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Full text of "Sendtnera : Mitteilungen der Botanischen Staatssammlung und des Instituts für Systematische Botanik der Universität München"

Sendtnera 



Mitteilungen der Botanischen Staatssammlung 

und des Instituts für Systematische Botanik 

der Universität München 



Herausgeber: 
Jürke Grau • Franz Schuhwerk 



Band 6 München 1999 



Sendtnera 



Mitteilungen der Botanischen Staatssammluns 

und des Instituts für Systematische Botanik 

der Universität München 




Herausgeber: 
Jürke Grau • Franz Schuhwerk 



Band 6 München 1 999 



Sendtnera (Mitteilungen der Botanischen Staatssammlung und des Instituts fur 
Systematische Botanik der Ludwig-Maximilians-Universität München) 
Band 6 



Erscheinungsdatum: 31.12.1999 



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 



AGERER, R.: Never change a functionally successful principle: The evolution of 
Boletales s.l. (Hymenomycetes, Basidiomycota) as seen fi-om below-ground 
features 5 

DÖBBELER, P.: Two new species of Bryonectria (Hypocreales, Ascomycetes) on 
bryophytes 93 

Erben, M.: Limonium nydeggeri - eine neue Art aus Südwestportugal 103 

Grau, J.: Eine neue Art der Gattung Alonsoa (Scrophulariaceae) aus Chile 109 

Hahn, C. & AGERER, R.: Systematical studies on Paxillaceae (Boletales, Basidio- 
mycota) 115 

PODLECH, D.: New Astragali from North Africa and Asia, mcluding some new 
combinations and remarks on some species 135 

PODLECH, D.: Typification of Astragalus species III. (Leguminosae) 175 

Ranjbar, M.: Some remarks on the genus Oxytropis (Fabaceae) from Iran 193 

Schuhwerk, F. & Lippert, W.: Chromosomenzahlen von Hieracium L. (Com- 

positae, Lactuceae) Teil 3 197 

TiLLlCH, H.-J. & Sill, E.: Morphologische und anatomische Studien an Hanguana 215 

Blume (Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit der Beschreibung 
einer neuen Art, Hanguana bogneri spec, nov 

New taxa and new combinations published in Sendtnera 6 (31.12.1999) 239 



Never change a functionally successful principle: 

The evolution of Boletales s.l. (Hymenomycetes, Basidiomycota) 

as seen from below-ground features* 



R. Agerer 



Summary: 

Agerer, R.: Never change a functionally successful principle: The evolution of Bo- 
letales s.l. (Hymenomycetes, Basidiomycota) as seen from below-ground features. - 
Sendtnera : 5-9 1 . 1 999. ISSN 0944-0 1 78. 

In the present study characteristics of substrate hyphae and of rhizomorphs have been 
applied as a completely new group of features to discern relationships of Boletales s.l. 
Rhizomorph structures have been shown to be very conservative. A so-called 'boletoid- 
rhizomorph type' is representative of Suillaceae, Rhizopogonaceae, Conio- 
phoraceae, Strobilomycetaceae, Paxillaceae, Pisolithaceae, Astraeaceae, Boletaceae 
and Sclerodermataceae. These rhizomorphs possess 'runner hyphae' where backward 
oriented ramifications grow towards the main hypha, after they have originated above 
the first simple septum or the first clamp of a side-branch. Frequently, these hyphae 
fork close to the main hypha into a distally and a proximally growing branch. The 
'runner hypha' and additional ones enlarge and become vessel-like. Gomphidiaceae, 
Tapinellaceae and Truncocolumellaceae have different types of rhizomorphs. Conio- 
phoroineae and Tapinellineae are proposed as new suborders of Boletales s.l. Omphalo- 
taceae remain provisionally placed in the Tricholomatales. The order Scleroderma- 
tales with its families Sclerodermataceae, Pisolithaceae and Astraeaceae are included in 
Boletales. 

Zusammenfassung: 

In den vorliegenden Untersuchungen werden, um verwandtschaftliche Beziehungen der 
Boletales s.l. zu klären, Charakteristika von Substrathyphen und von Rhizomorphen als 
völlig neue Gruppe von Merkmalen verwendet. Es konnte aufgezeigt werden, daß die 
Merkmale der Rhizomorphen sehr konservativ sind. Ein sog. 'boletoider-Rhizomor- 
phen-Typ' ist repräsentativ für Suillaceae, Rhizopogonaceae, Coniophoraceae, Strobi- 
lomycetaceae, Paxillaceae, Pisolithaceae, Astraeaceae, Boletaceae und Scleroderma- 
taceae. Diese Rhizomorphen besitzen 'Läufer-Hyphen' an denen rückwärts gerichtete 
Verzweigungen gegen die 'Läufer-Hyphen' wachsen, nachdem sie oberhalb des ersten 
einfachen Septums oder oberhalb der ersten Schnalle eines Seitenzweiges entsprungen 
sind. Häufig gabeln sich diese rückwärts gerichteten Hyphen nahe der 'Läufer-Hyphe' 
in einen distal und in einen proximal orientierten, weiter wachsenden Hyphenast. Die 
'Läufer-Hyphen' und weitere Hyphen verdicken sich und werden gefäßähnlich. Gom- 
phidiaceae, Tapinellaceae und Truncocolumellaceae zeigen abweichende Rhizomor- 
phen-Typen. Comophoroineae und Tapinellineae werden als neue Unterordnungen der 
Boletales s.l. vorgeschlagen. Die Omphalotaceae bleiben provisorisch in den Tricholo- 



Dedicated to Univ. -Prof Dr. Andreas Bresinsky, Regensburg, on the occasion of his 65' birthday. 



matales. Die Ordnung Sclerodermatales, mit den Familien Sclerodermataceae, Piso- 
lithaceae und Astraeaceae, wird in die Boletales einbezogen. 



Introduction 

The Boletales have attracted the interest of many mycologists, either with respect to their 
relation to Agaricales s.l., or with emphasis to the groupings within the order (comp. 
Bresinsky 1996). The discussion of the order Boletales became particularly fascinating, since 
pigment data were used to include fungal families in the Boletales with deviating types of 
fruitbodies, e.g. Coniophoraceae (Bresinsky 1973, Bresinsky & Bachmann 1971, Steglich 
etal. 1968, 1971) 

During the last decade five contributions concerning the systematics of the Boletales in a 
more comprehensive way (Besl & Bresinsky 1997, Bresinsky 1996, Hqiland 1987, 
Bruns et al. 1998, Singer 1986) were published. 

Singer ( 1 986) treats Boletales as a suborder within the Agaricales, and divides it into three 
families, viz. Boletaceae, Paxillaceae and Gomphidiaceae. Boletaceae are again subdivided into 
Gyroporoideae, Gyrodontoideae, Suilloideae, and Strobilomycetoideae. The taxa are primarily 
defined anatomically and morphologically. Almost exclusively pileate-stipitate members are 
included; neither resupinate forms (Coniophoraceae) nor gastroid species are considered (e.g. 
Rhizopogonaceae, Sclerodermataceae). 

HoiLAND (1987) applies chemical as well as anatomical characters with several character 
states, and uses parsimony and character compatibility analysis for phylogenetic conclusions. 
To distinguish plesiomorphic and apomorphic states, quite often Comer's Clavaria-theory 
(Corner 1964) is quoted to assume that Gomphus is a possible predecessor of the Boletales, 
and consequently, Gomphus is used as an outgroup. The families Paxillaceae, Gyrodontaceae, 
Xerocomaceae, Boletaceae, Strobilomycetaceae, Gomphidiaceae, Rhizopogonaceae, Chamo- 
nixiaceae and Coniophoraceae are included in this treatment. Gyrodontaceae (with Gyroporus 
and Gyrodon) are proposed to be included in the Paxillaceae; Chamonixia is regarded as related 
to Gyroporus, but treated as a separate family; Xerocomaceae are suggested to be combined 
with Boletaceae; Rhizopogonaceae are regarded as probable derivatives of Boletaceae, and 
Suillus is included in the Gomphidiaceae. 

Bresinsky (1996) puts emphasis on chemical characters, and in addition applies restriction 
fragment length polymorphism (RFLP) to several genera and species of the Boletales. His 
circumscription of Boletales includes gastroid, resupinate and pileate taxa. One main concern 
was the delimitation from other Hymenomycetes. Roughly four groups with a great deal of 
overlap have been distinguished. One group containing exclusively ectomycorrhizal fungi is 
provided only with pulvinic acid (F3) derivatives (e.g. Boletus, Chalciporus and Truncoco- 
lumella); a second shows the same pigment composition, but its constituents live saprotrophic 
as brown rot fiingi (e.g. Hygrophoropsis, Serpula and Tapinella); a third entity encompasses 
mostly ectomycorrhizal ftingi with cyclopentenones (F17-F20) as additional pigments to 
pulvinic acids (F3) (e.g. Gyrodon and Paxillus; Leccinum lacks pulvinic acids), and fourthly, 
again an ectomycorrhizal group with pulvinic acids, possesses terpenoids as accompanying 
compounds (F4-F14) (e.g. Gomphidius, Rhizopogon and Suillus). Others are regarded as 
closely related to these Boletales: Sclerodermatales {Scleroderma) and Omphalotaceae 
{Lampteromyces, Omphalotus) due to their pulivnic acids (F3), Scutigeraceae (Albatrellus) 
enriched with atromentin (Fl), the basic compound of pulvinic acids (F3) biosynthesis and 
cyclopentenones (F17-F20) and Strobilomycetaceae (Strobilomyces) which do not contain 



pulvinic acids, but DNA-characteristics indicate their membership of the Boletales. Further, 
Boletopsis appears in the scheme, but no explanation is given why this genus is regarded as 
being close to the Boletales. 

Besl & Bresinsky (1997) mainly put emphasis on the chemosystematics of Suillaceae and 
Gomphidiaceae, with the main result that both families are so closely related that they, 
together with Rhizopogonaceae, are united in a single suborder, Suillineae. Further suborders 
are: Paxillineae (with Coniophoraceae, Paxillaceae, Gyrodontaceae, and Omphalotaceae), 
Boletineae (Boletaceae) and Strobilomycetineae (Strobilomycetaceae). 

Bruns et al. (1998) applied sequences of a small region of the mitochondrial large subunit 
rRNA gene for phylogentic analysis of several Hymenomycetes, and included particularly 
relationships with ectomycorrhizal flingi. Within the Boletales they distinguished six groups. 
The boletoid group comprises species of Boletaceae and Strobilomycetaceae, the Paxillus 
group contains one species each of Chalciporus, Paxillus and Paragyrodon; in the 
Coniophoraceae-group cluster Tapinella, Serpula, Austropaxillus and Hygrophoropsis; 
separate groups are formed by Coniophora and by Gyrodon, Pisolithus, Phaeogyroporus and 
Gyrodon merulioides, respectively; a large cluster, the suilloid group, contains members of 
Gomphidiaceae, Rhizopogonaceae and Suillaceae as well as some genera previously not 
incorporated in these families, for instance, one species each of Melanogaster, Hymenogaster , 
and Truncocolumella. 

The present contribution considers a completely new complex of features to obtain new 
insights in the systematic relationships of and within the Boletales, namely characteristics of 
the substrate hyphae, i.e. the anatomy of rhizomorphs, and to some extent the structure of 
ectomycorrhizae. Thelephorales are also included, as both relationships have atromentin (Fl) 
as a basic compound for the synthesis of their pigments, and because Thelephoraceae and 
Gomphidiaceae have been suggested as closely related (Agerer 1991a). 





Fl: Atromentin (R' = R' = OH) pj: Thelephoric acid 



F3: Pulvinic acids 




F4: Boviquinone-3 (n=3) 
F5: Boviquinone-4 (n=4) 





F8: Methylenediboviquin9ne-3,3 (n = 3, m = 3) 
F9: Methylenediboviquinone-3,4 (n = 3, m = 4) 
FlO: Methylenediboviquinone-4,4 (n = 4, m = 4) 



F13: Bolegrevilol 




OAc 



F6: Diboviquinone-3,4 (n = 3, m = 4) 
F7: Diboviquinone-4,4 (n = 4, m = 4) 



F14: Suiliin 



OH 




.OH 




Fl 2: Tridentoquinone 



F16: 1,2,4-Trihydroxybenzene 

M. 




Fll: Bovilactone-4,4 y -oh 

HC 





OH HO 




F15: Gomphiiactone 



Fl 7: Gyrocyanin (R = H) F19: Chamonixin (R = H) 

F18: (^oporin (R = OH) F20: Involutin (R = OH) 



Important anatomical characters 

Three general groups of features are regarded as important for the evolution of Hymeno- 
mycetes and for interpretation of phylogenetic relations: the formation and reduction of 
clamps, evolution of substrate hyphae and occurrence of amyloidy. 



A. Transport structures, important targets of evolution 

Hyphae, as exploiting filaments, deplete their environment of nutrients close to their surface. 
Therefore, fiingi continuously have to grow, and a crucial need arises to translocate the 
incorporated substances within their hyphal systems. Three structures can be involved in 
transport: clamps, backward growing hyphae, and rhizomorphs. 

1. Clamps 

The possession of clamps has been repeatedly considered as a plesiomorphic character 
(BoiDiN 1971, Kühner 1980, Singer 1986). Since no conclusive arguments for this 
assumption are provided to date, some points are made herein. 

Hitherto, clamps apparently are almost exclusively considered from a one-sided point of view. 
Many references emphasise their role for a regular, conjugated division of nuclei, or at least 
they do not attempt to explain the reasons for the presence of clamps, due to the lack of 
clamps in so many Basidiomycota (Alexopoulos et al. 1996, Clemen^on 1997, Gäumann 
1964, Ingold & Hudson 1993). The efficiency of clamps for a regular and simultaneous 
division and transmission of dikarya is sometimes explained by the small diameter of hyphae 
and by the nuclei's mutual impediment during mitosis (Gäumann 1964). But detailed 
analyses of hyphal systems of different fungal relationships clearly show that there is no 
correlation between hyphal diameter and the presence or absence of clamps, although the 
nuclear diameter can be regarded as fairly constant for Hymenomycetes with the range of 1-2 
|im (comp. Agerer & Rambold 1998). 

For example, Byssocorticium lutescens Erikss. & Ryv. with hyphal diameters in the 
subhymenium of 2-3 |j.m forms clamps (Eriksson & Ryvarden 1973), whereas B. atrovirens 
(Fr.) Bond & Sing.: Sing, does not produce them, although it possesses the same hyphal 
dimensions (Brand 1991). There are some examples however, where a correlation of clamps 
with hyphal diameter could be given, but a causal interdependence has yet to be shown. For 
instance, Ceraceomerulius serpens (Fr.) Erikss. & Ryv. which possesses clamps, shows 2.5- 
3.5 |im thick hyphae (Eriksson & Ryvarden 1973), whereas the clampless hyphae of C. 
mbicundus (Litsch.) Erikss. are 3-5 |im thick (Eriksson & Ryvarden 1973). In some Athelia 
species, a further combination is realized. Here, within one and the same fruitbody, primarily 
thick, basal hyphae form clamps, whereas the thinner ones in the subhymenium are clampless 
(Eriksson & Ryvarden 1973). But the opposite situation is also known. Most clamp- 
bearing species of the genus Hyphoderma form li-A |im wide hyphae (Eriksson & Ryvarden 
1975), but the clampless hyphae of//, capitatum Erikss. & Ryv. are 2-3 ^m thick (Eriksson 
& Ryvarden 1975). Among others, the species of the genus Botryobasidium possess hyphae 
with 6-10 ^m diameter. They include members with clamps and without clamps (Langer G 
1994). The thinnest clampless hyphae possibly occur in the genus Dendrothele, for instance 
Dendrothele acerina (Fr.) Lemke, with diameters of 1-2 urn (Eriksson & Ryvarden 1973). 



10 



Most of the Hymenomycetes, at least when "The Corticiaceae of North Europe" are 
evaluated, form hyphae of 3-^ |im diam., regardless, whether clamps are formed or not. 
Although the dikaryotic condition of their clampless hyphae has not been shown in most of 
the species, their presence in all of the above mentioned taxa can very likely be inferred from 
the formation of four spores per basidium. 

Since, according to Alexopoulos et al. (1996), also some homokaryotic mycelia can form 
clamps, these specialized structures are not necessarily involved in synchronous nuclear 
division. A similar situation is known about the formation of whirl-clamps, where the suc- 
cessively originating clamps are not always accompanied by the division of nuclei, and, quite 
importantly, clamps are formed even without a conjugated division of nuclei (Clemenqon 
1997, Greis 1937). InStereum hirsutum (Willd.: Fr.) S.F.Gray however, several nuclei divide 
simultaneously, and the number of clamps corresponds with the number of nuclei involved 

(GÄUMANN 1964). 

In Coprinus disseminatus (Pers.: Fr.) S.F.Gray low-temperature treatment decreased 
hyphal diameter and depressed clamp formation (Butler 1981), although the hyphae re- 
mained dikaryotic. This indicates an independence of clamp formation from the simultaneous 
mitosis of the dikaryon and from hyphal diameter. In this dikaryotic mycelium, although 
compatible nuclei, i.e. different A allels (clamp cells are formed) and different B allels (clamp 
cell fuses with subterminal cell) are present (Casselton & KüES 1994), the formation of 
clamps is repressed by environmental conditions. 

A completely different approach to explain clamp formation, lets nuclei for the moment out 
of account and emphasizes primarily cytoplasmic transport as the main reason for the 
formation of backward oriented, hook-like outgrowths, the future clamps. 

As shown by Buller (1933) in sophisticated light microscopy observations based on the 
movements of cytoplasmic granules, the cytoplasm streams in all fungi investigated from older 
to younger portions of the hyphae, i.e. in most cases from the base to the growing hyphal tip. 
There was no support for the opinion that within one hypha bidirectional transport occurs. 
The mechanism for protoplasmic transport seems to be a turgor-driven mass flow of solutions 
(Jennings 1994), although this has been recently questioned by Heath & Steinberg (1999). 
A reversal transport, from tip to proximal parts, is evident when a tip forms an anastomosis 
with an older hyphal cell. Then the cytoplasm streams into this anastomosing tip down to 
older parts of the hypha (Buller 1933). 

Hyphal tips are not only sites of extension by production of wall material ('growth zone'), 
they are also, shortly behind the growth zone in the 'absorption zone', highly active areas of 
export of digesting enzymes and incorporation of ions and organic molecules (Ingold & 
Hudson 1993, Jennings & Lysek 1996). Hyphal tips show intensive lysis of organic ma- 
terial, and they are sites of absorption, as is clearly indicated by haustoria, which are in many 
cases transformed hyphal tips (Jennings & Lysek 1996).The molecules taken up have to be 
transported back to the 'storage zone' of the hyphae (Jennings & Lysek 1996), or further 
back to comsumption areas, and this apparently is flowing against the tip-directed counter- 
current of cytoplasm (Buller 1933). The mass flow towards hyphal tip is in fact important 
for extending hyphae, since cytoplasm is pushed forward the quicker the faster the hypha 
grows (Buller 1933), and hyphae can save cytoplasm in an economical way (Steinberg et 
al. 1998). But this flow cannot explain how the incorporated ions and molecules taken up in 
distal parts of the hyphae by endocytosis (Hoffmann & Mendgen 1998) are transported 
backward, against the counter-current mass flow of cytoplasm. Opposed streaming directions 
in one and the same hyphal compartment should at least occur down to those points, where 



11 



the first anastomosis occurs or where backward oriented branches can take charge of transport 
towards their tip, which is as a consequence oriented towards the base of the main hypha. On 
microtubules gliding vesicles have been demonstrated (Lehmler et al. 1997, Steinberg 1997, 
Steinberg & Schliwa 1995), and regarded as important for transport; the vesicles are shown 
to perform a retrograde movement (Steinberg et al. 1998). Recent studies provided evidence 
for a contemporary bidirectional transport of vesicles along microtubles in one and the same 
hypha, and at one and the same cytoplasmic site. Due to antiparallel orientation of micro- 
tubules, or to different kinds of motor molecules vesicles should be moved towards the hyphal 
tip as well as towards proximal cell regions (Steinberg., pers. comm.). 

Clamps of Basidiomycota are regarded as by-pass hyphae for the transport of nuclei 
(Hawksworth et al. 1985), and they can be formed in response to nuclear positions 
(Clemencon 1997). The hook of the later on formed clamp is undoubtedly a short, backward- 
oriented side-branch, with a Spitzenkörper, and vesicles have to be transported from the sub- 
apical Golgi bodies to the site of branch initiation (Heath 1994) and the nuclei to the position 
of clamp formation (Weber 1993). The role of microtubles in clamp formation (Ormerod et 
al. 1976, Shepherd et al. 1993), nuclei movement with changing intensity and reversal of their 
direction of movement (Suelmann et al. 1997, Weber I.e.) have been shown and lastly, the 
entry of a single nucleus into the hook, as Alexopoulos et al. (1996), Casselton & Kües 
(1994), iNGOLD & Hudson (1993) and Weber (1993) point out. Other authors however, refer 
to a daughter nucleus which enters the hook during mitosis (Clemencon 1997, Hawksworth 
et al. 1995). This course of events during clamp formation and the necessary transport of in- 
corporated molecules back to older hyphal compartments suggest a backward transport within 
cytoplasm (Shepherd et al. 1993). 

Nuclear divisions are mostly combined with the formation of septa, which are positioned 
between the pairs of daughter nuclei of the dikaryon, resulting in the two septa of a mature 
clamp (Girbardt 1979). Both have a central pore. The backward oriented hook which 
appeared during clamp formation is possibly not only a by-pass for migration of nuclei 
(Hawksworth et al. 1995), but also a general means to enable contemporary backward and 
forward cytoplasmic transport through the septa. Shepherd et al. (1993) found tubules 
moving in both directions through the dolipore septum. The movements appeared co- 
ordinated, and during the exchange between the terminal and the penultimate cells tubules 
transiently interconnected vacuoles in adjacent cells. Peristaltic movements appeared to 
transfer material beween them. If both the septa are used for transport, although tubules have 
been shown to move in the dolipores in both directions, a specialization of the dolipores of a 
clamp regarding transport directions could considerably facilitate bidirectional transport in a 
hypha. One septum could serve the forward pathway, the other could provide the by-pass for 
the backward transport. As the dolipores of the septa are very thin (Clemencon 1997), and 
parenthesomes encage the pore the countercurrents of transport will meet there in a 
bottleneck, and a piling up of vesicles could result. The inclusion of the second pore could 
relieve the cytoplasmic traffic. 

This hypothesis therefore considers clamps firstly as appropriate aids to prevent con- 
gestion of two countercurrent movements at a septum. Secondly, the clamp formation is not a 
means for unimpeded synchronous nuclear division, but should rather be understood as the 
necessary prerequisite for formation of two septa and hyphal compartmentation. However, 
the inclusion of the genes for clamp formation and fusion in the mating type loci (Casselton 
& Kües 1994) suggest a primary role of clamp formation in sexuality. A functional switch 
should have taken place. 



12 



Nuclear divisions also precede septa formation in simple septate hyphae, but only one septum 
is built. Bidirectional transport should therefore also be possible in simple septate hyphae. 
How this demand can be resolved, will be discussed further below. 

2. Backwardly growing hyphae 

Many Hymenomycetes do not establish clamps any more. Possibly others are primarily 
simple-septate. There are several examples in which the reduction of clamps is evident, and 
where the clamps can be reduced to different degrees. For instance, some species of the genus 
Suillus provide evidence for an almost complete reduction of clamps. Only the 'runner hypha' 
(rapidly growing main hypha with very distant septa and branches; different from 'leading 
hyphae' which grow rather slowly and form septa and branches in shorter distances), 
possesses clamps, which is later that hypha where the formation of rhizomorphs starts, e.g. 
Suillus variegatus (Swartz: Fr.) Kuntze (Raidl 1997). The question arises, why several fungi 
can deny formation of clamps. This question is particularly relevant when clamps are 
considered by-passes for a bidirectional transport. 

Since Falck (Falck 1912, Melin 1923) it has been repeatedly shown that backward 
growing ramifications of hyphae are a rather common feature of ectomycorrhizal (Agerer 
1995, Raidl 1997) and of saprotrophic fimgi (Agerer, in prep.). Agerer (1992b) and 
Cairney (1992) have hypothesized that this mode of ramification can facilitate bidirectional 
transport in a mycelial system. 

Most of the rhizomorphic fungi studied form backward oriented ramifications (e.g. Agerer 
1995, Agerer & Rambold 1998). Only a few exceptions are known: Cystoderma carcharias 
(Agerer, in prep.), Entoloma sinuatum (Bull.: Fr.) Kummer (Agerer 1997), Laccaria bicolor 
(R.Mre.) Orton (Raidl 1997/ L. amethystina (Bull.: Hooker) Murr. (Raidl & Agerer 
1992), Omphalotus olearius (DC: Fr.) Sing. (fig. 8), O. atraetopus (Kalchbr. apud Thiimen) 
Ch.Hahn (fig. 8) and Ripartites tricholoma (A. & S.: Fr.) P. Karst. (Agerer, in prep.). More 
often backward oriented branches are lacking in non-rhizomorphic fungi, e.g. Pseudotomentella 
tristis (P. Karst.) M.J.Larsen (Agerer 1994), Rozites caperatus (Pers.: Fr.) P. Karst. (Agerer 
1999a), Tomentella albomarginata (Bourd. & Galz.) M.J.Larsen (Agerer 1996a) and 
Tylospora asterophora (Bonord.) Donk (Raidl 1997). This possibly applies also to most 
species of the genus Inocybe. With the exception of Pseudotomentella tristis all the mentioned 
examples possess clamps. 

A possible explanation of the evolution of backward growing hyphae is deducible from 
observations by Niederpruem et al. (1971) on mutants of Schizophyllum commune Fr.: Fr. 
Pseudoclamps delimited by a neck septum and with a cross-wall in the main hyphal axis could 
grow out in different directions, forward as well as rearward, to form a new branch. Parag 
(1965) found uninucleate pseudoclamps growing out exclusively backward. It is tempting to 
speculate that, during evolution of Hymenomycetes, the origin of backward oriented ramifi- 
cations was based on such mutations. This course of events can very likely also explain the 
co-evolution of backward oriented ramifications and simple septa, realized, for instance, in 
Jaapia ochroleuca (Bres.) Narmf & J.Erikss. (figs. 13c, d). An outgrowing pseudoclamp 
lacking a neck septum could be the reason for the formation of backward oriented branches 
which refrain from clamp formation. Also young stages of whirl-clamps, not fused with the 
main hypha, can grow to normal hyphae (Kemper 1937). This was exclusively attributed to 
higher air moisture. 



13 



Backward oriented branches would be able to support or replace the by-pass effect of clamps 
for bidirectional transportation within the hyphae, regardless as to whether clamps are formed 
or not. Consequently, the different transport directions can be distributed over different 
hyphae, as Cairney (1992) hypothesized. This line of arguments can now be used for 
evolutionary considerations regarding specialization of substrate hyphae and evolution of 
rhizomorphs. 

3. Rhizomorphs: conserved structures with a high diversity 

As convincingly pointed out by Ober winkler (1985), the most primitive fruitbodies of 
Hymenomycetes belong to the corticioid type, i.e. resupinate with smooth hymenophore. 
Although such species are depicted at most exclusively with their hymenium, trama and 
hyphae connecting the fruitbodies to the substrate (e.g. Breitenbach & Kränzlin 1986, 
Eriksson & Ryvarden 1973), i.e. the 'superficial substrate hyphae', all species in addition 
grow within wood, leaves, etc., on which the fruitbodies are formed. These 'internal substrate 
hyphae' have to exploit the organic matter, and they have to transfer energy-rich compounds 
and nutrients. The mycelia use them for the formation of reproductive structures. The 
'internal substrate hyphae' are in particular targets of selection and evolution until they are 
functionally and anatomically adapted in their special ecological niche to the fungi's nutritional 
demands. 

'Internal substrate hyphae' growing in soft substrate or in cavities, and 'superficial sub- 
strate hyphae', can aggregate into linear structures, which can be differently assembled. Such 
multihyphal linear organs are generally known as rhizomorphs (Agerer 1987-1998, Cairney 
et al. 1991). According to Rayner et al. (1999) rhizomorphs "minimize the dissipative free 
surface of hyphae and is engergy-saving and even energy-yielding". Besides transport 
functions, this might be an additional advantage of hyphal aggregation to rhizomorphs. 

A screening of the series "Corticiaceae of North Europe" regarding 'superficial substrate 
hyphae' results in 91 genera, which exclusively form simple hyphae which are not bundled to 
any extent. Only 30 genera contain species which produce rhizomorphs. In total, only 65 
species of the 342 fungi treated in 'Corticiaceae of North Europe' are rhizomorphic. 

Most of these species form primitive rhizomorphs with often rather loosely bundled 
hyphae of uniform diameter. Quite infrequently some of the rhizomorph hyphae are formed as 
simple, thick-walled, sometimes slightly thinner skeletals. Only a few species show 
differentiated structures in so far as diameters of a few hyphae are enlarged. This latter 
situation is realized only in the following 19 species (a question mark indicates that literature 
data are not conclusive enough). Byssocorticium terrestre (Fr.) Bonord. & Sing. (Zak & 
Larsen 1978), Jaapia ochroleuca (figs. 13c, d), Kavinia alboviridis (Morgan) Gilbertson & 
Budington, K. himantia (Schw.) Erikss., Leucogyrophana pseudomollusca (fig. 18), L. 
mollusca (?) (Farm.) Farm. (Eriksson & Ryvarden 1976), Lindtneria trachyspora (Raidl, in 
prep.;, Phanerochaete calotricha (?) (F. Karst.) Erikss. & Ryv., P. galactites (?) (Bourd. & 
Galz.) Erikss. & Ryv., P. laevis (?) (Fr.) Erikss. & Ryv., P. sanguinea (?) (Fr.) Fouz. 
(Eriksson etal. 1978;, Phlebiella subflavidogrisea (Litsch.) Oberw. (Hjortstam et al. 1988), 
Phlebiellavaga{¥r.) Karst (Cairney & Clipson 1991), Ramaricium alboochraceum (Bres.) 
Jülich (Hahn et al. 2000, Eriksson & al. 1981), Steccherinum fimbriatum (Fers.: Fr.) J. Erikss. 
(Cairney et al. 1989;, Trechispora cohaerens (?) (Schw.) Jülich & Stalpers, T. microspora (?) 
(P. Karst.) Liberta, T. mollusca (Fers.: Fr.) Liberia and T. subsphaerospora (Litsch.) Liberta 
(Hjortstam et al. 1988). 



14 



Kavinia, Ramaricium and Trechispora species form at most ampullate hyphae in their rhizo- 
morphs. The most highly differentiated rhizomorphs occur in the few species Byssocorticium 
terrestre, Jaapia ochroleuca, Leucogyrophana mollusca (?), L. pseudomollusca, Lindtneria 
trachyspora, Phlebiella vaga, and Steccherinum fimbriatum. 

The first conclusion therefore is that Hymenomycetes with primitive fruitbodies lack 
rhizomorphs, or when rhizomorphs are formed, they are frequently only slightly differentia- 
ted. Apparently, those species with highly elaborated rhizomorphs are often related to species 
which possess fruitbodies of a more advanced type: Jaapia to Serpula (see below), Leuco- 
gyrophana to Hygrophoropsis (see below) and Lindtneria to Stephanospora (Oberwinkler 
& HoRAK 1979). As will be shown later for the order Boletales, and below for the Gomphales, 
the main rhizomorph organization can persist during evolution, even when the fruitbodies have 
evolved to a very complex structure. 

Many fungi of different relationships retain a primitive, undifferentiated organization of 
their rhizomorphs. Even when they have derived types of fruitbody. Other fungi do not form 
rhizomorphs at all, e.g. the genera Chroogomphus (Agerer 1990), Inocybe (Agerer 1995), 
some Tomentella species (Danielson & Prüden 1989), Pseudotomentella tristis (Agerer 
1994), Rozites caperatus (Agerer 1999a) and many Russula species 

A 'uniform-loose rhizomorph type' (fig. 1, designated as type A according to Agerer 
1987-1998) is represented by most of the rhizomorphic corticioid fungi, e.g. Amphinema bys- 
soides (Pers.: Fr.) J.Erikss. (Weiss 1991), Piloderma croceum J.Erikss. & Hjortst. (Brand 
1991) and Tomentella albomarginata (Agerer 1994). It is also known for example in Dermo- 
cybe (Agerer 1995, Agerer & Rambold 1998), Entoloma sinuatum (Agerer 1997), Gom- 
phidius roseus (L.) Fr. (Agerer 1991a), a few Tricholoma species (Agerer 1987, Agerer & 
Rambold 1998), and in Melanophyllum echinatum (Roth: Fr.) Sing. (Agerer, in prep.). 

The 'uniform-compact rhizomorph type' (fig. 1, type B according to Agerer 1987- 
1998) is, for instance, realized in most Lactarius species (Agerer & Rambold 1998), 
Bankera fuligineo-alba (Schmidt: Fr.) Pouz. (Agerer & Otto 1997), Cystoderma caracha- 
rias (Pers.: Seer.) Fay. (Agerer, in prep.), Entoloma clypeatum (L.: Fr.) Kummer (Agerer, 
unpubl.), Phellodon niger (Fr.: Fr.) P. Karst. (Agerer 1992c), Hydnellum peckii Banker apud 
Peck (Agerer 1993), Tometellopsis submollis (Svrcek) Hortst. (Agerer 1998b), Ripartites 
tricholoma (Agerer, in prep.), Omphalotus olearius (fig. 8) and O. atraetopus (fig. 8). 

A 'thelephoroid rhizomorph type' (fig. 1, included in rhizomorph type C (Agerer 
1987-1998)) can be designated as a further type. It is slightly differentiated with only the 
peripheral hyphae differing somewhat in diameter and structure. This type is typically 
shown, for example, by Thelephora terrestris Pers.: Fr. (Agerer & Weiss 1989), Tomentella 
ferruginea (Pers.: Fr.) Pat. (Raidl & Müller 1996), Gomphidius glutinosus (Schaeff) Fr. 
(Agerer 1991a). 

A 'russuloid rhizomorph type' (fig. 1, type E according to Agerer 1987-1998) is 
characteristic of some species of the genus Russula, e.g. Russula ochroleuca (Pers.) Fr. 
(Agerer 1986) and R. acrifolia Romagn. (Agerer et al. 1994). Some thickened hyphae with 
thin branches and preponderantly complete septa are irregularly distributed in the rhizo- 
morphs accompanied by ladder-like thick-walled hyphae with several septa in close distance 
(comp. Moyersoen 1996). 

The 'phlegmacioid rhizomorph type' (fig. 1, type D, according to Agerer 1987-1998) 
is similar to the 'russuloid rhizomorph'. A few randomly distributed slightly thicker hyphae, 
embedded in a matrix, can enlarge their septal pore; a distinct septal dissolution is mostly 
lacking. Such rhizomorphs are found in Cortinarius subg. Phlegmacium (e.g. Agerer 1988b) 



15 



or in Albatrellus ovinus (Schaeff: Fr.) Kotl. & Pouz. (Agerer et al. 1996b) and Polyporoletus 
sublividus Snell (Agerer et al. 1998).. 

Finally the boletoid rhizomorph type (fig. 1 , included in rhizomorph type F, according to 
Agerer 1987-1998), so characteristic of boletes will be discussed in detail further below. 

The fiingal relationship that will be discussed below has maintained principally identical 
rhizomorphs, but evolved fruitbodies of great diversity. It conveys an impression as to how 
conservative rhizomorph characters can be, suggesting a high value of these organs for 
discerning fungal relationships of higher-level taxa. 

Recently, it has been shown that the ectomycorrhizae, including rhizomorphs, of Gomphus 
clavatus (Pers.: Fr.) S.F.Gray and of several Ramaria species have ampullate hyphae in their 
rhizomorphs, produce acanthocystidia with yellowish contents and plasmatically yellowish, 
irregularly roundish, inflated, thin- walled cells (Agerer 1996b, 1998a). In addition, the eco- 
logical niche of their ectomycorrhizae is identical. They form dense mats predominantly in the 
mineral soil layer and become greyish or blackish when they senesce (Agerer & Rambold 
1998). Almost the same features could be found for Geastrum fimbriatum Fr. (Agerer & 
Beenken 1998). But in addition very thin, thick-walled dextrinoid hyphae could be detected 
on the mycorrhizal mantle and on the surface of the rhizomorphs of this species. Acantho- 
cystidia and the yellowish, inflated cells could however, only be found on the mycorrhizal 
mantle. It was concluded (Agerer & Beenken 1998) that all three genera are systematically 
rather closely related due to these unique features. This reasoning has been corroborated by 
results from comparison of DNA sequences (Hibbett et al. 1997a, 1997b), and by ultra- 
structural studies, for continuous parenthesomes could be found in all these genera (Hahn et 
al. 2000, Bauer, pers. comm.). Ramaricium alboochraceum (Bres.) Jülich, regarded as a mem- 
ber of Gomphaceae (Ginns 1979, Oberwinkler 1977a), shows the same organization of rhi- 
zomorphs; only the acanthocystidia are lacking (Hahn et al. 2000). As an additional feature, 
Ramaricium and Ramaria hyphae are covered by lens-shaped wall appositions (Agerer & 
Rambold 1998), which admittedly are lacking in the studied Gomphus and Geastrum species. 

Bresinsky (1996) and Pegler et al. (1993) regard the genus Gautieria as related to the 
order Boletales, apparently referring to the argument that, as many Boletales, Gautieria is 
parasitized by the genus Sepedonium (Besl et al. 1996, Pegler et al. 1993) and Gautieria and 
Chamonixia possess similar spores, as Smith & Singer (1959) had already pointed out. 
Oberwinkler (1977a) however, regarded Gautieriales as a separate gastroid order with un- 
known connections to other fiangi. Our own investigations clearly show that Gautieria spec, 
nov., from Chile (Palfner, in prep.) forms all the typical below-ground features of Ramaria: 
ampullate rhizomorph hyphae, acanthocystidia, roundish, thin-walled, plamatically yellowish 
cells, hyphae with lens-shaped appositions (fig. 4), and mat-like ectomycorrhizae which are 
formed in mineral soil layer, and become blackish when old. All these features shift the genus 
Gautieria close to the Ramaricium-Ramaria-Gomphus-Geastrum complex. Though ultra- 
structrual results are lacking yet, there cannot be much doubt that Gautieria is more a member 
of this complex, and not a member of Boletales, particularly as Bruns et al. (1998) could 
show that mitochondrial rDNA sequences indicate a monophyletic relationship of Gomphus, 
Ramaria, Kavinia and Gautieria. 

The diversity of the fruitbodies of this relationship could not be greater than demonstrated 
by these fungi. Ramaricium is typically corticioid, Ramaria is clavarioid, Gomphus forms 
stipitate-pileate fruitbodies, Geastrum is an epigeous earth-star with powdery spore masses, 
and Gautieria a hypogeous fungus with a chambered gleba until maturation and during spore 



16 



dispersal (Oberwinkler 1977a). Kavinia, also a member of this group (Bruns et al. 1998), 
contributes to this diversity with resupinate hydnoid fruitbodies. Whereas the spores of Ra- 
maricium. Ramaria, and Gomphus are very similar in their shape and in their brownish colour 
with rough surface, those of Geastrum are small, globose and warty, and the epispore of 
Gautieria is longitudinally ribbed. It is apparent that not only the fruitbodies and the spores 
changed their shape during evolution considerably, but that also the form and function of 
basidia as Gautieria and Geastrum have adapted the gastroid organization of the fruitbodies. 

The relationship which includes Gomphales, Geastrales and Gautieriales suggests that 
structures of the rhizomorphs comprise of highly conserved features. Their conservatism 
seems to be of a similar level as that of the parenthesome-dolipore type and of rDNA 
sequences. The reason why these structures are so highly conserved can only be speculated. 
The ampullate hyphae possibly enhance nutrient transport in the special ecologcial niche 
where mycorrhizae and hyphal mats are in close contact to mineral soil layers. There, the 
dense mats of hyphae often dry out the soil almost completely. Whether the yellowish 
contents of acanthocystidia and/or globular cells play a role for defence against soil organisms 
is as yet unknown. 

Similar rhizomorphs, but without the decisive acanthocystidia and without inflated cells 
with yellowish contents (they are also lacking on the mycorrhizal mantle), are typical of Hyä- 
num repandum L.: Fr. and H. rufescens Fr. (Agerer et al. 1996c, Raidl & Agerer 1992). 

Rhizomorphs with ampullate hyphae are henceforth, due to their characterization capacity 
of the Gomphales-Geastrales-Gauteriales complex, designated as 'ramarioid rhizomorph 
type' (fig. 1, included in rhizomorph type C according to Agerer 1987-1998) 

4. Structure and evolution of the 'boletoid rhizomorph type' (figs. 37, 38) 

The 'boletoid rhizomorph type' (fig. 1, included in tpye F (Agerer 1987-98)) is one of 
the most highly elaborated conducting organs of fungi. It is characterized firstly, when mature 
by very thick, often centrally arranged hyphae, with partially or even completely dissolved 
septa (= vessel-like hyphae). A fast growing 'runner hypha' with often very distant septa is 
the first hypha which increases considerably its diameter over its whole length; and this hypha 
is consistently the first one to dissolve septa. Exceptions are Jaapia ochroleuca (figs. 13c, d) 
and Leucogyrophana pseudomollusca (fig. 18a) with septa remaining complete. A similar final 
organization of rhizomorphs is also true for the 'agaricoid rhizmorph type'(fig. 1, also 
included in type F (Agerer 1987-98)), (fig. 5), but its ontogeny is completely different 
(Agerer, in prep.). Fast growing 'runner hyphae' are lacking, 'leading hyphae' can precede 
mycelial extension, but they are not the first to be enlarged, and when they become inflated, 
only a portion of their cells do so (fig. 38a). 

Secondly, in the 'boletoid rhizomorph type' backward oriented ramifications grow towards 
the main hypha, after they have originated above the first simple septum or the first clamp of 
a side-branch. There, in many species, it forks into two hyphae, one takes the growth 
direction of the main hypha, the other grows towards its proximal end (figs. 13d, 15a, 18d, 19a, 
22a, b, 23d, 24d, 26d, e, 27b, 28b, 29d-f, 30a, 31e, 34b, 35a, 36c). In some species however, 
no fork is formed. In the latter case the unramified, reversely oriented hypha grows directly 
towards the proximal end of the main hypha (figs. 13c, 23a,d, 24e, 30c,d, 32c, 33b, 35a, 36a). 
When young, these hyphae are distinctly thinner than the main hypha and keep in touch with 
it. Such hyphae in Serpula lacrymans (Wulf. In Jacq.: Fr.) Schroeter have been called 'Ran- 
kenfäden' by Falck (1912). The 'agaricoid rhizomorph' develops backward oriented hyphal 



17 

ramifications, too, but they do not originate from the proximal end of a side branch and do not 
grow towards the main hypha and do not keep in intimate contact with it over considerable 
distances. They are direct derivatives of a main hypha, and are of similar diameter. The same 
type of hyphal branching as in 'boletoid rhizmorphs' is known in some Tricholoma species 
and Amanita muscaria (L.: Fr.) Hooker (Raidl 1997). 

Thirdly, most species are characterized by peculiar nodes at branching points of rhizo- 
morphs (figs. 15c-e, 17b, 18a, 22c, 23g, 27a, b, 34c). Such thickenings can also be evident 
without ramification, e.g. in Serpula lacrymans. At any rate, the nodes occur at positions of 
septa or clamps of the main hypha. There, several thin hyphal branches divide repeatedly in 
an irregular manner resulting in a knot-like structure. Bent and tortuous shapes of these 
hyphae increase the distinctiveness of this structure. Such nodes are known, e. g. in Amanita 
muscaria (Raidl 1997), Leccinum scabrum (Bull.: Fr.) S.F.Gray (Müller &, Agerer 1990), 
Tomentella spec. (AzuL et al. 1999) and Tricholoma vaccinum (Pers.: Fr.) Kummer (Raidl 
1997). Hyphae form many anastomoses within this node, and also close to it. The anasto- 
moses probably facilitate transfer at this rhizomorph branching point, since it is known that 
vessel-like hyphae of rhizomorphs of some taxa, e.g. Suillus bovinus, are also repeatedly 
connected to the mantle by many short branches and anastomosing hyphae (Agerer 1990). 
Anastomoses with the main hypha can be found only as exceptions, for instance in Chalci- 
porus piperatus (Bull.: Fr.) Pat. (fig. 31f). 

Fourthly, conical side-branches are obvious in several species (Agerer & Rambold 1998), 
(figs.20c, 22g, 26h, 29h, 33c, 34e, 35b, 36d). The ontogeny of these structures is studied in 
detail in Paxillus involutus (Fr.) Batsch (Agerer 1988a). A hypha, branching off from a 
'runner hypha', becomes enveloped primarily at its base by thin hyphae from the main 
rhizomorph. leaving the distal part of the hypha naked over a longer distance (figs. 20c, 26h, 
33c, 34e). Besides members of the Boletales, these conical side-branches occur in Tomentella 
ferruginea (Raidl & Müller 1996), and very distinctly in Tomentella spec. (fig. 7e, f). 

'Boletoid rhizomorphs' are, from a phylogentic point of view, very old. They are already 
formed by species with a supposed primitive fruitbody type, for instance by Jaapia ochro- 
leuca. This extant species can help to understand the evolution of the 'boletoid rhizomorph 
type'. 

In Jaapia ochroleuca, long, tubular cystidia without septa (150-200 p,m long, according to 
Eriksson & Ryvarden 1976) which originate in the trama project through the hymenium 
(figs. 13a, 14a), but they can also grow radially over the substrate at the margin of the 
corticioid fruitbody (fig. 13b). Cystidia can reach extreme lengths of 360 |im (Rogers 1943). 
Apically thin-walled cystida are sometimes able to grow out, and continue their growth with a 
clamp and a normal thin-walled hyphal extension, as is shown for tubular cystidia in aculei of 
Hyphodontia floccosa (Bourd. & Galz.) J.Erikss. by Langer E. (1994). A similar procedure 
appears conceivable for the evolution of the rhizomorphs oi Jaapia ochroleuca predecessors. 
Tubular cystidia could have become longer and thicker, and at the clamps the second 
important characteristic of the boletoid rhizomorph appeared, namely backward growing 
hyphae above the first septum of a side-branch (figs. 13c, d). Although the fruitbody 
consistently possesses clamps, sidebranches and reversely oriented hyphae can forego of 
clamps and form simple septa (figs. 13c, d). The central 'runner hypha' is only enveloped 
with a few additional hyphae, and the rhizomorph is therefore barely differentiated. The septa 
do not dissolve, neither those of the 'runner hypha' nor the septa of the enveloping ones (figs. 
13c, d), although some 'internal substrate hyphae' show disintegrated septa (fig. 14c). 



Coniophora arida (Fr.) P.Karst, bears several clamps on its considerably long 'runner hypha' 
with very distant septa (figs. I5e, 16a). No simple septa occur in it. The whirl-clamps are 
formed consecutively, i.e. older 'runner hyphae' increase their number of clamps in the whirl 
consecutively in comparison to younger hyphae. This is in accordance with the observations 
of Kemper (1937). First side-branches can originate even on simple septa (fig. 15a), and are 
generally not formed by outgrowth of clamps (figs. 15e, 16c, d, 17e). Even thicker 
rhizomorphs do not show any sign of septal dissolution in the 'runner hyphae'; pores are at 
most only slightly enlarged (figs. 16a, b). The 'runner hypha, is the first to exploit the 
substrate, and is possibly also the first to transport substances back for support of hyphal 
growth. It is therefore remarkable that its septa are not dissolved. Whether the multiple 
clamps per septum provide the hypha with more transport facilities is difficult do demon- 
strate, but the enlarged and thick-walled 'runner hypha' suggests an effective transport, in 
spite of the missing septal disintegration. The simple septate enveloping hyphae however, dis- 
solve the septa (fig. 17a). The 'runner hypha' oi Coniophora arida, remind therefore of the 
'runner hypha' of Jaapia ochroleuca (figs. 13c, d). Leucogyrophana pseudomollusca 
possesses also very long 'runner hyphae' with distant clamps (fig. 18a), no whirl-clamps are 
formed, and the clamp septa are dissolved to variable degrees (fig. 18a). Enveloping hyphae 
show clamps and simple septa as well, and their septa are disintegrated to a higher extent (fig. 
18b), similar to those o^ Coniophora arida (fig. 17a). Hygrophoropsis aurantiaca (Wulf: Fr.) 
R.Mre. (figs. 20, 21) reveals the same ontogeny as L. pseudomollusca, but simple septa are 
apparently not formed. The 'runner hypha' has its clamps not so distant, and most of the 
septa are at least in part dissolved, resulting in very long vessel-hyphae with a possibly 
unimpeded transport system of more than 1.7 |im (fig. 21). Serpula lacrymans (fig. 19) rhizo- 
morph ontogeny has already been studied in detail by Falck (1912); his observations on early 
stages agree completely with our own. Mature rhizomorphs have a similar structure as those 
of Hygrophoropsis aurantiaca, but in addition skeletal hyphae are evident (Falck 1912, 
HoRNUNG & Jennings 1981). 

The 'boletoid rhizomorphs' oi Leucogyrophana pseudomollusca, Hygrophoropsis auran- 
tiaca and Serpula lacrymans are plesiomorphic regarding clamps in comparison to Jaapia 
ochroleuca and Coniophora arida, which frequently develop simple septa. Rhizomorphs with 
consistent clamps are also known, for instance in Alpova trappei Fogel (fig. 26), Austropaxillus 
boletinoides (Sing.) Bresinsky & Jarosch (Palftier, in prep.), Boletinus cavipes (Opat.) 
Kalachbr. (Raidl 1997, Treu 1990), Gyrodon lividus (Bull.: Fr.) Sacc. (Agerer et al. 1993), 
Gyroporus cyanescens (Bull.: Fr.) Quel. (Agerer 1999b), Melanogaster variegatus (Vitt.) 
Tul. (fig. 26), Paxillus involutus (Agerer 1988a), Phaeogyroporus beniensis Sing. & Dig. (fig. 
24), Pisolithus tinctorius (Mich.: Pers.) Coker & Couch (Agerer 1991b) and Scleroderma 
citrinum Pers. (Raidl 1997). 

Several species with 'boletoid rhizomorphs' have reduced the clamps in the rhizomorphs, 
but occasionally occur on the 'runner hypha', for example, in Rhizopogon roseolus (Corda in 
Sturm) Th.M.Fr. (Raidl & Agerer 1998), Scleroderma areolatum Ehrenb. (fig. 36), Suillus 
variegatus (Raidl 1997). Many species reduce clamps in rhizomorphs completely, e.g. Afro- 
boletus luteolus (Heinem.) Pegler & Young (fig. 23), Austroboletus gracilis (Peck) Wolfe (fig. 
28), Austropaxillus statuum (Speg.) Bresinsky & Jarosch (Hahn, unpubl.), Boletellus pruina- 
tus (Fr. & Hök) Klofac & Krisai-Greilhuber (fig. 29), Boletus erythropus (Fr.: Fr.) Pers. (fig. 
30), Chalciporus piperatus (Bull.: Fr.) Pat. (fig. 31), Chamonixia caespitosa Rolland (fig. 32), 
Leccinum scabrum (Bull.: Fr.) S.F.Gray (Müller &, Agerer 1990), Phylloporus rhodoxan- 
thus (Schw.) Bres. (fig. 33), Porphyrellus pseudoscaber (Seer.) Sing. (fig. 34), Pulveroboletus 



19 



cramesinus (Seer.) Sing. (fig. 35), Rhizopogon vinicolor A.H.Smith in Smith & Zeller (fig. 25), 
Strobilomyces floccopus (Vahl: Fr.) P. Karst, (fig. 22), Suillus tridentinus (Bres.) Sing. (Treu 
1990), Tylopilus felleus (Bull.: Fr.) P. Karst. (Uhl 1988a) and Xerocomus subtomentosus (L.: 
Fr.) Quel. (Palfner & Agerer 1995). 

For the evolution of the 'boletoid rhizomorph type' a progression can therefore be seen 
from rhizomorphs with consistent clamps to clamps only present on 'ruimer hyphae' to a 
complete loss of clamps. 

A further argument for an early evolution of this rhizomorph type comes from an un- 
identified parasite oi Strobilomyces floccopus rhizomorphs. A highly differentiated and very 
specialized 'soaking organ' could be found within the rhizomorphs (fig. 6). A clampless, thick- 
walled, brown hypha inflates subterminally balloon-like. Several very thin and bent hyphae 
grow out of the inflation and protrude between the rhizomorph hyphae. At the transition zone 
from the inflated part to the thick-walled hypha, an internal slightly dextrinoid calotte is 
formed into which a rostrum developed. This anatomically highly differentiated organ pro- 
bably also functions in a highly specialized manner. The dextrinoid calotte is perhaps a poorly 
dissolvable carbohydrate synthesized from soluble carbohydrates possibly soaked up by the 
thin, maccaroni-like hyphae. Due to this fixation, the soluble carbohydrates are re-moved from 
the osmotically effective pool to ensure further uptake of sugars. The rostrum might dissolve 
the carbohydrate again, and transport it away. Such a highly sophisticated haustorium should 
have a rather long evolutionary history, also indicating 'boletoid rhizomorphs' as ancient 
organs. Such an organ has previously never been found (comp. Jeffries & Young 1994). 



B. Amyloidy 

Amyloidy is a characteristic which has been used in systematics of Basidiomycota for a long 
time, particularly in the Agaricales s.l. (e.g. Singer 1986), and in the so-called Aphyllopho- 
rales (Jülich 1981, 1984). In Hymenomycetes, spores or hyphae can be amyloid (Jülich 
1981, 1984, Singer 1986), whereas in Urediniomycetes and Ustilaginomycetes, amyloid 
structures are unknown (comp. Jülich 1981, 1984). 

Several families of Hymenomycetes (family affiliation of genera according to Hawksworth 
et al. 1995) have genera with at least a few species with amyloid spores: Agaricaceae {Cysto- 
derma, Lepiota), Aleurodiscaceae {Acanthobasidium, Aleurocystidiellum, Aleurodiscus), Ama- 
nitaceae (Amanita), Amylocorticiaceae (Amyloathelia, Amylocorticium, Hypochniciellum, Irpi- 
codon, Melzericium, Plicatura), Asterostromataceae (Asterostroma), Atheliaceae (Hypo- 
chnopsis), Auriscalpiaceae (Auriscalpium, Dentipellis), Bondarzewiaceae (Bondarzewia), Bole- 
taceae (Tubosaeta, Heinemann & Rammeloo 1989), Clavariadelphaceae (Ceratellopsis), 
Clavicoronaceae (Clavicorona), Coriolaceae (Anomoporia, Heterobasidion, Postia, Tyromyces 
(Watling, pers. comm.)), Dichostereaceae (Dichostereum), Elasmomycetaceae (Elasmomyces, 
Gymnomyces, Macowanites, Martellia, Zelleromyces), Gloeocystidiellaceae (Boidinia, 
Gloeocystidiellum, Gloiodon, Laxitextum, Megalocystidium, Pseudoxenasma, Scytinostromella, 
Vesiculomyces), Hericiaceae (Arthomyces, Creolophus, Dentipratulum, Hericium, Mucronella), 
Hygrophoraceae (Neohygrophorus), Lachnocladiaceae (Scytinostroma, Vararia), Lentariaceae 
(Lentaria), Lentinellaceae (Lentinellus), Lentinaceae (Panus), Rhizopogonaceae (Rhizopogon, 
Smith & Zeller 1966), Russulaceae (Lactarius, Russula), Schizophyllaceae (Plicaturopsis), 
Scutigeraceae (Albatrellus), Stereaceae (Amylostereum, Laurilia, Stereum, Xylobolus), Tricho- 



20 



lomataceae {Armillaria, Baeospora, Cantharellula, Catathelasma, Clitocybula, Dermoloma, 
Dictyopanus, Fayodia, Filoboletus, Hydropus, Leucopaxillus, Melanoleuca, Mycena, 
Porpoloma, Pseudoarmillariella, Pseudoclitocybe, Pseudoomphalina, Xeromphalina), Typhu- 
laceae (Typhula) and Xenasmataceae {Aphanobasidium). 

Only a few genera are reported to possess species with amyloid hyphal walls: Albatrellus 
(Agerer et al. 1996a, Jülich 1984; Scutigeraceae), Amylocystis, Antrodia (Jülich 1984; Co- 
riolaceae), ßo/e/op5-/.s (Agerer 1992a; Bankeraceae, Stalpers 1993), Boletellus (Imler 1950; 
Boletaceae), Boletus sect. Luridi and Calopodes (Imler 1950, Singer 1986; Boletaceae); Bole- 
tinellus (Bruns, pers. comm.), Chroogomphus (Agerer 1990, Singer 1986; Gomphidiaceae), 
Deigloria (Singer 1986; Tricholomataceae), DentipelUs (Jülich 1984; Auriscalpiaceae), De- 
scolea, Descomyces (Agerer, in prep.; Bolbiliaceae), Gastroboletus (Miller 1971; Boleta- 
ceae), Gomphidius (Agerer 1991a, Singer 1986; Gomphidiaceae), Hericium (Harrison 
1964, Imler 1950, Jülich 1984; Hericiaceae), Macrolepiota (Agerer, in prep., Agaricaceae), 
Mycena {S>mGER 1986; Tricholomataeae), Omphalotus (Miller 1971, Singer 1986; Ompha- 
lotaceae), Paragyrodon (Imler 1950; Gyrodontaceae), Polyporoletus (Agerer et al. 1998; 
Scutigeraceae), Polyporus (Singer 1986; Polyporaceae), Pulveroboletus (Imler 1950, Bole- 
taceae), Pseudotomentella (Agerer 1994; Thelephoraceae), Rhizopogon (Agerer et al. 1996a; 
Rhizopogonaceae), Rozites (Agerer 1999a, Singer 1986; Cortinariacae), Scleroderma 
(Ingleby 1999; Sclerodermataceae), Thelephora (Agerer 1991a, Thelephoraceae), Tomen- 
tella {Agerer 1996a; Thelephoraceae), Tricholoma (Miller 1971; Tricholomataceae), Tubo- 
saeta (Heinemann & Rammeloo 1989; Boletaceae), Xerocomw^ (Imler 1950; Boletaceae). 

Sometimes (additional) amyloid hyphae can be found in fruitbodies of Rhizopogon, and in 
ectomycorrhizae of some Suillus species. But these hyphae originate from Chroogomphus and 
Gomphidius growing as foreign hyphae in these genera (Agerer 1990, 1991, Agerer et al. 
1996a). These hyphae are not considered in the above compilation, Gastroboletus possibly 
being an exception. 

Amyloidy could therefore be proven in 14 orders consisting of 36 different families of 
Hymenomycetes: Agaricales s.l. (Agaricaceae, Amanitaceae, Bolbiliaceae, Cortinariaceae, Hy- 
grophoraceae, Tricholomataceae), Boletales s.l. (Boletaceae, Gomphidiaceae, Gyrodontaceae, 
Omphalotaceae, Rhizopogonaceae, Sclerodermataceae), Bondarzewiales (Bondarzewiaceae), 
Cantharellales s.l. (Clavariadelphaceae, Scutigeraceae, Typhulaceae), Hericiales (Auriscalpia- 
ceae, Clavicoronaceae, Gloeocystidiellaceae, Hericiaceae, Lentinellaceae), Gomphales (Lenta- 
riaceae), Hymenochaetales (Asterostromataceae), Lachnocladiales (Dichostereaceae, Lachno- 
cladiaceae), Poriales (Coriolaceae), Polyporales (Polyporaceae), Russulales (Elasmomyceta- 
ceae, Russulaceae), Schizophyllales (Schizophyllaceae), Stereales (Aleurodiscaceae, Amylo- 
corticiaceae, Atheliaceae, Stereaceae, Xenasmataceae) and Thelephorales (Bankeraceae, Thele- 
phoraceae). Some of the mentioned orders form a natural relationship, viz. Bondarzewiales, 
Hericiales, Russulales (comp. Oberwinkler 1977a). The order Agaricales is according to 
Kühner (1980) possibly heterogeneous. As Tricholomatales and Pluteales are segregations of 
Agaricales s.l., 13 different relationships possess species with amyloid structures. 

Provided that the amyloid reaction is consistently based upon the same chemical com- 
pound, the question arises whether this feature has evolved convergently in Hymenomycetes 
several times, or whether it is a plesiomorphic character and has been lost repeatedly during 
evolution. 

Analysis of spore walls and amyloid hyphae of the Hericiales-Russulales complex (Dodd 
& McCracken 1972), and of some additional Hymenomycetes (Blackwell et al. 1985) 
revealed a wall bound, short-chain starch as the amyloid substance (Dodd & McCracken 



21 



1972). They suggested that the amyloid spore wall inhibits oxygen uptake resulting in a slow 
metabolism within the dormant spores, and the cold-water solubility of the short-chain com- 
pound should guarantee that the spores are able to germinate only after enough rainfall (Dodd 
& McCracken 1972). This hypothesis however, does not take into account that many 
spores have only amyloid warts or ridges and therefore no continuous amyloid protecting 
shield, and it does not explain why in some fungi walls of undifferentiated hyphae and of 
cystidia are amyloid. 

An additional hypothesis might claim the amyloidy primarily as a feature of hyphae. Since 
starch is a substance well courted, utilized by many organisms, it could, as a consequence of 
evolution, be withdrawn from substrate hyphae and transferred to cystidia or spores, to keep 
it from competing organisms. In this case, starch in spore walls could be a triggering food for 
microorganisms which, after biological degradation, might enable the spores to germinate. This 
hypothesis of starch retraction from hyphae is confirmed by partially amlyoid hyphae which 
show the blue reaction only in septa, or in central swellings of septa. This feature is realized 
by hyphae of mycorrhizal mantles or/and rhizomorphs in several species, e.g. Gomphidius 
species (Agerer 1990, 1991), Thelephora terrestris (Agerer 1991a), Albatrellus ovinus 
(Agerer et al. 1996b), Pseudotomentella tristis (Agerer 1994). Mycorrhizal hyphae are 
distinctly amyloid in, e .g. Albatrellus ovinus (Agerer et al. 1996b), Polyporoletus sublividus 
(Agerer et al. 1998), Rhizopogon subcaerulescens A. H. Smith (Agerer et al. 1996a), and 
Scleroderma sinnamarense Mont. (Ingleby 1999). In Rozites caperatus the whole gelatinous 
mantle is strongly amyloid, but soil hyphae only partially (Agerer 1999a). 

A possible, perhaps step-wise loss of amyloidy is shown when substrate hyphae are 
compared to stipe base hyphae and hyphae of remaining parts of the fruitbodies. For instance, 
all members of the Thelephoraceae with amyloid portions of soil or mycorrhizal hyphae (as 
above mentioned) do not show any amyloidy in their fruitbodies. Albatrellus ovinus and 
Polyporoletus sublividus have in comparison to mycorrhizal hyphae less distinctly amyloid 
hyphae in their stipe base (Agerer et al. 1996b, 1998). Gomphidius species ought to have no 
amyloid structures in their fruitbodies (Miller 1964, Singer 1986), but they show amyloid 
septa or some amyloid walls in mycorrhizal and soil hyphae (Agerer 1991a), and even in cap 
tissue of G. maculatus (Agerer 1991a). Chroogomphus has distinctly amyloid fruitbody, but 
soil hyphae and mycorrhizal hyphae are less amyloid (Agerer 1990). In some Boletus 
species, fruitbody plectenchyma are amyloid, e.g. in B. calopus Fr. and B. luridus (Engel et 
al. 1983), whereas the rhizomorphs are amyloid only in B. calopus (s. below). Although the 
mycorrhizae of/?, caperatus are distinctly amyloid, the soil hyphae are less amyloid, and even 
less amyloid than the stipe base hyphae (Agerer 1999a); the remaining fruitbody hyphae are 
only partially amyloid (Moser 1987). 

In this context, amyloid substrate hyphae of Thelephoraceae, Gomphidius, Scleroderma, 
and Polyporoletus sublividus, appear to be remnants of formerly more pronouced amyloidy. 
\n Albatrellus ovinus amyloidy has been retained in substrate and stipe base hyphae, in Bole- 
tus luridus amyloidy is, in contrast to B. calopus and Rhizopogon subcaerulescens, 
completely retracted to the fruitbody. 

Besides amyloidy, also the distribution of clamps over mycorrhizae, soil hyphae, stipe 
base and remaining parts of fruitbodies show that substrate hyphae, i.e. mycorrhizal hyphae 
in ectomycorrhizal species, are equiped with rather plesiomorphic characters (Agerer 1991a, 
Raidl 1997). But mycorrhizal mantles appear often in a more advanced state regarding 
reduction of clamps than substrate or emanating hyphae. This is apparent in ectomycorrhizae 



22 



of Rozites caperatus and of Inocybe spp., where mantle hyphae are clampless, but emanating 
and soil hyphae form them still (Agerer 1999a, Beenken et al. 1996). 

According to Swann & Taylor (1995) Hymenomycetes can based upon sequences of 18S 
rDNA be divided into two subclasses: subclass Tremellomycetidae characterized by extra- 
cellular amyloid substances (EAS) of their yeast stages, and Hymenomycetidae not producing 
such substances, if yeasts are formed at all (Prillinger et al. 1991, Taylor & Swann 1995). 
Studying the phylogenetic classification of pore-forming fungi utilising mitochondrial DNA, 
HIBBETT& Donoghue (1995) found several clusters, with Stereum at the base, followed by a 
cluster including Polyporus, which is a sister group of all other species with the Bondarzewi- 
ales-Hericiales-Russulales complex as a basal group. Stereum is known to form amyloid spores 
as is true of the Bondarzewiales-Hericiales-Russulales complex. The cluster containing Poly- 
porus (with amyloid stipe hyphae, see above) comprises no further examples of known amy- 
loidy. The heterogeneous assemblage of the sister group of the Bondarzewiales-Hericiales-Rus- 
sulales complex contains members of the Boletales. The basal clustering of Stereum and of the 
Bondarzewiales-Hericiales-Russulales complex might therefore indicate that amyloidy could 
be regarded as a rather old character (comp, also Hibbett et al. 1997b, Bruns et al. 1998). 
All these considerations lead to the conclusion that, firstly, amyloidy can most likely be 
regarded as plesiomorphic at least in the Hymenomycetidae and lost several times during evo- 
lution, but in other species amyloidy is, secondly, an erratic feature, which possibly hints at 
atavisms or an convergent development. As amyloidy is representative of Tremellomycetidae 
yeasts, it is tempting to speculate that amyloidy of Hymenomycetidae is derived from EAS of 
Tremellomycetidae. Rozites caperatus is particularly interesting with reference to its gela- 
tinous, strongly amyloid mantle, reminiscent of EAS of Tremellomycetidae. As amyloidy is 
hitherto unknown in the heterobasidomycetous lineage of the Hymenomycetidae and in the 
groups with continuous parenthesomes (comp. Bruns et al. 1998, Hahn et al. 2000, Hibbett 
et al. 1997b, Oberwinkler 1977a, Swann & Taylor 1995), amyloidy could perhaps also 
exclusively be a plesiomorphic feature of the homobasidiomycetous lineage of Hymenomyce- 
tidae with perforate parenthesomes. In either case, amyloidy seems to represent a plesio- 
morphic character in the orders of the present contribution inclusive of the Russulales, Heri- 
ciales, Bondarzewiales, and Stereales complex (Brums et al. 1998, Hibbett et al. 1997b). 



C. Plesiomorphic versus apomorphic anatomical characters 

Phylogenetic progressions within Boletales have already been compiled by Bresinsky & 
Wittmann-Bresinsky (1995), which can completely be accepted here. Some features have to 
be repeated. Others are added and arranged in an evolutionary sense from the plesiomorphic 
(left) to the apomorphic state (right). For evolution of ectomycorrhizal features compare also 
Agerer (1996c). \ 

Rhizomorphs lacking (outgroup) -> rhizomorphs present 

Rhizomorphs undifferentiated -^ rhizomorphs differentiated 

Rhizomorphs of the 'uniform-compact type' -> rhizomorphs of the 'phlegmacioid type' 

Rhizomorphs of the 'uniform-compact type' -^ rhizomorphs of the 'boletoid type' 

Backward oriented ramifications lacking -> backward oriented ramifications present 



23 



Rhizomorphs without a 'runner hypha' -^ 'runner hypha' present 

Backward oriented hypha above the septum of a 
hyphal sidebranch growing only towards proximal 

end of 'runner hypha' -^ this backward growing hypha forking 

into a distally and a proximally 
oriented branch 
Rhizomorphs without nodes -^ nodes formed 

Rhizomorphs with clamps throughout -> only running hypha with clamps -> 

-^ clamps completely lacking 
Rhizomorph septa complete -^ septa dissolving 

Rhizomorph marginal hyphae undifferentiated -^ with inflated cells 

Substrate hyphae amyloid -> substrate hyphae inamyloid 

Saprotrophic -> ectomycorrhizal 

Mycorrhizal mantle plectenchymatous -> pseudoparenchymatous 
Mycorrhizal mantle undifferentiated (type B, 

Agerer 1987-1998) -> with ring-like hyphal bundles (type A) 

Mycorrhizal mantle undifferentiated (type B) -^ with inflated cells (type F) 

Mycorrhizal mantle undifferentiated (type B) -> with cystidia (type D) 

Fruitbodies with exposed hymenium (ballistosporic) -> fruitbodies gastroid (statismosporic) 
Fruitbodies resupinate -> as consoles -> excentrically stipitate -> 

-> centrally stipitate 
Fruitbodies amyloid -> amyloidy lacking (-> amyloidy present) 

Set of metabolites complete -^ not complete 



The Relationships of Boletales s. I. 

A. The outgroup (fig. 2) 

Rozites caperatus is chosen as an outgroup, because this species combines several features 
which are regarded as plesiomorphic. 

Rhizomorphs are lacking, substrate hyphae form clamps, simple septa occur only in the 
mycorrhizal mantle. The walls of mantle hyphae and the gelatinous matrix are distinctly amy- 
loid, emanting hyphae are only in patches or in septa amyloid. The stipe base possesses 
amyloid hyphae, too; in the trama amyloid hyphae occur sporadically (Agerer 1999a). The 
formation of ectomycorrhizae however, is apomorphic. The fruitbody morphology is derived 
(Singer 1986) as are the ornamented, brown, dextrinoid spores (Singer 1986). Clamps in the 
fruitbody, the lack of cystidia and a regular gill trama (Singer 1986) are plesiomorphic 
characters. 

B. The Atromentin-Group (fig. 2) 

For Omphalotaceae, Thelephorales, and Boletales atromentin (Fl) is the uniting compound, 
which has been detected in several species (Besl & Bresinsky 1997, Gill & Steglich 
1987), and it holds a key position for the biosynthesis of cyclopentenones (F17-F20), thele- 
phoric acid (F2) and pulvinic acids (F3) (Besl & Bresinsky 1997). Atromentin (Fl) could 
not be found in Gomphidiaceae. Since atromentin is known as a precurser compound of pul- 



24 



vinic acids (F3) (Bresinsky 1977, Kämmerer et al. 1985), the absence of this substance from 
the pulvinic acids containing Gomphidiaceae is not of great importance. 

The occurrence of sesquiterpenoids in Omphalotaceae and of similar ones in Russulales was 
mentioned by Kämmerer et al. (1985) without any further systematic considerations. As 
amyloidy is characteristic of Russulales, and this relationship clusters with respect to DNA- 
sequences close to Boletales and Thelephorales (Bruns et al. 1998, Hibbett et al. 1997b), the 
Russulales should have also been included in the present comparison, but this would clearly be 
beyond the scope of the present contribution. The basal connection of Thelephorales and 
Boletales as shown in fig. 1 , might possibly have to be modified somewhat in this case. 

1. Omphalotaceae (fig. 2) 

In Omphalotuis oleahus, besides atromentin (Fl), a rich diversity of pigments could be 
found, characteristic are thelephoric acid (F2) (Gill & Steglich 1987), pulvinic acids (F3) 
(Gill & Steglich 1987), and the cyclopentenone gyroporin (F18) (Gill & Steglich 1987). 
Lampteromyces japonicus (Kawamura) Sing, contains in addition gyrocyanin (Fl 7) (Kämme- 
rer et al. 1985). This saprotrophic family causes a white rot, the test for phenoloxidases and 
peroxidases being positive (Kämmerer et al. 1985, Agerer et al., in prep.). 

The family Omphalotaceae was suggested by Bresinsky (Kämmerer et al. 1985), and 
founded basically on the presence of pulvinic acids (F3) and sesquiterpenes and on the pro- 
duction of a white rot. All Boletales are either ectomycorrhizal or cause a brown rot (Kämme- 
rer et al. 1985, Nilsson & Ginns 1979). The occurrence of thelephoric acid (F2), was 
apparently rated as of minor importance. A possible relation to Thelephoraceae was not 
discussed. 

The rhizomorphs of Omphalotus olearius and O. atraetopus belong to the 'uniform- 
compact type', clamps are present throughout, simple septa and backward ramifications of 
hyphae are lacking. Rhizomorphs are very infrequent and only formed outside the woody sub- 
strate (Agerer, pers. obs.). This features label Omphalotaceae as very primitive. The presence 
of a large set of pigments (thelephoric acid (F2), atromentin (Fl), pulvinic acids (F3), gyro- 
porin (F18)), is according to Bresinsky & Wittmann-Bresinsky (1995) also an indication of 
plesiomorphy. Amyloid hyphae in culture (Miller 1971, Singer 1986) corroborate this 
assumption. 

Since the rhizomorphs are of a very primitive type, without any apomorphic character, it is 
difficult to find relatives of this family, if the Boletales are not regarded as related. Such rhizo- 
morphs are also known in the Tricholomatales (e.g. Laccaria, Raidl & Agerer 1992; Ripar- 
tites, Agerer, in prep.; Cystoderma, Agerer, in prep.), and Agaricales ss. Kühner (Coprinus, Psa- 
thyrella, Agerer, in prep.). On the other hand, several Thelephorales form 'uniform-compact 
rhizomorphs'. But all produce backward branching hyphae, when rhizomorphs are formed. 

2. Thelephoric acid group (fig. 2) 

The order Thelephorales is equipped with thelephoric acid (F2) (Bresinsky & Renn- 
SCHMID 1971, Gill & Steglich 1987), but this compound can also be sporadically found in 
the Boletales, Omphalotaceae (s. above), and Coriolaceae (Gill & Steglich 1987, Besl & 
Bresinsky 1997). In addition, Thelephorales are traditionally characterized, as far as the 
families Thelephoraceae and Bankeraceae ss. Stalpers (Stalpers 1993) are considered, by 
mostly brown spores with a diversity of ornamentations, i.e. warts, double-warts, spines 
(Oberwinkler 1977a, 1977b, Stalpers 1993). The latter family was erected by Donk 
(1961) for genera with white spores, even in outline, and a fenugreek odour. Oberwinkler 



25 



(1977a) desired a more precise definition and delimitation of Bankeraceae, if considered 
separate from Thelephoraceae. Stalpers (1993) interprets this family in a wider sense and in- 
cludes all genera with stipitate fruitbodies. Based on the presence of thelephoric acid and great 
similarities in mycorrhizal features between Albatrellus ovinus and Thelephora terrestris, 
Agerer et al. (1996b) discussed a closer relationship between Scutigeraceae and Thelepho- 
rales. 

When below-ground-featiires of Thelephorales are included, five different groups can be 
distinguished. However, very limited data are available to attempt more detailed systematical 
conclusions. 

All members of Thelephoraceae are, as far as investigated, ectomycorrhizal and 
characterized by brown or brownish, rather thick-walled soil hyphae (Agerer 1996, Agerer 
& Weiss 1989, Agerer et al. 1995, Azul et al. 1999, Danielson & Prüden 1989, Jakucs et 
al. 1997, Jakucs & Agerer 1999a, Köljalg 1992, Raidl 8c Müller 1996); Pseudo- 
tomentella tristis is an exception (Agerer 1994). In ectomycorrhizal mantle structure this 
family is rather heterogeneous. Three main groups can be designated (fig. 2). One group is made 
up by some Tomentella species (Danielson & Prüden 1989) and Pseudotomentella tristis 
(Agerer 1994) which do not form backward oriented hyphal branches and rhizomorphs. 
They have a variety of ectomycorrhizal mantles. Pseudoparenchymatous mantles with cysti- 
dia are realized and plectenchymatous ones which lack both a hyphal pattern and cystidia. 
The second group possesses 'uniform-loose 'or 'thelephoroid rhizomorphs', but no nodes and 
conical side-branches. Representatives of this group are Thelephora terrestris (Agerer & 
Weiss 1989), Tomentella albomarginata {Agerer 1996a), T crinales (Fr.) M.J.Larsen (Köl- 
jalg 1992), T. galzinii (Burt) Bourd. & Galz. (Jakucs et al. 1997, Jakucs et al., in prep.). 
Plectenchymatous and pseudoparenchymatous mantles and cystidia can be found. The third 
group reveals nodulose rhizomorphs {Tomentella ferruginea (Pers.: Fr.) Pat., Raidl & 
Müller 1996; T. pilosa (Fr.) Bourd. & Galz., Jakucs & Agerer 1999a), and conical side- 
branches are in addition known from an unidentified Tomentella species (fig. 7; ''Quercirhiza 
nodulosomorphd", AzuL et al. 1999). Cystidia occur exclusively on mycorrhizal mantle and 
rhizomorphs of T. pilosa. , 

The rhizomorph ontogeny of some Thelephoraceae reminds one of that of the Boletales. 
This has been studied in detail in Thelephora terrestris (Agerer 1988a) and in '"Quercirhiza 
nodulosomorphd" (fig. 7). A backward oriented hypha is growing out above a clamp of a side- 
branch. It forks and one branch grows in proximal and the other in distal direction of the main 
filament. This is not a typical 'runner hypha' with distant clamps, and it does not inflate 
during later ontogenetic phases, possibly because these hyphae thicken their brown walls very 
early, and therefore impeding an enlargement. Köljalg (1996) lays emphasis on rhizomorphs 
of tomentelloid fungi. He demonstrated both monomitic and dimitic examples and regards this 
feature as important on genus or subgenus level. 

Some of the Thelephoraceae possess amyloid hyphal portions, seen in Thelephora 
terrestris (AGERER 1991a), Pseudotomentella tristis (Agerer 1994), Tomentella albomargi- 
nata (Agerer 1996a). Unfortunately this is a feature tested only by a few authors and for a 
very limited number of species. 

Apart from rhizomorphs of Thelephoraceae which show considerable heterogeneity, mantle 
types of ectomycorrhizae must also be regarded as possible determinants of relationships 
within this family. An example for a probably necessary systematical transfer from Thelepho- 
raceae to Bankeraceae based on mycorrhizal features is Tomentellopsis submollis (see below). 



26 



Bankeraceae, which have been examined for ectomycorrhizae and rhizomorphs all possess 
plectenchymatous mantles without pattern (Boletopsis leucomelaena, Agerer 1992a; Hydnel- 
lum peckii, Agerer 1993) or ring- or star-like arranged hyphal bundels {Bankera fuligineo- 
alba, Agerer & Otto 1997; Phellodon niger, Agerer 1992c; Sarcodon imbricatum, Agerer 
1991c). The Rhizomorphs are either of the 'uniform-compact type\Bankera fuliginea-alba, 
Agerer & Otto 1997, Phelldon niger, Agerer 1992c) or of the 'phlegmacioid type' {Bole- 
topsis leucomelaena, Agerer 1992a; Hydnellum peckii, Agerer 1993; Sarcodon imbricatum, 
Agerer 1991c). 'Phlegamcioid rhizomorphs' andring- or star-like hyphal bundles of mantles 
are hitherto unknown in Thelephoraceae. The mycorrhizae of Tomentellopsis submollis fit to 
the Bankeraceae with their 'uniform-compact rhizomorphs' and ringlike organized mantles 
(Agerer 1998b). 

Amyloidy is known only exceptionally in Bankeraceae. Amyloid points are found on 
substrate hyphae oi Boletopsis leucomelaena {Agerer 1992a). Imler (1950) reports on an 
"apparent amyloidy" of some Bankeraceae, but this amyloidy is based on intracellular amy- 
loid particles and not in cell walls. 

Interesting is the systematic position of Pseudotomentella tristis. As mentioned above, this 
species does not form thick-walled, dark brown substrate hyphae. It therefore is not well 
placed in the Thelephoraceae/Scutigeraceae-cluster of fig. 2, but would better be grouped 
within the Bankeraceae. A confirmation of such a position is provided by Bruns et al. (1998), 
who could show a closer connection with the Bankeraceae Sarcodon than with all Thele- 
phoraceae studied. With Pseudotomentella tristis, a member with amyloid septa and hyphal 
walls and lacking rhizomorphs, would be included in Bankeraceae. Generally, the rhizmorphs 
and mycorrhizae of Thelephoraceae and Bankeraceae are too little known to get a more 
detailed impression of their possible role in discerning relationships within this group. 

The mycorrhizal features of the Scutigeraceae ^/Zjüf/re//w5 ovinus (Agerer et al. 1996b), A. 
subrubescens (Pillukat, in prep.) and Polyporoletus sublividus (Agerer et al. 1998) are so 
simliar to those of Thelephora terrestris (Weiss & Agerer 1989) that an inclusion in 
Thelephorales appears justified (Agerer et al. 1996b) in spite of the smooth, colourless 
spores and their phlegmacioid rhizomorphs. Colourless (and warty) spores are however, also 
known in Bankeraceae. The pigment composition also hints at such a relationship, since thele- 
phoric acid (F2) could be detected (Agerer et al. 1996b), and atromentin (Fl) occurs (Gill & 
Steglich 1987). The synthesis of cyclopentenones indicates a possible relation to the 
Boletales (Bresinsky 1996), but pulvinic acids (F3) have not been found (Bresinsky 1996, 
Gill & Steglich 1987). Amyloid substrate hyphae (Agerer et al. 1996b) put Scutigeraceae 
close to Thelephoraceae. 

HiBBETTetal. (1997b) show a basal position of Hydnellum sp. and Thelephora sp. to the 
Boletales. Albatrellus syringae however, is in a different cluster, but not very distant from 
Hydnellum sp. and Thelephora sp. According to Bruns et al. (1998) Albatrellus and Poly- 
poropletus form a sister cluster to the Thelephorales. 

3. The pulvinic acid group (fig. 2, 3) 

Different pulvinic acids (F3) occur in this group (Gill & Steglich 1 987), although in some 
species they are lacking (comp. Strobilomycetaceae, Leccinum). A general feature are back- 
ward growing hyphal branches. 

The Boletales suborder Suillineae is basically only hold together by DNA data (Brums & 
SzARO 1990, Bruns et al. 1998, Kretzer & Bruns 1999). A great diversity of below-ground 



27 



features is realized. This suborder includes the Truncocolumellaceae, Gomphidiaceae, Suilla- 
ceae and Rhizopogonaceae. 

a. The group with 'uniform-compact rhizomorphs' (Truncocolumellaceae) (fig. 2) 
Truncocolumella, is regarded by Bresinsky (1996) together with Boletus and Xerocomus in 

one group. Truncocolumella citrina is the type species, whilst the second one, T. rubra, has 
been transferred to Gastroboletus (comp. Besl & Bresinsky 1997). Hawskworth et al. 
(1995) include this genus in Rhizopogonaceae. 

The rhizomorphs are undifferentiated of the 'uniform-compact type' (figs. 9, 10), unlike 
those of Rhizopogonaceae and Boletaceae (s. below). Undifferentiated rhizomorphs have also 
been described and depicted by Eberhart & Luoma (1996). Short-celled, thick-walled 
hyphae on thicker rhizomorphs and at the growing tip are hitherto unique. Clamps as a 
primitive feature are consistently present; amyloidy is lacking. Truncocolumella citrina forms 
ectomycorrhizae and chlamydospores (Eberhart & Luoma 1996). 

The spores of T. citrina are short-ellipsoid (Zeller 1939), thick-walled and dextrinoid (Smith 
& Singer 1959) whereas those of Rhizopogon are mostly thin-walled, elongate and mostly not 
dextrinoid (Martin 1996, Smith & Zeller 1966), they are amyloid in R. sect. Amylopogon 
(Smith & Zeller 1966). The shape and dextrinoidy of T. citrina spores reminds one of the 
spores of Tapinella atrotomentosa and T. panuoides (Hahn & Agerer 1999a); Tapinella spp. 
also produce chlamydospores (Hahn & Agerer 1999a). Chlamydospores are known in the 
tuberculate extomycorrhizae oi Rhizopogon vinicolor, too (Müller & Agerer, in prep.). 
The basal position of T. citrina in comparison to Rhizopogon is corroborated by sequences of 
the ITS region of rDNA (Kretzer et al. 1996), whereas in other DNA-studies a clear 
distinction from Rhizopogon and Suillus was impossible (Bruns et al. 1998, Kretzer & 
Bruns 1999). The primitive organization of the rhizomorphs of T. citrina separate Truncoco- 
lumellaceae systematically from Suillaceae and Rhizopogonaceae and Boletaceae (see below). 

b. The amyloid, episymbiotic group with 'thelephoroid' or lacking rhizomorphs 
(Gomphidiaceae) (fig. 2) 

The Gomphidiaceae is with respect to substrate hyphae a well defined taxon. Members 
form mycorrhizae with plectenchymatous mantles with differently, species-specifically shaped 
cystidia. Cystidia and thick-walled cells or septa of mantle or emanating hyphae and/or rhizo- 
morphs are amyloid (Agerer 1990, 1991). Emanating hyphae of mycorrhizae and rhizo- 
morph hyphae are often thick- walled and slightly brownish. Rhizomorphs are only produced 
by ectomycorrhizae of Gomphidius glutinosus and G. roseus and are at most slightly 
differentiated of the 'uniform-loose type' or of the 'thelephoroid type'(AGERER 1991a). Fruit- 
bodies lack rhizomorphs. Stipe base cystidia are similarly shaped to those of mycorrhizae and 
are slightly amyloid, even in Gomphidius glutinosus, G. maculatus and G. roseus (Agerer 
1991a). The pileus trama of G. maculatus is furnished with a few amyloid hyphae. Gom- 
phidius ought to be, in contrast to Chroogomphus, inamyloid (Singer 1986, Miller 1964). 

Similar mycorrhiza-cystidia are only known from Alhatrellus ovinus (Agerer et al. 1996b), 
A. subrubescens (Pillukat, in prep.), Thelephora terrestris (Agerer & Weiss 1989) and some 
Tomentella species (Agerer, unpubl.). The cystidia of Strobilomyces floccopus (fig. 22) and of 
Rhizopogon vinicolor (fig. 25) are produced on 'boletoid rhizomorphs', reveal a deviating 
ontogeny and lack amyloid reactions (see below). 

Clamps are frequent in mycorrhizal mantles and rhizomorphs of Gomphidiaceae, they 
become infrequent at the stipe base and are lacking in the fruitbody. This is an evident indi- 



28 



cation that features of mycorrhizae and rhizomorphs are plesiomorphic in comparison to 
those of fruitbodies (see above). The gradual reduction of amyloidy in Gomphidius from 
mycorrhizae to fruitbodies parallels this situation. 

A general feature of all Gomphidiaceae (five species tested, Agerer 1991a, 1996d) is their 
aibility to grow their hyphae within mantles and/or rhizomorphs of Rhizopogon and Suillus 
species (Agerer 1990, 1991, 1996d, Agerer et al. 1996a). Gomphidius glutinosus differs 
strikingly from G. maculatus, G. roseus, Chroogomphus helveticus (Sing.) Mos. and C. rutilus 
(Schaeff: Fr.) O.K.Miller, in growing in foreign rhizomorphs only inamyloid hyphae, whereas 
the latter four species produce amyloid, coil-like haustoria within cortical cells, which are 
enveloped by the Hartig net of the Suillus or Rhizopogon ectomycorrhizae. Primordia of 
Gomphidius glutinosus and of Chroogomphus helveticus ssp. tatrensis (Pilat) Kuthan & Sing, 
are found to be formed directly on rhizomorphs of a Suillus and Rhizopogon species, 
respectively (Agerer 1991a, 1990). 

Similar pigment compositions of Suillus (Suillaceae) and Gomphidius (Gomphidiaceae) 
urged to postulate a closer relationship between these two genera, and "based on the 
evidence..., the close relations between the genus Suillus and Gomphidiaceae (and Rhizopogon) 
cannot be disputed" (Besl & Bresinsky 1997). The authors provided detailed compilations 
of pigments of these two genera. 

Gomphidius glutinosus, G. maculatus and G. roseus have 1 ,2,4-trihydroxybenzene (F16) 
as a characteristic pigment. It is lacking in G. subroseus Kauffm. (Besl & Bresinsky 1997). 
This compound and its dimerisation products (e.g. gomphilactone (Fl 5)) could not be detected 
in Chroogomphus (Besl & Bresinsky I.e.). Chroogomphus species are, instead of, provided 
with boviquinones (Besl & Bresinsky I.e., Gill& Steglich 1987, Gill 1999: C. helveticus: 
boviquinone-3 and -4 (F4, F5); C. rutilus: boviquinone-3 and -4 (F4, F5), diboviquinone-3,4 
and -4,4 (F6, F7), methylenediboviquinone-3,3, -3,4- and -4,4 (F8-F10), and C. tomentosus 
(Murr.) O.K. Miller: 'boviquinones'), which lack Gomphidius (Besl & Bresinsky I.e.). 

The main argument (except for DNA-data, see above) to accept a close relationship 
between Gomphidiaceae and Suillus is the presence of boviquinones (F4-F10) in Chroo- 
gomphus and in Suillus. The pigment survey for Gomphidiaceae and Suillus given by Besl & 
Bresinsky (I.e.) indicate that boviquinones and the related prenylated phenols (suillin (F14), 
bolegrevilol (F13) and tridentoquinone (F12)) occur only infrequently in the 38 Suillus species 
tested. Bolegrevilol (F13) could be detected in S. granulatus (L.) Kuntze and S. grevillei 
(Klotzsch: Fr.) Sing., suillin (F14) in S. collinitus (Fr.) Kuntze, S. granulatus, S. bellinii (In- 
zenga) Watl. and S. variegatus, bovilactone (Fll) in S. americanus (Peck) Snell in Slipp & 
Snell, S. bovinus (L.) Kuntze and S. collinitus, boviquinone-3 (F4) in S. collinitus, bovi- 
quinone-4 (F5) in S. bovinus and S. spraguei (Berk. & M. CA. Curtis in Berk.) Kuntze, di- 
boviquinone-4,4 (F7) in S. americanus and S. bovinus, methyl-boviquinone-4,4 (FIO) in S. 
bovinus, tridentoquinone (F12) (related to boviquinone-4 (F5)) in S. tridentinus (Bres.) Sing. 
In summary, from 38 Suillus species tested, in 9 species boviquinones occur. In Boletinus 
asiaticus Sing, and B. paluster (Peck) Peck the related asiaticusins could be detected (Gill 
1999). Gastrosuillus laricinus (Singer & Both) Thiers and Boletinus cavipes (Fr.) Kalchr. lack 
boviquinones. Calculated for Suillaceae, boviquinones and their derivatives are found only in 
11 of 42 species whereas thelephoric acid (F2) is present in four of these 42 species. 

Boviquinone-4 (F5) in Suillus bovinus is formed by geranygeranylation of 3,4-dihydroxy- 
benzoic acid in position 5 of the phenyl nucleus, whereas boviquinone-3 (F4) of Chroogom- 
phus rutilus origiantes by famesylation of this compound in position 2 (Gill 1999). These are 
two different biosynthesis pathways. It would be necessary to compare the biosynthesis of. 



29 



the boviquinones of all Chroogomphus and of the boviquinone-positive Suillus species. 
Possibly, this difference could indicate a convergent evolution of boviquinones in Chroogom- 
phus and Suillus. 

Besl& Bresinsky (1997) isolated not further characterized 'lipolytic pigments', i.e. pre- 
nylated phenols and quinones (e.g. boviquinones (F4-F10)) in Suillus (not in Boletinus) and 
regard them as indicators of a relationship between Suillus and Gomphidiaceae/Rhizopo- 
gonaceae and they "have not been found in any single bolete species (i.e. Boletales with 
tubulate hymenophore) outside the genus Suillus" (Besl & Bresinsky 1997). (For further 
discussion see Suillaceae and Rhizopogonaceae, below). 

The value of Chroogomphus pigments in Suillus species is relativized by the episymbiotic 
feature of Gomphidiaceae. According to Agerer (1991, 1996d; Agerer, unpubl.) amyloid 
hyphae of Gomphidiaceae (of Chroogomphus and Gomphidius) could be found in rhizo- 
morphs and/or mycorrhizal mantles and roots of ectomycorrhizae of Suillus bovinus (epi- 
symbiotic with G. roseus), S. collinitus (C. rutilus), S. granulatus (C. rutilus), S. grevillei (G. 
maculatus), S. plorans (Rolland) Kuntze (C helveticus), S. sibiricus (Sing.) Sing. (C. helve- 
ticus) and S. variegatus (C. rutilus). The situation gets even more complicated by the fact that 
Gomphidiaceae hyphae can not only grow in rhizomorphs and mycorrhizae of Suillus, but 
they might also be found in fruitbodies. This could definitely be proven in fruitbodies of 
Rhizopogon subcaerulescens (Agerer et al. 1996a) and other Rhizopogon spp. (Smith & Zel- 
LER 1966). At least Gomphidius roseus and S. bovinus can form united stipe bases. But it is 
not known how far the G. roseus hyphae reach into the fruitbody of Suillus. A microscopical 
differentiation is impossible, as Gomphidiaceae hyphae become gradually inamyloid the 
denser they are embedded between those of Suillus (Agerer 1990). The great dependence of 
G. roseus hyphae for growth from Suillus bovinus is indicated by the fact that mycelial 
cultures of G. roseus could only be obtained when trama pieces of both species were 
coinoculated on agar plates (Agerer 1991a). 

Presently, the most favoured feature for delimitation and recognition of relationships are 
sequence studies of DNA. Bruns & Szaro (1990) and Bruns et al. (1998) report a close 
relationship between Boletaceae subf. Suilloideae and Gomphidiaceae with respect to 
sequences of nuclear and mitochondrial RNA genes. These results are consistent with those 
published by Bruns & Szaro (1992). Aslo ATPase 6 data indicate a closer relationship 
between Suillus and Brauniellula, a hypogeous Gomphidiaceae (Kretzer & Bruns 1 999). ITS 
sequences of nuclear rDNA indicate however, that the genus Suillus forms one group, basal to 
it appears Rhizopogon subcaerulescens, basal to R. subcaerulescens stands Truncocolumella 
citrina and again basal to it Chroogomphus vinicolor (Peck) O. K. Miller and Gomphidius 
glutinosus together (Kretzer et al. 1996). The position of Truncocolumella citrina between 
Suillus and Gomphidiaceae might indicate a greater separation of Suillus and Gomphidiaceae, 
as earlier suggested. But the DNA-tree (Kretzer et al. 1996) is unrooted or at least arbitrary 
rooted (Bruns, pers. comm.). Therefore this tree should not be overinterpreted; and ITS 
sequences are said to be only useful for differentiation of species and genera, whereas 
sequences of nuclear rDNA could be applied for higher level taxa analysis (Bruns et al. 1991). 
The plesiomorphic characters of the substrate hyphae (no or undifferentiated rhizomorphs, 
clamps) group Gomphidiaceae and Truncocolumellaceae together and basal to Suillaceae and 
Rhizopogonaceae which have highly developed rhizomorphs and almost completely reduced 
clamps (see below); and this conflicts with some DNA-data (s. below). Whether the great 
similarities between ectomycorrhizae of some Thelephoraceae, Scutigeraceae and Gomphidia- 



30 



ceae, are the consequence of a common ancestor or the result of convergent evolution is still an 
open question. 

c. The group with boletoid rhizomorphs (Suillaceae, Rhizopogonaceae) (fig. 2) 

'Boletoid rhizomorphs' are the most elaborated conducting organs of fungi (s. above) and 
are likely to be the reason for rapid evolution of fruitbodies and possibly a prerequisite for 
production for such voluminous fruitbodies as Boletus spp.(see also 'Boletales ceteri'). Two 
main functions can be attributed to these rhizomorphs with their rapidly growing 'runner 
hyphae'. They provide after Read (1992) "an extremely effective structure with which to 
scavenge for resources and the rapid onset of infection following contact between unifected 
roots" (or more general: substrate) "and the advancing mycelial front confirms ... a very high 
inoculum potential." In the case of saprotrophs, rhizomorphs are preferentially formed bet- 
ween larger substrate gaps, whereas dispersed mycelium prevails within a nutritive substrate 
(comp. Rayner et al. 1985). 

The families Suillaceae and Rhizopogonaceae appear well characterized by some special 
features of their 'boletoid rhizomorphs' and of their mycorrhizae. 'Boletoid rhizomorphs' have 
been proven for all Rhizopogon, Suillus and Boletinus species studied to date: Rhizopogon 
vinicolor (fig. 25), R. luteolus Fr. (Uhl 1988b), R. melanogastroides M. Lange (Raidl 1998), 
R. roseolus (Raidl & Agerer 1998), R. subcaerulescens (Agerer et al. 1996a), R. vulgaris 
(Vitt.) M. Lange (Jakucs et al. 1998j, Suillus bovinus (Duddridge et al. 1980), S. collinitus 
(Uhl 1988a), S. flavus (Raidl 1997, Treu 1990), S. granulatus (Raidl 1997), S. lahcinus 
(Berk.) Kuntze (Treu 1990), S. luteus (Raidl 1997), S. placidus (Bon.) Sing. (Agerer, un- 
publ.), S. plorans (Treu 1990), S. sibiricus Sing. (Treu 1990), S. tridentinus (Treu 1990), S. 
variegatus (Raidl 1997) and Boletinus cavipes (Raidl 1997, Treu 1990). All species, 
hitherto studied exactly enough, possess crystals, which reflect in Normarski's interference 
contrast microscopy light strongly (possibly oxalate crystals) and brownish exuded drops of 
pigment (e.g. Suillus laricinus, Treu 1990). Such crystals and drops of pigment occur also in 
Rhizopogon (e.g. Rhizopogon roseolus, Raidl & Agerer 1998). Drops of pigments are 
lacking in Boletinus cavipes (Treu 1990). The brownish drops of pigment are possibly the 
"lipophilic pigments" (prenylated phenols), isolated by Besl& Bresinsky (1997), which are 
"clearly the substances that form the characteristic incrustation of the fasciculated cystidia 
which are coloured by KOH .... and considered to be indicator substances for the genus Suillus 
(and for the Gomphidiaceae/ Rhizopogonaceae)" (Besl & Bresinsky 1997). A colouration of 
the exuded drops of pigment of the rhizomorphs in KOH is also apparent (Treu 1990). Both, 
drops of pigment and these crystals, are unknown in any other family of Boletales. 

The genera Suillus and Rhizopogon are both receptors of Gomphidaceae hyphae, which 
penetrate rhizomorphs, mantle and Hartig net (see Gomphidiaceae). This is possibly also true 
of Boletinus cavipes (Agerer, unpubl.). Gomphidaceae therefore similarly indicate fungal 
relationships as has been shown for the ascomycete Hypomyces chrysospermus Tul. 
(Sepedonium), which reveals fungi as being a member of the Boletales (Gill & Watling 1986, 
Helfer 1991). That also non boletalean fungi can be hosts of H. chrysospermus has already 
been discussed (see Gautieria, above.) 

Terpenoids are regarded as important pigments of Suillaceae and Rhizopogonaceae as well 
(Bresinsky 1996, Besl& Bresinsky 1997). Some Suillus species are characterized by bovi- 
quinones (F4-F10) and derivatives (comp. Gomphidiaceae). A specific derivative has been 
detected in Rhizopogon, the ansaquinone rhizopogone, a similar compound as tridentochinone 
(F12) from Suillus tridentinus (Bresinsky & Steglich 1989, Gill 1999, Gill & Steglich 



31 



1987). A close relationship between Suillus and Rhizopogon has been generally accepted 
(Bresinsky 1996, Besl& Bresinsky 1997, Bruns et al. 1998); but there is much to be said 
for and also against a close relationship between Suillaceae and Gomphidiaceae. 

The pros regard similarities in pigment composition (Besl & Bresinsky 1997), DNA- 
sequences (Bruns & Szaro 1990, Bruns et al. 1998) and ATPase 6 structrue (Kretzer & 
Bruns 1999). This has already been discussed in some detail above. In addition, the spores of 
Gomphidiaceae and Suillaceae are similarly shaped and reminiscent of Boletaceae. Their gill 
trama is bilateral, again as in Boletaceae (Singer 1986). The contras concern fundamental 
differences in the structrue of rhizomorphs and mycorrhizae, the lack of amyloidy in Suilla- 
ceae and the consistent presence of clamps in mycorrhizae of Gomphidiaceae. 

The 'thelephoroid' or 'uniform-loose' rhizomorphs in Gomphidiaceae are even more 
primitive than some of the genus Tomentella or Thelephora. No forking of backward growing 
hyphae could be found in Gomphidiaceae. The genus Chroogomphus lacks rhizomorphs on 
fruitbodies and on ecto mycorrhizae. The absence of rhizomorphs could be interpreted as a 
reduction, or as an original state. The 'thelephoroid' rhizomorphs of Gomphidius could have 
theoretically emerged by reduction from 'boletoid' ones. But two facts contradict this 
assumption. Firstly, if they should have developed by reduction, at least the earliest - in an 
ontogenetical and in a phylogenetical sense - characteristics of boletoid rhizomorphs should 
still be found, namely 'backward oriented ramifications growing towards a main hypha, after 
they have originated above the first clamp of a side-branch', and the runner hyphae. This is 
not the case. Runner hyphae of Suillus betray, due to their occasional clamps and their early 
formation during ontogeny, their plesiomorphic character (Raidl 1997) and can hence, after 
Heckel's 'biogenetic rule', be considered phylogenetically as most primitive. And there is no 
reasonable ground, why such a functionally successful organ, as a 'boletoid rhizomorph', 
should be reduced at all, even when these fungi have evolved to grow in rhizomorphs of Suilla- 
ceae and Rhizopogonaceae. The very complicated ontogeny of 'boletoid rhizomorphs' should 
be governed by several genes. A mutation could lead to a loss of this functionally highly 
sophisticated organ and would be detrimental for the evolution of these fungi. 'Boletoid rhizo- 
morphs' should have been retained in spite of the fruitbodies' access to nutrients via foreign 
rhizomorphs and mycorrhizae at least on the ectomycorrhizae of Gomphidius . Secondly, rhizo- 
morphs are very important for the function of the symbiosis in the ectomycorrhizae. Highly 
elaborated rhizomorphs have better access to nutrients of the substrate than 'thelephoroid' 
ones (Kammerbauer et al. 1989). There is again no explanation, why such a highly specia- 
lized symbiotic relation should be reduced in its functional capability by disorganization of its 
transport organs. 'Boletoid rhizomorphs' are structurally identical whether they are organs of 
fruitbodies or mycorrhizae. In Gomphidiaceae, typical rhizomorphs are retained on the ecto- 
mycorrhizae, but reduced on the fruitbodies, when they are developed on foreign rhizomorphs. 

Gomphidiaceae mycorrhizae form very frequent clamps in their mantles and rhizomorphs, 
they are only occasionally present on the runner hyphae of some Suillus species, but occur 
frequently in Boletinus. The loss of clamps and the lack of amyloidy, both indicate only a 
distant relation of Suillus to Gomphidiaceae as do the cystidia of Gomphidiaceae, unknown in 
Suillus. Several features of below-ground structures in Gomphidiaceae reveal apparently a 
'plesiomorphy syndrome' of independent characters. 

The episymbiotic life-style opened for the Gomphidiaceae a stabile ecological niche, with- 
drawn from competition by other fungi. This was likely the reason for rapid evolution to 
pileate-lamellate fruitbodies, in spite of their hardly differentiated or lacking rhizomorphs. 
And they contemporarily developed bilateral gill trama and fusoid spores. Convergent evo- 



32 



lution of this fruitbody type is not uncommon. Tapinellaceae and Paxillaceae provide in our 
context the best example. 

The conclusion is that, if there is a closer relationship between Gomphidiaceae and Su- 
illaceae as DNA-data suggest, Gomphidiaceae should be like Truncocolumellaceae, at the base 
of the cluster which is called the 'suilloid group' by Bruns et al. (1998) or 'suilloid radiation' 
by Kretzer & Bruns (1999). The terpenoids of Chroogomphus and of Suillus and the 
'lipophilic pigments' of Suillaceae and Gomphidiaceae could therefore be regarded, inspite of 
their infrequent occurrence, as characteristic of Boletales subord. Suillineae, if boviquinones in 
Suillus are genuine compounds and if there is no distinct difference in biosynthesis of bovi- 
quinones in Chroogomphus and Suillus (Gill 1999). 

The similar modifications of four different, independent DNA-regions of Gomphidiaceae, 
Suillaceae, Rhizopogonaceae (and Truncocolumellaceae) point at a monophyletic group 
(Bruns, pers. comm.), with the consequence that the 'boletoid rhizomorphs' of Suillaceae/ 
Rhizopogonaceae have evolved independently from those of 'Boletales ceteri' (see below, and 
fig. 2, 3). A convergent evolution of these four different DNA-regions and their parallel 
divergence in comparison to those of 'Boletales ceteri' is very unlikely (Bruns, pers. comm.). 
Convergent changes in DNA-sequences however - which should not be excluded per se - 
would on the one hand relativize the importance of DNA-data sets, and on the other, could 
designate 'boletoid rhizomorphs' as a character of a monophylum Boletales s.str., including 
only and all families with 'boletoid rhizomorphs'. The boletoid rhizomorphs of Suillaceae/ 
Rhizopogonaceae on the one hand and those of 'Boletales ceteri' (figs. 2, 3) might then parallel 
the situation in the Gomphales-Geastrales-Gautieriales complex, where identical rhizomorph 
structures indicate a close relationship (s. above; Hahn et al. 2000). But, as 'boletoid rhizo- 
morphs' are also known in Trichloma batschii Gulden (Raidl 1997), a convergent evolution 
of this rhizomorph type also within Boletales can not be excluded. 

Another example which proves rhizomorphs as owners of conservative characters is 
demonstrated by the Agaricaceae and their relatives. Hibbett et al. ( 1 997, 1 997b) found from 
DNA-sequences, close similarities between the Agaricaceae (Agaricus, Lepiota), and the Ly- 
coperdaceae {Calvatia and Lycoperdon) . The studies on rhizomorphs of Lepiota, Macro- 
lepiota, Leucoagaricus, Leucocoprinus, Agaricus, Podaxis, Calvatia and Lycoperdon, again 
revealed the same structures, which are different from those, for instance, of the Stropharia- 
ceae {Hypholoma and Stropharia), or the Coprinaceae (Coprinus, Psathyrella). All the above 
mentioned genera of Agaricaceae, Lycoperdaceae and Strophariaceae have 'agaricoid' rhizo- 
morphs. Dextrinoid, thick-walled marginal rhizomorph hyphae however, occur only in the 
Agaricaceae (Agerer, in prep.). Tulostoma, also found in the same DNA-cluster as Agarica- 
ceae and Lycoperdaceae, differs however, in lacking these dextrinoid hyphae (Agerer, in 
prep.). This difference has to be discussed in more detail, but is beyond the scope of this con- 
tribution. 

Rhizopogonaceae contains, according to Bresinsky (1996) and Pegler et al. (1993), the 
two genera Rhizpogon and Alpova. Trappe (1975) regards Alpova as a link between Rhizo- 
pogon and Melanogaster, and includes Alpova in Melanogastraceae. 

As stated above, Rhizopogon forms typical 'boletoid rhizomorphs'. Some species possess 
cysti-dia. They are either terminal and colourless {Rhizopogon subcaerulescens, Agerer et al. 
1996a) or they can be geniculate by including several portions of a branched hypha 
{Rhizopogon vinicolor, fig. 25). Alpova trappei (fig. 26) and Melanogster variegatus (fig. 27), 
the only species' of these genera studied in detail, form 'boletoid rhizomorphs', too. The 



33 



mycorrhizal mantles of Rhizopogon show a high diversity. Ring-Hke hyphal bundles can occur 
(e.g. R. subcaerulescens, Agerer et al. 1996a), the mantle can be built by multiply branched 
hyphae {R. luteolus, Uhl 1988b), or the whole mycorrhizal system can be enveloped by an 
irregular tomentum of cystidia-like hyphae (Zak 1971.). Alpova diplophloeus (Zeller & 
Dodge) Trappe & A.H.Smith forms ring-like hyphal bundles on the mantle surface 
(Massicotte et al. 1989), amorphous material and bulbous cystidia (Miller et al. 1988). The 
mycorrhizae of Melanogaster are not known in detail. Bresinsky (1996) places Melanogaster 
broomeianus close to Gyrodon and Paxillus due to the presence of involutin (F20) (Besl et al. 
1996), and includes it in the Paxillaceae (Bresinsky et al. 1999). In the investigations of 
Bruns et al. (1998) Melanogaster tuberiformis clusters with the 'sulloid group'. 

At least Rhizopogon sect. Amylopogon appears with respect to amyloid spores distinct 
form the other sections. However, Smith & Zeller (1966) could find in other sections 
amlyoid spores, at least in the early stages of spore development, cautioning them not to 
describe a separate genus (Smith 8c Zeller 1966). Amyloidy in this genus appears to be 
either an atavism, similar to that found in the genus Boletus (see below) or a hint for a closer 
relationship to Gomphidiaceae and hence a primitive character, although the Suillaceae, the 
possibly older family in comparison to Rhizopogonaceae, does not show any amyloidy. A 
more detailed discussion of the genus Rhizopogon is beyond the scope of this contribution, 
but the occurrence of thelephoric acid (F2) exclusively in Rhizopogon sect. Villosuli (Gill & 
Steglich 1987, Müller et al., in prep.) aspires after more detailed systematic studies. 

d. The group with 'phlegmacioid/agaricoid rhizomorphs' (Tapinellaceae) (fig. 2) 

Tapinella J.-E. Gilbert is a segregate of Paxillus which is meanwhile widely (e.g. Bresinsky 
et al. 1999, Redhead & Ginns 1985, Sutara 1992), but not generally accepted (e.g. Hawks- 
worth et al. 1995). Many differences justify the separate genus, as comprehensively dis- 
cussed by Hahn & Agerer (1999b): Tapinella possesses in contrast to Paxillus smaller 
spores, lacks cystidia, forms bidirectional not gelatinous gjU trama (unidirectional with 
gelatinous lateral strata in Paxillus), rhizomorph structure is 'phlegmacioid' or 'agaricoid' in 
Tapinella (boletoid in Paxillus), chlamydospores occur only in Tapinella but sclerotia are 
unknown, caulohymenium and involutin are lacking, and twin-clamps occur only in Tapinella. 
Paxillus is ectomycorrhizal in contrast to Tapinella which forms a brown rot. The similar habit 
of e.g. Paxillus involutus and of Tapinella atrotomentosa (Batsch: Fr.) Sutara is hence the 
result of convergent evolution. The frequent asymmetric shape of T. atrotomentosa is 
reminiscent of the less advanced fruitbody type still realized by T. panuoides. 

These differences suggest also the existence of a separate family Tapinellaceae. Tapinella 
comprises T. atrotomentosa (fig. 11) with agaricoid rhizomorphs whereas those of T. panuoi- 
des (Fr.: Fr.) Gilb, are phlegmacioid (Hahn & Agerer 1999b) and are therefore fundamentally 
different from those of Boletales proper. DNA-sequence data place both species on a common 
branch basal to all the species of Boletales tested (Bresinsky et al. 1999, Hibbett et al. 
1997b), and supports Locquin (1981, 1984) who proposed already this family name. Accor- 
ding to Bresinsky et al. (1999) this name is however, invalidly published. 

Whether the genus Pseudomerulius belongs to this family, which Jülich (1984) included in 
the Coniophoraceae, needs further studies. Similarities to T. panuoides can be seen in the same 
type of 'phlegmacioid rhizomorphs' of P. aureus (Fr.) Jülich (fig. 11), in the existence of 
knobby outgrowths with very thin hyphae, a gelatinous trama, and as a causal agent of brown 
rot (Jülich 1984). As the spores are thin-walled, not dextrinoid and at most slightly yellow 
there is no evident relation to Tapinella; furthermore, backward ramifying hyphae are lacking 



34 



in P. aureus (fig. 12). An affiliation to Coniophoraceae therefore can be rejected. Pulvinic acids 
(F3) have not been detected (comp. GiLL& Steglich 1987), but these compounds are lacking 
also in the Coniophoraceae member Jaap/a (Besl et al. 1986). 

e. The saprotrophic group with 'boletoid rhizomorphs' (Coniphoraceae) (figs. 2, 3) 

As discussed above, possibly the first 'boletoid rhizomorphs' which originated during 
evolution were similar to those found in Jaapia ochroleuca (fig. 13). Although pulvinic acids 
(F3) have not been detected (Besl et al. 1985), and the spores are colourless or at most 
slightly yellowish, their dextrinoidy places this genus in the proximity of the brown-rot genera 
Coniophora, Leucogyrophana and Serpula. The genus Hygrophoropsis, another brown-rot 
fungus (Lindeberg 1948) with colourless and dextrinoid spores, is suggested to be transferred 
from Paxillaceae or Hygrophoropsidaceae to Coniophoraceae (Bresinsky et al. 1999). Hence 
Paxillaceae comprises, after the exclusion of Hygrophoropsis and Tapinella, only ecto- 
mycorrhizal symbionts. Whether Jaapia ochroleuca is a brown-rot fungus has still to be sub- 
stantiated; both collections examined, grew on white-rotten wood (Agerer, pers. observ.). Even 
the suggested brown-rot fungus Tapinella atrotomentosa is able to form peroxidases, but 
phenoloxidases cannot be detected (Agerer, unpubl.). This composition of oxidative enzymes 
is shown by almost all ectomycorrhizal fungi amongst the Boletales s.l. (Agerer et al., in prep.). 

f. The ectomycorrhizal group with 'boletoid rhizomorphs'(figs. 2, 3) 

Strobilomycetaceae differ from the remaining families by the lack of pulvinic acids (F3) 
(Gill & Steglich 1987, Bresinsky 1996). They synthesize DOPA from tyrosine (Gill & 
Steglich 1987), and do not go further to atromentin (Fl) or pulvinic acids (F3). Several 
pulvinic acids containing species do not produce atromentin any more, although this 
compound is known as a semifinished product of pulivinic acids (Gill & Steglich 1987). At 
least they do not enrich it in the fruitbodies. The same could be possible for Strobilomyces. A 
second interpretation might be that most of the tyrosine, allocated to secondary metabolism, is 
consumed for the synthesis of DOPA. Both explanations consider therefore the lack of 
pulvinic acids for systematic affiliation of minor importance. 

The rhizomorphs are brownish grey and completely lack clamps: a combination of possibly 
apomorphic characters. Strobilomyces floccopus forms cystidia (figs. 22e, f), which originate 
internally, and are not formed as terminal ends of the outermost rhizomorph hyphae. Afro- 
boletus luteolus, has basically the same rhizomorphs. Typical cystidia are lacking, but terminal 
ends of peripheral hyphae are distally slightly inflated and their walls are somewhat thickened 
(fig- 23g). 

In spite of many attempts, it was not possible yet to prove whether Strobilomyces is ecto- 
mycorrhizal. But a tenacious resistance against growth in culture (Bresinsky 1996) is sup- 
posed as a hint for being mycorrhizal. Attempts to culture it were according to Bruns (pers. 
comm.) exceptionally successful. 

This family is characerized by thick-walled spores with reticulations or prominent ribs as 
ornament, and is therefore accepted as a separate family (Besl & Bresinsky 1997). Pegler & 
Young (1981) however, propose a very broad definition mainly based on spore-print colour 
and they include even the genus Suillus, an inclusion, which has not been accepted (e.g. 
Hawksworth et al. 1995, Hoiland 1987). 

Paxillaceae (i.e. excl. Hygrophoropsidaceae and Tapinellaceae, incl. Gyrodontaceae and 
Melanogastraceae, comp. Bresinsky et al. 1999) are, regarding to their rhizomorphs, difficult 
to distinguish from the other families with exception of the mostly consistent presence of 



35 

clamps. Some south hemispheric species are possibly an exception with an apomorphic lack 
of clamps. In addition, sclerotia which are formed frequently on rhizomorphs, and brownish 
colours are evident. 

Sclerotia are documented for Austropaxillus boletinoides (Palfner, in prep.), Boletinellus 
merulioides (Schwein.) Murrill (Cotter & Miller 1985), Gyrodon lividus (Bull.: Fr.) Sacc. 
(Agerer et al. 1993), Paxillus involutus (e.g. Laiho 1970), P. rubicundulus Orton, P. validus 
Ch. Hahn in Hahn & Agerer (Hahn & Agerer 1999a) and Phlebopus sudanicus Har. & Pat. 
(Thoen & Ducousso 1989). This is perhaps an apomorphic character which unites these 
genera, but this character has to be looked for in several more genera and species. In 
Coniophoraceae however, sclerotia are known, too, namely in Hygrophoropsis aurantiaca 
(e.g. Antibus 1989) and Leucogyrophana spp. (Ginns 1976). 

Clamps in below-ground hyphae have been proven in Boletinelleus merulioides (Cotter 8c 
Miller 1985), Gyrodon lividus (Agerer et al. 1993), Gyrodon cyanescens (Agerer 1999b), 
Melanogaster variegatus (fig. 27), Paxillus involutus (Agerer 1988a, Mleczko 1997^, P. ob- 
scurosporus Ch.Hahn, P. rubicundulus, P. validus (Hahn &. Agerer 1999a), Phaeogyro- 
porus beniensis (fig.24) and Phlebopus sudanicus (Thoen & Ducousso 1989). Clamps are 
lacking in rhizomorphs of Austropaxillus statuum (Hahn, unpubl.). 

According to Bresinsky (1996) and Besl et al. (1996), the members of Paxillaceae possess 
cyclopentenones (F17-F20), with the exception of Austropaxillus, a segregate from Paxillus 
(Bresinsky et al. 1999). It does not have involutin (F20), the typical cyclopentenone of 
Paxillus (Bresinsky et al. 1999). Therefore, this chemical character is not a consistent marker 
for Paxillaceae. 28S rDNA-sequences, distribute Paxillaceae with two separate clades amongst 
the Boletales, one clade with Paxillus, Gyrodon and Melanogaster, the other contains 
exclusively Austropaxillus species (Bresinsky et al. 1999). Austropaxillus forms a sister 
cluster to Serpula, and the Paxillus-Gyrodon-Melanogaster clade a sister group to Rhizo- 
pogon, Suillus and others. Only the latter cluster is a sister group to Coniophora- 
Leucogyrophana-Hygrophoropsis. Hence Paxillaceae overlap with respect to DNA-sequences 
different, anatomically defined groups. 

Sclerotia and clamps in brown rhizomorphs occur also in Pisolithus tinctorius (Agerer 
1991b, GoDBOUT«fe Fortin 1985, Weiss 1992). Clamps occur in Pisolithus aurantioscabrosus 
Watl., but sclerotia are unknown and the rhizomorphs are more orange (Watling et al. 1995). 
Rhizomorphs and mycorrhizae of Paxillus and Pisolithus are very similar (comp. Hahn & 
Agerer 1999a, Mleczko 1997, Watling et al. 1995, Weiss 1992) all possessing inflated 
cells on the rhizomorph margin. A relationship between Paxillaceae and Pisolithaceae is there- 
fore more likely than a relation between Pisolithaceae and Sclerodermataceae (s. below). Piso- 
lithaceae do not produce cyclopentenones (F17-F20) (Gill 199, Gill& Watling 1986, Gill 
& Steglich 1987) unlike most Paxillaceae, but these compounds are also lacking in Austropa- 
xillus as mentioned above. Unfortunately at present there are no DNA-comparisons between 
these two families. 

The main difference between Paxillaceae and Pisolithaceae regards the special shape of 
peridiole forming fruitbodies in Pisolithaceae, and lamellate (or tubular) hymenophores in 
Paxillaceae as well as the shape and ornamentation of the spores. Fruitbody as well as spore 
evolution is strongly dependent upon selection resulting from environmental influences. A 
similar situation with very high diversity of fruitbody types and spores could already be 
shown for the Gomphales-Geastrales-Gautieriales complex (see above). 



36 



All Boletaceae and Sclerodermataceae hitherto studied, posses 'boletoid rhizomorphs' (figs. 
37, 38): Austroboletus gracilis (fig. 28), Boletellus ananas (Curt.) Murr. (fig. 29a, b), B. prui- 
natus (fig. 29d-i), B. russellii (Forst.) Gilb. (fig. 29c), Boletus calopus (Agerer, unpubl.), B. 
edulis Bull.: Fr. (Gronbach 1988), B. erythropus (fig. 30), Chalciporus piperatus (fig. 31), 
Chamonixia caespitosa (fig. 32, Raidl 1999), Leccinum holopus (Rost.) Watl., L. scabrum, L 
variicolor Watl. (Müller & Agerer 1990), Phylloporus rhodoxanthus (fig. 33), Porphyrellus 
pseudoscaber (fig. 34), Pulveroboletus cramesinus (fig. 35), Scleroderma areolatum (fig. 36), 
S. bovista (Richter & Bruhn 1989), S. citrinum (Raidl 1997), S. sinnamarense (Ingleby 
1999), Tylopilus felleus (Uhl 1988a), Xerocomus armeniacus (Quel.) Quel. (Palfner & 
Agerer 1995), A: badius (Fr.) Kühn.: Gilb. (Gronbach 1988), X. chrysenteron (Bull.: St. 
Amans) Quel (Brand 1989) and A', subtomentosus (Palfner & Agerer 1995). 

All rhizomorphs are light coloured, mostly white, sometimes yellow. Crystals are not 
formed, but peripheral hyphae can be distinctly warty. Quite fi-equently, inflated, roundish 
cells, nodes and conical side-branches of rhizomorphs occur. Only the rhizmorphs of Ä calo- 
pus are amyloid, and very distinctly so. Scleroderma sinnamarense shows amyloid rhizo- 
morphs and ectomycorrhizal mantles (Ingleby 1999). 

Most species posses mycorrhizal mantles with ring-like arranged hyphal bundles (mantle 
type A, according to Agerer 1987-1998, Agerer & Rambold 1998): Boletellus pruinatus 
(Agerer, unpubl.). Boletus edulis (Gronbach 1988), B. luridus Schaeff: Fr. (Brunner et al. 
1992), Chalciporus piperatus (Agerer, unpubl.), Chamonixia caespitosa (Raidl 1999), 
Leccinum aurantiacum (Bull.: St. Amans) S.F.Gray (Godbout& Fortin 1985), L. holopus, L. 
scabrum, L.variicolor (Müller & Agerer 1990), Scleroderma bovista (Jakucs & Agerer 
1999a), S. citrinum (Waller et al. 1993), Tylopilus felleus (Uhl 1988a), Xerocomus arme- 
niacus (Palfner & Agerer 1995), X. badius (Gronbach 1988), X. chrysenteron (Brand 
1991) and X. subtomentosus (Palfner & Agerer 1995). 

All hitherto studied members of Boletaceae do not form clamps, neither in the rhizomorphs 
nor in the mycorrhizal mantle. The genus Scleroderma however, contains species with 
consistent clamps (5. citrinum, Raidl 1997, Waller et al. 1993), or clamps occurring 
occasionally only on the 'runner hyphae' (5. areolatum, fig. 36). 

With reference to pigments, Bresinsky (1996) grouped Leccinum, Chamonixia and Gyro- 
porus together. They contain cyclopentenones (F17-F20), in contrast to the remaining 
Boletaceae, Boletellus, Boletus, Chalciporus, Phylloporus, Tylopilus and Xerocomus. (But 
Leccinum does not contain pulvinic acids). Scleroderma produces pulvinic acids (F3) 
(Arnold et al. 1996). The consistent formation of clamps by Gyroporus cyanescens (Age- 
rer 1999b) indicates that this genus is rather a member of Paxillaceae than of Boletaceae. 

Porphyrellus pseudoscaber, frequently regarded as a member of Strobilomycetaceae 
(Moser 1987, Pegler & Young 1980, Hawksworth et al. 1995), belongs, when the light 
coloured rhizomorphs are taken into account, to Boletaceae. 

From the great similarities between rhizmorphs and ectomycorrhizae of Boletaceae and 
Sclerodermataceae and the presence of pulvinic acids (F3) in both families, Sclerodermataceae 
are transferred into Boletales. The roundish, warty or reticulated spores, known in Scleroder- 
mataceae (Sims et al. 1995), but not in Boletaceae is of minor importance, as spores with com- 
plicated spore ornamentation occur also in Strobilomycetaceae, a family generally accepted in 
the Boletales. The spore ornamentation can possibly change during gasteromycetation, as is 
obvious also for Geastrales and Gautieriales in comparison to the spores of the related 
Gomphales (s. above.) 



37 



Astraeus hygrometricus (Pers.) Morgan forms 'boletoid rhizomorphs', too (Gaie & Heine- 
MANN 1980, Richter & Bruhn 1989, Schramm 1966). The mycorrhizal structure is in- 
sufficiently known to draw conclusions on the mantle type. The consistent presence of 
clamps in rhizomorphs and the dark brown colour of hyphae (Schramm 1966) suggest rather 
a relation to Pisolithaceae than to Sclerodermataceae. 



Synopsis of the atromentin-containing relationships 

(emended under consideration of Besl & Bresinsky 1997, Bresinsky et al. 1999) 

Order (Tricholomatales Kühner) ? 

Family Omphalotaceae Bresinsky 
Order Thelephorales Comer: Oberw. 

Family Bankeraceae Donk emend. Stalpers 

Family Thelephoraceae Chev. 

Family Scutigeraceae Bond.& Singer: Singer 
Order Boletales Gilbert emend. Agerer 

Suborder Suillineae Besl & Bresinsky 

Family Truncocolumellaceae Agerer 

Family Gomphidiaceae R.Maire: Singer 

Truncocolumellaceae and Gomphidiaceae are exclusively placed in this suborder due to the 
DNA-data (see above). On anatomical reasons, both families should have been separated at least 
at suborder level. Such a rearrangement should be done after it has been shown that DNA- 
sequences of Gomphidiaceae and Truncocolumellaceae have evolved more rapidly in 
comparison to their functionally crucial anatomical features. In the latter case, both families 
would be basal to Suillaceae and Rhizopogonaceae 

Family Suillaceae (Singer) Besl & Bresinsky 

Family Rhizopogonaceae Dodge 
Suborder Tapinellineae Agerer 

Family Tapinellaceae Ch.Hahn 
Suborder Coniophoroineae Agerer & Ch.Hahn 

Family Coniophoraceae Ulbr. (incl. Hygrophoropsidaceae Kühner) 
Suborder Paxillineae Feltgen 

Family Paxillaceae R.Maire (incl. Gyrodontaceae (Singer) Keinem., incl. Melano- 

gastraceae Tul., excl. Chamonixiaceae Jülich) 

Family Pisolithaceae Ulbr. 

(Family Astraeaceae Zeller: Jülich) ? 
Suborder Strobilomycetineae Gilbert 

Family Strobilomycetaceae Gilbert 
Suborder Boletineae Rea emend. Gilbert 

Family Boletaceae Chev. (incl. Xerocomaceae Pegler & Young, incl. Chamo- 
nixiaceae Jülich) 

Family Sclerodermataceae Corda 

(Family Astraeaceae Zeller: Jülich) ? 



38 



New Taxa 

Coniophoroineae Agerer & Ch.Hahn, subordo nova 
Typus subordinis: Coniophoraceae Ulbr. 1928 

Diagnosis latina: Hymenomycetes. Basidiomycota. Sporis dextrinoideis; rhizomorphis 
boletoideis, saprotrophis. 

Tapinellineae Agerer. subordo nova 

Typus subordinis: Tapinellaceae Ch.Hahn in Hahn & Agerer (1999) Studien zur 
Systematik der Paxillaceae. Sendtnera 6: 1 15-133. 

Diagnosis latina: Hymenomycetes, Basidiomycota. Carposomata pileata, stipitata vel late- 
rale adnexa. Sporis brunneis, ellipsoideis, laevibus, dextrinoideis; chlamydosporis praesen- 
tibus; rhizomorphis unifomibus vel phlegmacioideis vel agaricoideis, non boletoideis. Invo- 
lutin deficiens, aceta pulvinica formantes. 

Truncocolumellaceae Agerer ~~^^ 

Typus familiae: Truncocolumella Zeller (1939) New and noteworthy Gasteromycetes, 
Mycologia 31:6. 

Diagnosis latina: Hymenomycetes, Basidiomycota. Carposomata gastroidea; columellis 
truncoideis vel dendroideis, glebis brunneis; sporis brunneis, laevibus, ellipsodeis, dextri- 
noideis, crassitunicatis; chlamydosporis praesentibus; rhizomorphis uniformibus. Involutin 
deficiens, aceta pulvinica formantes. 

LOQUIN (1984) mentioned the family Truncocolumellaceae and refers to his earlier publication LOQUIN 
(1981). Sinces intensive searches for the latter publication were not successftil and Truncocolumellaceae 
are not included in Hawksworth et al. (1995) as a valid family, Truncocolumellaceae are described and 
validly published in the present paper. 

Omphalotus atraetopus (Kalchbr. in Thiimen) Ch.Hahn, comb. nov. 

Basionym: Paxilliis atraetopus Kalchbr. in Thümen, F. (1878) Diagnosen zu Thümen's 
"Mycotheca universalis". Flora 61 : 87. - Lectotypus in M. 

According to Reid (1975) '"Paxillus" atraetopus is synonymous to Omphalotus olearius. 
The pallid spores, the paxilloid fruitbodies and the anatomy of the rhizomorphs (fig. 8h) 
confirm the categorization into Omphalotus. The difference in spore colour - Omphalotus 
olearius possess a white sporeprint, 'Taxillus" atraetopus on the other hand possess a 
yellowish sporeprint - is a reason to deny this conspecifity. The occurrence in the Capensis 
on native wood as a special habitat emphasize the seperation of ''Faxillus" atraetopus as an 
own species. Unfortunatly the original description is quite short and no recent collections are 
available. So the macroscopical characters are difficult to discuss. Possibly there are further 
distinctive features to distinguish these two species. Due to the difference in sporeprint 
colour, we believe that it is justified to suggest Omphalotus atraetopus as an own taxon on 
species level. 



Collection data of the used specimens 

Some 60 rhizomorphic genera of different families of Hymenomycetes have been studied to 
date and they contributed to the results of the present paper, but only the below mentioned 
genera are actually included. 



39 



Afroboletiis luteolus. Burundi, Rumonge, under Brachystegia microphylla in hill miombo, 
12.3.1991, Buyck 4238 (PC). - Agaric us bisporus. Germany, Bayern, Weilheim-Schongau, 
Grasleitener moor between Schöffau and Huglfing, eastern margin of the Schwein-Moos, ca. 
690 m asl, 21.10.1998, Hahn & Beenken LB1047 (M). - Alpova trappei: USA, Oregon, Sinn 
County, Willamette National Forest, Andrew's Experimental Forest, Mill creek 2, ca. 750 m 
asl, stand ca. 45-60 years old, 20.6.1995, Agerer RA] 2 199 (M). - Austroboletus gracilis: 
USA, Michigan, north-west comer of Montmorency County, 30.7.-28.8. 1967, Bresinsky 
67/1015 (M). - Boletellus ananas. USA, Florida, Gainesville, 26.8.1977, Bresinsky 77-36 
(M). - Boletellus pruinatus. Germany, Bayem, Regensburg, Vorderer Bayerischer Wald, 
Forstmühle, in forest district 'Rabenzipfl', 14.10.1997, Agerer RA12350 (M). - Boletellus 
russellii: USA, Florida, Gainesville, 26.8.1977, Bresinsky 77-35 (M). - Boletus erythropus: 
Germany, Bayern, Oberstdorf, on the Fellhom close to middle station of Fellhombahn, ca. 
1400-1500 m asl, 19.8.1998, Agerer RA12385 (M). - Chalciporus piperatus: Germany, 
Bayem, Regensburg, Vorderer Bayerischer Wald, Forstmühle, in forest district 'Rabenzipfl', 
\4.\0.\997, Agerer RAl 2348 (M). - Chamonixia caespitosa: Germany, Bayem, district Bad 
Tölz- Wolfratshausen, in the valley of the river Isar between Vorderriß and Wallgau, 47°33'N 
11°24'E, app. 800 m asl, 19.10.1998, Raid! SR720 (M). - Coniophora arida. Germany, 
Bayern, Regensburg, Forstmühle, in forest district 'Rabenzipfl, 31.7.1998, Hahn SR511 (M). 
- Gautieria sp.: Chile, X. region. Province Osomo, Cordillera de los Andes, National Park 
Puyehue, Refugio Antillanca, 40°42'S 72°18'W, ca. 1100 m asl, in Nothofagus pumilio wood 
with Drimys andina understorey, shortly below tree borderline, thin humus layer on vulcanic 
ash, 27.3.1998, Pal/her GP4901 (M). - Hygrophoropsis aurantiaca: Germany, Bayern, 
Aichach, zwischen Odelzhausen und Mering, im Höglwald near Tegembach, 20.9.1998, 
Agerer RAl 2408 (M). - Jaapia ochroleuca: Germany, Bavaria, AUgäu, Oberjoch, northern 
slope of Iseler, 1800-1900 m asl, on Pinus mugö, 4.9.1984, Oberwinkler F035659 (TUB). - 
Switzerland, Graubünden, Val brevere, on Larix, 1820 m asl, 9.9.1976, Oberwinkler FO24019 
(M). - Leucogyrophana pseudomollusca: Germany, Baden- Württemberg, Tübingen, Spitz- 
berg, 1.10.1977, Oberwinkler FO25068 (TUB). - Melanogaster variegatus: Italy, Emilia 
Romana, Parma, from Borgotaro direction Bardi, close to Borgotaro shortly behind Brunelli, 
close to the crossing direction Lame, Quercus-Castanea wood, Beenken LB1014 (M). - 
Omphalotus atraetopus: South Africa, Boschberg, near of Somerset-East, Febr. 1876, Mac 
Owen & Tuk 1216, type (M). - Omphalotus olearius: Italy, Emilia Romana, Parma, Tagli 
close to Borgotaro, 20.9.1994, Agerer RAl 2 132 (M). - Phaeogyroporus beniensis. Chile, 
Escuadron, Gregario, rara vez solitario, sobre suelo o entre corteza de tocones podridos de 
Pinus radiata, Garrido 196 (M). - Phylloporus rhodoxanthus. USA, Michigan, Mont- 
morency County, northwest comer, 30.7.-28.8.1967, Bresinsky 67/1173 (M). - Porphyrellus 
pseudoscaber: Poland, Gorce Mountains, western slope of Gore mountain, ca. 1200 m asl, in 
acid rocky clay soil, in mixed beech and spruce forest (Fagetum carpaticum), Mleczko 
RAl 2768 (M). - Pseudomerulius aureus. Czech Republic, Prachatice, Husinec, dam of 
Blanice, mountain Hradec, on laying pine branch, 13.9.1996, Hahn CH89/96. - Pulveroboletus 
cramesinus. Germany, Bayem, Hartschimmelhof, Goaslweide, MTB 80333/31, 
Cephalanthero-Fagetum, 17.9.1998, Karasch RAl 2407 (M). - "'Quercirhiza noduloso- 
morpha" (=Tomentella sp ): Portugal, Grandola, district Setubal, 38°5'N 8°33'W, on Quercus 
suber, 31.10.1998, myc. isol. Azul AAM3/98 (M). - Rhizopogon vinicolor: USA, Oregon, 
Sinn County, Willamette National Forest, Andrew's Experimental Forest, Mill creek 2, ca. 
750 m asl, stand ca. 45-60 years old, 20.6.1995, Agerer RAl 2202a (M). - Scleroderma 
areolatum. Italy, Emilia Romana, Parma, from Borgotaro direction Bardi, close to Borgotaro 



40 



shortly behind BruneUi, close to the crossing direction Lame, Quercus-Castanea wood, 
2.10.1998, Hahn RA] 2485 (M). - Serpula lacrymans. Strain collection of the Institut für 
Botanik, Regensburg, Besl 697. - Strobüomyces floccopus: Germany, Bayern, Regensburg, 
Vorderer Bayerischer Wald, Forstmühle, in forest district 'Rabenzipfl', 31.7.1998, Agerer 
RA12377 (M). - Tapineüa atrotomentosa: Germany, Bayern, Stamberg, Gauting, forests 
around Buchendorf, 22.9.1998, Agerer RA J 2420 (M). - Germany, Bayern, Regensburg, 
Forstmühle, Rabenzipfel, Hahn CH164/98. - Truncocolumella citrina. USA, Oregon, 
Jackson Co., Rogue River National Forest, just east of milepost 64 on highway 62 along 
southside of road, 42°54'N 122°18'W, ca. 1400 m asl, under Pinus contorta, Pseudotsuga 
menziesii, 15.10.1997, Castellano (Trappe 21123) (OSC). 

The many collections of fruitbodies with adhering rhizomorphs which have been provided by 
Ludwig Beenken, Christoph Hahn, Götz Palfner, Dr. Stefan Raidl/ München and Piotr Mleczko/ 
Krakow, and the valuable discussions with Christoph Hahn are highly appreciated. I am much obliged 
to Dr. Gero Steinberg's/ München support regarding cytoplasmatic transport. I thank Prof. Dr. Tom 
Bruns/ Berkeley, Prof Dr. Roy Watling/ Edinburgh, Dr. Gerhard Rambold/ München and Dr. Norbert 
Arnold/ München very much for their critical comments on the manuscript. D. Persoh/Miinchen 
constructed and helped to interpret some DNA-trees, this is kindly acknowledged. The loan of mate- 
rial from PC, TUB, and OSC is very much appreciated. The study was supported by the Deutsche 
Forschungsgemeinschaft (grant SFB 607, Project B7). 



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Prof Dr. Reinhard Agerer; Institut für Systematische Botanik der Universität München, 
Menzinger Straße 67, D-80638 München, Deutschland. 



Legend of figures 

Fig. 1: Schematical drawings and explanations of the different rhizomorph types 

(further explanations in the text). 
Fig, 2: Dendrogram showing relations between Omphalotaceae, Thelephoraies, and 
Boletaies (see also above: 'Plesiomorphic versus apomorphic anatomical 
characters'). The 'uniform-compact rhizomorph type' of Omphalotaceae, a 'full set of 
pigments', the saprotrophic behaviour and amyloid hyphae in culture are plesiomorphic 
(2). The sister group of Omphalotaceae is supported by different types of rhizomorphs 
and by different ecological states. A closer connection could only be expected to Banke- 
raceae with the same rhizomorph type, but with ectomycorrhizae and lacking pulvinic 
acids. - Ectomycorrhizae (1) are regarded as apomorphic in comparison to a saprotrophic 
life-style and have been developed at least three times. - The two sister clades, the 



48 



'thelephoric acid group' and the 'pulvinic acid group' (3), are connected by the common 
compound atromentin, the primary product of the synthesis of thelephoric acid and 
pulvinic acids. No pulvinic acids are known in the 'thelephoric acid group', whereas 
thelephoric acid has been detected in several species of the 'pulvinic acid group' (Suillaceae, 
Rhizopogonaceae, Tapinellaceae). - Backward oriented hyphal branches (4) are apo- 
morphic, resulting from their contribution in bidirectional transport and developed twice. - 
Brown thick-walled soil hyphae (5) are considered as apomorphic, providing resistence. - 
"Loss" of amyloidy (6) is apomorphic. It is repeatedly lost in the 'thelephoric acid group' 
(Bankeraceae, Thelephoraceae), but present in some species of Rhizopogon (Rhizo- 
pogonaceae), Boletus (Boletaceae), and Scleroderma (Sclerodermataeae). The occurrence at 
least in the two latter genera is interpreted as atavism. - (7) These three sister clades miss 
an apomorphic/ plesiomorphic differentiation. A broad range of rhizomorph types is 
demonstrated. Whether terpenoids are suited to be used as differentiation against the sister 
clades is uncertain, due to their infrequency in Suillaceae and Rhizopogonaceae (s. above). - 
The genus Rhizopogon is heterogeneous with respect to several features: amyloidy, for- 
mation of rhizomorph cystidia, and presence of thelephoric acid. This genus within Rhizo- 
pogonaceae might therefore be paraphyletic. - Loss of atromentin and pulvinic acids (8) 
appears apomorphic, although DOPA might prevent or mask the synthesis of pulvinic 
acids. The great diversity and elaboration of spore ornamentation and the formation of 
cystidia on the rhizomorphs are also regarded as apomorphic. It is uncertain, whether Stro- 
bilomycetaceae are ectomycorrhizal or saprotrophic. - (9) Scutigeraceae are characterized 
by phlegmacioid rhizomorphs, in Bankeraceae uniform, thelephoroid and phlegmacioid ones 
can occur. (Fl, F2, F3 refer to chemical formula of atromentin, thelephoric acid and pulvinic 
acids, respectively; comp, above). 
Fig. 3: Dendrogram showing relations within Boletales (see also above: 'Plesiomorphic 
versus apomorphic anatomical characters'). Ectomycorrhizae (1) are regarded as 
apomorphic in comparison to a saprotrophic life-style. - Loss of atromentin and pulvinic 
acids (2) appears apomorphic, although DOPA might prevent or mask the synthesis of 
pulvinic acids. The great diversity and elaboration of spore ornamentation and the 
formation of cystidia on the rhizomorphs are also regarded as apomorphic. It is uncertain, 
whether Stobilomycetaceae are ectomycorrhizal or saprotrophic. - The presence of clamps 
(3) is plesiomorphic and characteristic of all genera, with the exception of a few species of 
Austropaxillus. The loss in the latter genus is interpreted as an apomorphy in an otherwise 
plesiomorphic group. - Cystidia on the soil mycelium (4) are apomorphic. Possibly such 
cystidia are also present on mycelia of Melanogaster and Pisolithaceae, but so far not 
studied. It might therefore be a synapomorphy of Pisolithaceae and Paxillaceae. Cyclopen- 
tenones could be used for position (4), but they are less frequently present than mycelium 
cystidia; they are not found in Pisolithaceae and are lacking in Austropaxillus.- Ring-like 
bundles of hyphae of mycorrhizal mantles (5) are apomorphic due to a higher differentia- 
tion in comparison to the mostly rather undifferentiated mantles of Suillaceae and of 
Rhizopogonaceae, although ring-like hyphal bundles can also occur, e.g.. in R. subcae- 
rulescens with amyloid mantles. Possibly, this Rhizopogon group is more closely related to 
amyloid Boletus species (Boletaceae) or Scleroderma species (Sclerodermataceae). - (6) The 
position of Astraeaceae is as yet uncertain. Dark brown, clamp-bearing mycelia put 
however, Astraeaceae closer to the Pisolithaceae/ Paxillaceae group. Further studies are 
necessary. - Four monophyletic groups are evident: Co-niophoraceae, Strobilomycetaceae, 
Paxillaceae/Pisolithaceae, and Sclerodermataceae/Boletaceae. 



49 



Fig. 4: Gautieria spec: a. Emanating hyphae of a rhizomorph; note lens-shaped appositions 
of some thick-walled hyphae, and short, coral-like outgrowths (drawing divided into two 
parts; point of conitinuation in x - x). b. Crystal-bearing hyphae of a rhizomorph, below 
thick-walled ones, c, d. Acanthocystidia and thin-walled, inflated cells with yellowish 
contents of a rhizomorph surface, e. Optical section through a rhizomorph with ampuUate 
inflations. (GP4901, material fixed in FEA = mixture of formadelyde, ethanol and acetic 
acid, comp. Agerer 1987-1998). 

Fig. 5: Agaricus bisporus: a. Young rhizomorph with some inflating hyphae, no 'runner 
hypha' present; the thin, unthickened branches in their variable arrangement on the vessel- 
like hypha indicate the lack of a 'runner hypha'. b. Vessel-like hyphae with dissolved septa 
(drawing divided into two parts, point of continuation in x - x). c. Portion of a vessel-like 
hypha with dissolved septum and possibly proteinaceous crystals enveloped by a 
cytoplamic membrane. (LB 1047; material fixed in FEA). 

Fig. 6: Unknown parasite in rhizomorphs of Strobilomyces floccopus: a. Highly differentiated 
'soaking organ' A clampless, thickwalled, brown hypha inflated subterminally balloon-like. 
Several very thin and bent, 'maccaroni-like' hyphae grow out of it. At the transition zone 
of the inflated part to the thick-walled hypha, an internal, slightly dextrinoid calotte is 
formed, into which a rostrum is developed, b. Brown thick-walled hypha growing out of (or 
into) the rhizomorph of Strobilomyces floccopus; a conical structure is formed at the 
entrance (exit ?) to the rhizomorph.(In RA 12377, material fixed in FEA). 

Fig. 7. "Quercirhiza nodulosomorpha''' (= Tomentella sp.): a, b. Terminal end of rhizomorphs 
with backward oriented hyphae at the side-branch (drawing 'a' divided into two parts, 
point of continuation in x - x). c. d. Hyphae enveloped by thin, irregularly banched hyphae. 
e, f. Conical side-branch of a rhizomorph with a naked hypha.(AAM3/98, material fixed in 
FEA). 

Fig. 8. Omphalotus oleahus (a-g), O. atraetopus (h): Undifferentiated rhizomorphs of the 
'uniform-compact' type. a. Terminal end of a thin rhizomorph; growing hyphae become 
gradually enveloped from behind, b. Uniting rhizomorphs; backward ramifications of 
hyphae could not be found, c. Side-branch of a rhizomorph; some, evenly thick hyphae 
form the young branch, d. A hypha bows back within the rhizomorph. e. Undifferentiated 
rhizomorph with a reversely oriented clamp, f. Hypha with a side-branch, g. Formation of 
a rhizomorphal branch, h. Longitudinal cryo-section of a rhizomorph.( a-g from RA12132, 
h from type; herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 9. Truncocolumella cithna: a. Part of a hyphal system in a rhizomorph with backward 
oriented hyphal ramifications, b. Terminal end of some hyphae which form a thin, rhizo- 
morph. c. Terminal end of a thicker rhizomorph with thick- walled cells and anastomoses of 
terminal cells of neighbouring hyphae. d. Optical section through a rhizomorph with 
partially irregularly arranged hyphae, all hyphae of the same kind regarding diameter and 
wall thickness, only marginal hyphae with somewhat thicker walls, e. More central hyphae 
of a rhizomorph with slightly enlarged septal pores, no distinct dissolution of septa 
discernible. (Trappe 21 123, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 10. Truncocolumella citrina: a. Optical section through a thin rhizomorph, with a slightly 
inflated hypha containing browinsh, cystoplasmic substances, b. Point of ramification of a 
rhizomorph, in 'x' the side-branch emerges, c. Hypha of a thicker rhizomorph with back- 
ward oriented hyphal branching (clustered arrows represent main growth directions of 
hyphae at ramification points), d, e. Thick-walled terminal ends of hyphae of thicker rhizo- 



50 



morphs and of fruitbody-base, with oil drop contents, reminiscent of chlamydospores. 
(Trappe 21 123, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 11. Tapinella atrotomentosa: a. Terminal end of a thin rhizomorph; a first hypha 
becomes gradually enveloped by hyphae from behind; no typical 'runner hypha' is formed. 
b. Vessel-like hyphae with dissolved septa, right one with a backward oriented branch, c-f. 
Different modes of hyphal ramifications, g. Arthroconidia formed by a distal hyphal end. 
h. Intercalar chlamydospores. (Fig.' a' in two parts, point of continuation in x - x; fig. 'b' 
from RA 12420, herbarium material of fruitbody with adhering rhizomorphs; figs, a, c-h 
from CHI 64/98, culture). 

Fig. 12. Pseudomerulius aureus: a. Optical section through a rhizomorph; thickend, thick- 
walled, hyphae are embedded in a gelatinous matrix, b. Thickened hypha with partially dis- 
solved septa, c. Optical section of a rhizomorph; two hyphae have considerably thickened 
walls, causing an asymmetrical shape of the hyphal lumen, d. Hyphae of the fruitbody 
margin adnexed to the substrate; only forward oriented branches could be found, e-g. 
Several tiny hyphae are growing out from stout, irregularly shaped branches, h. Anasto- 
moses between two hyphae, one hypha shows reversely oriented clamps. (All figs, from 
CH98/96, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 13. Jaapia ochroleuca: a. Section through a fruitbody with a basidia, spores and 
cystidium (cystidium in two parts, point of continuation in x - x). b. Marignal area of a 
fruitbody with a long cystidium growing approximately radially (perhaps representing a 
young stage of rhizomorph development) with a thin-walled acompanying hypha (or young 
basidium). c, d. Ramifiacfion points of rhizomorphs (enveloping hyphae of the 'runner 
hyphae' not shown), e. 'Internal substrate hyphae' with intrahyphal hyphae, one with 
outgrowing hyphal tips. (Clustered arrows represent main growth directions of hyphae at 
ramification points; FO24019, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 14: Jaapia ochroleuca: a. Section through a fruitbody with 'superficial substrate- 
hyphae', basidia, spores and cystidia. b. Three 'internal substrate-hyphae' growing within 
a tracheid. c. 'internal substrate hypha' with a dissolved septum, d 'Internal substrate 
hypha' with reversely oriented cIamps.(F035659, herbarium material). 

Fig. 15. Coniophora arida: a. Ramification at a septum of a 'runner hypha'; only simple 
septa present, b. Young rhizomorph at a clamp of the 'runner hypha'; only one clamp 
formed, several enveloping hyphae with terminal cell. c. 'Runner hypha' with twin-clamps; 
beginning of a node formation, d. 'Runner hypha' with twin-clamps; one clamp is growing 
out to enveloping hyphae of a node. c. 'Runner hypha' with three clamps; side-branches 
formed by hyphae with simple septa, even at their base; beginning formation of a node. 
(Clustered arrows represent main growth directions of hyphae at ramification points; 
SR5 1 1 ; culture material). 

Fig. 16. Coniophora arida: a. Old 'runner hypha' of a thicker rhizmorph with lacking septal 
dissolution where clamps had been formed (drawing divided into several parts, points of 
continuation in x - x, y - y, etc.). b. 'Runner hypha', position of clamps; no septal 
dissolution present, e. 'Runner hyphae' with five clamps, d. Thickened 'runner hypha' 
with four clamps and the formation of a side-branch with a basal simple septum (left). 
(Clustered arrows represent main growth directions of hyphae at ramification points; 
SR51 1 in M; herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 17. Coniophora arida: a. One accompanying hypha of the 'runner hypha' with several 
dissolved septa (drawing divided into several parts, points of conitnuation in x - x, y - y , 
etc.). b. thickened 'runner hypha' with an inflation at the whirl-clamps; at least with four 



SI 



clamps and two side-branches with basal simple septa, and reversely oriented branches. 
(SR51 1, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 18: Leucogyrophana pseudomollusca: a. Central hypha of a thin rhizomorph, rhizo- 
morph approximately as thick as indicated at nodes, septa at nodes partially dissolved 
(drawing divided into several parts, points of continuation in v - v, x - x, etc.). b. Vessel-like 
hypha with dissolved septa, developed from an accomanying one of a 'runner hypha'. c. 
Dissolved septum of a vessel-like hypha. d. Point of ramification of a 'runner hypha'. e. 
Conical side-branch of a rhizomorph. (Clustered arrows represent main growth directions 
of hyphae at ramification points; FO25068, herbarium material of fruitbody with adhering 
rhizomorphs). 

Fig. 19: Serpula lacrymans: Very early ontogenetical stages of rhizomorph development, a-d. 
different modes of hyphal branching at a main hypha. (Drawing 'a'divided into several 
portions; point of continuation in w - w, x - x, etc.; clustered arrows represent main growth 
directions of hyphae at ramification points; Besl 697, culture). 

Fig. 20: Hygrophoropsis aurantiaca: a-b. Ramifications at branching points of a main hypha. 
c. Conical side-branch formed on a rhizomorph; a hypha becomes cone-like enveloped by 
hyphae. d. Vessel-like hypha at a branching point of a rhizomorph, optical section. 
(Drawing 'a" and 'b' in several parts; points of continuation in u - u, v - v; etc., clustered 
arrows represent main growth directions of hyphae at ramification points; RA 12408, 
material fixed in FEA). 

Fig. 21. Hygrophoropsis aurantiaca: One central hypha of a thin rhizomorph; rhizomorph 
approximately double as thick as the hypha; cells with possibly proteinaceous crystals. 
(Drawing divided into several portions; points of continuation in,, a - a, b - b, etc.; 
RA 12409, material fixed in FEA). 

Fig. 22. Strobilomyces floccopus: a-c. Ramifications of a 'ruimer hypha'. d. Cryo-section of a 
rhizomorph showing the basal portions of cystidia originating internally, e. Plan view of a 
rhizomorph with two cystidia originating on the rhizomorph surface, one cystidium drawn 
in two portions (point of continuation in x - x). f. Cryo-section of a rhizomorph with a 
cystidium. g. Conical side-branch of a rhizomorph. h. Vessel-like hyphae with dissolved 
septa. (Clustered arrows represent main growth directions of hyphae at ramification points; 
RA 12391, material fixed in FEA). 

Fig. 23.- Afroboletus luteolus: a-d. Ramifications of 'runner hyphae'. e. Thin rhizomorph with 
a thickened, central hypha with still undissolved septa, f. Vessel-like hypha with dissolved 
septa (drawing divided into two parts, point of continuation in x - x.). g. Surface view of a 
ramification point of a rhizomorph; note inflated, slightly thick- walled cells, and terminal 
inflations of cystidia-like hyphae (asterisk show the two rhizomorph branches). (Clustered 
arrows represent main growth directions of hyphae at ramification points; Buyck 4238; 
herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 24. Phaeogyroporus beniensis: a. Thin rhizomorph; drawing divided into several 
portions (points of continuation in p - p, q - q). b. One vessel-like hypha of a thick rhizo- 
morph drawing divided into several portions (points of continuation in x - x and y - y). c. 
Clamp area of a vessel-like hypha with partially dissolved septum, d-e. Ramifications of a 
main hypha. (Clustered arrows represent main growth directions of hyphae at ramification 
points; Garrido 196, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 25.- Rhizopogon vinicolor. a,b. Ramifications of a 'runner hypha'. c. Cystidium; note the 
geniculate appearance caused by unthickened, collapsed hyphal branches, d-f. Connections 
of cystidia with thin-walled hyphae. g. Cystidia close to thin hyphae with blue granules, h. 



52 



Vessel-like hyphae with dissolved septa. (Clustered arrows represent main growth 
directions of hyphae at ramification points; RA 12202a, tuberculate mycorrhizae fixed in 
FEA). 

Fig. 26. Alpova trappei: a-e. Ramifications of 'runner hyphae'. f. Surface view of a thicker 
rhizomorph with thick- walled, short, inflated, cells, g. Portions of vessel-like hyphae with 
dissolved septa, h. Conical side-branch of a rhizomorph terminating in two naked hyphae. 
(Clustered arrows represent main growth directions of hyphae at ramification points; 
RA12199, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 27: Melanogaster variegatus: a, b. Ramifications of thin rhizomorphs. c. Ramification of 
a 'runner hypha' with one dissolved septum, d. Portions of vessel-like hyphae with dis- 
solved septa, e. Surface view of a thicker rhizomorph with thick- walled, inflated cells. 
(Clustered arrows represent main growth directions of hyphae at ramification points; 
LB1014, material fixed in FEA). 

Fig. 28. Austroboletus gracilis: a. Thin rhizomorph, drawing divided into two portions 
(points of continuation in x - x). b, c. Ramifications of a main hypha. (Clustered arrows 
represent main growth directions of hyphae at ramification points; for fig. 'a' clustered 
arrows at the left side indicate hyphal growth directions above the vessel-like hypha, at the 
right side below it; Bres 67/1015, herbarium material of fruitbody with adhering 
rhizomorphs). 

Fig. 29. Boletellus ananas (a,b), B. russellii (c), B. pruinatus (d-i): a. First appearance of a 
backward oriented hypha at the base of a side-branch, b. Portion of a vessel-like hypha 
with dissolved septum, c. First appearance of a backward oriented hypha at the base of a 
side-branch, d-g. Ramifications of 'runner hyphae'. h. Conical side-branch of a rhizo- 
morph. i. Portions of vessel-like hyphae with dissolved septa. (Clustered arrows represent 
main growth directions of hyphae at ramification points; a,b Bres 77-36; c Bres. 77-35; d-i 
RA 123 50; a-c herbarium material of fruitbody with adhering rhizomorphs; d-i material fixed 
in FEA). 

Fig. 30. Boletus erythropus: a-e. Ramifications at a main hypha of a rhizomorph. f. Portions 
of vessel-like hyphae at partially dissolved septa. (Drawing 'a' in two parts; point of 
continuation in x - x; clustered arrows represent main growth directions of hyphae at 
ramification points; RA 123 85, material fixed in FEA). 

Fig. 31: Chalciporus piper atus: a-e. Ramifications of 'runner hyphae'. f. Thin rhizomorph 
with a vessel-like hypha showing a dissolved suptum; an anastomosis of enevloping 
hyphae with the vessel-like one is apparent (dotted circle). (Clustered arrows represent 
main growth directions of hyphae at ramification points; RA12389, material fixed in FEA). 

Fig. 32.' Chamonixia caespitosa: a-d. Ramifications at main hyphae of thin rhizomorphs; in 
'b' and 'd' backward oriented hyphae of two side-branches form an anastomosis, resulting 
in a triangular structure, e. Vessel-like hypha. (Clustered arrows represent main growth 
directions of hyphae at ramification points; SR720, material fixed in FEA). 

Fig. 33. Phylloporus rhodoxanthus: a, b. Ramifications of a main hypha. c. Conical side- 
branch of a rhizomorph. d. Vessel-like hypha of a rhizomorph with dissolved septa 
(drawing divided into two parts, point of continuation in x - x). e. Surface view of a thicker 
rhizomorph. (Clustered arrows represent main growth directions of hyphae at ramification 
points; Bres 67/1 173, herbarium material of fruitbody with adhering rhizomorphs). 

Fig. 34.- Porphyrellus pseudoscaber: a. Terminal end of an emanating hypha. b. Ramification 
of a 'runner hypha'. c. Vessel-like hypha within a node of a rhizomorph; note dissolving 
septum and formation of a side-branch (optical section), d. Vessel-like hyphae with dis- 



53 



solving septa, e. Formation of a conical side-branch of a rhizomorph. f. Surface view of a 
thicker rhizomorph with thick-walled, inflated cells and soil particles. (Clustered arrows 
represent main growth directions of hyphae at ramification points; RA12768, material fixed 
in FEA). 

Fig. 35. Pulveroboletus cramesinus: a. Branching types of a terminal end of a young rhizo- 
morph, depicted in three portions (points of continuation in x - x, and y - y). b. Conical 
side-branch of a rhizomorph. c. Vessel-like hyphae with dissolved septa. (Clustered arrows 
represent main growth directions of hyphae at ramification points; RA 12407, material fixed 
in FEA). 

Fig. 36: Scleroderma areolatum: a-c. Ramification of 'runner hyphae'. d. Young conical side- 
branch of a rhizomorph. e. Portions of vessel-like hyphae with dissolved septa, f. Bead-like 
inflated hyphae of a flat rhizomorph close to fruitbody base. (Clustered arrows represent 
main growth directions of hyphae at ramification points; RA12485, material fixed in FEA). 

Fig. 37: Schematic drawings of rhizomorphs formed without backward oriented ramifications 
(a), and with backwardly growing hyphae (b, c). Omphalotus is an example of 'uniform- 
compact rhizomorphs', Pseudomerulius of 'phlegmacioid' rhizomorphs; all are based ex- 
clusively on forward oriented hyphal branches (a). - Backward oriented hyphae which do 
not originate from the proximal end of a side branch and which do not grow towards the 
main hypha and do not keep in intimate contact with it over considerable distances (b) are 
characteristic of Truncocolumella citrina ('uniform-compact' rhizomophs), Tapinella atro- 
tomentosa and Agaricus bisporus ('agaricoid' rhizomorphs), and of Tapinella panuoides 
('phlegmacioid' rhizomorphs). - Backward oriented hyphal branches which grow towards 
the main hypha after they have originated above the first simple septum or the first clamp 
of a side-branch (c) are typical of all species of Boletales tested ('boletoid' rhizomorphs), 
exclusive of the families Gomphidiaceae, Truncocolumellaceae and Tapinellaceae. In many 
species they fork into two hyphae, one takes the growth direction of the main hypha, the 
other grows towards its proximal end. At the origin of the side-branches, node-like 
inflations are formed. 

Fig. 38: Comparison of mature 'agaricoid' and 'boletoid' rhizomorphs (those hyphae formed 
at the beginning of the ontogeny are depicted with thicker lines): In the 'agaricoid 
rhizomorph type'(a), the first hyphae to become inflated are those which are developed in 
the beginning of the rhizomorph ontogeny and usually only after a thicker hyphal bundle 
developed. A main hypha is enlarged as well as side-branches, but frequently not to their 
whole extent; step by step the diameter of additional hyphae enlarge and are added to the 
first ones. Rhizomorph side-branches can be connected to any hyphae of the main 
rhizomorph; a connection to those hyphae which are formed at the beginning is not 
necessarily realized. There are no preferential areas of side-branch development. - In the 
'boletoid rhizomorph type' (b) the first hypha to become inflated is the 'runner hypha' 
and its diameter is enlarged through its complete length. Inflated hyphae are added, 
preferentially those which grew alongside the 'runner hypha'. Side-branches of the 
rhizomorphs are directly connected to the 'runner hypha' via the firstly developed hyphal 
branch. There, additional thickened hyphae are added. Irregular branching in the ramification 
area of the original 'runner hypha' causes the node-like appearance. 



54 



backward 
growing 



forward 
growing 



r> agaricoid 

many vessel-like hyphae 

mainly central 

P;> phlegmacioid 

thick hyphae solitary, irregularly 
distributed, sometimes vessel-like 

f~^-^ ramariold 

ampullate inflations at septa 



thelephoroid p. p. 

central core of thicker, 
non-vessel-like hyphae 

uniform-compact 
uniform-loose 

^ clamp or 

simple septum 



RHy 



phlegmacioid 




boletoid 

backward oriented 
hyphae, originating 
above the first clamp 
or septum of a side- 
branch, grow toward 
the main hypha and 
keep in intimate 
contact with it; 
mostly forking 

runner-hyphae and 
vessel-like hyphae 



|~^thelephoroid p. p. 

as above 

runner-hyphae and 
vessel-like hyphae 
lacking 



russuloid 

thickened hyphae 
with complete septa 
and ladder-like hyphae 



<C3 



uniform-compact 

hyphae of similar 
diameter, dense 



< $^ y Mt^ uniform-loose 

hyphae of similar 
diameter, loose 




backward 
growing 
and 
Jorking 



LHy = 'Leading hypha' 
RHy = 'Runner hypha' 



Fig. 1 



55 





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Fig. 2 



56 



CoiMpphoraceae 





n 

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2. 2. t 






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sr 

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SB 
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Astcaeaceac 







5 3 



Sclerodermataceae 



Bö|ei»<eae 



Fig. 3 



57 




Fig. 4 



58 




Fig. 5 



59 




20^m 




Fig. 6 



60 




Fig. 7 



61 




Fig. 8 



62 




Fig. 9 



m 




64 




Fig. 11 



65 




Fig. 12 



m 




Fig. 14 



68 




Fig. 15 



69 




Fig. 16 



70 




Fie. 17 



71 



Fig. 18 





Fig. 19 




Fig. 20 



74 




Fig. 21 



75 




Fig. 22 



76 




Fig. 23 



78 




Fig. 25 



19 



Fig. 26 




80 




Fig. 27 




20fjm- 





82 




Fig. 29 



m 




Fig. 30 



84 




Fig. 31 



85 




Fig. 32 




Fig. 33 



8f 



Fig. 34 




88 




Fig. 35 



89 




Fig. 36 



90 



boletoid 




RHy 

A 



LHy Agaricus Q^> agaricoid 

Tapindia j -^^ agaricoid/phlegmacioid 



Gomphidius [T^^ theiephoroid 
Truncocolumdia \ 4^ uniform-corn p, 






uniform 



Omphalotus 




phlegmacioid 



Pseudomerulius 




4 



Jaapia 

Coniophora 

Leucogyrophana 

Serpula 

Hygrophoropsis 

Strobilomyces 
Afroboletus 

Gyrodon 

Phaeogyroporus 

Gyroporus 

Paxfllus 

Austropaxillus 

Mdanogaster 

Pisolithus 

Sufllus 
Boletinus 

Rhizopogon 
Alpova 

Austroboletus 

Chamonixia 

Lectin urn 

Boletdlus 

Boletus 

Tylopilus 

Pulveroboletus 

Chalciporus 

Phylloporus, 




Astraeus 
Scleroderma 



LHy = 'Leading hypha' 
RHy = 'Runner hypha' 

16.7.99 



Fig. 37 



91 



Jaapia 

Coniophora 

Leucogyrophana 

Serpula 

Hygrophoropsis 

Strobilomyces 
Afroboletus 




Gyrodon 

Phaeogyroporus 

Gyroporus 

PaxUlus 

Austropaxillus 

Mdanogaster 



Rhizopogon 
Alpova 

Austroboletus 

Chamonixia 

Leccinum 

Boletdlus 

Boletus 

Tylopilus 

Pulveroboletus 

Chalciporus 

Phyiloporus 

Astraeus 

Scleroderma 



Fig. 38 



93 



Two new species of Bryonectria (Hypocreales, Ascomycetes) 

on bryophytes 



P. DÖBBELER 



Abstract: 

DÖBBELER, P.: Two new species of Bryonectria (Hypocreales, Ascomycetes) on 
bryophytes. - Sendtnera 6: 93-102. 1999. ISSN 0944-0178. 

Two bryophilous species of the genus Bryonectria (Hypocreales, Ascomycetes) are 
described as new and illustrated. B. callicarpa from Europe characterized by orange- 
coloured perithecia infects Frullania dilatata (Hepaticae). B. phyllogena on Poly- 
trichum juniperinum (Musci) from Southern America and Europe has brownish 
perithecia covered by densely interwoven hyphae. In both species inconspicuous 
phialides representing the anamorphic state are recorded. 

Zusammenfassung: 

Zwei bryophile Arten der Gattung Bryonectria (Hypocreales, Ascomycetes) werden 
neu beschrieben und abgebildet. B. callicarpa aus Europa zeichnet sich durch 
orangefarbene Perithecien aus und infiziert Frullania dilatata (Hepaticae). B. 
phyllogena auf Polytrichum juniperinum (Musci) aus Südamerika und Europa hat 
bräunliche Perithecien, die von einem dichten Hyphenmantel bedeckt sind. Bei 
beiden Arten werden unauffällige Phialiden als anamorphe Stadien nachgewiesen. 



In the course of further studies on bryophilous ascomycetes two additional members of the 
genus Bryonectria Döbbeler (Bionectriaceae, Hypocreales) were detected by screening 
potential hosts. They are presented as new species which considerably extend our concept of 
the genus. 

Bryonectria callicarpa Döbbeler, spec. nov. (Fig. 1 , 2) 

Holotype: Spain, Prov. Gerona, Catalonian Pyrenees, surroundings of the main road N 260 
about 20 km west of Figueres in the direction of Olot, on the bark of trees, on Frullania 
dilatata, 20.8.1999, Döbbeler 7094 (Holotype M, isotype BCC). 

Etymology: callicarpus from Greek kallikarpos = with beautiful fruits, refers to the fruit- 
bodies. 



All measurements (except ascomata size) and illustrations were made from slide mounts in 
lactophenol cotton blue. 



94 



Ascomata 120-150 x 85-130 |im, conica vel subpyriformia, aurea ad luteola (in statu vivo), si- 
ne setis. Paries ascomatum lateraliter 12-20 |im crassus, e cellulis crassitunicatis irregularibus 
formatus. Paraphyses apicales praesentes. Hyphae inter ascos deficientes. Asci 52-70 x 8-10 
fim, cylindrici, in apice sine anulo, primo 8-spori, maturitate 4-spori. Ascosporae 12-14 x 4-5 
p,m, ellipsoideae, 2-cellulares, incoloratae sed in quaque cellula guttula luteola praeditae, 
episporio leniter cyanophilo. Hyphae 2,5-5 (im crassae, incoloratae, crassitunicatae, supra 
cellulas hospitis crescentes, appressoriis lateralibus, sessilibus, rotundatis, 5-8 |im diametien- 
tibus omatae. Status anamorphus: Phiales 5-8 x 2-3,5 ^m, ampul liformes, partem concavam 
interioremque texturarum hypharum 20-60 urn diametientium vestitae. Phialosporae per- 
minutae. 

Habitat parasitice in plantas vivas hepaticae Frullania dilatata. Ascomata saepe laxe 
aggregata, omnino in parte ventrali hospitis sita. 

Ascomata perithecial, (105-)120-150(-175) x 85-130(-150) p.m, conical to nearly pyri- 
form, orange red to yellowish, externally paler, dead herbarium material bleaching and finally 
colourless, without setae, surface uneven by shortly protruding groups of cells; no colour 
change in KOH; ascomata not cupulate when dry but laterally collapsing. Ascomatal wall in 
surface view normally without recognizable cells; in section wall laterally 12-20 |im thick, 
composed of irregular, thick-walled hyphal cells with small, reduced lumina; outer cells almost 
colourless, inner ones with yellowish guttules. Apical paraphyses arising from the inner wall 
of the middle and upper part of the ascomata, filiform, up to 60 |im long and 2-3 |im wide, 
only basally ramified, further up as periphyses lining the ostiolar canal; filaments rich in 
orange-yellow guttules. Interascal hyphae absent. Asci 52-70 x 8-10 p.m, cylindrical, straight 
or curved, the apex rounded and without a ring, foot short and thick; mature asci 4-spored. 
Ascospores (1 1-)12-14(-15) x 4-5 |j.m, ellipsoidal, 2-celled, not constricted at the septum, 
colourless, but with a large yellow lipid body in each cell and often a smaller one additionally, 
epispore slightly cyanophilic; 8 ascospore-initials are visible in young asci of which only 4 
reach maturity whereas the other 4 spores remain smaller, sometimes even insert a septum but 
do not develop further and finally degenerate. The distribution of the favoured spores within 
the asci is quite irregular. Hyphae 2.5-5 |im wide, colourless, thick-walled, with appressoria, 
growing superficially over the host cells, prefering the anticlinal cell walls, branched and 
anastomosing; in heavily infected leaves hyphae forming plates which may completely cover 
individual host cells. Appressoria 5-8(-10) |im diam, laterally arising from the hyphae, sessile, 
more or less circular in outline, preferably formed over the junctions of the anticlinal cell walls. 
Anamorphic state: subhemispherical aggregations of hyphal cells leaving a small hollow space 
between themselves and the leaf surface, seen from above roundish or slightly irregular, 20-60 
^im diam, about 20 ^im high, inner side of the dome-cells covered with 5-8 |im long and 2-3.5 
|am wide, bottle-shaped phialides, phialospores wedge- or rod-like, up to 4 x 0.5-1 |xm; 
ostiolar opening absent; the phialides producing structures are difficult to distinguish from 
perithecial primordia. 

Host: Frullania dilatata (L.) Dumort. 

Known distribution: Ireland, Germany, Spain 

Localization and biology: 

The fruit-bodies are restricted to the ventral side of the infected plants, that is the stems, 
underleaves and water sacs as well as the lateral leaves prefering the peripherical regions. They 



95 



are often loosely aggregated, laterally attached and irregularly oriented. The host shows a 
slightly reduced vitality and algae may be present. Nevertheless, a negative effect by the 
parasite seems to be unlikely. 

In the type material several apothecia of Pithya frullaniae Chalaud could be observed 
growing sporadically on dead parts of Frullania dilatata (see Chalaud 1942). Neither Bryo- 
nectria callicarpa nor P. frullaniae have been proven on associated plants of Radula 
complanata (L.) Dumort. 

Remarks: 

Even plants with many coloured ascomata are not recognizable as infected in dorsal view 
because the perithecia exclusively develop on the ventral side. It is indispensable to screen the 
Frullania-mats from the underside or to turn over individual plants or shoots. 

Two remarkable features of this species are the irregular orientation of the fruit-bodies 
pointing in all directions, combined with the hidden and therefore relatively protected position 
within the host-mats. Directed ascospore liberation into the open air apparently does not 
occur. The development of only four ascospores seems to indicate a reduced fertility though 
fourspored asci are known in other Bryonectria-species as well. Whether the loosely attached 
perithecia represent distribution units could not be proven. It is also unclear how the phialo- 
conidia produced within the dome-like cavities of hyphal cells are liberated. 

Bryonectria callicarpa is a distinctive species recognizable, apart form microscopic details, 
by its orange red to yellowish perithecia (at least in the vital state) borne on the ventral side of 
Frullania dilatata. All other species of Bryonectria so far named have uncoloured or hyaline 
perithecia. However, coloured perithecia have been observed several times in undescribed 
species of Bryonectria. Whether the coloration disappears completely in herbarium material is 
not known. The reddish Nectria egens Corner on the epiphyllous hepatic Leptolejeunea vitrea 
(Nees) Schiffn. from Malaya seems to be congeneric, judging from the detailed description and 
illustrations by Corner (1935). 

Further specimens examined: 

Ireland. Co Galway (H 16), Lough Corrib, Archipelago off N shore of Dooras Peninsula 
„Derry Rock North", 53°30'N 9°22'W, 0-3 m, 1.5.1998, Hertel 39525 b (M). 

Germany. Bavaria : Allgäu, Kreis Sonthofen, Hinterstein, 1000 m, 2^.1 .\9A9, Grützmann (M). 



Bryonectria phyllogena Döbbeler, spec. nov. (Fig. 3, 4) 

Holotype: Chile, VIII. Region del BioBio, Prov. de Nuble, Nevados de Chilian, Refugio 
Aserradero, ca. 1300 m, on Polytrichum juniperinum, 5.4.1987, Grau (M). 

Etymology: phyllogena from Greek phyllon = lamella and genos = origin, referring to the fruit- 
bodies arising from hyphae within the lamellar interspaces of the host leaves. 

Ascomata 190-250 x 150-200 ^m, conica vel pyriformia, dilute brunnea ad fiiliginea, sine 
setis sed hyphis adhaerentibus. Ostiolum zona pallida, 40-100 ^m diametienti circumdatum. 
Hyphae adhaerentes 15-50 x 3-4 p,m, luminibus reductis, irreguläres, stratum densum forman- 
tes. Paries ascomatum e cellulis nonnihil irregularibus crassitunicatis compositus, lateraliter 
usque ad 60 ^m crassus, stratum externum hypharum 10-30 \im crassum. Canalis ostioli peri- 
physibus vestitus. Hyphae inter ascos deficientes. Asci 75-95 x 10-13 ^im, subcylindrici, sine 
structura apicali, plerumque 8-spori. Ascosporae 10,5-14 x 6-7,5 |im, ellipsoideae, 2-cellu- 



96 



lares, incoloratae, episporio subcyanophilo. Hyphae 2,5-5 |im crassae, incoloratae, crassituni- 
catae, in spatiis inter lamellas supra cellulas hospitis repentes, appressoriis lateralibus, 
sessilibus, ovoideis vel late ellipticis, 8-12 x 6-8 p.m magnis, praeditae; hyphae saepe densae 
et pelliculas parvas formantes. Status anamorphus: Phiales 9-16 x 3-6 |im, ampuUiformes ad 
subcylindricae. 

Habitat parasitice in latere adaxiali foliorum musci Polytrichum juniperinum. Ascomata 
saepe aggregata et lateraliter coalescentia. 

Ascomata perithecial (175-)190-250(-300) x 1 50-200(-220) ^m, conical or pyriform, light 
to dark brown, rarely nearly colourless, overmature blackish-brown; ostiole surrounded by a 
light region, 40-100 ^m diam, basal parts of ascomata often also paler; without projecting 
setae, but surface uneven and tomentose with adjacent hair-like hyphae; no colour change in 
KOH. Adjacent hyphae about 15-50 x 3-4(-5) um, straight or flexuous, thick-walled with 
reduced lumina, generally light brown, irregularly arranged, forming a dense layer obscuring the 
ascomatal wall cells; apical hyphae elongated in a longitudinal direction, colourless, basally 
merging into the mycelial hyphae. Ascomatal wall in section laterally up to 60 |im thick, outer 
hyphal layer 10-30 ^m thick; wall cells somewhat irregular with branches and anastomoses, 
but mainly tangentially elongated, externally with strongly thickened walls, lumina 2-10(-15) 
|im long. Ostiolar canal lined by plasmatic hyphae up to 35 |im long and 2-3 ^m wide. Inter- 
ascal hyphae absent, but empty collapsed asci may be mistaken for paraphyses. Asci 75-95 
(-105) X (9-) 10-1 3 |im, subcylindrical, apical structures not discernible, with an attenuated 
foot; mature asci with (5, 6, 7) 8 spores, initially always 8-spored. Ascospores (9-) 10.5-1 4 
(-16) X 6-7.5(-8) |im, ovoid to broadly ellipsoidal, 2-celled, septum delicate, not constricted 
at the septum, colourless, episporium presumably finely rough and somewhat cyanophilic. 
Hyphae (2-)2.5-5(-6) |im wide, colourless, thick-walled, with appressoria, growing super- 
ficially and irregularly over the host cells or prefering the anticlinal cell walls, branched and 
anastomosing, often forming strands; extended hyphal plates of appressorium-like cells are 
formed in heavily infected leaves; mycelium restricted to the lamellar region of the leaf, 
especially within the interlamellar spaces. Appressoria 8-12(-13) x 6-8 |im, laterally arising 
from the hyphae, sessile, seen from above roundish or broadly elliptical, often slightly sinuate. 
Anamorphic state: phialidic cells bom laterally on the hyphae, 9-16 x 3-6 um, bottle-shaped 
to subcylindrical, often two or few phialides aggregated; sometimes difficult to distinguish 
from hyphae torn during preparation; phialospores not seen. 

Host: Polytrichum juniperinum Hedw. 

Known distribution: Poland, Chile 

Localization and biology: 

Ascomata of Bryonectria phyllogena are formed on leaves in the middle or lower parts of 
the host sometimes showing signs of beginning decomposition. They occupy exclusively the 
adaxial leaf side and arise from the interlamellar spaces or sit upon the lamellae. Ascomata are 
often aggregated and tend to coalesce laterally. Those situated on the lowermost part of the 
lamina are laterally attached and point upwards to the leaf apices, whereas the other ones are 
irregularly oriented when formed on the free-lying lamellae. When developing below the 
inflexed leaf margins they face towards the slit between them, or they bend the margins 
upwards in order to discharge the ascospores without hindrance into the open air. Heavily 
infected leaves may yield up to 45 ascomata including initials. Even in these cases no adverse 



97 



effect on the host can be observed. The species is habitually difficult to distinguish from the 
southern hemisphere Potridiscus polymorphus Döbbeler «& Triebel (Leotiales) which occurs on 
the same host species (Döbbeler & Triebel, in press). 

Remarks: 

Bryonectria phyllogena is not closely related to any other species of the genus. It has by far 
the largest fruit-bodies which additionally deviate by its brown colour and hyphal covering. 
Two further species are known on Polytrichaceae. B. biseptata Döbbeler on Dawsonia R.Br. 
from New Guinea and on Polytrichum Hedw. from Nepal is distinguished by asci with four 
biseptate spores (Döbbeler 1978, 1981). B. cuneifera Döbbeler on Polytrichastrum formo- 
sum (Hedw.) G.L.Sm. from Europe and B. cuneifera war. jamaicensis Döbbeler on Polytricha- 
delphus flexuosus (Müll. Hal.) Mitt, fi-om Jamaica has hyaline and completely smooth though 
verruculose perithecia up to 130 |im diam (Döbbeler 1978). 

At least a superficial resemblance exists with Ticonectria perianthii Döbbeler, a mono- 
specific hypocrealean genus attacking the perianths of the epiphyllous Radula flaccida 
Lindenb. & Gottsche (Döbbeler 1998). Ticonectria Döbbeler is characterized by a different 
excipulum structure, partly intercellularly growing hyphae, missing appressoria and a different 
anamorphic state. 

Further specimens examined: 

Poland. Pomerapia Occidentalis : distr. Chojnice, in pineto sicco ad lacum „Trzemeszno" 
prope pagum Mecikal, 3.10.1963, Lisowski (M, Bryoth. pel. 1725, sub Polytrichum juniperinum). 

Chile. In locis soli expositis, prope Corral, puerto de Valdivia, Krause (M). - Patagonia occ, 
in valle fl. Aisen, 18.2.1897, Dusen 588 (M). 

The genus Bryonectria comprises seven species which grow on systematically diverse 
bryophytes in various ecological habitats (Döbbeler 1998, Döbbeler & Hertel 1984). 
Although the host range of the genus is characterized by an assemblage of unrelated tropical 
and non-tropical mosses and hepatics, individual species are restricted to single host species or 
a group of related hosts. Accidental records from geographically remote areas of the world and 
other observations indicate that many more members of the genus exist. Presumably, 
Bryonectria will prove to be one of the most species-rich relationships within the Hypocreales 
and ascomycetes on bryophytes. 



I am grateful to Mr. Howard F. Fox (Dublin) and Dr. F. Schuhwerk (Munich) for comments on 
the manuscript. 



Literature 

Chalaud, G. 1942: Pityafrullaniae nov. sp., discomycete parasite de Frullania dilatata Dum. 

- Rev. Bryol. Lichenol., n. s. 13: 1 17-120. 
Corner, E.J.H. 1935: A Nectria parasitic on a liverwort: with further notes on Neotiella 

crozalsiana. - Gard. Bull. Straits Settlem. 8: 135-144. 
Döbbeler, P. 1978: Moosbewohnende Ascomyceten I. Die pyrenocarpen, den Gameto- 

phyten besiedelnden Arten. - Mitt. Botan. Staatssamml. München 14: 1-360. 
- 1981: Moosbewohnende Ascomyceten V. Die auf Dawsonia vorkommenden Arten der 



98 



Botanischen Staatssammlung München. - Mitt. Botan. Staatssamml. München 17: 393- 

473. 

1998: Ascomyceten auf der epiphyllen Radula flaccida (Hepaticae). - Nova Hedwigia 66: 

325-373. 

& Hertel, H. 1984: Drei neue moosbewohnende Ascomyceten aus der Subantarktis 

(Marion Island). - Sydowia 36 (1983): 33^5. 

& Triebel, D.: Potridiscus polymorphus (Leotiales) - a new ascomycete on Polytrichaceae 

(Musci) with palaeoaustral distribution. - Hoppea (in press). 



Dr. Peter Döbbeler, Institut für Systematische Botanik der Ludwigs-Maximilians-Universität 
München, Menzinger Straße 67, D-80638 München, Deutschland. 



99 




Fig. 1: Bryonectria callicarpa (M, holotype). a: Ascomata in outline; b: Asci in outline; c: 
Ascospores; d: Apical paraphyses; e: Immature (left) and mature asci. Scale bars: a: 100 
urn; b: 25 ^mi; c-e: 15 ^m. 



100 




Fig. 2: Bryonectria callicarpa (M, holotype). a: Phialides; b: Section through dome-like 
structure of anamorphic state with phialides on leaf surface; c: Hyphae with appressoria. 
Scale bar: a-c: 20 pm. 



101 




Fig. 3: Bryonectria phyllogena (M, holotype). a: Host leaf colonized by ascomata seen from 
above; b: Ascomata with paler apical region seen laterally or from above in outline; c: 
Ascospores; d: Hyphae covering the ascomatal wall; e: Asci. Scale bars: a: 1 mm; b: 150 \mv, 
c-e: 15 ^im. 



102 




Fig. 4: Bryonectha phyllogena (M, holotype). a: Lateral ascomatal wall in longitudinal section; 
b: Hyphae with phialides; c: Hyphae with appressoria; Scale bar: a-c: 15 |j,m. 



103 



Limonium nydeggeri - eine neue Art aus Südwestportugal 



M. Erben 



Zusammenfassung: 

Erben, M.: Limonium nydeggeri - eine neue Art aus Südwestportugal. - Sendtnera 
6: 103-107. 1999. ISSN 0944-0178. 

Limonium nydeggeri (Plumbaginaceae) wird als neue Art der portugiesischen 
Atlantikküste beschrieben. Sie ist ohne Zweifel in die nähere Verwandtschaft von 
Limonium ovalifolium zu stellen. Bekannt ist sie bisher nur aus der Umgebung des 
Cabo Säo Vicente (Algarve) und von dem ca. 270 km nördlich davon liegendem 
Cabo Carvoeiro (Estremadura). Die Art ist gut charakterisiert durch ihren auffal- 
lenden Habitus mit den dicht rosettig angeordneten, kleinen, kurzstieligen Blättern 
und den niedrigen, fiir die Gattung Limonium sehr dichten Infloreszenzen. 

Abstract: 

Limonium nydeggeri (Plumbaginaceae) is described as new species from the atiantic 
coast of Portugal. It is doubtless closely related to Limonium ovalifolium. This 
species is so far only known from the environs of Cabo Säo Vicente (Algarve) and 
c. 270 km north of this, at Cabo Carvoeiro (Estremadura). The new species is well 
characterized by its habit with densely rosulate, shortly petiolate and small leaves 
and by short, as throughout the genus Limonium, very dense inflorescences. 



Limonium nydeggeri Erben, spec. nov. 

Holotypus: Portugal, Algarve, Cabo de Säo Vicente, Kalkebene, 50 m, 18.5.1995, Nydegger 
33903 (MSB 73871; Isotypen: Herb. Erben, Herb. Nydegger). 

Abb.: 1-2. 

Planta perennis, glabra, oligocaulis. Caudiciili 0,5-5,0 cm longi, plerumque in parte su- 
periore dense ramosi. Folia dense ad densissime rosulata, florendi tempore persistentia, 
caeruleo-viridia, pruinosa, integra, 15^5(-50) mm longa et 8-18 mm lata, late oblanceolata, 
apice obtusa ad late acuta, pilifera, 1-3-nervata, in petiolum breve, 3-5 mm latum cito atte- 
nuata. Caules 3-14 cm longi, tenues, erecti, subrecti, in parte superiore ramosi. Inflorescentia 
subglobula ad semiglobula vel forma obtrullata. Rami fertiles, brevissimi, 5^0(-50) mm longi, 
oblique sursum spectantes, recti, in parte superiore ramosi. Spicae terminales, dense ad 
densissime dispositae, brevissimae, 3-10 mm longae. Spiculae 4,5-5,0 mm longae, 2-4-florae, 
densissime ad 10-15 in 1 cm dispositae. Bractea inferior tenuis, 1,2-1,8 mm longa et 1,2-2,0 
mm lata, triangulari-ovata, apice acuta, margine late membranaceo, parte centrali camosula. 



104 



acuminata, acumine marginem fere contingente. Bractea media tenuissima, 1 ,5-2,0 mm longa et 
1,0-1,3 mm lata, elliptica ad anguste ovata, apice asymmetrice emarginata. Bractea superior 
plerumque latior quam longior, 3,0-3,8 mm longa et 3,2-4,2 mm lata, latissime elliptica ad 
latissime obovata, apice depresse curvata, margine late membranaceo, parte centrali camosula, 
2,1-2,8 mm longa et 2,2-3,0 mm lata, latissime obovata, acuminata, acumine 0,4-0,6 mm 
longo, trianguläre, marginem non contingente. Calyx 3,5-4,2 mm longus, ex bractea superiore 
1,1-1,8 mm exsertus, plerumque glaber; dentes calycis ca. 0,8 x 1,5 mm, late ovati, undulati; 
costae tubi ad basim vel supra basim dentium calycis desinentes. Petala 5,5-6,0 mm longa et 
1,5-1,7 mm lata, cuneata, rubro-violacea. Corolla 4,0^,6 mm diametro. Numerus chromo- 
somaticus: 2n = 16. 

Pflanze ausdauernd, kahl, wenigstengelig. Stämmchen 0,5-5,0 cm lang, meist am oberen 
Ende dicht verzweigt. Blätter dicht bis sehr dicht rosettig angeordnet, zur Blütezeit nicht ver- 
welkt, blaugrün, oft weiß bereift, ganzrandig, 15-45(-50) mm lang und 8-18 mm breit; Spreite 
breit oblanzeolat, am oberen Ende stumpf bis breit spitz, mit einer 1-2 mm langen Stachel- 
spitze, 1-3 -nervig, rasch in einen kurzen, 3-5 mm breiten Stiel übergehend; seitliche Blatthälf- 
ten häufig leicht nach oben geklappt und Blattspitze leicht nach unten gebogen. Stengel 3-14 
cm lang, zart, aufrecht, nahezu gerade, in der oberen Hälfte oder häufig sogar erst im oberen 
Drittel verzweigt. Infloreszenz kugel- bis halbkugelformig oder im Umriß obtrullat. Äste fertil, 
sehr kurz, 5^0(-50) mm lang, schräg nach oben gerichtet, gerade, nur in der äußeren Hälfte 
verzweigt. Ähren dicht bis sehr dicht an den Enden der Äste sitzend, sehr kurz, nur 3-10 mm 
lang. Ährchen 4,5-5,0 mm lang, (l-)2^(-5)-blütig, sehr dicht stehend zu 10-15 pro Zenti- 
meter. Äußere Braktee zart, 1,2-1,8 mm lang und 1,2-2,0 mm breit, dreieckig-eiförmig, am 
oberen Ende spitz; Rand breit weißhäutig; zentraler Bereich etwas fleischig, mit einer fast bis 
zum Rand reichenden Spitze. Mittlere Braktee sehr zart, 1,5-2,0 mm lang und 1,0-1,3 mm 
breit, elliptisch bis schmal eiförmig, am oberen Ende asymmetrisch zweilappig. Innere Braktee 
normalerweise breiter als lang, 3,0-3,8 mm lang und 3,2-4,2 mm breit, sehr breit elliptisch bis 
sehr breit obovat, am oberen Ende flachbogig; Rand breit weißhäutig; zentraler Bereich etwas 
fleischig, 2,1-2,8 mm lang und 2,2-3,0 mm breit, sehr breit obovat, mit einer 0,4-0,6 mm 
langen, dreieckigen, nicht bis zum Rand reichenden Spitze. Kelch 3,5^,2 mm lang, die innere 
Braktee um 1,1-1,8 mm überragend, normalerweise völlig kahl; Kelchzähne ca. 0,8 x 1,5 mm, 
breit eiförmig, mehr oder weniger gefaltet; Rippen der Kelchröhre an oder etwas unterhalb der 
Kelchzahnbasis endend. Kronblätter 5,5-6,0 mm lang und 1,5-1,7 mm breit, keilförmig, 
rotviolett. Blütendurchmesser 4,0-4,6 mm. Chromosomenzahl: 2n = 16. 

Die hier beschriebene Art wurde Herrn Max Nydegger-Hügli (Basel) gewidmet, in Aner- 
kennung seiner großen Verdienste, die er sich mit der Erforschung der Flora des Mittelmeer- 
gebiets erworben hat. 

Die am Cabo Carvoeiro (Estremadura) gesammelten Exemplare besitzen im Gegensatz zu 
den an der Algarve vorkommenden Pflanzen immer spärlich behaarte Kelche und häufig etwas 
schmälere innere Brakteen und dicht gedrängte, kugelförmige Infloreszenzen. 

Limonium nydeggeri wurde bisher von mir als eine an den felsigen Standort angepasste 
Wuchsform der Art L. ovalifolium angesehen (Erben 1993). Langjährige Versuche haben aber 
gezeigt, dass auch die im Gewächshaus kultivierten Exemplare den charakteristischen Habitus 
der Wildpflanzen beibehalten. Somit erhöht sich die Zahl der in Portugal auftretenden, eng 



105 



zusammengehörenden Sippen aus der L. ovalifolium-Gruppe auf drei: L. nydeggeri, L ovali- 
folium (Poiret) O.Kuntze und L. lanceolatum (Hoffmanns. & Link) Franco. Als besonders 
charakteristische Merkmale dieser Gruppe sind ihre rosettig angeordneten, oblanzeolaten 
Blätter, ihre sehr dicht stehenden Ährchen und ihre kleinen Blüten zu nennen. Außerdem 
besitzen alle drei Arten die Chromosomenzahl 2n = 16. 

L. nydeggeri ist von den beiden anderen Sippen vor allem durch seine viel kleineren und im 
Verhältnis zur Länge viel breiteren Blätter, sowie durch die deutlich kürzeren Stengel und Äste 
zu trennen. L. lanceolatum fallt besonders durch seine großen, bis zu 20 cm langen Blätter und 
die bis zu 35 cm hohen, weit ausladenden Blütenstände auf. 

Untersuchte Auf Sammlungen: 

Portugal. Algarve : Distr. Faro, Cabo de Säo Vicente, roguedos, 19.4.1968, Borja, Manzanet & 
Demetrio (MAF, Herb. Erben) - Cabo de Säo Vicente, acantilados calizos, 80 m, 25.5.1979, Castro- 
viejo & Valdes-Bermejo 5240 (MA, Herb. Erben) - Sagres, beim Leuchtturm, 26.6.1990, Bogner 
2095 (Herb. Erben) - Cabo de Säo Vicente, westlich Sagres, Felsküste, 27.9.1990, Buttler (Herb. 
Buttler, Herb. Erben) - Bei Carrapateira an der Westküste, auf Sandstein, 27.6.1991, Mayer 103 
(M) - Cabo de Säo Vicente, 50 m, Kalkebene, 18.5.1995, Nydegger 33903 (MBS, Herb. Erben, 
Herb. Nydegger). - Estremadura : Distr. Leiria, Peniche, Cabo Carvoeiro, ca. 50 m, Felsküste, 12.8. 
1966, Merxmiiller 21670 & Grau (M) - Peniche, Cabo Carvoeiro, carreiro da Furninha, 39°2rN 
9°24'W, 40 m, rochers calcaires erodes, replat de haut de falaise maritime, avec Crithmum mariti- 
mum dominant, 18.5.1996, Lambinon 96/105 & Van Den Sande (M). 



Literatur 

Daveau, J. 1888: Plumbaginees du Portugal. - Bol. Soc. Brot. 6: 145-187. 

Erben, M. 1978: Die Gattung Limonium im südwestmediterranen Raum. - Mitt. Bot. Staats- 

samml. München 14: 361-631. 
- 1993: Limonium. - In: Castroviejo, S. et al. (eds.): Flora Iberica 3: 2-143. Madrid. 
HOFFMANNSEGG, J.C. & LiNK V., H.F. 1813-1820: Statice - In: Flore Portugaise 1 : 416^47. 
Pereira Coutinho, A.X. 1939: Statice. - In: Flora de Portugal 1 : 564-566. Lissabon. 



Dr. Matthias Erben, Institut für Systematische Botanik der Ludwig-Maximilians-Universität 
München, Menzinger Str. 67, D-80638 München, Deutschland. 



106 




Abb. 1 : L. nydeggeri, Habitus; Maßstab 1 cm. 



107 




Abb. 2a: Ährchen; b: Kelch; c: äußere Braktee; d: mittlere Brakteen; e: innere Braktee; f: Blatt 
{L. nydeggeri); g: Blatt {L. ovalifolium). Maßstäbe: a-e: 1 mm; f-g: 1 cm. 



109 



Eine neue Art der Gattung Alonsoa (Scrophulariaceae) aus Chile 



J. Grau 



Zusammenfassung: 

Grau, J.: Eine neue Art der Gattung Alonsoa (Scrophulariaceae) aus Chile. - Sendt- 
nera 6: 109-114. 1999. ISSN 0944-0178. 

Eine neue Art der Gattung Alonsoa, A. honoraria, aus dem Kleinen Norden Chiles 
wird beschrieben. Die bisher übersehene Art zeichnet sich durch eine intensiv kleb- 
rig-drüsige Behaarung und teilweise leicht gezähnte Kelchblätter aus. Die Art unter- 
scheidet sich deutlich von der bisher einzigen bekannten Art der Gattung aus Chile. 

Resumen: 

Se describe una especie nueva del genero Alonsoa, A. honoraria, del norte chico de 
Chile. La especie hasta ahora no advertida, se caracteriza por un indumento fuerte- 
mente glutinoso-glanduloso y tepalos en parte ligeramente dentadas. La especie 
distingue claramente de la unica especie del genero conocido hasta el momento en 
Chile. 



Die Gattung Alonsoa umfaßt etwa 20 Arten, die von Mexiko an der Ostseite Südamerikas 
entlang bis nach Chile verbreitet sind. Die meist roten Blüten sind um 1 80° gedreht, so daß im 
Gegensatz zu den meisten übrigen Scrophulariaceae die ursprüngliche Unterlippe das größte 
fahnenartige Kronblatt in der oberen Mitte der Blüte bildet. Die Kronblätter sind nur über 
einen kurzen Bereich miteinander verwachsen, die beiden untersten im schwächer entwickelten 
Teil der Blüte sogar getrennt. Die Dimension der Blüte und relative Größenunterschiede der 
einzelnen Kronblätter sowie der Staubblattpaare, die Stellung der Blätter, ob gegenständig oder 
in Dreierwirteln und schließlich ihre Gestalt, die von linealisch bis zu elliptisch reichen kann, 
bilden die hauptsächlichen Merkmale zur Charakterisierung der Arten. Alle bisher bekannten 
Arten sind im vegetativen Bereich kahl oder nur schwach drüsig und höchstens im Bereich der 
Blüten dichter und kurz drüsig behaart. 

Die Gattung, von der wenige Arten (De Wolf 1956) bisweilen als Sommerblumen in Eu- 
ropa gezogen werden, besitzt ihren größten Formenreichtum in Peru, wo Lopez ( 1 970) 1 7 Ar- 
ten unterscheidet. Die große vegetative Plastizität der schnellwüchsigen Arten schließt aller- 
dings nicht aus, daß manche der in Peru unterschiedenen Sippen zumindest nicht als eigene 
Arten getrennt bleiben können; die Gesamtartenzahl der Gattung könnte sich daher nach einer 
abschließenden Revision durchaus verringern. 



110 



Aus Chile war bisher nur eine Art bekannt, die ursprünglich als A. incisifolia Ruiz & Pav. 
geführt wurde (Munoz 1966), neuerdings als^. meridionalis (L.f.) Kuntze (Marticorena & 
QuEZADA 1985) bezeichnet wird. Diese letzte Art ist relativ weit verbreitet und ihr Areal 
würde, falls sich die Identität bestätigen läßt, von Chile bis nach Ecuador reichen. Allerdings 
weisen chilenische Pflanzen eher eine gröbere Zähnung auf, wie sie für A. incisifolia charak- 
teristisch ist. Die Frage nach der korrekten Benennung der typischen chilenischen Art wird 
also abschließend noch zu untersuchen sein. Wie viele Arten der Gattung kann A. meridionalis 
(dieser Name soll vorläufig bis zur endgültigen Klärung zunächst beibehalten werden) in Chile 
sowohl als einjährige Art oder als kurzlebige, basal verholzte Pflanze vorkommen. Die Art 
gehört zu den typischen Frühjahrsblühem von Coquimbo bis etwa Concepciön. 

Im nördlichen Teil ihres Areals in Chile hat sich die Gattung jedoch weiter differenziert. 
Nach den ausgedehnten Regenfällen des Jahres 1 997 konnte in der Provinz Choapa, südlich der 
kleinen Ortschaft Caimanes, eine Population gefianden werden, die augenscheinlich mit einer 
Reihe von Eigenschaften nicht in die Variationsbreite der typischen chilenischen Art fallt. Auf 
den ersten Blick unterschieden sich die Pflanzen durch ihre auffallend dichte und klebrige Be- 
haarung und ihren schlankeren Wuchs von der typischen chilenischen Art. Ein weiterer Un- 
terschied ergab sich bei der Kultur und einer auf diese Weise genaueren Untersuchung der Art. 

Die relativ hochgewachsenen Pflanzen standen in einem eingezäunten, noch unbeweideten, 
leicht abfallenden und mit einigen felsigen Partien besetzten, offenen Gelände ca. 5 km südlich 
des Ortsausgangs von Caimanes am sogenannten Camino de Los Tuneies. Die lockere Vegeta- 
tion mit einzeln Büschen von Acacia caven (Mol.) Mol. bestand unter anderem noch aus 
Sphacele salviae (Lindl.) Briq., Alstroemeria pulchra Sims, Phycella ignea Lindl., Leucocoryne 
coquimbensis F.Phil, und als besondere Seltenheit aus Schizanthus parvulus Sudzuki, der, so- 
weit bekannt, bislang nur in der Typusaufsammlung vorliegt (die übrigens aus der gleichen 
Gegend stammt). Diese zarte Art kletterte in der Begleitvegetation und überdeckte das Busch- 
werk mit einem zarten Netz. Sphacele und die neue Alonsoa wuchsen vorzugsweise am Fuß 
und zwischen den Felsen. 

Diese auffallende Art won Alonsoa, deren Eigenständigkeit weiter unten begründet wird, ist 
somit die zweite Art der Gattung für Chile. Ihren Namen erhält sie zu Ehren von Frau Honore 
Wamsler (Pöcking), die in vielfacher Weise den Botanischen Garten München-Nymphenburg 
unterstützt hat. 



Alonsoa honoraria Grau, spec. nov. 

Holotypus: Chile, IV Region, Prov. de Choapa, Camino de Los Tuneies, 5 km südl. Caima- 
nes, lockeres Buschwerk und Weiden, 2.11.1997, Ehrhart & Grau 97/1416 (M, Iso: 
CONC, SGO). 

Planta armua vel perennis ad 1 50 cm alta, herbacea vel in parte basali suffruticosa, ramosa, 
tota distincte stipitate glandulosa. Caules quadrangulares, superne indistincte octangulares vel 
teretes. Inflorescentia racemosa, bracteosa, pluriflora. Folia opposita, herbacea, lamina ovata 
pinnato-nervosa, duplicatim crenato-dentata, basalia ad 7,5 cm longa et 3 cm lata, ad 4 cm pe- 
tiolata, intermedia minora, breviter petiolata, summa parva, subsessilia. Bracteae florum par- 
vae, sessiliae, ± unilateraliter dentibus singulis omatae. Flores ad 10 mm pedicellati, resupinati. 



Ill 



Calyx ± inaequalis, laciniae superiores duo ovatae, unilateraliter ± distincte dentatae, laciniae 
inferiores tres anguste ovatae integerrimae. Corolla laete rubra in parte centrali immerso ob- 
scura, ad 1 7 mm longa et 1 5 mm lata, labio superiore e labiis lateralibus laxe reflexis. Labia 5 
corollae in parte centrali solum corollae connata, labia inferiora inter se non connata. Stamina 4 
ad 5 mm longa, filamentis glabris antheris flavis cordatis conniveis. Gynaeceum glabrum stylo 
ad 6 mm longo arcuato, capitato. Capsula ad 12 mm longa et 5 mm lata. Semina nigra 1 mm x 
0,7 mm. Numerus chromosomatum 2n = 56. 

Weitere untersuchte Aufsammlungen: 

Chile. IV Region. Prov. de Limari : Las Tunas; cerca de las rocas en la sombra, 380 m, 30°59'S 
71°20'W, 10.9.1950, Jiles 1798 (CONC, M) - Rio Ponio, entre los arbustos, 450 m, 30°40'S 
70°54'W, 8.10.1950, Jiles 1898 (CONC, M). 

Eine bis 150 cm hohe, ein- bis wenigjährige, basal verholzte (Sproß bei älteren Pflanzen bis 
2 cm im Durchmesser), reich verzweigte, dicht klebrig drüsige Pflanze. Alle grünen Pflanzen- 
teile sind dicht mit langgestielten, kopfigen Drüsen bedeckt. Auch die Blüten tragen auf der 
Unterseite der Kronblätter besonders im basalen Bereich die gleichen Drüsen. Nur die Staub- 
blätter und der Fruchtknoten sind drüsenfrei. Der deutlich geförderte Hauptsproß endet in 
einer traubigen Infloreszenz; die zahlreichen, ebenfalls Blütenstände bildenden Seitenzweige 
sind dem Hauptsproß untergeordnet. Die Infloreszenz ist reichblütig; meist sind je Blüten- 
stand jedoch nur zwei bis drei Blüten aufgeblüht, die rasch abfallen (bei der chilenischen Form 
von A. meridionalis sind die Blüten beständiger, es können daher bis zu sieben offene Blüten in 
einer Infloreszenz gefunden werden). Der Stengel ist vierkantig mit schmalen, schwach hervor- 
tretenden Rippen; im Bereich der Infloreszenz wird er undeutlich achtkantig bis schwach 
rundlich. Die Blätter stehen kreuzgegenständig und sind grob gekerbt-gezähnt, weich; erstaun- 
licherweise sind auch auf der Blattoberseite Spaltöffnungen angelegt. Die normalen Stengelblät- 
ter sind deutlich gestielt, der Stiel wird bis zu 4 cm lang, die Blattspreite ist spitzeiförmig und 
wird bis zu 7,5 cm lang und bis zu 3,5 cm breit (Abb. 2d-f). Im oberen Bereich des Sprosses 
werden die Laubblätter kleiner, ihr Stiel verschwindet und sie gehen allmählich in die schmäle- 
ren kürzeren und zunehmend asymmetrisch gezähnten Tragblätter der Einzelblüten über (Abb. 
2b-c). Die Blüten sind nach dem Grundprinzip der Alonsoa-^\ü\.Q gebaut (Abb. la-b); das 
breite, zentrale, obere Petalum der resupinierten Krone mißt 11x7 mm, die beiden seitliche 
Kronblätter werden bis zu 7 x 5 mm groß, die beiden unteren, nicht miteinander vereinigten 
Kronzipfel werden bis zu 5 x 3 mm groß. Insgesamt ist die Blüte leuchtend rot gefärbt. Der 
verwachsene Teil der Blüte bildet einen flachen, unten einseitig getrennten Becher, der dunkler 
gefärbt und mit einem blaßgrünen Rand am Kelch festgewachsen ist. Die beiden unteren klei- 
nen, nicht miteinander verwachsenen Kronzipfel hängen jedoch mit ihren schmal eingerollten 
Rändern so fest ineinander, daß sich die Krone ringförmig um den Fruchtknoten schließt. Im 
verwachsenen Teil sind die vier Staubblätter paarweise im oberen bzw. unteren Teil des Be- 
chers eingefügt. Die im Alonsoa typische leichte Asymmetrie des Kelches wird bei A. honora- 
ria besonders deutlich. Die beiden oben liegenden, das größte Kronblatt seitlich flankierenden 
Kelchblätter sind deutlich vergrößert und bei gut ausgebildeten Blüten des unteren Inflores- 
zenzbereichs an einer Seite deutlich gezähnt (Abb. Ic). Die drei übrigen Kelchblätter sind ganz- 
randig, schmäler und aimähemd gleich groß. Die dichte drüsige Behaarung der Außenseite des 
Kelches setzt sich auch nach innen fort. Der kahle Fruchtknoten ist schmal kegelförmig, bis 3 
mm lang; der aufwärts gebogene, bis 6 mm lange Griffel trägt eine kopfige Narbe. Die septizide 
Kapsel wird bis 12 mm lang, die Samen sind klein, schwarz und werden bis 1 x 0,7 mm groß. 
Die Chromosomenzahl beträgt 2n - 56. 



112 



Die aufalligste Eigenschaft der neuen Art ist ihre intensive, drüsige Behaarung, die in dieser 
Form bei keiner anderen bekannten Art auftritt. Die Blätter sind im Gegensatz zur weiteren 
chilenischen Art weich und erscheinen auch durch die amphistomatische Struktur nur wenig an 
den vorwiegend trockenen Standort angepaßt. Teilweise gezähnte Kelchblätter sind bisher 
(LOPEZ 1970) nur von Alonsoa auriculata Diels aus Peru bekaimt. Diese Art unterscheidet 
sich allerdings unter anderem deutlich durch ihre in Dreierwirteln stehenden Blätter. Insgesamt 
wächst Alonsoa honoraria schlanker als A. meridionalis in Chile. Sicherlich interessant wird 
bei Alonsoa die Klärung der Bestäubungsverhältnisse sein. Die Annahme von Lopez, daß 
Windbestäubung vorliege, kann angesichts der auffalligen Blüten nicht gefolgt werden. Aller- 
dings setzten in Kultur im Gewächshaus alle Kapseln an. Fakultative Autogamie erscheint 
daher wahrscheinlicher; welche zusätzlichen tierischen Bestäuber aufgrund der roten Blüten- 
farbe infrage kommen, kann im Augenblick nicht entschieden werden. 



Literatur 

De Wolf, G.P. 1956: Notes on cultivated Scrophulariaceae. - Baileya 4: 34-39. 
LOPEZ Guillen, J. 1970: El genero Alonsoa en el Peru. - Raymondiana 3: 155-246. 
Marticorena, C. & QuEZADA, M. 1985: Catalogo de la flora vascular de Chile. - Gayana, 

Bot. 42: 1-157. 
MuNOZ PiZARRO, C. 1966: Flores silvestres de Chile. Santiago de Chile. 



Prof Dr. Jürke Grau, Institut für Systematische Botanik der Ludwig-Maximilians-Universität 
München, Menzinger Str. 67, D-80638 München, Deutschland. 



113 




Abb. 1 a: Blütenkrone in der Aufsicht; b: Blüte von der Seite; c: Kelch von unten, Orientierung 
in der resupinierten Blüte, (alle Abbildungen nach kultiviertem Material der Typusauf- 
sammlung). Maßstab 5 mm. 



114 



100 \im 




;.je-5f»K„'r •-c/,x^vuv.„<t. \.^^^y)-,-f;4}>- v^;'^-? 




Abb. 2 a: Drüsenhaare der Blattoberseite; b und c: Tragblätter der Infloreszenz; d, e, f: Laub- 
blätter in absteigender Reihenfolge, Maßstab 2 cm. (alle Abbildungen nach kultiviertem Ma- 
terial der Typusaufsammlung). 



115 



Studien zur Systematik der Paxillaceae (Boletales, Basidiomycota) 



C. Hahn & R. Agerer 



Abstract: 

Hahn, C. & Agerer, R.: Systematical studies on Paxillaceae (Boletales, Basidio- 
mycota). - Sendtnera 6: 115-133. 1999. ISSN 0944-0178. 

The Separation of the genus Tapinella by GILBERT ( 1 93 1 ) and its acceptance by 
SUTARA (1992) is discussed and confirmed due to the consideration of hitherto less 
known features in Paxillus s.l. by structure of rhizomorphs, the anatomy of gill tra- 
ma, and features of mycelia in pure culture. The results indicate that Tapinella is a 
sistergroup to the remaining Boletales. The new family Tapinellaceae is proposed 
for this genus. 

Zusammenfassung: 

Die durch GILBERT ( 1 93 1 ) erfolgte und von SUTARA (1992) aufgegriffene Abtren- 
nung der Gattung Tapinella von Paxillus wird unter Einbeziehung von bislang wenig 
beachteten Merkmalen der Rhizomorphen, des Lamellenaufbaus und des Kulturmy- 
cels diskutiert und bestätigt. Die Ergebnisse deuten für Tapinella auf eine Stellung als 
Schwestergruppe zu den übrigen Boletales hin. Für dieses Genus wird die Familie 
Tapinellaceae beschrieben. 

Einführung 

In einer vorangegangenen Studie (Hahn & Agerer 1 999) wurde die Taxonomie und Ökologie 
des Paxillus-involutus-Yovn\QvkxQ\sts anhand europäischer und nordamerikanischer Aufsamm- 
lungen geklärt. In der nun vorliegenden Arbeit wird das Gattungskonzept von Paxillus Fr. und 
Tapinella Gilbert näher beleuchtet. 

Die Frage, ob die von Gilbert (1931) vollzogene und von Sutara (1992) detailliert be- 
gründete Abspaltung der Gattung Tapinella von Paxillus sinnvoll ist, wurde in der Vergangen- 
heit zwar kritisch diskutiert (z.B. Hahn 1996), doch rechtfertigen molekularbiologische Unter- 
suchungen (Fischer 1995, Bresinsky et al. 1999) dieses Vorgehen. 

In der vorliegenden Studie wurde der Versuch unternommen, neue Merkmalsbereiche 
(Rhizomorphen, Aufbau der Lamellentrama, Kulturmycel) zu untersuchen, um eine eventuelle 
Eigenständigkeit von Tapinella zu erhärten. 

Material und Methoden 

Morphologisch-anatomische Untersuchungen mit Hilfe des Lichtmikroskopes: Die mi- 
kroskopischen Untersuchungen wurden sowohl an Frischmaterial (wenn vorhanden) als auch 



116 



an Herbarbelegen vorgenommen. Analysen des Lamellenaufbaus (Querschnitte) wurden aus- 
schließlich anhand von getrocknetem Material durchgeführt, Untersuchungen der Sklerotien an- 
hand von Frischmaterial oder von frisch in FEA (Formalin : Ethanol 70% : Eisessig = 5 : 90 : 5) 
fixiertem Material. Rhizomorphen wurden so weit möglich anhand von Frischmaterial und 
frisch in FEA fixierten Proben studiert. Als Mikroskop stand ein Zeiss Standard 14 mit 
Nomarski-Interferenzkontrast zur Verfügung. 

Anfertigung der Zeichnungen: Die anatomischen Zeichnungen wurden mit Hilfe eines 
Zeichenspiegels (Firma Zeiss) vorgenommen. Die Endvergrößerung auf dem Papier betrug 
hierbei immer 2000x. Sporenzeichnungen wurden im Hellfeld erstellt, während Zeichnungen 
von Hyphenverbänden im Interferenzkontrast nach Nomarski durchgeführt wurden. 

Die Maßstabsbalken der Abb. 1^ und 6-8 entsprechen 20 |im, diejenigen der Abb. 5 und 9 
50 i^m. 

Isolation und Kultivierung von Pilzmycelien: Als Kulturmedien wurden Hefe-Malz- 
VoUagar, Hefe-Malz-Halbagar unter Zugabe von Tetracyclin sowie Wasser-Agar verwendet. 
An einer Sterilbank wurden von möglichst jungen und frischen Fruchtkörpem kleine Stücke 
aus der Huttrama direkt oberhalb der Lamellen entnommen und auf vorbereitete Petrischalen 
mit Agar übertragen. 

Material: 

Tapinella atrotomentosa (Batsch: Fr.) Sutara 

Deutschland. Bayern : Deggendorf, Grub, an Picea-abies-S,i\\m^i, 1 5. 10. 1998, Hahn CH 420/98 
(M) - Fürstenfeldbruck, Schöngeising, Holzhausen, 7.9.1998, //aA« CH 233/98 (M) - Königsdorf, 
zw. Eglsee und Babenstuben, 23.8.1998, Raidl CH 214/98 (M) - zw. Odelshausen und Mering, 
Höglwald bei Tegembach, \\.9.\992>,AgererCH 245/98 (M) - Regensburg, Forstmühle, Rabenzip- 
fel, 31.7.1 998, Hahn CH 164/98 (M) - Siegenburg, Dümbucher Forst, 21.7.1 998, Hahn CH 125/98 
(M) - Simbach, am Boden unter Picea abies, Pinus silvestris, Fagus sylvatica, 29.8.1998, Hahn 
CH 216/98 {U) - Starnberg, Tutzing, Pfaffenberg, 18.8.1996, Hahn, CH 30/96 (M); 17.8.1998, 
CH 199/98 (M); CH 201/98 (M). 

Tapinella panuoides (Fr.: Fr.) Gilbert vai. panuoides 

Deutschland. Bayern : Niederbayem, Passau, Hauzenberg, Kropfmühl, Graphitbergwerk, an Gru- 
benholz unter Tage (mit Übergängen zu fm. archeruntius), 29.5.1999, Hahn CH 36/99 (M). 

Italien: Südtirol: Mals, Matsch, beim Glieshof, ca. 1830 m, 6.8.1998, Agerer CH 183/98 (M). 

Tapinella panuoides var. ionipus (Quel.) Ch.Hahn 

Deutschland. Bayern : Deisenhofen bei Unterhaching, ca. 1 km westl. von Deisenhofen, in der 
Nähe der Keltenschanzen, auf Sägemehlhaufen, 6.9.1998, Garnweidner CH 242/98 (M). 



Ergebnisse 

Tapinella atrotomentosa. > ^ 

Rhizomorphen (Abb. 1, 2, 3a) bis ca. 1 mm dick, hoch differenziert („agaricoid" nach Agerer 
1999); gefäßartige Hyphen vorhanden und besonders auf den zentralen Bereich der Rhizo- 
morphen konzentriert, aber Verlauf unregelmäßig; Knotenbildungen (Agerer 1 999) fehlend, 
rückwärts gerichtete Verzweigungen vorhanden; Hyphen der Rhizomorphenperipherie dünn- 
wandig, 2,5^ um dick, parallel zur Rhizomorphenrichtung, von Hyphen tieferer Lagen nicht 
abweichend, Schnallen fast durchwegs vorhanden, einfache Septen selten; differenzierte Ober- 



117 



flächenzellen fehlend; Hyphen tieferer Lagen 2,5-6 |am dick, dünnwandig, zumeist parallel 
verlaufend; Schnallen durchwegs vorhanden, einfache Septen nicht beobachtet, z.T. aus den 
Schnallenbögen dünnere Hyphen auswachsend, selten Hyphen mit kurzen, seitlichen Aus- 
wüchsen; gefaßartige Hyphen 6-15 um dick, Zellwände verdickt (bis 3 [im), Schnallenbögen 
reduziert und vielfach nicht mehr zu beobachten oder nur angedeutet, Septen meist vorhanden, 
nur vereinzelt Septen aufgelöst. Sehr junge Rhizomorphen (Abb. 3a) ohne gefäßartige Hyphen, 
Zellwände bisweilen leicht verdickt, zumeist Schnallen vorhanden, vereinzelt aber auch 
einfache Septen beobachtet; Vorläuferhyphen fehlend. 

Freies Substratmycel (Abb. 3b): Hyphen 3-5 jam dick, Zellwände leicht verdickt, vor allem 
an Verzweigungsstellen Wände etwas deutlicher; Verzweigungen und Anastomosen häufig, 
einfache Septen wurden nicht beobachtet. 

Kulturmycel (Abb. 4): Hyphen 3-5 |im dick, meist dünnwandig, aber auch dickwandige 
Hyphen auftretend (und den Hyphen des Substratmycels gleichend), sowohl Schnallen als 
auch einfache Septen vorhanden; häufige Bildung von Chlamydosporen, deren Bildungszellen 
meist zumindest einseitig einfache Septen aufweisen, Chlamydosporen (Abb. 4) dickwandig 
(bis 2,5 |im Wandstärke), 8-38 x 4—10 um, sehr vielgestaltig, von langgestreckt zylindrisch bis 
hin zu zitronenfbrmig oder kugelig; Inhalt reich an kleinen Öltröpfchen, farblos oder blaß 
gelblich bis sehr hell bräunlich pigmentiert; Wände leicht bis sehr stark verdickt, farblos bis 
blaß gelblich; einsetzende Chlamydosporenbildung durch Konzentration des Cytoplasmas er- 
kennbar, im Verlauf der Bildung nicht alle Öltröpfchen der Erzeugerzelle in die Spore verlagert, 
freigesetzte Chlamydosporen zumeist blaß gefärbt. 

Lamellentrama (Abb. 5a, b): Hyphenverlauf jung unidirektional und leicht divergierend, 
ausgewachsene Lamellen mit bidirektionaler Hyphenanordnung; Subhymenium deutlich ausge- 
prägt, aus 3-5 |im dicken, dünnwandigen, sich verzweigenden Hyphen bestehend; Hyphen der 
lateralen Lamellentrama auch alt divergierend, dünnwandig, selten etwas dickwandig bzw. 
Wände z.T. in KOH-Lösung (z.B. 5%) deutlich aufquellend; Hyphen der zentralen Lamellen- 
trama unregelmäßig verflochten, jedoch mit zwei Vorzugsrichtungen, zum einen parallel 
zueinander und senkrecht zur Huttrama in Richtung Schneide verlaufend, zum anderen parallel 
zur Schneide und somit radial bezüglich Hut (bidirektionaler Hyphenverlauf); im Lamellen- 
längsschnitt (Schnitt parallel zu Lamellenflächen durch die Schneide) bidirektionaler Hyphen- 
verlauf besonders deutlich (Abb. 5b): Hauptverlaufsrichtung schneidenwärts gerichtet, hiervon 
durch Krümmung oder Verzweigungen einzelne Hyphen in ca. 90°-Winkel die Laufrichtung 
ändemd, sich mit anderen Hyphen streckenweise bündelnd, um wieder vereinzelt durch 
Rückkrümmung in alten, schneidenwärts gerichteten Verlauf einzumünden; Hyphen der zen- 
tralen Lamellentrama mit leicht verdickten, in KOH-Lösung deutlich quellenden Zellwänden, 
dies besonders im Hyphenquerschnitt zu erkermen 

Für weitere Fruchtkörpermerkmale sei auf Sutara (1992) verwiesen. 

Tapinella panuoides 

Rhizomorphen (Abb. 6, 7) bis ca. 1 mm dick, mäßig differenziert („phlegmacioid" nach 
Agerer 1999); gefäßartige Hyphen vorhanden und besonders auf den zentralen Bereich der 
Rhizomorphen konzentriert, aber Verlauf unregelmäßig, kaum zentral gebündelt; Knoten- 
bildungen (Agerer 1999) fehlend, rückwärts gerichtete Verzweigungen und Vorläuferhyphen 
nicht beobachtet; Hyphen der Rhizomorphenperipherie 2,5-4 ^m dick, dickwandig (Wand- 
stärke 0,5-1 |im), parallel zur Rhizomorphenrichtung, von Hyphen tieferer Lagen unterscheid- 
bar (dickwandiger, siehe Abb. 6), Schnallen fast durchwegs vorhanden, aber auch einfache 



118 



Septen nachweisbar; selten Doppelschnallen vorkommend; keine differenzierten Oberflächen- 
zellen vorhanden; Hyphen tieferer Lagen 2-4 ^im dick, dünnwandig, zumeist parallel ver- 
laufend; Schnallen durchwegs vorhanden; in allen Bereichen der Rhizomorphe (auch ober- 
flächennah) zusätzlich Skeletthyphen mit völlig oder nahezu reduziertem Lumen eingestreut, 
nur im Bereich der Septen geringe Restlumina regelmäßiger zu beobachten, septenlose Ver- 
zweigungen auftretend, Skeletthyphen farblos bis blaß gelbbräunlich pigmentiert, direkt aus 
undifferenzierten Hyphen hervorgehend; Schnallenbögen der Skeletthyphen meist erhalten und 
auifaUig; gefaßartige Hyphen 4-6 ^m dick, nur wenig dicker als umgebende, undifferenzierte 
Hyphen, daher nur wenig auffallend; gefaßartige Hyphen mit verdickten Zellwänden (bis 1 
)im). Schnallenbögen reduziert und vielfach nicht mehr zu beobachten oder nur angedeutet. 
Sehr junge Rhizomorphen (Abb. 7) ohne gefäßartige Hyphen, Zellwände nicht verdickt, 
Schnallen vorhanden, aber auch viele einfache Septen, zudem immer wieder Doppelschnallen 
vorhanden, hieraus auch Hyphen auswachsend; Anastomosen sehr häufig auftretend; sehr 
dünne, kurze, seitliche Auswüchse bei Hyphen der frühesten Rhizomorphenstadien 
auftretend; Rhizomorphen sich durch Zusammenwachsen und Bündeln von Einzelhyphen 
bildend, keine langgestreckten Vorläuferhyphen auftretend. 

Freies Mycel aus Holzsubstrat (Abb. 7): Hyphen 2-3 |im dick, Zellwände nicht verdickt; 
Verzweigungen und Anastomosen häufig, Zellen meist mit Schnallen, aber sehr häufig auch 
einfache Septen beobachtet, zudem auch Doppelschnallen vorkommend; häufig sich sehr dün- 
ne, rankenartige Auswüchse an den Substrathyphen bildend, sowohl septennah (aus dem 
Schnallenbogen oder ihm gegenüber) als auch hiervon unabhängig auftretend, häufig mehrere 
dieser Auswüchse aus einer Zelle entspringend, Auswüchse sehr häufig knotenartig verdreht, 
„Knäuel" bildend, hierbei immer wieder kurze, ebenso dünne Seitenästchen entspringend, sel- 
tener Auswüchse mit anderen Hyphen anastomisierend. 

Kulturmycel (Abb. 8): Hyphen 2,5-6 ^m dick, dünnwandig; sowohl Schnallen als auch ein- 
fache Septen vorhanden; häufige Bildung von Chlamydosporen, deren Bildungszellen meist 
einfache Septen aufweisen; Chlamydosporen (Abb. 8) sich in meist kurzzelligen, schnallen- 
losen Hyphenbereichen bildend, Chlamydosporen meist dickwandig (bis 3 [im Wandstärke), 
7-20 X 3-1 1 |im, sehr vielgestaltig, von langgestreckt zylindrisch bis hin zu zitronenförmig 
oder kugelig, Inhah farblos, blaß gelblich oder sehr hell bräunlich, reich an kleinen Öltröpfchen, 
Wände leicht bis sehr stark verdickt, farblos; einsetzende Chlamydosporenbildung durch 
Konzentration des Cytoplasmas erkennbar, im Verlauf der Bildung alle Öltröpfchen und 
erkennbaren Plasmabestandteile der Zelle in die Spore verlagert; freigesetzte Chlamydosporen 
zumeist blaß pigmentiert; alte, das Wachstum einstellende Kultur mit sehr großer Zahl an 
Chlamydosporen, Hyphen in diesem Stadium mit großteils einfachen Septen und nur hin und 
wieder auftretenden Schnallen. 

Lamellentrama (Abb. 9a, b): Hyphenverlauf jung unidirektional und leicht divergierend, 
ausgewachsene Lamellen mit bidirektionaler Hyphenanordnung; Subhymenium deutlich aus- 
geprägt, aus 2-A um dicken, dünnwandigen, sich verzweigenden Hyphen bestehend; Hyphen 
der lateralen Lamellentrama auch alt divergierend, dünnwandig, selten etwas dickwandig; 
Wände z.T. in KOH-Lösung (z.B. 5%) deutlich aufquellend; Hyphen der zentralen Lamellen- 
trama unregelmäßig verflochten, jedoch mit zwei Vorzugsrichtungen, zum einen parallel zuein- 
ander und senkrecht zur Huttrama in Richtung Schneide verlaufend, zum anderen parallel zur 
Schneide und somit radial bezüglich Hut (bidirektionaler Hyphenverlauf); im Lamellenlängs- 
schnitt (Schnitt parallel zu Lamellenflächen durch die Schneide) bidirektionaler Hyphenverlauf 
besonders deutlich (Abb. 9b): Hauptverlaufsrichtung schneiden wärts gerichtet, hiervon durch 
Krümmung oder Verzeigungen einzelne Hyphen in ca. 90°-Winkel die Laufrichtung ändernd. 



119 



sich mit anderen Hyphen streckenweise bündelnd, um wieder vereinzelt durch Rückkrümmung 
in alten, schneidenwärts gerichteten Verlauf einzumünden; Hyphen der zentralen Lamellen- 
trama mit leicht verdickten, in KOH-Lösung deutlich quellenden Zellwänden, dies besonders 
im Hyphenquerschnitt zu erkennen. 

Die aufgeführten Merkmale treten entsprechend auch bei T. panuoides var. ionipus auf, nur 
die Kulturmerkmale konnten hier nicht geprüft werden, da die untersuchte Ausfammlung nicht 
anwuchs. 

Für weitere Fruchtkörpermerkmale sei auf Sutara (1992) verwiesen. 



Diskussion 

Die Rhizomorphen von Tapinella atrotomentosa und T. panuoides weichen in ihrem Bau von 
den innerhalb der Boletales weit verbreiteten, hochdifferenzierten Rhizomorphen mit Knoten- 
bildungen ab (siehe Agerer 1999). 

Die Ontogenie typischer Boletales-Rhizomorphen (Agerer 1999, für Paxillus involutus 
siehe Agerer 1988) beginnt mit einer langgestreckten Vorläuferhyphe mit großen Septen- 
abständen. Im proximalen Bereich bildet diese Hyphe untergeordnete Seitenzweige, die wie- 
derum durch spezielle, weitere Verzweigungvorgänge (siehe Agerer 1 999) Knoten formieren, 
während sich die primäre Vorläuferhyphe in die erste gefaßartige Hyphe durch Verdickung und 
durch völlige oder teilweise Septenauflösung zu differenzieren anfangt. Durch eine Vielzahl 
von normalen sowie rückwärts gerichteten Verzweigungen wird die Rhizomorphe allmählich 
aufgebaut und verdickt. Die primären Seitenhyphen wiederholen diesen Vorgang. Weitere 
gefaßartige Hyphen differenzieren sich Schritt für Schritt. Dadurch entsteht ein System sich 
durch Knoten verzweigender Rhizomorphen mit gebündelten, zentralen, gefaßartigen Hyphen. 

Die Rhizomorphen von Tapinella entwickeln sich hingegen ohne die Bildung von Vor- 
läuferhyphen. Es kommen keine Knotenbildungen vor, rückwärts gerichtete Verzweigungen 
konnten lediglich bei T. atrotomentosa festgesteUt werden. Im Verlauf der Ontogenie differen- 
zieren sich zwar ebenfalls gefäßartige Hyphen, sie sind bei T. panuoides jedoch innerhalb des 
Rhizomorphenzentrums nur locker verteilt und verlaufen unregelmäßig (= „phlegmacioid", 
Agerer 1999). Zudem lösen sich die Septen nur vereinzelt auf, wie auch ihr Durchmesser 
kaum vergrößert wird. Tapinella atrotomentosa besitzt deutlicher zentral verlaufende gefäß- 
artige Hyphen, deren Durchmesser sich auch deutlich vergrößert („agaricoid" nach Agerer 
1999). Tapinella besitzt somit eine andere Rhizomorphenorganisation als die Mehrzahl der 
untersuchten Gattungen der Boletales (Agerer 1999). Lediglich die auch aufgrund ihrer para- 
sitischen Lebensweise ungewöhnlichen Vertreter der Gomphidiaceae sowie die Trunco- 
columellaceae weisen noch ursprünglichere Rhizomorphen auf (Agerer 1999). 

Skeletthyphen bei Rhizomorphen und die Bildung von rankenartigen, seitlichen Hyphen- 
auswüchsen sind spezielle Merkmale von T. panuoides, während die Doppelschnallen und das 
Auftreten von einfachen Septen für die Gattung Tapinella typisch sind. 

Aus der abweichenden Organisation der Rhizomorphen und der Lamellentrama wird ge- 
schlossen, daß Tapinella eine Schwestergruppe der übrigen Boletales ist (siehe auch Agerer 
1999). 

Die Rhizomorphen der Gattung Paxillus s.str. (/'ax///M5-/«vo/Mfw5-Formenkreis, siehe Hahn 
& Agerer 1999) unterscheiden sich abgesehen von ihrerer „boletoiden" Organisation von 
Tapinella auch in weiteren Merkmalen. In der Gattung Paxillus treten an der Oberfläche der 



120 



Rhizomorphen ausgeprägte Mycelcystiden mit apikal verdickten, kappenartigen Zellwänden, 
z.T. auch mit Papillen aus massiver Zellwand auf (Hahn & Agerer 1999). Zudem kommen 
aufgeblähte Zellen mit verdickten äußeren Zellwänden nestweise an der Rhizomorphenober- 
fläche vor. Ein weiterer Unterschied liegt in der Bildung von Kristallen bzw. punktierten 
Zellwänden bei Paxillus s.str. Die für Paxillus s.str. typischen Sklerotien (siehe z.B. Hahn & 
Agerer 1999) wurden bislang bei Tapinella nicht nachgewiesen. Hingegen sind sie auch von 
einigen anderen Paxillaceae bekannt, z.B. von Austropaxillus boletinoides (Singer) Bresinsky & 
Jarosch (Palfner, pers. Mitt.), Boletinellus merulioides (Schwein.) Murr. [Pantidou 1961a, als 
Gyrodon merulioides (Schwein.) Singer, Cotter & Miller 1985], Gyrodon lividus (Agerer 
et al. 1993), Phlebopus sudanicus (Har. & Pat.) Heinem. (Thoen & Ducousso 1989). 
Weiterhin wurden Sklerotien nachgewiesen bei Hygrophoropsis aurantiaca (Antibus 1989, 
Clemenqon 1997) und Leucogyrophana spp. Pouzar (Ginns 1976). 

In Kultur formt nur Tapinella und nicht Paxillus s.str. Chlamydosporen. Allerdings sind 
Chlamydosporen auch bei wenigen weiteren Boletales bekannt, so z.B. bei Coniophora 
(Ginns 1982), Buchwaldoboletus hemichrysus (Berk. & Curt.) Pilät [Pantidou 1961b, als 
Phlebopus sulphureus (Fr.) Singer], Gomphidius roseus (Fr.) Karst. (Agerer 1990, 1991), 
Rhizopogon vinicolor A.H.Smith (Müller & Agerer unpubl.) und Truncocolumella citrina 
(Eberhart & Luoma 1996). Die Bildung von Chlamydosporen erfolgte somit iimerhalb der 
Boletales mehrfach konvergent. 

Auch im Bereich der Fruchtkörper lassen sich Paxillus s.str. und Tapinella anhand einer 
Reihe von Merkmalen unterscheiden, wie bereits Sutara (1992) darlegte (siehe Tab. 1). Als 
weiterer, gewichtiger Unterschied ist der in dieser Untersuchung gefundene bidirektionale 
Aufbau der Lamellentrama bei Tapinella anzusehen. Dieser Organisationstyp ist bei Lamellen- 
pilzen die Ausnahme und tritt vor allem bei omphalinoiden Vertretern (Gerronema Singer, 
Phytoconis Bory, Haasiella Kotl. & Pouzar und Chrysomphalina Clemen9on) auf (Cle- 
MENCON 1997). Auch bei Lyophyllum decastes (Fr.) Singer ist nach Clemenqon (1997) in der 
Nähe der Schneide ein bidirektionaler Aufbau der Lamellentrama bekannt. Bei Vertretern der 
Boletales wurde dies bislang noch nicht beobachtet. 

In der Lamellentrama von Paxillus s.str. (siehe Hahn & Agerer 1999) kann zwar in den 
Lateralstrata z.T. eine gelatinöse Matrix auftreten, in die die Hyphen eingebettet sind, die Zell- 
wände quellen jedoch im Gegensatz zu Tapinella nicht in KOH auf Eine gelartige Matrix ist 
bei Tapinella noch nicht beobachtet worden. 

Obwohl die Fruchtkörper von Tapinella und Paxillus s.str. sich makroskopisch sehr ähneln, 
zeigen sich sowohl im Bereich des vegetativen Mycels (Rhizomorphen und Substratmycel), im 
Verhalten in Kultur (Chlamydosporen) wie auch in der Anatomie der Fruchtkörper (z.B. 
bidirektionale Lamellentrama) deutliche Unterschiede. Anhand der Fülle von unterschiedlichen 
Merkmalen und Merkmalsbereichen (siehe Tab. 1) sind die Fruchtkörper von Paxillus s.str. 
und Tapinella als Konvergenzen aufzufassen. 



121 



Tab. 1.: Vergleich won Paxillus s.str. mit Tapinella. Mit * gekennzeichnete Merkmale wurden 
bereits von Sutara ( 1 992) angegeben. 



Paxillus s.str. 


Tapinella 


Caulohymenium ausgebildet* 


Caulohymenium fehlend* 


Lamellentrama unidirektional (Hahn & 
AGERER1999) 


Lamellentrama bidirektional (Abb. 5a, b, 
9a, b) 


Lateralstrata der Lamellentrama z.T. 
gelatinös (Hahn & Agerer 1999)* 


Lamellentrama nicht gelatinös* 


Hymenialcystiden vorhanden 


Hymenialcystiden fehlend 


Keine Bildung von Doppelschnallen 


Doppelschnallen im Mycel fakultativ 
auftretend (Abb. 7) 


Nur Schnallen, keine einfachen Septen 
vorhanden 


Häufig einfache Septen auftretend 


Rhizomorphen mit Knotenbildung 
(Hahn & Agerer 1999), boletoid nach 
AGERER (1999) 


Rhizomorphen ohne Knotenbildung, 
agaricoid oder phlegmacioid nach Agerer 
(1999) 


Sklerotien vorhanden (z.B. Hahn & 
AGERER 1999) 


Sklerotien fehlend 


Keine Bildung von Chlamydosporen 


Bildung von Chlamydosporen im Mycel 
(in Kultur, Abb. 4, 8) 


Ektomykorrhiza-bildend* 


Lignicol (noch keine Ektomykorrhiza 
nachgewiesen)* 



Die vom Erstautor früher (Hahn 1996) geäußerte Vermutung, Tapinella könnte durch 
sekundären Übergang auf Holz sich aus Paxillus s.str. ähnlichen, noch Mykorrhiza bildenden 
Arten durch Reduktion der Fruchtkörper und Mykorrhiza hervorgegangen sein, kann daher als 
widerlegt angesehen werden, nachdem auch neuere molekularbiologische Arbeiten (z.B. 
BuESiNSKYet al. 1999, BRUNSet al. 1998) eine Stellung von Tapinella als Schwestergruppe zu 
den übrigen Boletales bestätigen. Wie sich zeigt, sind neben den heute populären DNA- 
Sequenzdaten auch anatomische Merkmale der Rhizomorphen sehr gut für die Bearbeitung 
systematischer Fragestellungen geeignet. Sie stellen zweifelsohne einen sehr konservativen 
Merkmalsbereich dar, was auch andere Untersuchungen (Agerer 1999, Agerer in Vorher., 
Hahn et al. 2000) bestätigen. 

Ob Tapinella atrotomentosa und T. panuoides wiederum untereinander monophyletischen 
Ursprungs sind, ist nach NiLSSON & GiNNS (1979) zweifelhaft, da sich die beiden Arten hin- 
sichtlich ihres Braunfäuletyps unterscheiden. Tapinella panuoides ist nach NiLSSON & Ginns 
(I.e.) im Gegensatz zu T. atrotomentosa in der Lage, in Kultur freie Cellulose abzubauen. 

Die erhobenen anatomischen Daten sprechen jedoch für eine verwandtschaftlich homogene 
Gattung Tapinella. Grundlegend ist der bidirektionale Aufbau der Lamellentrama, weiterhin die 
in Ontogenie und Bau sehr ähnlichen Chlamydosporen. Auch die bei Tapinella sterile Stielbe- 
kleidung, die auch nicht in Ansätzen das fiir Paxillus s.str. und viele Boletaceae durchgehende 
Caulohymenium (siehe z.B. Breitenbach & Kränzlin 1991, Sutara 1987, 1989, 1992) 
aufweist, sondern trichodermal organisiert ist, unterstreicht die Eigenständigkeit von Tapinella. 
Viele dieser gemeinsamen, aussagekräftigen Merkmale stellen Autapomorphien dar, und 
weisen auf die Monophylie von Tapinella hin (siehe auch Tab. 1). 



122 



Molekularbiologische Untersuchungen (Bresinsky et al. 1999) belegen die Monophylie von 
Tapinella mit einem Bootstrap- Wert von 99% (Sequenzierung des durch die Primerkom- 
bination LR0R-LR5 erhaltenen, 900 Basenpaar großen Abschnittes der kemcodierten 28S 
rDNA). 

Die Gattung Tapinella besteht somit zu Recht imd die belegten gravierenden Unterschiede 
zu Paxillus s. Str., die Tapinella ebenso von Austropaxillus und Gyrodon unterscheiden (Hahn 
unpubl.), erfordern die Errichtung einer separaten Familie Tapinellaceae: 

Tapinellaceae Ch.Hahn, fam. nov. 

Carpophora camosa, pileo excentrico vel laterale vel sessile, basidiomata rare campani- 
formia vel resupinata; hymenium lamelhformia; lamellae bifurcatae vel anastomisantes, decur- 
rentes; trama lamellarum aetate progresso bidirectionalis; cystidia deficientia; pilei- et stipiti- 
pellis trichodermiformes vel pilosa, hymenium stipitis deficiens; sporae ellipticae, laeves; 
Rhizomorphae complexae, hyphis vasiformibus parietibus incrassatis, nodi deficiens; cystidia 
mycelii deficiens; hyphae fibuligerae, rare bifibuligerae, sed septa simplicia interdum prae- 
sentia; mycelia chlamydosporigerae; habitat: lignicol. 
Typus: Tapinella Gilbert. 

LOQUIN (1984) erwähnt die Familie Tapinellaceae und bezieht sich auf seine frühere Pu- 
blikation (LOQUIN 1981). Da ausgedehntes Suchen nach dieser Veröffentlichung nicht er- 
folgreich war und die Tapinellaceae nicht in Hawksworth et al. (1995) als gültig publi- 
zierte Familie aufgeführt werden (s. auch BRESINSKY et al. 1999), werden die Tapinellaceae 
in der vorliegenden Arbeit gültig beschrieben. 

Folgende Umkombinationen sind durch die Anerkennung der Gattung Tapinella nötig: 

Tapinella panuoides var. ionipus (Quel.) Ch.Hahn, comb. nov. 

Basionym: Paxillus ionipus Quelet „1887", 1888, Bull. Assoc. France pour l'Avancement 
des Sciences: 588. 

„Paxillus'' ionipus unterscheidet sich von Tapinella panuoides nur hinsichtlich des violetten 
Basalmycels, eines violett gefärbten Stielfilzes und selten auch durch violette Farbtöne am Hut 
(im Übergangsbereich vom Stiel zum Hut). Wie auch Krieglsteiner (1982) feststellt, 
schwankt die Intensität der violetten Färbung, und es gibt Übergänge zu Tapinella panuoides s. 
Str. Eine Trennung dieser beiden Taxa auf Artniveau ist somit kaum zu begründen. 

Tapinella panuoides var. panuoides frn. acheruntius (Humboldt ex J.Schroet.) Ch.Hahn, 
comb. nov. 

Basionym: Paxillus acheruntius Humboldt ex Schroeter 1889 in Cohn: Kr.-Fl. von 
Schlesien, 3. Bd., 1. Hälfte, Pilze, p. 515. 

,J'axillus'' acheruntius wird nur als resupinate bis glockenförmig angeheftete Wuchsform 
unter Lichtausschluß (z.B. in Bergwerken, siehe Schroeter 1889 und Abbildung bei Lotsy 
1907: 717) angesehen und ist wohl nicht genetisch fixiert. Ein erst kürzlich vom Erstautor ge- 
sammelter Beleg von Tapinella panuoides in einem Bergwerk (CH 36/99 - siehe Material und 
Methoden) zeigte Übergänge von der gewöhnlichen, seitlich gestielten, muschelförmigen 
Fruchtkörperausprägung zu auf der Rückseite kurz gestielten, glockenförmigen und auch 
effuso-reflexen Fruchtkörpern. Besonders sehr junge Exemplare begannen häufig hängend 



123 



schüsselfbrmig zu wachsen. An der Fundstelle (Stollenbeginn, bereits unter Tage) war 
allerdings eine dauerhafte Kunstlichtquelle angebracht. Die effuso-reflexen Fruchtkörper ent- 
wickelten sich vor allem in stärker abgeschatteten Bereichen. Es liegt nahe, in T. panuoides fm. 
archeruntius eine reine Dunkelheitsform zu sehen. Aufgrund der potentiellen phylo- 
genetischen Bedeutung dieser im Extrem vöUig resupinat wachsenden Form soll das Taxon 
jedoch durch die Umkombinierung einen eigenen Namen als forma behalten. 



Der Erstautor dankt der Leitung des Graphitbergwerkes Kropfmühl für die Erlaubnis, Pilz- 
fruchtkörper unter Tage aufsammeln zu dürfen, sowie Herrn H. Forstinger (Ried/Innkreis, Öster- 
reich) für die Organisation dieser Exkursion, Herrn E. Gamweidner (Fürstenfeldbruck) für die 
Weitergabe frischen Belegmaterials mit Rhizomorphen {Tapinella panuoides var. ionipus), Herrn 
T. R. Lohmeyer (Taching) für die Beschaffung der Originalbeschreibung von ,yPaxillus ionipus". 



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Christoph Hahn & Prof. Dr. Reinhard Agerer; Institut für Systematische Botanik der 
Universität München, Menzinger Straße 67, D-80638 München, Deutschland. 



125 




Abb. 1: T. atrotomentosa, CH 201/98: Rhizomorphe, Kryotomschnitt bis in Rhizomorphen- 
mitte; gefäßartige Hyphen an dieser Stelle gebündelt. 



126 




Abb. 2: T. atrotomentosa, CH 201/98: Rhizomorphe, Zentralbereich (Kryotomschnitt); ein 
aufgelöstes Septum einer gefäßartigen Hyphen erkennbar. 



ia7 




Abb. 3: T. atrotomentosa, CH 214/98: Rhizomorphe und freies Mycel; a: junge Rhizomorphe, 
Zellwände leicht verdickt, stellenweise einfache Septen auftretend; b: freies Substratmycei, 
Anastomosenbildung, z.T. über Kontaktschnallen. 



128 





Abb. 4: T. atrotomentosa, CH 164/98: oben: Chlamydosporenbildung innerhalb junger Rhizo- 
morphen aus Kultur; unten: Einzelhyphen mit Chlamydosporen, eine Chlamydospore frei. 



129 




g 




Abb. 5: T. atrotomentosa, CH 216/98: Lamellentrama: a: Schnitt tangential zum Hutrand 
(Kryotomschnitt); b: Schnitt radial zum Hutrand durch Schneide(Kryotomschnitt). 



130 




Abb. 6: T. panuoides, CH 183/98: Rhizomorphe, Kryotomschnitt bis in Rhizomorphenmitte; 
gefäßartige Hyphen mit aufgelösten Septen. 



131 




Abb. 7: T. panuoides, CH 183/98: junge Rhizomorphen und freies Substratmycel aus mor- 
schem Holz: junge Rhizomorphen mit vielen Anastomosen; Doppelschnallen durch * ge- 
kennzeichnet, Verzweigungen aus Doppelschnalle mit • markiert; einzelne Hyphen mit 
dünnen Auswüchsen, an denen z.T. Sporen eines imperfecten Pilzes kleben. 



132 




Abb. 8: T. panuoides, CH 183/98: Chlamydosporenbildung in freiem Mycel aus Kultur. 



133 




■H(H 




Abb. 9: T. panuoides var. ionipus, CH 242/98: Lamellentrama: a: Schnitt tangential zum 
Hutrand (Kryotomschnitt); b: Schnitt radial zum Hutrand durch Schneide (Kryotomschnitt). 



135 



New Astragali and Oxytropis from North Africa and Asia, 
including some new combinations and remarks on some species 



D. PODLECH 



Abstract: 

PODLECH, D.: New Astragali from North Africa and Asia, including some new 
combinations and remarks on some species. - Sendtnera 6: 135-174. 1999. ISSN 
0944-0178. 

One new species from Morocco and 39 new species and subspecies from Asia of the 
genus Astragalus and one species of the genus Oxytropis {O. azerbaijanica) are 
described here. They belong to the following sections: Sect. Aegacantha: A. langta- 
nensis, A. nepalensis; Sect. Astragalus: A. diyarbakirensis, A. nigrocalycinus; Sect. 
Brachycarpus: A. bowes-lyonii,A. thomsonii;A. villosulus; Sect. Caprini: A. akhanii, 
A. antalyensis,A. avajensis, A. diversipilosus, A. glaberrimus, A. malatyaensis; Sect. 
Cytisodes: A. neyshaburensis; Sect. Dissitißori: A. fallacinus; Sect. Hypoglottidei: 
A. cedreticola; Sect. Incani: A. brevidentatus, A. longisubulatus, A. mercklinii subsp. 
farsicus, A. nevshehiricus, A. polhillii; Sect. Laxiflori: A. gigantostegius, A. karasa- 
rensis: Sect. Leucocercis: A. kentrophyllus; Sect. Malacothrix: A. bakirdgahensis, 
A. jacobsii, A. kashanensis, A. sultan-bulaghensis; Sect. Mikrophyton: A. mikrophy- 
toides; Sect. Onobrychoidei: A. arakliensis, A. demirizii, A. eskishehiricus, A. konya- 
ensis, A. pineticola; Sect. Rhabdotus: A. venulosus subsp. iraqensis; Sect. Sisyro- 
phorus: A. nigdeanus; Sect. Stereothrix: A. hakkariensis, A. montis-varvashti; Sect. 
Tapinodes: A. tetuanensis; Sect. Trachycercis: A. longivexillatus . Furthermore two 
new sections: Aberrantes and Mikrophyton and 9 new names and combinations are 
presented. 

Zusammenfassung: 

Eine neue Art der Gattung Astragalus aus Nordafrika und 39 neue Arten und Unter- 
arten aus Asien sowie eine neue Oxytropis- \tX {O. azerbaijanica) werden hier be- 
schrieben. Sie gehören folgenden Sektionen an: Sect. Aegacantha: A. langtanensis, 
A. nepalensis; Sect. Astragalus: A. diyarbakirensis, A. nigrocalycinus; Sect. Brachy- 
carpus: A. bowes-lyonii, A. thomsonii; A. villosulus; Sect. Caprini: A. akhanii, 
A. antalyensis,A. avajensis, A. diversipilosus, A. glaberrimus, A. malatyaensis; Sect. 
Cytisodes: A. neyshaburensis; Sect. Dissitiflori: A. fallacinus; Sect. Hypoglottidei: A. 
cedreticola; Sect. Incani: A. brevidentatus, A. longisubulatus, A. mercklinii subsp. 
farsicus, A. nevshehiricus, A. polhillii; Sect. Laxiflori: A. gigantostegius, A. karasaren- 
sis: Sect. Leucocercis: A. kentrophyllus; Sect. Malacothrix: A. bakirdgahensis, A. ja- 
cobsii, A. kashanensis, A. sultan-bulaghensis; Sect. Mikrophyton: A. mikrophytoides; 
Sect. Onobrychoidei: A. arakliensis, A. demirizii, A. eskishehiricus, A. konyaensis, 
A. pineticola; Sect. Rhabdotus: A. venulosus subsp. iraqensis; Sect. Sisyrophorus: A. 
nigdeanus; Sect. Stereothrix: A. hakkariensis, A. montis-varvashti; Sect. Tapinodes: 
A. tetuanensis; Sect. Trachycercis: A. longivexillatus. Ferner werden 2 neue Sektionen: 
Aberrantes and Mikrophyton und 9 neue Namen und Kombinationen mitgeteilt. 



136 



1. New species and sections 

Sect. Aberrantes Podlech, sect. nov. 
Typus sectionis: A. clausii C.A.Mey. 

Differt a sect. Alopecuroidei racemis laxis, pedicellis perlongis, calyce dentibus longis 

corollam superantibus, vexillo piloso, lamina carinae porrecta. 

. • • - . - ^t ' 

Plantae validae, erectae, dense longe albovillosae. Stipulae angustae, basi petioli adnatae. 
Racemi laxe multiflori, longe pedunculati. Pedicelli perlongi, bracteolis filiformibus instruct!. 
Calyx campanulato-tubulosus, dentibus filiformibus tubo aequilongis vel longioribus. Corolla 
calyce brevior. Petala breviter unguiculata. Vexillum dorse pilosum. Carina lamina porrecta. 

Plants tall, erect, long white-villous. Stipules narrow, adnate at the the base to the petiole. 
Racemes loosely many-flowered, with a long peduncle. Pedicels long, with filiform bracteoles. 
Calyx campanulate-tubular; teeth filiform, as long as the tube or longer. Petals shorter than the 
calyx, with a short claw. Standard hairy on upper side. Keel with straight blades. 

The two known species {A. clausii C.A.Mey., A. vassilczenkoi Berdyev) of this aberrant 
section have been up to now always put into the section Alopecuroidei DC. although the 
descriptions fit by no means the description of this section. Becht 1 978 without having seen 
the type thought A. clausii to be a synonym o{ A. vulpinus Willd., a very curious procedure. 

Astragalus langtangensis Podlech, spec. nov. e sect. Aegacantha 

Holotype: Nepal, [Langtang Himal], Langtang valley, 13000 ft., 24.6.1965, Achilling, 
Sayers & Bista 380 (K). 

Fruticulus spinosus ad 30 cm altus, ramosus. Stipulae membranaceae, 7-10 mm longae, li- 
neares vel anguste triangularibus, petiolo per 3 mm adnatae, margine albo-ciliatae. Folia juveni- 
lia imparipinnata, 9-1 1-juga 3-4 cm longa, petiolo 1-1,5 cm longo, rachide pilis patentibus 1-2 
mm longis obtecta. Rachides foliorum adultorum 3,5-7 cm longae, erectae, pungentes. Fo- 
liolum terminale minutum, demum deciduum. Foliola cetera obovata, 3-4 x 2-2,5 mm, supra 
glabra, subtus laxe longe subpatenter pilosa. Pedunculus ca. 5 mm longus, 2-florus. Bracteae 4-5 
mm longae. Calyx membranaceus, 11-12 mm longus, tubulosus, laxe ad densiuscule pilis 
flexuosis albis nigris immixtis patentibus ad 1 mm longis obtectus, dentibus subulatis 3—4 mm 
longis. Petala flava. Vexillum ca. 22 x 7 mm, apice incisa. Alae ca. 20 mm longae. Carina 1 7 
mm longa, laminis oblique ellipticis, apice breviter acutiusculi-rostrata. Legumina brevissime 
stipitata, oblique elliptica, 10-13 mm longa, 5-7 mm alta, 5-6 mm lata, ventro carinata, dorso 
applanata vel leviter et late depressa, apice subabrupte in rostrum tenuem 5-8 mm longum 
attenuatum, complete biloculare, valvis tenacibus, stramineis, tenuiter transversaliter rugosis, 
laxe pilis tenuibus albis ad 1 mm longus appressis ad subpatentibus obtectis. 

Plants fruticose, spiny, up to 30 cm tall, with a thick, branched caudex. Main stems up to 8 
mm in diameter, with blackish bark, densely covered with remnants of old stipules, glabrous, 
in upper part branched. Branches erect, very densely leafy, growth rate of the year 1-2 cm. 
Stipules membranous, 7-10 mm long, linear to narrowly triangular, adnate to the petiole for 3 
mm, otherwise free, white ciliate at the margins, otherwise glabrous. Leaves in the youth im- 



137 



paripinnate, 3-4 cm long, with a petiole 1-1.5 cm long, rachis flexible, loosely covered with 
slender, flexuous, spreading white hairs 1-2 mm long. Old leaves 3.5-7 cm long, the rachis 
erect, slender but rigid, pungent, brown-blackish, glabrescent to glabrous, sometimes without 
leaflets. Terminal leaflet minute, soon falling. The other leaflets in 9-1 1 pairs, obovate, 3-4 x 
2-2.5 mm, at the apex truncate to emarginate, minutely mucronulate, on upper side glabrous, 
on underside loosely covered with long, nearly spreading hairs, flat or complicated. Raceme 
with a peduncle ca. 5 mm long, 2-flowered. Bracts whitish- membranous, linear-acute, 4-5 mm 
long, white and black-hairy. Pedicels 3—4 mm long, erect, white and black-hairy. Calyx mem- 
branous, pale greenish-yellow, sometimes red-suffused, 11-12 mm long, tubular, loosely to 
rather densely covered with flexuous, spreading white and some black hairs up to 1 mm long; 
teeth subulate, 3-4 mm long. Petals yellow. Standard ca. 22 mm long, narrowly elliptic, ca. 7 
mm wide, at the apex incised, at the base gradually narrowed into the long claw. Wings ca. 20 
mm long; blades narrowly gibbous-elliptic, obtuse at the apex, 8x3 mm, auricle ca. 1 mm 
long, claw 11-12 mm long. Keel 17 mm long; blades obliquely elliptic, with gibbously curved 
lower edge and nearly straight upper edge, at the apex shortly acutish-rostrate; auricle 
indistinct, claw ca. 10 mm long. Staminal-tube obliquely cut. Ovary subsessile, white-hairy; 
style hairy only in basal part. Legums subsessile, obliquely elliptic, 10-13 mm long, 5-7 mm 
high, 5-6 mm wide, carinate ventrally, flat to slightly and widely depressed dorsally, at the 
apex subabruptly narrowed into the slender beak 5-8 mm long, fully bilocular; valves tough, 
straw-colored, thinly transversely rugose, loosely covered with appressed to nearly spreading, 
slender white hairs up to 1 mm long. 

Other specimens seen: 

India. Uttar-Pradesh : Kumaon, Distr. Almora, near Rilkote, Upper Johar, 11500-12000 ft., 
5.10.1 950, Awashti 1842 (M) - Kumaon, Jola, 3500 m, Strachey & Winterbottom (P). 

Astragalus nepalensis Podlech, spec. nov. e sect. Aegacantha 

Holotype: Nepal, Kali Gandaki, trail above Tukuceh, 8500 ft., 1.5.1967, A.Nicolson 82a 
(US). 

Fruticulus spinosus ad 30 cm altus. Caulis erectus, ad 5 mm crassus, ramosus, pilosus. 
Rami laterales in parte basali caulis remoti, ad 8 cm longi, intemodiis ad 5 mm longis, partim 
iterum ramosi, in parte apicali approximati, breviores, intemodiis quam stipulae brevioribus. 
Incrementum annuale ad 1 cm longum. Stipulae triangulares, 2-4 mm longae, petiolo per 1 mm 
adnatae, pilosae. Folia paripinnata, 3-4-juga. Rachides foliorum adultorum 2-4 cm longae, 
erecto-patentes, pungentes, laxe pilosae, foliolis delapsis. Folia juvenilia 1-2,5 cm longa, ra- 
chide flexibile. Foliola crassiuscula, elliptica vel obovata, 2-4 x 1-2 mm apice obtusa vel bre- 
vissime apiculata, supra sparse ad laxe subtus densiore subappresse pilosa, fere semper com- 
plicata. Pedunculi subnuUi, 2-4 flori. Bracteae ca. 2 mm longae. Calyx 9-10 mm longus, tubu- 
losus, dense albi-nigrescenti-villosus, dentibus 2-3 mm longis. Petala flava, in siccis purpura- 
scentes, glabra. Vexillum 18-19 x 7-8 mm, obovatum, apice incisum. Alae 17 mm longum, 
auricula 1-1,5 mm longa. Carina 16-17 mm longa. Legumina sessilia, ovata, 8-10 mm longa, ca. 
4 mm alta et 5 mm lata, rostro 2-3 mm longo, subbiloculare, valvis tenacibus stramineis, laxe 
pilis ad 1 mm longis subappressis ad ascendentibus albis et nigris obtectis. 

Plants fruticose, spiny, up to 30 cm tall. Stem erect, up to 5 mm in diameter, branched, 
slightly sulcate, with blackish bark, covered with persistent, appressed, dirty whitish or 



138 



greyish hairs. Lateral branches in lower part of the stem remote, up to 8 cm long, partly 
branched again, in upper part of the stem crowded, shorter, with stem internodes shorter than 
the stipules; growth rate of the year up to 1 cm long. Stipules triangular, 2-4 mm long, adnate 
to the petiole for 1 mm, otherwise free, loosely to densely appressed white-hairy. Leaves 
paripiimate; rachis 2-A cm long, obliquely erect, slender but pungent, the petiole as long or 
shorter than the leaflets-bearing part, the spiny end shorter than the uppermost leaflets; young 
leaves mostly shorter, 1-2.5 cm long, with flexible rachis. Leaflets thickish, elliptic to obovate, 
2-4 X 1-2 mm; at the apex obtuse to minutely apiculate, on upper side sparsely to loosely, on 
underside more densely appressed hairy, nearly always folded. Peduncles nearly absent, 
raceme 2-4-flowered. Bracts whitish-membranous, ca. 2 mm long, ovate-acute, white or 
greyish-hairy. Pedicels 2 mm long, black-hairy. Calyx 9-10 mm long, tubular, densely villous 
with white and black hairs; teeth narrowly triangular, 2-3 mm long, on innerside densely 
black-hairy. Petals yellow, in dry state purplish, glabrous. Standard 18-19 mm long, obovate, 
7-8 mm wide, at the apex incised, at the base gradually narrowed into th claw. Wings 1 7 mm 
long; blades narrowly oblong, obtuse at the apex, 6.5-7 x 2-2.5 mm; auricle 1-1.5 mm long, 
claw 10-1 1 mm long. Keel 16-17 mm long; blades obliquely elliptic, with widely rectangularly 
curved lower edge and nearly straight to slightly concave upper edge, acutish at the apex, 6 x 
3-3.5 mm; auricle short, claw 10-11 mm long. Staminal-tube silightly obliquely cut. Ovary 
subsessile, oblong, white-hairy; style glabrous. Legumes sessile, ovate, 8-10 mm long, ca. 4 
mm high and 5 mm wide, carinate ventrally, widely flat dorsally, therefore triangular in cross- 
section, at the apex abruptly narrowed into a straight beak 2-3 mm long, nearly fiilly bilocuiar; 
valves tough, straw-colored, loosely covered with subappressed to ascending white and black 
hairs up to 1 mm long. Seeds 4 in each locule, 2.5-3 x 2 mm, olive-brownish. 

Other specimen seen: 

Nepal. Muktinath Trek, Marpha, 8700 m, \3.4.\978,aiß'orcl G.Rice 19-78 (VS). 

Astragalus diyarbakirensis Podlech, spec. nov. e sect. Astragalus 

Holotype: Turkey, Prov. Diyarbakir, 12 km W Diyarbakir on road to Siverek, ± 600 m, 
11.5.1988, P. J. Cr/^6(K). 

Differt ab A. columnaris Boiss. planta graciliore, caule ad 4 (nee 8-12) mm crasso, stipulis 
margine ciliatis (nee glaberrimis), foliis 11-15 (nee 19-25) jugis, bracteis ca. 3 (nee 5-7) mm 
longis, ciUatis (nee glabris), calyce 7-8 (nee 1 1-12) mm longo, dentibus 1-1,5 (nee 5-6,5) mm 
longis, vexillo ca. 1 8 (nee 22-26) mm longo. 

Plants herbaceous, ca. 60 cm tall, very sparsely furnished with white hairs 0.4-1 mm long. 
Stem rather slender, ca. 4 mm thick, finely striate-sulcate, glabrous or with scattered spreading 
hairs. Stipules greenish, 5-10 mm long, those of lower leaves narrowly triangular, those of 
upper leaves subulate, alle shortly adnate to the petiole, ciliate at the margins. Leaves 15-20 
cm long; petiole 1-3 cm long, like the rachis slender, very sparsely furnished with spreading 
hairs. Leaflets in 11-15 pairs, remote, elliptic, rounded to slightly retuse at the apex, 8-12 x 
4-6 mm, on upper side glabrous, on underside sparsely covered with appressed to 
subappressed hairs. Racemes subsessile, loosely 2-5-flowered. Bracts whitish-membranous, 
linear-acute, ca. 3 mm long, ciliate. Pedicels 1-2 mm long. Calyx 7-8 mm long, only shortly 
ciliate at the upper margin and the teeth; teeth subulate, slightly unequal, 1-1.5 mm long. Pe- 



139 



tals yellow. Standard ca. 18 mm long; blade ca. 12 mm long, suborbicular, widely emarginate at 
the apex, at the base subabruptly narrowed into the short claw. Wings 16 mm long; blades 
narrowly oblong, obtuse at the apex, 10 x 2.5 mm; auricle ca. 2 mm long, claw 6 mm long. Keel 
14 mm long; blades obliquely oblong-elliptic, with gibbously curved lower edge and slightly 
concave upper edge, obtuse at the apex, 8x4 mm; auricle 1.5 mm long, claw 6 mm long. 
Staminatube obliquely cut. Ovary sessile, oblong, glabrous. Legumes unknown. 

This species resembles A. columnaris in being nearly glabrous, especially in the glabrous 
tube of calyx and ovary. Beside of the given differences yi. columnaris is endemic in south- 
western Turkey, while A. diyarbakirensis is from eastern Turkey. 

Astragalus nigrocalycinus Podlech, spec. nov. e sect. Astragalus 

Holotype: Türkei, A9 Erzurum: zwischen Oltu und Sihsor, 2 km nach der Abzweigung, 
1450 m, 23.5.1990, M Nydegger 45513 (MSB; Iso: BASBG). 

Plantae 40 cm altae vel altiores, pilis albis nigrisque 1-2 mm longis obtectae. Caules validi, 
pilosi. Stipulae 8-12 mm longae, a petiolo liberae, ciliatae. Folia 10-13 cm longa; petiolo 1-2 
cm longo sicut rachis albipiloso. Foliola 10-13 paria, ovata vel elliptica, supra glabra, subtus 
patule albipilosa. Racemi subsessiles, laxe 2-5-flori, axi nigripiloso. Bracteae lineares, 6-10 
mm longae, imprimis nigripilosae. Calyx 14-16 mm long, tubulosus, nigripilosus dentibus 5-6 
mm longis. Petala albida vel pallide flava. Vexillum ca. 30 mm longum, obovatum. Alae 26-27 
mm longae. Carina 22 mm longa. Legumina sessilia, elliptica, 20-25 mm longa, dense patule 
albipilosa. 

Plants at least 40 cm tall, covered with spreading white and black hairs 1-2 mm long. Basal 
parts missing. Stem angular-sulcate, somewhat flexuose, in middle part ca. 8 mm thick, hollow, 
loosely covered with white, below the nodes also with black hairs. Stipules greenish, narrowly 
triangular, ± long acuminate, 8-12 mm long, free from petiole, white and black-ciliate. Leaves 
10-13 cm long; petiole 1-2 cm long, like the rachis striate-sulcate, loosely white-hairy. Leaf- 
lets in 10-13 pairs, ovate to elliptic, at the apex widely rounded to truncate or slightly retuse, 
the terminal leaflet often emarginate, on upper side glabrous, on underside loosely ± spreading 
white-hairy. Racemes subsessile, along at least the upper half of the stem, loosely 2-5- 
flowered; axis 0.5-1 cm long, black-hairy. Bracts greenish, linear-acute, 6-10 mm long, pre- 
dominantly black-hairy. Pedicels 3-4 mm long, black-hairy. Calyx 14-16 mm long, tubular, 
straw-colored, loosely to rather densely spreadingly black hairy; teeth from triangular base 
subulate, 5-6 mm long, on innerside black-hairy. Petals whitish to pale yellow. Standard ca. 30 
mm long; blade obovate, 1 1 mm wide, widely rounded and emarginate at the apex, at the base 
subabruptly narrowed into the short claw. Wings 26-27 mm long; blades narrowly oblong, 
rounded to truncate at the apex, 12-13 x 3.5 mm; auricle 2 mm long, claw 13-14 mm long. 
Keel 22 mm long; blades obliquely oblong-curved, with in the middle gibbously curved lower 
edge and concave upper edge, obtuse at the apex, 9x4 mm; auricle 1 mm long, claw 1 3 mm 
long. Staminaltube straightly cut. Ovary sessile, oblong, densely white-hairy; style glabrous. 
Legumes sessile, elliptic seen from the side, 20-25 mm long, 9-12 mm high and 5 mm wide, 
distinctly laterally compressed, carinate ventrally and dorsally by the thickish nerv, obtuse at 
the apex with a narrowly triangular beak ca 3 mm long, fully bilocular; valves dark brownish. 



140 



lethery, finely rugulose, loosely to rather densely covered with fine, spreading white hairs. 
Seeds 4-5 in each locule. 

The only black-hairy species of the section in Turkey beside of A. caraganae, which is 
aberrant and isolated by its ± long pedunculated racemes. 

Astragalus bowes-lyonii Podlech, spec. nov. e sect. Brachycarpus 

Holotype: Pakistan, Chitral, Shandur, 36°04' N 72°30' E, 12200 ft, 23.7.1958, Bowes Lyon 
92 (E). 

Differt ab A. densifloro plantis acaulibus, pilis brevissimis 0,03-0,2 mm longis obtectis, sti- 
pulis brevioribus, foliis longioribus foliolis remotis, complicatis, racemis tempore fructifica- 
tionis distincte elongatis, calyce imprimis albipiloso, dentibus ejus anguste triangularibus nee 
subulatis, corolla rosea nee violacea, auriculis alarum ca. 1 mm nee 0,5 mm longis, seminibus 4 
nee 2. 

Plants cespitose, ca. 30 cm tall, acaulescent to subacaulescent, covered with very short, 
somewhat flattened, appressed hairs 0.03-0.2 mm long. Caudex with a pluricipital root-crown, 
branches short, diffusely branched. Stems, if present, very short, densely white hairy. Stipules 
greenish, 2-2.5 mm long, triangular or widely triangular, ciliate and white hairy, adnate to the 
petiole for ca. 0.5 mm, otherwise free or sometimes shortly connate behind the stem. Leaves 
7-13 cm long; petiole 2.5—4 cm long, like the rachis slender, loosely subappressed white hairy. 
Leaflets in 5-6 pairs, remote, narrowly elliptic, 5-12 x 1.5-3 mm, acute, on upper side 
glabrous, on underside loosely to rather densely appressed white hairy, always folded. 
Peduncles 1 2-24 cm long, slender, loosely covered with very short white but also with fewer 
black hairs. Racemes ovoid, densely many flowered, ± strongly elongated in fruit. Bracts 
whitish-greenish, 2-3.5 mm long, linear to narrowly ovate, white hairy. Flowers nearly sessile. 
Calyx ca. 4 mm long, campanulate, loosely to rather densely appressed white and black hairy 
but mostly predominantly white hairy; teeth narrowly triangular, ± as long as the tube. Petals 
pink. Standard 6-7 mm long; blade elliptic, 3-3.2 mm wide, widely incised at the apex, at the 
base gradually narrowed into the short claw. Wings 5.5-6 mm long; blades narrowly oblong, 
rounded at the apex, ca. 3.5 x 1 mm; auricle ca. 1 mm long, claw 2-2.5 mm long. Keel ca. 5 mm 
long; blades nearly triangular with widely rectangularly curved lower edge and nearly straight 
upper edge, round-tipped, ca. 2.5 x 1.8 mm; auricle ca. 0.5 mm long, claw 2.5 mm long. Stami- 
nal tube straightly cut. Ovary sessile, globose to ovoid, white hairy; style glabrous. Legumes 
sessile, ovoid, somewhat flattend dorsy-ventrally, 4-4.5 mm long, ca. 2 mm high and 3 mm 
wide, rounded at base and top, flattened ventrally, deeply and widely grooved dorsally, bi- 
locular; valves thin, smooth, ± densely covered with subappressed to ascending white and 
black hairs. Seeds two in each locule. 



Astragalus thomsonii Podlech, spec. nov. e sect. Brachycarpus 

Holotype: Tibet occ, reg. alpina, 12-15000 ft., Thomson (sub .4. densifloro) (MSB; Iso: K, 

M, W). 

Differt ab A. melanostachys Benth. et A. schugnanicus B.Fedtsch. quibus ob caules 
evolutos similis foliolis anguste ellipticis apice rotundatis (nee latioribus apice distincte retusa 



141 



vel emarginate), leguminibus obtriangularibus apice latissimis truncatis latioribus quam longis 
(nee apiee acuminatis vel rostratis, distincte longioribus quam latis. 

Plants up to 35 cm tall, covered with hairs 0.2-0.5 mm long. Caudex branched in upper 
part. Stems mostly several, erect, sulcate, loosely covered with subappressed to ascending 
white hairs, more densely hairy in young parts, glabrescent in old parts. Stipules 3-4 mm long, 
ovate to triangular, ± free from petiole, the lower ones highly up connate behind the stem, the 
upper ones only shortly cormate or free, glabrous. Leaves 2.5-6 cm long, the upper ones often 
sessile, petiole of lower leaves 1-3 cm long, like the rachis slender, sparsely covered with 
subappressed white hairs. Leaflets in 5-6 pairs, narrowly elliptic, 6-16 x 2-3 mm, narrowly 
rounded so subacute at the apex, on upper side glabrous, on underside in the youth densely 
appressed white hairy, later on loosely hairy. Peduncles 5-10 cm long, sparsely to loosely 
appressed white hairy, toward the raceme increasingly with ascending black hairs. Racemes 
ovoid, densely many-flowered, later on somewhat elongated, axis white and black hairy. 
Bracts whitish, linear, 2-3 mm long, white and black hairy. Flowers nearly sessile. Calyx 3- 
3.5 mm long, campanulate, with obliquely truncate orifice, densely appressed white and black 
hairy; teeth narrowly triangular, 1-1.5 mm long, white hairy on inner side. Petals color un- 
known. Standard 6-8 mm long; blade widely elliptic, 4-5 mm wide, widely emarginate at the 
apex, at the base abruptly narrowed into the short claw. Wings 5-6 mm long; blades narrowly 
oblong, somewhat dilated toward the truncate to widely rounded apex, 3-A x 1.3-2 mm; 
auricle 0.3-0.5 mm long, claw 2 mm long. Keel 4-4.5 mm long; blades obovate-oblong to 
obliquely obovate, with in upper third rectangularly curved lower edge and slightly convex 
upper edge, subacute at the apex, 2.5-2.8 x 1.5 mm; auricle minute, claw 2 mm long. Staminal 
tube straightly cut. Ovary very shortly stipitate, glabrous. Legumes nearly sessile, ca. 3.5 mm 
long, ca. 2 mm high, 5 mm broad, obtriangular, cuneate at the base, widely truncate at the apex 
with persistent style, widely flattened ventrally, narrowly and deeply grooved dorsally, with 
conspicuous transverse ridges, fully bilocular; valves thin, braun, with subappressed white and 
predonminant black hairs. Seeds singular in each locule, 2.5 x 2 mm, brovm, dull. 

Other specimens seen: 

India. Kashmir : Ladakh, Sassar, near border of Nubra, 1852, Thomson (K) - Tibet occ, Sassar 
pass, 15000', 13.8.1848, Thomson (K) - Zanskar, 14000', 1852, Thomson (K) - Zanskar, 
12000', 28.6.1848, Thomson (K). - Himachal-Pradesh : Lahul, Jaeschke 39 of 2nd collection (K). 

Astragalus villosulus Podlech, spec. nov. e sect. Brachycarpus 

Holotype: Afghanistan, Prov. Bamian, Koh-i-Baba, Kotal-i-Shatut zwischen Band-i-Amir 
und Panjao, Passhöhe, 3300 m, 10.7.1969, S. W.Breckle 2483 (MSB). 

Differt ab^. miserifloro Sirj. & Rech.f cui habitu similis tota planta pilis rigidis patentibus 
(nee apressis) obtecta, lamina alarum apicem truncatum vel emarginatum versus distincte dila- 
tata (nee anguste oblonga apice rotundata). 

Plants acaulescent, 8-14 cm tall, nearly all parts covered with 0.2-0.4 mm long, stiff, ± 
spreading hairs. Caudex shortly branched. Stipules brownish, triangular, ca. 2 mm long, nearly 
free from petiole, shortly connate behind the stem, ciliate at the margin or also hairy at the 
apex. Leaves 3-6 cm long; petiole as long or longer than the rachis, like the rachis loosely to 
rather densely white hairy. Leaflets in 4-5 pairs, remote, linear, 5-12 x 0.8-2 mm, on upper 



142 



side hairy only along the margins, on underside densely covered with spreading to ascending 
hairs, folded. Peduncles 5-12 cm long, slender, loosely to rather densely white, towards the 
top increasingly black hairy. Bracts whitish, narrowly triangular, 1-1.5 mm long, white or 
white and black hairy. Pedicels 0.3-0.5 mm long, hairy. Calyx 3 mm long, shortly campanu- 
late-tubular, covered with mostly basifixed white and black hairs up to 0.7 mm long; teeth 
narrowly triangular, ca. 1 mm long. Petals pale violet. Standard ca. 6 mm long, obovate, incised 
at the apex, gradually narrowed at the base, without distinct claw. Wings 4.5 mm long; blades 
spatulate, widened toward the truncate or slightly emarginate apex, 2.5 x 1.8 mm; auricle ca. 
0.7 mm long, claw 2 mm long. Keel 4 mm long; blades ± triangular, with widely curved lower 
edge and ± straight upper edge, 2.5 x 1.7 mm; claw 1.5 mm long. Staminal tube ± straightly 
cut. Ovary sessile, ovoid, hairy; style glabrous. Pod (unripe) sessile, 3 mm long, long white 
hairy. ■ 

Astragalus akhanii Podlech, spec. nov. e sect. Caprini 

Holotype: Iran, prov. Ham, 18 km SW of Salehabad, 17 km after Konjancham towards 
Shoor-Shirin, gypsum hills along Iran/Iraq frontier, 500-550 m, 1.5.1992, H. Akhani 8207 
(MSB). 

Differt ab A. leporino Boiss. & Noe indumento e pilis patentibus ad 1 mm longis (nee ad 3 
mm longis) consistente, stipulis petiole brevissime tantum adnatis ceterum liberis, leguminibus 
longiorubus latioribusque, ab A. gypsicola Maassoumi stipulis petiolo brevissime adnatis ce- 
terum liberis, foliis brevioribus foliolis minoribus acutis (nee rotundatis ad truncatis), racemis 
2-3-floris (nee 6-8-floris), leguminibus 1-25 (nee 30-40) mm longis. 

Plants acaulescent, with short, spreading, white hairs. Caudex up to 10 mm thick, at the 
apex shortly branched. Stipules whitish-membranous, ovate, longitudinally nerved, 10-12 mm 
long, the lower ones obtuse, the upper ones acute, adnate to the petiole for 2 mm, in upper 
part shortly ciliate at the margins, otherwise glabrous. Leaves 1 5-23 cm long; petiole 4-6 cm 
long, like the rachis striate-sulcate, rather densely covered with spreading hairs 0.3-1 mm long. 
Leaflets in 7-9 pairs, elliptic to widely elliptic, 10-22 x 8-17 mm, at the apex acutish or very 
shortly acuminate-mucronulate, on upper side loosely, on underside densely covered with 
ascendent to nearly spreading hairs 0.3-0.6 mm long. Peduncle 3-6 cm long, shortly 
spreadingly hairy. Racemes remotely 2-3 -flowered. Bracts whitish-hyaline, 6-8 mm longe, 
very narrowly triangular, hairy. Pedicels 5-8 mm long, spreadingly hairy. Calyx ca. 17 mm 
long, tubular, at the base slightly gibbous, at the orifice slighty obliquely truncate, straw- 
colored, rather densely covered with very thin, confiisely spreading hairs up to 0.7 mm longi; 
teeth subulates, (3-)4-6 mm long. Petals yellow. Standard 26-27 mm long; blade obovate, ca. 
8 mm wide, at the apex retuse, at the base subangularly narrowed into the long claw. Wings ca. 
25 mm long; blade narrowly oblong, slightly narrowed toward the obtusish apex, 10 x 2.3 mm; 
auricle ca. 1 mm long, claw 15 mm long. Keel 23 mm long; blade obliquely elliptic, with 
slightly but widely curved lower edge and nearly straight to slightly concave upper edge, at the 
apex obtuse, 9x4 mm: auricle 0.5 mm long, claw 15 mm long. Staminal-tube very obliquely 
cut. Ovary wit a stipe ca. 5 mm long, elliptic, white-hairy;, style glabrous. Legumen on a stipe 
4-5 mm long, elliptic, 16-25 mm long, 6-7 mm high and 13-15 mm wide, subcarinate 
ventrally, flattened to widely impressed, at the apex with a beak 3-4 mm long, incompletely 



143 



bilocular; valves coriaceous, pale brownish, densely covered with confusely spreading hairs 
0.3-1 mm long. Seeds 4x3 mm, pale brown, shiny. 

This new species belongs clearly to the A. caprinus-group (see Podlech 1988). It differs from 
A. leporinus Boiss. & Noe by the short spreading hairs up to 1 mm (not up to 3 mm) long, by 
stipules which are only very shortly adnate to the petiole and otherwise free and by longer 
and wider fruits. From A. gypsicola (wrongly gypsicolus) Maassoumi, from which 1 have seen 
no material it differs according to the description by stipules which are only shortly adnate to 
the petiole, by shorter leaves and smaller leaflets which are ± acute (not rounded or truncate) 
at the apex, by shorter peduncles of the only 2-3 (not 6-8)-flowered racems, the longer bracts, 
the standard which is retuse (not bilobed) at the apex and the distinctly shorter legumes. 

Astragalus antalyensis A.Duran & Podlech, spec. nov. e sect. Caprini 

Holotype: Turkey, [C3] Antalya, Akseki, Gukurköy yailasi, Yumrudikmeni mevki, 2200 
m, \6.1.\995, A.Duran 2792 (GAZI; Iso: MSB). 

Habitu, glabrietate et stylo piloso sub stigmate A. aegobromo Boiss. & Hohen, similis sed 
differt stipulis non cormatis, bracteis albidis angustis et leguminibus a latere compressis valvis 
tenuibus. 

Plantae subacaules, complete glabrae. Caules ad 4 cm longi. Stipulae albidae, 9-15 mm lon- 
gae, anguste ovatae, petiolo epr 3-7 mm adnatae. Folia 11-27 cm longa. Foliola 7-15-paria, 
ovata ad elliptica, 5-1 1 x 3-7 mm. Racemi laxi, pedunculo 1-2 cm longo suffulti. Bracteae 2-5 
mm longae. Pedicelli 7-10 mm longi. Calyx breviter tubulosus, ca. 10 mm longus, dentibus 2-3 
mm longis. Petala flava. Vexillum 20 mm longum, obovatum. Ovarium subsesssile, stylo sub 
stigmate linea e pilis brevibus consistente proviso. Legumina subsesssilie, oblique elliptica, 
10-15 mm longa, 5-5,5 mm alta, a latere compressa, ventro acute carinata, dorso leviter 
sulcata, fere complete bilocularia, valvis tenuibus. 

Plants ca. 30 cm tall, subacaulescent, completely glabrous. Caudex strongly divided, with 
many short, subterranean branches. Stems up to 4 cm long, angular-sulcate. Stipules whitish, 
9-15 mm long, narrowly ovate, acutish, adnate to the petiole for 3-7 mm. Leaves 1 1-27 cm 
long; petiole 3-9 cm long, like the rachis finely striate. Leaflets in 7-15 pairs, ovate to elliptic, 
widely rounded and minutely mucronulate at the apex, 5-11 x 3-7 mm. Peduncles 1-2 cm 
long. Racemes loosely several-flowered. Bracts whitish, narrowly triangular to linear-acute, 2- 
5 mm long. Pedicels 7-10 mm long, often tortuous. Calyx shortly tubular, ca. 10 mm long; 
teeth subulate, 2-3 mm long. Petals yellow. Standard 20 mm long; blades obovate, ca. 9 mm 
wide, widely emarginate at the apex, at the base gradually narrowed into the cuneate claw. 
Wings 1 8 mm long; blades narrowly oblong, rounded at the apex, 9x3 mm; auricle ca. 1 mm 
long, claw 9 mm long. Keel 14 mm long; blades triangular, with rectangularly curved lower edge 
and nearly straight upper edge, subobtuse at the apex, 6x4 mm; auricle short, claw 8 mm 
long. Staminal-tube straightly cut. Ovary subsessile, elliptic; style below the stigma with a line 
of short spreading hairlets. Pods subsessile, obliquely elliptic seen from the side, 10-15 mm 
long, 5-5.5 mm high and ca. 3 mm wide, strongly carinate ventrally, slightly grooved dorsally, 
attenuate at the apex into beak 2-3 mm long; valves straw-colored, thin. Fruits nearly 
completely bilocular with 3-4 seeds in each locule. 



144 



In habit, bareness and the pilose style below the stigma near to A. aegobromus Boiss. & 
Hohen., but differing strongly by the stipules not connate one to another, the whitish, narrow 
bracts and the laterally compressed, thin-walled fruits. 




Fig. 1 : Habit oi Astragalus antalyensis (holotype). Scale bar: 3 cm. 



145 



Astragalus avajensis Podlech, spec. nov. e sect. Caprini 

Holotype: Iran, prov. Hamadan, 10 km from Razan to Avaj, on the neck, 2200-2300 m, 
10.7.1994, Chehregani & S. Zarre 17893 (MSB). 

Differt ab A. kirpicznikovii Grossh. indumento e puis 0,1-1,5 mm longo (nee 1^ mm 
longo) consistente, pedicellis ca. 1 mm (nee 4-6 mm) longis, calycis tubo aequaliter piloso (nee 
basi glabro vel sparsissime, apicem versus densiuscule piloso), legumine angustiore, ca. 5 mm 
alto latoque (nee 6-10 mm alto et 8-10 mm lato), dorso distincte sulcato (nee late applanato). 

Plants 12-20 cm tall, subacaulescent, covered with thin, spreading white hairs 0.1-1.5 mm 
long. Caudex divided, with many short, subterranean branches. Stems if present up to 4 cm 
long, angular-sulcate, glabrous. Stipules whitish-membranous, ovate-triangular, 6-15 mm long, 
very shortly adnate to the petiole, glabrous or sparsely ciliate at the margins. Leaves 10-17 cm 
long; petiole 2-4 cm long, like the rachis loosely spreadingly hairy. Leaflets in 14-20 pairs, 
narrowly elliptic, rounded to truncate and minutely mucronulate at the apex, 8-15 x3-6 mm, 
glabrous on upper side, on lower side sparsely to loosely covered with short, flexuose, 
ascending to spreading hairs. Peduncles 1-2 cm long, rather densely covered with hairs up to 
1.5 mm long. Racemes densely 4-6-flowered. Bracts whitish-membranous, elliptic, 5-8 mm 
long, white-ciliate. Pedicels ca. 1 mm long. Calyx straw-colored, tubular, ca. 15 mm long, 
loosely to rather densely covered with spreading, soft white hairs up to 1 .5 mm long; teeth 
narrowly triangular, ca. 6 mm long. Petals yellow. Standard 25-26 mm long; blades obovate, 
ca. 9 mm wide, widely emarginate at the apex, at the base subabruptly narrowed into the 
cuneate claw. Wings 16-17 mm long; blades narrowly oblong, slightly spathulate-dilated 
towards the rounded apex, 9 x 3-3.5 mm; auricle ca. 1 mm long, claw 7 mm long. Keel 14-15 
mm long; blades elliptic, with curved lower and upper edge, straight, acute at the apex, 6-7 x 
3.5^ mm; auricle short, claw 8 mm long. Staminal-tube slightly obliquely cut. Pods with a 
stipe 1-2 mm long, obliquely oblong, with straight dorsal line and slightly curved ventral line, 
ca. 17 mm long, 5-6 mm high and ca. 5 mm wide, strongly carinate ventrally, deeply grooved 
dorsally, attenuate at the apex into a curved beak ca. 1 mm long; valves straw-colored, thin, 
loosely covered with fine, flexuose, tousled, ± spreading hairs. Fruits semibilocular with 2-3 
seeds in each locule, opening at the ventral suture. Seeds 5x4 mm, olive-green, smooth. 

Astragalus diversipilosus Podlech, spec. nov. e sect. Caprini subsect. Purpurascentes 

Holotype: Iran, Kurdistan, ad versuras 33 km NW Divandarreh versus Saqqez, 2100 m, 
29.6.1974, Rechinger 486 J Ob (M). 

Plantae acaules, albipilosae. Stipulae ca. 10 mm longae, per 2-3 mm petiolo adnatae, inferio- 
res glabrae, superiores ciliatae. Folia 7-1 1 cm longa; petiolo 1,5-2,5 cm longo, sicut rachis pilis 
patentibus 0,5-1,5 mm longis obtecto. Foliola 19-25 paria, subcontigua, supra laxe ad 
densiuscule pilis brevissimis obtecta, interdum glabrescentia, margine et subtus dense pilis 
brevissimis et laxe pilis ad 1 mm longis subpatentibus obtecta, saepissime complicata, anguste 
rubri-marginata. Racemi pedunculo 0,5-1 cm longo suffulti, 1-2 flori. Bracteae ca. 2 mm lon- 
gae, ciliatae. Pedicelli 5-6 mm longi. Calyx 12-14 mm longus, patentipilosus, dentibus trian- 
gularibus ca 3 mm longis. Corolla glabra, flava. Vexillum ca. 25 x 9 mm, lamina, triangulari- 
obovata, apice late retusi-rotundata, incisa, basi in unguem lamina ± aequilongum angustata. 
Alae ca. 23 mm longae; lamina triangulari-obovata, apice latissima. Carina 23-24 mm longa; 



146 



lamina in margine superiore dentibus minutis proviso. Legumen stipite 8-10 mm longo glabro 
suffultum, ellipsoideum, 13-18 mm longum, 7-8 mm altum et 3^ mm latum, semibiloculare; 
valvis laxe pilis brevissimis et distincte longioribus ad 1,5 mm longis obtectum. 

Plants acaulescent, white hairy. Caudex with a pluricipital root-crown, branches short, 
densely covered with remnants of old stipules. Stipules whitish-membranous, ovate or 
triangular, ca. 10 mm long, adnate to the petiole for 2-3 mm, otherwise free, the lower ones 
glabrous, the upper ones ± densely shortly ciliate. Leaves 7-11 cm long; petiole 1.5-2.5 cm 
long, like the rachis loosely to rather densely covered with thin but subrigid, spreading hairs 
0.3-1 .5 mm long. Leaflets in 19-25 pairs, mostly close together, 5-7 x 2-4 mm, on upper side 
loosely to rather densely covered with very short spreading hairs, sometimes glabrescent, on 
underside and at the margins loosely to rather densely covered with spreading very short hairs 
and loosely with hairs up to 1 mm long, narrowly reddish-margined, mostly folded. Peduncles 
0.5-1 cm long, spreadingly hairy. Racemes 1-2 flowered. Bracts whitish-membranous, ovate 
to narrowly triangular, ca. 2 mm long, ciliate. Pedicels 5-6 mm long, spreadingly hairy. Calyx 
12-14 mm long, tubular, loosely covered with thin, spreading hairs; teeth narrowly triangular 
to triangular, ca. 3 mm long. Petals glabrous, yellow. Standard ca. 25 mm long; blade triangular- 
obovate, ca. 9 mm wide, at the apex widely retuse to rounded, slightly emarginate, at the bae 
cuneately narrowed into the long claw. Wings ca. 23 mm long; blades triangular-obovate, 
widest at the retuse apex, 8 x 3.5 mm, above the auricle strongly constricted; auricle ca. 2 mm 
long, claw ca. 15 mm long. Keel 23-24 mm long; blades obliquely elliptic with in the middle 
gibbously curved lower edge and sigmoid-curved upper edge provided with minute teeth, 
toward the obtuse apex slightly narrowed, 10 x 4.5 mm; auricle short, claw 13-14 mm long. 
Staminal-tube straightly cut. Legumes on a glabrous stipe 8-10 mm long, elliptic, 13-18 mm 
long, 7-8 mm high and 3-4 mm wide, carinate ventrally, narrowly rounded dorsally, at the 
apex contracted into the beak 2-3 mm long, semibiloculare; valves thin, straw-colored, loosely 
covered with very short and with distinctly longer, spreading hairs up to 1.5 mm long. 

Astragalus glaberrimus Podlech, spec. nov. e sect. Caprini 

Holotype: Iran, Prov. Kerman, hills W of Deh Bakri, 2000-2500 m, 30.4.1975, BS. Parris 
75.215 (K). 

Differt ab A. zagrosicus Boiss. & Hausskn. folioUs latis, 4-8 x 3,5-8 (nee 8-18 x 4-11) 
mm, calyce 15-17 (nee 9-12) mm longo, vexillo 23-26 (nee 19-21) mm longo, ovario dense et 
longe albipiloso (nee glabro). v . 

Plants 15-17 cm tall, nearly acaulescent, with exception of the ovary completely glabrous. 
Stipules whitish-membranous, oblong, obtuse, 6-10 mm long, adnate to the petiole for ca. 2 
mm. Leaves 10-20 cm long; petiole 3-10 cm long, like the rachis slightly striate-sulcate. Leaf- 
lets in 11-16 pairs, widely elliptic to widely obovate, at the apex rounded to distinctly retuse, 
4-8 X 3.5-8 mm. Peduncle up to 1.5 cm long. Raceme 1-4-flowered. Bracts whitish-mem- 
branous, 5-7 mm longe, linear-acute. Pedicels 4.6 mm long. Calyx 15-17 mm long, tubular, 
straw-colored; teeth narrowly triangular, 4-5 mm long. Petals yellow. Standard 23-26 mm 
long; blade oblong, ca. 6 mm wide, at the apex emarginate, at the base long narrowed. Wings 
20-23 mm longe; blades narrowly oblong, obtuse at the apex, 8-9 x 2 mm; auricle oblong, ca. 
1.5 mm long. Keel 19-22 mm long; blades obovate-triangular, with widely curved lower edge 



147 



and sigmoid-concave upper edge, at the apex subobtuse, 6-7 x 3.5 mm; auricle wide, 0.3 mm 
long. Ovary with a stipe ca. 1 mm long, narrowly elliptic, densely white-hairy; style glabrous 
under the stigma. Legumes unkonwn. 

Astragalus malatyaensis Podlech, spec. nov. e sect. Caprini 

Holotype: Türkei, Prov. Malatya, 8 km E Darende gegen Malatya, 1080 m, Gipsboden, 
4.6.1987, M. Nydegger 42617 (MSB; Iso: BASBG). 

Differt ab A. ovino Boiss. stipulis petiolo brevissime, per 1-2 (nee 5-7) mm adnatae, foliis 
6-9 (nee 10-1 2-1 6)-jugis, lamina vexilli leviter oblongi-pandurata (nee obovata), auricula 
alarum ca. 3 mm (nee 1 mm) longa, ovario legumineque distincte stipitato (nee sessile). 

Plants glabrous, acaulescent or nearly so, only at the base of the leaflet, the petioles and the 
margins of the stipules with few minute, globose hairs. Stipules whitish-membranous, 
obliquely inserted at the stem, the lower ones oblong, obtuse, 10-12 mm long, the upper ones 
narrowly triangular, 15-20 mm long, all adnete to the petiole for 1-2 mm. Leaves 13-20 cm 
long; petiole 3-6 cm long, like the rachis slightly striate-sulcate, at the base up to 2 mm thick. 
Leaflets in 6-9 pairs, elliptic, 10-28 x 5-15 mm, at the apex rounded to acutish. Peduncle 1-5 
cm long. Racemes 3-4-flowered. Bracts 7-12 mm long, whitish-membranous or greenish at the 
apex, linear-acute. Pedicels 5-7 mm long. Calyx 15-17 mm long, tubulose, slightly obliquely 
cut at the orifice, straw-colored, toward the apex greenish; teeth narrowly triangular, 3.5-4 mm 
long. Petals yellow. Standard 25-28 mm long; blade recurved in the middle, slightly oblong- 
pandurate, ca. 10 mm wide, at the apex emarginate, at the base shortly cuneately narrowed. 
Wings 24-27 mm long; blades narrolwy oblong, at the apex acutish, 9-10 x 3 mm; auricle 
oblong-subacute, ca. 3 mm long, claw 15-17 mm long. Keel 22-24 mm long; blades obovate- 
triangular, with widely rectangularly curved lower edge and sigmoid-concave upper edge, at the 
apex obtusish, 7-8 x 4 mm; auricle 1 .5 mm long. Ovary with a stipe 2-3 mm long, style below 
the stigma glabrous. Legumes with a short, narrowly cuneate stipe, narrowly elliptic, ca. 40 
mm long, 13-15 mm high and 9-10 mm wide, narrowly rounded with thickish nerve ventrally 
and dorsally, at the apex shortly acuminate, incompletely bilocular; valves thick-coriaceous, 
strongly rugose. Seeds reniform, 6x5 mm, smooth. 

Astragalus neyshaburensis Podlech, spec. nov. e sect. Cytisodes 

Holotype: Iran, prov. Khorasan, on road from Neyshabur to Kashmar, 1 100 m, 14.5.1977, 
Dini & Arazm 33163 (W; Iso: MSB). 

Differt ab A. pseudorhacodes Gontsch. (sect. Macrocystodes), cui ob folias 2-jugas similis, 
pedunculo et rachide racemae pilis semipatulis (nee appressis), calyce breviore in fructo non 
inflato, leguminibus longioribus fere unilocularibus (nee bilocularibus) apice rostro ad 1 5 (nee 
2-3) mm longo provisis. 

Plants 20-25 cm tall, suffruticose, hairy. Stem branched at the base, 8-10 cm long, densely 
covered with symmetrically medifixed, appressed white, at the base of stipules also black 
hairs 0.6-0.8 mm long. Stipules whitish-membranous, from wide base triangularly acuminate, 
4-6 mm long, adnate to the petiole for 1-2 mm, behind the stem shortly vaginate-connate, 
densely appressed hairy. Leaves (4-) 10-1 2 cm long; petiole 2-5 cm long, like the rachis rigid. 



148 



hairy like the stem. Leaflets in 2 pairs, elliptic to obovate, (15-)20-30 x (8-) 12-23 mm, at the 
apex shortly acuminate to widely rounded, on both sides loosely to rather densely covered 
with appressed, mostly symmetrically medifixed white hairs 0.5-1.2 mm long. Peduncles 
rigid, 3-4 cm long, angular, loosely covered with ascending, often flexuose, predominantly 
black hairs 0.2-0.6 mm long. Raceme (in fruiting time) up to 8 cm long, with 8-12 fruits; axis 
with ascending, predominantly black hairs. Bracts whitish-membranous, narrowly triangular, 
4-5 mm long, mainly black hairy. Pedicels (in fruit) thick, ca. 3 mm long, black hairy. Calyx 
10-12 mm long, tubular, rather densely covered with conflisely ascending, asymmetrically to 
symmetrically bifurcate, flexuose, predominantly black hairs 0.1-0.6 mm long; teeth subulate, 
3-5 mm long. Petals unknown. Legumes obliquely erect to spreading, linear, slightly upcurved, 
40-50 mm long, 4 mm high and 3 mm wide, substipitately narrowed at the base, long acumi- 
nate at the apex into a slender beak 10-15 mm long, carinate ventrally, rounded to slightly 
grooved dorsally, nearly unilocular, with a septum up to 0.3 mm wide; valves rather densely 
covered with short, confusely ascending, flexuose black hairs up to 0.5 mm long and with 
ascending, strongly asymmetrically, straight, white hairs up to 1 mm long. ._^ 

This new species resembles A. pseudorhacodes Gontsch. from sect. Macrocystodes in 
having only 2 pairs of leaflets. It is different from the latter species by peduncles and rachis of 
the raceme covered with ascending (not appressed) hairs, by the shorter calyx not inflated in 
fruit, by the much longer und narrower fruit with white and black hairs (not villous with long 
stiff white hairs) and with a beak up to 15 (not 2-3) mm long and which is nearly unilocular 
(not bilocular). Most probably it is the same what Maassoumi (1998) has reported as A. 
pseudorhacodes. 

Astragalus fallacinus Podlech, spec, nov e sect. Dissitiflori 

Holotype: Türkei, A4 Cankiri, Kalecik - Cankiri, E Hisarcikkay, 850 m, Gipshügel, 5.6. 
1993, M Nydegger 47400 (MSB; Iso: BASBG). 

Differt ab A. aucheri Boiss. racemis tempore fructificationis elongatis, calyce laxe pilis albis 
subappressis et nigris brevioribus appressis obtecto (nee dense albivilloso), leguminibus bre- 
vioribus subcurvatis, ab A. nigrifructo Podlech & Ayta? leguminibus 16-20 (nee 40-50) mm 
longis fere mere albipilosis. 

Plants suffruticose, 15-20 cm tall. Stems of the year 8-10 cm long, densely covered with 
appressed white hairs 0.4-0.6 mm long. Stipules triangular, ca. 3 mm long, adnate to the 
petiole for ca. 1 mm, densely white-hairy, sometimes also with some black hairs. Leaves 4-7 
cm long; petiole 0,5-2,5 cm long, like the rachis rather densely hairy. Leaflets in 3^ pairs, 
linear, 8-18 x 1,5-2 mm, at the apex obtuse or acutish, on upper side loosely to rather 
densely, on underside more densely covered with white hairs ca. 1 mm long. Peduncles 6-12 
cm long, covered with white, toward the raceme also with black hairs. Racemes at first 
densely, later on loosely 4-7-1 4-flowered. Bracts 2-3 mm long, narrowly triangular, white 
and black hairy. Pedicels 1-2 mm long. Calyx 11-12(-14) mm long, tubular, at the base 
slightly gibbous, obliquely cut at the orifice, loosely covered with short, appressed, ± 
symmetrically medifixed black hairs and with subappressed to ascending, strongly asymmetri- 
cally medifixed white hairs ca. 1 mm long; teeth subulate, l-2(-4) mm long. Petals yellow. 
Standard 20-22 mm long; blade obovate, ca. 8 mm wide, at the apex slightly emarginate, at the 
base subangularly narrowed into the cuneate claw. Wings 20-21 mm long; blades obovate, at 



149 



the apex rounded, 6-7 x 3 mm; auricle curved, ca. 0,5 mm long, claw 13-14 mm long. Keel ca. 
1 6 mm long, blades obliquely obovate, with in upper part widely rectangularly curved lower 
edge and nearly straight upper edge, acutish at the apex, 5x3 mm; auricle 0,5 mm long, acute, 
claw ca. 1 1 mm long. Staminal-tube straightly cut. Ovary subsessile, linear, style long, 
glabrous. Legumes sessile, linear, 16-20 mm long, ca. 2 mm in diameter, slightly but distinctly 
curved, obstusely carinate ventrally, grooved dorsally, at the apex long acuminate-curved, with 
the persistent style, fully bilocular; valves covered with appressed white, sometimes also with 
few black hairs. 

Other specimen seen: 

Türkei. A5 Sinop : Boyabat - Kastemonu, 21 km W Boyabat, 280 m, Serpentin, 30.6.1987, 
Nydegger 42984 (MSB). 

Astragalus cedreticola A.Duran & Podlech, spec, no v. e sect. Hypoglottidei 

Holotype: Turkey. C3 Antalya, Akseki, Pinarbasi köyü, ^ukurköy yaylasi, Yumrudikmeni 
mevki, 2200 m, 4.7.1996, A.Duran 4071 (GAZI; Iso: MSB). 

Differt ab A. ovato DC. pilis (fructu excepto) ad 1 mm (nee 1,5-3,5 mm) longis obtectus, 
stipulis a petiolo liberis (nee per 1-3 mm adnatis), bracteis 5-7 mm (nee 8-15 mm) longis, 
calyce 11-13 mm longo dentibus 3,5—4 mm longis (nee 13-15 mm longo dentibus 5-7 mm 
longis), petalis caerulei-violaceis (nee flavis), vexillo ca. 18x7 mm, rhomboidei-elliptico (nee 
20-23 X 7-8 mm, oblongo), legumine pilis tenuibus flexuosis ad 3 mm longis et insuper pilis 
brevissimis obtecto (nee pilis longis tantum, in tuberculis insidentibus obtecto). 

Plants 20-30 cm tall, branched at the base, covered with thin soft, tousled, spreading hairs 
up to 1 mm long. Stems prostrate or ascending at the base or erect, rather densely covered 
with very short and up to 1 mm long white, below the nodes also black hairs. Stipules green, 
4-8 mm long, ovate, acute to shortly acuminate, free from the petiole, the lower ones vaginate- 
connate behind the stem often high up, the upper ones sometimes nearly free, all loosely 
covered with white and black hairs. Leaves 5-6 cm long; upper leaves with short petiole to 
subsessile, petiole of lower leaves 1.5-2.5 cm long, like the rachis sulcate at upper side, rather 
densely furnished with white hairs up to 0.5 mm long. Leaflets in 10-14 pairs, elliptic to 
obovate, 4-9 x 2-4 mm, widely retuse to widely emarginate at the apex, on upper side 
covered loosely to rather densely, on underside more densely with nearly spreading white 
hairs. Peduncles 4-10 cm long, finely striate, hairy like the stem. Racemes capitate to ovoid, 
densely 6-10-flowered, in fruit slightly elongated. Bracts green, narrowly triangular, 5-7 mm 
long, white and black hairy. Pedicels 0.5-1 mm long, hairy. Calyx 11-13 mm long, tubular to 
slightly inflated after anthesis, covered on the upper (adaxial) side predominantly with white, 
on the lower (abaxial) side with black spreading hairs; teeth subulate to linear-acute, 3.5-4 mm 
long, nearly glabrous on inner side. Petals bluish-violet. Standard 18 mm long; blade ca. 7 mm 
wide, rhombic-elliptic, widely emarginate at the apex, gradually narrowed into the cuneate 
claw at the base. Wings 14 mm long; blades nearly oblong, slightly gradually dilated toward the 
obliquely retuse apex, 7 x 2.5 mm; auricle wide, 0.5 mm long, claw 7 mm long. Keel 12 mm 
long; blades obliquely obovate, with in upper part widely curved lower edge and slightly 
curved upper edge, obtuse at the apex, 5x3 mm; auricle short, claw 7 mm long. Staminal-tube 
straightly cut. Ovary very shortly stipitate, elliptic, white-villous; style glabrous. Pods 



150 



sessile, obliquely elliptic, with nearly straight ventral (upper) edge and widely curved dorsal 
(lower) edge, 10-12 mm long, 3.5 mm high, 2.5 mm wide, carinate ventrally, deeply grooved 
dorsally, attenuate at the apex into a short beak; valves thin, densely lanate, with ± spreading, 
thin, soft, tousled white hairs up to 3 mm long and with very short hairs. Fruit nearly fully 
biloculare, with ca. 5 seeds in each locule. 

Other specimens seen: 

Turkey. C3 Antalya : Akseki, Pinarbasi köyü, Softalar yaylasi, 1800 m, Cedrus libani forest, 
4.6.1996, A.Duran 3900 (GAZl, MSB) - Akseki, Pinarbasi köyü, Cataloluk mevki, 1400 m, 
11.6.1995,/4. DMra«27/2(GAZI,MSB). 




Fig. 2: Astragalus cedreticola (holotype). Scale bar: 2 cm. 



151 



Astragalus brevidentatus Podlech, spec. nov. e sect. Incani 

Holotype: Türkei, B4 Ankara, zwischen Ankara und Aksaray, 10 km N Serefliko9hisar, 
970 m, steiniger Abhang, 1.4.1986, M. Nydegger 41023 (MSB; Iso: BASBG). 

Plantae ca. 5 cm altae, acaules, puis 0.4-0,7 rmn longis appressis obtectae. Stipulae 3^ mm 
longae, pilosae. Folia 1,5-3,5 cm longa, petiolo 0,5-1 cm longo, sicut rachis dense piloso. Fo- 
liola4-7 paria, anguste elliptica, acutiuscula, 2-6 x 0,4-1,6 mm, utrimque densiuscule pilosa. 
Racemi pedunculo ad 1,5 cm longo albipiloso suffulti, 2-5-flori. Bracteae 1,5-2 mm longae. 
Bracteolae minutae. Calyx ca. 12 mm longus, tubulosus, laxe vel apicem versus dense nigripilo- 
sus, dentibus subulatis 0,5-2 mm longis. Vexillum ca. 25 mm longum, lamina rhomboidei-obo- 
vata, ca. 10 mm lata. Alae ca. 21 mm longae. Carina ca. 20 mm longa. Ovarium stipite 1-2 mm 
longo suffultum, brevissime albipilosum, stylo glabro. Legumen ignotum. 

Plants ca. 5 cm tall, acaulescent, covered with ± symmetrically medifixed, appressed, in 
vegetative parts merely white hairs 0.4-0.7 mm long. Caudex strongly branched. Stipules 
straw-colored, 3^ mm long, ovate, acute, shortly adnate to the petiole, hairy. Leaves 1.5-3.5 
cm long, petiole 0.5-1 cm long, like the rachis densely hairy. Leaflets in 4-7 pairs, narrowly 
elliptic, acutish at the apex, 2-6 x 0.4-1.6 mm, on both sides rather densely hairy. Peduncle 
up to 1.5 cm long, densely white hairy. Raceme 2-5-flowered. Bracts whitish, 1.5-2 mm long, 
narrowly ovate, at the margins black ciliate. Pedicels 0.5 mm long. Bracteoles minute, linear, at 
the base of calyx. Calyx ca. 1 2 mm long, tubular, loosely or toward the teeth densely black 
hairy; teeth subulate, 0.5-2 mm long. Petals lilac? Standard ca. 25 mm long; blade rhombic- 
obovate, ca. 10 mm wide, at the apex narrowly rounded, at the base widely cuneately 
narrowed. Wings ca. 2 1 mm long; blades narrowly oblong, slightly curved, at the apex acutish 
and sometimes slightly obliquely emarginate, 9 x 2.5 mm; auricle ca. 1 mm long, claw ca. 12 
mm long. Keel ca. 20 mm long; blades obliquely obovate, with widely curved lower edge and 
slightly concave upper edge, at the apex obtusish, 8x3 mm; auricle short, claw ca. 12 mm 
long. Staminal-tube irregularely cut. Ovary with a stipe 1-2 mm long, very shortly white 
hairy; style glabrous. Legumes unknown. 

Astragalus longis ubulatus Podlech, spec. nov. e sect. Incani 

Holotype: Türkei, B7 Ercincan, 8 km S Kemaliye gegen Arabkir, bewaldetes Tal, 1040 m, 
22.6.1992, Nydegger 46374 (MSB; Iso: BASBG). 

Differt ab .4. pendula DC. calyce 16-18 mm (nee 9-1 1mm) longo, dentibus ejus 7-8.5 mm 
(nee 1-5 mm) longis, corolla calyce vix superante (nee ca. duplo longiore), legumine curvato 
20-22 mm longo (nee ± stricto, 12-18 mm longo). 

Plantae ad 45 cm altae, pilis 0,5-0,8 mm longis, ± symmetrice medifixis appressis obtectae. 
Stipulae 8-9 mm longae, basi per 3-^ mm petiolo adnatae, pilosae. Folia 18-22 cm longa. Fo- 
liola 7-10 paria, elliptica, 10-22 x 6-12 mm, utrimque pilosa. Racemi pedunculo 20-28 cm 
longo suffulti, dense multiflori, demum valde elongati. Bracteae 3-4 mm longae. Bracteolae 
minutae. Calyx 16-18 mm longus, tubulosus, albi- et nigri-pilosus, dentibus subulatis 7-8,5 
mm longis. Corolla calyce vix superans. Vexillum 1 8-20 mm longum, ellipticum, 6-7 mm la- 
tum. Alae 16-17 mm longae. Carina ca. 15 mm longa. Ovarium subsessile, albipilosum. Legu- 



152 



men pendulum, sessile, lineare, curvatum, 20-22 mm longum, ca. 3 mm altum et 2,5 mm latum, 
apice rostro recto ca. 2 mm longo, densiuscule appresse pilosum. 

Plants up to 45 cm tall, acaulescent, covered with appressed, ± symmetrically medifixed, in 
vegerative parts merely white hairs 0.5-0.8 mm long. Stipules 8-9 mm long, triangular, adnate 
to the petiole for 3-4 mm, ± densely hairy. Leaves 18-22 cm long; petiole 3-9 cm long, like 
the rachis densely hairy. Leaflets in 7-10 pairs, flat, elliptic, 10-22 x 6-12 mm, at the apex 
acutish or rarely obtuse, on both sides loosely to densely hairy. Peduncle 20-28 cm long, 
erect, slightly sulcate, loosely white hairy. Raceme at first densely many-flowered, later on 
elongated and up to 18 cm long, axis sulcate, sparsely white hairy. Bracts whitish-mem- 
branous, 3-4 mm long, linear-acuminate, white and black hairy. Flowers at first erect, later 
nearly spreading. Pedicels 1-2 mm long, hairy. Bracteoles minute, scarcely 1 mm long, at the 
base of calyx. Calyx 16-18 mm long, tubular, straw-colored, with 5 darker nerves, densely 
covered with white and also sparsely with black hairs; teeth subulate, 7-8.5 mm long. Petals 
pale violet ?, scarcely longer than the calyx. Standard 18-20 mm long, in upper part distinctly 
recurved; blade elliptic, 6-7 mm wide, at the apex narrowly rounded, at the base subangularly 
narrowed into the cuneate claw. Wings 16-17 mm long; blades linear, at the apex rounded, ca. 
7 X 1.5 mm; auricle up to 1 mm long, claw 9-10 mm long. Keel ca. 15 mm long; blades oblong- 
curved, with widely curved lower edge and concave upper edge, at the apex acutish, ca. 6 x 2 
mm; auricle acute, scarcely 1 mm long, claw ca. 9 mm long. Staminal-tube straightly cut. Ovary 
nearly sessile, narrowly elliptic, densely white hairy; style glabrous. Legumes pendulous, 
sessile, linear, curved, 20-22 mm long, ca. 3 mm high and 2.5 mm wide, carinate ventrally, 
slightly sulcate dorsally, at the apex with a straight beak ca. 2 mm long, rather densely 
appressed hairy. 

A. mercklinii Boiss. & Buhse subsp./ar^ic«* Podlech, subspec. nov. (sect. Incani) 

Holotype: Iran, Prov. Pars, inter Chiras et Persepolis, 800-1000 m, 2.5.1956, F. Schmid 
5514 (M; Iso: W). 

Differt ab subspeciei mercklinii indumento e pilis longioribus (0,6-1 mm versus 0,3-0,5 
mm longis), caudice saepissime simplice, irramoso, indistincte foliolis in (2-)4-6(-10) nee in 
(2-) 5-8(-12) paribus, ovatis, apice acutis et saepissime distincte acuminatis (nee ellipticis, 
ovatis vel late ovatis, apice late obtusis vel raro vix acutiuscuiis), bracteolis deficientibus vel 
raro evolutis, minutissimis, ad 0,5 mm longis, leguminibus longioribus ad 45 mm longis (nee ad 
30 mm longis). 

Differs from subsp. mercklinii by the indumentum of hairs 0.6-1 mm (not 0.3-0.5 mm) 
long, by fewer leaflets which are ovate, at the apex acute and mostly distinctly acuminate (not 
elliptic, at the apex widely obtuse), by the bracteoles absent or if rarely present, than minute, 
up to 0.5 mm long, and by the longer legumes up to 45 mm long (not up to 30 mm long). 

Other specimens seen: 

Iran. Lorestan : Azna, Shuhturun-kuh foothills, 1830 m, 6.5.2962, Furse 1775 (W). - Pars : Niris, 
28.3.1940, Koelz 14689 (US, W); 58 km W of Shiraz, 1600 m, 16.4.1964, Grant 15611 (W); SE 
Schiraz, 1 520 m, 30.5. 1 96 1 , Pabot 12446-E ( W). 



153 



Astragalus nevshehiricus Podlech, spec. nov. e sect. Incani 

Holotype: Türkei, B5 Nev'ehir, 2 km E Nev'ehir gegen Ürgüp, 1310 m, Hohlweg in einer 
Baumkultur, 7.6.1992, M. Nydegger 46186 (MSB; Iso: BASBG). 

Differt ab A. clavato DC. folioiis 3-5 (nee ca. 8) paribus, corolla violacea (nee viridiusculi- 
flava), legumine lineari, distincte curvato, piloso (nee lineari-clavato, subrecto, glabro). 

Plantae ca. 25 cm altae, pilis 0,3-0,6 mm longis obtectae. Stipulae 7-9 mm longae, basi per 
2 mm petiolo adnatae, laxe pilosae. Folia 7-16 cm longa, petiolo 3,5-8 cm longo. Foliola 3-5 
paria, remota, elliptica ad obovata, 10-22 x 6-12 mm, utrimque pilosa. Racemi laxe multiflori, 
tempore fructificationis valde elongati, pedunculo erecto 7-9 cm longo suffulti. Bracteae 2-3 
mm longae. Flores erecti. Bracteolae minutae. Calyx 9-12 mm longus, tubulosus, laxe nigripilo- 
sus, dentibus 2,5—4 mm longis. Corolla violacea. Vexillum 15-17 mm longum, 6-7 mm lata, 
elliptica, apicem versus lingulatim contracta. Alae 13-14 mm longae. Carina 12-13 mm longa. 
Legumina erecta vel subpatentia, stipite 2-3 mm longo suffulta, linearia, distincte ad semi- 
circulariter curvata, 22-25 mm longa et ca. 2 mm diametro, apice longe acuminate, dense ap- 
presse albipilosa. 

Plants ca. 25 cm tall, acaulescent, covered with appressed, ± symmetrically medifixed, in 
vegetative parts merely white hairs 0.3-0.6 mm long. Stipules 7-9 mm long, narrowly oblong- 
triangular, obtuse, adnate to the petiole for 2 mm, loosely hairy. Leaves 7-16 cm long; petiole 
3.5-8 cm long, like the rachis slender, loosely hairy. Leaflets in 3-5 pairs, elliptic to obovate, 
10-22 X 6-12 mm, at the apex acute to acutish or rarely rounded and minutely mucronulate, at 
both sides loosely to rather densely hairy. Peduncle erect, 7-9 cm long, sparsely to loosely 
covered with white, toward the raceme also with black hairs. Raceme loosely many-flowered, 
at fruiting time strongly elongated and up to 15 cm long. Bracts whitish-membranous, 2-3 mm 
long, narrowly triangular, sparsely black hairy. Flowers erect. Pedicels 2-4 mm long, black 
hairy. Bracteoles whitish, minute, scarcely 0.5 mm long, at the base of calyx. Calyx 9-12 mm 
long, tubular, loosely black hairy; teeth linear to subulate, 2.5-4 mm long. Petals violet. Stan- 
dard 15-17 mm long; blade 6-7 mm wide, elliptic, toward the apex subabruptly narrowed into 
a wide, tongue-like appendix, slightly emarginate, at the base widely cuneately narrowed. 
Wings 13-14 mm long; blades narrowly oblong, at the apex acutish, obliquely incised, in upper 
margin distinctly gibbous above the auricle, 6-7 x 2 mm; auricle ca. 1 mm long, claw 6-7 mm 
long. Keel 12-13 mm long; blades obliquely obovate, with widely curved lower edge and 
nearly straight upper edge, at the apex obtusish, ca. 5 x 2 mm; auricle wide, very short, claw 
7-8 mm long. Staminal-tube straightly cut. Ovary nearly sessile, sparsely white hairy; style 
glabrous. Legumes erect or ascending, with a stipe 2-3 mm long, linear, distinctly up to semi- 
circularly curved, 22-25 mm long and ca. 2 mm in diameter, carinate ventrally, sulcate dorsally, 
at the apex long acuminate, densely appressed white hairy. 

Astragalus polhillii Podlech, spec, nov e sect. Incani 

Holotype: Turkey, B9 Van: 18 km from Van to Er9ek, 2000 m, 5.6.1966, P. Davis 44366 
(K). 

Plantae 1 7-40 cm altae, pilis subappressis ad semipatentibus, saepe confuse flexuosis 0,2- 
1 mm longis obtectae. Stipulae 8-12 mm longae, petiole adnatae per 5 mm, basi pilosae. Folia 



154 



10-24 cm longa, petiolo 2,5-9 cm longo sicut rachis laxe piloso. Foliola 6-12 paria, lineari-el- 
liptica, 15-35 x 2-4 mm, apice acuminata, supra glabra vel sparse pilosa, subtus pilis confuse 
semipatenter ad 1 mm longis obtecta. Racemi longi, laxe multi-flori, pedunculo 12-20 cm longo 
suffulti. Bracteae 4-5 mm longae. Flores erecti. Bracteolae ca. 1 mm longae. Calyx 10-1 1 mm 
longus, tubulosus, albi-nigri-pilosus, dentibus 1-3 mm longis. Corolla dilute sulfurea. Vexillum 
20-21 mm longum, eliipticum, ca. 9 mm latum. Alae ca. 19 mm longae. Carina 16 mm longa. 
Ovarium stipite ca. 2 mm longo suffultum, lineare, albi-pilosum. Legumina ignota. 

Plants 1 7-^0 cm tall, covered with subappressed to ascending, symmetrically to slightly 
asymmetrically bifurcate white, in the inflorescence also black hairs. Caudex in upper part 
densely covered with blackish remnants of old stipules. Stipules 8-12 mm long, sometimes 
reddish-suffused, triangular (the lower) to narrowly triangular (the upper), adnate to the 
petiole for ca. 5 mm, at the base rather densely covered with appressed hairs, the free tips 
nearly glabrous. Leaves 10-24 cm long; petiole 2.5-9 cm long, loosely covered with 
subappressed to ascending, symmetrically to asymmetrically bifurcate, tousled hairs 0.2-1 
mm long. Leaflets in 6-12 pairs, linear-elliptic to more rarely narrowly elliptic, 15-35 x 2~4 
mm, acuminate to acuminate-mucronulate at the apex, on upper side glabrous or very sparsely 
to loosely hairy, minutely red-dotted, covered on underside with subappressed to confusely 
ascending hairs up to 1 mm long. Peduncle 1 2-20 cm long, finely striate-sulcate, loosely hairy 
like the rachis. Raceme loosely many flowered, 5-18 cm long, at fruiting time perhaps longer; 
axis with few black hairs. Bracts whitish-membranous, narrowly triangular, 4-5 mm long, 
sparsely black and white-hairy, at the margin ciliate with basifixed white hairs, in lower half 
with minute, subsessile, ovoid, glandlike trichomes. Pedicels ca. 1 mm long, black-hairy, erect, 
flowers erect. Bracteoles whitish, ca. 1 mm long, narrowly triangular, at the base of calyx. 
Calyx 10-11 mm long, tubular, obliquely inserted at the base, obliquely cut at the orifice, 
loosely covered with black and white, subappressed to slightly ascending, ± symmetrically 
medifixed hairs up to 0.8 mm long; teeth slightly unequal, narrowly triangular, 1-3 mm long. 
Petals pale sulphur-yellow. Standard 20-21 mm long; blade elliptic, ca. 9 mm wide, at the apex 
very shortly narrowed, rounded to slightly retuse, ± abruptly narrowed into the short claw. 
Wings ca. 19 mm long; blades narrowly oblong, rounded at the apex, 9 x 2.5 mm; auricle 2 mm 
long, claw 10 mm long. Keel 16 mm long; blades obliquely oblong-curved, with widely curved 
lower edge and concave upper edge, obtuse at the apex, 7x3 mm; auricle ca. 1 mm long, claw 
9 mm long. Staminaltube straightly cut. Ovary with a stipe ca. 2 mm long, linear, loosely con- 
fusely white-hairy. Legume unknown. 

Within the few species of the section with very narrow leaflets distinct by plants covered 
with subappressed to ascending, often tousled hairs, leaflets in 6-12 pairs, glabrous or very 
sparsely hairy on upper side, pale sulphur-yellow petals and shortly stipitate ovary. 

Astragalus gigantostegius Podlech, spec. nov. e sect. Laxiflori 

Holotype: Türkei, C4 Prov. Konya, S-Seite Krater Gölü E Karapinar, 1050 m, 30.5.1993, 
M. Nydegger 47338 (MSB; Iso: BASBG). 

Differt ab A. dictyolobus Bunge plantis validioribus ultra 60 cm altae (nee 1 5^0 cm altae), 
stipulis 18-22 (nee 7-15 mm) longis, foliolis 15-20 paribus (nee 17-28 paribus), anguste el- 



155 



lipticis vel anguste obovatis, 15-32 x 4-10 mm (nee ovatis ad elliptieis, 5-15 x 3-9 mm), 
racemis 5-6 cm longis, 8-10-floribus (nee 7-13 em longis, 15-30-45-floribus). 

Plants more than 60 em tall, eovered with thin, mostly spreading white hairs 2-5 mm long. 
Stem 60 em long or more, erect, somewhat flexuose, up to 7 mm in diameter, hollow, finely 
striate-suleate, loosely to rather densely hairy. Stipules green, narrowly triangular, long acumi- 
nate-subulate, 1 8-22 mm long, fi-ee from the petiole, long ciliate at the margins. Leaves 1 4-22 
cm long; petiole very short, 1-2 cm long, like the rachis deeply sulcate, loosely eovered with 
spreading hairs up to 5 mm long. Leaflets in 15-20 pairs, narrowly elliptic to narrowly ovate, 
rounded and shortly mucronulate to acute at the apex, 15-32 x 4-10 mm, on upper side 
glabrous, on underside loosely covered with ascending hairs up to 2.5 mm long. Racemes along 
most parts of the stem, subsessile, ovoid to oblong, 5-6 cm long, loosely 8-10-flowered; axis 
loosely spreadingly-hairy. Bracts green, filiform, 8-15 mm long, hairy. Pedicels 1-2 mm long, 
hairy. Calyx 13-16 mm long, shortly tubular at the beginning of anthesis, soon inflated and 
ovoid in fruit, straw-colored, rather densely covered with spreading hairs 4-5 mm long; teeth 
green, linear-acute to subulate, 5-10 mm long. Petals pale yellow. Standard 24 mm long; blade 
oblong-elliptic, slightly constricted below the middle, ca. 1 1 mm wide, widely rounded to 
truncate at the apex, abruptly and angularly constricted into the short, narrow claw. Wings ca. 
20 mm long; blades narrowly oblong, obliquely truncate and laterally subacute at the apex, 1 2 
x 4 mm; auricle 1 mm long, claw 8 mm long. Keel 16 mm long; blades obliquely elliptic, with 
in the middle moderately gibbously curved lower edge and nearly straight upper edge, obtuse 
at the apex, 7x4 mm; auricle ca. 1 mm long, claw 9 mm long. Staminaltube straightly cut. 
Ovary sessile, elliptic, hairy; style glabrous. Legumes sesssile, included in the calyx, elliptic 
seen from the side, ca. 10 mm long, 6 mm high and 3 mm wide, carinate ventrally and dorsally, 
abruptly narrowed at the apex into a recurved, subulate beak ca. 3 mm long, fully bilocular; 
valves pale brownish, thin but tough, somewhat rugulose, loosely covered with thin, white 
hairs 2-3 mm long. 

Astragalus karasarensis Podlech, spec. nov. e sect. Laxiflori 

Holotype: Anatolia, Prov. Sivas, B7 5 km E Karasar Ge9idi zwischen Kangal und Divrighi, 
1550 m, 15.6.1992, M Nydegger 46284 (MSB; Iso: BASBG). 

Differt dh A. phlomoides Boiss. foliolis 12-15 paribus (nee 15-32 paribus), 9-14 mm longis 
(nee 4-10 mm longis), subtus sparse ad laxe pilosis (nee subglabris), calyce 11-13 mm (nee 
15-17 mm) longo, dentibus ejus 2,5-5 mm (nee 6-8 mm) longis, vexillo 19-20 mm (nee 26-28 
mm) longo. 

Plants 1 5-25 em tall, covered with white hairs. Caudex ca. 5 mm in diameter, divided, with 
short branches. Stem 4-13 cm long, erect, somewhat flexuose, 2-3 mm in diameter, finely 
striate-suleate, loosely to rather densely covered with subappressed hairs 0.3-1 mm long and 
with subappressed to nearly spreading hairs up to 2(-3) mm long. Stipules green, narrowly 
triangular to linear, long acuminate, 8-12 mm long, nearly free from the petiole, short hairy. 
Leaves 8-15 cm long; petiole short, 1-3 em long, like the rachis sulcate, loosely hairy like the 
stem. Leaflets in 12-15 pairs, narrowly elliptic to narrowly ovate or elliptic to ovate, at the 
apex rounded to narrowly truncate, sometimes minutely mucronulate, 9-14 x 4-8 mm, 
glabrous on upper side, covered on underside sparsely to loosely, especially at the margins 



156 



and midvein with ascending hairs up to 0.5 mm long. Racemes 2-3 along the stem, subsessile, 
globular to shortly ovoid, 3-4 x 3 cm, rather densely many-flowered; axis sparsely hairy. 
Bracts green, filiform to linar-acute, 8-12 mm long, ciliate. Pedicels ca. 1 mm long, hairy. Calyx 
1 1-13 mm long, shortly tubular at the beginning of anthesis, soon inflated and ovoid in fruit, 
straw-colored, rather densely covered with ascending to spreading hairs up to 2 mm long; teeth 
green, narrowly triangular to linear-acute, 2.5-5 mm long. Petals yellow. Standard 19-20 mm 
long; blade obovate, scarcely constricted at the middle, ca. 1 mm wide, widely emarginate at 
the apex, at the base subangularly narrowed into the widely cuneate claw. Wings 1 8 mm long; 
blades narrowly elliptic, rounded at the apex, 8 x 3.5 mm; auricle 1.2-1.5 mm long, claw 10 
mm long. Keel 17-18 mm long; blades obliquely oblong-curved, with in the middle gibbously 
curved lower edge and concave upper edge, rounded at the apex, 7x4 mm; auricle 1 mm long, 
claw 10 mm long. Staminal-tube slightly obliquely cut. Ovary sessile, ovoid, white hairy; style 
glabrous. Legumes sessile, included in the calyx, obliquely elliptic seen from the side, with 
slightly curved dorsal line and strongly curved ventral line, rounded at the apex, ca. 6 mm long, 
4 mm high and 2 mm wide, subcarinate by the thickish nerve ventrally and dorsally, with a 
subulate straight beak ca. 2 mm long, fully bilocular; valves dirty straw-colored, thin but 
tough, loosely covered with subappressed hairs 0.5-1 mm long. 

Other specimens seen: 

Türkei. Sivas: B6N Tödürge Gölü, zwischen Zara und Hafik, 1350 m, 4.6.1990, Nydegger 45640 
(BASBG,MSB). 

Astragalus kentrophyllus Podlech, spec. nov. e sect. Leucocercis 

Holotype: Iran, Prov. Khuzestan: 25 km N of Haftgel towards Izeh, 550 m, 25.5.1973, M 
Iramhahr & M. Moussavi 15593-E (W). 

Differt dibA. ovoideo Sirj. & Rech.f. foliolis apice mucronula ad 0,5(-l) (nee 1-3) mm lon- 
gis, racema breviter pedunculata, legumine elliptica, primo pilis longioribus ad 0,8 (nee ad 2) 
mm longa, ab A. mucronifolio Boiss. foliolis 4-7 x 2-3 (nee 6-10 x 2-7) mm, apice mucronula 
adO,5(-l) (nee 1,5-3) mm longa provisis, pilis symmetrice medifixis 0,3-0,5 mm longis (nee 
arete asymmetrice medifixis ad 1 mm longis) obtectis, legumine a dorso compressa, 8 mm lata 
(nee a latere compressa, 3,5-5 mm lata). 

Plants spinose, cushion-like shrublets up to 15 cm tall, in most parts densely covered with 
appressed, ± symmetrically medifixed, 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 1-3 cm long, the dense indumentum of fine, basifixed white hairs up to 1 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 rachis up to 1 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(-l) mm long, on upperside 
densely, on underside loosely to rather densely covered with appressed, symmetrically medi- 
fixed hairs 0.3-0.5 mm long. Peduncles 1-1.5 cm long, curved, hairy like the rachis. Racemes 
loosely 3-flowered. Bracts whitish-hyalin, narrowly triangular, 2 mm long, ciliate. Pedicels at 



157 



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 orifice, 
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 midnerv. 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 3-3.8 
mm; auricle ca. 1 mm long, claw curved, 5-6 mm long. Keel 1 1-12 mm long; blades strongly 
obliquely elliptic-curved, with very widely rectangularly curved lower edge and concave upper 
edge, acute at the apex, 5-6 x 3-3.8 mm; auricle 0.5 mm long, claw 6 mm long. Staminaltube 
straightly to slightly obliquely cut. 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 in age. Seeds 3, nearly globular, 
5x4x3 mm, pale brownish, smooth. 

Other specimens seen: 

Iran. Ham : 10 km S. of Zarrin-Abad, Anaran Mountain, 1200-1300 m, 6.8.1989, Akhani 5540 
(MSB). - Khuzestan : II km W Haft-Kel, 300 m, 13.3.1959, Pabot 315 (G); 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, Iranshahr & Termeh 14964-E (W). - Pars : 4 km S Fahlian, 900 m, 
10.5.1959, Pabot s.n. (G); bei Kaserun, 8.5.1885, Stapf 1068 (WU); Kotal Komarech, 4.5.1885, 
Stapf 1143 {^NVi). 

Astragalus bakirdaghensis Podlech, spec, no v. e sect. Malacothrix 

Holotype: Türkei, B5 Kayseri, 8 km E Bakirdaghi gegen Tufanbeyli, 1330 m, 12.6.1987, M. 
Nydegger 42753 (MSB; Iso: BASBG). 

Plantae acaules ad subacaules, 20-25 cm altae, praecipue subappresse pilosae. Stipulae 
anguste triangulares ad lineares, 7-10 mm longae, petiolo adnatae per ca. 3 mm, breviter vagi- 
nanti-connatae, albi- et nigri-pilosae. Folia 10-17 cm longa, petiolo 3-5 cm longo, sicut rachis 
pilis appressis 0,05-0,4 mm longis et insuper pilis subappressis ad semipatentibus ad 0,8 mm 
longis obtecto. Foliola 14-18 paria, 5-12 x 1,5-3 mm, utrimque subappresse pilosa. Racemi 
pedunculo 13-18 cm longo subappresse piloso suffulti, densiuscule multiflori, 5-9 cm longi. 
Bracteae angustae, 5-9 mm longae, nigri-pilosae. Calyx ca. 14 mm longus, tubulosus vel leviter 
inflatus, densiuscule pilis semipatentibus ad patentibus albis nigrisque obtectus, dentibus 4-5 
mm longis. Corolla dilute flava. Vexillum 25 x 9 mm longum, lamina trullata, apicem versus 
late lingulatim attenuata. Alae 18 mm longae. Carina 17 mm longa. Legumina (immatura) sessi- 
lia, oblique elliptica, ca. 8 mm longa, pilosa, apice tenuiter rostrata. 

Plants acaulescent to nearly acaulescent, 20-25 cm tall, loosely to rather densely, mostly 
subappressed hairy. Stem, if present, up to 2 cm long, angularly-sulcate, densely appressed 
white hairy. Stipules whitish to greenish, narrowly triangular to linear-acute, 7-10 mm long, 
adnate to the petiole for ca. 3 mm, shortly connate behind the stem, glabrous at the base, with 
ciliate margins, in upper part rather densely covered with subappressed black and white hairs. 



158 



Leaves 10-17 cm long; petiole 3-5 cm long, like the rachis grooved at upper side, loosely 
covered with appressed white and sometimes few black hairs 0.05-0.4 mm long and sparsely 
with subappressed to ascending white hairs up to 0.8 mm long. Leaflets in 14-18 pairs, 
narrowly elliptic to narrowly obovate, 5-12 x 1.5-3 mm, obtuse to acute at the apex, loosely 
to densely covered at both sides with subappressed white hairs 0.5-1 mm long. Peduncle 13- 
1 8 cm long, striate-sulcate, loosely to rather densely covered with appressed to subappressed 
hairs 0.1-0.5 mm long and with much fewer hairs up to 1.5 mm long, in lower parts 
predominantly with white, toward the raceme with increasing black hairs. Racemes rather 
densely many-flowered, cylindric, 5-9 cm long, scarcely elongating at fruiting time. Bracts 
narrowly triangular-linear, acute, 5-9 mm long, black hairy. Pedicels 0.5-1 mm long, white- 
hairy. Calyx ca. 14 mm long, tubular to slightly inflated, obliquely cut at the orifice, rather 
densely covered with ascending to nearly spreading hairs 0.5-1.2 mm long, at the ventral 
(upper) side predominantly white, at the dorsal (lower) side predominatly black-hairy; teeth 
slightly unequal, the upper two narrowly triangular-subulate, ca. 4 mm long, the lower three 
subulate, ca. 5 mm long, glabrous on inner side. Petals pale yellow. Standard 25 mm long; blade 
trullate, 9 mm wide, widest in lower third, long and widely ligulately -narrowed upwards, 
scarcely emarginate at the apex, at the base abruptly narrowed into the long, cuneate claw. 
Wings 18 mm long; blades narrowly oblong, obtuse at the apex, 9 x 2.5 mm; auricle 1 mm long, 
claw 9 mm long. Keel 1 7 mm long; blades obliquely obovate, with in upper part widely curved 
lower edge and slightly concave upper edge, obtuse at the apex, 7 x 2.5 mm; auricle rounded, 
0.5 mm long, claw 10 mm long. Staminal tube straightly cut. Ovary sessile, elliptic, white 
hairy; style long, sparsely hairy in lower part. Legumes (immature) sessile, obliquely elliptic 
seen from the side, ca. 8 mm long, carinate ventrally, deeply grooved dorsally, subabruptly 
attenuate into a slender hairy beak. 

Astragalus jacobsii Podlech, spec. nov. e sect. Malacothrix 

Holotype: Iran, Prov. Azarbaijan Garbi, SW. side Lake Rezaiyeh, 36°56' N 45°52' E, 
1300-1400 m, fields and waste land near small salty lake, 21.5.1963, M. Jacobs 6645 (L; 
Iso: BG, E, GRO, K, MIN, W). 

Differt ab^. eriopodo Boiss. foliolis minoribus, 5-12 x 2,5 (nee 10-27 x 4-8) mm, supra 
glabris (nee sparse ad laxe pilosis), calyce 9-1 1 mm longo pilis ad 1 mm longis obtecto (nee 
11-13 mm longo pilis ad 2 mm longis obtecto), dentibus 2-2,5 (nee 3-6) mm longis, vexillo 
12-17 (nee 17-25) mm longo. 

Plants 40 cm tall, caulescent, covered with very fine, white, in the inflorescence also black 
hairs up to 1 mm long. Caudex repeatedly branched with slender, erect subterraneous 
branches. Stems several, up to 20 cm long, evenly foliate, finely striate, loosely covered with 
curly to flexuose, tousled hairs 0.1-1 mm long. Stipules whitish to greenish, 7-10 mm long, 
narrowly triangular, adnate to the petiole for ca. 2 mm, the lower distinctly, the upper 
indistinctly vaginate-connate behind the stem, all distinctly longitudinally nerved, sparsely to 
loosely hairy. Leaves 5-9 cm long; petiole 0.5-2.5 cm long, together with the rachis bisulcate 
at upper side, hairy like the stem. Leaflets in 6-9 pairs, elliptic, 5-12 x 2-5 mm, rounded at 
the apex, glabrous at upper side, loosely covered with flexuose-spreading hairs at lower side. 
Peduncles 2—4 in each stem, in the axils of the upper leaves 5-7 cm long, erect, angular-striate, 
hairy like the stem but with few spreading hairs up to 1 .5-2 mm long. Raceme cylindric, at 



159 



anthesis rather densely many-flowered, soon elongating and up to more than 1 5 cm long; axis 
hairy like the peduncle but with additional very short, spreading black hairs. Bracts whitish- 
greenish, 3-4 mm long, linear-acute to narrowly triangular, with long white and often with 
very short black hairs. Pedicels 0.5-1 mm long. Calyx reddish-suffused, 9-10 mm long, 
tubular, obliquely cut at the orifice, loosely covered with flexuose-spreading white hairs up to 
1 mm long and with short black hairs mainly toward the teeth; teeth narrowly triangular to 
subulate, 2-2.5 mm long, nearly glabrous at inner side. Petals dark salmon-red, fading bluish. 
Standard 12-17 mm long; blade 6-7 mm wide, rhombic, widest near the middle, in upper part 
triangularly narrowed toward the slightly emarginate apex, below the middle subabruptly 
narrowed into the long cuneate claw. Wings 1 1-15 mm long; blades narrowly oblong, rounded 
at the apex, 5-6 x 2 mm; auricle ca. 1 mm long, claw 6-9 mm long. Keel 10.5-14 mm long; 
blades obliquely oblong, with lower edge in upper part widely curved, and straight upper edge, 
subacute at the apex, 5 x 2.5 mm; auricle ca. 0.8 mm long, claw 7-9 mm long. Staminaltube 
straighly cut. Ovary with a stipe 1-2 mm long, narrowly elliptic, sparsely hairy; style 
glabrous. Pod (unripe) on a stipe ca. 3 mm long, narrowly elliptic, attenuate at both ends, 
densely covered with flexuose-spreading white hairs up to 4 mm long and with very short, 
spreading black hairs. 

Astragalus kashanensis Podlech, spec, nov., e sect. Malacothrix 

Holotype: Iran, Kashan (Mooteh protected region), ad biftircationem viae publicae versus 
Muteh (Mooteh) ducentis, ca. 140 km ab Esfahan boreo-occideentem versus, 1950 m, 
30.5.1974, K.H. Rechinger 46810 (W). 

Plantae subacaules, ca. 20 cm altae, pilis semipatentibus ad patentibus ad 2,5(-4) mm lon- 
gis, saepe in tuberculis minutis insidentibus obtectae. Caules ad 2 cm longi, pilis subappressis 
ad 1 mm longis obtecti. Stipulae 13-18 mm longae, per 3-5 mm petiolo adnatae, albi-nigri-pi- 
losae. Folia 12-18 cm longa, petiolo 3,5-6 cm longo, sicut rachis pilis ad 2,5 mm longis ob- 
tecto. Foliola 7-8 paria, remota, angusta, 12-20 x 2-3 mm, supra glabra, subtus longe pilosa. 
Racemi pedunculo 2,5-5 cm longo, sicut rachis piloso suffulti, densiuscule 10-30-flori. 
Bracteae angustae, 6-10 mm longae, albi-pilosae. Calyx 18-23 mm longus, tubulosus vel levi- 
ter ventricosus, pilis subpatentibus ad 4 mm longis obtectus, dentibus subulatis, 5-6 mm lon- 
gis. Vexillum 23 mm longum, lamina rhomboidei-elliptica, ca. 8 mm lata. Alae 21 mm longae. 
Carina alis aequilonga. Legumina subsessilia, elliptica, ca. 10 mm longa, pilis albis, oblique pa- 
tentibus ad 6 mm longis saepissime in tuberculis minutis insidentibus obtecta. 

Plants shortly caulescent, ca. 20 cm tall, loosely ftimished with rigid, ascending to 
spreading hairs mostly up to 2.5 mm long, often inserted onto little tubercles. Caudex slightly 
branched. Stems up to 2 cm long, sulcate, densely covered with subappressed white hairs up 
to 1 mm long. Stipules greenish, linear-acute to narrowly triangular, 13-18 mm long, adnate to 
the petiole for 3-5 mm, otherwise free from each other, loosely to rather densely white, at the 
base also black-hairy. Leaves 12-18 cm long; petiole 3.5-6 cm long, like the rachis loosely 
furnished with ascending to spreading white hairs up to 2.5 mm long. Leaflets in 7-8 pairs, 
remote, linear to narrowly ovate, often curved, folded, 12-20 x 2-3 mm, on upper side 
glabrous, on underside loosely to sometimes rather densely covered with subappressed to 
spreading white hairs up to 2.5 mm long. Peduncle one or more rarely two in each stem, 2.5-5 
cm long, rather thick, sulcate, straight, white-hairy like the rachis. Raceme ovoid to shortly 



160 



cylindric, rather densely 10-30-flowered; axis with long, nearly spreading white and few black 
hairs. Bracts linear-acute, 6-10 mm long, with long white, sometimes also with few black 
hairs. Pedicels ca. 1 mm long, white-hairy. Calyx 18-23 mm long, whitish-membranous, 
tubular to slightly ventricous, obliquely cut at the orifice, loosely furnished with nearly 
spreading white hairs up to 4 mm long; teeth subulate, 5-6 mm long, white-hairy also at the 
inner side. Petals colour unknown, in dry state dirty brownish. Standard ca. 23 mm long; blade 
ca. 8 mm wide, rhombic-elliptic, rounded at the apex, subabruptly narrowed into the long, 
narrowly cuneate claw. Wings ca. 21 mm long; blades narrowly oblong, rounded at the apex, 
ca. 8 X 2 mm; auricle ca. 0.5 mm long, claw 13 mm long. Keel ca. 21 mm long; blade obliquely- 
oblong, with widely curved lower edge and slightly concave upper edge, obtuse at the apex, ca. 
7x3 mm; auricle ca. 0.5 mm long, claw 14 mm long. Staminal-tube straightly cut. Ovary with 
a stipe ca. 1 mm long, ovoid, densely white hairy; style glabrous. Legume subsessile, elliptic, 
shortly attenuate at both ends, ca. 10 mm long, 3.5 mm high, compressed laterally, carinate 
ventrally, deeply grooved dorsally, the apex with a curved beak ca. 3 mm long, with ± 
persistent style; valves pale brown, densely covered with obliquely spreading white hairs up 
to 6 mm long, mostly inserted in little tubercles. Fruit completely bilocular, with 1-2 seeds in 
each locule. 



Astragalus sultan-bulaghensis Podlech, spec. nov. e sect. Malacothrix 

Holotype: Iran, Prov. Hamadan, Avadje, Sultan-bulagh mountains, 2000-2400 m, 
12.5.1987, A.A. Maassoumi & Mirhosseini 59388 (MSB; Iso: TARl). 

Differt ab A. spachiano Boiss. & Buhse caulibus pilis appressis ad 0,4 mm (nee ad 1 mm 
longis) obtectis, calycis indumento e pilis 0,3-1 mm longis, praecipue nigris consistente (nee e 
pilis brevissimis albis et insuper e pilis distincte longioribus albis et partim nigris consistente), 
indumento leguminis e pilis ad 4 mm longis subbasaliter affixis tantum consistente (nee e pilis 
ad 8 mm longis basaliter affixis et insuper pilis brevissimis consistente). 

Plants 7-20 cm tall, erect or prostrate, subacaulescent, furnished with white, in the 
inflorescence also with grey or blackish hairs. Caudex mostly slender, repeatedly branched, 
with short to more rarely elongated subterraneous branches. Stems very short, up to 3 cm 
long, densely covered with appressed white hairs 0.2-0.4 mm long. Stipules greenish, 4-6 mm 
long, those of lower leaves triangular to rarely ovate, sparsely hairy, those of upper leaves 
narrowly triangular, reflexed, ± densely subappressed hairy, all shortly (for ca. 1 mm) adnate 
to the petiole, behind the stem very shortly vaginate-connate. Leaves 2-12 cm long; petiole 1- 
4 cm long, like the rachis sulcate on upper side, loosely to rather densely covered with soft 
white hairs 0.1-0.5 mm long, which are in young leaves tangled, subappressed to rather 
ascending, in older leaves appressed. Leaflets in 11-16 pairs, elliptic, subacute or obtuse to 
rounded at the apex, 3-6 x 2-4 mm, at both sides densely covered with appressed hairs (0.3-) 
0.5-0.8 mm long. Peduncle 4-14 cm long, rather densely covered with appressed white, 
toward the raceme increasing black hairs. Raceme short, densely 4-11 -flowered, slightly 
elongating in fruit. Bracts 2-A mm long, narrowly elliptic, predominantly black hairy. Pedicels 
ca. 1 mm long. Calyx 1 1-13 mm long, tubular to slightly inflated, obliquely cut at the orifice, 
rather densely covered with tousled, spreading, predominantly black hairs 0.3-1 mm long, 
sometimes on the ventral (upper) side with often somewhat longer white hairs; teeth 2-3 mm 
lang, linear, hairy at iimer side. Petals drying dark violet. Standard 18-20 mm long; blade 



161 



slightly rhombic-elliptic, widest in the middle, ca. 8 mm wide, slightly retuse at the apex, at 
the base gradually narrowed into the wide, cuneate claw. Wings 15-17 mm long; blades 
narrowly oblong, rounded at the apex, 7.5-8 x 2.8 mm; auricle rounded, 1 .2-1 .8 mm long, claw 
8-9 mm long. Keel ca. 14 mm long; blades obliquely obovate, with in upper part widely, 
sometimes rectangularly curved lower edge and slightly concave upper edge, acutish at the 
apex, 6 X 2.5-3 mm; auricle 0.5-0.9 mm long, claw ca. 8 mm long. Staminal tube ± straightly 
cut. Ovary subsessile, pilose; style glabrous. Legumes subsessile, narrowly elliptic to 
spindlelike, attenuate at both ends, ca. 10 mm long, 3 mm high and wide, ventrally keeled, 
dorsally grooved, at the apex with a slightly recurved beak ca. 2 mm long; valves thin, tough, 
rather densely covered with spreading white hairs up to 4 mm long, subbasifixed with a short 
appendix. Fruit completely bilocular. 

Other specimen seen: 

Iran. Hamadan : Ecbatan sad to Yalfan village, 2000 m, 10.5.1987, Maassoumi 59334 (MSB). 

Astragalus sect. Mikrophyton Podlech, sect, nov 

Plantae acaulescentes vel breviter caulescentes, pulvinulos densos minutos formantes, pilis 
basifixis rigidis vel in foliolis pilis tenuissimis gossypineis obtectae. Stipulae seu petiolo 
adnate inter se liberae seu a petiolo liberae et ante petiolum alte inter se connatae. Foliola 2-3- 
paria. Flores singulae vel binae. Legumina sessilia vel subsessilia, parva, a latere compressa, 
ventro carinata, dorso carinata vel sulcata, incomplete ad complete bilocularia. Semina 1-3 pro 
loculo. Typus: A. mikrophyton Sirj. & Rech.f. 

A very isolated section within the basifixed hairy Astragali. 

Astragalus mikrophytoides Podlech, spec. nov. e sect. Mikrophyton 

Holotype: Afghanistan, Prov. Ghazni, in faucibus Say Khoshkak (Nawar Kotal) inter Okak 
et Behzud, ca. 33°53' N, 67°50' E, ca. 310 m, 6.7.1962, K.H. Rechinger J 7828 (W; Iso: 
MSB). 

Differt ab A. mikrophyton Sirj. & Rech.f. stipulis a petiolo liberis sed ante petiolum alte, 
saepe fere ad apicem inter se connatis, foliolis 2-paribus, planis, utrimque pilis rigidis ad 0,8 
mm longis obtectis, calyce laxe pilis patentibus albis 0,5-2 mm longis et nigris brevioribus 
obtecto, dentibus filiformibus 5 mm longis, leguminibus dorso distincte sulcatis, densiuscule 
pilis subappressis ad oblique patentibus brevibus ad 0,5 mm et insuper pilis longis patentibus 
3-4 mm longis obtectis, seminibus 3 pro loculo. 

Plants small, shortly caulescent, forming dense, small cushions up to 4 cm tall, covered 
with rigid, basifixed hairs. Caudex slender or up to 5 mm in diameter, with a pluricipital root- 
crown, with short to somewhat elongated, erect branches, at the apex often covered with slen- 
der persistent, erect old petioles. Stems absent or up to 2 cm long, densely covered with 
spreading white, below the nodes also black hairs 0.2-0.5 mm long. Stipules whitish-hyalin, 
3-4 mm long, free from the petiole, in front of the petiole connate nearly to the apex, covered 
sparsely to loosely, especially toward the margins with spreading white and black hairs 0.1- 
0.5 mm long. Leaves 1-2 cm long; petiole 0.5-1.2 cm long, slender, covered with rigid, white, 



162 



spreading bristles 0.2-0.6 mm long. Leaflets mostly in 2 pairs, obovate, 3-8 x 1.5-3.5 mm, at 
the apex rounded or more rarely subacute, flat, on both sides rather densely to densely covered 
with subappressed to ascending white hairs up to 0.8 mm long. Racemes nearly sessile, 1-2- 
flowered. Bracts whitish-hyaline, ca. 4 mm long, narrowly triangular, sparsely predominantly 
black-hairy. Pedicel ca. 2 mm long, spreadingly hairy. Bracteoles filiform, ca. 1 mm long, in the 
middle of pedicel. Calyx ca. 10 mm long, shortly tubular, the tube loosely covered with 
spreading white hairs 0.5-2 mm long and with much shorter black hairs; teeth filiform, ca. 5 
mm long, long spreadingly ciliate. Petals pink. Standard ca. 15 mm long; blade obovate, ca. 6 
mm wide, at the apex slightly emarginate, long gradually narrowed at the base. Wings ca. 14 
mm long; blades narrowly oblong, rounded at the apex, 9 x 2.2 mm; auricle indistinct, claw ca. 
5 mm long. Keel ca. 9 mm long; blades triangular-elliptic, with widely curved lower edge and 
slightly curved upper edge, obtuse at the apex, 4 x 2.5 mm; auricle indistinct, claw ca. 5 mm 
long. Staminal-tube slightly obliquely cut. Legumes sessile, elliptic, ca. 9 mm long, 4 mm high 
and 3 mm wide, carinate ventrally, grooved dorsally, at the apex with a slightly bent, straight 
beak 2-3 mm long, nearly completely bilocular; valves thin, straw-colored, rather densely 
covered with subappressed to ascending white and sometimes few black hairs 0.2-0.5 mm 
long and with ± spreading white halts 3-4 mm long. Seeds three in each locule, ca. 2.5 mm 
long. 

Astragalus arakliensis Podlech, spec. nov. e sect. Onobrychoidei 

Holotype: Türkei, A7 Trabzon, Arakli - Bayburt, 19 km S Daghbasi, 1900 m, 10.6.1988, 
M Nydegger 43284 (MSB). 

Differt ab A. mukusiense Rech.f. foliolis latioribus apice distincte inciso-emarginatis, calyce 
imprimis nigripiloso dentibus longioribus, ab A. pineticola Podlech bracteis brevioribus latiori- 
busque, calyce breviore tempore fructificationis ventricoso, vexillo breviore elliptico nee apice 
longe lingulatim attenuato. 

Planta 20 cm alta, pilis plerumque patentibus 0,3-1 mm longis, subsimplicibus aut in tu- 
berculis minutis aut sublateraliter cicatrice oblongo affixis obtecta. Caules erecti vel sub- 
prostrati, ad 15 cm longi, laxe pilis ± patentibus obtecti. Stipulae ad 6 mm longae, inter se alte 
connati-vaginantes, extus laxe pilis nigris obtectae, margine pilis mere albis rigidis in tuberculis 
distinctis insidentibus ciliatae. Folia 2-6 cm longa, rachide patenter pilosa. Foliola 10-13 paria, 
anguste elliptica, apice distincte incisi-emarginata, 4-10 x 2-4 mm, supra sparse, subtus laxe 
subappresse albipilosa. Pedunculi 2-6 cm longi. Racemi breves, 3-8 flori. Bracteae 3-5 mm 
longae, pilosae. Calyx 9-10 mm longus, tubulosus, tempore fructificationis leviter ventricosus, 
laxe pilosus, dentibus 2,5-3,5 mm longis. Corolla violacea. Vexillum 18-20 mm longum, 
anguste ellipticum. Alae ca. 15 mm longae. Carina ca. 13 mm longa. Ovarium sessile, albipilo- 
sum. Legumen (immaturum) sessile, anguste ellipsoideum, apice attenuatum, longe albipilo- 
sum. 

Plants cespitose, ca. 20 cm tall, covered with mostly spreading hairs 0.3-1 mm long, sub- 
basifixed, sitting on minute tubercles or with the subbasal, oblong cicatrice affixed to the 
tubercle. Caudex strongly branched. Stems several or many, erect or nearly prostrate, up to 15 
cm long, loosely covered with ± spreading white and especially at the nodes also with black 
hairs. Stipules greenish, the lower ones up to 3 mm long, the upper ones up to 6 mm long. 



163 



triangular, nearly free from the petiole, behind the stem high up connate-vaginate, on outer side 
loosely covered with black hairs, at the margins ciliate with rigid, merely white hairs sitting on 
little tubercles. Leaves 2-6 cm long; petiole 0.4-1.5 cm long, loosely covered with spreading, 
predominantly black hairs, rachis slender, covered with spreading, predominantly white hairs. 
Leaflets in 10-13 pairs, narrowly elliptic, 4-10 x 2-4 mm, at the apex distinctly emarginate- 
incised, on upper side sparsely, on underside loosely subappressed white hairy. Peduncles 2- 

6 cm long, loosely covered with spreading white, toward the raceme increasing black hairs. 
Racemes short, 3-8-flowered. Bracts whitish, narrowly ovate, 3-5 mm long, covered with 
nearly spreading black hairs and with white hairs partly sitting on little tubercles, at the 
margins also with few sessile or shortly stipitate gland-like hairs. Pedicels ca. 1 mm long, black 
hairy. Calyx 9-10 mm long, tubular, obliquely cut at the orifice, at fi-uiting time slightly 
inflated, loosely to rather densely covered with ascending, predominantly black hairs; teeth 
subulate, 2.5-3.5 mm long, white hairy on innerside. Petals glabrous, violet. Standard 18-20 
mm long; blade narrowly elliptic, at the apex distinctly incised, at the base cuneately 
narrowed. Wings ca. 15 mm long; blades narrowly oblong, at the apex rounded or slightly 
emarginate, 7 x 2.8 mm; auricle curved, ca. 1 mm long, claw ca. 8 mm long. Keel ca. 13 mm 
long; blades obliquely elliptic, with widely curved lower edge and sigmoid-concave to nearly 
straight upper edge, at the apex acutish, ca. 6 mm long; auricle acute, ca. 1 mm long, claw ca. 7 
mm long. Staminal-tube straightly cut. Ovary sessile, densely white hairy; style glabrous. 
Legumen (unripe) sessile, narrowly ellipstic, long white hairy. 

Astragalus demirizii R.Kramer & Podlech, spec. nov. e sect. Onobrychoidei 

Holotype: Türkei, Ankara, Tuz Gölü, E Ufer, 14 km S Sereflikochisar, 899 m, Halophyten- 
flur, 20.5.1986, U. Kramer 52 (BSB-VO; Iso: BSB-VO, MSB). 

Plantae 20-50 cm altae, dense albi-pilosae. Caules 10-45 cm longi, dense pilis ca. 1 mm 
longis obtecti. Stipulae 2-A mm longae, inter se connatae, pilosae. Folia (2-)6-10 cm longa; 
petiolo brevissimo, rachide dense pilosa. Foliola 6-9 paria, subsessilia, (5-)7-12 x (2-)3-5,5 
mm, elliptica vel obovata ad anguste obovata, utrimque appresse pilosa. Racemi pedunculo 3- 

7 cm longo, piloso suffulti, 20-50-flori. Bracteae 1,5-2,5 mm longae, ciliatae. Calyx 6,5-8,5 
mm longus, tubulosus, appresse ad patenter pilosus, dentibus triangularibus 1 mm longis. Co- 
rolla glabra, incamata. Vexillum 15-19 mm longum, rotundato-rhombiforme, 6-9 mm latum. 
Alae 14-18 mm longae. Carina 12-14 mm longa. Ovarium sessile, pilosum. Legumina sessilia, 
elliptica, 7-10 mm longa, 3 mm alta et 2 mm lata, ventro carinata, dorso rotundata, pilis 
flexuosis, 2-3 mm longis obtecta, incomplete biloculare. 

Plants procumbent to ascending, 20-50 cm tall, densely covered with mostly ± 
symmetrically medifixed white hairs. Caudex up tp 10 mm in diameter. Stems 10^5 cm long, 
sulcate, equally foliate, densely covered with appressed hairs ca. 1 mm long. Stipules mem- 
branous, 2-4 mm long, nearly free fi-om petiole, behind the stem vaginate-connate up to two 
third, with two triangular tips, sparsely covered with asymmetrically medifixed hairs 0.5-0.8 
mm long, glabrescent, ciliate at the margins. Leaves (2-)6-10 cm long; petiole very short, 0.2- 
0.5 cm long, like the rachis densely covered with appressed, symmetrically medifixed hairs ca. 
0.5 mm long. Leaflets in 6-9 pairs, narrowly elliptic to narrowly ovate, (5-)7-12 x (2-)3-5.5 
mm at the apex obtusish to acutish, on both sides covered with appressed hairs 0.5-1 mm 
long. Peduncles 3-7 cm long, sulcate, densely covered with appressed hairs ca. 0.5 mm long. 



164 



Racemes densely 20-50-flowered, at fruiting time elongating up to 7 cm. Pedicels ca. 1 mm 
long. Flowers erect. Bract membranous, 1.5-2.5 mm long, narrowly triangular, sparsely hairy, 
at the margins cilate with strongly asymmetrically bifurcate hairs ca. 0.5 mm long. Calyx 
tubular to slightly inflated, 7.5-9.5 mm long, covered with appressed, symmetrically medi- 
fixed white hairs and with much fewer spreading, asymmetrically medifixed hairs 0.5-1 mm 
long; teeth ca. 1 mm long, triangular, in innerside covered with asymmetrically medifixed and 
basifixed hairs 0.5 mm long. Petals glabrous, pink, fading dark violet. Standard 13-19 mm long; 
blade 6-9 mm wide, obtusely rhombic, at the apex emarginate, gradually narrowed at the base. 
Wings slightly shorter than the standard, 13-18 mm long; blades narrowly oblong, at the apex 
obliquely emarginate, 6-9 x 1.5-2.5 mm; auricle ca. 1 mm long, claw 7-9 mm long. Keel 12-14 
mm long; blades obliquely narrowly ovate, with nearly straight upper edge, at the apex 
acutish, 4.5-6 x 2-2.5 mm; auricle ca. 0.5 mm long, claw 7.5-9 mm long. Staminal-tube 
straightly cut. Ovary sessile, covered with somewhat flattened white hairs; style hairy at the 
base. Legumes with a stipe up to 1 mm long, narrowly elliptic, 7-10 mm long, ca. 3 mm high 
and 2 mm wide, carinate ventrally, rounded dorsally, with a strongly curved beak ca. 1 mm 
long, incompletely bilocular; valves straw-colored, densely covered with subbasifixed, straight 
to flexuouse hairs 2-3 mm long. Seeds ca. 3 in each locule, ca. 1 mm long, brown, rugulose. 

Other specimen seen: 

Türkei. Ankara : Tuz Gölü, 14 km S Sereflikochisar, 905 m (Strandwall am Seeufer, Sanddüne) 
Halophytenflur, 9.6. 1986, Kürschner 2332 (BSB-VO). 

Astragalus eskishehiricus Podlech, spec. nov. e sect. Onobrychoidei 

Holotype: Türkei, B3 Eskishehir, Cifteler - Eskishehir, 1 km S Hamidiye, 930 m, 15.6.1993, 
M. Nydegger 47470 (MSB). 

Plantae ca. 30 cm altae, indumento e pilis appressis symmetrice ad leviter asymmetrice 
medifixis, 0,2-0,5 mm longis consistente. Caulis ad 30 cm longus sparse albi-nigri-pilosus. Sti- 
pulae 2,5-3 mm longae, triangulares, petiolo vix adnatae, inter se liberae. Folia 7-10 cm longa, 
subsessilia, rachide sparse ad laxe albi-nigri-piloso. Foliola 10-13 paria, anguste elliptica, apice 
obtusa, 5-14 x 1,5-3 mm, supra glabra subtus laxe albipilosa. Pedunculi 7-12 cm longi, ut in 
caule pilosi. Racemus dense 10- ad multiflorus. Bracteae 0,5-1,5 mm longae. Calyx 4-5 mm 
longus, campanulatus, albi-nigripilosus, dentibus subulatis, 1-2 mm longis. Corolla glabra, 
violacea. Vexillum 14-15 mm longum, lamina ovata, ca. 6 mm lata. Alae 10-11 mm longae. 
Carina ca. 7 mm longa. Ovarium sessile, albipilosum, stylo glabro. Legumen (immaturum) 
anguste ovoideum, apice angustatum, dense subappresse albivillosum. 

Plants ca. 30 cm tall, covered with appressed, symmetrically to slightly asymmetrically 
medifixed hairs 0.2-0.5 mm long. Stem up to 30 cm long, erect or at the base ascending, 
sparsely covered with white and black hairs. Stipules 2.5-3 mm long, triangular, scarcely 
adnate to the petiole, free from each other, at the margins white and black ciliate. Leaves 7-10 
cm long, nearly sessile; rachis sparsely to loosely white and black hairy. Leaflets in 10-13 
pairs, narrowly elliptic, 5-14 x 1.5-3 mm, obtuse at the apex, on upper side glabrous, on 
underside loosely white hairy. Peduncles 7-12 cm long, hairy like the stem. Raceme short or 
elongated, densely 10-many-flowered. Bracts whitish, ovate, 0.5-1.5 mm long, black ciliate. 
Pedicels ca. 1 mm long. Calyx 4-5 mm long, campanulate, with slightly obliquely cut orifice, 
loosely white and black hairy; teeth subulate, unequal, the two upper ca. 1 mm long, the three 



165 



lower up to 2 mm long, all white hairy on innerside. Petals glabrous, violet. Standard 14-15 
mm long; blade ovate, ca. 6 mm wide, at the apex rounded, at the base shortly cuneately 
narrowed. Wings 10-1 1 mm long; blades narrowly oblong, at the apex slightly emarginate, 7 x 
2 mm; auricle curved, ca. 0.8 mm long, claw ca. 4 mm long. Keel ca. 7 mm long, blades 
obliquely elliptic, with widely curved lower edge and nearly straight upper edge, at the apex 
acutish, 4x2 mm; auricle minute. Ovary sessile, white hairy; style glabrous. Legumes (unripe) 
narrowly ovoid, densely subappressed white hairy. 

Astragalus konyaensis Podlech, spec. nov. e sect. Onobrychoidei 

Holotype: Türkei, C4 Konya, Kizilyaka - Hadim e Kaplandi, 1330 m, 22.6.1990, M. Ny- 
degger 45772 (MSB; Iso: BASBG). 

Species inter alia sectionis insignis ob indumentum caulium, rachidium pedunculorumque e 
pilis appressis ± symmetrice medifixis et insuper densiuscule pilis ad triplo longioribus, semi- 
patentibus ad subappressis, valde asymmetrice medifixis consistentem, ob denies calycis ri- 
gide pectinatim ciliatos et ob petala longa, subcaerulea. 

Plants up to 35 cm tall, with a double indumentum of medifixed white, at the base of 
stipules also with black hairs. Stem up to 30 cm long, adscending to erect, densely covered 
with appressed, ± symmetrically medifixed hairs and rather densely with subappressed to 
ascending, strongly asymmetrically medifixed hairs three times longer. Stipules 7-12 mm long, 
narrowly triangular or from a triangular base long acuminate, scarcely adnate to the petiole, 
behind the stem shortly to high up connate-vaginate, densely white, at the base also black 
hairy. Leaves 10-15 cm long; petiole 1-5 cm long, together with the rachis hairy like the stem. 
Leaflets in 5-8 pairs, narrowly elliptic to narrowly obovate, 6-23 x 2.5-7 mm, at the apex 
obtusish to acutish, densely covered with rigid, subappressed to ascending, strongly asymme- 
trically medifixed to nearly symmetrically medifixed hairs up to 2(-3) mm long, often folded 
when young. Peduncle strong, 9-19 cm long, hairy like the stem. Raceme nearly globose, 
densely many-flowered. Bracts narrowly triangular, 8-1 1 mm longe, densely subappressed 
hairy. Flowers nearly sessile. Calyx 11-15 mm long, campanulate-tubular, ± densely sub- 
appressed hairy; teeth 6-9 mm long, subulate, densely pectinately covered with rigid, nearly 
spreading hairs. Petals bluish- violet, glabrous. Standard 18-20 mm long, 8-10 mm wide, 
trullate to nearly rhombic, at the apex widely rounded and slightly emarginate, at he base 
shortly cuneately narrowed. Wings 13-15 mm long; blades obliquely narrowly obovate, at the 
apex rounded, 7.5-9 x 3-4 mm; auricle ca. 1 mm long, claw 5-6 mm long. Keel 12-13 mm 
long; blades obliquely elliptic, with widely curved lower edge and sigmoid-concave to nearly 
straight upper edge, acutish at the apex, 6-7 x 3-3.5 mm; auricle minute, claw ca. 6 mm long. 
Staminal-tube straightly cut. Ovary sessile, densely white hairy; style glabrous. Legumes un- 
known. 

A very conspicuous species easily recognizable by the double indumentum of stems, 
rachides and peduncles, the rigidly ciliate calyx teeth and the long, bluish petals. 

Other specimens seen: 

Türkei. C4 Konya : Belören - Hadim, 10 km N Hadim, 1370 m, 28.7.1992, Nydegger 46879 
(BASBG, MSB). 



166 



Astragalus pineticola Fodlech, spec. no\. e sect. Onobrychoidei 

Holotype: Türkei, C5 l9el, Arslanköy - Camliyayla W (^amliyayla, 700 m, Föhrenwald 
[Pinetum], 13.5.1987, M Nydegger 42329 (MSB; Iso: BASBG). 

Differt ab A. setuloso Boiss. & Balansa indumento e pilis extremiter asymmetrice medifixis 
partim tuberculis minutis affixis et simplibus immixtis consistente, calyce longiore, vexillo 23- 
25 mm longo, apice longe lingulatim attenuate. 

Plantae 25-30 cm altae, dense pilis extremiter asymmetricis ad subsimplicibus albis sub 
nodis insuper nigris, partim appressis partim subapressis ad oblique patentibus 0,3-1 mm 
longis obtectus. Stipulae 3-6 mm longae, a petiolo liberae, inter se ad dimidium longitudinis 
connati-vaginantes. Folia 7-12 cm longa. Foliola (7-) 10-1 5 paria, elliptica, apice distincte et 
anguste emarginato-incisa, 5-1 1 x 2-A mm, utrinque laxe subappresse albipilosa. Racemus pe- 
dunculo 9-10 cm longo suffultus, brevis, dense ca. 10 florus. Bracteae lineari-acutae, 6-7(-8) 
mm longae. Calyx 12-13 mm longus, tubulosus, pilis albis nigrisque oblique patentibus laxe 
obtectus, dentibus subulatis 4-5 mm longis. Corolla glabra, violacea. Vexillum 23-24 mm Ion- 
gum, lamina medio rhomboidei-dilatata, ca. 8 mm lata, apicem longe lingulatim attenuata. Alae 
ca. 16 mm longae. Carina ca. 15 mm longa. Ovarium sessile, dense albipilosum. 

Plants 25-30 cm tall. Stem ascending, densely covered with extremely asymmetrically 
medifixed and subbasifixed, partly appressed, partly ascending white hairs 0,3-1 mm long. 
Stipules greenish, the lower ones 3-4 mm, the upper ones 5-6 mm long, narrowly triangular, 
free from the petiole, behind the stem connate-vaginate up to the middle, loosely hairy with 
white hairs which are partly and especially at the margins sitting on little tubercles. Leaves 7- 
12 cm long, petiole 1,5-2,5 cm long, like the rachis covered with ascending to spreading hairs 
partly sitting on little tubercles. Leaflets in (7-)10-15 pairs, elliptic, at the apex distinctly et 
narrowly incised, 5-1 1 x 2-4 mm, on both sides loosely covered with subappressed hairs. 
Peduncle 9-10 cm long, hairy like the stem but with increasing black hairs toward the raceme. 
Raceme short, densely ca. 10-flowered. Bracts whitish, linear-acute, 6-7(-8) mm long, with 
black and white nearly spreading hairs partly sitting on little tubercles and at the margins with 
few sessile to shortly stipitate gland-like hairs. Calyx 12-13 mm long, tubular, loosely 
spreadingly black and white hairy; teeth subulate, 4-5 mm long, with ascending black hairs and 
especially at the margins with white hairs partly sitting on minute tubercles. Petals glabrous, 
violet. Standard 23-24 mm long; blade in the midlle rhombic-dilated, ca. 8 mm wide, toward 
the apex long ligulately narrowed, at the base long cuneately narrowed. Wings ca. 1 6 mm long; 
blades narrowly oblong, rounded at the apex, 7x2 mm; auricle curved, ca. 1 mm long, claw ca. 
9 mm long. Keel ca. 15 mm long; blades obliquely elliptic, with widely rounded lower edge and 
nearly straight upper edge, at the apex minutely acuminate, ca. 6 x 3 mm; auricle straight, 
acute, ca. 0.5 mm long, claw ca. 9 mm long. Ovary sessile, ovoid, densely white hairy; style 
hairy in basal part. Legumes unknown. 



167 



Astragalus venulosm Boiss. subsp. iraqensis Podlech, subspec. nov. (Sect. Rhabdotus) 

Holotype: Iraq, ad confines Turciae prov. Hakan, in ditione pagi Sharanish, in saxosis 
cacuminis mentis Zawita a Zakho septentrionem versus, 2000 m, 4.-9.7.1957, K.H. 
Rechmger]0967{W). 

Figure: Townsend& Guest (1974). 

Differt ab subspecie typico indumento e puis conspicue brevioribus consistente, in caule ad 
0,7 (nee ad 2,5) mm longis, in pedunculo pilis 0,3-1 mm longis tantum (nee insuper pilis 1,5- 
2,5 mm longis), leguminibus altioribus latioribusque (11-13 x 6-8 x 5-6 mm nee 10x5x3 
mm) dorso ventroque obtuse carinatis (nee acute earinatis) et indistincte numero foliolarum, 
(3-)7-14(-17) paria (nee (6-)12-17 paria). 

Astragalus nigdeanus Podlech & Ekici, spec. nov. e sect. Sisyrophorus 

Holotype: Turkey, C4 Prov. Karaman, Koros'in ust kesimleri. 1750 m, 22.6.1998, M.Ekici 
2093 (GAZI; Iso: MSB). 

Differt ab A. aydosensi Pesmen & Erik foliolis angustioribus 4-5 (nee 1-2) paribus, ab A. 
pelligero Fenzl foliolis angustioribus, ab utraque corolla violacea et pilis longis calycis partim 
in tuberculis cavis conspicuis insidentibus. 

Plants 5-7 cm tall, suffruticulose, branched at the base, with medifixed white, at the 
stipules and calyx also with black hairs. Stipules 4-5 mm long, adnate to the petiole for 2 mm, 
behind the stem vaginate-connate nearly up to the top, toward the apex rather densely white 
and black hairy. Leaves 1.0-2.5 cm long; petiole nearly as long as the blade, like the rachis 
densely covered with short, somewhat flexuose, subappressed hairs and mostly also with 
asymmetrically medifixed, straight, spreading hairs up to 2 mm long. Leaflets in 4—5 pairs, 2-8 
X 0.5-2 mm, narrowly obovate, at the apex obtuse, the upper ones often close together, all on 
both sides densely sericeous with subappressed hairs. Racemes with a peduncle 1.5-2.5 cm 
long and hairy like the rachis, densely globose, many-flowered. Bracts narrowly ovate- 
acuminate, ca. 6 mm long, loosely white and sometimes also black hairy. Pedicels ca. 1 mm 
long. Calyx widely campanulate-tubular, 8-10 mm long, densely covered with short, some- 
what flexuose, white, toward the teeth predominantly black hairs and with asymmetrically 
medifixed, spreading white hairs up to 2,5 mm long and partly sitting on distinct, hollow 
tubercles; teeth subulate, 3-4 mm long. Petals violet, standard and wings hairy. Standard ca. 17 
mm long, ca. 7 mm wide, narrowly rhomboid-elliptic, at the apex narrowed into tonguelike 
obtuse appendix, at the base cuneately narrowed, densely subappressed hairy on the backside. 
Wings ca. 1 3 mm long; blades narrowly oblong, rounded at the apex, ca. 7 x 2 mm, on outer 
side loosely covered with long hairs; auricle 0.5 mm long, claw filiform, 6 mm long. Keel ca. 1 1 
mm long; blades obliquely elliptic, with widely curved lower edge and nearly straight upper 
edge, acutish at the apex, 5x2 mm; auricle minute, claw ca. 5 mm longe, connate in upper half 
Ovary covered with very long, nearly spreading white hairs; style short, glabrous. Legumes 
unknown. 



168 



Other specimens seen: 

Turkey. C4 Karaman : Koros' in ust kesimleri. 1750 m, 22.6.1998, Ekici 2098 (GAZl, MSB). - 
C5 Nigde : Camardi, von Cukurbag gegen Emli Bogazi, 1650 m, 15.6. 1987, Nydegger 42802 (BASBG, 
MSB). 

Astragalus hakkariensis Podlech, spec. nov. e sect. Stereothrix 

Holotype: Türkei, C 10 Prov. Hakkari, 5 km W Esendere gegen Yüksekova, 1720 m, 
24.7.1983, MyVvJegger75¥27 (MSB; Iso: BASBG). ,., .■ ~ , ■ 

Differt ab A. sphaeranthus Boiss. pilis pro parte, imprimis in calyce tuberculis 
insidentibus, petalis conspicue longioribus, vexillo 22-25 mm longo, anguste oblongo (nee 16- 
17 mm longo, elliptico), legumine minore, appresse piloso (nee semipatenter piloso), 
seminibus 1 pro loculo (nee 2-3 pro loculo). 

Plantae 12-20 cm altae, caespitosae, pilis subrigidis ad l(-2) mm iongis obtectae. Caules 
dense pilis patentibus subflexuosis obtecti. Stipulae 5-9 mm longae, petiolo breviter adnatae, 
breviter vaginanti-connatae, pilosae. Folia 4— 7 cm longa, petiolo 1-2 cm longo, rachide piloso. 
Foliola 8-11 paria, anguste elliptica, 5-12 x 1,5-5 mm, utrimque dense subappresse pilosa. 
Racemi pedunculo 1,5-5 cm longo, patule piloso suffulti, globosi, dense multi-flori. Bracteae 
lineares, 8-15 mm longae, pilosae. Calyx 13-16 mm longus, turbinati-tubulosus, in parte basali 
sparse ad laxe, in parte apicali densiuscule pilis Iongis in tuberculis insidentibus obtectus, den- 
tibus subulatis, 7-1 1 mm Iongis. Petala pallide flava vel lactea. Vexillum 22-25 x 6-7 mm, 
anguste oblongi-ellipticum. Alae 17-18 mm longae. Carina 14-15 mm longa. Legumina sessilia, 
elliptica, ca. 8 mm longa, 3 mm alta et 2 mm lata, pilis subappressis 0,2-0,3 mm Iongis ob- 
tecta. Semina una pro loculo. 

Plants 12-20 cm tall, cespitose, covered with subrigid white hairs, in vegetative parts up to 
1 mm long. Caudex up to 6 mm in diameter, repeatedly branched, with short subterraneous 
branches. Stems several, 9-12 cm long, ofl;en branched, densely covered with spreading, often 
somewhat tousled hairs 0.3-1 itmi long. Stipules membranous with greenish tip or completely 
greenish, 5-9 mm long, triangular to narrowly triangular, adnate to the petiole for ca. 1 mm, 
shortly vaginate-connate behind the stem, loosely hairy, the marginal hairs often sitting on 
little tubercles. Leaves 4-7 cm long; petiole 1-2 cm long, together with the rachis hairy like the 
stem. Leaflets in 8-1 1 pairs, narrowly elliptic, 5-12 x 1.5-5 rrmi, rounded to slightly retuse at 
the apex, densely covered on both sides with subappressed hairs, flat. Peduncle 1 .5-5 cm long, 
striate-sulcate, densely covered with spreading white hairs 0.3-1.5(-2) mm long. Raceme 
globose, densely many-flowered, 2.5-4 cm in diameter. Bracts greenish, narrowly triangular- 
linear, 8-15 mm long, covered with spreading white hairs often sitting on little tubercles. 
Flowers subsessile. Calyx 13-16 mm long, turbinate-tubular, in lower part of the tube 
sparsely to loosely, in upper part and at the teeth rather densely covered with long spreading 
hairs sitting on distinct tubercles; teeth subulate, 7-1 1 mm long, hairy all around. Petals all 
pale yellowish or creamy white, upper part of wings and keel sometimes fading pale violet on 
drying or all whitish, tinged with violet. Standard 22-25 mm long, narrowly oblong-elliptic, 6- 
7 mm wide, widely rounded or minutely incised at the apex, scarcely narrowed at the base. 
Wings 17-18 mm long; blades narrowly oblong to slightly dilated toward the rounded apex, 8- 
9x3 mm; auricle ca. 1 mm long, claw ca. 9 mm long. Keel 14-15 mm long; blades obliquely 
elliptic, with widely curved lower edge and nearly straight upper edge, shortly acuminate at 



169 



the apex, 5-6 x 3 mm; auricle minute, claw 8-9 mm long. Staminaltube straightly cut. Ovary 
sessile, appressed white-hairy; style glabrous. Pod sessile, elliptic, ca. 8 mm long, 3 mm high 
and 2 mm wide, carinate ventrally, narrowly grooved dorsally, at the apex with a subulate beak 
ca. 1 mm long, fully bilocular; valves thin, straw-colored, densely covered with subappressed 
hairs 0.2-0.3 mm long. Seeds only one in each locule, elliptic, 4x2 mm. 

Other specimens seen: 

Turkey. Hakkari : 10 km from Bajirge to Yüksekova, 1900 m, \8.6.\966, Davis 45208 (E) - 2\ 
km from Bajirge to Yüksekova, 2230 m, 18.6.1966, Davis 45234 (E) - Pass on Bajirge - Yiikse-kova 
road, 2500 m, 30.6.1967, Watson & al. 2962 (E, K) - dto., 2200 m, 30.6.1975, Rechinger 53839 
(M, W). 

Astragalus montis-varvashti Podlech, spec. nov. e sect. Stereothrix 

Holotype: Iran, Mazanderan: Elburz, Elika, Varvasht mountain, 3500^100 m, 4.8.1972, F. 
Terme ]5240-E{W). 

Differt ab A. capito stipulis duplo longioribus, 4—8 mm (nee 2-4 mm) longis, basi pilis par- 
tim nigris provisis, pilis pedunculi longioribus ad 2 mm (nee ad 1 mm) longis, bracteis 
conspicue majoribus, 9-13 mm (nee 4—7 mm) longis, calyce ca. 15 mm longo dentibus 7-9 mm 
(nee 11-12 mm longo, dentibus 5-7 mm longis), petalis calyce aeqilongis (nee longioribus). 

Plants dwarf, cespitose, 4-12 cm tall, covered with somewhat rigid, predominantly white 
hairs up to 2 mm long. Caudex up to 10 mm in diameter, repeatedly branched, with numerous 
short subterraneous branches. Stems of the year 1-4 cm long, densely covered with sub- 
appressed white, near the nodes also with black hairs up to 0.5 mm long. Stipules mem- 
branous, pale brownish, 4-8 mm long, adnate to the petiole for up to 1 mm, vaginate-connate 
behind the stem up to the midle, the free tips 2-A mm long, linear, acute, appressed white and 
sometimes also black hairy. Leaves 1.5-5 cm long; petiole 0.5-1.5 cm long, like the rachis 
rather densely covered with subappressed to spreading white and some black hairs 0.3-1 mm 
long. Leaflets in 9-12 pairs, elliptic, 4—9 x 1.5-3.5 mm, obtuse to acutish at the apex, densely 
subappressed white hairy on both sides, often folded. Peduncle 0.5-6 cm long, densely 
covered with tousled, ± spreading white hairs, toward the raceme also with increasing black 
hairs 0.2-1 mm long. Raceme ovoid-globose, densely many-flowered. Bracts greenish, 
narrowly ovate, acuminate, 9-13 mm long, rather densely covered with subappressed to 
nearly spreading white and black hairs. Flowers nearly sessile. Calyx infundibuliform-tubular, 
ca. 1 5 mm long, densely covered with ± spreading white hairs and sometimes with few shorter 
black hairs; teeth linear-subulate, 7-9 mm long, hairy on inner side. Petals whitish-creamy 
with hlac keel, in dry state often dirty violet. Standard ca. 1 5 mm long, not surpassing the 
calyx; blade elliptic, ca. 7 mm wide, emarginate at the apex, gradually narrowed into the short 
cuneate claw. Wings ca. 1 3 mm long; blades narrowly oblong, obliquely emarginate at the apex, 
6x2 mm; auricle ca. 1 mm long, claw 7 mm long. Keel 1 1 mm long; blades obliquely obovate, 
with widely curved lower edge and slightly concave upper edge, acutish at the apex, 5 x 2.2 
mm; auricle short, claw 6 mm long. Staminaltube straightly cut. Ovary subsessile, narrowly 
elliptic, appressed white-hairy; style glabrous. Legumes unknown. 



170 



Astragalus tetuanensis Podlech, spec, nov.e sect. Tapinodes 

Holotypus: Morocco, Prov. Tetouan, cerca de Bab Taza, 1000 m, en un pradito de anuales, 
substrato calizo, 30.5.1981, Castroviejo, Fdez. Casas, Muhoz Garmendia & Susanna FC 
5J79(MA;Iso: MA). 

Differt ab A. froedinii Murb. indumento e pilis ad 1,5 mm longis (nee 0,1-0,3 mm) consi- 
stente, foliolis 4-7 (nee 9-14) paribus, bracteis 2-3 (nee 1-1,5) mm longis, vexillo albido (nee 
pallide violaeeo), ab A. depresso L. caulibus pilosis (nee glabris), foliis brevibus subsessilibus 
(nee ad 22 cm longis petiolo bene evoluto), foliolis 4-7 (nee 5-15) paribus, racemo 4-8-floro 
(nee 1 5-25-floro), bracteis 2-3 mm (nee 3-6 mm) longis, ovario glabro (nee piloso). 

Plants 6-15 cm tall, cespitose, covered with rather thick but thin-walled, tousled but 
somewhat rigid, spreading white, only in the inflorescence with few black hairs up to 1.5 mm 
long, at the base of stipules, bracts and bracteoles also with minute sessile, ovoid, gland-like 
hairs. Caudex with a pluricipital rootcrown. Stems several, up to 15 cm long, loosely or in 
upper part rather densely hairy. Stipules greenish-membraneous, 3-4 mm long, obliquely 
inserted at the stem, adnate to the petiole for ca. 1 mm, ciliate at the margin. Leaves 2-3 cm 
long, nearly sessile; rachis sparsely to loosely-hairy. Leaflets in 4-7 pairs, elliptic to obovate- 
triangular, at the apex retuse to distinctly and widely emarginate, 3-5 x 2-3 mm, on upper 
side glabrous, on underside sparsely and mainly at the midrib and margins appressed hairy. 
Peduncle 0.5-3 cm long, rather densely spreadingly hairy. Raceme capitate to subumbellate, 
4-8-flowered. Bracts whitish-membraneous, 2-3 mm long, narrowly triangular, mainly white- 
hairy. Pedicels 1-2 mm long, subglabrous. Flowers at anthesis erect, later on spreading and 
then nodding. Bracteoles rarely present, ca. 1 mm long, laterally at the base of calyx. Calyx 
pale greenish-membraneous, ca. 5 mm long, campanulate, sparsely covered with appressed 
white and some black hairs; teeth narrowly triangular, ca. 2 mm long. Petals whitish. Standard 
ca. 9 mm long; blade obovate, ca. 4 mm wide, at the apex widely incised, at the base cuneately 
narrowed. Wings 8 mm long; blades obovate, at the apex widely rounded, 5 x 2.5 mm; auricle 
ca. 1 mm long, narrow, claw ca. 3 mm long. Keel 7 mm long; blades obliquely elliptic-triangu- 
lar, with in the middle very widely curved lower edge and nearly straight upper edge, acutish 
at the apex, 4 x 2.5 mm; auricle ca. 0.3 mm long, claw 3 mm long. Staminaltube slightly 
obliquely cut. Ovary sessile, glabrous. Pods unknown. 

Astragalus longivexillatus Podlech & Ekici, spec. nov. e sect. Trachycercis 

Holotype: Türkei, BIO Agri, 5 km N Doghubayazit gegen Igdir, 1570 m, Brachland, 
18.5.1990, M Nydegger 45450 (MSB; Iso: BASBG). 

Plantae acaules, pilis medifixis 0,3-0,6 mm longis subappressis obtectae. Stipulae 5-10 mm 
longae, basi petiolo per 1-2 mm adnatae, basi dense apicem versus sparse pilosae. Folia 3,5- 
10 cm longa, petiolo 1,5-4 cm longo, sicut rachis laxe ad densiuscule piloso. Foliola 3-8-paria, 
remota, anguste elliptica, 3-12 x 1-5 mm, supra glabra, subtus sparse, interdum in nervo me- 
diano tantum pilosa. Racemi pedunculo 1-1,5 cm longo suffulti, laxiuscule 2-6-flori. Bracteae 
2-5 mm longae, albi-nigripilosae. Calyx anguste tubulosus, 11-13 mm longus, pilis brevibus 
imprimis nigris albis immixtis obtectus, dentibus subulatis, 2-3,5 mm longis. Corolla flava. 
Vexillum 26-29 mm longum, lamina anguste elliptica, 9-10 mm lata. Alae 25-27 mm longae. 



171 



Carina 20-22 mm longa. Ovarium stipite ca. 1 mm longo suffultum, albi-pilosum. Legumen 
ignotum. 

Plants acaulescent, with medifixed, subappressed hairs 0.3-0.6 mm long. Caudex branched. 
Stipules greenish, 5-10 mm long, the lower ones shorter than the upper ones, oblong-acute, 
adnate to the petiole for 1-2 mm, at the base densely, toward the apex more sparsely hairy. 
Leaves 3.5-10 cm long; petiole 1.5-4 cm long, like the rachis loosely to rather densely hairy. 
Leaflets in 3-8 pairs, remote, narrowly elliptic, 3-12 x 1-5 mm, at the apex rounded to 
obtusish, on upper side glabrous, on underside sparsely hairy or sometimes only hairy at the 
midvein. Peduncle 1-1.5 cm long, densely white, toward the raceme predominantly black 
hairy. Raceme loosely 2-6-flowered. Bracts white-membranous, 2-5 mm long, narrowly 
triangular, white and black hairy. Pedicels 2-3 mm long, black hairy. Calyx narrowly tubular, 
11-13 mm long, slightly obliquely cut at the orifice, loosely to rather densely covered with 
short, predominantly black and some white hairs; teeth subulate, 2-3.5 mm long, with often 
densely ciliate, revolute margins. Petals yellow. Standard 26-29 mm long; blade narrowly 
elliptic, 9-10 mm wide, at the apex slightly emarginate, at the base gradually long narrowed. 
Wings 25-27 mm long; blades narrowly oblong, at the apex acutish, ca. 10 x 2.5-3 mm; auricle 
0.5-0.7 mm long, claw 13-15 mm long. Keel 20-22 mm long; blades oblong-curved, with 
widely curved lower edge and concave to nearly straight upper edge, at the apex acutish, ca. 7 
X 2.5-3 mm; auricle short, claw 14-16 mm long. Staminal-tube straightly cut. Ovary with a 
stipe ca. 1 mm long, white hairy; style glabrous. Legumes unknown. 

Other specimen seen: 

Türkei. B9 Agri : zwischen Agri und Doghubayazit, bei der Abzweigung nach Sulu9am, 1 780 m, 
Brachland, 17.5.1 990, Nydegger 45434 (MSB). 

Oxytropis azerbaijanica Podlech e sect. Janthina 

Holotype: Iran, Prov. Azerbaijan orient.: in jugo inter Marand et Sufian, 1600-1750 m, 
6.6.1971, K//. Rechinger 41162 (M; Iso: W). 

Differt ab ^. karjaganii Grossh. foliolis longioribus, 8-17 (nee 5-10) mm longis, petalis 
longioribus, mucrone carinae longiore, leguminibus distincte curvatis nee oblongis rectis. 

Plant ca. 20 cm tall, acaulescent, covered with thin, mostly spreading white hairs up to 1 .5, 
in fruit up to 2 mm long. Caudex divided, with short branches, densely covered with remnants 
of old leaves. Stipules whitish-hyaline, from narrowly triangular base long linear-acuminate, 
7-10 mm long, adnate to the petiole for ca. 4 mm, rather densely covered with ascending hairs 
up to 0.6 mm long. Leaves 9-13 cm long; petiole 4-7 cm long, like the rachis with a clearly 
double indumentum, densely covered with subappressed, confusely curly hairs 0.1-0.4 mm 
long and with fewer straight, ascending to nearly spreading hairs up to 2 mm long. Leaflets in 
7-9 pairs, narrowly ovate to narrowly elliptic, 8-17 x 3-5 mm, acute at the apex, on both 
sides densely covered with mostly straight, subappressed to slightly ascending hairs up to 2 
mm long. Peduncles 9-10 cm long, erect, hairy like the rachis. Racemes (in fruit) rather loosely 
8-10-flowered. Bracts whitish, linear-acute, ca. 5 mm long, white and black hairy. Pedicels ca. 
2 mm long, white and black hairy. Calyx ca. 10 mm long, campanulate, rather densely covered 
with mostly spreading white hairs up to 1.5 mm long and with shorter, often curly and more 
appressed black hairs; teeth linear, acute, 6-7 mm long. Petals colour unknown. Standard 14- 



172 



15 mni long; blade widely elliptic. 8-9 mm wide, at the apex widely retuse to emarginate, at 
the base subabruptly and shortly narrowed. Wings 8-9 mm long; blades narrowly oblong, 
slightly dilated toward the obliquely emarginate apex, 10-1 1 x 4 mm; auricle wide, ca. 1 mm 
long, claw 3.5—4 mm long. Keel ca. 13 mm long; blades oblong-obovate, with in upper part 
slightly curved lower edge and slightly concave upper edge, 6 (without beak) x 3 mm, at the 
apex obliquely obtuse with a slightly curved beak ca. 2 mm long; auricle ca. 1 mm long, claw 
4-5 mm long. Staminal-tube obliquely cut. Ovar>' shortly stipitate, long white hairy. Legumes 
on a stipe ca. 2 mm long, erect to spreading, oblong, distinctly curved, 15-18 mm long, ca. 5 
mm high and wide, carinate ventrally, very deeply grooved dorsally, at the base nearly 
rounded, at the apex gradually narrowed into a straight beak 1-2 mm long, unilocular; valves 
thin but tough, straw-colored, densely covered with thin, mostly flexuose, ± spreading white 
hairs up to 2 mm long. 



2, New combinations and new names within Astragalus 

Astragalus andreji-sytinii Podlech, nom. nov. - Based on Astragalus eugenii Grossh. in Fl. 

URSS 12: 884. 1946, nom. illeg., non A. eugeniae B.Fedtsch., Hot. Mater. Gerb. Bot. Inst. 

Komarova Akad. Nauk SSSR 8: 166. 1940. 
Astragalus austroiranicus Podlech. nom. nov. - Based on Astragalus minutissimus Freyn & 

Bornm.. Bull. Herb. Boiss. 5: 588. 1897, nom. illeg., non Weddell, Chlor. Andina 2: 257. 

1861. 
Astragalus belangeri (Kuntze) comb. nov. - Basionym: Tragacantha belangen Kuntze, 

Revis. Gen. 2: 940. 1891 = A. kuntzei Sheld., Minnesota Bot. Stud. 1: 123. 1894 h A. te- 

nellus Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 120 et I.e. 15(1): 206. 

1869, nom. illeg.. non Pursh, Fl. Amer. Sept. 2: 473. 1814. 
Astragalus shahbazanicus Podlech, nom. nov. - Based on Astragalus drepanophorus 

Bomm., Repert. Spec. Nov. Regni Veg. 50: 174. 1941, nom. illeg., non Griseb., Abh. K. 

Ges. Wiss. Goettingen 24: 102. 1879. 
Astragalus kirilovii Podlech, nom. nov. - Based on Astragalus intermedius Kar. & Kir., Bull. 

Soc. Imp. Naturalistes Moscou 15: 340. 1842, nom. illeg., non Host, Fl. Austr. 2: 361. 

1831. 
Astragalus mongholicus Bunge var. dahuricus (DC.) Podlech, comb. nov. - Basionym: 

Phaca alpina L. var. dahurica DC, Prodr. 2: 273. 1825 = A. membranaceus Bunge 1869, 

Mem. Acad. Imp. Sei. Saint Petersbourg 15(1): 30. 1869, nom. illeg., non Moench, Meth.: 

167. 1794. 

A. membranaceus. which is either accepted as a species or treated as a synonym of A. 

penduliflorus Lam, is in fact only a variant of A. mongholicus with sparsely hairy ovaries 

and pods. Because of its widely sympatric distribution it is not possibly to accept the two 

variants as subspecies. 

The following two combinations are made to adapt the nomenclature of New Worlds species 
to the subspecies-concept used by most taxonomists in the Old World: 



173 



Astragalus laxmannii Jacq. subsp. robustior (Hook.) Podlech, comb. nov. - Basionym: A. 
adsurgens Pall. var. robustior Hook., Fl. Bor.-Amer. 1: 149. 1831 = A. adsurgens Pall, 
subsp. robustior (Hook.) Welsh, lova State Coll. J. Sei. 37: 357. 1963. 

Astragalus laxmannii Jacq. subsp. viciif alius (Welsh) Podlech, comb. nov. - Basionym: A. 
adsurgens subsp. viciifolius Welsh, Iowa State Coll. J. Sei. 37: 357. 1963 = A. viciifolius 
Hulten, Ark. Bot. 33(B): 1. 1947, nom. illeg. [non^. viciaefolius DC.]. 

Astragalus juratzkanus is a widespread species of Sect. Dissitiflori in the Kopet Dagh range 
and in northeastern Iran. When describing this species Freyn & Sintenis have separated 
within all the cited collections of Sintenis without any sense several specimens as the basis 
for new taxa. So they have described 16 different taxa of this somewhat polymorphic 
species. After examination of all the type material only one species and one slightly 
different subspecies can be recognized. 
To validate the name A. juratzkanus Freyn & Sint. for the species I add all synonyms: 

Astragalus juratzkanus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 567. 1905 

= A. neilreichianus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 571. 1905 

= A. variifolius Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 784. 1905 

- A. variifolius var. homoiophyllus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 786. 1905 

= A. xanthoxiphidium Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 788. 1905 = A. xanthoxiphi- 
dium subsp. accrescens Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 796. 1905 = A. xan- 
thoxiphidium subsp. latifoliolatus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 1012. 1905 

= A. xanthoxiphidium subsp. accrescens var. latilobus Freyn & Sint., Bull. Herb. Boiss., ser. 
2,5:797. 1905 

= A. xanthoxiphidium subsp. campylopus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 797. 
1905 

= A. xanthoxiphidium subsp. curvicaulis Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 1014. 
1905 

= A. xanthoxiphidium subsp. holoxanthus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 1013. 
1905 

= A. xanthoxiphidium subsp. obscurus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 799. 1905 

= A. xanthoxiphidium subsp. rectus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 797. 1905 

= A. xanthoxiphidium subsp. rectus var. elongatus Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 
798. 1905 

= A. xiphidioides Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 786. 1905 

= A. transhyrcanus Popov, Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 10: 10. 
1947 

The following subspecies differs from the type subspecies only by the legumes densely 
covered with asymmetrically medifixed white hairs up to 1.2 mm long, with the longer point 
straight to curved or even hooked and with much fewer distinctly shorter, ± symmetrically 
medifixed black hairs. In all other characters it falls within the variability of the species. 

A. juratzkanus subsp. brotherusii (Freyn & Sint.) Podlech, comb. nov. - Basion.: A. xan- 
thoxiphidium Freyn & Sint. subsp. brotherusii Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 
1015. 1905 

- A. ufraensis Freyn & Sint., Bull. Herb. Boiss., ser. 2, 5: 568. 1905 



174 

References \v ^ «, ^, 

Becht, R. 1978: Revision der Sektion Alopecuroidei DC. der Gattung Astragalus. - Phanerog. 

Monogr. vol. 10. Vaduz. 
Maassoumi, A.A. 1998: New findings on the genus Astragalus in Iran. - Iran. J. Bot. 7: 221- 

226. 
PoDLECH. D. 1988: Revision von Astragalus L. sect. Caprini DC. (Leguminosae). - Mitt. 

Bot. Staatss. München 25: 1-924 
TowNSEND, C.C. & Guest, E. (eds.) 1974: Flora of Iraq, vol. 3. Glasgow. 



Prof. Dr. Dietrich Podlech, Institut für Systematische Botanik der Ludwig-Maximilians- 
Universität München, Menzinger Straße 67, D-80638 München, Deutschland. 



175 



Typification of Astragalus species III. (Leguminosae) 



D. PODLECH 



Abstract: 

PODLECH, D.: Typification of Astragalus species III. (Leguminosae). - Sendtnera 6: 
175-191. 1999. ISSN 0944-0178. 

221 hitherto untypified taxa of the genus Astragalus L. from several herbaria are 
typified here. 

Zusammenfassung: 

221 bisher noch nicht typifizierte Sippen der Gattung Astragalus L. aus verschie- 
denen Herbarien werden hiermit typifiziert. 



In two papers on typification of Astragalus-taxa (Podlech & Sytin 1996; Podlech 1998) 
the bulk of hitherto untypified taxa of the genus Astragalus L. have been typified. In this pa- 
per most of the remaining taxa will be typified with the exception of some species, the types 
of which should be selected from herbaria in the former Soviet-Union (Erevan, Tiflis, Baku, 
Tashkent, Tomsk and others). They can be typified only after visiting the mentioned herbaria. 

A. acantherioceras Rech.f & Kaie, Biol. Skr. 9(3): 147. 1858. - Lectotype (designated here): 
Afghanistan, Shindand, 1 100 m, 26.4.1949, Keie 4450 (W!; Iso: C!). 

A. acrocarpus Freyn & Sint., Österr. Bot. Z. 44: 64. 1 894. - Lectotype (designated here): 
[Turkey] Paphlagonia, Tossia, ad Giaur tschai, 26.6. (flor.), 5.7. (fruct.) 1892, Sintenis 4485 
[the flowering and fruiting specimens were always distributed together under one number] 
(BRNM!: the fruiting plant; Iso: B!, BM!, BP!, BR!, BRNM!: the flowering plants, G!, 
GOET!, HBG!, JE!, K!, LD!, LE!, M!, MSB!, OXF!, P!, PR!, W!, WU!). 

A. afghanomontanus Sirj. & Rech.f, Biol. Skr. 9(3): 157. 1958. - Lectotype (designated 
here): Afghanistan, Deh Kundi, Sar-i-Nil, 7.6.1949, Edelberg 1965 (W!; Iso: C!). 

A. ahouicus Parsa var. pumilus Parsa, Fl. Iran 9: 36. 1966. - Syntypes: Golpayegan, 19.5. 
1957; [Iran, Markasi] Khomayn, 12.5.1957, Parsa 10012. Lectotype (designated here): 
Parsa 10012 (K!). 

A. ajfreidii Aitch. & Baker, J. Linn. Soc, Bot. 19: 157. 1882. - Syntypes: [Afghanistan] at 
the spurs of Mt. Sikaram, 12.6.1880, Aitchison 101; and the Larkarai pass, 10000-12000', 
12.6.1880, Aitchison 373. Lectotype (designated here): Aitchison 101 (K!; Iso: C!, P!). 

A. amarus Pall. 1800, Sp. Astragal.: 8, t. 6. - Lectotype (designated here): Arsagar in deserto 
Caspico, 1793, Pallas (BM!: best congruent with the plate; Iso: BM!, K!, LE!). 



176 



A. amoenus Fenzl, Pug. PI. Nov. Syr.: 4. 1842. - Lectotype (designated here): [Turkey] in 
alpibus Tauri occid., aestate 1836, Kotschy 124 (W!; Iso: B!, BM!, BP!, CGE!, G!, G- 
BOIS!, K!, LE!, M!, MSB!, OXF!, P!, PRC!, W!, ZT!). Note: The indication of a holo- 
type in Fl. Turkey is erroneous. 

A. anacamptoides Sirj. & Rech.f., Biol. Skr. 9(3): 140. 1958. - Lectotype (designated here): 
Afghanistan, Pul-Alam, 7000', 15.6.1937, Koelz 11830 (W!; Iso: C!, E!). 

A. anatolicus Boiss. 

- var. parviflorus Turrill, Kew Bull. 1924: 320. 1924. - Syntypes: Gallipoli, Angadere, 
Biyick Yakajik Tepe, 250 m, 29.4.1923, Ingoldby 29 (K!); dto., Ingoldby 119 (K!); 
Gallipoli, Angadere, 22.-24.7.1923, Ingoldby 443 (K!); Gallipoh, Kilia, 24.4.1924, Durham 
101. Lectotype (designated here): Durham 101 (K!). 

A. ancistrocarpus Boiss. & Hausskn. 

- var. brachycarpus Eig 1955, Syst. Stud. Astrag. Near East: 123. - Syntypes: Iraq, Kirkuk, 
Kani, Domlan hills, 30.4.1933, Guest 4339 & 4363 (K!). Lectotype (designated here): 
Guest 4339 {K\). 

- var. stepposus Eig 1955, Syst. Stud. Astrag. Near East: 122. - Syntypes: N'lraq, N Beled 
Sindjar, 27.4.1933, Eig & Zohary; 30 km W Hassetche, at the foot of Jebel Abd el Aziz, 
28.4.1933, Eig & Zohary; top of Jebel Abd el Aziz, 29.4.1933, Eig & Zohary (HUJ). 
Lectotype (designated here): N'lraq, N Beled Sindjar, 27.4.1933, Eig & Zohary (HUJ!, Iso: 
K!,W!). 

A. andalanicus Boiss. & Hausskn. 1872, in Boissier, Fl. Or. 2: 345. - Lectotype (designated 
here): in monte Andalan supra Sihna Persiae occ, IX. 1867, Haussknecht 25 (G-BOIS!; Iso: 
BM!,K!,W!). 

A. angustatus Boiss. 

- subsp. subtrijugus Bomm., Bull. Herb. Boiss., ser. 2, 5: 843. 1905. - Lectotype (desig- 
nated here): [Iran] inter Rescht et Kaswin, prope Patschinar, 500-600 m, 12.5., Bornmüller 
65/<^(B!;Iso: JE!,LE!). 

A. angustifoliolatus K.T.Fu, Bull. Bot. Res. North-East Forest. Inst. 2(1): 123. 1982. - 
Lectotype (designated here): [China] Yunnan, Lijiang Xian, prope Hortus Botanicus, 2750 
m, 8.8.1962, Zhongdian Exped 613 (PE!; iso: KUN!, PE!). 

A. anserinaefolius Boiss. 

- var. elegantifolius Parsa 1966, Fl. d'Iran 9: 38 - Syntypes: Sirjan, III.-IV.1956, Parsa ? 
20038 K!: foto MSB!; dto., 200/0. Lectotype (designated here): Parsa ? 200/0 (K!). 

A. anthylloides Lam. 

- var. villiger Bomm. 1940, Repert. Spec. Nov. Regni Veg. Beih. 89: 178 - Syntypes: 
Tossia, 24.5.1892, Sintenis 3887 (LD!); Türkei, Paphlagonia, in collibus ad oppidulum 
Koc-hisar (Ilkhas), 1000 m, 23.6.1929, Bornmüller 14092. Lectotype (designated here): 
Bornmüller 14092 (B!; Iso: HBG!, LD!, Z!). 

A. arbelicus Bomm. 1937, Beih. Bot. Centralbl. 57(2): 281. - Syntypes: Kurdistan, E Erbil 
am Kuh-i-Sefm beim Dorf Schaklawa, 1200 m, 21.5.1893, Bornmüller 1181; Umgebung von 
Riwandus, nahe der persischen Grenze auf dem Händarin, 1300 m, 21.6.1893, Bornmüller 
1180 (B!). Lectotype (designated here): Bornmüller 1181 (B!). 

A. arenicola Pomel 1874, Bull. Soc. Sei. Phys. Algerie 11: 184. - Syntypes: Arbaouat, Pomel 
(MPU); Brezina, Pomel (MPU); Mala, Pomel (MPU); Mechaignen, Pomel (MPU). Lecto- 
type (designated here): Arbaouat, Pomel (MPU!). 

A. argolicus Hausskn. 1893, Mitth. Thüring. Bot. Vereins n.s. 5: 81. - Syntypes: in olivetis 
pr. Athenas, Haussknecht; Eleusin, Haussknecht; prope Naupliam, Haussknecht. Lecto- 



177 



type (designated here): M. Palamidi, supra Naupliam, IV. 1885, Haussknecht (JE!; Iso: K!, 
LE!,W!,WU!). 

A. argyroides Beck 1886, Denkschr. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl. 51: 341. - 
Syntypes: prope Manjil ad Bustanak, 9.5.1882, Pichler (W!, WU!); ad Schawerin [Schä- 
berin] prope Hamadan, 12.6.1882, Pichler. Lectotype (designated here): ad Schawerin 
[Schäberin] prope Hamadan, 12.6.1882, Pichler (with fruits) (WU!; Iso: W!). 

A. argyrophyllus Boiss. & Gaill. 1872, in Boissier, Fl. Or. 2: 358. -Syntypes: ad Ouadi el 
Kam Antilibani et in monte Gebel el Kharbe prope Damascum, Gaillardot 47 Obis; dto., 
1764 (G-BOIS!: without flowers); in Syria bor., prope Aintab, 16.6.1865, Haussknecht (G- 
BOIS!). Lectotype (designated here): Gaillardot 470bis (G-BOIS!; Iso: G-BOIS!). 

A. atricapillm Bomm. 1905, Bull. Herb. Boiss., ser. 2, 5: 753. - Lectotype (designated here): 
Elburs occ, supra Häsartschal alpium Tacht-i-Soleiman, 4200 m, 29.6.1902, Bornmüller 
6854 (B!; Iso: BP!, G!, JE!, LE!, ?!, W!). 

A. atrosanguineus Murb. 1922, Acta Univ. Lund., n.s. 18(3): 63. - Lectotype (designated 
here): Grand Atlas ä Amismiz, 1000 m, 1 .5 .\92\ , Murbeck {LD\; Iso: LD!). 

A. australis (L.) Lam. 

- prol. buscesciikv. 1916, Bot. Közlem. 15: 12 - Type: in rupibus calcareis montis Bucsecs 
comit. Brassö Hungariae austro-orientalis. Lectotype (designated here): 2.8.1886, Csato 
(BP!). 

- var. canescens Beauverd 1903, Bull. Herb. Boiss., ser. 2, 3: 456. - Syntypes: Val di Ollo- 
mont alia Balne (Val di Aoste), 2100-2300 m, 28.7.1902, Vaccari (G!); Val de Cogne ä 
Granson, 2300 m (Val d'Aoste), 7.8.1902, Vaccari (with drawing!). Lectotype (designated 
here): Val de Cogne ä Granson, 2300 m (Val d'Aoste), 7.8.1902, Vaccari (with drawing!) 
(G!). 

A. austriacus Jacq. 1762, Enum. Stirp. Vindob.: 263. -Lectotype (designated here): [Austria] 

Brühl, leg. Hayne (W: Hb. Jacquin!) 
A. bachtiaricus Bunge 

- var. leiocarpus Bomm. 1910, Beih. Bot. Centralbl. 27(2): 332. - Syntypes: Sultanabad, m. 
Kuh Sefid Khane, VII. 1903, Strauss (B!); in monte Karagan, ditionis Hamadan, VI. 1902, 
Strauss. Lectotype (designated here): in monte Karagan ditionis Hamadan, VI. 1902, Strauss 
(B!). 

A. bahrakianus Grey- Wilson 1974, Kew Bull. 29: 45. - Lectotype (designated here): Afgha- 
nistan, Wakhan, 2 miles east of the Dahlez Pass, west of Bahrak, ca. 3800 m, 26.6.1971, 
Grey-Wilson & Hewer 1454 (K!, flowering; iso: K!, fmiting). 

A. baldaccii Degen 1896, Österr. Bot. Z. 46: 415. - Lectotype (designated here): in monte 
Nimercka jugo Karajan supra Diovisda distr. Pogoni Albaniae, 8.7.1896, Baldacci 115 
(MSB!; Iso: BM!, BR!, K!, LE!, P!, PR!, PRC!, WU!, Z!). 

A. basianicus Boiss. & Hausskn. 1872, in Boissier, Fl. Or. 2: 361. - Syntypes: in Meso- 
potamia ad Derbent i Basian, V.1867, Haussknecht 33 (344) (G-BOIS!, JE!); et in monte 
Pir Omar Gudrun, 4000', Haussknecht. Lectotype (designated here): in monte Pir Omar 
Gudrun, 4000', VI. 1867, Haussknecht 342 (G-BOIS!; Iso: G-BOISS!, JE!, LE!, W!) 

A. beckii Bomm. 1908, Mitth. Thiiring. Bot. Ver., n.s. 23: 5. - Lectotype (designated here): 
Prov. Gilan, supra Rudbar, 7.5.1902, Bornmüller 6770 (B!: with diagnosis; Iso: B!, BM!, 
BP!, BREM!, E!, G!, HBG!, JE!, K!, LD!, LE!, MSB!, P!, PR!, W!, WU!, Z!, ZT!). 

A. bezudensis Sirj. & Rech.f 1958, Biol. Skr. 9(3): 128. - Lectotype (designated here): Afgha- 
nistan, Bezud, 3000 m, 4.6.1949, Edelberg 1852 (W!; Iso: C!) 



178 



A. bhotanensis Baker 

- var. montigenus Hand.-Mazz. 1933, Symb. Sin. 7: 558. - Lectotype (designated here): 
[China] unter dem Pass Hwayanggo, am Weg von Lidjiang nach Yungning, 3500-3700 m, 
12.7.1915, Handel-Mazetti 7027 (WU!; Iso: E!, W!). 

A. bijugus Sirj. & Rech.f. 1958, Biol. Skr. 9(3): 160. - Lectotype: Afghanistan, Nil Kotal, 

3000 m, 18.8.1948, /:0/e 5227 (W!; Iso: C!). 
A. bornmuelleri ('bornmiilleri') Freyn 1890, Österr. Bot. Z. 40: 403. - Lectotype (designated 

here): Türkei, Amasia, 400-500 m, 13.6.1889, Bornmüller 89 (BRNM!; Iso: B!, BM!, BR!, 

BRNM!,G!,LE!,PRC!, W!). 
A. brachypetalus Traut v. 

- subsp. brachyanthus Freyn & Sint. 1904, Bull. Herb. Boiss., Ser. 2, 4: 449. - Syntypes: 
Kopet Dagh, Suluklü, 4.7.1900, Sintenis 729 (BRNM!, LD!: foto MSB!); et 730a. Lecto- 
type (designated here): Sintenis 730a (LD!; Iso: B!, BP!, BREM!, BRNM!, E!, G!, JE!, K!, 
L!, LE!, MSB!, P!, PR!, PRC!, STU!, WU!, Z!) 

- - var. erythranthm Freyn 1904, Bull. Herb. Boiss., Ser. 2, 4: 450. - Syntypes: Sülüklü, 
2.8.1900, Sintenis 730b; Karakala, in monte Sundsodagh, 2.6.1901, Sintenis 1889 (BP!, 
BRNM!, E!, G!, K!, LD!, LE!, P!, ST!, WU!, Z!). Lectotype (designated here): Sintenis 
730b (BRNM!; Iso: BM!, BP!, BREM!, BRNM!, G!, HBG!, JE!, L!, LD!, LE!, M!, P!, 
PR!, PRC!, UPS!, Z!). 

A. bracteosm Klotzsch 1862, in Klotzsch & Garcke, Bot. Erg. Waldem. Reise: 160, tab. 5, 
nom. illeg. [non Boiss. & Noe]. - Type: Himalaya, Hoffmeister (B: destroyed). Neotype 
(designated here): tab. 5 in Klotzsch & Garcke, Bot. Erg. Waldem. Reise: 160. 

A. brevidens Freyn & Sint. 1904, Bull. Herb. Boiss., Ser. 2, 4: 1 1 18. - Lectotype (designated 
here): Turkmenia, Suluklü, 1.7.1900, Sintenis 738 (BRNM!; Iso: B!, BM!, E!, JE!, K!, LD!, 
LE!,P!,STU!). 

A. campylanthoides Bomm. 1908, Mitth. Thüring. Bot. Ver. n.s. 23: 19. - Lectotype (desig- 
nated here): dit. Hamadan in montibus Karagan, IV. 1902, Strauss (B!; Iso: JE!, LE!). 

A. canaliculatus Willd. 1813, Enum. Hort. Berol. Suppl.: 52. - Syntypes: [cuh. in Hort. 
Berol.] (B- WILLD 13997/1 & 2). Lectotype (designated here): (B-WILLD 13997/1!). 

A. carniolicus A.Kern. 1896, in sched. impr. ad Fl. Austro-Hung. 7: 3. - Lectotype (desig- 
nated here): Camiolia, Nanos, 1250 m, Mulley in Fl. Exs. Austr.-Hung. 2408 (WU!; Iso: B!, 
BR!, BRNM!, C!, G!, H!, K!, L!, LD!, LE!, M!, MSB!, MW, P!, PE!, PR!, REG!, W!, Z!, 
ZT!). 

A. cerasinus Baker 1880, J. Linn. Soc, Bot. 18: 47. - Syntypes: Kurram, Zabardastkalla and 
Alikhel, 18.4.1879, Aitchison 109 (K!, LE!); dto., 30.4.1879, Aitchison 191 (LE!, K!); 
10.6.1879, Aitchison 556; Aitchison, 476 (LE!). Lectotype (designated here): Aitchison 556 
(K!;Iso:LE!) 

A. chamaephaca Freyn 1890, Österr. Bot. Z. 40: 402. - Lectotype (designated here): Amasia, 
prope Khaousa, 350-500 m, 3.5.1889, Bornmüller 99 (BRNM!; Iso: B!, BM!, BP!, BR!, 
E!, G!, GOET!, HBG!, JE!, K!, LD!, M!, MSB!, OXF!, P!, PR!, PRC!, STU!, W!, WU!, 
ZT!). 

A. chlorocarpus Griseb. 1843, Spicil. Fl. Rumel. Bith. 1: 50. - Syntypes: Macedonia boreali 
ad viam inter Komanova et Strazin, Friedrichsthal; m Thracia borali pr. Philippopolin, 
Friwaldsky. Lectotype (designated here): in Thracia boreali pr. Philippopolin [= Plovdiv], 
FnvaW^Aj/ (GOET!; Iso: G-BOIS!). . , •• 



179 



A. chlorotaenius Freyn & Bomm. 1892, Österr. Bot. Z. 42: 12. - Lectotype (designated 
here): Pontus australis, Jatmisch, prope Tokat, 1400 m, 31.5.1890, Bornmüller 2117 (B!; 
Iso: B!, BRNM!). 

A. chorassanicm Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 83 in clave [et 
I.e. 15 (1): 145. 1869]. - Lectotype (designated here): Prov. Chorassan prope Sebsewar et 
Nischapur, VI. 1858, Bunge & Bienert (P; Iso: G-BOIS!: prope Nischapur, K!, LE!). 

A. concretus Benth. 1835, in Royle, Illustr. Bot. Himal. Mount.: 199. - Lectotype (designated 
here): Emodo ad Gossainthan, Wallich Cat. 5931 (K!: Hb. Bentham; Iso: BR!, CGE!, E!, 
G!, K!: Hb. Bentham & Hb. Hooker, LE!, M!). 

A. cottonianus Aitch. & Baker 1888, Trans. Linn. Soc. London, Bot. 3: 51. - Lectotype 
(designated here): Afghanistan, Badghis, 2., 4.5.1885, ^//c/iwo« 377 (Kl; iso: BM!, G!, K!, 
LE!). 

A. craibianus N.D.Simpson 1915, Notes Roy. Bot. Gard. Edinburgh 8: 240. - Lectotype 
(designated here): Western China, 3900 m, VII. 1904, Wilson 3424 (K!; Iso: P!). 

A. cyclophyllon Beck 1886, Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 51: 339. - 
Lectotype (designated here): inter Hamadan et Malajir [am Malaia-Passe inter Hamadan et 
Malaia, 31.5.1882], Pichler (WU!; Iso: JE: fragm.!, Wl, WU!, K!: in monte Elwend Media). 

A. danieli-kochii Freyn 1905, Bull. Herb. Boiss., ser. 2, 5: 558. - Lectotype (designated 
here): in prato Alaiense, 2300 m, 27.6.1898, Paulsen 589a (C!; Iso: BRNM!, LE!). 

A. decumbens Kom. 1914, Repert. Spec. Nov. Regni Veg. 13: 230, nom. illeg. [non A.Gray]. - 
Syntypes: China, prov. Kansu, secus fl. Taoho prope Dsiao-tcheng, 89°50', 28.5.1875, 
Potanin; Prov. Se-tshuan, circa Ta-tsien-lu, 9.7.1893, Potanin. Lectotype (designated here): 
China, prov. Kansu, secus fl. Taoho prope Dsiao-tcheng, 8950', 28.5.1875, Potanin (LE!). 

A. depress us L. 

- subsp. atlantis Maire 1937, Bull. Soc. Hist. Nat. Afr. Nord 28: 352. - Syntypes: Entifa, au 
Dj. Takreda, Ibrahim (P!); Ourika pres le Tizi-n-Chiker, 2000-2300 m, Maire. - Lecto- 
type (designated here): Ourika, entre Timiahi et Chiker, 2000 m, 13.7.1921, Maire (MPU!; 
Iso: P!). 

- var. medius Cuatrec. 1929, Trab. Mus. Ci. Nat. Barcelona 12 [Estudios sobre la Flora y la 
Vegetacion del Macizo de Mägina]: 334. - Syntypes: Mägina Oriental, sobre Pena Gragera, 
1900 m, 1.7.1925, Cuatracasas; Canada del Tejuelo sobre Cano del Aguadero, 1800 m, 
25.6.1926, Cuatrecasas (MA!); Mägina occidental, en Penade Jaen, 2000 m, 4.7.1926, 
Cuatrecasas; torcales superiores deMägina, 2000-2100 m, 26.6.1926, Cuatrecasas; 
Carades, vert. NW, 1900-2000 m, 17.6.1926 Cuatrecasas. Lectotype (designated here): 
Torcales superiores de Mägina, 2000-2100 m, 26.6.1926, Cuatrecasas (MA!). 

A. dichroanthus Freyn & Sint. 1892, Österr. Bot. Z. 42: 13. - Lectotype (designated here): 

Armenia turcica, Keban-Maden in montosis ad Denislii, 21.6.1889, Sintenis 819 (BRNM!; 

Iso: LD!). 
A. distinctissimus Rech.f & Edelb. 1958, Biol. Skr. 9(3): 86, nom. illeg. [non Eig]. - Lectotype 

(designated here): Afghanistan, Zentral-Nord-Gebirge, VII. 1953, Edelberg 2310 (C!; Iso: 

W!: fragm.). 
A. dolichopodus Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 1120. - Lectotype (designated 

here): Pamir, prope lac. Jashil Kul, 3800 m, 22.7.1898, Paulsen 866 (BRNM!; Iso: 

BRNM!, C!, LE!). 
A. durandianus Aitch. & Baker 1888, Trans. Linn. Soc. London, Bot. 3: 54. - Lectotype 

(designated here): Khorasan, on the pass to the south of Bezd, 6000', 18.6.1885, Aitchison 

688 (K!; Iso: BM!, C!, G!, LE!, W!). 



180 



A. echinatus Murray 1770, Hort. Bot. Stirp.: 222. - Type: non indicatus; Neotype (desig- 
nated here): [misit] Gouan (Linn. 926.33, LINN). 
A. edinburghensis Ponert 1973, Feddes Repert. 83: 632. - Lectotype (designated here): 

Turkey, Hakkari, Zap Gorge, 15 km from Hakkari to Cukurca, 1200 m, Davis 44793 (E!; 

Iso: K!). 
A. elegans Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 55 [et I.e. 15(1): 89. 

1869]. - Syntypes: Persia, Adserbidshan prope Tabris, Szovits 362 (G-BOIS!, P!); Sseid- 

abbad baud procul ab urbe Tabris, Bunge. Lectotype (designated here): Sseid-abbad baud 

procul ab urbe Tabris, Bunge (P!; Iso: G-BOIS!). 
A. enantiotrichus Freyn 1905, Bull. Herb. Boiss., ser. 2, 5: 561. - Syntypes: Pamir, prope 

lacum Jashil Kul, 3800 m, 21.7.1901, Paulsen 849 (C!, LE!); dto., 13.8.1898, Paulsen 1109. 

Lectotype: Pamir, Jashil Kul, 3800 m, 13.8.1898, Paulsen 1109 (C!; Iso: BRNM!, LE!). 
A. euphraticus Freyn 1895, Bull. Herb. Boiss. 3: 186. - Lectotype (designated here): Armenia 

turcica, Egin, ad Euphratem, V.1890, Sintenis 2296 (BRNM!; Iso: B!, E!, K!, LD!, P!, PR!, 

PRC!,W!,WU!). 
A. exscapus L. 

- f. scaposus Beck 1900, in Rchb., Icon. Fl. Germ. Helv. 22: 105, tab. 156 II. - Lectotype 
(Iconotype) (designated here): plate in Reichenbach, I.e. tab. 156 II. 

A. fialae Degen 1900, Österr. Bot. Z. 50: 242. - Syntypes: Treskavica Planina Bosniae, 
27.7.1893, Fiala; in Cemagorae mont. Dibala supra Kat. Kostica distr. Kuci, 10.7.1898, 
Baldacci 188 {U\, PR!, PRC!, W!, WU!). Lectotype (designated here): Treskavica Planina 
Bosniae, 27.7.1893, Fiala (WU!). 

A. flavescens Boiss. 1843, Diagn. pi. orient., ser. 1, 2: 64. - Syntypes: in reg. alpina Tmoli 
[Tmoleus circa Bogdagh], VII. 1842, Boissier; supra Philadelphiam, Boissier; et supra 
Sardes loco Bozdagh, Boissier (G-BOIS!, M!). Lectotype (designated here): in reg. alpina 
Tmoh, VII.1842, Boissier (G-BOIS!; Iso: G!, G-BOIS!, GOET!, HBG!, JE!, K!, LIV!, 
MSB!,P!,TUB!, W!,ZT!). 

A. ßexirachis Rech.f. & Edelb. 1958, Biol. Skr. 9(3): 89. - Lectotype (designated here): 
Afghanistan, Deh Kundi, Sar-i-Nil, 3000 m, 7.6.1949, Edelberg 1946 (W!; Iso: C!). 

A /orre^/ii N.D.Simpson 1913, Notes Roy. Bot. Gard. Edinburgh 7: 124. - Lectotype (desig- 
nated here): western flanc of the Lichiang range, Yunnan, lat. 27°25' N, 11000', VI. 1910, 
Forrest 6041 (E!; Iso: BM!, K!, W!: fragm.). 

- var. minor H.T.Tsai & T.T.Yu 1940, Bull. Fan Mem. Inst. Biol., Bot. Ser. 9: 260. - 
Lectotypus (designated here): S.Sikang, Shiancheng, Tungzung to Tungwah, 3000 m, 
20.9.1937, T.T.Yu 13401 (KUN!; Iso: KUN!, PE!). 

A. fuliginosus Beck 1886, Denkschr. Kaiserl. Akad. Wiss. Math.-Naturwiss. Kl. 51: 340. - 

Lectotype (designated here): ad vias versus Merdjil [Manschil], 26.4.1882, Pichler (WU!; 

Iso: JE!: fragm., K!, W!, WU!). 
A. gerntanicopolitanus Bomm. 1931, Magyar Bot. Lapok 30: 64. - Syntypes: Paphlagonia 

australis, in valle Cakmakli-dere prope Cankri (Germanicopolis olim), 800-900 m, 

6.6.1929, Bornmüller 14018; et 3.7.1929, Bornmüller 14098. Lectotype (designated here): 

Bornmüller 14098 (B!; Iso: ZT!). 
A. glaucophyllodes Bomm. & Woronov 1914, Vestn. Tiflissk. Bot. Sada 34: 2. - Lectotype 

(designated here): Prov. Kars, distr. Kaghysman, prope Zarab-chan, 3.6.1913, Woronow 

725/J(B!;Iso:LE!). ■ 



A. glycyphyllus L. 

- f. bosniacus Beck 1896, Ann. K.K. Naturhist. Hofmus. 11: 75. - Syntypes: bei Visoko, 
Formanek; urn Gorazda, Formanek; dto.. Beck; am Kmur bei Foca, For mane k; Prokosko 
jezero auf der Vranica Planina, VIII. 1894, Schwarz (PRC!); Herzegovina, auf der Ivan 
Planina, Sündermann; Vrabacbei Konjica, Formanek; Montenegro, bei Skrobotusa, 
Szyszylovicz. Lectotype (designated here): in silva Skrobotusa, 2.8.1886, Szyszylowicz (W!). 

A. goeznensis Eig 1955, Syst. Stud. Astrag. Near East: 38. - Syntypes: Türkei, Goezne (NW 
Mersina), 1000-1100 m, 16.8.1931, Eig & Zohary; dto., 1200-1300 m, 17.8.1931, Eig & 
Zohary. Lectotype (designated here): Goezne (NW Mersina, 1200-1300 m, 17.8.1931, Eig 
& Zohary (UV J;lso: El, Wl). 

A. gombaultii Eig 1955, Syst. Stud. Astrag. Near East: 127. - Syntypes: N' Lebanon, Jebel 
Sannin, 2000 m, 20.7.1930, Gombault 465; ibidem, 1.6.1879, Peyron [957] (G!); ibid., 
6.7.1897, Bornmüller; ibid., 19.7.1935, Mouterde; S' Lebanon, Tom Niha, 13.3.1877, Ball. 
Lectotype (designated here): Gombault 465 (P!). 

A. granatensis Lange 1865, Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1865: 177 
[et in Pugillus: 372], nom. illeg. [non Lam.]. - Syntypes: Sierra Elvira fl. granat., 24.4.1852, 
Lange (C!); et Sierra de Alfacar prope Granada, 27.4.1851, Lange (C!, MSB!, P!); Guejar 
ad rupes (Sierra Nevada), 16.4.1852, Lange. Lectotype (designated here): Guejar ad rupes 
(Sierra Nevada), 16.4.1852, Lange (C!; Iso: C!). 

A. grandiflorus Freyn 1892, Österr. Bot. Z. 42: 46, nom. illeg. [non L. nee Bunge]. - Lecto- 
type (designated here): Armenia turcica, Kharput, ad Schuschnas, 8.6.1889, Sintenis 720 
(BRNM!; Iso: B!, BR!, JE!, LD!, P!). 

A. grisebachianus Aitch. & Baker 1888, Trans. Linn. Soc. London, Bot. 3: 52. - Syntypes: 
Afghanistan, Harirud valley, 10.5.1885; et 6.6.1885, Aitchison 419 (LE!). Lectotype 
(Becht 1978): Harirud valley, 10.5.1885 et 6.6.1885, Aitchison 419 (K!; Iso: BM!, G!): 
Lectotype (designated here): Harirud valley, 10.5.1885, (K!; Iso: K!). Note: The both 
collections are sometimes distributed together (BM!, G!, K!. The so-called lectotype by 
Becht 1978 is such a mixed sheet. 

A. hamulosus H.Lev. 1914-15, Fl. Kouy-Tcheou: 226. - Syntypes: [China, prov. Guizhou] 
Hini-Fou, ruisseaux riziere, 20.4.1897, Bodinier 1580; Tin Lan, Cavalerre (E!, MSB!, P!). 
Lectotype (designated here): Bodinier 1580 (P!; Iso: E!, MSB!). 

A. hancockii 1880, Bull. Acad. Imp. Sei. Saint-Petersbourg 26: 471. - Syntypes: China, 
[Prov. Hubei] prov. Petschili, m. Siao-wu-tai-shan, 3600-5000', 1876, Hancock (LE!); 
China, prov. Petschili, m. Siao-wu-tai-shan, Aufstieg vom Kloster Tieh lin sze, VI. 1879, 
Möllendorf. Lectotype (designated here): China, prov. Petschili, m. Siao-wu-tai-shan, 
Aufstieg vom Kloster Tieh lin sze, 3600-5000', VI.1879, Möllendorf (LEI; Iso: LE!, PE!). 

A. hedinii Ulbr. 1905, Bot. Jahrb. Syst. 35: 679. - Lectotype (designated here): locality on 
the sheet not to read; according to Ostenfeld & Paulsen: Northern Tibet, Mandarlik, 
VII. 1900, Hedin (C!; the original material at Beriin is lost). 

A. henryi Oliv. 1891, in Hooker's Icon. PI. 20: t. 1959. - Lectotype (designated here): China, 
Prov. Hupeh, Fang District, Henry 6902 (K!; Iso: BM, E!, K!, LE!, P!) 

A. hirsutus Vahl 

- subsp. paraglobosus Ponert 1973, Feddes Repert. 83: 631. Lectotype (designated here): 
[Turkey] Kayseri, Bakir Da. above Kisge, P. Davis 19340 (E!; Iso: K!) 

- var. hispidus Freyn & Bomm. 1892, Österr. Bot. Z. 42: 48. - Syntypes: Tokat, 800-900 
m, 13.5.1889, Bornmüller 106; Sivas, in monte Tschamlii-bel supra Yeni-chan, 1500 m, 
1.6.1890, Bornmüller (Z!); Amasia, in monte Akdagh, 1200-1900 m, 22.5.1890, Born- 



182 



müller 2102 (B!); Pontus galaticus, in mte. Abadschi Dag, 1500 m, 14.5.1890, Bornmüller 
2103 (B!). Lectotype (designated here): Bornmüller 106 (B!; Iso: BRNU!, G!, JE!, LE!, 
OXF!) 

A. holdichianus Aitch. & Baker 1888, Trans. Linn. Sog. London, Bot. 3: 54. - Syntypes: 
Afghanistan, Badghis, 29.4.1885 (fl.), Aitchison; 19.5.1885, Aitchison 347. Lectotype 
(designated here): Aitchision 347 (K!: fruiting plant; Iso: BM, C!, G!, K!, LE!, P!, W: 
fragm.!). 

A. hololeios Bornm. 1934, Bot. Jahrb. Syst. 66: 226. - Lectotype (designated here): Kabul, in 
monte Babur, 2000 m, 1928-1929, Manger (JE!; Iso: B!). 

A. humillimus Freyn 1894, Österr. Bot. Z. 44: 61, nom. illeg. [non A.Gray]. - Syntypes: 
Paphlagonia, Kastemuni, ad Taschly Chan, 6.5.1892, Sintenis 3853 (B!, BRNM!, LD!); 
Tossia, prope Tossia Dewrendi in jugo Kutschuk-Ilkas-Dagh, 4.6.1892, Sintenis 4127\ ad 
Schakylar, 16.7.1892, Sintenis 4688 (B!, BRNM!, JE!, LD!, MSB!). Lectotype (designated 
here): Sintenis 4127 (BRNM!; Iso: B!: erroneously 4124, BP!, BR!, FR!, G!, GOET!, 
HBG!, K!, L!, LD!, M!, MSB!, OXF!, P!, PR!, STU!, W!, WU!, Z!). 

A. huthianus Freyn & Bornm. 1898, Bull. Herb. Boiss. 6: 605. - Lectotype (designated here): 
Prov. Kerman, mont. Kuh Laiesar prope Laiesar, 2900-3000 m, 23.7.1892, Bornmüller 
5759 (B!; Iso: BRNM!). 

A. ibrahimianus Maire 1922, Bull. Soc. Hist. Nat. Afr. Nord 13: 38 - Syntypes: Djebel 
Touchka, montagne au sud de la ville de Maroc [= Marrakech], 1 1.6.1875, Ibrahim; Djebel 
Ouensa, montagne au sud de la ville de Maroc, VI. 1874, Ibrahim (BREM!, K!, LE!, P!); in 
jugo Tagherat ditionis Reraya, 3300-3500 m, 22.7.1922, Maire (LD!, MPU!, P!); in monte 
Tachdirt, 3200 m, 11.7.1921, Maire (MPU!, P!); in jugo Chiker ditionis Ourika, 2400 m, 
\4.7.\92\, Maire (MPU!, P!). Lectotype (designated here): Djebel Touchka, montagne au 
sud de la ville de Maroc [= Marrakech], 11.6.1875, Ibrahim (MPU!; Iso: BP!, BR!, 
GOET!, K!, LE!, MSB!, PRC!, WU!). 

A. jagnobicus Lipsky 

- var. shtivensis Sirj. & Rech.f. 1958, Biol. Skr. 9(3): 52. - Syntypes: [Afghanistan] Nuristan, 
Shtive, 2600 m, 16.6.1948, Edelberg 1033 (C!, W!); Weran, Nau, 3000 m, 21.7.1948, 
Edelberg 1474; Netschingel, 15.7.1949, Edelberg 2001 (C!, W!). Lectotype (designated 
here): Edelberg 1474 (W!; Iso: C!). 

A. juratzkanus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 567. - Lectotype (designated 

here): Turkmenia, Kisil-Arwat, 3.5.1901, Sintenis 1623a (BRNM!; Iso: JE!, LD!, LE!). 
A. kialensis N.D.Simpson 1915, Notes Roy. Bot. Gard. Edinburgh 8: 242. - Lectotype 

(designated here): [China] Szechuen, Tongolo (Principaute de Kiala), Ta-tsien-lu, 

10.8.1893, Soulie 332 (K!; Iso: G!, MSB!, P!). 
A. kourosianus Parsa 1947, Kew Bull. 1947: 19. - Lectotype (designated here): Hormud, 

Fars, 5.4.1939, Parsa ? 670 (K!; Iso: TEH, fide Maassoumi: pers. comm.). 
A. krajinae Domin 1931, Acta Bot. Bohem. 10: 48 [et in sched. impr. ad Fl. Chechosl. exs. no. 

270]. - Lectotype (designated here): Rossia subcarpatica orientali-septentrionalis, montes 

Svidovec, ad rupes Dragobrat, 1650-1700 m, 11.7.1931, M.Deyl in Fl. Chechosl. exsicc. 

270 (PRC!; Iso: B!, BP!, H!, HBG!, K!, LD!, LE!, MSB!, PRC!, W!). 
A. lanatus Labill. 

- var. denniensis Gombault 1953, Bull. Soc. Bot. France 100: 331 - Lectotype (P!; Iso: P! 
A. laüstylus Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 1106. - Lectotype (designated here): 

Pamir, prope lacus Jashil-Kul, 3800 m, Paulsen 814 (BRNM!; Iso: C!, LE!). 



im 



- subsp. aridus Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 1107. - Lectotype (designated 
here): Pamir, prope lacum Jashil-Kul, 3800 m, 22.7.1898, Paulsen 865 (BRNM!; Iso: C!, 
LE!). 

A. leucothrix Freyn & Bomm. 1891, Österr. Bot. Z. 41: 406. - Lectotype (designated here): 

Pontus australis, Amasia, ex gr. Logman et Kirklar ad arcem, 600-800 m, 28.5.1890, 

Bornmüller 1840 (B!; Iso: B, BRNM!, G!, JE!, W!, WU!). 
A. linearifolius Pers. 1807, Syn. 2: 336. - Type (In LD no specimen of Persoon). Neotype 

(designated here): prope Rudo ad fluv. Lim, 370-400 m, VI. 1910, Maly in herb. norm. ed. 

Dörfler 5254 (M!; Iso: B!, PR!, W!). 
A. lipskyanus Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 755. - Syntypes: Pamir, ad lacus 

Jashil-Kul, 28.7.1898, Paulsen 977 (BRNM!, C!; et ibidem, 8.8.1898, Paulsen 1075. 

Lectotype (designated here): Paulsen 1075 (C\). 
A. longicuspis Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 121 in clave [et 

I.e. 15(1): 211. 1869]. - Lectotype (designated here): ad urbem Urmiah Prov. 

Aderbeidschan, 10.4.1828, Szovits 68 (LE!; Iso: G-BOIS!, H!, K!: teste Bunge, LE!). 
A. longidens Freyn 1895, Bull. Herb. Boiss. 3: 181. - Lectotype (designated here): Armenia 

turcica, Egin, ad Ischek-Meden, 18.6.1890, Sintenis 2658 (BRNM!; Iso: B!, E!, G!, K!, 

LD!,MSB!,P!, WU!). 
A. longilobus E.Peter 1937, Acta Horti Gothob. 12: 47. - Lectotype (designated here): Kansu 

austr.occ, upper Tebbu country, slopes at foot of Shimen, 12000', July-Aug. 1925, Rock 

7i075(E!;Iso:,K!,P!,PE!). 
A. luteolus H.T.Tsai & T.T.Yu 1936, Bull. Fan Mem. Inst. Biol., Peiping, Bot. Ser. 7: 23. - 

Lectotype (designated here): NW.Szechuan, Sung-pan hsien, 21.8.1928, W.P.Fang 4313 

(PE!: with flowers; Iso: E!, K!, P!, PE!: with fruits). 
A. lycius Boiss. 

- f acutiflorus Hub.-Mor. 1940, Repert. Spec. Nov. Regni Veg. 48: 284. - Lectotype 
(designated here): Vilajet Burdur in Pisidien, Dirmil-Tefenni, 1250 m, 9.6.1938, Reese. Renz 
& Huber 5531 (G!; Iso: G!). Note: Homotypic with A. mesogitanus Boiss. f glabrifolius 
Hub.-Mor. 

A. macedonicus Heldr. & Charrel 1892, in schedis impr. ad Heldreich Herb. Graec. Norm. no. 

1136. - Lectotype (designated here): Macedonia, in m. Korthiati prope Thessalonicam, 

4000', 19.5.1891, Charrel in Heldreich Herb. Graec. Norm. 1136 (MSB!; Iso: B!, BRNM!, 

G!, JE!, K!, LD!, LE!, MPU!, PR!, PRC!, P!, STU!, W!, Z!) [in WU no specimen!]. 
A. macropterus DC. 1825, Prodr. 2: 283. - Syntypes: In Dahuria et Sibiria Scythica, Schangin 

(G-DC!, LE!); ex Dahuria misit Fischer. Lectotype (designated here): ex Dahuria misit 

Fischer (leg. Pausner) (G-DC!; Iso: LE!). 
A. macrostegius Rech.f 1958, Biol. Skr. 9(3): 37. - Lectotype (designated here): Nuristan, 

Minjan pass, 12000', 26.7.1937, Koelz 12716 (W!; Iso: US). 
A. maculatus Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 119 [et I.e. 15(1): 

206. 1869], nom. illeg. [non Lam.]. - Lectotype (designated here): inter Nischapur et 

Meschhed, 1858, Bunge & Bienert (LE!; Iso: G-BOIS!). 
A. maddenianus Baker 1876, in Hooker f., Fl. Brit. India 2: 127. - Syntypes: Kumaon, 

\QQQQ-\\QQQi\ Madden, Edgeworth. - Lectotype (designated here): Himalaya, 10-11000', 

Badrinath, Madden (K!). 
A. medullaris Boiss. 1849, Diagn. pi. orient., ser. 1, 9: 66. - Lectotype (designated here): in 

regno Cabulico, Griffith 1061 (K: Hb. Hooker!; Iso: G!, G-BOIS!, K!: Hb. Hooker) - distr. 

no. 75i2(C!,CGE!,L!,ZT!). 



184 



A. mendax Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 770. - Lectotype (designated here): 
Olgin-Lug in mont. Alai, 2600 m, 20.6.1898, Paulsen 438 (BRNM!; C!, LE!: fragm.). 

A. microcalycinus Siij. & Rech.f. 1958, Biol. Skr. 9(3): 163. - Lectotype (designated here): 
Afghanistan, Kabul, 6000', 15.5.1937, Koelz 11456 (W!; Iso: US). 

A. minutissimus Freyn & Bomm. 1897, Bull. Herb. Boiss. 5: 588, nom. illeg. [non Weddell]. - 
Syntypes: Persia, Kerman, Kuh-i-Häsar (inter Kerman et Bandar-Abbas) prope pag. 
Tschähasar-tagh, 3000 m, 8.8.1892, Bommüller 3762a (B!) [Bornmüller 3762 with exactly 
the same text of 3762a on label in B!, BRNM!, G!, K!, LE!, WU!]; mt. Kuh-Lalesar prope 
pag. Lalesar, 3000 m, 12.7.1892, Bornmüller 3761. Lectotype (designated here): Born- 
müller 3761 (BRNM!; Iso: B!). 

A. moellendorfii Bunge 

- var. kansuensis E.Peter 1937, Acta Horti Gothob. 12: 50. - Lectotype (designated here): 
[China] Kansu centr., Lien-hwa-shan [between Taochow and Titao, 11500 ft., 14.- 
20.7.1925, Rock 12718 (E!; Iso: K!, W!). 

A. muelleri Steud. & Höchst. 1827, Flora 10: 72. - Lectotype (designated here): Ins. Veglia, 

V.1826 Müller (TUB: herb. Hochstetter! as/i. mülleri nobis; Iso: K!, W!). 
A. ntuliensis Hand.-Mazz. 1933, Symb. Sin., Pt. 7: 554. - Lectotype (designated here): China, 

Prov. Setschwan, Tschako SW Muli, auf dem gegen Dschungdien führenden Rücken, 3950 

m, 5.8.1915, Handel-Mazzetti 7454 (WU!; Iso: E!, W!). 
A. murrü Huter 1889 in schedis imprim. ad exsicc. ed. Baenitz [et in Österr. Bot. Z. 55: 29. 

1905]. - Lectotype (designated here): Tirolia septentrionalis ad "Neuen Schönberg", 

VI. 1889, Murr, in exsicc. ed. Baenitz, (W!; Iso: BP!, BRNM!, GOET!, HBG!, JE!, K!, L!, 

PR!, WU!,ZT!). 
A. nanellus H.T.Tsai & T.T.Yu 1936, Bull. Fan Mem. Inst. Biol., Peiping, Bot. Ser. 7: 25. - 

Lectotype (designated here): Western Szechuan, Fu~pien hsien, 3200 m, 18.6.1930, 

F.T. Wang 21356 (PE!; Iso: PE!, W!: fragm.). 
A. narbonensis Gouan 

- subsp. atlatiticus Ball 1873, J. Bot. 11: 306. - Lectotype (designated here): in regione 
inferiori Atlantis Majoris, in convalle Ait Mesan, ca. 1200 m, Ball (K!; Iso: BM!, G!, K!, 
LE!). 

- var. glabrescens Coss. 1852, Not. PI. Crit.: 161. - Syntypes: Sierra Nevada, loco dicto 
Cortijo de la Vibora, VII. 1851, Bourgeau 1143; in provincia Giennensi (province de Jaen) 
prope Trusala, 1851, Blanco 255. Lectotype (designated here): Bourgeau 1143 (P!; Iso: 
GOET!, P!). 

A. neilreichianus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 571. - Lectotype (desig- 
nated here): Turcomania, Kisil Arwat, 3.5.1901, Sintenis 1623b (BRNM!; Iso: JE!, LD!, 
LE!). 

A. nigritus Sirj. & Rech.f 1958, Biol. Skr. 9(3): 43. - Lectotype (designated here): 
Afghanistan, Farakulum, 2800 m, 19.7.1948, Köie 2569 (W!; Iso: C!). 

A. nitidulus Hand.-Mazz. 1913, Ann. Nat. Hofmus. Wien 27: 78. - Lectotype (designated 
here): Rücken EI Hilu zwischen Sabcha und Tibne ober Der et Sor am Euphrat [inter 
Meskene et Der-ez-Sor], 250-350 m, 29.3.1910. Handel-Mazzetti 545 (WU!; Iso: B!: 
fragm., W!). 

A. «ovi« Winterl 1788, Index Hort. Bot. Pest: 15, fig. 13. - Type: not indicated. Lectotype 
(Iconotype) (designated here): Fig 13 in Index Hort. Bot. Pest. 15. 



185 



A. nuristanicus Sirj. & Rech.f. 

- var. elasoonensis Sirj. &, Rech.f. 1958, Biol. Skr. 9(3): 79. - Type: Nuristan, Elasoon, 

Edelberg [s.n.]. Lectotype (designated here): [Afghanistan] Elasoon, 7.7.1949, Edelberg 

] 746 iW\;lso:C\). 
A. olufsenii Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 450. - Syntypes: Sufi-Kurgan in mt. 

Alai, 18.6.1898, Paulsen 421 (BRNM!, C!); in prato Alaiensi, 27.6.1898, Paulsen 589b 

(C!); Pamir, ad lacus Jashil-Kul, 3800 m, 28.7.1898, Paulsen 973 (BRNM!, C!); prope lac. 

Bulung Kul, 3800 m, 25.8.1898, Paulsen 1168. Lectotype (designated here): Paulsen 1168 

(BRNM!; Iso: C!). 
A. oophorus Freyn 1905, Bull. Herb. Boiss., ser. 2, 5: 1019. - Lectotype (designated here): 

Pamir, Jashil-Kul, 3800 m, 4.8.1898, Paulsen 1026 (BRNM!; Iso: C!, LE!). 
A. orophacoides Freyn 1905, Bull. Herb. Boiss., ser. 2, 5: 563. - Syntypes: Pamir, ad Sary 

Mullah, 4100 m, 5.7.1898, Paulsen 681 (C!, BRNM!); Pamir, ad Shatshan prope fl. 

Murghab, 3800 m, 11.7.1898, Paulsen 730. Lectotype (designated here): Paulsen 730 

(BRNM!; Iso: C!,LE!). 
A. orthanthus Freyn 1905, Bull. Herb. Boiss., ser. 2, 5: 1018, nom. illeg. [non A.Gray]. - 

Syntypes: Pamir, prope fl. Kara-Su, 12.7.1898, Paulsen 754 (C!, LE!); Pamir, lac. Jashil 

Kul, 13.8.1898, Paulsen 1103. Lectotype (designated here): Paulsen 1103 (BRNM!; Iso: 

C!,LE!). 
A. orthocarpus Boiss. 1849, Diagn. pi. orient., ser. 1, 9: 68. - Lectotype (designated here): 

[Afghanistan] in regno Cabulico, Griffith 1060 = distr. no. 1520 (K: Hb. Hooker! Iso: K!, 

P!)- 
A. orthodontus Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 11(16): 101 [et I.e. 

15(1): 176. 1869]. - Lectotype (designated here): [Turkey] Cilicia or., monte Jool-dagh 

prope Baatsch, 4800', Kotschy 233 (P!; Iso: G-BOIS!, K!, L!, MSB!). 
A. paghmanensis Sirj. & Rech.f. 1958, Biol. Skr. 9(3): 100. - Lectotype (designated here): 

Afghanistan, Paghman, 7000 ft., 12.5.1937, Koelz 11390 (W!; Iso: US). 
A. panjaoensis Sirj. & Rech.f 1958, Biol. Skr. 9(3): 154. - Lectotype (designated here): 

Afghanistan, Panjao, 2700 m, 1.8.1948, Koie 2748 (W!; Iso: C!). 
A. paphlagonicus Freyn & Sint. 1894, Österr. Bot. Z. 44: 65, nom. illeg. [non Bunge]. - 

Syntypes: Paphlagonia ad Tossia, prope Kawak-Tscheschme, 7.6.1892, Sintenis 4126 

[without fruits] (B!, BP!, BRNM!, JE!, LD!, P!, PR!, WU!); ad Hadschi Ahmed, 

17.6.1892, Sintenis 4126b [with fruits]. Lectotype (designated here): Sintenis 4126b 

(BRNM!; Iso: G!, LD!, PR!, PRC!, W!, Z!). 
A. patulepilosus Sirj. & Rech.f 1958, Biol. Skr. 9(3): 166. - Lectotype (designated here): 

Afghanistan, Shindand, 1400 m, 1949, Koie 4331 (W!; Iso: C!). 
A. peterfii Jav. 1916, in Sched. Fl. Hung. Exsicc. 4: 38. - Lectotype (designated here): supra 

pag. Magyarszovät, versus pag. Olahgyeres, comit. Kolozs Hungariae orientalis, 350^00 

m, 15.5. et 1.6.1916 fl., 30.7.1916 fr., Peterfi (BP!: the specimen signed as lectotype; Iso: 

BP! et in Fl. Hung. exs. 363: BP!, BR!, BRNU!, C!, H!, K!, L!, LD!, LE!, MSB!, PE!, PR!, 

PRC!, W!,Z!,ZT!). 
A.petroviciiVQ\Qn. 1903, Sitzungsber. Königl. Böhm. Ges. Wiss. Prag, Math.-Naturwiss. CI. 

Jahrg. 1903(28): 4. - Lectotype (designated here): Serbia ad Nish in m. Suha planina, 

VII. 1887, Bornmüller (PRC!; Iso: B!, MSB!, W!). 
A. pityusarum Bornm. 1931, Magyar Bot. Lap. 30: 62. - Lectotype (designated here): 

Bithynia, Vesir-han, in monte Tepaja, 300-400 m, 25.5.1929, Bornmüller 14099 (B!; Iso: 

B!, BM!, BP!, G!, HBG!, K!, LD!, LE!, MSB!, P!, PR!, W!, Z!). 



186 



A. podocarpus C.A.Mey. 

- var. leucotrichus Bomm. & Gauba 1935, Repert. Spec. Nov. Regni Veget. 39: 100. - 
Lectotype (designated here): Elburs, Keredj Tal bei Sirah, 2700 m, 31.8.1935, Gauba 625 
(B!subno49ö;Iso:B!,W!). .. ^ • / ■ ^^■-'^'' 

A. podolobus Boiss. & Hohen. ^vK '^^■^^^j^x-^'iJ'^"''''^ 

- f. albovillosus Sirj. 8c Rech.f. J^l, Sitzungber. Öserr. Akad. Wiss., Mathem.-Naturw. KL, 
Abt. I, 170:50.- Lectotype (designated here): Khorasan, Montes Hazar Masdjid, inter 
Gash et Tolgor, 7.-10.6.1948, Rech.f. 5145b (W; Iso: B!, E!, G!, G-Aellen!, K!). ^ /^ \ 

A. polychromus Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 454. - Lectotype (designated here): 

Pamir, ad flum. Muskol, 4300 m, 2.6.1898, Paulsen 665 (C!; Iso: BRNM!, LE!). 
A. praecox Baumg. 1816, Enum. stirp. Transsilv. 2: 362. - Syntypes: juxta Kolosvar; Deva; 

Karoly-Fejervar; sz. Ersebeth; Keresd; Segesvar; Prodt; Berethalom; et ad Szasz-Kesd; 

Köhalom. Lectotype (designated here): in herbidis montosis saxosis ad Scycsvoro [= 

Segesvar] versus Wziter Berg, IIL1809, Baumgarten (REG!). 
A. prattii N.D.Simpson 1915, Notes Roy. Bot. Gard. Edinburgh 8: 244. - Lectotype 

(designated here): Szechuen, Tachien-lu, 2700-4050 m, Pratt 573 (K; Iso: BM!, G!). Note: 

The statement of Wenninger 1991: Holotype: Wilson 3420 is wrong! 
A. pseudocylindraceus Bornm. 1915, Magyar Bot. Lapok 14: 51 - Lectotype (designated 

here): Kurdistania Turcia, pr. Egin ad "Habnus-Erkek", 3.6.1890, Sintenis 2538 (Bl: foto 

K!; Iso: B!: foto MSB!, JE!, MSB!, W!, WU!) 
A. pubescens Schrank 1792, Prim. fl. Salisb.: 2. - Neotype (designated here): Salzburg, (M!: 

hb. Schrank, as A. pilosus). 
A. rawlinsianus Aitch. & Baker 1888, Trans. Linn. Soc. London, Bot. 3: 51. - Syntypes: 

Badghis, 1.5., 19.5. (P!: in fruit) et 20.5.1885 (BM!: in fruit), Aitchison 357. Lectotype 

(designated here): [1.5.1885], ^/Yc/jwoa? J57(K!: the flowering plant; Iso: BM!, C!, FI!, G!, 

K!, LE!, P!, W!). Note: The flowering and fruiting plants are almost distributed together. 
A. retufoliatus Y.C.Ho 1981, Bull. Bot. Res. North-East. Forest. Inst. 1(3): 1 14. - Lectotype 

(designated here): Yunnan, Yü beng-za, 3600 m, 21.6.1937, T.T. Yü 8683 (PE!; Iso: KUN!, 

PE!). 
A. rimarum Bomm. 1905, Bull. Herb. Boiss., ser. 2, 5: 755. - Lectotype (designated here): 

Elburs occ, in montibus inter Asadbar et Gerab, in jugo Gerdene Bary, 2900 m, 1.7.1902, 

Bornmüller 6856 (B!; Iso: B!, BM!, BP!, G!, K!, LE!, P!, W!). 
A. rochelianus Heuff 1853, Flora 36: 622. - Lectotype (designated here): In rupestribus infra 

Drukova legionio Vallachio banaticae, 1843, Heuffel (BP!). 
A. roomer/ Simonk. 1892, Termeszettud. Közl. (Pötfiiz.) 19: 138. - Lectotype (designated 

here): N Tolgyes-szorostol [ravine] loco "Vereskö" [near Borszek], ca. 1150 m, 5.8.1891, 

Roemer (BP!; Iso: BP!,.LE!, M!). 
A. saadius Parsa 1966, Fl. Iran 9: 77. - Lectotype (designated here): [Iran] Sirjan, VI. 1957, 

Parsa?{Yi\,\so:Yi\). 
A. samarkandinus Freyn 1904, Bull. Herb. Boiss., ser. 2, 4: 763. - Lectotype (designated 

here): [Uzbekistan] prope Samarkand, ad Balan Nur, 6.5.1898, Paulsen 107 (C!; Iso: 

BRNM!). 
A. sarae Eig 1955, Syst. Stud. Astrag. Near East: 40. - Syntypes: Iraq, Jebel Sindjar, 1 100 m, 

1200 m & 1500 m (HUJ!), 27.4.1933, Eig & Zohary; Iraq, Jebel Sindjar, 1060 m, Eig & 

Zohary. Lectotype (designated here): Iraq, Jebel Sindjar, 1060 m, Eig & Zohary (HUJ!; Iso: 

E!,K!,W!). 



187 



saxatilis Freyn & Bomm. 1891, Österr. Bot. Z. 41: 495. - Syntypes: Amasia, Abadschi- 

Dagh, 1100-1200 m, 11.5.1890, Bornmüller 1865 (B!, BM!, BRNM!, G!, JE!, LD!, LE!, 

OXF!, W!); Sana-Dagh, 14.5.1890, 700-900 m, Bornmüller 2114 (B!); Maghmur-Dagh, 

4.5.1890, Bornmüller 2115; Armenia turcica, Gumuschkhane, inter Beschklissa et Ardas, 

30.4.1890, Sintenis 2104. Lectotype (designated here): Sintenis 2104 (BRNM!; Iso: BP!, 

E!, G!, JE!, LD!, P!, PR!, W!, WU!). 

saxicola Ulbr. 1922, Repert. Spec. Nov. Regni Veget. Beih. 12: 423. - Type: Tschili, Hsiau 

Wu tai schan, 3050 m, Limpricht 2552. Lectotype (designated here): [an den höchsten 

Felsgraten des Pe tai, 3250 m, 17.7.1915], Limpricht 2552 (WU!; Iso: K!). 

saxifractor Rech.f. & Gilli 1958, Biol. Skr. 9(3): 133. - Syntypes: Afghanistan, Tangi 

Gharu, 1500-1700 m, 11.5.1951, Gilli 1518; and 22.6.1951, Gilli 1519 (W!). Lectotype 

(designated here): Gilli 1518 (W!). 

scholerianus Bomm. 1908, Mitth. Thüring. Bot. Vereins, n.s. 23: 3. - Syntypes: [Turkey, 

Konya] Phrygia, Akscheher, montis Sultan-dagh, 1700-1800 m, 21.6.1899, Bornmüller 

4411 (B!, BP!, BREM!, BRNM!, G!, JE!, K!, P!, W!); [Konya] Sultan-dagh supra pagum 

Permata, 1600 m, 25.6.1899, Bornmüller 4410. Lectotype (designated here): Bornmüller 

4410 (B!; Iso: BP!, BR!, BREM!, BRNM!, BRNU!, E!, G!, HBG!, JE!, K!, LE!, M!, 

MSB!,P!,PRC!, W!, WU!). 

semnanensis Bomm. & Rech.f. 1940, Repert. Spec. Nov. Regni Veg. 48: 123. - Lectotype 

(designated here): Iran, Prov. Damghan-Semnan, Sorcheh [Surkheh] bei Semnan, ca. 1400 m, 

29.6.1937, Rechinger 1249 (W!; Iso: B!, BM!, K!). 

senilis Bornm. 1905, Bull. Herb. Boiss., ser. 2, 5: 755. - Lectotype (designated here): inter 

Rescht et Kaswin, in trajectu Charsan, 2000 m, 13.5.1902, Bornmüller 6729 (B!; Iso: B!). 

serafschanicus Freyn 1904, Bull. Herb. Boiss., ser 2, 4: 767. - Lectotype (designated here): 

Serafschan, prope Samarkand, in steppa ad Balan Nur, 6.5.1898, Paulsen HI (BRNM!; 

Iso:C!,LE!). 

sessiliceps Bornm. 1908, Mitth. Thüring. Bot. Vereins, n.s. 23: 17. - Lectotype (designated 

here): Persia, inter Kermanschah et Nehawend, ad Dscham-Tueh, 14.5.1904, Strauss (B!; 

Iso: JE!). 

smithianus E.Peter 1938, Acta Horti Gothob. 12: 52. - Lectotype (designated here): China, 

Szechuan, Dongrergo, 20.7.1922, Smith 3927 (W!; Iso: LD!). 

stenoceras C.A.Mey. 

var. macranthus Bunge 1866, Bull. Soc. Imp. Naturalistes Moscou 39(2): 25. - Lectotype 

(designated here): Songaria, inter Arganaty et Kessykauss, 2.-3.5., Semenow (P!; Iso: LE!). 

, stenostachys Beck 1886, Denkschr. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl, 2. Abt. 
51: 333. - Lectotype (designated here): [Iran] prope Tschitschian, in itinere ad Nehawend, 
29.5.1882, Pichler (WU!; Iso: B!, W!, WU! - LE!: in agro Ecbatanensis). 

: stereocalyx Bornm. 1908, Mitth. Thüring. Bot. Vereins, n.s. 23: 15. - Lectotype 
(designated here): Phrygia, mt. Sultan-dagh ditionis opp. Akscheher, in jugo Teke-dagh, 
1600-1700 m, 25.6.1899, Bornmüller 4409 (sub. nomine A. nidus-avis) (B!; Iso: B!, BP!, 
BR!, BREM!, BRNM!, E!, G!, HBG!, K!, L!, LD!, LE!, M!, MSB!, P!, PR!, PRC!, STU!, 
W!, WU!,Z!). 

. stevenianus DC. 1825, Prodr. 2: 285. - Syntypes: in Asia minore, C.Rostan; in Iberia et 
Caucaso, Steven. Lectotype (designated here): in Iberia et Caucaso, Steven (G-DC!; Iso: H!: 
[circa Tiflin],P!). 



A. 5/o/aiii Nabeiek 1923, Spisy Prir. Fak. Masarykovy Univ. 35: 84. - Lectotype (designated 
here): Mesopotamia mont. Tur Abdin, ad pag. Takjan inter Geziret-ibn Omar et Midiat, 
900 m, 2.7.1910, //aZ?eMiÖ7<5(SAV!;Iso:SAV!). 

A. subspinosm Hayek & Siehe 1914, Ann. Nat. Hofmus. Wien 28: 163. - Lectotype 
(designated here): Antitaurus, Vil. Konia, Masmutli-Dagh, 1800 m, Ende Juni 1906, Siehe 
2ö5(W!;Iso: JE!,LE!,Z!). 

A. subumbellatus Klotzsch 1862, in Bot. Ergeb. Waldem. Reise: 159, tab. 3. - Type: Hima- 
laya, //o^^ewrer (B, destroyed; Fragm: P!). Neotype (Iconotype) (designated here): Tafel 
3, Bot. Ergeb. Waldem. Reise in connection with the fragment in P!: Hb. Bunge). 

A. suluklensis Freyn & Sint. 1904, Bull. Herb. Boiss., ser. 2, 4: 451. - Lectotype (designated 
here): [Kopet Dagh] Suluklü, 1.7.1900, Sintenis 737 (LD!; Iso: B!, BP!, BRNM!, G!, JE!, 
LD!,LE!,P!). 

A. superfluus Rech.f & Koie 1958, Biol. Skr. 9(3): 156. - Lectotype (designated here): 
Afghanistan, Herat, 1 100 m, 29.4.1949, Koie 4454 (W!; Iso: C!). 

A. syhensis Freyn 1905, Bull. Herb. Boiss., ser. 2, 5: 559. - Lectotype (designated here): 
Ferghana, Syk-Bulak ad fl. Langar, 16.6.1898, Paulsen 351 (BRNM!; Iso: C!, LE!). 

A. tauricola Boiss. 

- var. niveus Barbey 1885, Bull. Soc. Vaud. Sei. Nat. 21: 221. - Lectotype (designated here): 
insula Carpathos ad Menites, 24.5.1883, Pichler 240 (G!; Iso: G!, G-BOIS!, K!, LE!, 
MSB!,P!, WU!). 

A. tempskyanus Freyn 1890, Österr. Bot. Z. 40: 442. - Lectotype (designated here): Amasia, 
400-500 m, 18.5.1889, 5or«mw//er 700 (BRNM!; Iso: B!, BR!, G!, GOET!, JE!, K!, LE!, 
MSB!, OXF!, P!, PR!, PRC!, STU!, W!, WU!). 

- subsp. crassicarpus Freyn & Sint. 1894, Österr. Bot. Z. 44: 64. - Lectotype (designated 
here): Paphlagonia ad Tossia, ad Schakylar, 16.7.1892, Sintenis 3663b (BRNM!; Iso: BP!, 
JE!, MSB!, P!, PR!). 

- subsp. unguiculatus Freyn 1894, Österr. Bot. Z. 44: 63. - Lectotype (designated here): 
Paphlagonia ad Tossia, ad Kawak-Tscheschme, 10.5. (flor.) et 7.6. (fruct.) 1892, Sintenis 
3663 (BRNM!; Iso: B!, BP!, BR!, BRNM!, JE!, LE!, MSB!, P!, PR!, PRC!, W!, WU!). 

A. tenuiscapus Freyn & Bomm. 1897, Bull. Herb. Boiss. 5: 590. - Lectotype (designated 
here): Kerman, Kuh-Lalesar, 3900^000 m, 15.7.1892, Bornmüller 3777 (B!; Iso: BRNM!). 
A. tongolensis Ulbr. 

- var. glaber E.Peter 1937, Acta Horti. Gothob. 12: 49. - Lectotype (designated here): Tibet 
orient., Amnyi Machen range [W of Yellow River: valley beyond Draknak nira, 13000', 
VII.1926, Rock 14420 (E!; Iso: E!, K!, PE!). 

- var. lanceolato-dentatus E.Peter 1937, Acta Horti Gothob. 12: 49. - Lectotype (designated 
here): Szechuan occ, Tsipula, 4000 m, 5.8.1922, Smith 4175 (W!). 

A. transcaspicus Freyn & Bomm. 1900, Mem. Herb. Boiss. 13: 16. - Lectotype (designated 
here): Transcaspica, prope Usun Ada, 20.4.1896, Brotherus 1016 (BRNM!; Iso: G!, H!). 

A. triqueter Bomm. & Gauba 1935, Repert. Spec. Nov. Regni Veg. 39: 106, nom. illeg. [non A. 
triquetrus A.Gray]. - Syntypes: Elburs, über Keredj, 17.5.1934, Gauba (B!); et 10.6.1934, 
Gauba (B!); et 18.5.1935, Gauba 372a. Lectotype (designated here): Gauba 372a (B!). 

A. ufraensis Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 568. - Lectotype (designated 
here): Krasnowodsk, ad Ufra, 21.4.1901, Sintenis 1566 (BRNM!; Iso: B!, BM!, BP!, 
BRNM!, E!, G!, H!, JE!, K!, L!, LD!, LE!, M!, MSB!, MW, P!, PR!, PRC!, STU!, W!, 
WU!). 



189 



A. uhlwormianus Freyn & Bomm. 1890, Österr. Bot. Z. 40: 441. - Lectotype (designated 

here): Pontus australis, prope Khaousa, 400-500 m, 3.6.1889, Bornmüller 98 c (B\; Iso: B!, 

BRNM!, LE!). 
A. ulothrix Beck, 1886, Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 51: 340. - 

Lectotype (designated here): inter Maniam et Kaebuterchan, 15.5.1882, Pichler (WU!; Iso: 

JE! fragm., WU! - G!: in monte Karaghan). 
A. unifoliolatus Sirj. & Rech.f. 1953, Anz. Österr. Akad. Wiss. Math.- Naturwiss. Kl. 1953 

(90): 116, nom. illeg. [non Bunge]. - Lectotype (designated here): Iran, Pashmshurum, 

Bakhtiari, 29.4.1940, Koelz 15115 (W!; Iso: US). 
A. utriger Pall. 

- var. ovatus DC. 1802, Astragalogia: 217. - Lectotype (designated here): 'A. orientalis 
acaulos ferme flore luteo', Tournefort (P-TRF!; Iso: B-WILLD 14050!, P-LA!). 

A. vandasii Velen. 1893, Sitzungsber. Königl. Böhm. Ges. Wiss. Prag, Math.-Naturwiss. CI. 

1892 (37): 24. - Lectotype (designated here): Mt.. Rhodope ad Backovo, V.1893, Vele- 

novsky <& Stribrny (PRC: no. 858C!; Iso: Bl, BR!, G!, PRC: no. 858A!, 858B!). 
A. variifolius Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 784. - Syntypes: Kasandschik, 

inter Usun-su et Uschak, 28.4.1901, Sintenis 1594a; dto., Sintenis 1594c (BRNM!, LD!); 

Kisil-Arwat, 3.5.1901, Sintenis 1623c (BM!, BP!, BRNM!, E!, K!, LD!, LE!, PR!, PRC!). 

Lectotype (designated here): Sintenis 1594a (BRNM!; Iso: B!, BRNM!, G!, JE!, LD!, LE!, 

MSB!,P!,STU!, WU!). 

- var. homoiophyllus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 786. - Syntypes: Ka- 
sandschik, 28.4.1901, Sintenis 1594d\ Kisil-Arwat, 3.5.1901, Sintenis 1623/ {B\, BRNM!, 
LD!); bei Stara Karakala, 26.5.1901, Sintenis [1623g] (LD!). Lectotype (designated here): 
Sintenis 1594d(LD\; Iso: BRNM!; JE!, LE!). 

A. variistipula Turrill 1929, Kew Bull. 1929: 227. - Syntypes: N. Persia, near Tabriz, V.1927 

(flowering), Gilliat-Smith 1901; & VI. 1927 (fruiting), Gilliat-Smith 2274 (Kl). Lectotype 

(designated here): Gilliat-Smith 1901 (K!). 
A. venostanus Fritsch 1922, Exkursionsfl. Österr., ed. 3: 277, in clavi. - Type not indicated. 

Lectotype (designated here): Südtirol, Laas im Vintschgau, 1869, A.Kerner (WU!; Iso: 

WU!,L!). 
A. versipilus Rech.f. & Kaie 1958, Biol. Skr. 9(3): 168. - Lectotype (designated here): 

Afghanistan, inter Herat et Shindand, 1400 m, 8.5.1949, Koie 3944 (W!; Iso: C!). 
A. vesicarius L. 

- var. multißorus Cuatrec. 1929, Trab. Mus. Ci. Nat. Barcelona 12 [Fl. Mägina]: 335. - Syn- 
types: Sierra Mägina: Penascos del Barranco del Tejuelo, 1800 m, 30.6.1925, Cuatrecasas 
(MA!); Mägina, vert. SE, hacia El Gollzino, 1800 m, \.1.\925, Cuatrecasas (K!); Penascos 
del Centenillo, 1800 m, 9.7.1925, Cuatrecasas (MA!); Almaden, vert. N., 1850m, 19.7. 1925, 
Cuatrecasas; Almaden, en Rejön del Peregrino, 1700 m, 1.7.1926, Cuatrecasas; Carceles, 
vert. NW., 1950 m, 17.6.1926, Cuatrecasas; Mägina Central, 2000 m, 26.6.1926, Cuatre- 
casas; Canada del Tejuelo, 1700 m, 25.6.1926, Cuatrecasas (MA!). Lectotype (designated 
here): [Spain] Sierra de Mägina, Carceles NW, 1950 m, 17.6.1926, Cuatrecasas (BC!). 

A. vicioides Ledeb. 

- \ar. parvißora Trautv. 1860, Bull. Soc. Imp. Naturalistes Moscou 33(1): 498. - Syntypes: 
prope Karkaraly, 19.7.1840, Schrenk; in promontorio montium Tastau Songoriae, 1840, 
Schrenk (P!). Lectotype (designated here): prope Karkaraly, 19.7.1840, Schrenk (LE!). 

A. virgaeformis Sirj. & Rech.f 1958, Biol. Skr. 9(3): 139. - Lectotype (designated here): 
Afghanistan, Hauz-i-Mahiha, 2500 m, 1 1.7.1948, Kßie 2383 (W!; Iso: C!). 



190 



A. viridiformis Sirj. 1941, Repert. Spec. Nov. Regni Veg. 50: 285. - Lectotype (designated 
here): Armenia turcica, Kharput, Schuschnas, 5.6.1889, Sintenis 581 (WU!; Iso: BRNM!). 

A. viridissimus Freyn «fe Sint. 1892, Österr. Bot. Z. 42: 48. - Lectotype (designated here): 
Pontus, Trapezunt, inter Mataradjik et Hamskiöi, 28.4.1890, Sintenis 2082 (BRNM!; Iso: 
JE!, LD!). 

A. vulpinus Willd. 1802, Sp. PI. 3: 1259. - Typonym: A. alopecuroides auct: Pall., Sp. 
Astragal.: 9. 1800, non L. - Syntypes: Sibiria, in Hb. Willdenow 13962/1-2. Lectotype 
(designated here): Sibiria, (B- Willd 13962/21; Iso: B- WILLD 13962/1!, M!, W!). 

A. weirianus Aitch. & Baker 1888, Trans. Linn. Soc. London, Bot. 3: 54. - Lectotype (desig- 
nated here): Badghis [probably near Gal-icha W of Gulran], 29.4.1885, Aitchison 336 (K!; 
Iso: BM!, C, G!, LE!, P!). Note: The statement of Podlech & Maassoumi (1989) 
"holotype in K" is erroneous. 

A. wettsteinianus Freyn & Sint. 1895, Bull. Herb. Boiss. 3: 183. - Lectotype (designated 
here): Armenia turcica, Gümüschkhane, in monte Darsosdagh, 6.6.1894, Sintenis 5735 
(BRNM!; Iso: B!, BM!, BP!, BR!, E!, G!, JE!, K!, L!, LD!, LE!, M!, MSB!, P!, PR!, 
PRC!,STU!,W!,WU!). 

A. xanthinus Freyn & Bomm. 1892, Österr. Bot. Z. 42: 47; et Bull. Herb. Boiss. 3: 184. 
1895. - Lectotype (designated here): Pontus australis, mt. Yildiss-Dagh, 2300 m, 8.6.1890, 
Bommüller 2098 (B!; Iso: PRC!). 

A. xanthoxiphidium Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 788. - Type: As there 
is no type for the species, one must be selected out of the subspecies. I select the type of 
subsp. accrescens. Lectotype (designated here): Aschabad, IV.V.1900, Sintenis 101c (LD!; 
Iso:B!,LE!,MSB!,P!). 

- subsp. accrescens Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 796. - Lectotype 
(designated here): Aschabad, IV.V.1900, Sintenis 101c (LD!; Iso: B!, LE!, MSB!, P!). Note: 
As there are no collections and no type given for the subspecies I select as lectotype the 
collection given for var. angustilobus . 

- - var. angustilobus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 796. - See type for 
ssp. accrescens. 

- - var. latilobus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 797. - Syntypes: 
Aschabad, 20.4. et 12.5.1900, Sintenis lOle [acording the printed label it should be var. 
angustilobus] (BP!, BRNM!, G!, JE!, LD!, P!, STU!, W!, WU!); et Sintenis lOlf. 
Lectotype (designated here): Sintenis 101f(LD\- Iso: BRNM!, LD!). 

- subsp. brotherusii Yreyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 1015. - Syntypes: 
prope Kodsch Transcaspiae, 1.5.1896, Brotherus 1031; Krasnowodsk, 15.4.1901, Sintenis 
1567. Lectotype (designated here): Sintenis 7567 (LD!; Iso: B!, BRNM!, JE!, LD!, LE!). 

- subsp. campylopus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 797. - Lectotype 
(designated here): Karakala, in monte Sundsodagh, 18.5. et VI. 1901, Sintenis 1707a (LD!; 
Iso: B!, BM!, BP!, BRNM!, E!, G!, JE!, K!, LD!, LE!, MSB!, P!, PRC!, STU!, WU!). 

- subsp. curvicaulis Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 1014. - Syntypes: 
Aschabad, 20.4. et 18.5.1900 [in printed labels 12.5.1900], Sintenis 101a. Lectotype 
(designated here): Sintenis 101a (LD!; Iso: LD!; plants from both dates have been mostly 
distributed together: B!, BRNM!, JE!, LE!, STU!). 

- subsp. holoxanthus Freyn &. Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 1013. - Lectotype 
(designated here): Aschabad, 18.5.1900, Sintenis 101b (LD!; Iso: BM!, BRNM!, E!, K!, 
LD!). 



191 



- subsp. laüfoliolatus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 1012. - Lectotype 
(designated here): Aschabad, 12.5.1900, Sintenis Wie (LD!; Iso: BRNM!, JE!, LD!, LE!). 

- subsp. obscurus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 799. - Lectotype 
(designated here): Aschabad, 20.4.1900, Sintenis lOld (BRNM!; Iso: BRNM!, LE!, 
MSB!). 

- subsp. rectus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 797. - Syntypes: Kisil 
Arwat, Karakala, in monte Sundsodagh, 18.5.1901, Sintenis 1707b; Suluklü, 7.7.1900, 
Sintenis 7i<5 (BRNM!, LD!). Lectotype (designated here): Sintenis 1707b (LD!; Iso: B!, 
BM!, BP!, BRNM!, E!, G!, JE!, K!, LD!, LE!, STU!). 

- - var. elongatus Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 798. - Lectotype (desig- 

nated here): Kisil Arwat, Karakala, in monte Sundsodagh, 18.5.1901, Sintenis 1707c 
(LD!;Iso:LD!,P!). 
A. xiphidioides Freyn & Sint. 1905, Bull. Herb. Boiss., ser. 2, 5: 786. - Syntypes: 
Kasandschik, inter Usun-su et Uschak, 28.4.1901, Sintenis 1594b; Kisil-Arwat, 3.5.1901, 
Sintenis 1623e. Lectotype (designated here): Sintenis 1594b (BRNM!; Iso: LD!). 
Ä. xiphocarpus Bunge 1868, Mem. Acad. Imp. Sei. Saint Petersbourg 1 (16): 27 [et I.e. 15(1): 
32. 1869]. - Lectotype (designated here): Sikkim, Khasrou, 9000', Hooker (K!: Hb. 
Hooker; Iso: CGE!, K!: Hb. Bentham& Hb. Hooker, L!, GOET!: 10000-11000'). 
A. xylorrhizus Freyn & Sint. 

- var. angustus Freyn & Sint. 1894, Österr. Bot. Z. 44: 28. - Syntypes: Paphlagonae ad 
Tossia, ad Kawak Tscheschme, 7.6.1892, Sintenis 4128; ad Tschinonbaba, 19.7.1892, 
Sintenis 4128b. Lectotype (designated here): Sintenis 4128 (BRNM!; Iso: B!, BM!, BP!, 
BR!, G!, GOET!, JE!, K!, L!, LE!, M!, MSB!, OXF!, PR!, PRC!, W!, WU!). 

A. zubairensis Eig 1955, Syst.'Stud. Astrag. Near East: 72. - Lectotype (designated here): 76 
km W Basra and about 60 km from Ez-Zubair, 8.4.1933, Eig & Zohary (HUJ!; Iso: E!, 
HUJ!). 



References 

Becht, R. 1978: Revision der Sektion Alopecuroidei DC. der Gattung Astragalus L. - 

Phanerog. Monogr. 10: 1-227. 
Ostenfeld, C. & Paulsen, O. 1922: Flowering Plants from Inner Asia. - In: Hedin, S.: 

Southern Tibet, vol. 6, 3. 
PoDLECH, D. 1998: Typification oi Astragalus species II. Species mainly from the herbaria of 

Paris (P) and Geneva (G). - Sendtnera 5: 247-263. 

- & Maassoumi, A.A. 1989: Two new species oi Astragalus from Iran and their affinities to 
A. weirianus from Afghanistan. - Kit Tan (ed.): The Davis and Hedge Festschrift: 71-79; 
Edinburgh. 

- & Sytin, a. 1996: Typification of Russian and some other species o^ Astragalus L. - 
Sendmera3: 149-176. 

Wenninger, J. 1991: Revision von Astragalus L. sect. Chlorostachys Bunge, sect. 
Phyllolobium Bunge und sect. Skythropos Bunge (Leguminosae). - Mitt. Bot. Staatssamml. 
München 30: 1-196. 

Prof. Dr. Dietrich Podlech, Institut für Systematische Botanik der Ludwig-Maximilians- 
Universität München, Menzinger Straße 67, D-80638 München, Germany 



193 



Some remarks on the genus Oxytropis (Fabaceae) from Iran 



M. Ranjbar 



Abstract: 

Ranjbar, M.: Some remarks on the genus Oxytropis (Fabaceae) from Iran. - Sendt- 
nera6: 193-196. 1999. ISSN 0944-0178. 

The Iranian representatives of the genus Oxytropis DC. are examined. Altogether 32 
species are recognized. O. Aellenii and O. shirkuhi are treated as synonyms of 
O. iranica and O. chrysocarpa respectively. O. khorasanica is proposed as a new 
name for O. gracillima and 8 species are excluded from Iran. The species occuring in 
Iran are keyed out. 

Zusammenfassung: 

Die iranischen Vertreter der Gattung Oxytropis DC. werden revidiert; 32 Arten 
werden anerkannt. O. Aellenii und O. shirkuhi werden als Synonyme von O. iranica 
bzw. O. chrysocarpa aufgefaßt. O. khorasanica ist ein neuer Name für O. gracillima; 8 
Arten werden aus der iranischen Flora ausgeschlossen. Die im Iran vorkommenden 
Arten von Oxytropis werden geschlüsselt. 



Introduction 

Oxytropis is a large genus of small herbs, mainly present in high pastures and rocky places. It 
is separated from Astragalus largely by tradition and for convenience; the main difference is in 
the shape of the keel (Lock & Simpson, 1991). According to the most recent morphological 
classification of the papilionoid tribe Galegeae (POLHILL, 1981), the closest relatives of Oxy- 
tropis include Meristotropis Fisch. & C.A.Mey., Astragalus L., Caragana Fabr., Chesneya 
Lindl. ex Endl., and all members of the subtribe Astragalinae. 

The genus Oxytropis is one of the most complex genera in the Fabaceae. For the most part 
this genus was introduced as a sister group of the genus Astragalus. Boissier (1872) in Flora 
Orientalis placed 8 species under two sections, namely Phacoxytropis and Euoxytropis and 
Vassilczenko (1984) in Flora Iranica placed 40 species under two subgenera, namely Oxy- 
tropis and Euoxytropis. These species and records were introduced only by one specimen and 
differential characters between them were very artificial. 

Recent taxonomic studies were carried out by Ranjbar (submitted) on the Iranian herbarium 
specimens of Oxytropis in the herbarium of the Research Institute of Forests and Rangelands 
(TARI). The author has also identified the materials of this genus in the herbaria of Plant Pests 



194 



Diseases Research Institute (IRAN) and University of Hamadan (HUH). Some of the species 
were considered to be synonyms, and thus some records were excluded from the Flora of Iran. 
Now Oxytropis is represented by 32 species in Iran. The identification of the species of the 
genus is very complex, difficult and often confused. In spite of that the work on plants of 
genus Oxytropis done by Vassilczenko (1984) in Flora Iranica, a new revision for the genus 
in Iran is still needed. Obviously a complete revision of the genus requires more specialized 
investigations which would also include species outside of Iran. This article contains 1 new 
name, 3 synonyms and a new key for the Iranian species of the genus. 



Nomenclatural changes and corrections of records 

1 . New name 

Oxytropis khorasanica Ranjbar, nom. no v. 

= O. gracillima Vassilcz., Fl. Ir. 157: 101-164 (1984), non Bunge in Mem. Acad. Imp. Sei. 
St.-Petersbourg ser. 7, 22: 160 (1874). 

2. New synonyms 

Oxytropis iranica Vassilcz., Nov. Syst. PI. Vase. Leningrad 17: 180 (1980). '' ; 

= O. Aellenii Vassilcz., Nov. Syst. PI. Vase. Leningrad 17: 195 (1980). 

Oxytropis chrysocarpa Boiss., Diagn. PI. Or. Nov. Ser. 1, 6: 34 (1845). 

= O. shirkuhi Vassilcz., Bjull. Mosk. Obsc. Isp. Prir., Otd. Biol. 93 (3): 97-102 (1988). 

3. Doubtftil records 

Oxytropis jezdii Vassilcz. 

This species was introduced by Vassilczenko (1988: 100-101) from central Iran, but no 
specimen was seen by the author. It may be that the new species was based on an incorrect 
identification of O. chrysocarpa Boiss. . ,^ , 

Oxytropis wendelboi Vassilcz. 

This species was recorded by Vassilczenko (1984: 152) from N of Iran, but no specimen 
was seen by the author. It may be that the record was based on an incorrect identification 
of O. szovitsii Boiss & Buhse. 

4. Incorrect records , 

Oxytropis immersa (Baker) Bunge ex B.Fedtsch. 

This name was recorded by Vassilczenko (1984: 129-130) from NE of Iran. The correct 
name for specimens under this name is O. iranica Vassicz, therefore Vassilczenko 's record 
is certainly incorrect. 

Oxytropis hirsutiuscula Freyn. and Oxytropis caraganetorum Vassilcz. 

These species were cited by Vassilczenko (1984: 118, 120) from central Iran and 
specimens were seen by the author. Correct identification of them are O. heratensis Bge. 



195 



Oxytropis czapan-daghi B.Fedtsch. 

This species was cited by Vassilczenko (1984: 133-134) from NE of Iran. This specimen 
has been studied by the author, and the correct name for it is O. khorasanica Ranjbar. 



Key to the species 

1 . Plant caulescent, racemes borne on leafy stems 2 

- Plant acaulescent, racemes arising from leafy base of plant 1 

2. Plant covered with appressed hairs 3 

- Plant covered partly with spreading hairs 4 

3. Pods covered with appressed, short, white and black hairs O. heratensis 

- Pods covered with, soft, long, more or less spreading white and black hairs 

O. thaumasimorpha 

4. Leaflets in 5-8 pairs 5 

- Leaflets in 9-16 pairs 7 

5. Calyx teeth longer than tube, pod covered with long and white hairs O. alavae 
Calyx teeth 2 to 3 times shorter than tube, pod covered with pubescent hairs 6 

6. Peduncles 2 times longer than leaves, standard 5-7 mm long O. sojakii 

- Peduncles many times shorter than leaves, standard 21-24 mm long O. strausii 

7. Pods erect to horizontal O. kotschyana 

- Pods deflexed 8 

8. Leaflets 9-10 pairs O. rechingeri 
Leaflets 12-16 pairs 9 

9. Calyx covered with white and black hairs, keel beak 3^ mm long O. kopetdaghensis 

- Calyx covered with white hairs, keel beak 1.5 mm long O. assadliensis 

10. Pods covered with short appressed hairs 1 1 

- Pods covered with soft, long and white hairs 1 6 

1 1 . Pods covered with short, white and black pubescent hairs O, savellanica 

- Pods covered with white pubescent hairs 1 2 

12. Pods bladder-like, ovate-globular O. persica 
Pods not bladder-like, oblong-elliptic 1 3 

13. Pods covered with short, white pubescent hairs 14 

- Pods covered with long, white pubescent hairs 1 5 

14. Calyx campanulate, teeth calyx linear-lanceolate, 4-6 mm long O. bicornis 
Calyx tubular, teeth calyx filiform, 2-2.5 mm long O. pusilloides 

15. Peduncles 10-12 cm long, petioles 4-6 cm long O. zangolehensis 

- Peduncles 3-5 cm long, petioles 1-2 cm long O. takhti-soleimanii 

16. Calyx covered with short, white and black pubescent hairs 1 7 

- Calyx covered with short, white pubescent hairs 24 

17. Pods deflexed O. surmandehi 
Pods erect 1 8 

18. Calyx teeth 2-2.5 times shorter than tube 1 9 
Calyx teeth equal to or longer than tube 20 

19. Pods covered with semi-appressed white hairs O. iranica 
Pods covered with soft, long and white hairs O. kermanica 

20. Standard 9-10 mm long 21 



196 



- Standard 12- 13 mm long 22 

21. Standard oblong-elliptic O. hypsophila 
Standard orbicular O. binaludensis 

22. Leaflets 5-7 pairs O. karjaginii 

- Leaflets 7-16 pairs 23 

23. Leaflets 7-10 pairs, standard emarginate at the apex O. aucheri 

- Leaflets 12-16 pairs, standard rounded at the apex •( -- O. szovitsii 

24. Calyx teeth shorter than tube O. khorasanica 

- Calyx teeth equal to or longer than tube > 25 

25. Standard orbicular or obovate or rhomboid ' ' 26 
Standard elliptic or oblongo-elliptic s 28 

26. Standard rhomboid O. cinerea 

- Standard orbicular or obovate - - 27 

27. Standard orbicular, peduncles shorter than leaves O. rhodontha 

- Standard oblong or obovate, peduncles equal to leaves ■ O. chrysocarpa 

28. Standard emarginate at the apex 29 
Standard rounded at the apex 3 1 

29. Leaflets 5-8 pairs, calyx teeth equal to tube O. masanderanensis 

- Leaflets 9-14 pairs, calyx teeth 1.5 times tube 30 

30. Calyx teeth 1.5 times longer than tube O. suavis 

- Calyx teeth equal to tube O. rudbariensis 

3 1 . Leaflets 5-10 pairs, peduncles longer than leaves O.kuchanensis 

- Leaflets 10-12 pairs, peduncles shorter than leaves O. neo-rechingeriana 



I would like to thank Prof. Dr. Dietrich Podlech (Institute for Systematic Botany, University 
of Munich) for critical review of the manuscript. I wish to thank the Regius Keeper of the 
national botanic garden, Iran for making the herbarium facilities available for study. Also I am 
much in debted to Mrs. R. Karamian for improving the english text. 



References 

BoissiER, E. 1872: Flora Orientalis. Vol. 11. Basel. 

Bunge, A. 1874: Species Generis Oxytropis DC. - Mem. Acad. Imp. Sei. Saint-Petersbourg. 

Ser7,22: 1-166. 
Candolle, A.P. de. 1825: Prodromus systematics regni vegetabilis. Vol. 2. Paris. 
Lock, J.M. & Simpson, K. 1991 : Legumes of west Asia. Royal Botanic Gardens, Kew. 
Polhill, R.M. 1981: Galegeae. - In: Polhill, R.M. & Raven, P.H. (eds.): Advances in 

legume systematics, 357-363. Royal Botanical Garden, Kew. 
Ranjbar M. 1997: Oxytropis. - In: AssADi, M. et al. (eds.): Flora of Iran (submitted). A 

revision of genus Oxytropis DC. in Iran. 
Vassilczenko, I.T. 1984: Oxytropis. - In: Rechinger, K.H. (ed.): Flora Iranica. No. 165: 

101-164. 
- 1988: New Asiatic species of genus Oxytropis DC. - Bjull. Mosk. Obsc. Isp. Prir., Otd. 

Biol. 93(3): 97-102. 

Massoud Ranjbar, Department of biology. Herbarium division. University of Buali sina, 
P.O.Box 65175/41 11, Hamadan, Iran. E-mail: Ranjbar(^basu.ac.ir 



197 



Chromosomenzahlen von Hieracium (Compositae, Lactuceae) Teil 3. 



F. Schuhwerk & W. Lippert 



Zusammenfassung: 

Schuhwerk, F. & Lippert, W.: Chromosomenzahlen von Hieracium L. (Compo- 
sitae, Lactuceae) Teil 3. - Sendtnera 6: 197-214. 1999. ISSN 0944-0178. 

48 Sippen des Subgenus Hieracium überwiegend aus Mitteleuropa wurden zytolo- 
gisch untersucht. Von den untersuchten Sippen (zusammengestellt in Tab. 1) waren 
70% triploid, 22% tetraploid und 8% diploid. Der Anteil der Tetraploiden ist in 
Mitteleuropa nur etwa halb so groß wie auf der Balkan-Halbinsel. Unter den Tetra- 
ploiden sind der größere Teil sog. Zwischenarten, unter den Triploiden etwa die 
Hälfte. Für H. doUineri ssp. dollineri wird eine revidierte Darstellung der Verbrei- 
tung in Bayern skizziert. 

Veränderungen in Kultur wurden insbesondere bei den Blattformen und -Zähnungen, 
sowie beim Indumentbesatz festgestellt. Die Variationsrichtung dieser Veränderun- 
gen war dabei bei verschiedenen Sippen teilweise gleichsinnig, teilweise gegensätz- 
lich. Diese Modifikabilität sollte bei der systematischen Bewertung einiger Merk- 
malskomplexe berücksichtigt werden. 

Abstract: 

In this third part of our studies on the cytology of Hieracium 48 taxa of the 
subgenus Hieracium from Central Europe are investigated. 70% of the taxa investi- 
gated (compiled in tab.l) were triploid, 20% tetraploid and 8% diploid. The part of 
tetraploids in this study of mainly Central European taxa is about half as much as 
in taxa from the Balkan Peninsula. The majority of the tetraploids but only the 
halfpart of the triploids are intermediate species ('Zwischenarten'). 
The variability of plants under cultivation was also investigated. Modifications 
have been observed mainly in the shape and serration of the leaves and in the 
amount of the indument. The direction of variation of these modifications in 
different taxa partly was the same and partly contrary. This modificability should 
be regarded at the systematic valuation of some character complexes. 



1. Einleitung 

Die zytologischen Kenntnisse bei Hieracium sind immer noch ungenügend: von vielen Taxa 
(auch den sog. Sammelarten) ist noch keine Chromosomenzahl bekannt, von anderen sind zu 
wenige Zählungen publiziert, als daß sich gesicherte Aussagen machen ließen. Allerdings sind 
bei dem in dieser Folge behandelten Subgenus Hieracium die zytologischen Verhältnisse 
einfacher als beim Subgenus Pilose IIa und abgesehen von den diploiden Sippen auch die 
Möglichkeit eingeschränkt, aus den Chromosomenzahlen Rückschlüsse auf die Sippenstruktur 



198 



und das Fortpflanzungsverhalten zu ziehen (Bräutigam & Bräutigam 1996). In der vor- 
liegenden Arbeit werden teilweise noch durch Merxmüller untersuchte Aufsammlungen, teils 
Ergebnisse der von uns fortgeführten Untersuchungen vorgestellt. Es werden Sippen des 
Subgenus Hieracium überwiegend aus Mitteleuropa behandelt. 



2. Material und Methoden 

Die teils lebend gesammelten, teils aus ausgesäten Achänen gewonnenen Pflanzen wurden im 
ungeheizten Gewächshaus kultiviert. Nach der Entnahme von Wurzelspitzen wurden die 
Pflanzen umgetopft und anfangs im Gewächshaus, später im Freiland in Erde eingesenkt 
weiterkultiviert. 

Die Zählungen wurden an Wurzelspitzen vorgenommen. Ca. 0,2-0,5 cm lange Wurzelspit- 
zen wurden 2-3 Stunden in wässriger Lösung von 0,002 mol Hydroxychinolin vorbehandelt. 
Nach 15 Minuten Hydrolyse in In HCl bei 60°C folgte die Färbung mit Orcein-Eisessig und 
die Anfertigung von Quetschpräparaten. 



3. Hauptteil 

Allgemeine Ergebnisse 

Mit fast genau 70% stellen die Triploiden erwartungsgemäß den Großteil der untersuchten 
Sippen. Bei etwa 8% unserer Zählungen erwiesen sich die untersuchten Sippen als diploid. 
Nicht überraschend war dies bei zwei Herkünften von H. umbellatum. Das ebenfalls diploide 
H. pseudocorymbosum wird als mit H. umbellatum verwandt vermutet. Die diploide Pflanze 
von H. racemosum wuchs in einer Population mit triploiden Pflanzen. Etwa 22% der 
untersuchten Sippen waren tetraploid. Bei Untersuchungen von Hieracien der Balkan-Halb- 
insel war dieser Anteil etwa doppelt so groß (Schuhwerk «fe Lippert 1998). Wie dort sind der 
größere Teil der Tetraploiden sog. Zwischenarten. Dies ist allerdings keine zwingende 
Parallele, sondern allenfalls eine unterschiedliche Schwerpunktverteilung: die untersuchten Tri- 
ploiden sind etwa zur Hälfte Haupt-, zur andern Hälfte Zwischenarten. 

Für zwei in Süddeutschland weit verbreitete Sammelarten zeichnen sich inzwischen Ten- 
denzen ab. H. glaucinum wurde bisher ausnahmslos, H. bifidum ganz überwiegend als triploid 
festgestellt. Dies gih für jeweils mehrere untersuchte Unterarten. 

Auch bei den hier untersuchten Pflanzen waren in Kultur Veränderungen zu beobachten. 
Ein Teil der Pflanzen bildete erst in Kultur eine Grundblattrosette (//. jurassicum, H. lingels- 
heimii, z.T. H. umbellatum); bei H. dollineri war bei der schattig gewachsenen kultivierten 
Pflanze die Grundblattrosette stärker betont. Bei H. bupleuroides entwickelten die Pflanzen in 
Kultur teilweise weniger (nur vier statt bis sieben), dafür etwas breitere (7-8 statt 5-6 mm) 
Stengelblätter. Zwei Sippen von H. glaucinum (ssp. glaucinum und ssp. similatum) ent- 
wickeln in Kultur nicht den typischen gestutzten oder schwach herzförmigen Blattspreiten- 
Grund, sondern einen keilig verschmälerten. Entsprechend der typischen Blattfolge von 
Hieracium bei Wildpflanzen wies höchstens das jeweils älteste Blatt den kennzeichnenden, 
gestutzten oder schwach herzförmigen Grund auf. Bei einigen Sippen war die Zähnung des 
Blattrandes bei den kultivierten Pflanzen verschiedener Jahre unterschiedlich intensiv (//. 
lycopifolium, H. pseudocorymbosum, H. lingelsheimii), teils war sie reduziert (//. racemosum 
ssp. leiopsis), teils in Kultur stärker ausgeprägt (//. dollineri, H. caesium ssp. caesium, H. 



199 

glaucinum ssp. basalticum). Auch die Blattfleckung kann bei kultivierten Pflanzen im Ver- 
gleich mit den entsprechenden Wildaufsammlungen in unterschiedlicher Weise variieren: 
schwächer oder gamicht z.B. bei H. caesium ssp. carnosum, H. franconicum, unterschiedlich 
bei H. glaucinum ssp. recensitum, H. bifidum ssp. subcaesium, deutlicher und stärker dagegen 
z.B. bei H. bifidum ssp. canitiosum und ssp. eriopodoides. 

Bei H. pseudocorymbosum ssp. petryanum bilden nur die kräftigsten Pflanzen an der 
Spitze des Korbstandes eine deutliche Dolde aus. Mikrocephalie (d.h. Hüllenlängen bei den 
Wildpflanzen um 5 mm) verschwand in Kultur bei H. bifidum ssp. subcaesium und H. glau- 
cinum ssp. fraternum. 

Der Indumentbesatz an verschiedenen Organen war schon früher in Kultur als sehr variabel 
beobachtet worden. Der Stemhaarbesatz an den Blättern nahm bei H. glaucinum ssp. 
basalticum ab, war bei H. lycopifolium unterschiedlich, blieb bei H. pseudocorymbosum gleich, 
während er bei H. racemosum ssp. leiopsis und H. franconicum ausgedehnter war. Stemhaar- 
besatz an den Hüllen nahm z.B. bei H. racemosum ssp. leiopsis, H. franconicum und bei H. 
jurassicum in Kultur zu. Bei H. atratum ssp. schroeterianum schwankt die Drüsenmenge an 
den Blatträndem bei den kultivierten Pflanzen der verschiedenen Jahre deutlich, weniger 
diejenige auf den Blattflächen. 

In gegensätzlicher Richtung variieren kann auch die Färbung der Drüsen an der Hülle. Bei H. 
atratum ssp. atratum sind bei kultivierten Pflanzen die Drüsenköpfchen heller und die helle 
Strecke am Schaft länger. Auch die Haare dort weisen in Kultur einen längeren hellen Spitzen- 
teil auf Bei H. saxifragum ssp. dufftii dagegen sind an den kultivierten Pflanzen die bei der 
Wildaufsammlung gelbköpfigen, zarten Drüsen an der Hülle gelegentlich etwas kräftiger und 
dunkler. 

Bei H. lycopifolium bleibt die schwache Bewimperung der Ligulaezähne bei einzelnen 
kultivierten Pflanzen aus. 

Auch diese Beobachtungen mahnen zur Vorsicht bei der Bewertung verschiedener Merk- 
malsausbildungen bei Hieracien, vor allem bei nur quantitativ unterschiedlichen. Weitere Beob- 
achtungen der Veränderungen in Kultur müssen zeigen, ob die teilweise gegensätzlichen 
Variationsrichtungen spezifisch sind für Sippen oder Verwandtschaftsgruppen, oder ob sie 
z.B. von unterschiedlichen klimatischen Bedingungen in den einzelnen Kulturjahren abhängen. 



Einzelergebnisse 

Vorbemerkungen zur Art der Darstellung 

Alle zitierten Belege sind im Herbarium der Botanischen Staatssammlung München (M) nie- 
dergelegt. Falls eine Pflanze nur lebend gesammeh wurde, also kein Herbarbeleg der Wildauf- 
sammlung vorhanden ist, wird dies vermerkt. Alle Pflanzen wurden im Botanischen Garten 
München kultiviert (betreut von A. Hartmann). Nach der Kultur-Nummer zitierte Belege 
stammen von diesen kultivierten Pflanzen. Um die Fundortsangaben etwas zu kürzen, steht 
bei mehreren Aufsammlungen innerhalb derselben Art ein „-" für die Wiederholung 
gleichlautender Angaben (wie Regierungsbezirk, Landkreis u.a.) der zuvor genannten Auf- 
sammlung. Die Chromosomenzahlen wurden für jede getrennt aufgeführte Kultur-Nummer 
getrennt festgestellt. 

In der Benennung der Sippen folgen wir in der Regel den Bearbeitungen Zahns (im 
„Pflanzenreich" 1921-1923, bzw. in der „Synopsis" 1922-1938). Bisher fiir Hieracium 
veröffentlichte Chromosomenzählungen wurden vor allem den bekannten Indices enmommen 
(zusammengestellt bei Schuhwerk 1996). 



200 



Hieracium alpinumL. ssp. alpinum 

Unsere Zählung bestätigt die bisher aus Mitteleuropa vorliegenden, bei denen für die 
Sammelart ebenfalls 2n = 27 beobachtet wurde. 

2n = 27 

Deutschland. Bayern : Chiemgauer Alpen, Geigelstein, vom Sattel nördlich des Geigelstein- 
gipfels zur Roßalm, MTB 8239/4, versauerte Rasen auf Kössener Schichten, 20.7.1986, Lippert 
21526, = Kultur-Nr., Herbarbelege vom Mai 1988. 

Hieracium alpinum L. ssp. halleri (Vill.) Zahn 

Wie bei Chrtek (1997) für slowakische Pflanzen festgestellt, erwiesen sich auch alpische 
Herkünfte als triploid. Wie er es für die alpischen Populationen als recht häufig feststellt, sind 
auch bei den untersuchten Pflanzen die Ligulae röhrig. In Kultur entwickeln die Pflanzen 
gelegentlich zwei oder sogar drei Körbe. 

2n = 27 

Österreich. Tirol : Tuxer Voralpen, Weidener Hütte oberhalb Nafing, Anstieg Halslspitze- 
Nurpenjoch-Rastkogel, 2200 2400 m, 1979 Albertshofer, Kultur-Nr. zunächst 1009, dann 1010 
bzw. H-97 = H-98, Herbarbelege vom Juli 1982 und Sommer 1984. 

Hieracium amplexicaule L. ssp. berardianum (Arv.-Touv.) Zahn 

Die von Zahn später illegitim ssp. petraeum (Hoppe) Zahn genannte Sippe ist von der 
ebenfalls haarlosen oder nur arm behaarten ssp. amplexicaule abgesehen von der 
unterschiedlichen Achänenfarbe (schwarzbraun bzw. rotbraun) nicht leicht unterscheidbar. 
Auch chorologisch verhalten sich beide recht ähnlich: Zahn nennt die ssp. berardianum 
allerdings etwas häufiger aus den Ostalpen und einzig von den Dinariden. Ihre Chromo- 
somenzahl war bisher unbekannt; sowohl triploide wie tetraploide Pflanzen waren bei der 
Sammelart bzw. anderen Unterarten schon beobachtet worden (Schuhwerk 1996). Junge 
Blätter der kultivierten Pflanze sind bei Nr. 2284 etwas behaart im Gegensatz zu den völlig 
haarlosen der Wildaufsammlung. 

2n = 27 

Österreich. Tirol : Osttirol, Venedigergruppe, Aufstieg zur Sajathütte, 1985 Albertshofer, 
Kultur-Nr. 2284, Herbarbeleg vom Juli 1988. 

Hieracium atratum Fr. ssp. atratum ^^^ 

Bei der Wildaufsammlung ist der Schaft der Drüsen an Korbstielen und Hülle dunkel und 
nur direkt unter dem dunkelgelben Köpfchen hell. Bei kultivierten Pflanzen sind die Drüsen- 
köpfchen hellgelb und die helle Strecke am Schaft ist länger. Auch die Haare dort weisen in 
Kultur einen längeren hellen Spitzenteil auf. Nach bisherigen Zählungen gibt es bei der Sam- 
melart tri- und tetraploide Sippen, die Chromosomenzahl der Unterart war bisher unbekannt. 

2n-27 

Italien. Trentino : Prov. Trento, Madonna di Campiglio, Valle di Nambrone, am Crozzi 
deirUomo, 2060 m, MTB 9730/3, Silikatfels und Erlengebüsch, 1.8.1995 Gottschlich 30595 & 
Nydegger 34260, Kulüir-Nr. 97-64, Herbarbeleg vom 15.7. 1998 (2 Zählungen). 

Hieracium atratum Fr. ssp. schroeterianum Zahn 

Zahn (1938: 188) schreibt bei dieser Sippe, die Drüsenbekleidung erinnere an H. inty- 
baceum, schließt daran aber die in eine gänzlich andere Richtung weisende Vermutung an, es 
handle sich vielleicht um „//. atratum > bocconei oder alpinum < diaphanoides'\ Diese 
gegensätzlichen Verwandtschaftsbeziehungen und die starke Aufspaltung der Sippe durch 
Zahn könnten aber auch darauf hindeuten, daß es sich um ein durch das Merkmal „Drüsen- 



201 



reichtum" zusammengehaltenes künstliches Taxon handelt (Gottschlich in litt.)- Populations- 
beobachtungen im AUgäu (Österreich, Vorarlberg, Bregenzer Wald, Kleines Walsertal; westlich 
oberhalb Baad, oberhalb der oberen Spital- Alpe; saurer Rasen; ca. 2000-2050 m. MTB 
8626/43, 30.7.1997 Schuhwerk 97/253a et al.) machten es dort jedoch wahrscheinlich, daß es 
sich um eine aus H. picroides und H. murorum entstandene Sippe handelt, die zur Art 
aufgestuft werden sollte, falls diese Entstehungsweise molekularbiologisch erwiesen werden 
könnte. Molekularbiologisch könnte auch Homogenität bzw. Heterogenität der Sippe über- 
prüft werden. Ihre Chromosomenzahl war bisher unbekannt. Die Drüsenmenge an den Blatt- 
rändem schwankt bei den kultivierten Pflanzen der verschiedenen Jahre deutlich, weniger 
diejenige auf den Blattflächen. Die Zähnung der Blattränder ist nur bei kräftigen Pflanzen 
deutlich. Aus "Arvenwäldem im Scarltal" stammt eine von Zahn bei der Erstbeschreibung 
(Zahn 1906: 390) unter zahlreichen Syntypen zitierte Aufsammlung von C.J. Schröter. 

2n = 27 

Schweiz. Graubünden : Engadiner Dolomiten, Val S-charl oberhalb Alp Astras in Richtung Plan 
Mattun, ca. 2200 m, sehr trockener Wuchsort neben Zwergsträuchem, 12.8.1992, Pahl 5610, 
det. F. Schuhwerk. Kultur-Nr. 94-85, Herbarbelege vom 14.6. (Dubl. GLM, Hb. Gottschlich) und 
17.7.1996, 17.6., 6. und 20.8.1997 und 27.7.1998. 

Hieracium bifidum Kit. ex Homem. ssp. canitiosum (Dabist.) Zahn 

Die Chromosomenzahl der in Bayern nach den Angaben Zahns auf die Alpen beschränkten 
Unterart war bisher unbekarmt. Die bei der Freilandpopulation schwache Blatt-Fleckung ist 
bei den kultivierten Pflanzen in der Regel deutlicher und stärker. 

2n = 27 

Deutschland. Bayern : Regierungsbezirk Niederbayem, Landkreis Landshut, Mittleres Isartal, 
NW-Hang südöstlich von P. 375 südöstlich Aumühle unterhalb Landshut, MTB 7439/21, offene 
Stelle in lockerem Brachypodium-Sesleria-Rasen, 21.5.1993, Schuhwerk & Stein (nur lebend, 
Blätter schwach gefleckt), Kultur-Nr. 2811, Herbarbelege vom 14.6.1996, 13.6. und 26.8.1997. 

Hieracium bifidum Kit. ex Homem. ssp. eriopodoides (Zahn) Zahn 

Die Chromosomenzahl der zur grex pseudodollineri (Murr & Zahn) Zahn gehörigen 
Unterart war noch unbekannt. Diese Gruppe von Unterarten verbindet nach Zahn H. bifidum 
und H. dollineri, doch mögen auch Beziehungen zu H. oxyodon bestehen. Auf Beziehungen 
zur sect. Drepanoidea (= sect. Glauca) weist jedenfalls auch der zur Bestimmung allerdings 
kaum verwendbare, nur bei starker Binokular- Vergrößerung erkennbare abweichende Typus 
der Sternhaare an Hülle und Korbstielen hin. Er vermittelt mit kürzeren Stielen zu demjenigen 
der sect. Drepanoidea, der „pubes farinosa" Grisebachs (1852:4). Die Blätter - bei der Wild- 
aufsammlung ungefleckt - können bei kultivierten Pflanzen schwach gefleckt sein. 

2n = 27 

Deutschland. Bayern : Regierungsbezirk Oberbayem, Landkreis Miesbach, Mangfallgebirge, 
Kreuzberg Ostseite, MTB 8337/43, 23.6.1993, Mayer [93/J176, det. F. Schuhwerk, Kultur-Nr. 94- 
65, Herbarbelege vom 7. (Dubl. in: GLM, Hb. Gottschlich), 18.6.1996 und 29.6.1998. - Landkreis 
Berchtesgadener Land, an der Straße nach Oberjettenberg bei der Abzweigung zum Dolomit- Werk, 
MTB 8342/24, 13.6.1996, Lippen 27190 = Kultur-Nr., Herbarbelege vom 29.6.1998. 

Hieracium bifidum Kit. ex Homem. äff. ssp. psammogenes Zahn 

Unterarten der stark behaarten grex psammogenes Zahn sind im Fränkischen Jura sehr 
selten und auf dessen Südostteil beschränkt. Von der eigentlichen ssp. psammogenes 
unterscheidet sich die vorliegende Aufsammlung allerdings durch bis zu 5 mm lange, vor allem 
aber direkt über der Basis 0,2 (0,15-0,25) mm dicke, gekrümmte, borstige Haare an Blattrand 



202 



und -Unterseite. Daher hält Meyer (1999: 8) mit Gottschlich diese Sippe, die im Kipfenber- 
ger Gebiet ein kleines Areal besiedelt, für eine unbeschriebene Unterart aus dem Formenkreis 
von//, wiesbaurianum. Dajedoch andere auf diese Sammelart weisende Merkmale fehlen und 
es noch keinerlei breiter angelegte Meßreihen der Blattbehaarung in den fraglichen Verwandt- 
schaftskreisen gibt, soll die Aufsammlung hier vorläufig noch H. bifidum zugeordnet werden. 
Die Blatt-Oberseiten neigen in Kultur zum Verkahlen, auch der Haarbesatz der Hülle kann sich 
etwas verringern. 

2n = 27 

Deutschland. Bayern : Fränkische Alb, Felsen gegenüber Grosdorf bei Kipfenberg, 470 m, 
MTB 7034/1, 16.5.1990, Schuhwerk 90/1012, Bräutigam, Krach & Lippert, Kultur-Nr. 90/1012- 
2, Herbarbelege vom Juli 1993 und 3.7.1996. Kultur-Nr. 90/1012-5, ebenfalls triploid, ging nach 
der Zählung ein. Kultur-Nr. 92-5, Herbarbelege vom Juli 1993 (Dubletten in: GLM, Hb. Gott- 
schlich) und 11.6.1996. 

Hieracium bifidum Kit. ex Hörnern, ssp. stenolepis (Lindeb.) Zahn 

Die tetraploide Stufe nennt für die Sammelart bisher nur Moore (1982: 285, ohne Quellen- 
angabe). Die im südöstlichen Frankenjura vorkommende Sippe zeichnet sich innerhalb der 
äußerst vielgestaltigen ssp. stenolepis aus durch recht derbe, blaugrüne, regelmäßig gefleckte 
Blätter, die unterseits und gelegentlich auch oberseits mit Stemhaaren besetzt sind. Es bleibt 
nachzuprüfen, ob hier nicht eine eigene Lokalsippe vorliegt. 

2n = 36 

Deutschland. Bayern : Regierungsbezirk Oberpfalz, Landkreis Regensburg, Fränkischer Jura, 
Weißer Stein beim Wuzenfels südlich P. 401,6 im Bachmühltal westlich Deuerling, 400 450 m, auf 
Weißjurafelsen unter Kiefern, MTB 6937/33, 15.7.1990, Schuhwerk 90/1338 (nur lebend, 3 di- 
rekt nebeneinander wachsende Rosetten), Kultur-Nr. 90/1338 C, Herbarbeleg vom 6.8.1997. 

Hieracium bifidum Kit. ex Hornem. ssp. subcaesium (Fr.) Zahn 

Die als „microcephal" gesammelte Wildpflanze Nr. 92-50 weist in Kultur 10 mm lange, also 
völlig normal große Hüllen auf. Bei den kultivierten Pflanzen ist die Fleckung der Blätter 
einmal nur sehr schwach, das andere Mal deutlicher ausgeprägt. 

2n = 27 

Schweiz. Wallis : Penninische Alpen, Val dAnniviers, in den ersten Kehren östlich Chippis, 
750 m, Kartierungsfeld 724, MTB 9711/1, 9.6.1988, Gottschlich 10445, Kultur-Nr. 92-50, Her- 
barbelege vom 11.6.1993 (Dubletten in: Hb. Gottschlich), und 10.6.1996. 

Hieracium bupleuroides C.C. Gmel. 

Die Aufsammlung Nr. H-92 war noch von Merxmüller als ssp. leviceps Nägeli & Peter var. 
subepilosum Zahn (Hülle haar- und drüsenlos) bestimmt worden. Abgesehen von Formen, die 
mit relativ schmalen, spitzen Blättern und kleineren Hüllen bereits zu //. glaucum vermitteln 
(und denen Syntypen der ssp. laeviceps, Hieracia Naegeliana 307 und 308, entsprechen), 
zeichnet sich jedoch in Bayern keine klare Gliederung der schon mehrfach als triploid 
nachgewiesenen Sammelart ab. Daher wird Nr. H-92 ebenso wie die von Merxmüller als ssp. 
schenkii Nägeli & Peter var. juratense (Gremli) Zahn bestimmte Nr. H-84 hier nur unter der 
Sammelart geführt. 

2n-27 

Deutschland. Bayern : Regierungsbezirk Oberbayem, Landkreis Reichenhall, Schuttrinne am 
nordwestlichen Ufer des Thumsees, MTB 8242/4, 10.8.1978 Lippert 16638-1 & Merxmüller, 
Kultur-Nr. H-92, Herbarbelege vom Juli 1982 und Sommer 1984. - Regierungsbezirk Oberfranken, 
Landkreis Forchheim, Streitberg, an Felsen der „Klararuh", MTB 6133/3, 1978, Merxmüller & 
Lippert (nur lebend), Kultur-Nr. H-84, Herbarbeleg vom Sommer 1984. 



203 



Hieracium bupleuroides C.C. Gmel. ssp. bupleuroides 

Während die Wildaufsammlung 7 bis ca. 5-6 mm breite Stengelblätter aufweist, besitzen die 
kultivierten Pflanzen nur 4 bis 7 8 mm breite Blätter. In Kultur sind die Hüllen ein wenig 
kleiner und vor allem heller, da die Papillen auf den Hüllschuppen hell- bis weißspitzig sind; 
auch fehlen den kultivierten Pflanzen die an der Wildpflanze ganz spärlich vorhandenen 
Drüsen an der Hülle. 

2n = 27 

Österreich. Tirol : Bezirk Innsbruck, Wettersteingebirge, in der Leutaschklamm südwestlich 
Mittenwald oberhalb des Elektrizitätswerkes, 1025 m, MTB 8533/32, Felsspalten an der Straße, 
Substrattiefe teilweise unterschiedlich, 5.8.1997, Schuhwerk 97/338, Kultur-Nr. 3211, Herbarbeleg 
vom 23.7.1998. 

Hieracium caesium Fr. ssp. caesium 

Die Chromosomenzahl der in Bayern auf den südöstlichsten Fränkischen Jura beschränkten 
und stark gefährdeten Sippe war bisher unbekannt. Bei der kultivierten (halbschattig ge- 
wachsenen) Pflanze nimmt die Haarbekleidung an der Hülle und die Stemhaarbekleidung an 
den Korbstielen gegenüber der Wildaufsammlung etwas ab. Demgegenüber ist in Kultur die 
Zähnung der Blätter erheblich deutlicher: Zähne der Grundblätter bis über 5 mm lang statt 
höchstens 2 mm, am Stengelblatt bis über 1 statt 2-4 mm. Die Form der Zähne bleibt aber 
konstant. 

2n = 36 

Deutschland. Bayern : Regierungsbezirk Niederbayern, Landkreis Kelheim, Fränkischer Jura, 
Felsen westlich unterhalb der Burg Randeck, nördlich des Rathauses von (Neu-)Essing, ca. 450 m, 
MTB 7036/4, 24.6.1996, Zahlheimer & Schuhwerk 96/56, Kultur-Nr. 97-6, Herbarbeleg vom 
12.8.1998. 

Hieracium caesium Fr. ssp. carnosum (Wiesb. ex Dichtl) Zahn 

Die erst bei Dichtl (1884: 57) als Art gültig publizierte Sippe (Gottschlich in litt.) stellt 
möglicherweise einen Endemiten des niederösterreichischen Alpenrandes dar. Die bei Zahn 
(1935: 678) mit „?" aufgeführten, auf Vollmann zurückgehenden bayerischen Funde sind in M 
jedenfalls nicht belegt. Die bei der Wildaufsammlung deutlich gefleckten Blätter sind bei den 
kultivierten Pflanzen kaum bzw. nicht gefleckt. 

2n = 36 

Österreich. Niederösterreich : Alpenrandberge an der Thermenlinie, Hochberg bei Perchtolds- 
dorf, Lücken in felsigen, Sesleria-Te\c\\QX\ Pinus «/g/-a- Wäldern am NE-Hang, Blätter gefleckt, 
13,6.1990, Schuhwerk 90/1101 = Kultur-Nr., Herbarbelege vom 26.8.1997 und 23.6.1998. 

Hieracium dollineri Sch.-Bip. ex Neilr. ssp. dollineri 

Die Chromosomenzahl der von Zahn illegitim ssp. eriopodum (A. Kemer) Zahn genannten 
Sippe und auch der Sammelart war noch unbekannt. Herbarrevisionen in M ergaben, daß in 
Bayern nur diese Unterart vorkommt, streng auf die Täler von Isar und Loisach beschränkt. In 
den Alpen fehlt die Art. Bei der (schattig gewachsenen) kultivierten Pflanze ist die Grund- 
blattrosette stärker betont und auch die Blattrand-Zähnung ausgeprägter als bei der 
Wildaufsammlung. 

2n = 27 

Deutschland. Bayern : Oberbayem, Landkreis Garmisch-Partenkirchen, Niederwerdenfelser 
Land, Isarauen zwischen Mittenwald und Krün, MTB 8533/2, lichter Schneeheide-Kiefemwald, 
18.7.1993, Lorenz [93H/]5, Kultur-Nr. 94-53, Herbarbelege vom 1.7.1998. 



204 



Hieracium franconicum (Griseb.) Zahn 

Unsere Zählung bestätigt eine schon früher in München an Material von der Schwäbischen 
Alb durchgeführte. Bei den kultivierten Pflanzen beider Aufsammlungen sind die Hülle und die 
Blattunterseiten reicher mit Stemhaaren besetzt als bei der Originalaufsammlung. Bei Nr. H-96 
sind die Blätter der kultivierten Pflanze im Gegensatz der Originalaufsammlung nicht gefleckt 
und die Hüllen ärmer behaart. 

2n = 27 • "^ 

Deutschland. Bayern : Regierungsbezirk Oberfranken, Landkreis Forchheim, Gipfelregion und 
Westhänge des Bodenstein (Ehrenbürg) nördlich Schlaifhausen, 450-530 m, MTB 6232/4, 14.6. 
1978, Merxmüller & Lippert 16540 (ES), Kultur-Nr. H-79, Herbarbeleg vom Sommer 1984. - 
Merxmüller & Lippert 16538 (E2), Kultur-Nr. H-96, Herbarbelege vom Juli 1981 und Sommer 
1984. 

Hieracium glaucinum Jord. grex cinerascens (Jord.) 

Die Aufsammlung weicht von ssp. cinerascens (Jord.) Soö ab durch die oberseits 
verkahlenden Blätter und unterscheidet sich von ssp. fraternum (Sudre) Soö durch die am 
Rand der Blätter bleibende borstige Behaarung und die fehlende Zähnung. Auffallig ist die dem 
Boden angepreßte Stellung der Blattrosette. 

2n = 27 

Deutschland. Bayern : Niederbayem, Landkreis Kelheim (oder Oberpfalz, Landkreis Neu- 
markt), Felsen über Altmühlmünster, MTB 7035/2, Mai 1982, Prager Hi. 2/82 (nur lebend), 
Kultur-Nr. H-189, Herbarbeleg vom Sommer 1984. 

Hieracium glaucinum Jord. ssp. basalticum (Sch.-Bip.) J. Duvign. (//. praecox Sch.-Bip. 
ssp. basalticum (Sch.-Bip.) Zahn) 

Die Chromosomenzahl dieser charakteristischen Unterart war bisher unbekannt. In Kultur 
verschwinden bei Nr. 23482 die Stemhaare auf den Blattunterseiten zum Teil; auch die 
Behaarung an der Hülle nimmt etwas ab. Bei Nr. 23468 ist die Zähnung des Blattrandes bei der 
Wildaufsammlung wenig ausgeprägt und auf den Spreitengrund beschränkt, bei den kultivierten 
Pflanzen aber sehr deutlich. 

2n = 27 

Deutschland. Rheinland-Pfalz : Nordpfälzer Bergland, Donnersberg, Ruine Wildenstein im 
Wildensteiner Tal, 9.6.1987, Lippert 23482 = Kultur-Nr. A, B und C, Herbarbelege vom 
17.7.1989 (A) und Juli 1988. - Pfälzerwald, am Weg zur Ruine Landeck, 8.6.1987, Lippert 23468 
= Kultur-Nr., Herbarbeleg vom Juli 1988. 

Hieracium glaucinum Jord. ssp. cinerascens (Jord.) Soö (//. praecox Sch.-Bip. ssp. 
cinerascens (Jord.) Zahn 

Unsere Aufsammlung bestätigt eine bisher übersehene Zählung von Delcourt, die in 
Anmerkung auf dem Etikett zu H. praecox ssp. cinerascens var. expallescens (Sudre) Zahn 
(Soc. Ech. pl. vasc. fasc. 15 n° 7035) angeführt wird. Von Schuhwerk & Lippert (1998) 
wurde die Sippe auch als tetraploid nachgewiesen. Nach den Angaben Zahns (1935: 299 f ) 
war die Unterart aus Bayern bisher nicht bekannt. 

2n = 27 

Deutschland. Bayern : Oberfranken, Landkreis Forchheim, Gipfelregion und Westhänge des 
Bodensteins (Ehrenbürg) nördlich Schlaifhausen, 450-530 m, MTB 6232/4, 14.6.1978, Merx- 
müller & Lippert (nur lebend), Kultur-Nr. H-82, Herbarbelege vom Juli 1981, 28.5.1982 und 
Sommer 1984. 



205 



Hieracium glaucinum Jord. ssp. fraternum (Sudre) Soo (//. praecox Sch.-Bip. ssp. 
fraternum Sudre) 

Die Wildaufsammlung von 90/1035 war mikrocephal mit Hüllenlängen von 4 bzw. 5 mm; 
die kultivierten Pflanzen weisen normale Hüllenlängen von 10-11 mm auf. Die Aufsammlung 
unterscheidet sich von anderen bayerischen Belegen der Sippe durch den nicht so stark 
gestutzten Spreitengrund. 

In = 21 

Deutschland. Bayern : Unterfranken, Landkreis Haßberge, Steinbruch auf der rechten Seite des 
Ebelsbachtales auf der Höhe von Melm bei Neubrunn nördhch Gleisenau, rechts des Baches, MTB 
5930/31, halbschattiger Kiefernwald auf alter Steinbruchhalde, 25.5.1990, Schuhwerk 90/1035, 
Meierott & Eisner; Kultur-Nr. 90/1035, 2. Zählung unter Kultur-Nr. 95/90/1035, Herbarbelege 
vom 10.6., 16., 25., 29.7. und 6.8.1996, 6., 13.6. und 22.9.1997 (zur Verteilung in Soc. Ech. pl. 
vasc. fasc. 28 vorgesehen). 

Hieracium glaucinum Jord. ssp. glaucinum (H. praecox Sch.-Bip. ssp. glaucinum (Jord.) 
Zahn) 

In Kultur (besonders im Sommer) entwickeln die Pflanzen häufiger Blätter mit keihg 
verschmälertem statt gestutztem Spreitengrund. Außerdem sind die Hüllen etwas kleiner und 
die Haare daran werden weniger bzw. fehlen. 

2n = 27 

Deutschland. Bayern : Unterfranken, Landkreis Main-Spessart, südlicher Ausläufer des Stein- 
berges 1,5 km südöstlich Karlstadt (Weinlage „Stettener Stein"), nordwestexponierter Hang im 
Einschnitt nördlich P. 273, ca. 260 m, MTB 6024/42, ± offener Rasen mit Saumarten über Mu- 
schelkalkschotter, zusammen mit Helianthemum apenninum x ovatum, 1.6.1995, Meierott <& 
Schuhwerk 95/50 a, Kultur-Nr. 96/36, Herbarbelege vom 2.6. (Dubl. in: GLM, Hb. Gottschlich), 
20. und 26.8.1997. 

Hieracium glaucinum Jord. ssp. prasiophaeum (Arv.-Touv. & Gautier) (//. silvaticum L. 
grex praecox ssp. prasiophaeum (Arv.-Touv. & Gautier) Zahn, H. praecox Sch.-Bip. ssp. 
gougetianum (Gren. & Godron) Zahn) 

Eine gültige Kombination für diese Sippe unter H. glaucinum gibt es nocht nicht, doch 
sollten vor einer formellen Umkombination die jeweiligen Typen der Basionyme eingesehen 
werden. Immerhin stellen Arvet-Touvet und Gautier ihr H. prasiophaeum in eine „groupe 
Olivacea", während Zahn (1900: 1781) ausdrücklich feststellt, daß sie „mit den Cerinthoidea 
nichts zu tun hat". Die triploide Stufe war für diese Unterart bereits von Mills & Stach 
(1974) festgestellt worden. In Kultur sind die Hüllen gelegentlich etwas kleiner oder auch 
größer, Stemhaare und Drüsen daran etwas vermindert, während die spärliche Hüllenbehaarung 
fast ganz verschwindet. Die Blattspreiten werden etwas breiter, die Fleckung verschwindet bei 
Gewächshauspflanzen, kann aber bei trocken gewachsenen Freilandpflanzen stärker sein, bei 
denen wiederum die Blattzähnung stark reduziert ist. 

2n = 27 

Deutschland. Bayern , Oberfranken, Landkreis Forchheim, Gipfelregion und Westhänge des 
Bodensteins (Ehrenbürg) nördlich Schlaifhausen, 450-530 m, MTB 6232/4, 14.6.1978, Merxmul- 
ler & Lippert 16536 (E 1) (Dubl. in MSB 20055), Kultur-Nr. H-78, Herbarbelege vom Juli 1981, 
Mai 1982 und Sommer 1984. - Merxmüller & Lippert (nur lebend), Kultur-Nr. H-81, Herbarbelege 
vom Juli 1981 und April 1982. - Landkreis Lichtenfels, Staffelberg bei Staffelstein, MTB 5932/1, 
Felsspalten, 26.6.1982, Lippert 18268, KulUir-Nr. H-195, Herbarbelege vom Sommer 1984. 



206 



Hieracium glaucinum Jord. ssp. recensitum (Jord. ex Boreau) Gottschlich (//. praecox Sch.- 
Bip. ssp. recensitum (Jord. ex Boreau) Sudre 

Die Unterart weist vor allem am Spreitengrund recht variable Blattformen auf; ihre 
Chromosomenzahl war bisher noch nicht ermittelt. Die bei der Wildaufsammlung von Nr. 
3100 lebend hell lauchgrünen, stark und deutlich gefleckten Blätter sind bei der kultivierten 
Pflanze rein grün und ungefleckt. Bei Nr. 92-35 sind die Belege von 1996 deutlich gefleckt, 
diejenigen von 1993 dagegen nur schwach bzw. gamicht. 

2n-27 

Deutschland. Bayern : Oberpfalz, östlicher Landkreis Neumarkt, SW-Hang des Anteiberges (P. 
534,8) südlich Ransbach, um 470 m, MTB 6636/34, Halbtrockenrasen (Mesobromion), ehemals 
unter lichtem Kiefernschirm, nahe dem Fundort von H. wiesbaurianum glaucocinerascens, 5.7. 
1995, Meyer & Schuhwerk 96/141, Kultur-Nr. 3100, Herbarbeleg vom 30.7.1998. - Unter- 
franken, Landkreis Würzburg, Südteil des Kaimut nördlich Homburg, MTB 6223/2, Juni 1990, 
Meierott 90/283 (Hb. Meierott), Kultur-Nr. 92-35, Herbarbeleg vom Juli 1993 (Dubl. in Hb. 
Gottschlich) und 11.7.1996. 

Hieracium glaucinum Jord. ssp. similatum (Jord. ex Boreau) Gottschlich (//. praecox Sch.- 
Bip. ssp. similatum (Jord. ex Boreau) Zahn) 

Von typischen im Freiland gesammelten Belegen dieser Sippe, deren Chromosomenzahl 
noch nicht bekannt war, weichen die kultivierten Pflanzen in ihrer Blattform ab. Ihr Spreiten- 
grund ist schwach keilig verschmälert, höchstens das jeweils älteste Blatt weist den 
kennzeichnenden, gestutzten oder schwach herzförmigen Grund mit beiderseits 1-2 Zähnen 
auf Bei Wildpflanzen dagegen ist allenfalls beim jüngsten Blatt der Spreitengrund etwas 
verschmälert. Zahn unterscheidet allerdings innerhalb der Unterart Varietäten mit sogar 
länglich-lanzettlichen Blättern mit verschmälertem Grund, die nach Gottschlich (in litt.) jedoch 
abgetrennt werden müssen. 

2n = 27 

Deutschland. Bayern : Landkreis Neustadt a. d. Aisch-Bad Windsheim, oberhalb einer Wald- 
wiese südöstlich Markt Sugenheim, MTB 6428/2, auf Blasensandstein-Verwitterung, 18.9.1987, 
Subal (nur lebend), Kultur-Nr. 2491, 2492, 2493, 2494, Herbarbelege jeweils Juli 1988. 

Hieracium glaucum All. ssp. isaricum (Nägeli ex J. Hofm.) Nägeli & Peter 

Die zahlreichen aus den bayerischen Alpen und ihrem Vorland genannten Unterarten von 
H. glaucum lassen sich auf eine Sippe ohne Stemhaare auf den Blättern und eine weiter als bei 
Zahn gefaßte ssp. isaricum mit Stemhaaren auf den tendenziell etwas breiteren Blättern 
aufteilen. Diese Unterart, deren Chromosomenzahl bisher noch nicht bekannt war, kommt nur 
in den nördlichen und nordöstlichen Kalkalpen vor und erreicht z.B. die Zentralalpen nicht 
mehr. In Bayern sind die beiden Sippen allerdings chorologisch nicht getrennt und kommen 
gelegentlich auch zusammen an den gleichen Fundorten vor. 

2n = 27 

Deutschland. Bayern : Regierungsbezirk Oberbayem, Landkreis Traunstein, im Fischbachtal 
südlich Laubau südwestlich Ruhpolding, MTB 8241/4, Felsschutt, 18.6.1996, Lippert 27232 (nur 
lebend), Kultur-Nr. 3089, Herbarbeleg vom 30.7.1998. 

Hieracium harzianum Zahn 

Die Chromosomenzahl der morphologisch zwischen H. franconicum und H. laevigatum 
stehenden, auf der Ehrenbürg im nördlichen Fränkischen Jura endemischen Art war bisher 
unbekannt. Mit nur wenigen Stengelblättem entspricht die kultivierte Pflanze der von Zahn 
unterschiedenen ssp. pseudofranconicum Zahn. 



207 



2n = 36 

Deutschland. Bayern : Oberfranken, Landkreis Forchheim, südlichster Felskopf an der Ost- 
seite des Rodenstein an der Ehrenbürg bei Forchheim, MTB 6232/44, 17.6.1989, Wagenknecht, 
Schuhwerk et al. (nur lebend), Kultur-Nr. 2625, Herbarbelege vom 7.7.1998. 

Hier actum humile Jacq. ssp. pseudocottetU (Zahn) Zahn 

Der mäßige Stemhaarbesatz der Korbstiele, nach Zahn das einzige durchgreifend von der 
Typus-Unterart differenzierende Merkmal, kann bei einzelnen Korbstielen der gleichen Pflan- 
ze unterschiedlich sein. Da in den bayerischen Alpen auch Mischpopulationen mit ssp. humile 
beobachtet wurden, dürfte die Bewertung der Sippe als Varietät (wie von Zahn ursprünglich 
ebenfalls eingestuft) angemessener sein. Ihre Chromosomenzahl war bisher unbekannt. 

2n = 27 

Deutschland. Bayern : Karwendelgebirge, am Oberen Lichteck beim Schafreuter (Vorkar- 
wendel), 1965 m, südexponierter Felsabbruch eines Plattenkalk-Schichtpakets, Felsflur mit alpi- 
nem Charakter {Draba tomentosa, Festuca alpina und Poa cenisia in der Nähe), MTB 8434/4, 
11.8.1991, Saitner B 28, det. F. Schuhwerk, Kultur-Nr. 92-22, Herbarbeleg vom Juli 1993 (Dubl. 
in: GLM, Hb. Gottschlich). 

Hieracium illyricum Fr. ssp. illyricum 

Die Chromosomenzahl auch der - morphologisch zwischen H. porrifolium und H. laevi- 
gatum stehenden - Sammelart war bisher unbekannt. 

2n = 36 

Slowenien. Julische Alpen, Baca-Tal, Straße von Petrovo brdo nach Zg. Sorica, 900 m, Wald- 
rand, 1991, E. & M. Pahl 2878, det. F. Schuhwerk, Kultur-Nr. 92-40, Herbarbeleg vom Juli 1993. 

Hieracium jurassicum Griseb. ssp. jurassicum {H. juranum Fr. ssp. juranum) 

Die Chromosomenzahl der Sammelart war bislang unbekannt. Bei den kultivierten Pflanzen 
ist der Sternhaarbesatz der Hüllen etwas dichter als bei der Wildaufsammlung. Erst in Kultur 
bilden die Pflanzen eine Rosette aus drei bis fünf Grundblättem. 

2n = 27 

Österreich. Kärnten : Bezirk Spittal a. d. Drau, oberhalb Innerfragant, Ode. Flattach, Hohe 
Tauem, Sadniggruppe, am Aufstiegsweg zur Fraganter Hütte bei der Unfallstelle des Hüttenwirtes, 
ca. 1450 m, MTB 9044/14, Fichtenwaldsaum, 15.8.1994, Schuhwerk Nr. 94/832 et a\., Kulttir-Nr. 
96-39, Herbarbelege vom 11.8.1997, 7. und 17.7.1998. 

Hieracium laevigatum Willd. 

Die Aufsammlung gehört trotz relativ zahlreicher, kräftiger Drüsen an der Hülle nicht zu 
ssp. gothicum (Fr.) Zahn. In Kultur verringert sich die Drüsenzahl etwas und die Drüsen 
werden heller. 

2n = 27 

Deutschland. Bayern : Regierungsbezirk Unterfranken, Rhön, südsüdöstlich Bischofsheim, 
Irenkreuz gegen Neustädter Haus, 650 m, MTB 5626/11, 29.8.1996, Lippert 27395, Kultur-Nr. 
97-42, Herbarbeleg vom 12.8.1998. 

Hieracium laevigatum Willd. ssp. perangustum (Dahlst.) Zahn 

Die vorliegende Aufsammlung weicht durch gelbbräunliche, aber nicht dunkle Griffel etwas 
von der Beschreibung dieser Unterart ab, deren Chromosomenzahl bisher unbekannt war; die 
Sammelart war bisher schon als di- und triploid bekarmt. 

2n = 27 



208 

Deutschland. Baden- Württemberg : Regierungsbezirk Südbaden, Landkreis Waldshut, Gemeinde 
Ibach, Südschwarzwald, an dem die Mulde „Loh" westlich begrenzenden Rücken wenig südlich P. 
1086,7 nordwestlich Ober-Ibach, 1080 m, MTB 8214/32, brachliegender, saumartenreicher 
Magerrasen, 23.8.1993, Schuhwerk 93/434 & L Schuhwerk (Dubl. (3) in: GLM, NY, Hb. 
Gottschlich), Kultur-Nr. 2852, Herbarbeleg vom 27.7.1998. 

Hieracium lingelsheimii Pax (//. goemoerense Borbäs ex Zahn) 

Die Chromosomenzahl der morphologisch zwischen H. bupleuroides und H. laevigatum 
stehenden, in der Niederen Tatra endemischen Art war bisher unbekannt. Während die Pflanze 
der Wildaufsammlung keine Grundblätter aufweist, besitzen die kultivierten Pflanzen teilweise 
eine Grundblattrosette. Die Länge der Blattzähne schwankt bei den kultivierten Pflanzen. 

2n = 36 

Slowakei. Streydoslovensk : Niedere Tatra, Poprad - Cervena Skala, an der Straße, 24.8.1991, 
Pahl 432], det. F. Schuhwerk, Kultur-Nr. 94-82, Herbarbelege vom 6. und 29.8.1996, 29.7.1997 
(Dubl. in: Hb. Gottschlich), 20. und 23.7.1998. 

Hieracium lycopifolium Froel. ssp. lycopifolium 

Die Chromosomenzahl der in Deutschland auf den Südwesten beschränkten Sammelart war 
noch unbekannt. Die schwache Bewimperung der Ligulaezähne bleibt bei einzelnen kulti- 
vierten Pflanzen aus. Teils stärker, teils schwächer als bei der Wildaufsammlung ist bei den 
kultivierten Pflanzen der Stemhaarbesatz der oberen Stengelblätter. Auch die charakteristische 
Blattrand-Zähnung ist bei den kultivierten Pflanzen unterschiedlich intensiv. 

2n = 36 

Deutschland. Baden- Württemberg : Südbaden, Landkreis Waldshut, Südschwarzwald, Schwarza- 
tal am Abhang unter dem alten Porphyrsteinbruch oberhalb P. 459,8 am Ostufer des Witznauer 
Beckens, MTB 8315/12, 470 m, lichtes Gebüsch aus Quercus petraea auf Porphyrschutt, 
24.9.1981, Schuhwerk 81/680, Kultur-Nr. H-151, Herbarbelege vom Sommer 1982 und 1984, 
September 1987. 2. Zählung unter Kultur-Nr. 94-100, Herbarbelege vom 25.10.1995 (Dubl. in Hb. 
Gottschlich), 29.8.1996 (8 Dubl.) und 24.8.1998 (Dubl. in GLM). 

Hieracium maculatum Schrank ssp. maculatum 

Die Typus-Unterart ist die in Bayern häufigste und am weitesten verbreitete Sippe der 
Sammelart, für die bisher nur die triploide Stufe nachgewiesen war. 

2n = 36 

Deutschland. Bayern : Regierungsbezirk Oberpfalz, Landkreis Regensburg, Scheuchenberg 
östlich über Sulzbach/Donau, MTB 6939/3, wärmeliebender Eichenwald auf Gneis (Luzulo- 
Quercetum), Juli 1980, Schuhwerk 80/405, Kultur-Nr. H-121, Herbarbelege vom Juli 1982 und 
Sommer 1984. 

Hieracium nigrescens Willd. ssp. cochleare (Huter) Zahn 

Die Sammelart ist bisher als tri- und tetraploid nachgewiesen, die Chromosomenzahl der 
Unterart war noch nicht bekannt. 

2n = 27 

Deutschland. Bayern : AUgäuer Alpen, Fellhom - Schlappolt, Grat, Übergang Seslerietalia - 
Nardion, 1989, Meyer, Kultur-Nr. 2675, Herbarbeleg vom 13.6.1997. 

Hieracium piliferum Hoppe ssp. hololeptum Nägeli & Peter 

Die zur glanduliferum-Gmppe gehörige Sippe unterscheidet sich von ssp. glanduliferum 
nur durch gelbe Griffel (benachbarte Wildaufsammlung, Merxmüller 33395 & Angerer); die 
Bestimmung stammt noch von Merxmüller. Für die Sammelart war die tetraploide Stufe bisher 
nicht nachgewiesen. Die im Gewächshaus kultivierte Pflanze unterscheidet sich von der Wild- 



209 



aufsammlung Merxmüller 33395 & Angerer aus dem gleichen Gebiet durch deutlich geringeren 
Drüsenbesatz des Schaftes, wobei die Drüsen länger und heller sind. Die Haarbekleidung der 
Hülle ist bei der kultivierten Pflanze weniger dicht und die Haare sind heller. 

2n = 36 

Schweiz. Tessin : oberstes Maggiatal, Naret, ca. 2000 m, August 1982, Merxmüller & Angerer 
(nur lebend), Kultur-Nr. H-203, Herbarbeleg vom Sommer 1984. 

Hieracium platyphyüum Arv.-Touv. ssp. maranzae (Murr & Zahn) Zahn 

Die Chromosomenzahl sowohl der Sammel- wie der Unterart war noch unbekannt. In Kul- 
tur werden die Blätter zum Teil breit oval und erinnern an das mutmaßlich verwandte H. 
racemosum ssp. leiopsis, ohne jedoch dessen charakteristische Blattrand-Zähnung aufzu- 
weisen. Von Zahn in der Hieraciotheca Europaea (Nr. 700) ausgegebene Pflanzen der ssp. 
hostianum Wiesb. ex Zahn unterscheiden sich von der vorliegenden Aufsammlung durch die 
stärkere Behaarung von Stengel und Blättern. 

2n = 27 

Italien. Südtirol : Zillertaler Alpen, Terenten, beim Friedhof, 1210 m, MTB 9136/4, Fichten- 
wald, 14.9.1992, Gottschlich 21163, Kultur-Nr. 94-76, Herbarbelege vom 25.10.1995, 29.8.1996, 
20.8.1997 (4 Dubl.) und 12.8.1998. 

Hieracium pseudocorymbosum Gremli ssp. petryanum Zahn 

Die Chromosomenzahl der in Deutschland nur im Südostschwarzwald (Schwarzatal) vor- 
kommenden Art war bisher unbekannt. Die für eine sogenannte 'Zwischenart' ungewöhnliche 
Diploidie (vgl. Merxmüller 1975) könnte auf eine Entstehung aus tetraploidem H. 
lycopifolium als Pollen-Donator und im Gebiet wohl wie üblich diploidem H. umbellatum 
hinweisen, doch stehen Untersuchungen hierzu noch aus. Die einzelnen Populationen der 
Sippe liegen in dem kleinen Wuchsgebiet jedoch nicht immer in direkter Nachbarschaft von H. 
lycopifolium. Blattrand-Zähnung und Blattbreiten sind bei den kultivierten Pflanzen etwas 
unterschiedlich. Der Stemhaarbesatz der oberen Stengelblätter - etwas schwächer als bei H. 
lycopifolium - bleibt in Kultur meist erhalten. Nur die kräftigsten Pflanzen bilden an der Spitze 
des Korbstandes eine deutliche Dolde aus. 

2n=18 

Deutschland. Baden- Württemberg : Südbaden, Landkreis Waldshut, Südschwarzwald, Schwarza- 
tal am Abhang unter dem alten Porphyrsteinbruch oberhalb F. 459,8 am Ostufer des Witznauer 
Beckens, 470 m, MTB 8315/12, wärmeliebendes Gebüsch aus Quercus petraea auf Porphyrschutt, 
22.9.1990, Schuhwerk 90/1445 & Gottschlich, Kultur-Nr. 94-101, Herbarbelege vom 26.8. und 
25.9.1997, 24.8.1998 (Dubl. in GLM). 

Hieracium racemosum Waldst. & Kit. ssp. leiopsis Murr & Zahn 

Die Chromosomenzahl der von Zahn später illegitim in ssp. leiobium Murr & Zahn 
umbenannten Sippe war noch unbekannt; für die Sammelart war die diploide Stufe bisher nicht 
nachgev^esen. Ob die von Zahn hierher gerechneten westalpischen und Südtiroler Funde 
tatsächlich zu dieser Sippe gehören, bleibt an Belegen nachzuprüfen. Im mittleren Tiroler 
Inntal besiedelt sie ein nach derzeitiger Kenntnis ca. 50 km langes Areal etwa zwischen Stams 
und Jenbach. Stichproben ergaben, daß vor allem im Innsbrucker Nahbereich zwar manche der 
von Zahn genannten Wuchsorte vernichtet sein dürften; in der weiteren Umgebung gibt es 
jedoch weit mehr als die dort aufgeführten Vorkommen. Durch die lang gefransten Ränder der 
Gruben im Korbboden fällt die Sippe etwas aus dem Rahmen der Sammelart H. racemosum. 
Der Stemhaarbesatz auf den Unterseiten der oberen Stengelblätter - bei der Wildaufsammlung 
von Nr. 94-1 10 auf den Mittelnerv beschränkt - greift bei den kultivierten Pflanzen auch auf 



210 



die Fläche und die Blattoberseite über. Auch die Hülle ist bei den kultivierten Pflanzen etwas 
reicher an Stemhaaren. Die charakteristische Blattzähnung kann in Kultur bei kümmerlich 
gewachsenen Pflanzen reduziert sein. 

2n=18 

Österreich. Tirol : Bezirk Innsbruck, südliches Mittelgebirge, um „MäusI" im Zimmertal 
südlich gegenüber Hall i. Tirol, ca. 700 m, MTB 8735/1, Straßenböschungen, 26.9.1993, Bräuti- 
gam & Schuhwerk 93/526 (als H. cf. platyphyllum Arv.-Touv., rev. G. Gottschlich 1998), Kultur- 
Nr. 94-111, Herbarbelege vom 25.10.1995 (Dubl. in GLM) und 25.9.1997. 
2n = 27 

- Gleicher Fundort, gleiche Daten, Nr. 93/527 - Bg. 2, Kultur-Nr. 94-110, Herbarbelege vom 
29.8.1996 (4 Dubl.), 26.8.1997 und 24.8.1998. 

Hieracium rigidiceps S. Braut. & V. Braut. 

Die Chromosomenzahl der aus Sachsen beschriebenen, aus Bayern bisher nur in Nord- 
bayern selten nachgewiesenen Art war unbekaimt. In Kultur nimmt sowohl die Hüllen- 
behaarung als auch der Sternhaarbesatz auf den Blattunterseiten ab; die Zähnung der Blätter 
wird deutlicher. 

2n = 27 

Deutschland. Bayern : Unterfranken, Landkreis Rhön-Grabfeld, westlich Oberweisenbrunn/ 
Neuwildflecken gegen die Schwedenschanze, MTB 5525/4, Muschelkalk, Wegrand, 26.6.1988, 
Lippen 23767 = Kultur-Nr., Herbarbeleg vom Juli 1990. 

Hieracium sabaudum L. ssp. sublactucaceum Zahn 

Die Chromosomenzahl der nach Zahn (1938: 557) „ohne Grenze in ssp. vagum überge- 
henden" Sippe war noch unbekannt. Die Behaarung der Blattunterseiten bei der Wild- 
aufsammlung verliert sich in Kultur weitgehend. 

2n = 27 

Deutschland. Bayern : Oberfranken, Landkreis Hof/Wunsiedel, Martinlamitzer Forst am 
Großen Kornberg westlich Selb, MTB 5838/11, Kiefemwaldsaum, Oktober 1993, Horbach 93/ 
1731, Kultur-Nr. 94-79, Herbarbelege vom 25.10.1995 (3 Dubl.) und 26.8.1997 (8 Dubl.). 

Hieracium sabaudum L. ssp. vagum (Jord. ex Boreau) Zahn 

2n = 27 

- Niederbayem, Landkreis Freyung-Grafenau, bei Perlesreuth, MTB 7246/2, 1978, Bauer (nur 
Achänen), Kultur-Nr. H-88, Herbarbelege vom 15.8.1982 und Sommer 1984. 

Hieracium saxifragum Fr. ssp. dufftii Zahn 

Die Chromosomenzahl auch der Sammelart war bisher unbekannt. Anschließend an ihr 
Thüringer Hauptareal waren von der H. lachenalü bereits sehr nahe stehenden Sippe wenige 
primäre Wuchsorte im Frankenwald bekannt. Nach neueren Funden von H.D. Horbach (M) 
konnte sie sich vermutlich von einem von diesen - der Wojaleite bei Wurlitz - ausgehend und 
wohl mit Serpentinschotter als Wegebaumaterial verschleppt, ein kleines Sekundärareal an 
Wegrändern, in Steinbrüchen u. ä. erobern. An den Belegen kultivierter Pflanzen sind die bei 
der Wildaufsammlung gelbköpfigen, zarten Drüsen an der Hülle gelegentlich etwas kräftiger 
und dunkler. 

2n = 36 

Deutschland. Bayern : Oberfranken, Frankenwald, Petersgrat gegenüber Lamitzmühle am 
rechten Ufer der Sächsischen Saale nördlich Joditz, am Fuß des nördlichen Grates, MTB 5637/1, 
felsiger Rain am Wegrand in Hangfußwald aus Fagus syhatica und Carpinus betulus, 9.8.1986, 



211 

Schuhwerk 86/620, Kultur-Nr. 87-35, Herbarbelege vom Juli 1988, 17.7.1989, 20.7.1990, Juli 
1992 (Dubl. in: GLM, Hb. Gottschlich) und Juli 1993. 

Hieracium schmidtii Tausch ssp. comatulum (Jord. ex Boreau) Gottschlich (//. pallidum Biv. 
ssp. comatulum (Jord. ex Boreau) Zahn 

Die Chromosomenzahl dieser weitverbreiteten Unterart von H. schmidtii war bisher 
unbekannt. Zahn nennt (1935: 245, alle unter H. pallidum) außer dieser Sippe für den 
Rabenstein auch (ebd. 261) die ssp. jovimontis Zahn, für das Höllental (ebd. 249) ssp. 
graniticum (Sch.-Bip.) Zahn und gibt (1929: 1269) „bei Stehen ähnlich ssp. candicans 
(Tausch) Zahn" an, womit ebenfalls das Höllental-Vorkommen gemeint sein dürfte. Keine 
dieser Sippen außer ssp. comatulum wurde im Höllental und am Rabenstein gefunden und 
Belege für diese Angaben gibt es zumindest in M nicht. Für das Vorkommen im Höllental war 
dies schon von Schack ( 1 94 1 : 8 1 ) festgestellt worden. 

2n = 27 

Deutschland. Bayern : Oberfranken, Landkreis Hof, Höllental nördlich Naila, Felsrinne süd- 
östlich unter dem Hirschsprungfels, MTB 5636/1, bemooste Felsplatte unter lichtem Gebüsch, 
10.7.1986, Buttler 29400 & Schuhwerk 86/335, Kulttir-Nr. 86-42, Herbarbeleg vom Juli 1988. - 
Unterfranken, Landkreis Rhön-Grabfeld, Rabenstein-Gipfel und -Südhang westlich Oberweißen- 
brunn, MTB 5525/3, 26.6.1988, Lippert 23797 = Kultur-Nr., Herbarbeleg der Wieder-Aussaat 94- 
108 vom 25.7.1996. 

Hieracium schmidtii Tausch ssp. kalmutinum (Zahn) Gottschlich (//. pallidum Biv. ssp. 
kalmutinum (Zahn) Zahn ex Kneucker) 

Die Chromosomenzahl der ökologisch von den übrigen Unterarten von H. schmidtii sehr 
abweichenden Sippe war bisher unbekaimt. 

2n = 27 

Deutschland. Bayern : Unterfranken, Landkreis Main-Spessart, Kalbenstein nördlich Karlstadt 
zwischen Klettergarten und Grainberg südwestlich unter dem Rosenholz, MTB 6024/2, Trocken- 
rasen, 13.6.1986, Schuhwerk (nur lebend), Kultur-Nr. 86-17, Herbarbeleg vom Juli 1988. 

Hieracium sparsum Friv. ssp. grisebachii (A. Kern.) Zahn 

Triploidie war bisher nur für griechische Herkünfte von H. sparsum s.str. bekannt 
(Schuhwerk & Lippert 1998). Das z.B. von Sell & West (1976) davon unterschiedene H. 
silesiacum Krause, zu dem nach diesen Autoren auch die ssp. grisebachii zu stellen ist, war 
bisher als tetraploid bekaimt (Chrtek 1996). 

2n = 27 

Österreich. Tirol : Ötztaler Alpen, unterhalb Obergurgl, Tratten bei Poschach, 1900 m, MTB 
9132/1, Felsband mit Laserpitium halleri, Veronica fruticans, 15.8.1982, Schuhwerk 82/292, 
Kultur-Nr. 87-32, Herbarbelege vom Juli 1990 und 22.7.1997, 2. Zählung unter Kultur-Nr. 94-1 14 
(Wiederaussaat im Jahr 1990 entnommener Achänen), Herbarbelege vom 15.7.1996. 

Hieracium umbellatum ssp. brevifolioides Zahn 

Unsere Zählung bestätigt die von Strid «fe Franzen (1981: 842) an griechischem Material 
ermittelte Zahl. Als sexuelle Sippe zeigt die Unterart teilweise eine beträchtliche Variabilität, 
z.B. bezüglich der relativen Blattbreiten oder der Zähnung des Spreitenrandes. Bei den kulti- 
vierten Pflanzen (Gewächshaus wie Freiland) sind die Hüllen heller als bei der Wildauf- 
sammlung, grün statt schwärzlich. 

2n= 18 

Österreich. Tirol : Bezirk Innsbruck, südliches Mittelgebirge, am Blasienberg westlich Vols, 
Gemeinde Axams auf der Kuppe bei Wasserbehälter, ca. 650-680 m, Waldsaum, MTB 8733/24, 



212 



27.9.1993, Bräutigam & Schuhwerk 93/534, Kultur-Nr. 94-113, Herbarbelege vom 22.9.1997 
und 24.9.1998. 



Hieracium umbellatum L. ssp. umbellatum 

Die Herbarbelege von Nr. H-45 der einzelnen Jahre variieren etwas in der Färbung der 
Hüllblätter; zum Teil weisen sie eine deutliche Grundblattrosette auf. Daß die Blattränder der 
Wildaufsammlung von Nr. 2635 kaum angedeutet gezähnt sind, bleibt auch bei den kultivierten 
Pflanzen erhalten. 

2n=18 

Deutschland. Bayern : Regierungsbezirk Schwaben, Landkreis Aichach-Friedberg, sandiger 
Hohlweg bei Happach ca. 5 km südöstlich Altomünster, Sept. 1974, Grau (nur lebend), Kultur-Nr. 
H-45, Herbarbelege von 1974, September 1978 und Sommer 1984. - Regierungsbezirk Oberpfalz, 
Landkreis Regensburg, Fränkischer Jura, knapp südlich unter der Burg von Kallmünz, 430 m, MTB 
6837/2, Festuca pallens-YehhändQT, 30.7.1989, Schuhwerk 89/85, Hüllblätter abstehend bis spar- 
rig zurückgekrümmt, Griffel gelb, deutlich früher blühend als andere Pflanzen von H. umbellatum 
am gleichen Wuchsort, Kultur-Nr. 2635, Herbarbeleg vom 22.9.1997. 

Tab. 1: Chromosomenzahlen und Herkunft der untersuchten Sippen. Name unterstrichen: 
Chromosomenzahi bzw. diese Ploidiestufe für das betreffende Taxon bisher unbekannt. 



Taxon 


Chromo- 
somenzahl 


Anzahl der 
Zählungen 


Herkunft 


alpinum ssp. alpinum 


2n = 27 


1 


Deutschland Bayern 


alpinum ssp. halleri 


2n = 27 


2 


Österreich Tirol 


amplexicaule ssp. berardianum 


2n = 27 


1 


Österreich Tirol 


atratum ssp. atratum 


2n = 27 


2 


Italien Trentino 


atratum ssp. schroeterianum 


2n = 27 


1 


Schweiz Graubünden 


bifidum ssp. canitiosum 


2n = 27 


1 


Deutschland Bayern 


bifidum ssp. eriopodoides 


2n = 27 


2 


Deutschland Bayern 


bifidum äff ssp. psammogenes 


2n = 27 


3 


Deutschland Bayern 


bifidum ssp. stenolepis 


2n = 36 


1 


Deutschland Bayern 


bifidum ssp. subcaesium 


2n = 27 


1 


Schweiz Wallis 


bupieuroides 


2n = 27 


2 


Deutschland Bayern 


bupleuroides ssp. bupieuroides 


2n = 27 


1 


Österreich Tirol 


caesium ssp. caesium 


2n = 36 


1 


Deutschland Bayern 


caesium ssp. carnosum 


2n = 36 


1 


Österreich Niederösterreich 


dollineri ssp. dollineri 


2n = 27 


1 


Deutschland Bayern 


franconicum 


2n = 27 


2 


Deutschland Bayern 


glaucinum ssp. basalticum 


2n = 27 


2 


Deutschland Rheinland-Pfalz 


glaucinum grex cinerascens 


2n = 27 


1 


Deutschland Bayern 


glaucinum ssp. cinerascens 


2n = 27 


1 


Deutschland Bayern 


glaucinum ssp. fraternum 


2n = 27 


2 


Deutschland Bayern 


glaucinum ssp. glaucinum 


2n = 27 


1 


Deutschland Bayern 



213 



Taxon 


Chromo- 
somenzahl 


Anzahl der 
Zählungen 


Herkunft 


glaucinum ssp. prasiophaeum 


2n = 27 


3 


Deutschland Bayern 


glaucinum ssp. recensitum 


2n = 27 


2 


Deutschland Bayern 


glaucinum ssp. similatum 


2n = 27 


4 


Deutschland Bayern 


giaucum ssp. isaricum 


2n = 27 




Deutschland Bayern 


harzianum 


2n = 36 




Deutschland Bayern 


humile ssp. pseudocottetii 


2n = 27 




Deutschland Bayern 


illyricum ssp. illyricum 


2n-36 




Slowenien 


jurassicum ssp. jurassicum 


2n = 27 




Österreich Kärnten 


laevigatum 


2n = 27 




Deutschland Bayern 


laevigatum ssp. perangustum 


2n = 27 




Deutschland Baden 


lingelsheimii 


2n = 36 




Slowakei Niedere Tatra 


lycopifolium ssp. lycopifolium 


2n = 36 


2 


Deutschland Baden 


maculatum ssp. maculatum 


2n = 36 




Deutschland Bayern 


nigrescens ssp. cochleare 


2n = 27 




Deutschland Bayern 


piliferum ssp. hololeptum 


2n = 36 




Schweiz Tessin 


platyphyllum ssp. maranzae 


2n = 27 




Italien Südtirol 


pseudocorymbosum ssp. petryanum 


2n= 18 




Deutschland Baden 


racemosum ssp. leiopsis 


2n= 18 




Österreich Tirol 


racemosum ssp. leiopsis 


2n = 27 




Österreich Tirol 


rigidiceps 


2n = 27 




Deutschland Bayern 


sabaudum ssp. sublactucaceum 


2n = 27 




Deutschland Bayern 


sabaudum ssp. vagum 


2n = 27 




Deutschland Bayern 


saxifragum ssp. dufftii 


2n = 36 




Deutschland Bayern 


schmidtii ssp. comatulum 


2n = 27 


2 


Deutschland Bayern 


schmidtii ssp. kalmutinum 


2n = 27 




Deutschland Bayern 


sparsam ssp. grisebachii 


2n = 27 


2 


Österreich Tirol 


umbellatum ssp. brevifolioides 


2n= 18 




Österreich Tirol 


umbellatum ssp. umbellatum 


2n= 18 


2 


Deutschland Bayern 



Auch diesmal erhielten wir bei unseren Untersuchungen vielerlei Hilfe und Unterstützung. Prof 
Dr. J. Grau gestattete die Kultur unserer Pflanzen im Gewächshaus und im Freilandteil des Institu- 
tes für Systematische Botanik der LMU München im Botanischen Garten München, förderte un- 
sere Arbeiten in vielfacher Hinsicht und war stets aufgeschlossen für unsere Sonderwünsche. A. 
Hartmann betreute unsere Pflanzen in bewährter Weise. Die Chromosomenzählungen führten mit 
großer Sorgfalt M. Erben, G. Heubl, C. Kern, G. Kühlhom, Be. und Bi. Lippert, B. Treppenhauer, 
R. Vogt sowie vor allem 1. Sebek und E. Vosyka durch. Viele Helfer brachten Hieracien von ihren 
Exkursionen mit oder führten uns zu Fundorten bemerkenswerter Arten: E. Albertshofer, A. Bau- 
er, G. Gottschlich, J. Grau, H.D. Horbach, J. Krach, W. Lorenz, A. Mayer, L. Meierott, N. Meyer, 



214 

M. Nydegger, E. und M. Pahl, L. Prager, A. Saitner, W. Subal und J. Wagenknecht. G. Gottschlich 
(Tübingen) korrigierte mit großer Sorgfalt unser Manuskript. Ihnen allen danken wir herzlich. 



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Hieracium L. subgenus Pilosella (Hill) S.F. Gray by flow cytometric DNA analysis.-Fol. 

Geobot. Phytotax. 31: 315-321. 
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the Sudeten Mts. and the Western and Ucrainian Eastern Carpathians. - Fragm. Florist, et 

Geobot. 41(2): 783-790. 

- 1997: Taxonomy of the Hieracium alpinum group in the Sudeten Mts., the West and the 
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DiCHTL, A. 1884: Ergänzungen zu den „Nachträgen zur Flora von Nieder-Österreich". - 

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geographica. Sect. I. Revisio specierum Hieracii, in Europa sponte crescentium. Göttingen. 
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Arnolds Habichtskraut {Hieracium wiesbaurianum ssp. arnoldianum) in der Altmühl-Alb. 

Vervielfältigtes Manuskript, Hemhofen-Zeckem. 
Mills, J.N. & Stage, CA. 1974: Chromosome numbers of British plants, 2. - Watsonia 10: 

167-168. 
Moore, D.M. 1982: Flora Europaea check-list and chromosome index. Cambridge. 
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Bayern. - Ber. Bayer. Bot. Ges. 25: 80-85. 
Schuhwerk, F. 1996: Published chromosome-counts in Hieracium. - http:// www. botanik. 

biologie.uni-muenchen.de/botsamml/projects/chrzlit.html 

- & Lippert, W. 1998: Chromosomenzahlen won Hieracium L. (Compositae, Lactuceae) Teil 
2. - Sendtnera 5: 269-286. 

Sell, P.D. & West, C. 1976: Hieracium. - In: Tutin, T.G. et al. (eds.): Flora Europaea vol 4: 

358^10, Cambridge. 
Strid, A. & Franzen, R. 1981: In: Chromosome number records LXXIIl (ed. Ä. Löve). - 

Taxon 30(4): 829 842. 
Zahn, K. H. 1900-1902: Hieracium. - In: Hallier, E. & Wohlfarth, R. (eds.), W.D.J. 

Kochs Synopsis der deutschen und schweizer Flora, ed. 3, 2: 1697-1931. Leipzig. 

- 1906: Die Hieracien der Schweiz. - Neue Denkschr. Schweiz. Natiu-f. Ges. 40(4): 165-728. 
Separat gedruckt 568 S., Basel. 

- 1929: Hieracium. - In: Hegi, G. (Hrsg.): Illustrierte Flora von Mitteleuropa Band Vl/2: 
1182-1351, München. 

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mitteleuropäischen Flora 12(2) 790 S., 12(3). Leipzig. 



Dr. Franz Schuhwerk und Dr. Wolfgang Lippert, Botanische Staatssammlung München, 

Menzinger Straße 67, D-80638 München. 

E-mail: schuhw(^botanik.biologie.uni-muenchen.de 



215 



Systematische Studien zur Morphologie und Anatomie von Hanguana 

Blume (Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit der 

Beschreibung einer neuen Art, Hanguana bogneri spec. nov. 



H.-J. Tillich und E. Sill 



Zusammenfassung: 

Tillich, H.-J. & Sill, E.: Morphologische und anatomische Studien an Hanguana 
Blume (Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit der Beschreibung 
einer neuen Art, Hanguana bogneri spec. nov. - Sendtnera 6: 215-238. 1999. ISSN 
0944-0178. 

Vertreter der lange Zeit in einer Familie zusammengefaßten Gattungen Hanguana 
und Flagellaria sind nur selten in Kultur und bezüglich vieler morphologischer und 
anatomischer Details noch sehr ungenügend bekannt. Für die vorliegende Arbeit 
standen vier verschiedene Herkünfte von Hanguana sowie eine Herkunft von Fla- 
gellaria indica L. zur Verfügung, die im Botanischen Garten München kultiviert 
werden. Von Hanguana werden Infloreszenz- und Blütenbau sowie die Anatomie des 
Blattes und der Wurzel beschrieben. Die bisher ebenfalls unbekannte Anatomie der 
Wurzel von Flagellaria wird erstmals untersucht. Ein blühendes männliches Exem- 
plar von Hanguana aus Sarawak wird als neue Art, Hanguana bogneri Tillich & 
Sill spec. nov. beschrieben. Die hier untersuchten Merkmale stützen die Stellung der 
Flagellariaceae in den Poales und der Hanguanaceae als basaler Seiten2:weig der 
Commelinales. 

Abstract: 

Members of the genera Hanguana and Flagellaria, being combined in one family 
for a long time, are rarely found in cultivation. The knowledge of a number of mor- 
phological and anatomical details is yet insufficient. For the present study four 
different origins of Hanguana and one specimen of Flagellaria indica L. have 
been available, all cuhivated in the Botanical Garden Munich. With respect to 
Hanguana new details of the morphology of inflorescence and flower and the ana- 
tomy of leaf and root are presented. The root anatomy of Flagellaria is described 
for the first time. A flowering male plant of Hanguana from Sarawak is described 
as a new species, Hanguana bogneri Tillich & Sill spec. nov. The investigated 
characters are supporting a close relationship of Flagellaria to Gramineae and 
Restionaceae, and a basal position of Hanguanaceae in the Commelinales. 



Einleitung 

Die Hanguanaceae Airy Shaw ( 1 965) sind eine kleine südostasiatisch-indomalaische Pflanzen- 
familie mit der einzigen Gattung Hanguana Blume. Bislang sind nur zwei Arten beschrieben: 
H. malayana (Jack) Merr. (syn. Susum anthelminticum Blume) und H. major Airy Shaw. 



216 



Hanguana war zunächst mit den Gattungen Flagellar ia L. und Joinvillea Gaudich. ex Brongn. 
& Gris in der Familie der Flagellariaceae Dumortier (1829) zusammengefaßt, die mittlerweile in 
die drei monogenerischen Familien der Flagellariaceae s.str., Hanguanaceae Airy Shaw und 
Joinvilleaceae Tomlinson & Smith aufgespalten worden ist (Airy Shaw 1965; Tomlinson & 
Smith 1970). 

Alle drei Gattungen sind nur selten gesammelt worden und noch seltener in Kultur anzutref- 
fen. Folglich ist die Kenntnis des Merkmalsbestandes noch sehr lückenhaft. Früchte, Samen 
und Keimpflanzen von Hanguana und Flagellaria sind vor kurzem genauer untersucht worden 
(Tillich 1996). Der Infloreszenz- und Blütenbau, insbesondere bei der diözischen Gattung 
Hanguana, ist dagegen nur oberflächlich bekannt. Dahlgren et al. (1985) beschreiben z.B. im 
Zentrum männlicher Hanguana-^WxiQn nur „callosities" von morphologisch ungeklärter Natur. 

Auch die Kenntnisse des anatomischen Baues von Hanguana und Flagellaria sind noch 
sehr unvollkommen. Erste Daten über H. malayana und Flagellaria indica lieferten Solere- 
DER und Meyer (1929). In einer ausführlicheren Untersuchung zur Anatomie der Flagellaria- 
ceae s.l. untersuchte Smithson (1956) vor allem einige Blattmerkmale und betonte die isolierte 
Stellung von Hanguana. 

Tomlinson (1969) bezieht sich auf die Untersuchungen von Smithson (1956) und stellt 
ebenfalls die isolierte Stellung der Gattung Hanguana gegenüber Flagellaria und Joinvillea 
heraus. Über den Bau der Wurzeln ist bisher überhaupt nichts bekannt. 

Im Botanischen Garten München sind H. malayana, H major und drei weitere Herkünfte 
der Gattung in Kultur, die sich deutlich von den beiden bisher beschriebenen Arten und z.T. 
auch voneinander unterscheiden. Während H malayana in offenen, nassen bis überfluteten 
Sumpfgebieten große und hochwüchsige Bestände bilden kann, leben andere, offenbar artver- 
schiedene Populationen als niedrige Einzelpflanzen oder in kleinen Trupps im Unterwuchs 
von schattigen Wäldern der malayischen Halbinsel sowie auf Sumatra und Borneo. Die weni- 
gen bisherigen Kenntnisse der Anatomie beziehen sich nur auf H malayana. Die Verschieden- 
heit im Habitus und im Lebensraum zwischen den Hanguana-Herkünften legt es nahe, die 
Anatomie auf breiterer Grundlage vergleichend zu untersuchen. Besonders interessant sind in 
dieser Hinsicht die Wurzeln, seit Seubert (1996 ff.) und Hofreiter (1997) zeigen konnten, 
daß die Wurzelanatomie bei den Monocotylen von großem systematischen Wert ist. Deshalb 
wurde auch die Wurzelanatomie von Flagellaria indica in diese Unt er suchun^n mit einbezogen. 

Die Analyse bisher nicht oder nur ungenügend bekannter Merkmale ist bei den hier unter- 
suchten Sippen besonders notwendig. Es ist noch immer unklar, welches ihre nächsten Ver- 
wandten bzw. Schwestergruppen sind. Der korrekte Platz im System der Monocotylen wurde 
z.B. für Hanguana im Laufe von drei Jahrzehnten bei den Asparagales (speziell in der Nähe 
von Lomandraceae oder Asteliaceae: Airy Shaw 1965, Dahlgren et al. 1985), irgendwo in 
den „Commelinoids" (Rudall & Caddick 1994, Rudall & Cutler 1995), bei den Zingi- 
berales (Stevenson & Loconte 1995) und den Commelinales (Tillich 1996) vermutet. 



Hanguana bogneri H.-J.Tillich & E.Sill, spec. nov. 

Holotypus: Malaysia, Prov. Sarawak, bei der Orang-Utan-Station Semenggoh, J. Bogner 
94/2211; kultiviert im Botanischen Garten München; Herbarbeleg der blühenden Pflanze 
vom 21. Oktober 1996 (M). 

Planta erecta, ca. 30 cm alta. Folia altema, lanceolata-acuminata, ca. 20 cm longa et 5-6 cm 
lata. Inflorescentia terminalis, pleiothyrsoidea vel pleiostachya. Flores dioici. Flos masculinus 
ca. 5 mm diametro, tepala 3+3, flava, exteriora minutissima, interiora ca. 2 mm longa, cucullata. 



217 

Stamina 3+3, filamentis basin versus dilatatis, antheris basifixis, introrsis, fissuris longitudi- 
nalibus. Pollen flava. Flos centro stylulis sterilibus brevibus, nectaribus tribus bifidis circum- 
datis provisus. Flores femini in spica compositi. Fructus baccatus, albus, uniseminatus. 

Blühende Pflanze (Typusexemplar) ca. 30 cm hoch; Blätter lanzettlich, ca. 20 cm lang, 5-6 
cm breit, die beiden Spreitenhälften etwas ungleich breit. Spreite lang zugespitzt, mit uni- 
facialer Spitze, basal in einen kurzen Blattstiel verschmälert und dann in eine kurze, stengel- 
umfassende Scheide übergehend, tief dunkelgrün mit mattem, samtartigen Glanz. Blüten diö- 
zisch, in terminalen Blütenständen. Die männliche Infloreszenz ist ein polyteler Pleiothyrsus. 
Blüten ohne Tragblätter, an der Basis jedes Thyrsus in dreiblütigen Wickeln, diese nach distal 
zu Einzelblüten verarmend. Die Infloreszenzäste (Paracladien) stehen jeweils in Gruppen in 
der Achsel eines Tragblattes. Männliche Blüten ca. 5mm breit, mit drei winzigen äußeren und 
drei deutlich größeren, hellgelben inneren Tepalen mit Kapuzenspitze. Stamina 3+3, Filamente 
an der Basis verbreitert, Antheren basifix, intrors, mit Längsschlitzen sich öffnend. Pollen gelb, 
mit Pollenkitt. Im Zentrum der Blüte drei kleine, sterile Styluli, umgeben von drei zweilappi- 
gen Nektarien. Weibliche Blüten in einer zusammengesetzten Ähre. Die Früchte sind weiße, 
einsamige Beeren. 



Material und Methoden 

Die Pflanzen für die vorliegenden Untersuchungen werden im Botanischen Garten München 
kultiviert und haben folgende Herkünfte: 

Hanguana major Airy Shaw: Malaysia, Provinz Sarawak, J. Bogner 82/237. Die Pflanze 
wurde zunächst unter dem Namen H. malayana (Jack) Merr. kultiviert und unter diesem 
Namen auch von Sill (1997) untersucht. Erst nach dem erstmaligen Blühen im Dezember 
1 997 konnte die Pflanze korrekt bestimmt werden. 

Hanguana bogneri spec, nov.: Malaysia, Prov. Sarawak, nahe der Orang-Utan-Station 
Semenggoh, J. Bogner 94/22 1 1 . Am Standort wurden nur Früchte gesammelt. Früchte, Samen 
und die daraus angezogenen Keimpflanzen sind von Tillich (1996) beschrieben worden. Bei 
SiLL (1997) wurde die einzige erfolgreich weiter kultivierte Pflanze mit dem Arbeitsnamen 
Hanguana „Sarawak" bezeichnet. 

Hanguana „Borneo": Malaysia, Provinz Sarawak, kultiviert im Botanischen Garten von 
Semenggoh, Bogner 94/1302. 

Hanguana „Singapur": Singapur, Bukit Timah Nature Reserve, R. Brett s. n. 24.08.1993. 
Am Standort wurden nur Samen gesammelt. In den Royal Botanic Gardens, Kew, wurden 
daraus Jungpflanzen angezogen, zwei Pflanzen erhielt der Botanische Garten München. 

Belegexemplare aller Hanguana-Sippen sind in der Botanischen Staatssammlung München 
(M) hinterlegt. 

Flagellaha indica L.: Herkunft unbekannt, seit Jahrzehnten im Botanischen Garten Mün- 
chen kultiviert. 

Flagellaria guineensis Schum.: Süd- Afrika, KwaZulu-Natal, St. Lucia Nature Reserve, M. 
Weigend & K. Drießle 98/69: FAA-fixierte Infloreszenzen. 

Die Blattanatomie von Hanguana wurde zunächst mit Handschnitten untersucht. Vom 
gesamten Material wurden außerdem Paraffin-Dürmschnitte von Blättern und Wurzeln herge- 
stellt, mit Standardreagenzien gefärbt und zu Dauerpräparaten verarbeitet. Diese sind in der 



218 



Präparatesammlung des Instituts für Systematische Botanik hinterlegt. 

Die morphologischen und anatomischen Zeichnungen wurden mit Hilfe einer Zeichenein- 
richtung für Mikroskope angefertigt. 

Die für eine zelluläre Zeichnung zu kleinen Leitbündel sind in den Tafeln halbschematisch 
dargestellt, wobei das Xylem kariert, das Phloem mit dicken Punkten und die Leitbündel- 
scheide mit radial verlaufenden Linien gezeichnet wurden. Bei sehr kleinen Leitbündeln wurden 
die mehr oder weniger reduzierten Leitgewebe vollständig weggelassen und nur die Leitbündel- 
scheide mit radialen Linien angedeutet. Der Punkt, an dem sich die Linien treffen, liegt nicht 
genau in der Mitte, sondern stets etwas xylemwärts, da sich die Leitbündelscheide vor dem 
Xylem erst später differenziert und daher weniger stark ausgeprägt ist. 



Infloreszenz- und Blütenbau 

Hanguana: Der männliche Blütenstand von H. bogneri ist ein polyteles System mit thyr- 
sischen Floreszenzen (Abb. IB, 2C, lOA). Bei dem dieser Beschreibung zugrunde liegenden 
Exemplar stehen unterhalb der Hauptfloreszenz die Paracladien in sechs Etagen. Die Para- 
cladien entspringen zu mehreren der Achsel eines Tragblattes. Ihre genaue Verkettung ist noch 
unklar. Die in einer Achsel stehende Gruppe scheint aber jeweils aus einem medianen Achsel- 
sproß und einer Serie kollateraler akzessorischer Sprosse zu bestehen. Die seitlich neben dem 
primären Achselsproß stehenden Paracladien besitzen jedenfalls keine Spur von Tragblättem, 
die auf eine proximale Verzweigung des Primansprosses schließen lassen körmten. Die Größe 
der Tragblätter an der Hauptachse nimmt von der Basis der Infloreszenz an aufwärts rasch ab. 
Während das unterste noch einem kleineren Laubblatt gleicht, haben die obersten nur noch die 
Gestalt von Schuppenblättem. Auch die Anzahl der in einer Blattachsel stehenden Paracladien 
sinkt in acropetaler Richtung von sieben auf zwei (Abb. IB). 

Die sitzenden Blüten haben weder Trag- noch Vorblätter. An der Basis der Floreszenzen 
stehen sie jeweils in dreiblütigen Wickeln. Weiter aufwärts verarmen die Cymen zu Zweier- 
gruppen, und der distale Teil jeder Floreszenz ist in den meisten Fällen auf die Gestalt einer 
Ähre verarmt. 

Das Perigon der männlichen Blüte besteht aus drei winzigen äußeren und drei deutlich 
größeren, hellgelben, an der Spitze kapuzenförmigen Tepalen (Abb. 2B). Die Stamina sind 
durch bandförmige, zur Basis hin verbreiterte Filamente sowie durch tetrasporangiate, introrse, 
mit Längsschlitzen öffnende Antheren gekennzeichnet. Die Filamente berühren sich an ihrer 
verbreiterten Basis gegenseitig, zuweilen sind sie sogar paarweise ein Stück weit aufwärts ver- 
wachsen. Der Pollen ist gelb und durch reichlichen Pollenkitt verklebt. Die kugeligen Pollen- 
kömer sind spinulos und inaperturat. Im Zentrum der Blüte findet sich ein rudimentäres 
Gynoeceum mit drei kleinen Styluli. Das Gynoeceum ist basal zu einer sechslappigen, diskus- 
artigen Scheibe verbreitert. Jedes Carpell trägt beiderseits seines Griffelrudiments je einen 
großzelligen Auswuchs. Die Auswüchse zweier benachbarter Carpelle neigen sich zueinander 
(Abb. lOB), sie haben wahrscheinlich die Funktion von Nektarien. 

Vom weiblichen Blütenstand dieser Sippe ist nur der von J. Bogner gesammelte fruchtende 
Zweig bekannt (vgl. Tillich 1996). Er läßt erkennen, daß die Blüten in ährigen Floreszenzen 
stehen. 

Ein vollständiger weiblicher Blütenstand konnte von H. major untersucht werden. Hier liegt 
ein polyteles System mit ährigen Floreszenzen vor. Der Verzweigungsgrad ist gering, nur die 
unteren Paracladien bringen je ein nahe ihrer Basis entspringendes Paracladium zweiter Ord- 



219 



nung hervor. Die Blüten sitzen einzeln an der relativ dicken Achse in der Achsel sehr kleiner 
Tragblätter (Abb. 2D, IOC). Jede Blüte besitzt ein transversales Vorblatt mit einer rudimen- 
tären Knospe, die ihrerseits ein transversales Vorblatt entwickelt (Abb. 2E). Die Ähren sind 
demnach als verarmte Thyrsen zu interpretieren. 

In der weiblichen Einzelblüte dominiert der voluminöse Fruchtknoten. Er trägt drei sitzen- 
de, unregelmäßig in kurze Äste verzweigte, gelblich-grüne Narben (Abb. lOD). Die kleinen Te- 
palen liegen dem Gynoeceum dicht an. Vor den drei inneren Tepalen stehen rudimentäre Stami- 
na (Abb. 2F). Äußere Tepalen, Tragblätter und Infloreszenzachsen sind locker mit Büschel- 
haaren besetzt, die inneren Tepalen und der dunkelgrüne Fruchtknoten sind kahl. Die 
auffallend dicke Fruchknotenwand besteht aus saftreichem Gewebe, in jedem der drei Fächer 
entwickelt sich in zentralwinkelständiger Position eine waagerecht stehende, atrope Samenan- 
lage. Diese überwölbt kuppeiförmig die massive Plazenta und deutet damit schon die spätere 
Gestalt des Samens an. Die Micropyle wird auf einem kleinen Buckel über die Oberfläche der 
Samenanlage emporgehoben (Abb. lOE). 

Flagellaria. Der Blütenstand ist eine reich verzweigte polytele Synfloreszenz mit ährigen 
Floreszenzen (Abb. 1 A). Der Verzweigungsgrad reicht bis zu Paracladien vierter Ordnung. Die 
ungestielten Blüten sitzen an jedem Paracladium dicht gedrängt in dessen distalem Abschnitt. 
Jede Blüte besitzt ein winziges Tragblatt, Vorblätter fehlen. Die einzelnen Floreszenzen be- 
stehen bei F. indica in der Regel aus fünf bis acht kopfig gedrängten Blüten, bei F. guineensis 
aus 9-15 Blüten, welche durch ihre größere Anzahl eher den Eindruck einer gestreckten Ähre 
vermitteln (Abb. 3C, D). Die Infloreszenzäste beliebiger Ordnung sind dorsiventral stark ab- 
geflacht, so daß sie lateral zwei scharfe Kanten besitzen (Abb. 3A). Die Paracladien haben 
keine Tragblätter, entspringen aber oberhalb eines kleinen Wulstes an ihrer abaxialen Basis, der 
eventuell als ein Tragblattrudiment zu interpretieren ist. Er erinnert an ähnliche Strukturen in 
Blütenständen der Gramineen. Das unterste Paracladium kann allerdings der Achsel des ober- 
sten Laubblattes entspringen, bleibt dann aber gegenüber den aufwärts folgenden mehr oder 
weniger stark gehemmt. An der adaxialen Basis jedes Paracladiums befindet sich eine leichte 
Anschwellung, für die Appel & Bayer (1998) eine Funktion beim Abspreizen der In- 
floreszenzäste vermuten. 

Die Blüten besitzen ein Perigon aus zwei dreizähligen Kreisen sehr zarter, durchscheinender 
Tepalen. Die Antheren werden an langen Filamenten weit aus der Blüte herausgeschoben. Die 
drei freien Griffel überragen die Tepalen nur wenig (Abb. 3B). Sie erreichen ihre volle Länge 
aber bereits vor der Streckung der Filamente, so daß sie kurzzeitig über die Antheren 
hinausragen (Abb. 3C). Der Pollen ist trocken und wird bei Erschütterung reichlich verstäubt. 



Blattbau in der Gattung Hanguana 

Morphologie: Die rosettig angeordneten, parallelnervigen Blätter sind linealisch-lanzettlich bis 
eiförmig-zugespitzt. Die kleine unifaciale Blattspitze vertrocknet rasch. Der Blattstiel ist teils 
deutlich, teils nur unscharf von der Blattspreite abgesetzt. Die Blattscheide umfaßt die Achse. 
H. major unterscheidet sich durch ihre Größe erheblich von den anderen Pflanzen. Ihre Blätter 
sind linealisch-lanzettlich, etwa 120 cm lang und 15 cm breit. Die Blätter der drei übrigen 
Pflanzen sind wesentlich kleiner. H. „Borneo" besitzt die schmälsten Blätter. Sie sind linea- 
lisch-lanzettlich, etwa 35 cm lang und 3,5 cm breit, mit nur undeutlich abgesetztem Blattstiel 
und als einzige durch anthocyanhaltige Hypodermiszellen, vor allem auf der Blattunterseite, 
rot überlaufen. Die Blattformen von H. bogneri und H. „Singapur" sind einander sehr ähnlich. 



220 



Sie sind beide etwa 20 cm lang und 5-7 cm breit, eiformig-zugespitzt und haben einen deutlich 
abgesetzten Blattstiel (Abb. 2A). Die Blätter von H. bogneri sind jedoch leicht an der tief dun- 
kelgrünen Färbung der Blattoberseite und dem samtartigen Glanz ihrer gesamten Oberfläche zu 
erkennen. 

Anatomie: Die Epidermis ist stets einschichtig und von einer deutlichen Cuticula über- 
zogen. Deren Oberfläche ist auf der Lamina bei drei der vier Pflanzen lediglich leicht gewellt 
(Abb. 4B), bildet bei H. bogneri jedoch auffallende Cuticularleisten, die im wesentlichen 
transversal verlaufen und daher besonders gut im Längsschnitt (Abb. 4C, lOF), aber auch in 
der Epidermisaufsicht (Abb. 4D) zu erkennen sind. Diese sind verantwortlich fur den matten 
Glanz und die sich weich anfühlende Oberfläche. Die Epidermiszellen aller vier Pflanzen sind 
dickwandig und longitudinal gestreckt, die antiklinen Wände sind reich getüpfelt (Abb. 5A, B). 

Stomata kommen, unregehnäßig verteilt, hauptsächlich auf der Blattunterseite vor (Abb. 
5A). Auf der Blattoberseite sind sie nur vereinzelt, besonders im Bereich der Mittelrippe, zu 
fmden Die Sp alt öffnungsap parate sind tetracytisch. Die zum Spalt weisende Wand der Schließ- 
zellen ist stark verdickt und in eine äußere und eine innere Randleiste ausgezogen (Abb. 4A). 

Bei allen untersuchten Pflanzen tragen beide Seiten des Blattes weiße, büschelig verzweigte 
Haare , die stellenweise verfilzen können. Abaxial bleiben sie meist länger erhalten und ver- 
zweigen sich weitaus stärker als adaxial, wo oftmals nur noch die Haarbasen zu finden sind 
(Abb. 5C, D). Die Äste eines Büschelhaares sind stets mehrzellig, wobei die distalen Haar- 
zellen dünnwandiger sind als die nahe der Haarbasis gelegenen. Die aus mehreren gedrungenen 
Zellen aufgebaute Haarbasis entspringt einer leicht vergrößerten Fußzelle in einer Einsenkung 
der Epidermis. Um die Vertiefung herum sind die Epidermiszellen kurz, unregelmäßig bis ring- 
förmig angeordnet und werden zur Fußzelle hin immer kleiner. 

Die adaxiale Hypodermis ist im Bereich der Lamina großzellig, einschichtig und enthält kein 
Chlorophyll (Abb. lOF). Über den longitudinal verlaufenden, großen Leitbündeln ist sie klein- 
zelliger und kann sogar unterbrochen sein (Abb. 6B, D). Nur bei H. „Borneo" konnte im Ge- 
gensatz zu den anderen Pflanzen nirgends eine Unterbrechung der adaxialen Hypodermis fest- 
gestellt werden (Abb. 4A). Bei H. major ist die Unterbrechung oftmals schwer zu erkennen, da 
nur eine einzige Zelle zwischen Epidermis und Leitbündel liegt. Diese gleicht in ihrer Form 
jedoch eher einer Palisadenzelle als einer Hypodermiszelle (Abb. 6C). Die abaxiale Hypo- 
dermis der Lamina besteht aus kleineren, chloroplastenfreien Zellen, ist stets einschichtig und 
bildet im Bereich der Blattscheide größere Zellen aus als an der adaxialen Seite. Die adaxialen 
Hypodermiszellen der Spreite sind in Aufsicht leicht transversal gestreckt und meist sechs- 
eckig, die abaxialen hingegen etwas longitudinal gestreckt und eher rechteckig (Abb. 5 A, B). 

Das Mesophyll läßt sich in der Spreite mehr oder weniger deutlich in Palisadenparenchym 
und Schwammparenchym gliedem (Abb. 6A, B). Diese Schichtung ist bei H. „Singapur" am 
schwächsten ausgeprägt, da die Palisadenzellen im Blattquerschnitt relativ kurz und breit sind 
(Abb. 6D). Auch bei H. major weist der Bau des Palisadenparenchyms Unregelmäßigkeiten 
auf, bedingt durch Zellen, die sich in ihrer Größe stark unterscheiden und oftmals nicht anti- 
klinal gestreckt sind (Abb. 6C). Das Palisadenparenchym aller vier Pflanzen ist im Blattquer- 
schnitt stets niedriger als das Schwammparenchym, macht aber mindestens etwa ein Drittel 
des Mesophylls aus. Es ist meist zweischichtig und enthält den überwiegenden Teil der 
Chloroplasten. Seine Zellen sind im oberflächenparallelen Schnitt mehr oder weniger rund und 
durch relativ große Interzellularräume voneinander getrennt. Das Schwammparenchym besteht 
aus großen Zellen, die unregelmäßig gestaltet oder nahezu sternförmig sein können und dann 
untereinander über armartige Fortsätze verbunden sind. Die Interzellularen sind groß und an- 
nähernd rund. 



22T 



Der Blattrand der Spreite und des Stiels krümmt sich leicht nach abaxial. Das Mesophyll der 
Lamina reicht nur bei H. major bis zum Blattrand, ist dort aber nicht mehr in Palisaden- und 
Schwammparenchym differenziert und enthält kaum noch Interzellularen (Abb. 6F). Bei den 
übrigen Herkünften erreicht das Mesophyll nicht den Randbereich, so daß die aufeinander lie- 
genden Epidermen einen schmalen Hautsaum bilden (Abb. 6E). Der Rand der Blattscheide ist 
bei allen untersuchten Pflanzen ein farbloses, dünnes Häutchen, das sich nach adaxial krümmt, 
nur von den Epidermen gebildet wird und rasch vertrocknet (Abb. IC). 

Die Leitbündel sind in der Mittelrippe in drei übereinander liegenden Systemen angeordnet, 
wobei die abaxiale und die mittlere Serie aus normal orientierten Bündeln besteht. Die Bündel 
der adaxialen Serie sind teils schräg, teils invers orientiert und enden im basalen Drittel der 
Spreite (Abb. 7A, B). Ihr Durchmesser und der Grad ihrer anatomischen Differenzierung 
nehmen von der Blattscheide über den Blattstiel bis zu ihrem distalen Ende in der Spreite kon- 
tinuierlich ab. Außerhalb der Mittelrippe fehlen die inversen Bündel. Dafür treten die Bündel 
der mittleren Serie, die in der Mittelrippe in mehreren Ebenen angeordnet sein können, im Be- 
reich der Lamina an die adaxiale Hypodermis heran. Die Bündel der abaxialen Serie der Mitte 1- 
rippe verlaufen in der Lamina innerhalb des Schwammparenchyms. In der Nähe des Blatt- 
randes fehlt schließlich auch die abaxiale Serie (Abb. 6E, F). 

Das Xylem enthält selbst in den stärksten Bündeln keine Tracheen, sondern nur englumige 
Tracheiden. Die Bündel der mittleren Serie bzw. die adaxialen Bündel der Lamina besitzen eine 
doppelte Scheide. Der irmere, sklerenchymatische Anteil ist je nach Position des Bündels 
mehrere Zellagen dick oder nur schwach ausgebildet und oft über dem Xylem stärker als über 
dem Phloem entwickelt. Die äußere Leitbündelscheide ähnelt einer Endodermis und besteht aus 
u-förmig verstärkten Zellen (Abb. 4E). 

Die Bündel der abaxialen Serie sind deutlich kleiner als die der mittleren und werden vor 
allem in der Spreite stark reduziert. Sie besitzen dort nur noch ein wenigzelliges Phloem, das 
Xylem fehh meist vollständig. Eine Gliederung in innere und äußere Bündelscheide ist hier 
lediglich vor dem Phloem zu erkennen. Auf der adaxialen Seite wird die Scheide nur aus einer 
Lage großlumiger, unverstärkter und longitudinal gestreckter Zellen gebildet (Abb. 4F, 1 OF). 

Im Mesophyll der Lamina verlaufen zusätzlich kleine Leitbündel in transversaler Richtung, 
die in Xylem und Phloem mit normaler Orientierung gegliedert sind. 

Bei Frischmaterial sind die Leitbündelscheiden im Bereich der Blattscheide und des Blatt- 
stiels zum Teil auffallend farbig. Die Zellwände der endodermisartigen, äußeren Scheide kön- 
nen dunkelbraun, die der inneren Sklerenchymscheide orange gefärbt sein. Die Intensität der 
Färbung hängt von der Lage des Leitbündels ab. Je weiter sich ein Bündel proximal und rand- 
wärts befindet, desto kräftiger ist seine Farbe. Richtung distal imd zentral des Blattes werden 
die Leitbündel zunehmend blasser. 

Über der Mittelrippe liegen, wie schon erwähnt, die wenigen adaxialen Stomata. Die ad- 
axiale Hypodermis ist hier meist zweischichtig, über Leitbündeln manchmal nur einschichtig. 
Die abaxiale Hypodermis ist stets einschichtig. Es gibt keine Differenzierung in Palisaden- und 
Schwammparenchym (Abb. 7A). Als Assimilationsparenchym sind nur noch die unmittelbar 
an die Hypodermen grenzenden Schichten ausgebildet. Das übrige interzellularenreiche Meso- 
phyll ähnelt dem Schwammparenchym und enthält nur sehr wenige Chloroplasten. Die Zellen 
sind isodiametrisch, armartige Fortsätze sind nur stellenweise angedeutet. 

Als besonders auffallendes Merkmal sind die lysigenen Kanäle zu erwähnen, die bei allen 
untersuchten Pflanzen in Längsrichtung durch die Mittelrippe der Blattscheide ziehen und bei 
H. bogneri und H. „Borneo" sogar bis weit in den Blattstiel hinaufreichen (Abb. 7D, E). Sie 
enthalten einen bei Feuchtigkeit stark quellenden, farblosen und klebrigen Schleim, der sich mit 



222 



Astrablau im mikroskopischen Präparat mehr oder weniger stark blau anfärbt. 

Nur bei H. major wurden SiHkatkristalle gefunden. Diese kommen als Kristallgries vor allem 
in parenchymatischen, xylemwärts liegenden Zellen der Bündelscheiden von kleinen, reduzier- 
ten Leitbündeln und manchmal auch in Mesophyllzellen vor. Einzelne große Silikat körp er finden 
sich häufig in der adaxialen Hypodermis (Abb. 4G), seltener auch in der abaxialen Hypoder- 
mis, im Mesophyll oder in einer äußeren Leitbündelscheide. Es sind meist tailliert-quader- 
förmige Einzelkristalle, seltener auch sternförmig verwachsene Säulen. 

Außer in der Epidermis und den endodermisartigen Bündelscheiden kommen in allen Gewe- 
ben mit Granula gefüllte Sekretzellen vor, deren Inhalt in Handschnitten von Frischmaterial 
graubraun gefärbt ist und bei denen es sich nach Tomlinson (1969) um Tanninzellen handelt. 
Der Zellinhalt färbt sich bei Paraffin-Dünnschnitten nach der Standardfarbung mit Astrablau 
und Safranin rot bis violett, bei H. bogneri auch blau an. In Handschnitten von Frischmaterial 
von H. bogneri und H. „Singapur" wurden außerdem Sekretzellen ohne Granula beobachtet, 
die farblich von blaßgelb über orange bis zu rotbraun variieren. Hierbei handelt es sich 
möglicherweise um junge, noch unreife Tanninzellen. 



Wurzelanatomie der Gattung Hanguana 

Die Wurzeln aller untersuchten Pflanzen dieser Gattung stimmen in wesentlichen Merkmalen 
überein. Die Rhizodermis ist stets einschichtig ausgebildet und bleibt auch an der älteren 
Wurzel noch lange erhalten. Ihre Zellen sind dünnwandig und nur zum Teil zu Wurzelhaaren 
ausgewachsen. Im Dünnschnitt fällt die flaschenförmige Gestalt der Basis der angeschnittenen 
Wurzelhaare auf (Abb. 8A). An die Rhizodermis schließt sich eine zwei- bis funfschichtige 
Exodermis an. Ihre Zellen sind unterschiedlich groß und können verholzen (Abb. 8A). 

Das interzellularenreiche Rindenparenchym beginnt sich frühzeitig aufzulösen (Abb. 8A). 
Bei älteren Wurzeln sind von ihm nur noch eine einzige Schicht Parenchymzellen unterhalb der 
Exodermis und eine zwei Zellen breite Lage über der Endodermis erhalten. Dazwischen bleiben 
nur einige radiale Speichen übrig, die im Wurzelquerschnitt zwei bis drei Zellen breit sind. 

Die Endodermis besteht aus u-förmig verstärkten Zellen (Abb. SB, C). Sie ist von einzelnen 
parenchymatischen Durchlaßzellen unterbrochen, die nur bei H. „Borneo" nicht beobachtet 
werden konnten. Die Wände der Endodermiszellen älterer Wurzeln sind stark verdickt und 
umschließen nur noch kleine Restlumina. 

Der Perizykel besteht aus ein bis drei Zellagen und kann verholzen (Abb. SB, C). 

Das polyarche Leitbündel enthält je nach Dicke der Wurzel zwölf bis 36 Protoxylem- und 
ebensoviele Protophloempole (Abb. SD). Die Xylemprimanen sind meist gut erkennbar. Es er- 
folgt ein stetiger Übergang von Protoxylemtracheiden zu Metaxylemtracheen, wobei die ein- 
zelnen weitlumigen Metaxylemtracheen jeweils mehreren Protoxylempolen zuzuordnen sind. 
Das Protophloem ist nicht vom Metaphloem abgrenzbar. Das Phloem bildet entweder rund- 
liche Nester zwischen den Protoxylempolen (Abb. SC), oder es zieht sich als schmales radiales 
Band zwischen den Metaxylemtracheen bis nahe an das Markgewebe heran (Abb. SB). In 
diesem Falle kann es auch durch Gruppen sklerenchymatischer Zellen in einzelne Inseln auf- 
geteilt sein (Abb. SD). Das gesamte Leitsystem ist in einen breiten, mehr oder weniger stark 
verholzten Ring eingebettet. 

Das Markgewebe ist parenchymatisch bis sklerenchymatisch ausgebildet und enthält bei 
den dicken Wurzeln von H. major zahlreiche Interzellularräume (Abb. SD). 

Die Wurzelgewebe sind frei von Silikatkristallen . 



223 

Sekretzellen sind im Rindenparenchym, Perizykel, Leit- und Markgewebe zu finden, wobei sie 
im Zentralzylinder weitaus häufiger vorkommen als in der Rinde. In den dicken Wurzeln von 
H. major können die das Phloem begleitenden Tanninzellen miteinander verschmelzen und 
radial verlaufende Sekretlakunen bilden. 



Wurzelanatomie von Flagellaria indica 

Die einschichtige Rhizodermis bleibt im Gegensatz zu Hanguana an älteren Wurzeln nur in 
Resten erhalten. Ihre Zellen sind etwas dickwandiger, wesentlich schmaler und überwiegend zu 
Wurzelhaaren ausgewachsen (Abb. 9A). 

Die Exodermis besteht aus etwa drei Schichten kleiner und unverstärkter Zellen (Abb. 9A). 

Die Zellen des angrenzenden drei- bis vierschichtige Sklerenchymrings besitzen dagegen 
stark verdickte Zellwände. Das interzellularenreiche Rindenparenchym ist aus unterschiedlich 
großen Zellen aufgebaut. In seinen peripheren Interzellularen befindet sich ein Sekret, das sich 
mit Safranin im mikroskopischen Präparat rot anfärbt (Abb. 9A). In der Mitte des Rindenpar- 
enchyms bilden sich rhexigen radiale Lakunen (Abb. 9B, lOG). In der die Endodermis umge- 
benden Schicht können einzelne schwach bis stark verholzte Zellen vorkommen (Abb. 9C). 

Die Endodermis besteht, im Gegensatz zu der von Hanguana, aus radial gestreckten, extrem 
o-förmig verdickten Zellen mit winzigem Restlumen (Abb. 9C). Durchlaßzellen konnten nicht 
beobachtet werden. 

Der Perizykel ist zwei- bis vierschichtig und stark skierotisiert (Abb. 9C). 

Das polyarche Leitbündel besitzt je 60-80 Protoxylem- und Protophloempole (Abb. 9G). 
Die Protoxylempole sind als relativ dünnwandige Xylemprimanen und kleine Tracheiden gut 
erkennbar. Die großen, runden Tracheen des späten Metaxylems sind in zwei bis drei Ringen 
unregelmäßig konzentrisch um das Markgewebe angeordnet. In den zwischen den Xylempolen 
liegenden Phloemnestem stellt der kleine periphere, besonders kleinzellige Bereich das Proto- 
phloem dar, während der Hauptanteil vom klein- bis sehr großzelligen Metaphloem eingenom- 
men wird (Abb. 9C). Tracheen, Tracheiden imd Phloem sind in ein breites Band verholzter 
Xylemparenchymzellen eingebettet (Abb. 9D). 

Das interzellularenreiche Markgewebe ist parenchymatisch und kann inselartig zwischen 
die am weitesten zentral gelegenen Tracheen hinein reichen. Es hat demzufolge eine sehr un- 
regelmäßige Querschnittsform (Abb. 9D, lOG). 

In den Wurzeln von F. indica wurden keinerlei Kri stall einschlüsse gefunden. 

Sekretzellen kommen in den Wurzeln von F. indica ebenfalls nicht vor. Worum es sich bei 
der Substanz handelt, die sich in den äußeren Interzellularräumen des Rindenparenchyms 
befindet, ist nicht bekannt (Abb. 9A). 



Diskussion 

Die Gattungen Flagellaria und Hanguana weichen in allen bisher detailliert untersuchten 
Merkmalsbereichen so sehr voneinander ab, daß ihre einstige Zusammenfassung in einer 
gemeinsamen Familie heute nicht mehr ernsthaft in Erwägung gezogen werden kann. 

Infloreszenz- und Blütenbau: Der reich verzweigte Blütenstand von Flagellaria mit 
seinen ährigen Floreszenzen deutet einige Beziehungen zu den Gramineen und den Restiona- 
ceen an. An die Gramineen erinnert das Fehlen von Tragblättem an den Infloreszenzästen, 



224 



während jede Einzelblüte über ein Tragblatt verfugt. Dazu kommen die gedrängte Anordnung 
der Blüten in relativ wenigblütigen Ähren am Ende von Infloreszenzästen und die Wind- 
blütigkeit. Im Gegensatz zu den Poaceen entsprechen die Blüten mit ihrem trimer-penta- 
zyklischen Bau aber noch dem vollständigen Monocotylen-Diagramm. Unter den Restionaceen 
sind erst wenige detaillierte Infloreszenzanalysen bekannt. Große Ähnlichkeit zu Flagellaria 
weist aber der männliche Blütenstand von Leptocarpus similis Edgar auf (vgl. Kircher 1986). 

Die Hanguana-\nüoxQszQnztr\ weichen im Verzweigungsmuster und auch habituell stark 
von den Blütenständen der Flagellaria und aller übrigen Familien der Poales ab. Thyrsische 
Floreszenzen mit einer Vielzahl von Sonderformen sind dagegen bei den Commelinaceae und 
den Familien der Zingiberales weit verbreitet. Die Abbildung eines als Susum anthelminticum 
bezeichneten weiblichen Exemplars aus Sumatra bei Maury (1889) macht deutlich, daß auch 
weibliche Infloreszenzen von Hanguana einen ähnlich hohen Verzweigungsgrad wie die hier 
beschriebene männliche Infloreszenz von H. bogneri erreichen können. Die Abbildung ist von 
Takhtajan (1982) sowie Bayer et al. (1998) mit der Bezeichnung H. malayana übernommen 
worden. Die Artzugehörigkeit ist aber fraglich, da Habitus und Blattgestalt nicht mit H. malay- 
ana übereinstimmen. 

Die Blüten von Hanguana unterscheiden sich nicht nur durch ihre Zweihäusigkeit, sondern 
auch in Bezug auf alle Detailmerkmale von denen der Gattung Flagellaria. Unklar bleibt aber 
noch, wie die Hanguana-^XnXtn bestäubt werden. Sie sind optisch wenig auffallend, die Pro- 
duktion von Nektar in den männlichen Blüten und der Besitz von Pollenkitt machen jedoch 
eine zoogame Bestäubung wahrscheinlich. Mit welchen Mitteln die weiblichen Blüten den Be- 
stäuber anlocken, ist aber bisher nicht erkennbar. Maury (1889) beschreibt die Farbe des 
Fruchtknotens des von ihm untersuchten Exemplars zwar als „rose-verdätre", bei H. major 
bleibt er aber rein grün. Früchte werden reichlich gebildet (vgl. Tillich 1996). 

Der Fruchtansatz bei Flagellaria ist dagegen meist nur sehr gering. Die oft großen Bestände 
entstehen vor allem durch reichliche vegetative Vermehrung (Matolweni & McLellan 
1997). 

Vegetative Anatomie von Hanguana. Über die bisher vorliegenden spärlichen Daten von 
Smithson (1956) hinaus können jetzt die gattungstypischen Merkmale des Blattbaues deutlich 
herausgestellt werden. Besonders kennzeichnend sind longitudinal gestreckte Epidermiszellen 
mit stark getüpfelten, antiklinen Zellwänden, vereinzelt auch auf der Blattoberseite anzutref- 
fende, tetracytische Stomata und mehrzellige, verzweigte und in die Epidermis eingesenkte 
Haare, die von Solereder und Meyer (1929) irrtümlicherweise als Drüsenhaare bezeichnet 
wurden. Desweiteren können die unter beiden Epidermen liegenden, chlorophyllfreien Hypo- 
dermen genannt werden, die auf der adaxialen Seite transversal, auf der abaxialen Seite longitu- 
dinal gestreckt sind. Eine Differenzierung in Palisaden- und Schwammparenchym in der 
Lamina ist dagegen nicht bei allen Arten deutlich ausgeprägt, auch für H. malayana gibt Smith- 
son (1956) ein relativ einheitliches, papillöses Mesophyll an. In der Mittelrippe der Blatt- 
spreite, im Blattstiel und in der Blattscheide findet sich überall ein gleichförmiges Mesophyll. 
Unter der adaxialen Hypodermis liegt eine Serie inverser Leitbündel. In fast allen Geweben 
finden sich femer graubraune, granulagefüllte Sekretzellen. Auch die bisher nur von Arber 
(1925) in einer Zeichnung angedeuteten, von späteren Autoren nicht wieder erwähnten 
Schleimkanäle in der Blattscheide gehören zum Merkmalsbestand der Gattung. 

Die Wurzeln von Hanguana weisen ebenfalls eine Reihe sehr charakteristischer, durch- 
gehend vorhandener Merkmale auf. Dazu gehört die relativ langlebige Rhizodermis und ein sich 
frühzeitig auflösendes Rindenparenchym, so daß schließlich nur wenige, konzentrische Zell- 
schichten im äußeren und inneren Rindenparenchym erhalten bleiben. Femer sind die u-formig 



225 



verstärkten Endodermiszellen, ein polyarches Leitsystem mit im Querschnitt ovalen Tracheen 
und inselartig bis radial -bandförmig gestaltetem Phloem sowie die im Zentralzylinder gehäuft 
vorkommenden Sekretzellen zu nennen. Im Gegensatz zu einer irrtümlichen Angabe bei Dahl- 
GREN et al. (1985) findet sich bei Tomlinson (1969) kein Hinweis auf das Vorkommen von 
leiterförmig durchbrochenen Tracheen-Endplatten. 

Differentialmerkmale der Hanguana-Herkünfte: Alle untersuchten Hanguana-Her- 
künfte stimmen in zahlreichen Merkmalen des Blattbaues überein. Daneben weisen sie aber 
auch eine Reihe von anatomischen und morphologischen Differentialmerkmalen auf, mit deren 
Hilfe sie klar unterscheidbar sind. Die wichtigsten Trennmerkmale sind in Tabelle 1 zusam- 
mengestellt. 

Tabelle 1 : Blattanatomische Differentialmerkmale der untersuchten Hanguana-Herkünfte 





H. major 


H. bogneri 


H. „Singapur" 


H. „Borneo" 


Form der Lamina: 

-linealisch-lanzettlich 

-eiförmig-zugespitzt 


+ ' 


+ 


+ 


+ 


Blattlänge: 
> 100 cm 
<50cm 


+ 


+ 


+ 


+ 


Cuticularleisten 


- 


+ 


- 


- 


Deutliches 
Palisadenparenchym 


- 


+ 


- 


+ 


Laminarand mit 
Mesophyll 


+ 


- 


- 


- 


Deutlicher Blattstiel 


+ 


+ 


+ 


- 


Blattstiel mit 
Sekretkanälen 


- 


+ 


- 


+ 


Silikatkristalle 


-1- 


- 


- 


- 


Hypodermis mit 
Anthocyan 


- 


- 


- 


+ 



Vergleich von Hanguana und Flagellaria: Bei der Gegenüberstellung der Wurzelana- 
tomie der Gattung Hanguana einerseits imd Flagellaria indica andererseits fallen zahlreiche 
Unterschiede auf 

Die Rhizodermis von Flagellaria geht im Gegensatz zu der von Hanguana bald zugrunde. 
Zwischen der Exodermis und dem parenchymatischen Rindengewebe wird bei Flagellaria ein 
mehrschichtiger Sklerenchymring angelegt, der bei Hanguana gänzlich fehlt. Schon Lindinger 
(1906) hatte für einige Familien, darunter die Flagellariaceae, einen Mantel aus Sklerenchym- 
fasem in der äußeren Rinde erwähnt, allerdings ohne eine Angabe darüber, auf welche Gattung 
sich diese Beobachtung bezieht. Im Rindenparenchym von Flagellaria finden sich darüber 
hinaus radial verlaufende, rhexigene Lakunen, wohingegen sich das Rindenparenchym von 
Hanguana weitgehend gleichmäßig oder bis auf radiale Speichen auflösen kann. Weitere 
wesentliche Trermmerkmale zwischen den beiden Gattungen sind in Tabelle 2 zusammen- 
gestellt. 



226 



Tabelle 2: Wesentliche Unterschiede im Wurzelbau von Hanguana und Flagellar ia 





Hanguana 


Flagellaria 


Rhizodermis 


relativ lange und gut erhalten 


nur in Resten erhalten 


Sklerenchymring im äußeren 
Rindengewebe 


fehlt , 


vorhanden 


Endodermiszellen 


u-fbrmig verstärkt 


o-förmig verstärkt 


Anzahl der Xylem- bzw. 
Phloempole 


<40 


>60 


Form der Tracheen im 
Querschnitt 


oval 


rund 


Markgewebe 


Querschnitt kreisförmig 


Querschnitt unregelmäßig, 
einzelne Markparenchym- 
nester zwischen den Tracheen 


Vorkommen von Sekret 


intrazellulär, vor allem im 
Zentralzylinder 


interzellulär, im äußeren 
Rindengewebe 



Die systematische Stellung der Flagellariaceae und der Hanguanaceae: Die Stellung 
der Flagellariaceae s.str. und Joinvilleaceae in den Poales ist heute weitgehend unumstritten 
(Dahlgren et al. 1985, Linder & Kellogg 1995). Weniger klar ist jedoch die Stellung der 
Hanguanaceae, die in den vergangenen Jahrzehnten bei den Aspar^^les (Dahlgren et al. 1985), 
Zingjberales (Stevenson & Loconte 1995) und den Commelinoiden (Rudall & Caddick 
1994) bzw. Commelinales (Tillich 1996) angesiedelt waren. 

Aus den Asparagales können die Hanguanaceae inzwischen sicher ausgeschlossen werden. 
Zumeinen ist mittlerweile Stärke im Endosperm nachgewiesen (Tillich 1996), zum anderen 
weist das Vorkommen von fluoreszierenden phenolischen Substanzen in den Zellwänden deut- 
lich zu den „commelinoiden" Familien (Rudall & Caddick 1994). Hanguana zeichnet sich 
durch eine charakteristische Kombination von ursprünglichen und abgeleiteten Merkmalen aus. 
Unter den abgeleiteten Merkmalen sind die Synfloreszenzstruktur, die Diözie und der sehr 
eigenartige Samenbau zu nermen. Das Fehlen von Tracheen in den Sproßorganen ist dagegen als 
sehr ursprünglich zu werten. Innerhalb der Commelinales würde Hanguana auf Grund dieses 
Merkmals eine eher basale Stellung einnehmen, da die übrigen Familien (Commelinaceae, 
Mayacaceae, Rapateaceae und Xyridaceae) sämtlich Tracheen in ihren Sproßorganen besitzen. 
Die Angabe von Dahlgren et al. (1985), wonach im Sproß von Cartonema keine Tracheen 
vorkommen, bedarf der Überprüfung. Tomlinson (1969) beschreibt jedenfalls im Xylem 
„many files of wide tracheal elements". 

Stevenson & Loconte (1995) werten eine Reihe von Merkmalen als Synapomorphien mit 
den Zingiberales und stellen die Hanguanaceae als Schwestergruppe neben die Zingiberales. Die 
hierfür herangezogenen Merkmale sind aber wenig überzeugend. Kieselkörper und „bambu- 
soide" Blattstiele kommen auch in den Commelinales vor, und die palmate Venation der Han- 
gwflWüt-Blätter ist nur vage definiert. Die Zingiberales besitzen andererseits eine Reihe von 
durchgehenden Merkmalen, durch welche sie von Hanguana entfernt werden, so z.B. epigyne 
Blüten und große Lakunen in den Blättern. Auch die verzweigten Haare von Hanguana unter- 
scheiden sich von allen Haartypen innerhalb der Zingiberales. Die vielfältigen Haartypen der 
Commelinaceae regen viel eher zu einem Vergleich mit Hanguana an. So besitzt z.B. die Gat- 
tung Palisota ebenfalls verzweigte Haare. Unter Berücksichtigung aller bisher bekannten Merk- 
male fügen sich die Hanguanaceae am besten als basaler Seitenzweig bei den Commelinales ein. 



227 

Für die Bereitstellung des Materials danken wir Dr. Paula J. Rudall, Royal Botanical Gardens, Kew, 
sowie Josef Bogner, Botanischer Garten München. Für die großzügige Hilfe bei der digitalen 
Bildbearbeitung der Farbtafel danken wir Prof. Dr. G. Wanner, Botanisches Institut, Universität 
München. 



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- 1981: A new species of //üi«gMür«a from Borneo. -Kew Bull. 35: 819-821. 

Appel, O. & Bayer, C. 1998: Flagellariaceae. - In: Kubitzki, K. (ed.): The Families and 

Genera of Vascular Plants. Vol. IV: Flowering Plants. Monocotyledons. Alismatanae and 

Commelinanae (except Gramineae), 208-21 1. Berlin. 
Arber, a. 1925: Monocotyledons: a morphological study. - Cambridge. Reprint 1961. Wein- 

heim. 
Bayer, C, Appel, O. & Rudall, P.J. 1998: Hanguanaceae. - In: Kubitzki, K. (ed.): The 

Families and Genera of Vascular Plants. Vol. IV: Flowering Plants. Monocotyledons. 

Alismatanae and Commelinanae (except Gramineae), 223-225. Berlin. 
Dahlgren, R.M.T., Clifford, H.T. & Yeo, P.F. 1985: The Families of the Monocoty- 
ledons. - Berlin. 
Hofreiter, a. 1997: Systematische Wurzelanatomie der Commelinaceae R. Brown (1810) 

(Monocotyledoneae). - Diplomarbeit, Institut für Systematische Botanik, LMU-München. 
Kircher, P. 1986: Untersuchungen zur Blüten- und Infloreszenzmorphologie, Embryologie 

und Systematik der Restionaceen im Vergleich mit Gramineen und verwandten Familien. 

Dissertationes Botanicae 94. - Berlin. 
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. 
Lindinger, L. 1906: Zur Anatomie und Biologie der Monokotylenwurzel. - Beih. Bot. Cen- 

tralbl. Abt. I, 19, Heft 2: 321-358. 
Matolweni, L.O. & McLellan, T. 1997: Genetic diversity of an exploited species, Flagel- 

laria guineensis . - S. African J. Bot. 63: 294-298. 
Maury, J. 1889: Susum anthelminticum. - Revue Horticole 61: 16-1%. 
Rudall, P.J. &. Caddick, L.R. 1994: Investigation of the presence of phenolic compoimds in 

monocotyledonous cell walls, using UV fluorescence microscopy. - Ann. Bot. (London) 

74:483^91. 
Rudall, P.J. & Cutler, D.F. 1995: Asparagales: a Reprisal. - In: Rudall, P.J., Cribb, P. J., 

Cutler, D.F. & Humphries, C.J. (eds.): Monocotyledons: systematics and evolution, 

157-168. Kew. 
Seubert, E. 1996a: Root anatomy of palms. II. Calamoideae. - Feddes Repert. 107: 43-59. 

- 1996b: Root anatomy of palms. III. Ceroxyloideae, Nypoideae, Phytelephantoideae. - 
Feddes Repert. 107: 597-619. 

- 1997: Root anatomy of palms. I. Coryphoideae. - Flora 191: 81-103. 

- 1998a: Root anatomy of palms. IV. Arecoideae. part 1. general remarks and descriptions on 
the roots. - Feddes Repert. 109: 89-127. 

- 1998b: Root anatomy of palms. V. Arecoideae. part 2. systematic implications. - Feddes 
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228 



Sill, E. 1997: Systematische Anatomie der Vegetationsorgane der Hanguanaceae Airy Shaw 
(1965) (Monocotyledoneae). - Diplomarbeit, Institut für Systematische Botanik, LMU- 
München. 

Smithson, E. 1956: The comparative anatorry of the Flagellariaceae. - Kew Bull. 1 1 : 491-501 . 

SoLEREDER, H. & Meyer, F.J. 1929: Systematische Anatomie der Monokotyledonen, Heft 4 
Farinosae: 3-6. Berlin. 

Stevenson, D.W. & Loconte, H. 1995: Ciadistic analysis of monocot families. - In: Ru- 
DALL, Cribb, P.J., Cutler, D.F. & Humphries, C.J. (eds.): Monocotyledons: systematics 
and evolution, 543-578. Kew. 

Takhtajan, A. (ed.) 1982: Plant Life. Vol. 6. Angiosperms. Monocotyledons. Moskau, 
(russisch). 

Tillich, H.-J. 1996: Seeds and seedlings in Hanguanaceae and Flagellariaceae (Mono- 
cotyledons). - Sendtnera 3: 187-197. 

ToMLiNSON, P.B. 1969: Commelinales - Zingiberales. - In: Metcalfe, CR. (ed.): Anatomy 
of the Monocotyledons, Vol. III. Oxford. 

- & Smith, A.C. 1970: Joinvilleaceae: a new family of monocotyledons.- Taxon 19: 887-889. 

Prof. Dr. Hans-Jürgen Tillich, Elisabeth Sill, Institut fur Systematische Botanik der 
Ludwig-Maximilians-Universität München, Menzinger Strasse 67, D-80638 München. 





Abb. 1 : Synfloreszenzbau, schematisch. A: Flagellaria indica, proximales Paracladium; B: Han- 
guana bogneri, männliche Synfloreszenz. Akzessorische Paracladien als Pfeile dargestellt. 



229 




Abb. 2: Hanguana. A-C: H. bogneri (männlich). A: Laubblatt. B: Blüte. C: Floreszenz. D-F: 
H. major (weiblich). D: Distaler Abschnitt einer Floreszenz. E: Blüte. F: inneres Tepalum, 
adaxial, mit Staubblattrudiment, tb: Tragblatt, vb: Vorblatt, vbl: Vorblatt der Achsel- 
knospe. Maßstabbalken; A: 5 cm; B: 3 mm; C: 1 cm; D: 3 cm; E, F: 5 mm. 



230 




Abb. 3: Flagellaria. A-C: F. indica. A: Proximaler Infloreszenz-Ast. B; Einzelblüte. C: Flores- 
zenz, bei der bis auf drei alle Blüten abpräpariert wurden, um die acropetale Aufblühfolge 
und die Protogynie zu veranschaulichen. D: F. guineensis, Floreszenz. g: Griffel, tb: Trag- 
blatt. Maßstabbalken: A: 1 cm; B-C: 2 mm; D: 5 mm. 



231 









C3 




r7TTiwr<5 



Abb. 4: Hanguana, Blattanatomie. A: H. „Borneo", abaxiale Epidermis, quer, mit Spalt- 
öffnungsapparat. B-C: adaxiale Epidermis, antikline Wände mit Tüpfeln, Längsschnitte. B: 
H. „Singapur" ohne Cuticularleisten. C: H. bogneri mit Cuticularleisten. D-F: H. bogneri. 
D: Epidermis mit Cuticularleisten in Aufsicht. E-F: Leitbündel der Lamina, quer. E: 
adaxiales Bündel. F: reduziertes abaxiales Bündel. G: H. major, adaxiale Hypodermis, quer, 
mit Silikatkristallen. Cu: Cuticularleiste. PNz: Nebenzelle. S: Sekretzelle. Sz: Schließzelle. 
Tu: Tüpfel. Interzellularen punktiert. Maßstabbalken: A-C: 50 |xm. D-G: 100 um. 



232 




Abb. 5: Hanguana bogneri, Blattanatomie. A-B: Zellmuster von Epidermis und Hypodermis 
in Aufsicht. A: abaxial, mit tetracytischen Stomata (dem Spalt zugewandte Zellwand der 
Schließzellen punktiert). B: adaxial. C-D: Epidermis, quer, mit Haarbasis. C: abaxial, mit 
reich verzweigtem Büschelhaar. D: adaxial, mit kurzem, wenig verzweigten Haar. Fz: Fuß- 
zelle. Interzellularen punktiert. Maßstabbalken: A-D: 100 p.m. 



233 







Abb. 6: Hanguana, Blattanatomie. A-D: Lamina, quer. A-B: deutliches Palisadenparenchym. 
A: H. „Borneo", mit nicht unterbrochener Hypodermis. B: H. bogneri. C-D: ohne deut- 
liches Palisadenparenchym. C: H. major. D: H. „Singapur". E-F: Laminarand, quer. E: 
H. „Borneo", mit Hautsaum aus zwei aufeinanderliegenden Epidermen. F: H. major, das 
Mesophyll reicht bis zum Blattrand. Leitbündel schematisiert. Interzellularen punktiert. S: 
Sekretzelle. Maßstabbalken: A-F: 300 M-hi. 



234 




Abb. 7: Hanguana, Blattanatomie, Querschnitte. A-B: H. bogneri. A: Mittelrippe der Lamina 
mit inversen, stark reduzierten Leitbündeln im adaxialen Mesophyll. B: Mittelrippe der 
Blattscheide, adaxialer Ausschnitt mit inversen Leitbündeln und lysigenem Schleimkanal. C: 
H. „Singapur", Blattscheidenrand. D-E: Mittelrippe des Blattstiels. D: H. „Singapur", ohne 
Schleimkanäle. E: H. bogneri, mit Schleimkanälen. Leitbündel schematisiert. Interzellularen 
und Schleimkanäle punktiert. S: Sekretzelle. Maßstabbalken: A-B: 300 ^im, C-E: 100 fim. 



235 




-Rp 



B>^-^^ 




k.' )W( '/$ 














Abb. 8: Hanguana, Wurzelanatomie, Querschnitte. A: H. „Borneo", Wurzelperipherie mit 
Rhizodermis und Exodermis, Rindenparenchym in Auflösung. B-C: innere Rinde und peri- 
pherer Zentralzylinder. B: H. major mit Phloemband zwischen Xylemstrahlen. C: H. „Bor- 
neo" mit Phloemnestem zwischen Xylempolen. D: H. bogneri, Zentralzylinder, schema- 
tisch, mit von Sklerenchym umgebenem Leitgewebe (Phloem punktiert) und kaum verholz- 
tem Markparenchym. En: Endodermis, Ex: Exodermis, M: Markparenchym, Pz: Perizykel, 
Rh: Rhizodermis, Rp: Rindenparenchym, S: Sekretzelle, T: Tracheide, Tr: Trachee, Wh: 
Wurzelhaar, X: Xylemprimane. Maßstabbalken: A-C: 100 um, D: 300 |j.m. 



236 




B 






Abb. 9: Flagellaria indica, Wurzelanatomie, Querschnitte. A: Wurzelperipherie mit Rhizo- 
dermis, Exodermis, Sklerenchymring und Rindenparenchym, Interzellularen teilweise mit 
Sekret gefüllt (schwarz). B: Rindenparenchym mit beginnender radialer Lakunenbildung. C: 
innere Rinde und peripherer Zentralzylinder. D: Zentralzylinder, schematisch (Ausschnitt), 
Leitsystem von^ Sklerenchym umgeben (Phloem punktiert), mit einzelnen Parenchym- 
nestem, Markgewebe unverholzt. En: Endodermis, Ex: Exodermis, M: Markparenchym, P: 
Parenchymnester, Pz: Perizykel, Rh: Rhizodermis, Rp: Rindenparenchym, Sk: Skleren- 
chymring, T: Tracheide, Tr: Trachee, Wh: Wurzelhaar X: Xylemprimane. Maßstabbalken: 
A-C: lOO^im, D:300^im. 



237 




<Ja''^uffMt}i^YU^^^^^^S^S^^'^ '■•jx-fi-v -i^y^tkLP^M 



238 



Abb. 10 (Farbtafel): Hanguana und Flagellaria. A-B: H. bogneri (männlich). A: blühende 
Pflanze. B: Blüte, zwei der drei großen Nektarien sind sichtbar. C-E: H. major (weiblich). 
C: ein Paracladium mit ähriger Floreszenz. D: Blüte. E: Samenanlage, die sich kuppelförmig 
über die massive Plazenta wölbt, die Pfeile weisen auf den Rand der Samenanlage, die 
Micropyle befindet sich in der durch den Stern markierten Erhebung. F: H. bogneri, Längs- 
schnitt durch die Blattspreite mit großen adaxialen Hypodermiszellen, einem längs geschnit- 
tenen Leitbündel der abaxialen Serie und einem quergeschnittenen Transversalbündel. G: 
Flagellaria indica, Wurzel-Querschnitt. Maßstabbalken: A: 10 cm, B, E, G: 1 mm, C: 5 cm, 
D: 5 mm, F: 300 ^im. 



239 



New taxa and new combinations published in Sendtnera 6 (3 1 . 1 2. 1 999) 

Alonsoa honoraria Grau, spec. nov. 

Astragalus sect. Aberrantes Podlech, sect. nov. 

Astragalus akhanii Podlech, spec. nov. 

Astragalus andreji-sytinii Podlech, nom. nov. 

Astragalus antalyensis A.Duran & Podlech, spec. nov. 

Astragalus arakliensis Podlech, spec. nov. 

Astragalus austroiranicus Podlech, nom. nov. 

Astragalus avajensis Podlech, spec. nov. 

Astragalus bakirdaghensis Podlech, spec. nov. 

Astragalus belangen (Kuntze) Podlech, comb. nov. 

Astragalus bowes-lyonii Podlech, spec. nov. 

Astragalus brevidentatus Podlech, spec. nov. 

Astragalus cedreticola A.Duran & Podlech, spec. nov. 

Astragalus demirizii R.Kramer «fe Podlech, spec. nov. 

Astragalus diversipilosus Podlech, spec. nov. 

Astragalus diyarbakirensis Podlech, spec. nov. 

Astragalus eskishehiricus Podlech, spec. nov. 

Astragalus fallacinus Podlech, spec. nov. 

Astragalus gigantostegius Podlech, spec. nov. 

Astragalus glaberrimus Podlech, spec. nov. 

Astragalus hakkariensis Podlech, spec. nov. 

Astragalus jacobsii Podlech, spec. nov. 

Astragalus juratzkanus subsp. brotherusii (Freyn & Sint.) Podlech, comb. nov. 

Astragalus karasarensis Podlech, spec. nov. 

Astragalus kashanensis Podlech, spec. nov. 

Astragalus kentrophyllus Podlech, spec. nov. 

Astragalus kirilovii Podlech, nom. nov. 

Astragalus konyaensis Podlech, spec. nov. 

Astragalus langtangensis Podlech, spec. nov. 

Astragalus laxmannii Jacq. subsp. robustior (Hook.) Podlech, comb. nov. 

Astragalus laxmannii Jacq. subsp. viciifolius (Welsh) Podlech, comb. nov. 

Astragalus longisubulatus Podlech, spec. nov. 

Astragalus longivexillatus Podlech & Ekici, spec. nov. 

Astragalus malatyaensis Podlech, spec. nov. 

Astragalus mercklinii Boiss. & Buhse subsp. farsicus Podlech, subspec. nov. 

Astragalus mikrophytoides Podlech, spec. nov. 

Astragalus sect. Mikrophyton Podlech, sect. nov. 

Astragalus mongholicus Bunge var. dahuricus (DC.) Podlech, comb. nov. 

Astragalus montis-varvashti Podlech, spec. nov. 

Astragalus nepalensis Podlech, spec. nov. 

Astragalus nevshehiricus Podlech, spec. nov. 

Astragalus neyshaburensis Podlech, spec. nov. 

Astragalus nigdeanus Podlech & Ekici, spec. nov. 

Astragalus nigrocalycinus Podlech, spec. nov. 

Astragalus pineticola Podlech, spec. nov. 



240 



Astragalus polhillii Podlech, spec. nov. 

Astragalus shahbazanicus Podlech, spec. nov. 

Astragalus sultan-bulaghensis Podlech, spec. nov. 

Astragalus tetuanensis Podlech, spec. nov. 

Astragalus thomsonii Podlech, spec. nov. 

Astragalus venulosus Boiss. subsp. iraqensis Podlech, subspec. nov. 

Astragalus villosulus Podlech, spec. nov. 

Bryonectria callicarpa Döbbeler, spec. nov. 

Bryonectria phyllogena Döbbeler, spec. nov. ., 

Coniophoroineae Agerer & Ch.Hahn, subordo nova 

Hanguana bogneri H.-J.Tillich & E.Sill, spec. nov. 

Limonium nydeggeri Erben, spec. nov. 

Omphalotus atraetopus (Kalchbr. in Thiimen) Ch.Hahn, comb. nov. 

Oxytropis azerbaijanica Podlech, spec. nov. 

Oxytropis khorasanica Ranjbar, nom. nov. 

Tapinella panuoides var. ionipus (Quel.) Ch.Hahn, comb. nov. 

Tapinella panuoides var. panuoides fm. acheruntius (Humboldt ex J.Schroet.) Ch.Hahn, comb. 

nov. ,1 

Tapinellaceae Ch.Hahn, fam. nov. 
Tapinellineae Agerer, subordo nova 
Truncocolumellaceae Agerer, fam. nov. 



Hinweise für die Autoren 

♦ Die Sendtnera (früher Mitteilungen der Botanischen Staatssammlung München) veröffentlicht wissenschaftliche Original- 
arbeiten und Kurzmitteilungen in deutscher und englischer Sprache (andere Sprachen nur nach Rücksprache) aus dem Gesamt- 
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behandeln. 

♦ Manuskripte und dazugehörige Abbildungen sind an eine der folgenden Adressen zu senden; 

Prof. Dr. J. Grau Dr. F. Schuhwerk 

Institut für Systematische Botanik der LMU-München Botanische Staatssammlung München 

Menzinger Straße 67 Menzinger Straße 67 

D-80638 München D-80638 München 

W (089)17861-254 Fax: (089)172638 "B (089)17861-240 

♦ Die Manuskripte müssen den an wissenschaftliche Arbeiten gestellten Anforderungen genügen. Auf klare Aussagen, Über- 
sichtlichkeit und größtmögliche Straffung des Textes ist zu achten. Der Titel muß kurz und dem Inhalt angemessen sein; er sollte 
Begriffe enthalten, die eine Indexierung erleichtern. Arbeiten, die diesen Anforderungen nicht genügen, müssen zurückgewiesen 
oder abgeändert werden. Die Herausgeber werden die eingereichten Manuskripte von Fall zu Fall an auf das behandelte Gebiet 
spezialisierte Experten weiterleiten. Die Entscheidung über Annahme oder Ablehnung der Arbeit bzw. über notwendige 
Änderungen wird den Autoren von den Herausgebern mitgeteilt. 

Mit der Annahme des Manuskriptes zur Publikation gehen die Verlagsrechte an die Botanische Staatssammlung München über; 
das geistige Eigentum an der veröffentlichten Arbeit bleibt hiervon unberührt. 

♦ Die Länge der Artikel sollte 50 Druckseiten nicht überschreiten. Die Veröffentlichung längerer Beiträge ist möglich, muß aber 
frühzeitig mit den Herausgebern abgesprochen werden. 

♦ Die Manuskripte müssen den nachfolgenden Hinweisen entsprechend abgefaßt, sorgfältig korrigiert und in ihrer endgültigen 
Form eingereicht werden. Autoren, die ihre Arbeit in einer anderen als ihrer Muttersprache verfassen, werden dringend gebeten, 
den Text vor der Abgabe durch eine Person mit entsprechenden Sprachkenntnissen überprüfen zu lassen. 

♦ Die Manuskripte sollten auf einem IBM-kompatiblen Computer mit Word für Windows oder Word, bzw. auf Apple Macintosh 
erstellt und auf Diskette (3,5 Zoll) mit zugehörigem Ausdruck eingereicht werden. 

Abbildungen müssen notwendig sein, in sinnvollem Zusammenhang mit dem Text stehen und im Original mit einem in der 
Legende erläuterten Maßstabs-Balken versehen sein. Tabellen und Abbildungen [Zeichnungen, Fotos als S/W-Hochglanz- 
abzüge, Diagramme, Karten (mit geographischen Koordinaten)] sind jeweils fortlaufend zu numerieren. Sie müssen inclusive der 
zugehörigen Legende in den Satzspiegel der Druckvorlage (16 x 25 cm) passen. Zeichnung und Beschriftung müssen auch nach 
der ca. 82%-igen Verkleinerung zum Druck noch alle Details erkennen lassen (Strichstärke beachten). 

♦ Jeder Arbeit ist eine kurze Zusammenfassung in der Sprache der Arbeit, bei deutschsprachigem Text eine weitere in Englisch, bei 
fremdsprachigen Arbeiten eine weitere in Deutsch voranzustellen. 

♦ Wissenschaftliche Sippennamen sind in Übereinstimmung mit dem "International code of botanical nomenclature" zu ver- 
wenden und bis incl. Gattung kursiv zu schreiben. Die Schreibweise der Autoren ist mit Brummit & Powell abzugleichen. Bei 
ausführlichen Zitaten von Sippennamen und Synonymen sind Einzelwerke nach Stafleu et al., Zeitschriftentitel nach 
"Botanico-Periodicum-Huntianum" abzukürzen. 

♦ Bei den Artbeschreibungen sind die Einzelmerkmale in Komplexe zusammenzufassen. Auf größtmögliche Straffung und gute 
Vergleichbarkeit der einzelnen Beschreibungen untereinander ist zu achten. 

♦ Herbarbelege sollten nur in erforderlichem Ausmaß zitiert werden (repräsentative Belege). Etiketten-Texte sind zu standar- 
disieren und in folgender Sequenz zu zitieren: Staat. Politische Untereinheit/Region : Fundort, (geographische Koordinaten, 
Höhe, Angaben zu Standort und Vergesellschaftung), Datum, Sammler Sammel-Nummer (Herbarium, evtl. Herbar-Nummer - 
Abkürzungen nach "Index herbariorum"). 

♦ Bestimmungsschlüssel sind nach folgendem Muster zu erstellen: 

1 Nadeln zu 5 in Kurztrieben gebüschelt 2 

- Nadeln zu 2-3 in Kurztrieben gebüschelt 4 

2 Nadel 8-15 cm lang Pinus nigra 

- Nadeln 3-8 cm lang 3 

♦ Literaturhinweise im Text sind durch Angabe des Autorennamens und Erscheinungsjahres zu zitieren, z.B. Huber (1969). In das 
Literaturverzeichnis sind nur im Text aufgeführte Arbeiten aufzunehmen. 

GoTTSBERGER, G. 1977: Some aspects of beetle pollination in the evolution of flowering plants. - PI. Syst. Evol., Suppl. 1: 

253-283. 

Grisebach, A.H.R. 1839; Genera et species Gentianearum. Stuttgart. 

Hoffmann, O. [1890-]1894: Compositae. - In; Engler, A. & Prantl, K. (eds); Die natüriichen Pflanzenfamilien 4(5); 87- 

387. Leipzig. 

Huber, H. 1991: Leitfaden durch die Ordnungen und Familien der Bedecktsamer. Stuttgart. 

Maheshwarl S.C. & Baldev, B. 1958: A contribution to the morphologic and embryology of Commelina forskalaei Vahl. - 

Phytomorphology 8; 277-298. 

♦ Die Anschrift der Verfasser ist mit Titel und ausgeschriebenem Vornamen anzugeben. 

♦ Nach der Einrichtung als Druckvorlage erhalten die Autoren einen Ausdruck ihrer Arbeit, der innerhalb von zwei Wochen auf 
Fehler hin durchgesehen und zurückgesandt werden muß. Inhaltliche Änderungen gegenüber dem eingesandten Manuskript sind 
jedoch nur in Ausnahmefällen möglich. 

♦ Die Autoren erhalten 50 Sonderdrucke ihrer Arbeit kostenlos geliefert. Sind mehr als 50 Sonderdrucke gewünscht, so muß dies 
mit den Herausgebern vereinbart und zum Selbstkostenpreis berechnet werden. 

♦ Annahmeschluß für die Sendtnera Band 7; 30.6.2000. 



Inhalt 



AGERER, R.: Never change a functionally successful principle: The evolution of 
Boletales s.l. (Hymenomycetes, Basidiomycota) as seen from below-ground 
features 5 

DÖBBELER, P.: Two new species of Bryonectria (Hypocreales, Ascomycetes) on 
bryophytes 93 

Erben, M.; Limonium nydeggeri - eine neue Art aus Südwestportugal 103 

Grau, J.: Eine neue Art der Gattung Alonsoa (Scrophulariaceae) aus Chile 109 

Hahn, C. & Agerer, R.: Systematical studies on Paxillaceae (Boletales, Basidio- 
mycota) 115 

PODLECH, D.: New Astragali from North Africa and Asia, including some new 
combinations and remarks on some species 135 

PODLECH, D.: Typification of Astragalus species III. (Leguminosae) 175 

Ranjbar, M.: Some remarks on the genus Oxytropis (Fabaceae) from Iran 193 

Schuhwerk, F. & Lippert, W.: Chromosomenzahlen von Hieracium L. (Com- 

positae. Lactuceae) Teil 3 197 

Tillich, H.-J. & Sill, E.: Morphologische und anatomische Studien an Hanguana 215 

Blume (Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit der Beschreibung 
einer neuen Art, Hanguana bogneri spec, nov 

New ta.xa and new combinations published in Sendtnera 6 (31.12.1999) 239 





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