82 Annalen des Naturhistorischen Museums in Wien, B, 111

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R ie k e n B., 2 0 0 3 : Arachne und ihre Schwestern. Eine Motivgeschichte der Spinnen von den "Naturvölkermärchen" bis zu den "Urban Legends" - Waxmann, Münster, NewYork, München, Berlin: 1 - 2 8 7 .

R o t t e r D., 2006: Einfluss der Verbuschung auf die Artengemeinschaften auf Heißländen in der Unteren Lobau. - Wissenschaftliche Reihe Nationalpark Donau-Auen Heft 22. 33 pp.

R ü z ick a V & B u c h a r J., 2008: Dodatek lce katalogu pavoukü Ceske republiky 2001-2007 [Supplement to the Catalogue o f Spiders of the Czech Republic 2001-2007]. - Sbornik Oblastniho muzea v Moste, fada prirodovednä, 2007/2008: 3-32.

S c h ra n k F. vo n P., 1781: Enumeratio insectorum austriae indigenorum. - Augustae Vindeli- corum, 552 pp. (Araneae, pp. 526-534).

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H p p n e r , H ö r w e g & M i l a s o w s z k y : Bibliographische Checkliste der Spinnen (Araneae) Wiens 83

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84 Annalen des Natuvhistorischen Museums in Wien, B, 111

Buchbesprechung

S y s o e v A. & S c h i l e y k o A. 2009. Land Snails and Slugs of Russia and Adjacent Countries.

Pensoft (Pensoft Series Faunistica 87, ISSN 1312-0174), Sofia, 454 pp.,ISBN 978-954-642-474-7, € 80,-(http://pensoft.net/newreleases/index.htm)

Die publizistische Tätigkeit der russischen Zoologen ist mit dem Zerfall der Sowjet­union nicht zum Erliegen gekommen. Im Gegenteil, sie ist uns näher gerückt. Nicht nur, dass die meisten Veröffentlichungen am globalen Markt erhältlich sind, sind sie wesent­lich besser erschließbar, weil viele auf Englisch verfasst sind. Das 2009 erschienene Buch über die Landschneckenfauna erstreckt sich laut Titel auf Russland und angren­zende Gebiete, reicht aber in seiner Bedeutung weit über diese Grenzen hinaus. Das immer noch riesige Russland hat ja beträchtliche Anteile an unterschiedlichen tiergeo­graphischen Regionen. Die Publikation ist also nicht nur für jene von großem Wert, die an den Schnecken innerhalb der genannten Grenzen interessiert sind, sondern auch für alle, die im Kaukasus, in so distinkten Regionen wie Ostasien und Osteuropa, Skandina­vien oder an den Gestaden des Schwarzen oder Kaspischen Meers Interesse haben. Sie umfasst eigentlich große Teile der Paläarktis.Die Fotografien des Erstautors sind brillant und auch der Druck ist von hinreichender Qualität, wenngleich ein wenig mehr Perfektion zu ertragen gewesen wäre. Alexander Sysoev ist bereits als Autor zahlreicher fachspezifischer Publikationen hervorgetreten, darunter umfassende Kataloge der russischen Molluskenfauna. Der fachlich malakologische Input des Zweitautors ist ebenfalls zweifelsohne von höchstem Gewicht. Anatoly A. Schileyko ist Autor zahlreicher Fachpublikationen, zu nennen und zu empfehlen sei an dieser Stelle sein "Opus magnum" der "Treatise on recent Terrestrial Pulmonate Molluscs", der in 14 Bänden Ordnung in die Vielfalt der weltweit vorkommenden Landschnecken bringt. Aus diesem ihrem reichen Erfahrungsschatz schöpfend, haben die beiden in dem vorliegenden Bildband eine schöne Ordnung in die Schneckenwelt des größten Landes der Welt gebracht. Als Grundlagen für Abbildungen und Beschreibungen dienten hier, wie in vergleichbaren Werken, die wissenschaftlichen Sammlungen von naturkundli­chen Museen. Russland hat mit Moskau und St. Petersburg zwei hervorragende Samm­lungen als Basislager sozusagen. Schileyko ist allerdings auch als Spezialist internatio­nal. Seine Kenntnis der weltweit wichtigsten Forschungssammlungen erstreckt sich von Washington über Paris bis Wien.

Das Buch ist gleichzeitig reich und sparsam. Zusätzlich zu den vielen farbigen Abbil­dungen, geben die Kurzportraits zu den einzelnen Arten Auskunft über die Typusloka­lität, den Standort der Typen, die Schalenmaße und die Verbreitung der Tiere. Auch eine erstaunlich umfangreiche Synonymie ist im Angebot. Und manchmal werden wir auch noch mit kommentierenden Anmerkungen zu einzelnen Arten versorgt. Das Werk gibt außerdem Aufschluss über Erstbeschreibungen und die wichtige historische wie aktu­elle Fachliteratur. Was fehlt, sind Beschreibungen und Unterscheidungen der Arten. Aber das wäre wohl zu viel verlangt, bei einer Fauna, die fast 800 Arten und Unterarten behandelt. Ein schönes, ein inhaltsreiches und ein kompaktes Buch. Dieses gehört in die Bibliothek jedes Malakologen und natürlich jedes faunistisch-systematisch orientierten Institutes, das über den Tellerrand der eigenen regionalen Tierwelt hinausschaut.Dr. Helmut SattmannDritte Zoologische Abteilung, Naturhistorisches Museum Wien, Burgring 7, 1010 Wien, Austria

Ann- Naturhist. Mus. Wien, B 85-99 Wien, März 2010

New taxa of Gundelia (Compositae) from Armenia

E. Vitek*, G. Fayvush**, K. Tamanyan** & B. Gemeinholzer***

A bstract

Two new taxa, Gundelia aragatsi and G. aragatsi ssp. steineri are described. Gundelia was mainly treated as monospecific genus - this status cannot be maintained. The differences of the new taxa, the status of a probable further new species, Gundelia sp. (Garni), and the important characters for future research are dis- cussed.

Key W ords: Flora of Armenia; Compositae, Gundelia, G. tournefortii, G. rosea, G. aragtsi, G. aragatsi ssp. steineri; new species, systematics, taxonomy.

K urzfassung

Zwei neue Taxa, Gundelia aragatsi und G. aragatsi ssp. steineri werden in der bisher meist als monospezi­fisch betrachteten Gattung Gundelia beschrieben. Die Unterschiede, der Status einer möglichen weiteren unterscheidbaren Art, Gundelia sp. (Garni), sowie die für die weitere Erforschung wichtigen Merkmale werden diskutiert.

Introduction

Gundelia tournefortii was known in A rm enia from a restric ted area around Garni (Avetisisan 1995). In 2006 a non professional botanist, Alfred Steiner, m em ber o f a hiking tour group led by the authors, found a new population o f Gundelia near Eche- gnadzor, c. 85 km SE o f the earlier known area. Later in the same year the first author found another population o f Gundelia on Mt. Aragats, c. 85 km WNW o f the earlier known area. These populations are so different in their habit to the known one, that they could not be subsummarized under G. tournefortii, the only accepted name in the rele­vant floras for this area (Vasil 'chenko I.T. 1961, K upicha 1984, R echinger 1989, Avetisian 1995, SOFIEVA 1961).

Material and methods

Material has been collected on the same localities in 2006, 2007, 2008 and 2009 at dif­ferent times of the year. Specimens are deposited in W 1 and ERE, duplicates also in

Dr. Ernst Vitek, Naturhistorisches Museum Wien, Botanische Abteilung, Burgring 7, A-1010 Wien, Austria - ernst.vitek@nhm-wien.ac.atDr. George Fayvush, Dr. Kamilla Tamanyan, Institute for Botany, Academy of Sciences, Botanic Garden, Acharian str. 1, Yerevan 0063, Armenia - gfayvush@yahoo.com, ktamanian@yahoo.com

***Dr. Birgit Gemeinholzer, Botanischer Garten und Botanisches Museum Berlin Dahlem, Freie Univer­sität Berlin, Königin-Luise-Straße 6-8, 14195 Berlin, Germany.

1 Abbreviations for herbaria follow Index herbariorum: http://sweetgum.nybg.org/ih/

86 Annalen cles Naturhistorischen Museums in Wien, B, 111

other herbaria. The material was morphologically evaluated. Furthermore, leaf material for DNA analysis has been transferred to Silicagel in the fleld and handed over to B. Gemeinholzer (Berlin). Total genomic DNA was isolated from 5 Armenian samples as well as from 10 dried herbarium specimens covering big part of the distribution ränge of the genus (details for specimens in Appendix 1). 24 mg per sample of mostly leaf mate­rial was taken. The samples were crushed and DNA was then extracted using Quiagen DNeasy Mini Kit and following Standard procedure.

The ITS region was amplified in two overlapping parts using the primers ITS-A and ITS-C (B la ttn e r 1999) for ITS 1, ITS2-D, ITS-B (B la t tn e r 1999) for ITS 2 or ITS-A and ITS-B for the complete region. The following protocol during the Polymerase Chain Reaction (PCR) was used: initial denaturation 2 min at 94°C, denaturation 20 sec at 94°C, annealing 45 sec at 52°C, elongation 1 min at 72°C (40 cycles) and final extension 10 min at 72°C. PCR was carried out with a reaction volume of 11.5jul core mix plus 1 jlxI DNA (1:10, 1:50 or 1:100 dilution depending on usability). The reaction volume con- tained 8.05 [i\ ddH20, 1.25 (il lOx buffer (Biothemi) and dNTP’s (Fermentas), 0.25 |il BSA (BioLabs), 0.25 jul o f each primer (10pmol/|il) and 0.04 jul lu Taq-Polymerase (Biotherm). The PCR products were purified with Milipore DNA purification Kit (Roth) and then cycle sequencing was carried out using CEQ DCTS Quick Start Kit (Beck- mann-Coulter) following the Standard procedure. As sequencer a CEQ8000 (Beckmann- Coulter) was used. The sequences were edited in ChromasLite2000 (Technelysium Pty. Ltd., Helensvale, Australia) and aligned by hand using BioEdit (H a ll 1999). As not for all sequences the 5.8S rDNA was amplified and there are no expected mutations any- way, this region was excluded for calculation. The alignments is available from B. Gemeinholzer upon request. The trees were reconstructed using MrBayes 3.1.2 (Ron- quist & H uelsenbeck 2003) and PAUP 4.0b 10 (Swofford 2002). A Bayesian Analysis was performed using gamma distribution rate Variation among sites and 3 million gen- erations of the MCMC chains in two independent runs, trees saved every 100 genera- tions. The first 30 000 trees were discarded as burn-in for the analysis then reached sta- tionarity. All other trees sampled were used to calculate a strict consensus tree.

The datasets were also analysed by using Maximum Parsimony. All heuristic searches were conducted in Paup 4.0b 10 with equal weights, 1000 closest sequence additions and tree bisection-reconnection (TBR) branch swapping, permitting 10 trees to be held at each step. An evaluation of the trees was performed by using bootstrap analysis with 1000 replicates, equal weights, TBR swapping, MulTrees option in effect and 10 trees held at each step. The strict consensus tree of Bayesian inference was compared to the bootstrap 50% majority rule consensus tree. Trees were drawn using TreeView (Page1996).

Results of DNA-analyses

The ITS region was amplified for a total o f 15 individuals from throughout the genus distribution ränge. For calculations an already published sequence of Gundelia tourne- fortii (AY504691) with unknown providence was added as well as Catananche caerulea (AJ633465, AJ633466, AJ633467), C. caespitosa (AJ633468), C. arenaria (DB472), Scorzonera purpurea ssp. purpurea (AJ633477), S. crispatula (AJ633486), S. villosa

V it e k , F a y v u s h , T a m a n y a n & G e m e i n h o l z e r : New taxa o f Gundelia (Compositae) from Armenia 87

53/0.79

rC66/ 1.00

64/0.87

100/1.00

100/1.0087/1.00

99/0.97 PC100/1.00

100 /1.0061/0.82

96/0.98

100/ 1.00

PC

66/0.56 C

*G. aragatsi ssp. steineri Armenia FN582290*G. aragatsi ssp. steineri Armenia FN5 82289*G. aragatsi Armenia FN582288*G. aragatsi Armenia FN5 82283*G. sp. Armenia FN582287*G. tournefortii Turkey FN582281*G. tournefortii Iran FN582284*G. tournefortii Iran FN582277*G. tournefortii Iran FN582285*G. tournefortii Iran FN582278*G. tournefortii Afghanistan FN582275*G. tournefortii AY504691*G. tournefortii Iran FN582282*G. tournefortii Afghanistan FN582279*G. tournefortii Azerbaijan FN582280*G. tournefortii Turkey FN582286Scorzonera purpurea AJ633477Scorzonera crispatula AJ633486Tragopogon dubius AJ633503.1Scorzonera villosa AJ633482Scolymus maculatus AJ633469Catananche caerulea AJ633465Catananche caerulea AJ633467Catananche caerulea AJ633466*Catananche arenaria FN582275Catananche caespitosa AJ633468Warionia saharae A Y 190608

Fig. 1: Majority rule consensus tree of the MP analysis with bootstrap values > 50 presented first and posterior probabilities > 0.50 second. * are new results (vouchers see appendix 1).

(AJ633482), Tragopogon dubius (AJ633503), using Warionia saharae (AY190608) as outgroup. All calculations confirmed Gundelia to be monophyletic with strong statisti- cal support (100% bootstrap support/1.00 likelihood support).

Within Gundelia the analysis of the ITS region revealed little differentiation. The Arme- nian specimen form a separate cluster together with one yellow flowered accession from Turkey branching off basally. However the whole clade is only supported by veiy low bootstrap and likelihood support (53%/0.79). The Armenian taxa of Gundelia feature close relationships.

The low genetic differentiation within the analyzed genetic marker reflects the limited variability of the ITS region within the genus due to recent differentiation or possibly pinpoints to a much more complex evolutionary picture of hybridization and adaptation. For further analyses additional material, especially doubtless G. rosea and material from the type area of G. tournefortii needs to be screened with molecular methods discrimi- nating on a smaller scale.

8 8 Annalen des Naturhistorischen Museums in Wien, B, 111

Fig. 2: a) Gundelia tournefortii, Karacadag bei Diyarbakir, SE-Turkey; b-c) Iran, Tehran, ca. 20 km to Karadj, Alborz mountains. - a) © G. Pils, b) © F. Ghahremaninejad.

V it e k , F a y v u s h , T a m a n y a n & G e m e i n h o l z e r : New taxa of Gundelia (Compositae) from Armenia 89

Fig. 3: Gundelia sp., Annenia, near Garni: a) habitus, note the glabrous leaves; b) infructescence; c) single disseminule aggregated of 6 flowers; d) inflorescence with campanulate flowers, closed at ± noon, e-f) flowers, opening in ± late afternoon.

9 0 Annalen des Naturhistorischen Museums in Wien, B, 111

Fig. 4: Gundelia aragatsv. a) habitat mountain steppe; b-c) probable pollinators, b) honey bee, Apis mellifera, c) pollen feeding beetles, Phyllopertha horticola; d) disseminules aggregated of three flowers; e) rare orange flowers; f) widely open dark brown flowers.

V ite k , F a y v u s h , T a m a n y a n & G e m e i n h o l z e r : New taxa o f Gundelia (Compositae) from Armenia 91

Fig. 5: Gundelia aragatsi ssp. steineri: a) locality in shrubby steppe; b) disseminules aggregated of three flowers; c) unripe infuctescence; d) inflorescence with campanulate flowers; e-f) dark to blackish brown flowers opening late afternoon, g) rare orange flowers. - d) © A. Steiner.

92 Annalen des Naturhistorischen Museums in Wien, B, 111

Results of morphological analyses

Searching for a correct name for the three populations of Gundelia in Armenia it was realised, that the Linnean name of G. tournefortii was not fixed by a type. This was done by V it e k & J a r v is (2007), choosing material from Rauwolf as type and such connecting the name to cream flowering Gundelias in Aleppo (Syria) (R a u w o l f 1583).

Differences in flower colours have been used to distinguish Gundelia tournefortii f. pur- purascens B o r n m . (= G. purpurascens B o r n m ., nom. inval. in sched.) and var. asper- rima T r a u t v . Later G. rosea has been described ( A l -T a e y & H o s s e in 1984). Other attempts to distinguish different units within Gundelia concentrated on the shape of leaves, form and length of bracts: var. armata F r e y n & S in t ., var. tenuisecta Boiss. (= G. tenuisecta (Boiss.) F r e y n & S i n t .). But these characters are obviously variable in both, G. tournefortii and G. rosea (A l -T a e y & H o s s e in 1984) and therefore proved not to be useful for distinguishing taxonomic units. A s consequence G. rosea was reduced to synonymy (e.g. R e c h in g e r 1989) and Gundelia treated as monospecific genus with the only variable species G. tournefortii.The three populations in Armenia differ in many characters from each other and from G. tournefortii and G. rosea (Table 1): Hairiness of leaves and inflorescence, flower colour and shape, observed respectively assumed pollinator, size and aggregation o f dis- seminule. Therefore the two taxa, which cannot be assigned to any earlier known species are described here as new.

The third one has similar flower colour to G. rosea, but the drawing o f the fruit o f G. rosea in A l-T aey & H o s s a in (1984) show double size and different form - therefore also this population could constitute a well defined species. We refrain at the moment to describe it, unless Gundelia glabra M ille r , a taxon described from "near Baibout in Armenia" (now Bayburt, Turkey) put in synonymy to G. tournefortii by K u p ic h a (1984), is clarified and typified - the epithet "glabra" and the description " acanthi aculeati folio, capite glabro" would perfectly fit the plants from the surroundings o f Garni (c. 14 lcm E o f Yerevan). The habitat is a forbs-grasses steppe with Aegilops cylindrica, Avena sterilis, Bromus danthoniae, Bromus tectorum, Festuca valesiaca, Hordeum bulbosum, Koeleria macrantha, Secale montanum, the forbs Acroptilon repens, Anchusa orientalis, Cephalaria syriaca, Consolida orientalis, Falcaria vulgaris, Glaucium elegans, Hypecoum pendulum, Picnomon acarna, Reseda lutea, Roemeria refracta, Xeranthe- mum squarrosum, and spring ephemeroids Iris elegantissima, Merendera trigyna.

The authors discussed several times which taxonomic status should be given to the new Gundelia from Mt. Aragats. The dark brown flowers - especially in connection with the rare orange flowered plants - could be interpreted as extreme coloured variant of the yel­low G. tournefortii. The findings of the differences in the aggregation of the flowers and the fraits (Fig. 6) were the last impetus to describe it as distinct species: G. aragatsi.

Gundelia aragatsi V itek, Fayvush, Tamanyan & Gemeinholzer, sp.n.

Descriptio: Herba perennis, lactifera, crassicaulis. Folia altema, pinnatisecta usque ad ± incomplete pinnatisecta, spinoso-dentata, superiora decrescentia et minus dissecta, arachnoidea. Inflorescentiae solitariae, usque ad 8 x 5 cm (bracteis inclusis), dense

V ite k . F a y v u s h , T a m a n y a n & G e m e i n h o l z e r : New taxa of Gundelia (Compositae) from Armenia 93

* ♦ * * ♦ < » i

» M u t t

*■

> * 4

^

* | ^ ^ ^ 0 Ä

t n % # i ■

i $

% 0 H $f 1§ 1§ %« % f t M v

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Fig. 5: Ripe disseminules of a) Gundelia aragatsi, Teher 11.7.2007, b) G. aragatsi ssp. steineri, Oganesian & al. 07-0055, c) G. sp. (Armenia), Teher 11.7.2007.

arachnoideae. Bracteae spinoso-marginatae, inflorescentiam vix excedentes, apice in spinam induratam protractae. Flores explicate aperientes, fusci ad pulli, rariter auran-tiaci. Fractus fere 3 (-4) floribus compositi, usque ad 12 mm longi.

A G. toumefortii et G. rosea floribus fuscis et forma fructuum, ab G. sp. (Gami) indu-mento arachnoideo, floribus fuscis et forma fructuum differt.

Differs of G. toumefortii and G. rosea by the dark brown flowers and the form of the fruits composed only of three flowers, differs of G. sp. (Gami) by the dark brown flow­ers, the form of the fruits and the arachnoid hairs on leaves and inflorescence.

Distribution: Mt. Aragats, southwestem flank, around villages Baysiz and Kakavadzor; observed area with more than 2 km diameter, but probably also further north and north- east (Fig. 7).

Type: Armenia, Aragatsotn province, Mt. Aragats SW-slope, track between Avtona water reservoir and Kakavadzor, 1880 m s.m ., 40°22'5"N/44°2'49"E, 2007-06-23 G. Fayvush, K. Tamanyan, H. Ter-Voskanyan, E. Vitek 07-1373 [holotype ERE, isotypes W (four sheets) 2009-18514-2009-18517, B, BC, BM, BRNU, E, MO, MSB, NY, WU].

Other material: Armenia, Aragatsotn province, SE slopes of Mt. Aragats, track from water reservoir below Avtona to Kakavadzor, 1890 m s.m ., 40°22'5"N/44°2'49"E, 2006-06-05 E. Vitek 06-1218 [ERE, W 2009-18510-2009-18511, B]; - Armenia, Ara­gatsotn province, SE slopes of Mt. Aragats, road from Kakavadzor to Baysiz, 1760 m

94 Annalen des Naturhistovischen Museums in Wien, B, 11 ]

T ab. 1: C o m p a r iso n o f ch aracters o f th e A rm en ia n taxa o f Gundelia w ith G. tournefortii and G. rosea.

C haracter

material

G. aragatsi (F ig . 4 )

Armenia, S flank o f Mt. Aragats

G. aragatsi ssp . steineri (F ig . 5)

Armenia, near Echegnadzor

leaves

inflorescence

hairy

densely hairy

densely hairy’1

densely hairy

Flowercolour

shape o f upper part

opening o f flower

observed flower visitors / probable pollinators

flower aggregate of

ripe fruits** of

cypselaform

outside blackish brown, inside dark brown, very rarely dark orange

spread

widely open during the day

beetles (pollen feeders),Phyllopertha horticola (H. Schönmann, pers. comm.); honey bee, Apis mellifera (H. Zettel, pers. comm.)

3 (-5) flowers

3 flowers

(Fig. 6)± triangular9-12 x 6-9 x 3-5 mm

outside greyish (rusty) brown to blackish brown, inside blackish brown, very rarely dark orange

campanulate

closed during the day, opening afternoon (or dark sky)

? (possibly night butterflies)

3 flowers

3 flowers

(Fig. 6)± triangular9-12 x 7 -9 x 3-5 mm

transverse section

habitat

altitude, m s.m.

naiTow ovate

mountain steppe

1700-1900

narrow ovate

shibliak

1000-1100

* Long, soft, multicellular hairs, for the evolutionary value of this character see Karis & al. 2001.** In the field in rare cases different numbers o f flowers aggregating to one disseminule could be observed,

especially in the lowermost and uppermost parts o f inflorescences. The analysis o f ripe fruits later in the year showed a nearly constant number.

V it e k , F a y v u s h , T a m a n y a n & G e m e i n h o l z e r : New taxa o f Gundelia (Compositae) from Armenia 95

O f G. tournefortii and G. rosea n o fresh and l iv in g m a ter ia l c o u ld b e in v e s t ig a te d . T h e data given in floras can b e u sed o n ly w ith doubt - o b v io u s ly there is m ore than o n e taxon (se e d iscu ss io n ).

G. sp. (G arn i) (F ig . 3 )

Armenia, near Garni

glabrous

± glabrous, with very few hairs on bracts

G. tournefortii (F ig . 2 )

based mainly on the type and photos

hairy to densely hairy

hairy to densely hairy

G. rosea

extracted from A l -T a e y & H o s s a in ( 1 9 8 4 )

?

?

dark pink to purple, rarely whitish (N e r s e s y a n , pers. comm.)

creamy to yellow (green) bright to light pink inside, deep purple outside

campanulate to slightly spread

closed during the day,opening late afternoon (or dark sky)

spread

widely open during the day

? (possibly night butterflies)

6(-9) flowers

6 flowers

5-7 (_?)*** flowers

5-7 flowers ***

more than 5-8 (-11)

?

obconical11-15 x 7-11 x 7_11 mm

± circle

± humid steppe

1 5 0 0 -1 6 0 0

obconical1 0 -1 5 x 7 x 7 m m

( A l - T a e y & H o s s a in 1 9 84 ,

M e i k l e 1 9 8 5 )

± circle

dry slopes, sometimes (semi-) ruderal, weed in fields ( 0 - ) 1 0 0 -2 5 0 0

(P il s 2 0 0 6 , M e ik l e 1 9 8 5 )

obovoid 15-20 mm

cf. ± circle

pine woodland, on red rocky soil 8 0 0 -9 0 0 (- ? )

***The complex structure o f the inflorescence of Gundelia with disseminules aggregated o f 5 -7 flowers is described by C l a s s e n - B o c k h o f f & al. 1989. It is supposed that the investigated material from SW- Turkey is G. tournefortii

96 Annalen des Naturhistorischen Museums in Wien, B, 1} j

s.m ., 40°21,4 r ,N /4 4 °3 '4 r,E, 2006-06-05 E. Vitek 06-1233 [ERE, W 2009-18503- 2009-18504]; - Armenia, Aragatsotn province, Mt. Aragats SW-slope, above road Kakavadzor - Baysiz, 1720 m s.m., 40o21'41"N/44°3l41"E, 2007-06-23 G. Fayvush, K. Tamanyan, H. Ter-Voskanyan, E. Vitek 07-1405 [ERE, W 2009-18512]; - Armenia, Aragtsotn province, SE slope o f Mt. Aragats, NE of Baysiz, 1860 m s.m., 40°22'14"N/ 44°03'47"E, 2009-06-10 G. Fayvush, K. Tamanyan, E. Vitek 09-0839 [ERE, W 2009- 18588-2009-18589]; - Armenia, Aragtsotn province, SE slope of Mt. Aragats, E of Baysiz, 1810m s.m., 40°2r46"N/44°04'H"E, 2009-06-10 G. Fayvush, K. Tamanyan, E. Vitek 09-0840 [ERE, W 2009-18505-2009-18507]; - (fruits) Armenia, Aragatsotn pro­vince, SE slopes of Mt. Aragats, track from water reservoir below Avtona to Kakavad­zor, 1875 m s.m., 40°21'59"N/44o2'44"E, 2007-07-11 S. Ter-Pogoshyan [ERE, W 2009- 18773, B, BC, E, NY],

The habitat on Aragats mountain is an intensively grazed mountain meadow-steppe with tragacanth elements: Gundelia aragatsi, Astragalus microcephalus, Achillea mille- folium, Dactylis glomerata, Poa bulbosa, Scutellaria orientalis, Cerinthe minor, Lotus caucasicus, Rindern lanata, Cousinia brachyptera, Anisantha tectorum, Alkanna orien­talis, Prangos ferulacea, Asperula glomerata, Lappula barbata, and rare and very sparse shrubs: Cotoneaster integerrimus, Rhamnus cathartica, Crataegus meyeri.

The name remembers the locality on Mt. Aragats.

The population near Echegnadzor is without any doubt closely related to G. aragatsi - also confirmed by the low genetic differentiation (Fig. 1) - but they should be differed taxonomically. Based on the marked differences in habitat and pollinator (opening time of flowers), and some morphological differences (e.g. shape and colour of flowers) the rank of subspecies seems to be adequate, and therefore these plants are described as Gundelia aragatsi ssp. steineri.

Gundelia aragatsi ssp. steineri V itek , Fayvush, Tamanyan & G em einholzer ssp.n.

Descriptio: Herba perennis, lactifera, crassicaulis. Folia altema, pinnatisecta usque ad ± incomplete pinnatisecta, spinoso-dentata, superiora decrescentia et minus dissecta, arachnoidea. Inflorescentiae solitariae, usque ad 8 x 5 cm (bracteis inclusis), ± dense arachnoideae. Bracteae spinoso-marginatae, inflorescentiam vix excedentes, apice in spinam induratam protractae. Flores campanulati, aperientes post meridiem, umbrini. Fructus fere 3 floribus compositi, usque ad 12 mm longi.

A Gundelia aragatsi floribus umbrinis aperientibus post meridiem, altitudine distributio- nis et habitatione differt.

Differs of G. aragatsi by the campanulate flowers, opening in the aftemoon (or at dark sky), and in growing in a different altitude and habitat.

Distribution: Only the type locality known, one population in an area of c. 100 x 200 m (Fig. 7).Type: Armenia, Vayots Dzor province, mainroad to south Armenia, W of Yeghegnadzor, SE of crossroad to Erechgnadzor, slope S of river, 1050 m.s.m, dry rocky slope, 39°44'21"N/45015'3"E, 2009-06-01 G. Fayvush, K. Tamanyan, E. Vitek 09-0700 [holo­type ERE, isotypes W 2009-18513, B, BC, E, MO, MSB, NY, WU].

V iti-'K, F a y v u s h , T a m a n y a n & G e m e i n h o l z e r : New taxa of Gundelia (Compositae) from Armenia 97

Fig. 7: M ap o f A r m e n ia w ith the f lo r is t ic p r o v in c e s (T a k h ta ja n 1 9 5 4 ) and th e lo c a lit ie s o f Gun­delia aragatsi ( ★ ) , G. aragatsi ssp . steineri ( K ) and Gundelia sp . ( + ) b a se d on sp e c im e n s and fie ld n o tes . F or b etter o r ien ta tion Y er ev a n ( □ ) and M t. A raga ts ( ^ are sh o w n .

Other material: Armenia, Vayots Dzor province, mainroad to south Armenia, W of Yeghegnadzor, SE of crossroad to Erechgnadzor, 2006-07-26 H.Ter-Voskanyan, A. Aslanyan [ERE]; - Armenia, Vayots Dzor province, mainroad to south Armenia, W of Yeghegnadzor, SE of crossroad to Erechgnadzor, slope S of river, 1050 m.s.m, dry rocky slope, 39044'21"N/45°15'3"E, 2007-06-11 M. Oganesian, H. Ter-Voskanyan, E. Vitek07-0055 [ERE, W 2009-18501-2009-18502, B, MSB, NY].

The habitat in the gorge of Arpa river is a Shibliak formation, in which Gundelia aragtsi ssp. steineri is a main dominant of the community, accompanied by shrubs: Atraphaxis spinosa, Eurotia ceratoides, Rosa spinosissima, and herbs: Achillea millefolium s.L, Aegilops cylindrica, Artemisia fragrans, Melica transsilvanica, Stachys inflata, Thymus kotschyanus.

The name is given in honour of Alfred Steiner (Oberwaltersdorf, A ustria), who detected this population.

Discussion

Earlier attempts for distinguishing taxa within Gundelia using flowers colour, the length of the bracteose leaves in the inflorescence and the form of the leaves failed. These char- acters are variable and obviously the leaf characters are not correlated with natural taxo- nomic units. Therefore Gundelia for long time was seen as monospecific genus. The description of Gundelia rosea has been ignored or this taxon was treated as synonym to G. toumefortii. The here reported new results show that Gundelia is not monospecific, there are clearly distinguishable species within the genus. The important characters seem to be flower colour, shape and size of fruits and possibly also the differences of the

98 Awicilen des Naturhistorischen Museums in Wien, B, 111

indumentum. The ITS tree shows several clades, but the material cannot be correlated with these morphological characters at the moment, because for many of the herbarium collections part of this information is lacking. Future collected material with full Infor­mation will allow a clearer picture on the evolution of Gundelia.

Acknowledgements

We thank the other members of the Annenian collecting team: M. Oganesian and H. Ter-Voskanian (Yerevan) for support during field work and S. Ter-Poghosyan (Yerevan) for collecting ripe fruits. A. Steiner (Ober­waltersdorf), F. Ghahremaninejad (Tehran) and G. Pils (Spittal/Drau) allowed the use o f photos, H. Schön­mann and H. Zettel (Vienna) determined the pollinators, B. Elci (Ankara) provided plant material from Turkey, and H. Fleischer-Notter (Berlin) assisted in the laboratory.

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A ppendix 1: M ater ia l u se d for D N A -a n a ly se s . T h e c o m p le te data o f th e sp e c im e n s from W can be fou n d in V irtu a l H erbaria (h ttp ://h erb a r iu m .u n iv ie .a c .a t/d a ta b a se /sea rch .p h p ).T axon database number voucher origin annotationG. aragatsi ssp. steineri 1 FN582290 W 2009-0018501 ArmeniaG. aragatsi ssp. steineri 2 FN582289 W 2009-0018501 ArmeniaG. aragatsi FN582288 W 2009-0018514 ArmeniaG. aragatsi FN582283 W 2009-0018510 ArmeniaG. sp. FN582287 W 2009-0018774 ArmeniaG. toumefortii FN582281 W 1976-0007273 TurkeyG. toumefortii FN582284 W 2001-0011207 IranG. toumefortii FN582277 W 1965-0015396 IranG. toumefortii FN582285 W 2001-0010125 IranG. toumefortii FN582278 W 1965-0017153 IranG. toumefortii FN582276 W 1994-0005375 Afghanistan var. armataG. toumefortii AY504691G. toumefortii FN582282 W 1958-0000263 IranG. toumefortii FN582279 W 1976-0001491 AfghanistanG. toumefortii FN582280 W 1975-0005807 AzerbaijanG. toumefortii FN582286 B TurkeyScorzonera purpurea AJ633477 GAT Poland cultivatedScorzonera crispatula AJ633486 GAT France cultivatedTragopogon dubius AJ633503.1 GAT Germany cultivatedScorzonera villosa AJ633482 GAT Germany cultivatedScolymus maculatus AJ633469 GAT Netherlands cultivatedCatananche caerulea AJ633465 GAT Germany cultivatedCatananche caerulea AJ633467 GAT France cultivatedCatananche caerulea AJ633466 GAT Netherland cultivatedCatananche arenaria FN582275 B MoroccoCatananche caespitosa AJ633468 GAT Germany cultivatedWarionia saharae A Y 190608

A p p e n d ix 2: M ateria l o f Gundelia sp . e sp e c ia l ly u sed for co m p arison :Armenia, Kotayk province, E o f Yerevan, road from Geghadir to Gami, 1555 m S .m ., 40°8'23"N/ 44°39'45"E, 2007-06-18 K. Tamanyan, M. Oganesian, E. Vitek 07-0738 [ERE, W 2009-18499-2009- 18500, B]; - (fruits) Armenia, Kotayk province, E o f Yerevan, road from Atsavan to Garni, 1533 m s.m., 40o08'25"N/44°39'44"E, 2007-07-11 S. Ter-Pogoshyan [ERE, W 2009-18774, B, NY],