteiner Forst, Sandsteine am Schutzengelsteinbruch, Arnold, Sept. 1887, [on Micarea peliocarpa], det. D. T. — Oberbayern: Berchtesgaden, Schneib- steingipfel, Rauchenberger, [no date], [on Lecanora epibryon], det. D. T. — The latter specimen might represent a new taxon; see ZHURBENKO & SANTESSON (1996). Dactylospora parasitaster (Nyl.) Arnold Oberbayern: Ruine ober Erlingshofen, Eichstätt, Arnold, June 1869, [on Chromatochlamys muscorum], lichenicolous or bryophilous. Dactylospora tegularum (Arnold) Hafellner Oberbayern: Auf Ziegeln der Kirchhofmauer in Tutzing, Miinchen, Arnold & Gmelch, Sept. 1890, sub Buellia tegularum, Arnold, Lich. Exs. 1512, [on Caloplaca arenaria], holotype, (AR- NOLD 1891: 130, HAFELLNER 1979: 149) — Ziegel der Kirchhofmauer in Buchendorf, Miinchen, Arnold, Sept. 1891, [on Caloplaca arenaria], three specimens, (ARNOLD 1892: 26, HAFELLNER 1979:-451): 218 Echinothecium reticulatum Zopf Schwaben: Landkreis Füssen, Sandstein in Moorwiese bei Zell, Schröppel, May 1951, [on Xan- thoparmelia conspersa], det. D. T. Endococcus perpusillus Nyl. Syn. Endococcus macrosporus (Arnold) Nyl. Oberfranken: Quarzfelsen bei Pegnitz in Oberfranken, Arnold, 1869, [on Rhizocarpon distinc- tum]. — Schwaben: Bregenzer Wald, Südgrat des Fellhorn bei Oberstdorf, Poelt, Sept. 1964, sub Tichothecium stigma, [on Rhizocarpon geographicum]. — A number of additional specimens from Bavaria in M; see TRIEBEL (1989: 97). Endococcus propinquus (Körb.) D.Hawksw. Oberbayern: Kalkstein eines Steinhaufens zwischen Dettenhausen und Egling, München, Arnold, Aug. 1889, sub Tichothecium gemmiferum, [on Verrucaria nigrescens]. — A number of additional specimens from Bavaria in M; see TRIEBEL (1989: 102). Endococcus rugulosus Nyl. Mittelfranken: Quarzblock zwischen Neuhaus und Krottensee in der Oberpfalz, Arnold, Sept. 1882, sub Tichothecium gemmiferum, [on Aspicilia cinerea], two specimens. Epicladonia sandstedei (Zopf) D.Hawksw. Oberbayern: Bad Reichenhall, Kugelbachwald, Schönau, July 1929, sub Diplodina sandstedei, [on Cladonia fimbriata] — Berchtesgaden-Hintersee, Seeklause, auf einem Felsblock, Schönau, Aug. 1928, sub Diplodina sandstedei, [on Cladonia fimbriata]. Epigloea urosperma Döbbeler Oberbayern: Landkreis Garmisch-Partenkirchen, Murnauer Moos, feuchte Wiese an der Straße zwischen Aschau und Eschenlohe, G. & P. Döbbeler, Aug. 1983, [on Placynthiella uliginosa s.l.], holotype. — For several additional specimens from Bavaria in M see DÖBBELER (1994: 278). *]llosporiopsis christiansenii (B.L.Brady & D.Hawksw.) D.Hawksw. Schwaben: Allgäu, Bezirk Pfronten, bei Zell, Schröppel & Suessenguth, Sept. 1951, sub /llo- sporium roseum, [on Physcia stellaris], det. D. T. — Allgäu, Buxheim bei Memmingen [or] Blai- chach bei Immenstadt, Rueß, [?] 1899, sub /llosporium roseum, [on Phaeophyscia orbicularis], specimens from two localities intermixed, one is Phaeophyscia with I. christiansenii, det. D. T. — Oberbayern: Großhesselohe bei München, Schnabl, Oct. 1897, sub /llosporium roseum, [on Physcia tenella), det. D. T. Illosporium carneum Fr. Oberfranken: Erlangen, 500 m SSW Unterailsfeld bei Gößweinstein, Mägdefrau, Apr. 1941, [on Peltigera spec.]. — Oberbayern: Waldboden beim Buchhof unweit Starnberg bei München, Ar- nold, Sept. 1896, Arnold, Lich. Monac. Exs. 456, [on Peltigera didactyla], (ARNOLD 1897: 39) — Landkreis Fürstenfeldbruck, Haspelmoor, am Südwestrand, Doppelbaur, Apr. 1951, [on Peltigera rufescens| — Berchtesgaden, am Abstieg vom Trischhübel zum Wimbachtal, Schoenau, Aug. 1920, [on Peltigera didactyla]. — Five additional specimens from various regions of Bavaria in M. Lichenoconium erodens M.S.Christ. & D.Hawksw. Schwaben: Landkreis Oberallgäu, Hindelang, Oberjoch, westlich des Berghauses Iseler, Triebel 375, Sept. 1983, [on Hypogymnia physodes| — Landkreis Oberallgäu, Hindelang, Oberjoch, Kematsrieder Moor nordwestlich des Ortes, Triebel 418, Sept. 1983, [on Hypogymnia physodes]. — Oberbayern: Forstenrieder Park an der südlichen Stadtgrenze von München, Triebel 48, Feb. 1982, [on Ramalina pollinaria] — Forstenrieder Park an der südlichen Stadtgrenze von München, Schwaiger 750, Sept. 1985, [on Hypogymnia physodes] — Landkreis Eichstätt, Affental W bis 219 NW von Walting, Triebel 1544 & Rambold, Aug. 1985, [on Ramalina pollinaria] — Schlierseer Berge, Landkreis Rosenheim, zwischen Klamm-Alm und Asten (ca. 8 km NE Bayrischzell), Wunder 3142, Okt. 1985, [on Parmelia sulcata], accompanied by Lichenopuccinia poeltii, Sclerococcum parmeliae; separated, see also Tremella hypogymniae. Lichenoconium lecanorae (Jaap) D.Hawksw. Oberbayern: Landkreis Miesbach, Weidfilz c. 2.5 km W of Königsdorf, Triebel & Rambold 5815, July 1990, [on Lecanora spec.] — Forstenrieder Park südlich München, Schwaiger 666, Sept. 1985, [on Lecanora conizaeoides] — Landkreis Weilheim, Paterzeller Eibenwald N Paterzell, Triebel 477, Sept. 1984, [on Lecanora albella]. — Niederbayern: Bayerischer Wald, Landkreis Regen, Gipfel des Großen Falkensteins NE Zwiesel, Schuhwerk 895/4b, May 1989, [on Lecanora cf. polytropa]. — Two additional specimens from the surroundings of München; see also Tremella hypogymniae. Lichenoconium pyxidatae (Oudem.) Petr. & Syd. Oberbayern: Forstenrieder Park, südlich von München, Schwaiger 938, Sept. 1985, [on Cladonia fimbriata] — Forstenrieder Park, südlich von München, Schwaiger 1282, Oct. 1985, [on Cladonia cf. digitata] — Ammergauer Alpen, Friedergrieß etwa 1,5 km nordöstlich Griesen, Döbbeler, Sept. 1985, [on Cladonia pyxidata ssp. pocillum]. Lichenoconium usneae (Anzi) D.Hawksw. Schwaben: Landkreis Füssen, Bleckenau am Weg zum Rothmoos, Schröppel & Doppelbaur, June 1963, sub Coniothyrium usneae, [on Usnea spec.]. — Oberbayern: Oberammergau, Schnabl, Aug. 1896, [on Melanelia exasperata], holotype of Coniothyrium imbricariae Allescher, det. D. Hawksworth, (ALLESCHER 1897: 18). Lichenodiplis lecanorae (Vouaux) Dyko & D.Hawksw. Oberbayern: Espen zwischen Roggenstein und Buchheim, München, Arnold, May 1896, sub Tichothecium microcarpon, [on Caloplaca cerina], rev. D. T. — An einem Lattenzaun in Untersendling, München, Arnold, Nov. 1890, sub Tichothecium microcarpon, [on Caloplaca spec], ev. D. T. **Lichenopeltella coppinsii Earland-Bennett & D.Hawksw. Oberbayern: Kalkstein eines Steingerölles im Buchenwalde unterhalb Pullach, München, Ar- nold, Oct. 1888, sub Arthopyrenia lichenum, [on Verrucaria spec.], rev. D. T., (ARNOLD 1891: 131) — Kalksteine im Buchenwalde oberhalb Delling westlich von Weßling, München, Arnold, June 1889, sub Arthopyrenia lichenum, [on Verrucaria muralis], two specimens, rev. D. T., (ARNOLD 1891: 131) — Kalksteine im Buchenwalde oberhalb Delling bei Weßling, München, Arnold, June 1889, sub Arthopyrenia lichenum, [on Verrucaria spec.], rev. D. T., (ARNOLD 1891: 131) — Kalk- steine des Waldweges zwischen dem Bahnwärterhaus und [dem Forsthaus] Wörnbrunn im Grün- walder Park, München, Arnold, April 1890, sub Arthopyrenia lichenum, [on Verrucaria spec.], rev:/D,T. *Lichenopeltella maculans (Zopf) Höhn. Oberpfalz: Kleiner Osser über Lam, Poelt, June 1965, [on Umbilicaria hirsuta], conf. M. Matzer. ** Lichenopuccinia poeltii D.Hawksw. & Hafellner Oberbayern: Schlierseer Berge, Landkreis Rosenheim, zwischen Klamm-Alm und Asten (ca. 8 km NE Bayrischzell), Wunder 3142, Okt. 1985, [on Parmelia sulcata], det. D. T., accompanied by Lichenoconium erodens, Sclerococcum parmeliae; separated. 220 Lichenosticta alcicorniaria (Linds.) D.Hawksw. Oberbayern: München, Großhesselohe, Schnabl, 1894, [on Cladonia fimbriata], holotype of Aposphaeria cladoniae, det. D. L. Hawksworth, (ALLESCHER 1896: 32) — Landkreis Berchtes- gadener Land, Gemeinde Ramsau, Nordseite des Hochkalter-Massivs, Wunder 3990, Aug. 1985, [on Cladonia coniocraea], det. J. Hafellner — Landkreis Berchtesgadener Land, Gemeinde Ramsau, zwischen Grieshütte und Wimbachschloß, Wunder 3972, Aug. 1985, [on Cladonia spec.], det. J. Hafellner — Berchtesgaden, Eckerfirst (Hoher Göll), Schoenau, Aug. 1920, [on Cladonia digi- tata}, det. D. T. ** Lichenostigma cosmopolites Hafellner & Calatayud Niederbayern: Bayrischer Wald, Umgebung von Regen, Britzelmayr 55], [no date], [on Xanthoparmelia somloensis], det. D. T. — Bayrischer Wald, without more definite locality data, Gattinger, Sept. 1846, [on Xanthoparmelia somloensis], det. D. T. *Lichenostigma elongatum Nav.-Ros. & Hafellner Unterfranken: Hammelburg, Vill, 1889, [on Lobothallia radiosa], det. D. T. — Hammelburg, Vill, May 1890, [on Lobothallia radiosa] — Mainfranken, Landkreis Gemünden, Kalkfelsen im Retzbachtal, Klement 04843, 1956, [on Lobothallia radiosa], det. D. T. *Lichenostigma rugosum G.Thor Schwaben: Allgäu, Landkreis Füssen, Illasberg bei Roßhaupten, K/ement, 1952, [on Diplo- schistes scruposus], det. D. T. — Landkreis Füssen, Zell, auf Flyschsandstein im Moor zwischen Pfronten und Zell, Schröppel, May 1951, [on Diploschistes scruposus], det. D. T. Marchandiomyces aurantiacus (Lasch) Diederich & Etayo Oberfranken: Bayreuth, de Thümen, Mar. 1876, [on lichen indet.], conf. P. Diederich (cf.). — Oberbayern: Oberammergau, Dickelschwaige, Allescher, Sept. 1889, [on lichen indet.], conf. P. Diederich (cf.). Marchandiomyces corallinus (Roberge) Diederich & D.Hawksw. Oberbayern: Bei Großhessellohe, München, Schnabl, Sept. 1896, [on Parmelia sulcata], (AR- NOLD 1897: 39), det. P. Diederich — Bei Großhessellohe, München, Schnabl, Sept. 1896 , [on Melanelia spec.], sub /llosporium corallinum, Arnold, Lich. Monac. Exs. 455, (ARNOLD 1897: 39), det. P. Diederich — Wasserburg am Inn, Innpromenade, Schoenau, Sept. 1942, sub Illosporium roseum, [on Hypogymnia physodes, Parmelia sulcata], det. P. Diederich. Merismatium discrepans (J.Lahm) Triebel A number of specimens from Bavaria in M are cited by TRIEBEL (1989: 194). Merismatium peregrinum (Flot.) Triebel Oberbayern: Amphibolitblock zwischen Buchhof und Haarkirchen, Starnberg, München, Arnold, Sept. 1895, [on Rimularia badioatra], det. D. T., (HERTEL & RAMBOLD 1990: 167) — Gneisblock im Walde links der Straße zwischen Percha und Neufahrn, Starnberg, München, Arnold, Sept. 1890, [on Rimularia badioatra], det. D. T., (HERTEL & RAMBOLD 1990: 167). — Nieder- bayern: Böhmerwald, Höllbachspreng am Falkenstein bei Zwiesel, Schuhwerk 1121/1, Apr. 1979, [on Rimularia gibbosa], det. D. T. — One additional specimen on Rimularia gibbosa in M is cited in HERTEL & RAMBOLD (1990: 172). Merismatium scammoecum Lettau Mittelfranken: Landkreis Hersbruck, Kalkfelsen oberhalb Enzendorf im Pegnitzthale, Arnold, July 1868, lectotype and isolectotype of Polyblastia discrepans f. dilatata. — For additional specimens from Bavaria in M see TRIEBEL (1989: 194). 221 Microcalicium arenarium (Hampe ex A.Massal.) Tibell Oberfranken: Eckersdorfer Schlucht bei Bayreuth, Arnold, May 1861, sub Cyphelium are- narium, Arnold, Lich. Exs. Lich. Jur. 205, [on Psilolechia lucida] — Bayreuth, bei Eckersdorf, Arnold, [no date], [on Psilolechia lucida], (ARNOLD 1862: 306). — Niederbayern: Bayerisch-Böh- mischer Wald, Landkreis Wolfstein, Lesesteinhaufen nördlich Grainet, Poelt, Aug. 1971, sub Coniocybopsis arenaria, [on Psilolechia lucida] — Bayerischer Wald, Landkreis Passau, im Gaißa- Tal, nahe Tiefenbach bei Passau, nördlich der Gaißa-Mühle, Huber, Apr. 1978, sub Microcalicium arenicola, Hertel, Lich. Alp. 322, [on Psilolechia lucida] — Bayerischer Wald bei Sankt Oswald, Krempelhuber, [no date], sub Calicium citrinum, [on Psilolechia lucida]. ** Minutoexcipula tuerkii Hafellner Schwaben: Allgäuer Alpen, Schrecksee, Grummann 4208 dupl., Aug. 1951, [on Pertusaria glo- merata], det. D.T, accompaniedby Muellerella lichenicola. ** Monodictys cellulosa S.Hughes Oberbayern: Berchtesgadener Land, S von Bad Reichenhall, Ramsau, knapp WNW der Mündung des Lattenbaches in die Ramsauer Ache, Wittmann, Mar. 1995, [on Parmelina tiliacea], dei): T. Muellerella lichenicola (Sommerf. ex Fr. : Fr.) D.Hawksw. Oberfranken: Pottenstein in Oberfranken, Wagner, [no date], sub Pharcidia, [on Caloplaca holocarpa s.l.], rev. D. T. — Additional specimens from Bavaria in M are cited in TRIEBEL (1989: 158), see also Minutoexcipula tuerkii. Muellerella pygmaea (Körb.) D.Hawksw. var. pygmaea Mittelfranken: Bei Dietenhofen in Franken, Rehm, [no date], [on Lecidea fuscoatra s.l.]. — Oberbayern: Dachziegel in Solndorf bei Feldkirchen, München, Arnold, July 1891, sub Ticho- thecium pygmaeum, [on Lecidea fuscoatra] — Landkreis Berchtesgadener Land, Gemeinde Schö- nau am Königssee, zwischen Jenner und Schneibsteinhaus, Türk & Wunder, Oct. 1984, [on Caloplaca proteus], at least three additional specimens on this host from the Bavarian Alps — Steinhaufen zwischen Dettenhausen und Egling, Miinchen, Arnold, July 1891, sub Tichothecium pygmaeum, {on Lecidea fuscoatra], four specimens from the same locality and host, one of this is Arnold, Lich. Monac. Exs. 201 (see also TRIEBEL 1989: 164). Muellerella pygmaea var. athallina (Müll.Arg.) Triebel Specimens from Bavaria in M are cited in TRIEBEL (1989: 170). Muellerella pygmaea var. ventosicola (Mudd) Triebel Oberbayern: Steinhaufen zwischen Dettenhausen und Egling, München, Arnold, Sept. 1890, sub Tichothecium pygmaeum, [on Rhizocarpon distinctum], three additional specimens from the same locality with different dates — Gneisstein im Waldgehänge südlich von Haarkirchen, München, Arnold, Sept. 1890, sub Tichothecium pygmaeum, [on Rhizocarpon reductum]. Nectriopsis lecanodes (Ces.) Diederich & Schroers Mittelfranken: Waldgraben bei Ezelheim in Franken, Rehm, Oct. 1869, Rehm, Ascomyc. 38, with Scutula dedicata (Karsteniomyces-state). ** Nectriopsis parmeliae (Berk. & M.A.Curtis) M.S.Cole & D.Hawksw. Schwaben: Pfronten, Rue’, Nov. 1897, sub Pleonectria lichenicola, [on Physcia stellaris], rev. Br! 222 Opegrapha physciaria (Nyl.) D.Hawksw. & Coppins Mittelfranken: Bei Obernesselbach in Franken, Rehm, 1867, sub Celidium varium, Rabenhorst, Lich. Europ. Exs. 785, [on Xanthoria parietina], with Xanthoriicola physciae, (ARNOLD 1885: 221) — In Laubwäldern bei Sugenheim in Franken, Rehm, Nov. 1866, sub Celidium varium, Arnold, Lich. Exs. Lich. Jur. 335, [on Xanthoria parietina], with Xanthoriicola physciae, (ARNOLD 1885: 221). — Oberbayern: Auf Buchen am Waldsaume ober den Anlagen bei Eichstätt, Arnold, Oct. 1868, sub Celidium varium, Arnold, Lich. Exs. Lich. Jur. 335b, [on Xanthoria parietina], mostly Xanthoriicola physciae, (ARNOLD 1868: 523, 1885: 221). — Two further specimens from the same localities in M. Opegrapha pulvinata Rehm Oberbayern: Chiemgauer Alpen, Weitalm am Hochgern, Steiner, Aug. 1950, [on Dermato- carpon miniatum] — Kalkblöcke zwischen Taubensee und Rauher Nadel, Bayerische Alpen, Arnold, Sept. 1872, [on Dermatocarpon spec.]. Opegrapha rupestris Pers. A large number of specimens from Bavaria is kept in the lichen herbarium. The following three specimens from Bavaria have been distributed in exsiccatae. Oberfranken: Zwischen Tüchersfeld und Pottenstein in Oberfranken, Arnold, [no date], sub Opegrapha saxicola, Rabenhorst, Lich. Europ. Exs. 334. — Oberbayern: Chiemseer Berge, zwischen Karkopf und Hochries südlich oberhalb Grainbach, Hertel, Aug. 1974, sub Opegrapha saxicola, Hertel, Lich. Alp. 331 — Im Laubwalde der Anlagen bei Eichstätt, Arnold, Aug. 1864, Arnold, Lich. Exs. Lich. Jur. 286, (ARNOLD 1884: 661). Opegrapha zwackhii (A.Massal. ex Zwackh) Källsten Oberfranken: Oberfranken, Schönfeld, Kaiser, [no date], [on Phlyctis argena], det. B. J. Coppins. Phacopsis oxyspora (Tul.) Triebel & Rambold var. oxyspora Oberbayern: Im Walde vor der Vorderen Oesteralpe bei Partenkirchen, Arnold, Aug. 1875, [on Platismatia glauca]. — For a second specimen from Bavaria see under Abrothallus parmeliarum; see also ARNOLD (1897: 38). Phacopsis oxyspora var. fusca Triebel & Rambold Two specimens from Bavaria in M; see TRIEBEL et al. (1995: 80). Phacopsis vulpina Tul. Oberbayern: Berchtesgaden, Funtensee und Feldalmgrube, Rauchenberger, [no date], [on Letharia vulpina], (KREMPELHUBER 1861: 275) — Funtenseetauern, Sendtner, 1856, [on Letharia vulpina], det. D. T. Phaeopyxis punctum (A.Massal.) Rambold, Triebel & Coppins Oberfranken: Keuperregion bei Bayreuth, Walther, Feb. 1863, sub Nesolechia punctum, Arnold, Lich. Exs. Lich. Jur. 252, [on Cladonia spec.]. — This and additional specimens from Bavaria in M see RAMBOLD & TRIEBEL (1990: 385). Phaeospora parasitica (Lönnr.) Arnold Oberbayern: Landkreis Bad Tölz-Wolfratshausen, im Lainbachtal südöstlich Benediktbeuren, Feuerer 12354a, 12354c, Aug. 1982, [on Rhizocarpon umbilicatum], two specimens — Landkreis Miesbach, Wendelstein, Nordseite kurz unterhalb des Gipfels, Feuerer 1155, Aug. 1982, [on Rhizocarpon umbilicatum]. 2233 Phaeospora rimosicola (Leight. ex Mudd) Hepp Oberfranken: Oberhalb Pottenstein in Oberfranken, Wagner, Sept. 1867, sub Xenosphaeria rimosicola, Arnold, Lich. Exs. Lich. Jur. 379, [on Rhizocarpon petraeum], two specimens (AR- NOLD 1885: 224). — Schwaben: Landkreis Oberallgäu, Hochgrat am Weg zum Rindalphorn, Feuerer 11847 & Höhne, May 1982, sub Phaeospora parasitica, [on Rhizocarpon petraeum]. — Ober- bayern: Auf dem Hochgern ober Wessen in den bayerischen Alpen, Arnold, Aug. 1869, [on Rhizocarpon petraeum], (ARNOLD 1870b: 236) — Begraster Abhang, Westseite des Blombergs bei Tölz, Arnold, Aug. 1881, sub Phaeospora rimosicola, Arnold, Lich. Exs. Lich. Jur. 379b, [on Rhizocarpon cf. subconcentricum]. — A number of additional specimens from Bavaria in M. Phaeosporobolus usneae D.Hawksw. & Hafellner Oberbayern: Nationalpark Berchtesgaden, Watzmannmassiv, ca. 200 m nordwestlich der Kli- mastation oberhalb der Mitterkaseralm, Hadatsch 66, Sept. 1991, [on Usnea subfloridana (det. R. Türk)], det. D. T. — Nationalpark Berchtesgaden, Steinernes Meer, Stuhlgrabenkogel, Wunder 4139 & Türk, Sept. 1985, [on Letharia vulpina], det. D. T. — Berchtesgadener Alpen, über dem Funtensee, Poelt, Sept. 1948, [on Letharia vulpina], det. D. T. — Im Karolinengeräumt im Forstenrieder Park, München, Arnold, Sept. 1890, [on Usnea subfloridana (det. P. Clerc)], det. D. T. — Ammergauer Alpen, Weg von der Notkarspitze nach Farchant, Poelt, Oct. 1977, [on Usnea spec.], det. D. T. — See also Phoma cytospora. *Phoma cytospora (Vouaux) D.Hawksw. Schwaben: Allgäu, Pfronten, Hochalm, Schröppel, Nov. 1954, [on Parmelia saxatilis], accompanied by Phaeosporobolus usneae, det. D. T. **Phoma denigricans Hafellner Oberbayern: Alpspitze, Krempelhuber, 1846, [on Lecanora epibryon], det. D. T. — Watzmann, Krempelhuber, 1855, [on Lecanora epibryon], det. D. T. — Nationalpark Berchtesgaden, Stei- nernes Meer, Viehkogel, Gipfel, Wunder & Türk, Sept. 1985, [on Lecanora epibryon], det. D. T. Phyllosticta lichenicola Allescher Oberbayern: München, Ebenhausen, April 1895, Schnabl, [on Parmotrema chinense], holo- type, (ALLESCHER 1896: 32, ARNOLD 1897: 39, HAWKSWORTH 1981: 83 "nomen ambiguum"). Plectocarpon lichenum (Sommerf.) D.Hawksw. Unterfranken: Bei Ruppertshiitten (Lohr) im Spessart, Rehm, July 1877, sub Celidium stic- tarum, Rehm, Ascomyc. 424, [on Lobaria pulmonaria]. — Oberfranken: Bei Waldstein im Fich- telgebirge, Laurer, [no date], sub Celidium stictarum, Hepp, Flechten Europ. 590, [on Lobaria pulmonaria]. — Oberbayern: Lauterbachthal bei Kreuth, Kühn, Sept. 1872, sub Celidium stic- tarum, Rabenhorst, Fungi Europ. Exs. 1648, [on Lobaria pulmonaria] — Bei Raitenbuch, Eich- stätt in Bayern, Arnold, 1862, sub Celidium stictarum, Rabenhorst, Lich. Europ. Exs. 657, [on Lobaria pulmonaria], (ARNOLD 1885: 221). — A number of additional specimens from Bavaria collected in the 19"" century, but only four specimens collected in the 20" century in M; see also ARNOLD (1897: 38). Polycoccum arnoldii (Hepp) D.Hawksw. Oberfranken: Auf steinigem Boden des Brand bei Hetzelsdorf in Oberfranken, 1857, [on Diploschistes muscorum], (ARNOLD 1858: 702, KREMPELHUBER 1861: 276). — Oberbayern: Auf steinigem Boden, zwischen Winterhof u. Ruppertsbuch, bei Eichstätt, Arnold, [no date], sub Phaeospora (Sphaeria?) arnoldi, Hepp, Flechten Europ. 701, [on Diploschistes muscorum], two isotypes, (ARNOLD 1858: 702) [The host is originally given as “Urceolaria scruposa f. iridata” . This is a synonym of Diploschistes muscorum, see LUMBSCH (1987: 604)] — Auf Erde, kahle Bergeshöhen bei Eichstätt, Arnold, 1856, [on Diploschistes muscorum], (ARNOLD 1858: 702) — Kahle Bergeshöhen bei Eichstätt, Arnold, 1857, [on Diploschistes muscorum], (ARNOLD 1858: 702) — Steiniger Boden bei Nassenfels unweit Eichstätt, Arnold, Oct. 1858, [on Diploschistes mus- corum], (ARNOLD 1858: 702, KREMPELHUBER 1861: 276). — Two additional specimens in M. 224 *Polycoccum cladoniae Diederich & D.Hawksw. Oberbayern: Rottach b. Tegernsee, ?Krempelhuber, Oct. 1859, [on Cladonia digitata], det. D. T. — Berchtesgaden, Wimbachtal, Schneelahnerwald, Schoenau, Aug. 1929, [on Cladonia digi- tata], det. D. T. Polycoccum marmoratum (Kremp.) D.Hawksw. Oberfranken: Kalkfelsen ober der Oberfellndorfer Straße bei Streitberg, Arnold, July 1868. — Oberbayern: Kalkfelsen bei Hüting, Eichstätt, Arnold, Aug. 1860, 3 syntypes of Tichothecium marmoratum, (ARNOLD 1861: 265) — Dolomit unterhalb der Römerschanze zwischen Piesenhard und der Feldmühle bei Eichstätt, Arnold, 1860, syntype of Tichothecium marmoratum, (ARNOLD 1861: 265) — Kalkfelsenwand gegenüber Kunstein bei Eichstätt, Arnold, June 1862, sub ?Ticho- thecium marmoratum, Arnold, Lich. Exs. Lich. Jur. 246 — Kalksteine im Walde des Arzberges ober Beilngries im Altmühltal, Arnold, 1867 — Kalkfelsen am Ufer des Taubensees ober Wessen in den bayerischen Alpen, Arnold, Sept. 1868, (ARNOLD 1869: 268). — A number of additional specimens collected by F. Arnold mostly in the surroundings of Eichstätt in M (see also ARNOLD 1861: 265, 1885: 157). Polycoccum microsticticum (Leight. ex Mudd) Arnold Oberbayern: Steinhaufen zwischen Dettenhausen und Egling, München, Arnold, July 1891, sub Polycoccum microsticticum, Arnold, Lich. Monac. Exs. 200a, [on Acarospora fuscata], the specimen in M has also Carbonea vitellinaria on Candelariella vitellina, another specimen from the same locality was collected by F. Arnold in Sept. 1890, (ARNOLD 1891: 132, 1892: 26) — Steinhaufen zwischen Deining und Egling, München, Arnold, Aug. 1888, [on Acarospora fuscata], one additional specimen with similar locality data in M. ** Polycoccum sporastatiae (Anzi) Arnold Schwaben: Gaultsandsteinfelsen der oberen Hochalpe der Gottesackerwände im Allgäu, Rehm, 1860, [on Sporastatia testudinea], det. D. T. — Hochalpe im Allgäu, Rehm, 1860, [on Sporastatia testudinea], det. D. T. Pronectria robergei (Mont. & Desm.) Lowen Mittelfranken: Waldgraben bei Ezelheim in Franken, Rehm, Oct. 1869, sub Nectriella robergei, Rehm, Ascomyc. 37, [on Peltigera rufescens], det. R. Santesson. — Oberbayern: Schöngeising, Le- derer, Sept. 1894, sub Nectria lichenicola, [on Peltigera didactyla] - Am Saume des Buchenwaldes zwischen Schöngeising und dem Jexhof, München, Arnold, June 1894, sub Nectria lichenicola, Arnold, Lich. Monac. Exs. 373, [on Peltigera didactyla], det. R. Santesson, two specimens of this exsiccata and one additional specimen from the same locality (ARNOLD 1897: 39). Pyrenidium actinellum Nyl. Schwaben: Obermädeli-Alpe, Allgäu, Schwarze Milz [borderland to Austria, Tirol], Rehm [on Solorina crocea], det. D. T. — Oberbayern: zwischen Buchendorf und Leutstetten, München, Arnold, 1892, sub Phaeospora granulosae, Arnold, Lich. Exs. 1564, [on Trapeliopsis granulosa], holotype, (ARNOLD 1897: 39, TRIEBEL 1989: 68) — Buchendorfer Gemeindewald, München, Arnold, 1892, sub Phaeospora granulosae, [on Trapeliopsis granulosa], (ARNOLD 1897: 39, TRIEBEL 1989: 70). ** Sclerococcum parmeliae Etayo & Diederich Schwaben: Allgäu, Hohen Schwangau, in der Blockenau, Schröppel, Okt. 1954, [on Parmelia saxatilis], det. D. T. — Oberbayern: Schlierseer Berge, Landkreis Rosenheim, zwischen Klamm-Alm und Asten (ca. 8 km NE Bayrischzell), Wunder 3142, Okt. 1985, [on Parmelia sulcata], det. D. T., accompanied by Lichenoconium erodens, Lichenopuccinia poeltii, separated. 225 Sclerococcum sphaerale (Ach.) Fr. : Fr. Oberfranken: Auf Granit auf dem Schneeberge im Fichtelgebirge, Gümbel, 1860, sub Cyphelium corallinum, [on Pertusaria corallina]. — Mittelfranken: Auf Keupersandsteinfelsen im Rügländer Thale bei Adelmannsdorf (Dietenhofen) in Franken, Rehm, 1855, sub Cyphelium corallinum, [on Pertusaria corallina]. — Oberpfalz: Oberpfälzer Wald, Landkreis Neunburg v. W., Gneisblöcke im Blockgewirr bei Warnthal, Poelt, May 1965, [on Pertusaria corallina]. — Niederbayern: Böh- merwald, Gipfel des Großen Rachel bei Zwiesel, C. & J. Poelt & Ullrich, Sept. 1963, sub Coniothecium pertusariicola, [on Pertusaria corallina], rev. D. T. Scutula dedicata Triebel, Wedin & Rambold A number of specimens from Bavaria in M; see TRIEBEL et al. (1997: 327). Scutula epiblastematica (Tul.) Rehm See TRIEBEL et al. (1997: 330). Scutula heeri (Hepp) Trevis. See TRIEBEL et al. (1997: 332). Scutula krempelhuberi Korb. Oberbayern: Berchtesgaden, Krempelhuber, 1854, [on Solorina saccata]. — Two additional collections on Solorina might belong to this species, WEDIN & TRIEBEL (in prep.). Scutula miliaris (Wallr.) Trevis. See TRIEBEL et al. (1997: 336). ** Skyttea cismonicae Hafellner Oberbayern: Zwischen Vordergraseck und dem Eckbauern bei Partenkirchen in Siidbayern, Ar- nold, Sept. 1874, [on Loxospora cismonica]. Host lichen distributed sub Haematomma cismonicum, Arnold, Lich. Exs. Lich. Jur. 141b. Lichenicolous fungus separated, det. P. Diederich & D. T. ** Sphaerellothecium minutum Hafellner Oberpfalz: Ossa [= Osser], Lederer, 1893, [on Sphaerophorus fragilis], det. D. T. — Nieder- bayern/Oberpfalz: Arber, Lederer, 1892, [on Sphaerophorus fragilis], det. D. T. Sphaerellothecium propinquellum (Nyl.) Cl.Roux & Triebel Three specimens from Bavaria in M; see ROUX & TRIEBEL (1994: 531). Sphinctrina anglica Ny). Schwaben: Nebelhornbahn bei Oberstdorf, Allgäu, Schauer, Oct. 1962, sub Sphinctrina micro- cephala, conf. O. Löfgren. — Oberbayern: Holzapfelskreuth, Lederer, 1892, sub Sphinctrina microcephala, conf. O. Lofgren — Bei Eichstätt, Arnold, 1862, sub Sphinctrina microcephala, Arnold, Lich. Exs. Lich. Jur. 245, conf. O. Léfgren, two specimens of this exsiccata, (ARNOLD 1885: 61) — Ostlich GroBhadern, Miinchen, Arnold, 1862, sub Sphinctrina microcephala, Arnold, Lich. Monac. Exs. 193, conf. O. Léfgren, two specimens of this exsiccata. — A number of additional specimens from Bavaria in M; see also ARNOLD (1862: 395) and HERTEL et al. (2001). Sphinctrina turbinata (Pers. : Fr.) De Not. Mittelfranken: Dietenhofen, Rehm, 1855, [on Pertusaria pertusa], conf. O. Léfgren, (KREM- PELHUBER 1861: 273). — Oberbayern: Forstenrieder Park südlich München, Schwaiger 1467, May 1986, [on Pertusaria coccodes] — Zwischen Schloß Pahl und Kerschlach, Oberbayern, Poelt, June 1957, sub Sphinctrina gelasinata, [on Pertusaria coronata], conf. O. Löfgren — Bei Gauting, München, Hiendlmayr, Aug. 1864, sub Calycium turbinatum, [on Pertusaria pertusa], conf. O. 226 Löfgren. — A number of additional specimens from Bavaria in M; see also ARNOLD (1858: 700, 1885: 61, 1897: 32) and HERTEL et al. (2001). ** Stigmidium bellemerei Cl.Roux & Nav.-Ros. Oberpfalz: Ruine Ehrenfels bei Beratzhausen in der Oberpfalz, Arnold, Sept. 1879, sub Arthopyrenia lichenum, Arnold, Lich. Exs. 820, [on Lecania suavis], det. D. T. Stigmidium cerinae Cl.Roux & Triebel Schwaben: Allgäu, Aggenstein-Gipfel, Klement, 1952, [on Caloplaca cerina var. chloroleuca}, (ROUX & TRIEBEL 1994: 48]). Stigmidium congestum (Körb.) Triebel Six specimens from Bavaria in M; see ROUX & TRIEBEL (1994: 486). Stigmidium fuscatae (Arnold) R.Sant. Mittelfranken: Brauner Jura eines Steinbruchs nördlich von Treuchtlingen, Eichstätt, Arnold, Apr. 1870, [on Acarospora fuscata], holotype, (ARNOLD 1872: 572, 1874a: 174, 1885: 223). — Oberpfalz: Brauner Jura des Erzberges bei Amberg, Arnold, Sept. 1861, [on Acarospora fuscata], two specimens, (ARNOLD 1872: 572, 1885: 223) — Brauner Jura auf dem Erzberge bei Amberg, Arnold, 1862, [on Acarospora fuscata], (ARNOLD 1872: 572, 1885: 223). Stigmidium hageniae (Rehm) Hafellner Mittelfranken: Bei Obernesselbach in Franken, Rehm, 1870, sub Sphärella hageniae, Rehm, Ascomyc. 32, [on Anaptychia ciliaris], isotype — Bei Obernesselbach in Franken, Rehm, 1870, [on Anaptychia ciliaris! — Bei Sugenheim in Franken, Rehm, 1868, sub Pharcidia hageniae, Arnold, Lich. Exs. Lich. Jur. 398, [on Anaptychia ciliaris]. Stigmidium lichenum (Arnold) Triebel & P.Scholz, comb. nov. Bas. Arthopyrenia lichenum Arnold, Flora 59: 567 (1876) Pharcidia lichenum Arnold, Flora 55: 572 (1872) — not val. publ. (ICBN Art. 52.1 — superflous) Pharcidia lichenum Arnold, Verh. K.K. Zool.-Bot. Ges Wien 22: 302 (1872) — not val. publ. (ICBN Art. 32.1 — without descr.) Pharcidia lichenum (Arnold) Arnold ex G.Winter, Rabenh. Kryptog.-Flora Deutschl., 2. Aufl., Pilze 1(2): 343-344 (1885) (ICBN Art. 33.3 ex 7 — valid comb.) Oberbayern: Ober Wasserzell bei Eichstätt, Arnold, Apr. 1876, sub Arthopyrenia lichenum, Arnold, Lich. Exs. Lich. Jur. 692a, [on Verrucaria maculiformis], lectotype and isolectotype designated here (ARNOLD 1876: 567, 1885: 222) — Kalksteine ober Wasserzell, Eichstätt, Arnold, 1876, sub Arthopyrenia lichenum, (ARNOLD 1876: 567, 1885: 222) — Auf Solnhofer Schiefer- platten bei Eichstätt, Arnold, May 1886, sub Pharcidia lichenum, Rehm, Ascomyc. 893 — Ober- halb Wasserzell bei Eichstätt, Arnold, May 1886, sub Arthopyrenia lichenum, Arnold, Lich. Exs. Lich. Jur. 692b, (ARNOLD 1885: 222). - A number of additional specimens collected by F. Arnold in M. **Stigmidium mycobilimbiae Cl.Roux, Triebel & Etayo Oberbayern: Kalkhornsteinwand auf dem Hochgern in den Bayerischen Alpen, Arnold, Aug. 1869, [on Mycobilimbia accedens], det. D. T. Stigmidium psorae (Anzi) Hafellner Oberbayern: Berchtesgaden, Steinernes Meer, bei der Schönbichlalpe, Schönau, Aug. 1929, [on Psora decipiens], (TRIEBEL 1989: 84). 2217 Stigmidium rivulorum (Kernst.) Cl.Roux & Nav.-Ros. Oberbayern: Kalk-Glimmersteine im Bache östlich beim chinesischen Turm, München, Arnold, July 1897, [on Verrucaria aquatilis], (ARNOLD 1901: 13) — Kalksteine im Bache beim chinesi- schen Turm im englischen Garten, München, Arnold, July 1897, sub Arthopyrenia rivulorum, Arnold, Lich. Monac. Exs. 486b, [on Verrucaria aquatilis], (ARNOLD 1901: 13) — Steine im Bache beim chinesischen Turm im englischen Garten, München, Arnold, Sept. 1897, sub Arthopyrenia rivulorum, Arnold, Lich. exs. 1566b, [on Verrucaria aquatilis], (ARNOLD 1901: 13). Stigmidium schaereri (A.Massal.) Trevis. Three specimens from Bavaria in M; see ROUX & TRIEBEL (1994: 507). ** Stigmidium squamariae (de Lesd.) Cl.Roux & Triebel Schwaben: Allgäu, Lkrs. Füssen, Pfronten, im Aachtal unterhalb des Kienberges, Schröppel, June 1953, [on Lecanora muralis], det. D. T. — Allgäu, Lkrs. Füssen, Zell, Moor am Sulzbach, Schröppel, Aug. 1951, [on Lecanora muralis], occuring together with Cercidospora macrospora, ate IDR AR Stigmidium tabacinae (Arnold) Triebel Syn. Stigmidium glebarum (Arnold) Hafellner Oberfranken: Dolomitfelsen ober der Waidmannsgeseeser-Schlucht bei Pottenstein in Ober- franken, Arnold, Sept. 1886, sub Arthopyrenia glebarum on Thalloid. caeruleonigric., Arnold, Lich. Exs. 1196, [on Toninia physaroides], holotype and isotype of Stigmidium glebarum. *Syzygospora physciacearum Diederich Oberbayern: Isaranlagen gegen Thalkirchen, Allescher, Nov. 1878, sub /llosporium roseum, [on Physcia tenella], det. P. Diederich. *Taeniolella punctata M.S.Christ. & D.Hawksw. Schwaben: Giinzburg/ Donau, Donauauen "Leibi", Doppelbaur F 46, March 1963, [on Graphis scripta], det. D. T. — Oberbayern: Menterschwaige bei München, Krempelhuber, 1959, [on Graphis scripta], det. D. T. Taeniolina scripta (P.Karst.) P.M.Kirk Oberbayern: Oberschleißheim, nördlich München, Naturwaldreservat Fasanerie, Korbiniani- Hölzl, Hadatsch, Sept. 1990, [on Lepraria incana], det. D. T. Thelocarpon lichenicola (Fuckel) Poelt & Hafellner Oberfranken: An Sandsteinen des braunen Jura in einem Waldhohlwege am Wege von Banz nach Altenbanz in Oberfranken, Arnold, Sept. 1882, sub Thelocarpon excavatulum, Arnold, Lich. Exs. 960, isotype, (ARNOLD 1885: 167), det./conf. G. Salisbury 1972, J. Kocourkova-Horakova 1997 — An Sandsteinen des braunen Jura, Waldhohlwege bei Schloß Banz in Oberfranken, Arnold, Sept. 1882, sub Thelocarpon excavatulum, Arnold, Lich. Exs. 960, ? holotype, det./conf. J. Kocourkovä-Horäkovä 1997. (The label of this specimen is handwritten with drawings of F. Arnold in contrast to the same number of this exsiccata cited above). — Oberfranken: Sandsteine beim Schutzengelsteinbruch im Veldensteiner Forste, Neuhaus, Oberpfalz, Arnold, [no date], det. ]. Kocourkova-Horakova 1997. Tremella coppinsii Diederich & G.Marson Oberbayern: Tal des Schwarzbaches zwischen dem Wirtshaus Wachterl und Kaltbach, ca. 10,5 km südlich von Bad Reichenhall, Berchtesgadener Land, Wittmann, Feb. 1995, [on Platismatia glauca]. 228 Tremella hypogymniae Diederich & M.S.Christ. Schwaben: Landkreis Sonthofen, 6,5 km SE of Oberstaufen, along trail below Spitzlerberg, at the right side of the creek Weißach between lower station of the Hochgratbahn and crossing with trail to Lanzenbach, Triebel & Rambold 5934, May 1991, [on Hypogymnia physodes], with Lichenoconium lecanorae and L. erodens on Tremella hypogymniae, (DIEDERICH 1996: 94), det. P. Diederich. Vouauxiella lichenicola (Linds.) Petr. & Syd. Schwaben: Allgäuer Alpen, Landkreis Oberallgäu, Hindelang, Oberjoch, Straßenrand westlich des Kematsrieder Moores, Triebe! 405, Sept. 1983, [on Lecanora chlarotera]. — Oberbayern: Land- kreis Eichstätt, Walnußbaumallee beim Ziegelhof ca. 2 km NE Eichstätt, Triebel & Rambold 1532, Aug. 1985, [on Lecanora chlarotera] — Biehl bei Benedictbeuern, an der Straße von Kimsee, Allescher, Sept. 1885, sub Pharcidia epicymatia, [on Lecanora chlarotera], vid. C. Roux — Am nördlichen Waldrande zwischen Obersradling und Großhesselohe, Boll, Feb. 1889, sub Pharcidia congesta, [on Lecanora chlarotera], vid. C. Roux — Bei Ebenhausen, München, Arnold, May 1894 [on Lecanora cf. carpinea], (ARNOLD 1897: 38), growing together with Arthonia subvarians. — Niederbayern: Bayerischer Wald, am Helmberg südöstlich Münster bei Straubing, Poelt, June 1965, [on Lecanora chlarotera], growing together with Stigmidium congestum. — See also ROUX & TRIEBEL (1994: 486). Vouauxiella verrucosa (Vouaux) Petr. & Syd. Oberbayern: Südbayern, Petershausen, nördlich des Gehöfts Weingarten, Rambold 760, June 1982, [on Lecanora chlarotera] — Berchtesgadener Land, zwischen Hinterbrand und Vorderbrand, Wunder 2057, Aug. 1983, [on Lecanora chlarotera], det. D. T. (cf.). ** Vouauxiomyces ramalinae (Nordin) D.Hawksw. Oberbayern: Forstenrieder Park südlich München, Schwaiger 782, Sept. 1985, [on Ramalina spec], det) D: T. Vouauxiomyces truncatus (de Lesd.) Dyko & D.Hawksw. Oberbayern: Forstenrieder Park südlich München, Schwaiger 1454, 1466, May 1986, [on Flavoparmelia caperata], one additional collection from Forstenrieder Park 1986 — Forstenrieder Park, Arnold, June 1895, [on Flavoparmelia caperata] — Uferpromenade von St. Bartholomä am Königssee, Berchtesgadener Land, Wunder 2235, Apr. 1984, [on Flavoparmelia caperata]. Xanthoriicola physciae (Kalchbr.) D.Hawksw. Schwaben: Ammergauer Alpen, Landkreis Füssen, südlich Buching, Schröppel & Poelt, Mar. 1963, sub Coniosporium physciae, Poelt & Steiner, Lich. Alp. 198, [on Xanthoria parietina]. — Oberbayern: Bei Großhesselohe, München, Schnabl, Sept. 1896, sub Coniosporium physciae, Ar- nold, Lich. Monac. Exs. 454, [on Xanthoria parietina], two specimens of this exsiccata, (ARNOLD 1897: 39), conf. T. Feuerer — Landkreis Erding, c. 4 km SSE of Moosburg, parking lot of the motor highway E 53 (A 92), Rambold, Mar. 1998, sub Xanthoriicola physciae, Triebel, Microf. Exs. 350, [on Xanthoria parietina] — Landkreis Landsberg am Lech, Pappelallee nördlich Schwif- ting, Feuerer 2736, Mar. 1979, [on Xanthoria parietina]. -— A number of additional specimens from Upper Bavaria in M, see map in FEUERER & HÖHNE (1980: 132). Xenonectriella lutescens (Rehm) Weese Oberpfalz: Straßenböschung des Kreuzbergs zwischen Hahnbach und Vilseck in der Oberpfalz, Arnold, Sept. 1882, sub Pleonectria lutescens, Arnold, Lich. Exs. 963, [on Solorina saccata], (ARNOLD 1885: 222) — Kreuzberg bei Vilseck, Oberpfalz, Arnold, Sept. 1882, sub Nectria luiescens, Rehm, Ascomyc. 681, [on Solorina saccata], isolectotype and one additional specimen from the same locality, (ARNOLD 1885: 222). 229 Zwackhiomyces dispersus (J.Lahm ex Körb.) Triebel & Grube Oberbayern: Am Waldfußweg unterhalb Grünwald, München, Arnold, May 1889, sub Cerci- dospora, [on Protoblastenia rupestris], (TRIEBEL 1989: 120). Zwackhiomyces immersae (Arnold) Grube & Triebel Oberbayern: Südbayern, Werdenfelser Land, Umgebung Garmisch-Partenkirchen, Partnach- klamm, unteres Ende der Schlucht, Rambold, July 1983, [on Clauzadea monticola], (TRIEBEL 1989: 120). ** Zwackhiomyces inconspicuus Grube & Hafellner Mittelfranken: Sugenheim in Franken, Rehm, 1867, sub Arthopyrenia aspiciliae, [on Lecanora dispersa], rev. D. T. Appendix Two specimens of Chaenothecopsis rubescens housed in the lichen collection in M are cited here. This is in addition to SCHMIDT (1970: 147) who notifies the occurence of this species in Bavaria without citing any locality. Chaenothecopsis rubescens Vain. Unterfranken: Haßfurt, in der Altach an einer Eiche, Vill, 1894, sub Calycium parietinum, det. A. Schmidt 1984. — Oberbayern: Alte Eiche im Wald der Donau-Auen zwischen Gerolfing und Dünzelau bei Ingolstadt, Arnold, May 1867, det. A. Schmidt 1969. We are very grateful to H. Hertel, W. Lippert and F. Schuhwerk (all München) for help with reading handwritten labels and clarifying localities. D. L. Hawksworth (Madrid, Spain) is thanked for reviewing the manuscript. The nomenclatural advice of R. Santesson (Uppsala, Sweden) is greatly acknowledged. P. Diederich (Strassen, Luxembourg) and M. Grube (Graz, Austria) supported us by determing some critical specimens. E. Hofmann (Miinchen) kindly checked the English of an earlier version. References ALLESCHER, A. 1896: Diagnosen einiger neuer, im Jahre 1895 gesammelter Arten bayerischer Pilze aus der Abteilung der Fungi imperfecti. — Ber. Bayer. Bot. Ges. 4: 31-40. — 1897: Diagnosen einiger neuer, meist im Jahre 1896 gesammelter Arten bayerischer Pilze, nebst Bemerkungen über einige kritische Arten. — Ber. Bayer. Bot. Ges. 5: 13-25. ARNOLD, F. 1858: Die Lichenen des fränkischen Jura. (Schluss.). — Flora 41: 691-702. — 1861: Die Lichenen des fränkischen Jura. (Schluss.). — Flora 44: 257-268. — 1862a: Die Lichenen des fränkischen Jura. — Flora 45: 305-313. — 1862b: Die Lichenen des fränkischen Jura. (Schluss.). — Flora 45: 385-396. — 1865: Die Lichenen des fränkischen Jura. — Flora 48: 596-599. — 1868: Die Lichenen des fränkischen Jura. — Flora 51: 520-524. — 1869: Lichenologische Fragmente. 5. Zwei Tage in Wessen. — Flora 52: 251-255, 257-269. — 1870a: Lichenologische Fragmente. 6. Bei Partenkirchen. — Flora 53: 1-10,1-23. — 1870b: Lichenologische Fragmente. 9. Auf dem Hochgern. — Flora 53: 225-236. — 1872: Die Lichenen des fränkischen Jura. — Flora 55: 569-573. 230 — 1874a: Lichenologische Fragmente. 16. (Schluss.). — Flora 57: 173-175. — 1874b: Lichenologische Fragmente. 17. I. Der Taubensee. II. Die Kampenwand. III. Der Wallberg. — Flora 57: 376-384, 449-455. — 1876: Die Lichenen des fränkischen Jura. — Flora 59: 564-567. — 1877a: Lichenologische Fragmente. 20. I. Partenkirchen. II. Kampenwand. — Flora 60: 281-286. — 1877b: Lichenologische Fragmente. 20. III. Parasiten. — Flora 60: 298-302. — 1884a: Die Lichenen des fränkischen Jura. — Flora 67: 65-96. — 1884b: Die Lichenen des fränkischen Jura (Fortsetzung.). — Flora 67: 645-664. — 1885a: Die Lichenen des fränkischen Jura. (Fortsetzung.). — Flora 68: 49-80. — 1885b: Die Lichenen des fränkischen Jura. (Fortsetzung.). — Flora 68: 143-176. — 1885c: Die Lichenen des fränkischen Jura. (Schluss.). — Flora 68: 211-246. — 1891: Zur Lichenenflora von München. — Ber. Bayer. Bot. Ges. 1, Anh.: 1-147. — 1892: Zur Lichenenflora von München. — Ber. Bayer. Bot. Ges. 2, Anh.: 1-76. — 1897: Zur Lichenenflora von München. — Ber. Bayer. Bot. Ges. 5, Anh.: 1-45. — 1901: Zur Lichenenflora von München. — Ber. Bayer. Bot. Ges. 8, Anh.: 1-24. DIEDERICH, P. 1996: The lichenicolous Heterobasidiomycetes. — Biblioth. Lichenol. 61: 1-198. DÖBBELER, P. 1994: Epigloea urosperma (Ascomycetes) — ein neuer Flechtenparasit. — Sendt- nera 2: 277-282. FEUERER, T. 2001a: Checkliste der Flechten und flechtenbewohnenden Pilze Bayerns. Version 1.1.2001. — http://www .rrz.uni-hamburg.de/biologie/ialb/herbar/bay_f2.htm. — 2001b: Checkliste der Flechten und flechtenbewohnenden Pilze der Landeshauptstadt München. Version 1.1.2001. — http://www.rrz.uni-hamburg.de/biologie/ialb/herbar/m_rl 1. htm. — & Houne, N. 1980: Beitrag zur Floristischen Kartierung von Flechten in Bayern. 1. Die Gattung Xanthoria. — Ber. Bayer. Bot. Ges. 51: 123-132. GRUBE, M., MATZER, M. 1997: Taxonomic concepts of lichenicolous Arthonia species. — In: TURK, R. & ZORER, R. (eds.): Progress and problems in lichenology in the Nineties — IAL 3. — Biblioth. Lichenol. 68: 1-17. HAFELLNER, J. 1979: Karschia. Revision einer Sammelgattung an der Grenze von lichenisierten und nichtlichenisierten Ascomyceten. — Beih. Nova Hedwigia 62: 1-248. HAWKSWORTH 1981: The lichenicolous Coelomycetes. — Bull. Br. Mus. Nat. Hist. Bot. S. 9: 1-98. HERTEL, H. 1969: Arthonia intexta Almqu., ein vielfach verkannter fruchtkörperloser Flechten- parasit. — Ber. Dtsch. Bot. Ges. 82: 209-220. — & RAMBOLD, G. 1990: Zur Kenntnis der Familie Rimulariaceae (Lecanorales). — In: JAHNS, H.M. (ed.): Contributions to lichenology in honour of A. Henssen. — Bibl. Lichenol. 38: 145-189. — & SCHWAIGER, J., VORWERK, B. 2001: Die Flechtenflora der Staatsforste am Südrand Münchens, einst und jetzt. — In: BEsL, H. & SCHÖNFELDER, P. (eds.): Bresinsky-Festschrift. — Hoppea 61: 445-524. KEISSLER, K. v. 1930: Die Flechtenparasiten. — In: ZAHLBRUCKNER, A. (ed.): Dr. L. Raben- horst’s Kryptogamen-Flora von Deutschland, Österreich und der Schweiz. Zweite Auflage. Die Flechten (Lichenes) 8: 1-712. Leipzig. KREMPELHUBER, A. v. 1861: Die Lichenen-Flora Bayerns. — Denkschr. Königl. Bayer. Bot. Ges. 4(2): 1-317. 231 LumBscH, H.T. 1987: Eine neue Subspecies in der Flechtengattung Diploschistes aus der Südhemisphäre. — Herzogia 7: 601-608. RAMBOLD, G., TRIEBEL, D. 1990: Gelatinopsis, Geltingia and Phaeopyxis: three helotialean genera with lichenicolous species. — Not. R. Bot. Gard. Edinb. 46: 375-389. REHM, H. 1890: Die Pilze Deutschlands, Oesterreichs und der Schweiz. III. Abtheilung: Ascomyceten: Hysteriaceen und Discomyceten. — In: WINTER, G. & REHM, H. (eds.): Dr. L. Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz. Zweite Auflage. Pilze 1(3): 337-400. Leipzig. — 1891: Die Pilze Deutschlands, Oesterreichs und der Schweiz. III. Abtheilung: Ascomy- ceten: Hysteriaceen und Discomyceten. — In: WINTER, G. & REHM, H. (eds.): Dr. L. Raben- horst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz. Zweite Auflage. Pilze 1(3): 401-608. Leipzig. Roux, C., TRIEBEL, D. 1994: Revision des espéces de Stigmidium et de Sphaerellothecium (champignons lichénoles non lichénisés, Ascomycetes) correspondant a Pharcidia epicymatia sensu Keissler ou a Stigmidium schaereri auct. — In: Roux, C. (ed.): Hommage scientifique a G. Clauzade. — Bull. Soc. Linn. Provence 45: 451-542. SCHMIDT, A. 1962: Die Gattung Cyphelium in Bayern. — Ber. Bayer. Bot. Ges. 35: 113-119. — 1970: Anatomisch-taxonomische Untersuchungen an europäischen Arten der Flechten- familie Caliciaceae. — Mitt. Staatsinst. Allg. Bot. Hamburg 13: 111-166. ScHoLz, P. 2000: Katalog der Flechten und flechtenbewohnenden Pilze Deutschlands. — Schriftenr. Vegetationskd. 31: 1-298. TRIEBEL, D. 1989: Lecideicole Ascomyceten. Eine Revision der obligat lichenicolen Ascomyce- ten auf lecideoiden Flechten. — Biblioth. Lichenol. 35: 1-278. — 2001: [first posted on 01-02-01; most recent update: 01-02-21]. Search Bibliography of Exsiccatae. — Botanische Staatssammlung München: http://www.mycology.net/exsiccata/ ExsiccataFind.cfm. — München, Germany. — & RAMBOLD, G., ELIX, J.A. 1995: A conspectus of the genus Phacopsis (Lecanorales). — Bryologist 98: 71-83. — & Wepm, M., RAMBOLD, G. 1997: The genus Scutula (lichenicolous ascomycetes, Lecanorales): species on the Peltigera canina and P. horizontalis groups. — In: TIBELL, L. & HEDBERG, I. (eds.): Lichen studies dedicated to Rolf Santesson. — Symb. Bot. Ups. 32(1): 323-337. WIRTH, V. 1994: Checkliste der Flechten und flechtenbewohnenden Pilze Deutschlands — eine Arbeitshilfe. — Stuttg. Beitr. Naturkd., S. A (Biol.) 517: 1-63. ZHURBENKO, M., SANTESSON, R. 1996: Lichenicolous fungi from the Russian Arctic. — Herzogia 12: 147-161. Zopr, W. 1897: Untersuchungen über die durch parasitische Pilze hervorgerufenen Krank- heiten der Flechten (Erste Abhandlung). — Nova Acta Leopoldina 70: 97-192. Dr. Dagmar TRIEBEL, Botanische Staatssammlung München, Menzinger Strasse 67, D-80638 München, Germany. E-mail: triebel@botanik.biologie.uni-muenchen.de Dr. Peter SCHOLZ, Paetzstr. 37, D-04435 Schkeuditz, Germany. E-mail: UfU.Bioindikation@t-online.de Ay i} vo > i) 1 -. 1 Ti N M J ay r Lab IE i if ; Wit ‘if et N Pi} i i a8 Bit ot, +) , yy eae” A A I EN ae Spas eee aia el Ni: " MW taken, www IPNE Nm mw Y Epinions inn ‘in bile vw } a ee A My Aue ET Monk 1h upp ee i aa Mimi De ESEL DIE dnt tale) FG ye Han ‘ 2 an (uh ' ale j ‚tt eh via a on : ae 2 "CR 0.77" | fi ete ed | i | € lad ER CH | 2ace " anh ‘ltd | EL nn rn ‘eae ; | 2’ = | Hay? AR ae iy) ı! (i = m Leer Ina hat een le) i id ell ; eo Le perce (shes mare mol we rt 2 er een > Vv uw e en ae ee TE a Va e i. re Ka! Mie Dur Seb. PER 233 Taxonomic Revision of Astragalus sect. Acanthophace (Fabaceae) Sh. ZARRE & D. PODLECH Abstract: ZARRE, SH. & PODLECH, D.: Taxonomic Revision of Astragalus sect. Acanthophace (Fabaceae). — Sendtnera 7: 233-251. 2001. ISSN 0944-0178. Sect. Acanthophace Bunge of the genus Astragalus L. is revised taxonomically. The newly described sect. Lamprocarpa Maassoumi is reduced to synonymy under sect. Acanthophace. The systematic position of Astragalus pseudoangustifolius Sirj. & Rech.f. and A. cryptocarpus DC. is clarified through selection of epitypes. A. cystosus Zarre & Podlech and A. hezarensis Zarre & Podlech are described as new species here for the first time. Zusammenfassung: Eine Revision von Astragalus L. sect. Acanthophace Bunge wird vorgelegt. Die erst kürzlich beschriebene Sekt. Lamprocarpa Maassoumi wird als Synonym zu Acan- thophace gestellt. Die systematische Position von A. pseudoangustifolius Sirj. & Rech.f. und A. cryptocarpus DC. wird durch die Wahl von Epitypen fixiert. A. cystosus Zarre & Podlech und A. hezarensis Zarre & Podlech werden als neue Arten beschrieben. Introduction Sect. Acanthophace Bunge is revised here in the frame work of preparing a treatment of the thorny Astragalus groups for the Flora Iranica. A complete revision of this section and sect. Aegacantha Bunge was published by I. DEML in 1972. However, due to the poor availibility of material to the author at the time and the unclear borders between the sections of the thorny Astragali, her work was a preliminary one and it was necessary to revise the section again. In the famous work of BUNGE (1868/1869) this section was classified under subgen. Phaca Bunge, characterized by large pods rupturing the calyx. The current systematic position of sect. Acanthophace is, however, within subgen. Astragalus (see PODLECH 1982). It has been shown that this section is a basal group within thorny Astragalus and there are several symplesiomorphies shared by the species belonging to this section. Presence of flattened ribbon-like and papillose hairs, large ligneous pods and a standard not sharply differentiated to claw and limb are some such symplesiomorphies (ZARRE 2000). This relatively small section is heterogeneous, so the attribution of some species to the section is doubtful. For this reason MAASSOUMI (1995) separated A. ovigerus [as a new 234 species A. lamprocarpus| under a new section, namely sect. Lamprocarpa Maassoumi. But the correctness of this treatment has been questioned (ZARRE 2000) because of paraphyletic nature of such a classification. Astragalus sect. Acanthophace is closely related to A. sect. Megalocystis Bunge and sect. Adiaspastus Bunge. A. lalesarensis Bornm. plays a transitional role between all three sections. Vegetative form and shape of petals in this species are very similar to A. hezarensis (sect. Acanthophace), while its red colored and inflated fruiting calyx reminds one of A. murinus (sect. Megalocystis). Moreover, it is similar (and most probably closely related) to A. sahendi (sect. Adiaspastus) in many respects. The appearance of paraclades on short lateral branches at the base of each leaf is a rare character in the thorny Astragali which is developped in A. sclerocladus of sect. Acanthophace. This character occurs otherwise in sect. Poterion Bunge, which is also more or less a basal group within the thorny Astragali (see ZARRE 2000). This study is mainly based on herbarium material. Several sheets have been examined for each species, received on loan from the following herbaria (abbreviations according to HOLMGREN & al.): B, BM, E, G-BOIS, G-DC, JE, K, M, M, P, IRAN, TARI, TUH, UPS, US, W and WU. Moreover, during several excursions in Iran, some of the species were studied in the field by one of the authors (ZARRE). Flower dissections were made from several specimens and added to the sheets after examination. Taxonomic Account Astragalus L sect. Acanthophace Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 45. 1868. Lectotype (DEML 1972): A. schistocalyx Bunge. = A. sect. Lamprocarpa Maassoumi, Iran. J. Bot. 6: 199. (1994) 1995. Type (monotypic): A. lamprocarpus Maassoumi. Subshrubs or cushion-forming plants, loosely or densely branched at base. Caudex up to 3 cm in diam., greyish brown to black. Hairs mostly flattened and ribbon-like, papillose on surface, white in vegetative parts, mixed with grey and black ones in the inflorescences. Stipules chartaceus, yellowish-white, adnate to petiole for ca. 1/3-1/2 of their length, otherwise free from each other or only shortly connate at the base, triangular-ovate, the free acute tips with only one main nerve. Leaves paripinnate; rhachides thick and rigid, hairy, whitish green; leaflets + thick, mostly small, green or greyish green, hairy, mostly soon glabrescent, sometimes completely glabrous. Inflorescences lateral, few—10-flowered racemes. Peduncle shorter than leaves, appressed hairy. Bracts chartaceous, with one main nerve. Bracteoles sometimes present, adnate to the calyx, singular or in pairs. Pedicels up to 2.5 mm long. Calyx tubular, chartaceus, with 14-16 parallel indistinct longitudinal, thin nerves, sparsely to densely hairy, ruptured by the pod; teeth subulate. Corolla pink to purple or bluish to violet, more rarely creamy to yellowish, claw of wings and keel only at base adnate to staminal tube. Standard (10-)12-28 mm long, not sharply differentiated into claw and blade, constricted at the middle, lower half narrower than upper. Wings slightly shorter than standard, blade narrowly oblong to narrowly obovate, claw as long as or longer than blade. Keel slightly shorter than wings, blades obovate-triangular, with widely to rectangular-curved lower edge and + straight upper edge. Ovary shortly stipitate, hairy; style hairy at the base or in lower half. Fruits bilocular, or only in A. hezarensis and A. ovigerus unilocular, laterally or 235 dorsi-ventrally compressed, elliptic-oblong to ovate-elliptic or widely elliptic. Seeds olive green to brown with black spots, rugose on surface, elliptic-kidney-shaped. Diagnostic key to the species of Astragalus sect. Acanthophace E Short lateral shoots present at the base of each leaf on the main stem though irregularly; paraclades borne on the short lateral shoots; calyx hairs exclusively white 8. A. sclerocladus Bunge Short lateral shoots absent; paraclades borne directly on the main stem; calyx hairs white and black, only in A. /ycioides some specimens with exclusively white hairy calyx 2 Ovary and fruits unilocular; mature fruits 15-38 mm long 3 Ovary and fruits bilocular; mature fruits 7-15 mm long 4 Mature fruits 15-18 mm long; leaflets 5-7 pairs; calyx greyish green to creamy 5. A. hezarensis Zarre & Podlech Mature fruits 30-38 mm long; leaflets 7-10(-15) pairs; calyx often flushed with red 7. A. ovigerus Boiss. Young stems + glabrous; stipules glabrous or only at margins ciliate 5 Young stems densely hairy; stipules hairy allover 7. Bracts linear-subulate, 1.5-2.5 mm long and 0.5-1 mm wide 9. A. stenostegius Boiss. & Hausskn. Bracts ovate, 2-4 mm long and 1.5-2.5 mm wide 6 Leaflets in 4-9 pairs, sparsely to loosely covered on underside with ascending hairs; bracts 2-4 mm long; calyx sparsely to loosely covered with ascending white and black hairs; standard 12-17 mm long; legumes 8-12 mm long 4. A. horridus Boiss. Leaflets in 6-11 pairs, glabrous or with few appressed hairs at the midvein; bracts 1-2.5 mm long; calyx very sparsely covered with strongly appressed black hairs; standard 16-21 mm long: Legumes 6-7 mm long 1. A. cryptocarpos DC. Calyx teeth 1/2—1/1 as long as the tube; standard 11-14 mm long 3. A. hemsleyi Aitch. & Hemsl. Calyx teeth 1/5—1/3 as long as the tube; standard 14-25 mm long 8 Fruits laterally compressed, 2-3 mm wide; young rhachides dense. Standard glabrous 6. A. lycioides Boiss. Fruits dorsi-ventrally compressed, 5—9 mm wide; young rhachides remote. Standard hairy along the midline on dorsal surface 2. A. cystosus Zarre & Podlech The species Astragalus cryptocarpos DC., Astragal.: 187. 1802. Holotype: “Tragacantha orientalis humilli- ma foliis viciae, costae purpureae innascentibus, Tournefort cor. 29, Armenia” (P (hb. Vaillant!; iso: G-DC!, LE (fragm.)! [all without flowers and fruits]. Epitype (here designated): Turkey, [B9 Bitlis] Hanemir Da, N Seite von Oboskü Köyü aus, 2320 m, 10.8.1987, Engel 108a (MSB!). = Astragalus rechingeri Sirj., Ann. Naturhist. Mus. Wien 49: 269. 1939. Holotype: Kurdistan, Agherov Da. (S der Ebene von Pesandasht, 2900 m, 24.6.1936, Frödin 77 (UPS!: photo K; iso: W!: photo MSB!). Dwarf cushion-forming shrublets, 15-20 cm tall, densely branched at the base, mostly very sparsely furnished with white and black hairs 0.2-1 mm long. Stems from a prostrate base ascending, older parts without remnants of last years stipules and rachides, parts of the current year 1-6 cm long, in first year 1-2.5 mm diam., glabrous below stipules. Stipules whitish, chartaceous, 3-6 mm long, adnate to the petiole for 2-3 mm, otherwise free from each other, triangular- acuminate, glabrous or sometimes with few hairs at the margins. Leaves 1.4-7 cm long; rhachides crowded, + thin, flexible, straight or curved, glabrous or with few appressed white hairs; terminal spine 1/3-1/l as long as the uppermost leaflets; petiole 0.6-2.2 cm long (ca. 1/3-1/2 as long as rhachis). Leaflets in 6-11 pairs, green, 3-8 x 0.7-1.5 mm, narrowly oblong, or rarely narrowly obovate, mostly folded, at the apex acute or obtuse, minutely mucronulate, glabrous or on underside with few appressed hairs at the midrib. Peduncles 0.2-0.5 cm long, sparsely covered with some black or rarely mixed with white, + spreading hairs. Racemes loosely 2-5-flowered; flowering axis 0.2-0.5 cm long. Bracts chartaceous, whitish, 1-2.5 mm long and 0.5-1 mm wide, narrowly triangular-acuminate, deciduous, glabrous or sometimes with few tubercle-based hairs at margins. Pedicels 1-2 mm long, with few subappressed black hairs. Bracteoles absent. Calyx dark grey or brownish becoming creamy, tubular, ruptured by the pod, 6.5-10 mm long and 1.5-3.5 mm diam., very sparsely covered with strongly appressed black hairs, at the teeth more densely hairy and with some white hairs mixed in; teeth (1.5—)3 mm long, subulate. Petals pinkish-white, keel sometimes with bluish violet blades. Standard 16-21 mm long and 5-8 mm wide, without clearly differentiated claw, oblong, shallowly constricted at middle, lower part as wide as the upper part, retuse at the apex. Wings 14-18 mm long; blade narrowly oblong, at the apex obtuse or very minutely mucronulate, 6.5-10 x 2-3 mm; auricle 0.7-1 mm long, claw 8.5—9 mm long. Keel 13-15 mm long; blades ovate-oblong or rarely elliptic, 5-6 x 2.5-3 mm; claw 8.5-9 mm long. Ovary very shortly stipitate, densely sericeous except for base, ventral and dorsal edges; style glabrous. Legumes shortly stipitate, obliquely elliptic with straight ventral edge and strongly curved dorsal edge, 6-7 mm long, 2.5-4 mm wide and 3-5.5 mm high, laterally compressed, with a straight beak 0.5—1 mm long, bilocular; valves sparsely appressed hairy, becoming glabrous. Seeds immature, single in each locule. Distribution: E Turkey. Specimens seen: Turkey: Prov. Bitlis/Van: mt. 10 km SE Pelli, 2450 m, 8.7.1954, Davis & Polunin D.22582 (BM, E). — Prov. Van: distr. Satak, Kavussahap Dag, 2750 m, 27.7.1954, Davis & Polunin D.23075 (BM, E) — Tatvan to Geva, Kusgunkiran Gegidi, 2250 m, 11.8.1987, Engel 110 (MSB). Note: The type material of A. cryptocarpos both in G-DC and P consists only of some sterile shoots, without flowers and fruits. That is the reason why this species was treated as doubtful by BUNGE (1867/68) and Boıssıer (1872). The original description was most probably prepared on the basis of a poor specimen with only one flower and pod. Since the type material associated with this name is ambiguous, it was necessary to select an epitype (see GREUTER & al. 1994, article 9.7) congruent with the original description and material of this name. After typification it became clear that A. rechingeri must be reduced to synonymy under this species. The latter was treated as a member of sect. Acanthophace in Flora of Turkey by CHAMBERLAIN & MATTHEWS (in Davis, Flora of Turkey 1970) and has been transferred into sect. Acidodes Bunge (a synonym of sect. Adiaspastus Bunge, see ZARRE 2000) by DEML (1972). The type material of A. rechingeri is a young plant without any mature pod. But the new collections of this species by Engel near the type locality confirmed that this species has bilocular fruits, and therefore belongs to the sect. Acanthophace sensu DEML (1972) and not to sect. Acidodes. Bilocular pods with only one seed in each locule and almost glabrous leaves and stipules are the features which were cited as characteristic in the original descriptions of both A. cryptocarpos and A. rechingeri. Moreover, there is no other species of A. sect. Acanthophace distributed in Turkey. Therefore, the synonymy of A. rechingeri under A. cryp- tocarpos is well confirmed. The type material of A. rechingeri as well as other specimens belonging to A. cryptocarpos show the indumentum type typical for A. sect. Acanthophace: the hairs are appressed, ribbon- like and densely papillose. As it was mentioned recently (ZARRE 2000), such indumentum is also characteristic for subtribe Coluteinae of the tribe Galegeae. Astragalus cystosus Zarre & Podlech, spec. nov. Holotype: Iran, Prov. Khorassan, Montes Yoghatay in declivibus 25 km N Sabzevar, 1500-1600 m, 16.6.1975, K.H. Rechinger 53642 (M!; iso: W!). Figure: Fig. 1. Valde affinis A. /ycioides sed differt leguminis dorsi-ventraliter (nec lateraliter) compressis, 5-9 mm (nec 2-3 mm) latis et rhachidibus remotis (nec densis). Fruticulosus, spinosus, 15-35 cm altus, pilis subbasifixis, albis nigrisque 0,1-1,5 mm longis. Stipulae chartaceae, 1-4 mm longae, triangulares. Folia 1,3—6,5 cm longa; rhachides + remotae, dense sericeae. Foliola 4-9 juga, 1-4 mm longa et 2-5 mm lata, obovata, retusa, appresse sericea. Racemi pedunculo 0,3-2,2 cm longo, dense appresse pilosi suffulti, laxe 2-6 (-9)-flori. Bracteae chartaceae, anguste triangulares, 1-2 mm longae, dense appresse piloseae. Pedicelli ca. 2 mm longi. Bracteoleae saepe evolutae, minutae. Calyx tubulosus, flavidi- rubellus, 8-13 mm longus, sparse ad dense appresse pilosus. Petala rosea ad violacea. 238 Vexillum 16-25 mm longum et 5-9 mm latum. Legumina coriacea, bilocularia, valde inflata, dorsi-ventraliter compressa, 10-15 mm longa et 5-9 mm lata. Dwarf spiny subshrubs, 15-35 cm tall, densely or rarely loosely branched at the base, furnished with mostly flattened and ribbon-like, in vegetative parts white, in inflorescences mostly also black hairs 0.1-1.5 mm long. Stems 1.5—2.5 mm in diam. in first year, densely appressed hairy, gradually glabrescent. Stipules chartaceous, yellowish, mostly reflexed from the middle, 1-4 mm long, adnate to the petiole for ca. 1/2-2/3 their length, triangulate or + oblong-acuminate, densely hairy, partly glabrescent. Leaves 1.3-6.5 cm long; rhachides + remote, thin, rigid, densely hairy or partly glabrescent, terminal spine 2-5 times as long as the uppermost leaflets; petiole ca. 1/3 as long as the rhachis. Leaflets in 4-9 pairs, mostly complicate, rarely flattened, greyish green to green, 1-4 x 1-2.5 mm, obovate, at the apex shallowly retuse, on both sides sparsely to densely appressed hairy, partly glabrescent. Peduncles 0.3-2.2 cm long, densely appressed hairy. Racemes loosely 2-6(-9)-flowered. Bracts narrowly triangular, 1-2 mm long, chartaceous, densely appressed hairy. Pedicels ca. 2 mm long, appressed hairy. Bracteoles mostly present at the base of calyx, single or rarely in pairs, ca. | mm long, subulate, hairy. Calyx yellowish, sometimes flushed with red, 8-13 mm long and 2-3 mm in diam., sparsely to densely covered with appressed or rarely ascending hairs; teeth 1.5-3 mm long, mostly unequal, subulate. Petals pink to violet. Standard hairy on midrib at the dorsal-abaxial surface, 16-25 mm long and 5-9 mm wide, without sharply differentiated claw, + elliptic-pandurate, shallowly constricted at the middle, at the apex retuse. Wings slightly shorter than standard; blade narrowly oblong, obtuse at the apex, 7-9 x 2 mm; auricle 0.5—0.8 mm long. Keel 14-19 mm long; blades 5.5-8 x 2-2.5 mm. Legumes with a stipe ca. 1 mm long, 10-15 mm long, 5-9 mm wide and 3-6 mm high, dorsi-ventrally compressed, bilocular; valves coriaceous. Seeds 4-12. Distribution: NE Iran. Specimens seen: Iran. Prov. Khorasan: Esfarayen versus Sabzevar, 20 km from Sabzevar, 1600-1700 m, 27.6.1975, Termeh s.n. (IRAN) — Montes Yoghatay in declivibus 25 km N Sabzevar, 1500-1600 m, 16.6.1975, Rechinger 53642 (M, W). Note: The material desribed here as the new species “A. cystosus” was attributed to A. pseudoangustifolius Sir}. & Rech.f. by MAASSOUMI (1995). Although the specimen cited as the reference for such a treatment (Assadi & Mozaffarian 35753 in TARI) was also collected from Kuh-e Bizg, like A. pseudoangustifolius, it differs in some respects from the type material of the latter species. The most important difference in vegetative parts between A. cystosus and the type of A. pseudoangustifolius is the loose habit of the former and its remotely positioned rhachides. Since the type material of A. pseudoangustifolius is young and lacks any fruit, an epitype was selected for this specimen (see under A. lycioides). This epitype falls in all respects, especially with its laterally compressed fruits, within the variability of A. /ycioides. Therefore, A. pseudoangustifolius is reduced here to synonymy under A. /ycioides. So, there was no name for the species with largely inflated and dorsi-ventrally compressed fruits and it is therefore newly described here. 239 Fig. 1. Astragalus cystosus (after Termeh s.n.): — a: the whole plant, b: rhachis segment with leaflets, c: calyx, d: standard, e: wing, f: keel, g: staminal tube, h: fruit. Astragalus hemsleyi Aitch. & Baker, J. Linn. Soc., Bot. 19: 158. 1882 = A. canispinus Boiss., Fl. Or. Suppl.: 181. 1888, illeg. [homotypic with A. hemsleyi]. Syntypes: [Afghanistan] Hariab District, 1880, J.E.T. Aitchison 61/80; dto., Sergal to Biankhel, 5.6.1879, J.E.T. Aitchison 1215 (K!). Lectotype (ALI 1961): Hariab District, Afghanistan, 1880, J.E.T. Aitchison 80 (K!; iso sub 61/80: BM!, C!, G-BOIS!, K!, LE!, P!). 240 Figures: RECHINGER, K.H.: Symbolae Afghanicae 3 — Biol. Skr. 9(3): Fig. 131, 132. 1957; DEML, I.: Revision der Sektionen Acanthophace Bunge und Aegacantha Bunge der Gattung Astragalus L. — Boissiera 21: p. 185, 186, figs. 1,2 & 25. 1972. Dwarf spiny subshrubs, 15—25 cm tall, densely or rarely loosely branched at the base with white, only in the calyx also with greyish-black hairs 0.1-0.6 mm long. Stem 1.5-2.5 mm diam. in first year, densely appressed sericeous, gradually glabrescent. Stipules chartaceous, yellowish, 2-4 mm long, adnate to the petiole for ca. 1/2 of their length, triangular or + oblong- acuminate, sparsely hairy, soon glabrescent, ciliate at margins. Leaves 2-5 cm long; rhachides + dense, thin, rigid, densely hairy, glabrescent with age; terminal spine 24 times as long as the uppermost leaflets; petiole ca. 1/5-1/3 as long as the rhachis. Leaflets in 6-9 pairs, slightly complicate or flat, greyish green to green, 1-3 x 1-2 mm, widely obovate to nearly orbicular, shallowly retuse at the apex, on upper side glabrous, on underside sparsely adpressed hairy, especially at midvein and margins. Peduncles 0.1-0.2 cm long, hairy. Racemes loosely 2—4- flowered. Bracts chartaceous, narrowly triangular, 1-2 mm long, appressed hairy. Pedicel 1-2.5 mm long, hairy. Bracteoles sometimes present at the base of calyx, single or rarely in pairs, 0.2-1 mm long, subulate, hairy. Calyx yellowish-white, mostly flushed with red, 6-9 mm long and 2-3 mm in diam., sparsely to densely covered with appressed to spreading hairs; teeth 2-4.5 mm long, mostly unequal, subulate. Petals pink to red. Standard 10-14 mm long and ca. 5 mm wide, without sharply differentiated claw, + elliptic-pandurate, shallowly constricted at middle, retuse at the apex. Wings slightly shorter than standard; blade narrowly oblong, obtuse at the apex, 5-6.5 x 2 mm; auricle 0.3-0.4 mm long. Keel 9-10 mm long; blades ca. 4-5 x 2 mm. Legumes with a stipe ca. 0.5 mm long, narrowly elliptic, ca. 8 mm long and 3.5 mm high, laterally compressed, bilocular; valves coriaceous, + densely covered with appressed to ascending hairs. Seeds 3-8. Distribution: Afghanistan and Pakistan. Specimens seen: Afghanistan. Prov. Paktia: Latakary Pass zwischen Alikhel und Dobandi, 3300 m, 5.7.1991, Volk 71/489 (MSB) — oberhalb Qasim Khel, 3250 m, 24.6.1969, Freitag 5772 (Hb. Freitag) — Hariab District, 1880, Aitchison 61/80 (BM, G-BOIS, K, LE, P) — Sergal to Biankhel, 5.6.1979, Aitchison 1215 (K). Pakistan. Kurram: in valle Kurram, 2700-3000 m, Harsukh 14830 (E). — Quetta: Urak-Tal, 12 Meilen W. Quetta, V.1958, Repp (W) — Ziarat, 2440 m, 18.5.1989, Lace (E). — Kalat: in monte Harboi pr. Kalat, 2700 m, R.R. Stewart 772 (E). ??: Surkai Zangy, 19.5.1956, Peddie 124 (K). Note: This species is endemic around the Afghanistan-Pakistan border. Another thorny species of Astragalus which shows a similar distribution pattern is A. diopogon from sect. Megalocystis. Since the fruits of the latter sometimes rupture the calyx (see TIETZ & ZARRE 1994), a feature characterizing A. sect. Acanthophace, it can be confused with A. hemsleyi. However, the exclusively white hairy calyx of A. diopogon (in contrast to the white and black hairy one in A. hemsleyi) and its relatively large bracts (3-9 mm, against 1-2 mm in A. hemsleyi) can be used as diagnostic characters for separating them. 241 Astragalus horridus Boiss., Diagn. pl. orient., ser. 2, 2: 66. 1843 = Tragacantha horrida (Boiss.) Kuntze, Revis. Gen. 2: 945. 1891. Syntypes: Asia minor, Aucher 1253 (P!); Persia prope Ispahan, Aucher-Eloy 1280; Persia occidentali,. Aucher-Eloy 1281 (G-BOIS!: photo M!, K!: foto M!, P!); [Iran] in montibus Ghilani, Aucher-Eloy 4394 (LE!, P!). Lectotype (DEML 1972): Persia prope Ispahan,. Aucher-Eloy 1280 (G-BOIS!: foto MSB!; iso: G!, K!: foto M!, LE!, OXF!, P!). = A. chionobius Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11 (16): 46. 1868 [et l.c. 15(1): 74. 1869] = Tragacantha chionobia (Boiss.) Kuntze, Revis. Gen. 2: 944. 1891. Lectotype (PODLECH 1998): Persia, Kuh-Delu, 10.6.1842, Kotschy 474 (P!: foto MSB!; iso: G!, G-BOIS!: foto MSB, K!, M!, P!, ZT!). = A. chionobius var. hirtus Boiss., Fl. Or. 2: 313. 1872. Lectotype (PODLECH 1998): [Iran] Esfahan, ad nives Kellal et Sebse Kuh, 1868, H.C. Haussknecht (G-BOIS!; iso:G!, K!, W!). Figures: DEML, I.: Revision der Sektionen Acanthophace Bunge und Aegacantha Bunge der Gattung Astragalus L. — Boissiera 21: p. 183, 186, Fig. 9-12, 16, 17, 22, 23. 1972. Dwarf spiny subshrubs, 15-35 cm tall, densely or loosely branched at the base with in vegetative parts white, in the inflorescenses mostly also with black hairs 0.1-1 mm long. Stems 1.5-3 mm diam. in first year, glabrous. Stipules membranous, yellowish-white, 3-6 mm long, adnate to the petiole for ca. 2/3 of their length, partly or completely clasping the stem, mostly ovate-acuminate, ciliate at margins and tip, glabrous otherwise. Leaves 1.7-6.5 cm long; rhachides + dense, thin, rigid, sparsely to loosely furnished with appressed to ascending hairs, glabrescent with age; terminal spine 2-5 times as long as the uppermost leaflets; petiole ca. 1/3 as long as the rhachis. Leaflets in 4-9 pairs, slightly complicate to flat, green, 3-7 x 1-2.5 mm, narrowly obovate, acutish to shallowly retuse at the apex, on upper side hairy only at the margins, on underside very sparsely to loosely furnished with ascending hairs. Peduncles 0.2-0.7 cm long, appressed hairy. Racemes loosely 4-9-flowered. Bracts whitish, membranous, 2-4 mm long and 1.5-2.5 mm wide, ovate, sparsely hairy. Pedicels 1-2 mm long, hairy. Bracteoles sometimes present, at the base of calyx, single or rarely in pairs, 1-2.5 mm long, ciliate. Calyx yellowish, mostly flushed with red, 7-12 mm long and 2-5 mm in diam., sparsely to loosely covered with appressed to ascending black and white hairs; teeth 1.5-3(-4) mm long, mostly unequal, subulate or linear. Petals pink to purple. Standard 12-17 mm long and 4.5-7 mm wide, without clearly differentiated claw, elliptic to + elliptic- pandurate, somtimes shallowly constricted at middle, retuse at the apex. Wings 11-15 mm long; blade narrowly oblong, obtuse at the apex, 5.5-7 x 2 mm wide; auricle ca. 0.5-0.8 mm long. Keel 10-14 mm long; blades + semicircular,with widely curved lower edge and straight upper edge, 4-6 x 2-2.5 mm. Ovary densely hairy. Legumes with a stipe 0.5-1 mm long, narrowly elliptic, 8-12 mm long and 3-4 mm high, laterally compressed, bilocular; valves coriaceous. Seeds 4-8. Distribution: C, W and SW Iran. Specimens seen: Iran. Prov. Boyer Ahmadi (Kohgiluyeh): Kuh Dena, 3600 m, ix.1955, Remaudiére 5242 (IRAN). — Prov. Chahar Mahal Bakhtiari: Zard-Kuh, above Kuhrang valley, 4265 m, 5.8.1966, Archibald 2957 (E, K) - Zard Kuh, 3500-4000 m, 10.8.-3.9.1975, Carls (W) — Gandomkar, 242 Kuh-e Garreh, 2300-2700 m, 9.6.1973, Iranshahr & Moussavi 15557 (IRAN, W) — Ardal, Zard Kuh, 2700-3200 m, 14.-15.6.1973, /ranshahr & Moussavi 15599 (W) — Kouhrang Dam, versus Sarcheshmeh Karoun, 2300-2400 m, 18.6.1973, /ranshahr & Moussavi 15603 (IRAN, W) — Dul- Kala, near Kuhrang, c. 2500 m, 7.6.1966, Zhumer 1035 (BG). — Prov. Esfahan: prope Ispahan,. Aucher-Eloy 1280 (G, G-BOIS, K, LE, OXF, P) — Qashqai region, Kuh-e Surmandeh (Kuh-e Alijug), Semirom, in declivibus boreo-orientalibus, 2700-3900 m, 7.6.1974, Rechinger 47550 (M, W). — Prov. Fars: Kuh-Delu, 10.6.1842, Kotschy 474 (G, G-BOIS, K, M, P, ZT). — Prov. Lorestan: in mte. Schuturunkuh, VII.1908, Strauss (B) & VIII.1903, Strauss (B, K). — Prov. Markazi: Sultanabad, in monten Raswend, 15.7.1892, Strauss (B) — dto., VIII.1899, Strauss (B) — ibid., 26.7.1903, Strauss (B) — dto., 4.7.1909, Strauss (B). — Not traced: Djoubulak, IX.1898, Strauss (B). Note: A. horridus is endemic to the central Zagros mountain range. Membranous stipules are one of the important diagnostic characters of this species, a character which relates it to A. chartostegius Boiss. & Hausskn. of A. sect. Adiaspastus. Interestingly, the latter is also distributed over the same area, though more frequent. The bilocular ovary and fruit of A. horridus (against the unilocular one in A. chartostegius) is an important feature which can be used for separating them. The specimen Rechinger 47550 differs from other examined material in having spreading- hairy leaves. Moreover, the type of indumentum on calyx shows a considerable variability in this species: some specimens have spreading-hairy calyx and other appressed ones. A similar situation we have regarding A. lycioides with respect to A. schistocalyx. The specimen /ranshahr & Moussavi 15603 differs from other specimens in having a yellowish white corolla and calyx. It is most probably only an albino morph and cannot represent a new taxon. Astragalus hezarensis Zarre & Podlech, spec. nov. Holotype: Iran, Prov. Kerman, Kuh-e Hezar, 20 km SW Rayen, NE ridge leading to summit, 3800 m, 1.6.1977, J.R. Edmonson & A.G. Miller 1574 (W!; iso: E!). Figures: fig. 2. Affinis A. ovigero sed differt leguminis brevioribus, 15-18 mm (nec 30-38 mm) longis, foliolis 5-7 jugis (nec 7-10 jugis)et calyce griseo-viridi vel lacteo (nec rubello); ab A. lalesarensis (sect. Megalocystis) pilis albi-nigrisque (nec tantum albis) obtectus et calyce griseo-viridi vel lacteo (nec rubello); ab A. sahendi (sect. Adiaspastus) stipulis 5-8 mm (nec 13-17 mm) longis, foliolis 1.5-6 mm (nec 13-17 mm) longis, bracteis 3-4 mm (nec 7-10 mm) longis et leguminibus 11-18 mm (nec 9-11 mm) longis. Plantae suffruticosae, 15-25 cm altae, pilis basifixis albi-nigrisque, 0,5-1,5 mm longis instructae. Stipulae chartacae, 5-8 mm longae, late ovatae, ciliatae. Folia ad 5,5 cm longa. Foliola 5-7-juga, ad 6 mm longa et 3 mm lata, obovato-elliptica, plerumque plana, sparse ad dense appresse sericea. Racemi pedunculo 0,3-1 cm longo suffulti, laxe 1—2-flori. Bracteae membranaceae, 3-4 mm longae et ca. 1,5 mm latae. Bracteoleae minutae. Calyces 11-15 mm longi et 3-5 mm diam., pilis patentibus densiter obtecti, dentibus 2-3 mm longis. Vexillum 243 22-27 mm longum et 9-12 mm latum. Legumina a latere compressa, coriacea, 11-18 mm longa et 5-8 mm alta, unilocularia. Dwarf spiny subshrubs, 15-25 cm tall, + loosely branched at base, with mostly flattened, rarely cylindrical, acute white or in the inflorescences mostly also with black hairs 0.1-1.5 mm long. Stems 1-3 mm diam. in first year, densely appressed hairy, soon glabrescent. Stipules chartaceous, yellowish-white, 5-8 mm long, adnate to the petiole for 1/2—2/3 of their length, widely ovate, shortly acuminate, ciliate at margins and tip, glabrous otherwise. Leaves 1.1-5.5 cm long; rhachides + dense, thick, rigid, densely appressed hairy, later on partly glabrescent; terminal spine 1/3-1/10 as long as the uppermost leaflets; petiole 1/3-1/2 as long as the rhachis. Leaflets in 5-7 pairs, + flat, greyish green, 1.5—6 x 1-3 mm, obovate to elliptic, at the apex obtuse, with a minute mucro 0.2—0.8 mm long, on upper side + glabrous, on underside sparsely to loosely appressed hairy, soon glabrescent. Peduncles 0.3-1 cm long, densely hairy. Racemes loosely 1—2-flowered.. Bracts + membranous, 3-4 mm long and ca. 1.5 mm wide, narrowly triangular, sparsely hairy. Pedicels 2-3 mm long, hairy. Bracteoles 1.5-2.5 mm long, subulate. Calyx greyish green, sometimes obliquely gibbous at the base, 11-15 mm long and 3-5 mm diam., rather densely covered with spreading hairs; teeth 2-3 mm long, mostly unequal, from a triangular base subulate. Petals white (yellow when dried), keel with purple blades. Standard 22-27 mm long and 9-12 mm wide, without sharply differentiated claw, + obovate-elliptic, shallowly retuse at the apex. Wings 18-24 mm long; blade narrowly oblong-elliptic, obtuse at the apex, 9-13 x 2.5—4 mm; auricle ca. 0.6-1 mm long. Keel 15-19 mm long; blades oblong-obovate, 7-8 x 4 mm. Ovary shortly stipitate, style hairy in lower third. Legumes shortly stipitate, narrowly oblong-elliptic, 11-18 mm long and 5-7 mm high, laterally compressed, with a beak 2-3 mm long, unilocular; valves sparsely to densely hairy. Distribution: S Iran (only known from Hezar mountains around Kerman). Specimens seen: Iran. Prov. Kerman: Kuh-e Hezar, 20 km SW Rayen, NE ridge leading to summit, 3800 m, 1.6.1977, J.R. Edmonson & A.G. Miller 1574 (E, W) — Raien, Gharieh-ye Mir-Shady, Kuh-e Hezar, 3000-3450 m, 21.5.1976, Moussavi 35026 (IRAN, W). Note: A. hezarensis is closely related to A. lalesarensis which has been treated as a member of A. sect. Megalocystis (see TIETZ & ZARRE 1994). No fruiting specimen of A. lalesarensis was available to the earlier authors and it is possible that its fruit is also similar to those of A. hezarensis. However, the calyx in A. lalesarensis is distinctly inflated (a feature characteristic of A. sect. Megalocystis). A. hezarensis is a narrowly endemic species which occurs in the same area as A. lalesarensis. The possession of white-hairy, inflated calyces contrary to black and white hairy, non-inflated calyces separates A. lalesarensis from A. he- zarensis. The close relationship between these species is another evidence for affinity of A. sect. Acanthophace and A. sect. Megalocystis. Most probably A. sect. Megalocystis originated several times from earlier species of A. sect. Acanthophace and is therefore polyphyletic. A. hezarensis is also very closely related to A. sahendi of A. sect. Adiaspastus (for a detailed description see ZARRE, 2000), but differs from it in having shorter stipules and larger fruits. Moreover, A. hezarensis has papillose hairs, against A. sahendi with striate hairs. Aditionally, the fruits, stipules and bracts of A. sahendi are larger than those of A. hezarensis. 244 Fig. 2. Astragalus hezarensis: — a: the whole plant, b: rhachis segment with leaflets, c: calyx, d: bracts, e: standard, f: wing, g: keel, h: staminal tube, i: ovary, j: fruit. — After Moussavi 35026 (IRAN). 245 Astragalus lycioides Boiss., Diagn. pl. orient., ser. 1, 2: 66. 1843 = Tragacantha lycioides (Boiss.) Kuntze, Revis. Gen. 2: 946. 1891. Holotype: Persia prope Isfahan, Aucher-Eloy 1270 (G-BOIS!: foto M!; iso: G!, K!, P!). = A. dendridium Fisch., Bull. Soc. Imp. Naturalistes Moscou 26(2): 427. 1853. Syntypes: M. Sufi pr. Isfahan, G. Kapherr (H!); in M. Korud pr. Isfahan, Bode. Lectotype (designated here): in m. Korud pr. Isfahan, Bode (LE!; iso: G-BOIS!: foto MSB!). = A. leptacanthus Fisch., Bull. Soc. Imp. Naturalistes Moscou 26(2): 432. 1853 = Tragacantha leptacantha (Boiss.) Kuntze, Revis. Gen. 2: 946. 1891. Type: Persia, prope Ssof N Isfahan, 22.5.1847, F.A. Buhse 1450 (iso: G!, G-BOIS!: foto M!). = A. schistocalyx Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 46. 1868 [et l.c. 15(1): 75. 1869]. Syntypes: inter Nischapur et Mesched prov. Chorassan Persiae, fr. 13.7.1856 A. von Bunge & T. Bienert (L!, LE!, P!); fl. 12.6.1858, A. von Bunge & T. Bienert. Lectotype (PODLECH, 1998): [Iran] inter Nischapur et Mesched prov. Chorassan Persiae, fl., 12.6.1858, A. von Bunge & T. Bienert (P!; iso: G-BOIS!, GOET!, K!: photo M!, P!, W!). = A. syrtschensis Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 46. 1868 [et Lc. 15(1): 74. 1869]. Lectotype (designated here): [Iran] Syrtsch inter Chabbis et Kerman, iv.1859, A. von Bunge & T. Bienert (P!; iso: G-BOIS!, GOET!, K!: foto MSB!, L!, LE!, M!, P!: photo M!, W!). = A. schistocalyx var. bizgimontanus Sir}. & Rech.f., Ann. Naturhist. Mus. Wien 58: 63. 1951. Holotype: [Iran] Khorasan, in Monte Kuh-e Bizg, 2000 m, E. Gauba & Mirdamadi 527 (W!). Epitype (designated here): Iran, prov. Khorasan: in monte Kuh-e Bizg, 4.-6.7.1937, K.H. Rechinger 1442 (W!) = A. pseudoangustifolius Sirj. & Rech.f., Anz. Österr. Akad. Wiss., Math.-Naturwiss. Kl. 90: 119. 1953. Holotype: [Iran] Khorasan, Kuh-e Bizg near Turbat-e Shaik Djam, 17.6.1885, J.E.T. Aitchison 666 (C!; iso: K!, P!: photo M!, W!). Epitype (designated here): Iran, prov. Khorasan: in monte Kuh-e Bizg, 4.-6.7.1937, K.H. Rechinger 1442 (W!). Figures: FISCHER, F.B.: “Synopsis Astragalorum Tragacantharum” — Bull. Soc. Imp. Naturalistes Moscou 27, tab. J. 1854; DEML, I.: Revision der Sektionen Acanthophace Bunge und Aegacantha Bunge der Gattung Astragalus L. — Boissiera 21: p. 186, figs. 15, 26 (both as A. /ycioides) and p. 185, 186, fig. 3, 4 13, 14, 27, 28 (all as A. schistocalyx). 1972. Dwarf spiny subshrubs, 15-35 cm tall, densely or rarely loosely branched at the base with mostly flattened and ribbon-like, in vegetative parts white, in the inflorescence mostly white and black mixed hairs 0.1-1 mm long. Stems 1.5-2.5 mm diam. in first year, densely covered with appressed to ascending hairs 0.1-0.5 mm long, glabrescent with age to glabrous in the next year. Stipules chartaceous, yellowish, mostly deflexed from the middle, 1-4 mm long, adnate to the petiole for ca. 1/2-2/3 of their length, triangular or + oblong-acuminate, densely covered with appressed to ascending hairs. Leaves 1.3-6.5 cm long; rhachides + dense, thin, rigid, densely appressed hairy, gradually glabrescent; terminal spine 2-5 times as long as the uppermost leaflets; petiole ca. 1/3 as long as the rhachis. Leaflets in 4-9 pairs, mostly complicate, rarely flat, greyish green to green, 1-4 x 1-2.5 mm, obovate, retuse at the apex, on upperside glabrous or sparsely hairy, on underside more densely hairy. Peduncles 0.3-2.2 cm long, with appressed to ascending hairs. Racemes loosely 2-6(-9)-flowered. Bracts narrowly 246 triangular, 1-2 mm long, sparsely hairy or only ciliate at the margins. Pedicels ca. 2 mm long, hairy like the peduncle. Bracteoles mostly present at the base of calyx, single or rarely in pairs, 0.5—1.5 mm long, subulate, hairy. Calyx yellowish, often flushed with red, 8-13 mm long and 2-3 mm in diam., sparsely to rather densely covered with appressed to spreading white and greyish-black, rarely only with white hairs up to 1.2 mm long; teeth 1.5-3 mm long, mostly unequal, subulate. Petals pink to violet. Standard hairy on midrib at the dorsal surface, 16-25 mm long and 5-9 mm wide, without sharply differentiated claw, obovate to + elliptic- pandurate, mostly shallowly constricted at middle, retuse at the apex. Wings (12—)14—21 mm long; blades narrowly oblong, obtuse at the apex, 7-9 x 2 mm; auricle 0.5-0.8 mm long. Keel (11-)14-19 mm long; blades 5.5-8 x 2-2.5 mm. Legumes with a stipe ca. 1 mm long, narrowly elliptic, 10-11(-13) mm long, 2-3 wide and 3-6 mm high, laterally compressed, bilocular; valves coriaceous, densely covered with appressed to spreading hairs. Seeds 4-12. Distribution: NE, C and S Iran. Specimens seen: Iran. Prov. Esfahan: Karkas mts. S Natanz, 2500 m, 28.6.1973, Andersen & Jensen 7295 (E, L) — inter Isfahan et Teheran, Kohrud, V.1859, Bunge & Bienert (K, P) — S Kashan, Kuh-e Karkas, 27.5.1974, I/ranshahr s.n. (IRAN) — Natanz, 1500 m, 19.5.1975, Iranshahr 40962 (IRAN, W) — Natanz, above the village Abianeh, 2600 m, 29.7.1993, Maassoumi & al. 71919 (MSB) — before Meymeh, on the road of Soh (Sof), before Delhor village, 2100 m, 4.9.1993, Maassoumi & Zarre 72013 (MSB) — Kashan, Kamow, Kuh-e Gar-gash, 2400-3200 m, 5.7.1984, Moussavi & Tehrani 41584 (W) — Kuh-e Karkas, in saxosis prope stationem Radar, 2880 m, Rechinger 1974 (M, W) — Kashan, Natanz, 1500 m, Rechinger 52056 (M, W) — Taleghan, c. 30 km SW Ardestan, 2000 m, 15.5.1974, Wendelbo & Foroughi 11530 (W) — Kuh-e Karkas, near television mast N side of pass, N Tarq, 2700 m, 14.5.1974, Wendelbo & Foroughi 11428 (W) — Kuh-e Karkas area, just N Targ, 15.5.1974, Wendelbo & Foroughi 11456 (W) — prope Isfahan, Aucher-Eloy 1270 (G, G-BOIS, K, P) — in m. Korud pr. Isfahan, Bode (G-BOIS, LE) — prope Ssof N Isfahan, 22.5.1847, Buhse 1450 (G, G-BOIS). — Prov. Kerman: Kuh-e Jebal Barez, Kuh-e Sarzeh, 5 km NE Garraghan, E of road from Jiroft to Deh Bakri, 3000-3500 m, 13.6.1977, Assadi & al. 2131 (E) — in reg. alpina montis Kuh-i Dschupar, 3000-3400 m, 10.6.1892, Bornmiiller 3718 (B, WU) — in monte Kuh-i-Nasr, 3800 m, 4.7.1892, Bornmüller 3719 (B, G) - in monte Kuh-i Sirdsch, 2700 m, 22.5.1892, Bornmiiller 3720 (B, WU) — in monte Kuh-i Lalesar, 3800-3900 m, 17.7.1892, Bornmiiller 3721 (B) — dto., 2650-3000 m, 24.6.1976, Moussavi & Tehrani 36794 (IRAN, W) montes Kuh-e Jebal Barez in vicinitate vici Deh Bakri, 1700-2700 m, 2.5.1973, Sojak 3967 (PR) — Syrtsch inter Chäbbis et Kerman, iv.1859, Bunge & Bienert (G- BOIS, GOET, K, L, LE, M, P, W). — Prov. Khorasan: Neyshabur, Sheykh Abol-Hassan, Binaloud, 1500-2250 m, 30.-31.7.1976, Termeh & Tehrani 35139 & 35144 (IRAN, W) — in monte Kuh-e Bizg, 4.-6.7.1937, Rechinger 1442 (K, W) — in Monte Kuh-e Bizg, 2000 m, Gauba & Mirdamadi 527 (W) — Kuh-e Bizg near Turbat-e Shaik Djam, 17.6.1885, J.E.T. Aitchison 666 (C, K, P, W) — environs de Mughan et versant nord de la Kuh-i Binalud, 1900-2300 m, Schmid 6259 (E, W), 6260 (W) — montes Kuh-e Binalud, ca. 20 km bor.-orient. ab oppido Neyshabur, 2300-2800 m, 14.6.1977, Sojak 7662 (PR) — inter Nischapur et Mesched, fl., 12.6.1858, Bunge & Bienert (G-BOIS, GOET, K, P, W). — Prov. Yazd: Shirkuh, 20 km SSW Taft, 2700 m, 25.5.1977, Aryavand & al. 1427 (E) — Schir Kuh, 3960 m, 24.7.1932, Balls 138 (K) — Deh-Bala, Shirkuh mt., 3500 m, 21.6.1975, Foroughi & Assadi 17963 (G) — Taft, Gharieh-ye Hedesh, Deh- bala, Shir-kuh, 2800-3300 m, 10.6.1976, Moussavi & Tehrani 36789 (IRAN, W) — dto., 2300-2600 m, 12.6.1976, Moussavi & Tehrani s.n. (IRAN) — dto. 16.7.1976, Moussavi & Tehrani 35025 (IRAN, W) Note: The holotypes of both A. pseudoangustifolius and A. schistocalyx subsp. bizgi- montanus are very young plants without mature fruits. Since the pod structure is very important in A. sect. Acanthophace, it was necessary to select epitypes as reference material 247 for these species. The specimen Rechinger 1442 in W fits in all respects the description of both names and was collected like the others two from the famous NE Iranian mountain “Kuh- e Bizg”. Therefore Rechinger 1442 is selected here as an epitype for both the others. DEML (1972) gave three differences for separating A. schistocalyx from A. Iycioides: appressed hairs on calyx of A. schistocalyx (against spreading ones in the latter), peduncles 1—- 3 (-7) mm long (against 5-22 mm long in A. /ycioides), and racemes with 2-5 flowers (not up to 8 as in A. Jycioides). However the examined specimens show a high variability in all these characters, which does not support the separation of two species. For example the specimens collected from the area about Natanz and Ardestan (Prov. Esfahan), i.e. Andersen & Jensen 7295 (E, L), Rechinger 52056 (M, W) and Wendelbo & Foroughi 11530 (W), and also a specimen from Prov. Khorassan, namely Sojak 7662 (PR), possess long peduncles up to 12 mm and inflorescences with 8-12 flowers, but appressed calyx hairs. Moreover, some specimens collected from an area near the type locality of A. lycioides (Soh), i.e. Maassoumi & Zarre 72013 (M) and Wendelbo & Foroughi 11456 (W), show the subappressed type of calyx hairs and are in all other respects intermediate between A. /ycioides and A. schistocalyx. Therefore, the name A. schistocalyx is reduced here to synonymy under A. /ycioides. Astragalus ovigerus Boiss., Diagn. Pl. orient., ser. 1, 2: 67. 1843 = Phaca ovigera (Boiss.) Boiss., Diagn. Pl. orient., ser. 1, 6: 35. 1846. Holotype: Persiae alpibus Zerdkou, Aucher 1277 (G-BOIS!; iso: BM!, G!, K!, P!). = A. lamprocarpus Maassoumi (1994) 1995: Iran. J. Bot. 6: 199 (1994) 1995. Holotype: [Iran] Lorestan, Ghali Kuh (E of the pass on road Aligodarz-Shoulabad), 3100-3600 m, 1.7.1977, Runemark & Lazari 26506 (TARI!). Figures: DEML, I.: Revision der Sektionen Acanthophace Bunge und Aegacantha Bunge der Gattung Astragalus L. — Boissiera 21: p.185, 186, fig. 5, 22-23. 1972. Dwarf spiny subshrubs, 15-30 cm tall, densely branched at base, with mostly flattened, rarely cylindric-acute, in vegetative parts white, in the inflorescenses white and black or sometimes merely black hairs 0.1-1.5 mm long. Stems of the current year 1-3.5 mm diam., up to 1 cm long, appressed hairy, soon glabrescent. Stipules chartaceous, yellowish-white, 5-8 mm long, adnate to the petiole for 1/2-2/3 of their length, widely ovate-acuminate, densely appressed hairy. Leaves 2-9 cm long; rhachides + dense, thick, rigid, densely appressed hairy, gradually glabrescent, in the second year glabrous; terminal spine 1/1—3/1 as long as the uppermost leaflets; petiole 1/3-1/2 as long as the rhachis. Leaflets in 7-10(-15) pairs, + flat, greyish green, 1.5-9 x 1-3 mm, narrowly ovate to narrowly obovate or obovate to elliptic, at the apex obtuse with a minute mucro 0.2-0.8 mm long, on both sides loosely to densely covered with appressed to ascending hairs. Peduncles 1-3 cm long, densely hairy. Racemes loosely 1-3-flowered. Bracts chartaeous, 3-4 mm long and ca. 1.5 mm wide, narrowly triangular, hairy. Pedicels 2-4 mm long, predominantly black hairy. Bracteoles rarely present, ca. 2 mm long, subulate. Calyx greyish green, flushed with red, tubular, 11-14 mm long and 3-5 mm in diam., densely covered with spreading, predominantly black hairs; teeth 1.5-2.5 mm long, triangular. Petals reddish to violet. Standard 22-27 mm long and 8-12 mm wide, 248 without distinctly differentiated claw, + obovate-elliptic, scarcely constricted at middle, shallowly retuse at the apex. Wings 18-24 mm long; blade, narrowly oblong, obtuse at the apex, 9-13 x 2.5-4 mm; auricle 0.6-1 mm long. Keel 15-19 mm long; blades oblong-curved, obtuse at the apex, 7-8 x 4 mm. Staminal tube obliquely cut at the mouth, filaments with 6-7 mm long free ends.Ovary shortly stipitate, sparsely hairy; style hairy in lower third. Legumes ovoid, 30-37 mm long, carinate ventrally, shallowly grooved dorsally, shortly acuminate at the apex, sparsely hairy to glabrescent, unilocular. Specimens seen: Iran. Prov. Chahar Mahal Bakhtiari: Persiae alpibus Zerdkou, Aucher 1277 (BM, G, G-BOIS, K, P). — Prov. Lorestan: Kuh-i-Shuturan, 2440 m, 26.5.1941, Koelz 17824 (US) — Ghali Kuh (E of the pass on road Aligodarz-Shoulabad), 3100-3600 m, 1.7.1977, Runemark & Lazari 26506 (TARI!). Note: The holotype of A. /amprocarpus was only for a short time available for investigation and therefore no detailed analysis could be made. Nevertheless no differences could be found between it and A. ovigerus. Moreover, the specimen Koelz 17824 (US) was collected from the same area as the holotype of A. Jamprocarpus and agrees with A. ovigerus. In the original description (MAASSOUMI 1994), A. lamprocarpus is not compared with A. ovigerus, but it fits in all respects the description of the latter. A. ovigerus and A. hezarensis are the only species of A. sect. Acanthophace which possess unilocular pods. Moreover, their fruits represent the larger ones within the section. However, these features are not enough to separate them as a section distinct from A. sect. Acanthophace as MAAssouMI (1995) did with A. sect. Lamprocarpa. For a more detailed discussion see ZARRE (2000: page 22). A. ovigerus is a paleoendemic species which shows a disjunct distribution pattern. Moreover, the species is very rare in the distribution area and does not form dense populations. Astragalus sclerocladus Bunge, Mém. Acad. Imp. Sci. Saint Pétersbourg 11(16): 46. 1868 [et l.c. 15(1): 75. 1869] = Tragacantha scleroclada (Boiss.) Kuntze, Revis. Gen. 2: 947. 1891 = A. schistocalyx Bunge subsp. sclerocladus (Bunge) I.Deml, Boissiera 21: 34. 1972. Syntypes: env. de Kohrud, C.P. Bélanger 693 (P!); inter Sof et Kohrud, v.1859, A. von Bunge & T. Bienert. Lectotype (PODLECH 1998): [Iran] inter Sof et Kohrud [inter Isfahan et Teheran], v.1859, A. von Bunge & T. Bienert (P!: photo MSB!; iso: G-BOIS!, GOET!, K!: photo MSB!, L!, LE!, W!). Spiny shrubs, 20-60 cm tall, densely or rarely loosely branched at the base with mostly flattened and ribbon-like, only white hairs 0.1-0.8 mm long. Stems 1.5-3 mm diam. in the first year, densely appressed hairy, glabrescent with age. Stipules chartaceous, yellowish, mostly reflexed from the middle, 1-2 mm long, adnate to the petiole for ca. 1/2 of their length, widely triangular-acuminate, densely appressed hairy. Leaves 0.5-4 cm long; rhachides + remote, thick, rigid, densely hairy, sericeous, gradually glabrescent and glabrous in the second year; terminal spine 1-3 times as long as the uppermost leaflets; petiole ca. 1/4-1/3 as long as the thachis. Leaflets 3-6 pairs, mostly flat, greyish green to light green, 3-8 x 2-5 mm, obovate 249 to orbicular or subcordate, rarely narrowly obovate, retuse at the apex, sparsely appressed hairy, especially at the midrib and the margins, soon glabrescent. Peduncles 0.1-0.5 cm long, densely appressed hairy. Racemes loosely 2—6-flowered. Bracts narrowly triangular ovate, 1-2 mm long and 0.7-1 mm wide, chartaceous, densely appressed hairy. Pedicels 1-2 mm long, hairy. Bracteoles rarely present, in pairs at the pedicel, ca. 0.8 mm long, subulate, hairy. Calyx yellowish, mostly fushed with red in upper part, 6-9 mm long and 2-4 mm in diam., sparsely to rather densely appressed white hairy; teeth 1-3 mm long, mostly unequal, subulate. Petals pink to red. Standard 12-18 mm long and 5-6.5 mm wide, without sharply differentiated claw, + elliptic-pandurate, mostly shallowly constricted at middle, retuse at tip. Wings slightly shorter than standard; blade narrowly oblong, obtuse at the apex, 6-8.5 x 2 mm; auricle 0.4-0.6 mm long. Keel 10-14 mm long; blades 5-6.5 x 2-2.5 mm. Legumes shortly stipitate, elliptic, 6-8(-10) mm long and ca. 4 mm high, laterally compressed, bilocular; valves coriaceous, densely appressed hairy. Seeds 3-9. Specimens seen: Iran. Prov. Esfahan: Kuh Barsuk, Kohrud-Gebirge, 27.6.1906, Strauss (B) — In m. Kuh-i Kohrud, 10.5.1908, Bornmüller (B) — env. de Kohrud, C.P. Bélanger 693 (P) — inter Sof et Kohrud [inter Isfahan et Teheran], V.1859, Bunge & Bienert (G-BOIS, GOET, K, L, LE, P, W) - Kashan, Niassar, 23.5.1970, /ranshahr 14662 (IRAN) — ca. 2 km from deviation of Ghamsar, on the road to Ghohrud, 1800 m, 4.9.1993, Maassoumi & Zarre 72018 (MSB, TARI) - In m. Kashan, on the road to Ghohrud, 12 km to Ghohrud, 2150 m, 11.3.1990, Zarre 69116 (MSB, TARI). Note: A. sclerocladus is a narrowly endemic species known only from the submontane regions around Ghohrud village south of Kashan (centre of Iran). In this region A. sclerocladus forms relatively dense populations and is one of the dominant species. This species has been reduced to subspecies rank by DEML (1972), in spite of considerable differences between the two taxa. In ist vegetative form, A. sclerocladus is the largest species of A. sect. Acanthophace. Moreover, it is the only species of the section whose paraclades are borne on the short lateral branches and not directly on the main stem: a character which relates this species to A. sect. Poterion Bunge (another basal section within the thorny Astragali). Beside these features, the obovate and deeply retuse leaflets of A. sclerocladus separate it from A. Iycioides. Because of the above, MAASsouM!'s (1994) opinion in considering A. sclerocladus as a distinct species from A. lycioides is well supported. Astragalus stenostegius Boiss. & Hausskn., Fl. Or. 2: 314. 1872 = Tragacantha stenostegia (Boiss.) Kuntze, Revis. Gen. 2: 948. 1891. Lectotype (designated here; DEML 1972 as holotype): [Iran/Iraq] Mt. Avroman et Schahu Kurdistaniae, 1000-12000', VII.1867, H.C. Haussknecht (G-Boiss!: photo MSB!; iso: G-BOIS!: photo MSB!, JE!, K!, P!). = A. spinellus Boiss. & Hausskn., Fl. Or. 2: 315. 1872 = Tragacantha spinella (Boiss.) Kuntze, Revis. Gen. 2: 948. 1891. Holotype: [Iran/Iraq] Mt. Avroman et Schahu Carduchiae Persiae, VII.1867, H.C. Haussknecht (G-Boiss!: photo K, MSB!). Figures: DEML, I., Revision der Sektionen Acanthophace Bunge und Aegacantha Bunge der Gattung Astragalus L. — Boissiera 21: p. 185, 186, Fig. 6-8, 19 & 24. 1972. 250 Dwarf spiny subshrubs, 15-35 cm tall, densely or loosely branched at the base with mostly flattened, ribbon-like, in vegetative parts white, in the inflorescens mixed with black hairs 0.1-1 mm long. Stems 1.5-2.5 mm diam. in first year, glabrous. Stipules membranous, yellowish-white, 3-6 mm long, adnate to the petiole for ca. 1/2 of their length, narrowly triangular, ciliate at margins and apex, glabrous otherwise. Leaves 1.7-4 cm long; rhachides + dense, thin or rarely thick, rigid, densely covered with appressed to spreading hairs up to 0.8 mm long, glabrescent with age and glabrous in the next year; terminal spine 1-3 times as long as the uppermost leaflets; petiole ca. 1/3 as long as the rhachis. Leaflets in 4-7 pairs, slightly complicate to flat, green, 3-8(-10) x 1-2.5 mm, narrowly ovate to narrowly obovate, at the apex rounded to shallowly retuse, on both sides sparsely + appressed hairy. Peduncles 0.2-1 cm long, with appressed to spreading hairs. Racemes loosely 3—6-flowered. Bracts narrowly triangular to subulate, 1.5-2.5 mm long and 0.5-1 mm wide, membranous, sparsely white hairy. Pedicels 1-1.5 mm long, hairy. Bracteoles absent. Calyx yellowish, sometimes flushed with red, 5-9 mm long and 1.5-3 mm diam., tubular-campanulate, densely covered with spreading hairs up to | mm long; teeth 1.5-3 mm long, mostly unequal, subulate or linear. Petals pink to violet. Standard 10-13 mm long and 4-5 mm wide, without sharply differentiated claw, + elliptic-pandurate to obovate, mostly shallowly constricted at middle, retuse at the apex. Wings 8-11 mm long; blades narrowly oblong, obtuse at the apex, 4.5-6 x ca. 1.5 mm; auricle 0.3-0.6 mm long. Keel 7-9 mm long; blades ca. 4.5 x 2 mm. Legumes (immature), ca. 8-10 mm long, laterally compressed, bilocular. Specimens seen: Iran. Prov. Hamadan: In monte Elwend, VI.1902, Strauss (B). — Prov. Kordestan: Mt. Avroman et Schahu Kurdistaniae, 3050-3650 m, VII.1867, Haussknecht (G-BOIS, JE, K, P). — Prov. Lorestan: In m. Kuh-i Gerru, VII.1902, Strauss (B, JE) — dto., 2.8.1908, Strauss (B, JE, P, WU). — Prov. Markazi: in m. Raswend, VIII.1899, Strauss (B). Note: The collection of Strauss from Monte Elwend (B) differs from other specimens in having more robust and longer spines, larger calyx (9-13 mm long) and standard 20-24 mm long. Moreover, the calyx is exclusively black hairy in this specimen, in contrast to other examined sheets with high proportion of white hairs too. The shape and size of bracts, however, match the type of A. stenostegius. More material and studies are needed for the final systematic positioning of this specimen. Most probably it is a representative of a new taxon. DEML (1972) expressed her doubt on the identity of A. spinellus and A. stenostegius. The type material of both species, which was collected from the borders between Iran and Iraq and probably from the same locality, shows only one difference regarding the size of leaves which are very small in A. spinellus. The material collected by Strauss from Kuh-i Gerru (near Nehawend, W Iran) in both herbaria B and JE is intermediate in this respect and therefore A. spinellus is reduced here to synonymy. The present study was supported partially by the project number 512/1/438 “Biosystematical Studies in Tribe Galegeae (Fabaceae)” of the Tehran University. We thank the curators of various institutes for the loan of herbarium material. 251 References BoissIER, E. 1872: Flora orientalis vol. 2. Genf. BunGE, A. 1868-1869: Generis Astragali species gerontogeae. - Mém. Acad. Imp. Sci. Saint Pétersbourg, ser. 7, 11(16): 1-140 et 15(1): 1-245. Davis, P.H. (ed.) 1972: Flora of Turkey and the East Aegean Islands vol. 3. Edinburgh. DECANDOLLE, A.P. 1802: Astragalogia, nempe Astragali, Biserrulae et Oxytropidis, nec non Phaca, Coluteae et Lessertiae historia iconibus illustrata. Paris. DEML, I. 1972: Revision der Sektionen Acanthophace Bunge und Aegacantha Bunge der Gattung Astragalus L. — Boissiera 21: 1-235. GREUTER, W. & al. (eds.) 1994: International Code of Botanical Nomenclature (Tokyo Code), Adopted by the Fifteenth International Botanical Congress, Yokohama, August-September 1993. — Regnum Veg. vol. 131. HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990: Index herbariorum I. The herbaria of the world, ed. 8. — Regnum Veg. vol. 20. MAASSOUMI, A.A. (1994) 1995: Additiones to the genus Astragalus (Papilionaceae) in Iran. — Iran. J. Bot. 6: 197-214. PODLECH, D. 1982: Neue Aspekte zur Evolution und Gliederung der Gattung Astragalus L. (Leguminosae). — Mitt. Bot. Staatssamml. Miinchen 29: 461-494. — 1998: Typification of Astragalus species II. Species mainly of the herbaria of Paris (P) and Geneva (G). — Sendtnera 5: 265-263. TIETZ, S. & ZARRE, SH. 1994: Revision von Astragalus L. sect. Megalocystis Bunge (Fabaceae). — Sendtnera 2: 287-363. ZARRE, SH. 2000: Systematic Revision of Astragalus L. sect. Adiaspastus, sect. Macro- phyllium and sect. Pterophorus. — Englera 18: 1-205. Dr. Shahin ZARRE, Department of Biology, Faculty of Sciences, Tehran University, Enghelab Ave., Tehran, Iran. Prof. Dr. Dietrich PopDLEcH: Institut für Systematische Botanik der Ludwig-Maximilians- Universität München, Menzingerstr. 67, D-80638 München, Germany. b { Wh un ° i j , 7} BEN \ ot " nets | ne un re hl . Nur OW Pate ® 1) 4 | et ge eee koe ae CO ee A a u LM Pi ne. u u N A in Pie th) wt hel am KLEIDER ; " Cae N ae ay id , : I i ‘ af pe ‚ ave ra m We oui ot DEN a VIE T EEE ad la nth zer I2 : j u | of N ur Er Lh wi kg Dy te Dr VAR a | : ” vile oh vy Nee Maser: oe LP een & u dal 7 _s net Me eer anion a, a . pi aren Bein. | Ma toe ger up Nu ah | Ze 253 New taxa and new combinations published in Sendtnera 7 (31.8.2001) Aaronsohnia pubescens (Desf.) Bremer & Humphries subsp. maroccana (Ball) Förther & Podlech, comb. nov. Astragalus angorensis Podlech, nom. nov. Astragalus anguranensis Podlech & Maassoumi, spec. nov. Astragalus argentocalyx [Ali ex] Podlech, spec. nov. Astragalus austrotibetanus Podlech & L.R.Xu, spec. nov. Astragalus bashmaghensis Maassoumi & Podlech, spec. nov. Astragalus behbehanensis Podlech, spec. nov. Astragalus bingoellensis Podlech, spec. nov. Astragalus bozakmanii Podlech, spec. nov. Astragalus bradosticus Maassoumi & Podlech, spec. nov. Astragalus cobresiiphila Podlech & L.R.Xu, spec. nov. Astragalus conaensis Podlech & L.R.Xu, spec. nov. Astragalus cystosus Zarre & Podlech, spec. nov. Astragalus damxungensis Podlech & L.R.Xu, spec. nov. Astragalus dickorei Podlech & L.R.Xu, spec. nov. Astragalus doabensis Podlech, spec. nov. Astragalus doghrunensis Maassoumi & Podlech, spec. nov. Astragalus fruticulosus Podlech, spec. nov. Astragalus hezarensis Zarre & Podlech, spec. nov. Astragalus keredjensis Podlech, nom. nov. Astragalus L. sect. Pseudotapinodes Podlech & L.R.Xu, sect. nov. Astragalus lanzhouensis Podlech & L.-R.Xu, spec. nov. Astragalus maowensis Podlech, & L.R.Xu, nom. nov. Astragalus marivanensis Podlech & Maassoumi, spec. nov. Astragalus montis-karkasii Podlech, spec. nov. Astragalus obtusifoliolus (S.B.Ho) Podlech & L.R.Xu, comb. nov. Astragalus olurensis Podlech, spec. nov. Astragalus owirensis [Ali ex] Podlech, spec. nov. Astragalus parrisii Podlech, nom. nov. Astragalus poluninii Podlech, spec. nov. Astragalus pravitzii Podlech, spec. nov. Astragalus pseudopersicus Podlech & Maassoumi, spec. nov. Astragalus pseudotauricola (Ponert) Podlech, comb. et stat. nov. Astragalus goturensis Podlech, spec. nov. Astragalus rasmontii Podlech, spec. nov. Astragalus recurvatus Podlech, spec. nov. Astragalus rhododendrophila Podlech & L.R.Xu, spec. nov. Astragalus saadatabadensis Podlech, spec. nov. Astragalus sata-kandaoensis Podlech, spec. nov. Astragalus sherriffii Podlech, spec. nov. Astragalus spitzenbergeri Podlech, spec. nov. Astragalus tecti-mundi Freyn subsp. orientalis Podlech, subsp. nov. Astragalus tibeticola Podlech, spec. nov. Astragalus tsangpoensis Podlech & L.R.Xu, spec. nov. Astragalus wakhanicus Podlech, spec. nov. Astragalus zaraensis Podlech, spec. nov. Centaurea pungens Pomel subsp. austromaroccana Forther & Podlech, subspec. nov. Drimia maritima (L.) W.T.Stearn subsp. maura (Maire) Férther & Podlech, comb. nov. Erodium maroccanum (Maire) Förther & Podlech, comb. nov. Hyacinthoides hispanica (Mill.) Rothm. subsp. algeriensis (Batt.) Förther & Podlech, comb. et stat. nov. 254 Limonium bollei (Wangerin) Erben, comb. nov. Limonium capitis-eliae Erben, spec. nov. Limonium caralitanum Erben, spec. nov. Limonium carisae Erben, spec. nov. Limonium comosum Erben, spec. nov. Limonium fesianum Erben, spec. nov. Limonium malfatanicum Erben, spec. nov. Limonium multifurcatum Erben, spec. nov. Limonium portovecchiense Erben, spec. nov. Passiflora tina R.Boender & T.Ulmer, spec. nov. Pulicaria antiatlantica Förther & Podlech, spec. nov. Stigmidium lichenum (Arnold) Triebel & P.Scholz, comb. nov. Zephyra compacta C.Ehrhart, spec. nov. Hinweise für die Autoren + Die Sendtnera (früher Mitteilungen der Botanischen Staatssammlung München) veröffentlicht wissenschaftliche Originalarbeiten und Kurzmitteilungen in deutscher und englischer Sprache (andere Sprachen nur nach Rücksprache) aus dem Gesamtgebiet der Systematischen Botanik. 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Tie at et i LOLA 3 5185 0 57 Inhalt BOENDER, R. & ULMER, T.: Passiflora tina spec. nov., a new species of Passiflora, subgenus Astrophea (Passifloraceae) from Beuador. ......... 02,2... 000.202 DAVIES, A.M.R. & FACHER, E.: Achene hairs and their diversity in the genus Chaetanthera Ruiz & Pav. (Mutisieae, Asteraceae)... 2.20... nee DÖBBELER, P.: Polytrichadelphus magellanicus sensu lato (Musci) and its asco- mycetes — different fungi on different hosts........:.:...... sess ee EHRHART, C.: Zephyra compacta (Tecophilaeaceae) — eine neue Art aus Chile... . ERBEN, M.: Bemerkungen zur Taxonomie der Gattung Limonium VIL........... FÖRTHER, H. & PODLECH, D.: Contributions to the Flora of Northern Africa I, New ee — HERTEL, H.: Floristic and taxonomic notes on saxicolous lecideoid lichens....... KÖBELE, C.P. & TILLICH, H.-J.: Die Infloreszenzen der Juncaceae.............. PODLECH, D.: Contributions to the knowledge of the genus Astragalus L. (Legumi- nosaey VIER. ne PODLECH, D. & MAASSOUMI, A.A.: New species of Astragalus L. sect. Hymeno- Steps Dom WAM Wis ais ae ye e Se eos ee ae eee TRIEBEL, D. & SCHOLZ, P.: Lichenicolous fungi from Bavaria as represented in the Botanische Stastssammlung München... ... 2020s nhs Soe oe ee ZARRE, SH. & PODLECH, D.: Taxonomic Revision of Astragalus sect. Acantho- Phase (Eabäcese)..... 0a. ee nen i ed ee ee New taxa and new combinations published in Sendtnera 7 (31.8.2001).......... Gens : Seer 137 163 203 211 233 253