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PHYTON
ANNALES REI BOTANICAE
VOL. 49, FASC. 2 PAG. 145±320 29. 3. 2010
New Classification of
Allium
L. subg.
Melanocrommyum
(WEBB &BERTHEL.) ROUY (
Alliaceae
) Based on Molecular
and Morphological Characters
By
Reinhard M. FRITSCH*), Frank R. BLATTNER*), and Maia GURUSHIDZE*)
With 7 Figures
Received September 30, 2009
Key words: Allium subgenus Melanocrommyum, Alliaceae. ± Classification,
distribution, molecular marker systems, morphology, phylogenetic relationships,
taxonomy.
Summary
FRITSCH R. M., BLATTNER F. R . & G URUSHIDZE M. 2010. New classification of Al-
lium L. subg. Melanocrommyum (WEBB &BERTHEL.) ROUY (Alliaceae) based on mo-
lecular and morphological characters. ± Phyton (Horn, Austria) 49(2): 145±220, with
7 figures.
Allium subgenus Melanocrommyum from Eurasia is taxonomically complicated
with different and contradictory taxonomic treatments proposed in the past. A com-
prehensive study, covering nearly all existing taxonomic groups and their entire
geographic distribution, by sequencing the nuclear rDNA internal transcribed spacer
145
*) Dr. Reinhard M. FRITSCH (corresponding author), Dr. Frank R. BLATTNER,Dr.
Maria GURUSHIDZE, Leibnitz-Institut fuÈ r Pflanzengenetik und Kulturpflanzenfor-
schung (IPK), Corrensstraûe 3, 06466 Gatersleben, Germany; e-mail: fritschr@ipk-
gatersleben.de
Phyton (Horn, Austria) Vol. 49 Fasc. 2 145±220 29. 3. 2010
region (ITS) confirmed the monophyly of the subgenus, while most traditionally used
sections were either para- or polyphyletic. Addition of ten more species and 150 new
accessions to that dataset confirmed the earlier results and underlined that multiple
rapid radiations occurred within different monophyletic groups of the subgenus.
About 40 well separated molecular clades were recognized, and taxonomically dis-
cussed and affiliated, but their phylogenetic relations remained often unresolved.
The splits of some large ªclassicalº sections was unavoidable. Therefore the sections
Asteroprason R. M. FRITSCH and Procerallium R. M. FRITSCH as well as the subsec-
tions Humilicognata R. M. Fritsch, Diffusoumbellata R. M. FRITSCH,Keratoprason
R. M. FRITSCH and Pharmakoprason R. M. FRITSCH were newly described, the sub-
sections Decipientia (OMELCZUK)R.M.FRITSCH and Stellata (F. O. KHASS.&
R. M. FRITSCH)R.M.FRITSCH were raised to sectional level, four subspecies were
raised to species level, and the name Allium jaubertii R. M. FRITSCH was created for
an illegitimate species name.
Altogether 160 species and subspecies were accepted (plus 53 unclear or synon-
ymous names) in the updated conspectus of A. subg. Melanocrommyum. They were
affiliated to 20 sections and 22 subsections (sect. Acanthoprason WENDELBO and sect.
Melanocrommyum WEBB &BERTHEL. were only subdivided in alliances). Lectotypes
of 17 species were designated or validated, and neotypes for A. isfairamicum O.
FEDTSCH., A. suworowii REGEL,A. viridiflorum POBED., and epitypes for A. bakhtiar-
icum REGEL and A. derderianum REGEL were designated.
Zusammenfassung
FRITSCH R. M., BLATTNER F. R . & G URUSHIDZE M. 2010. New classification of Al-
lium L. subg. Melanocrommyum (WEBB &BERTHEL.) ROUY (Alliaceae) based on mo-
lecular and morphological characters. [Neue Klassifikation von Allium L. subg.
Melanocrommyum (WEBB &BERTHEL.) ROUY (Alliaceae) basierend auf molekularen
und morphologischen Merkmalen]. ± Phyton (Horn, Austria) 49(2): 145±220, mit 7
Abbildungen.
Diese eurasisch verbreitete Untergattung ist taxonomisch schwierig. Bisherige
Gliederungen sind sehr unterschiedlich und widersprechen einander. Eine bereits
publizierte, umfassende Untersuchung fast aller beschriebenen taxonomischen
Gruppen aus dem gesamten Verbreitungsgebiet analysierte die Sequenzen der ITS-
Region (internal transcribed spacer Region der nuklearen rDNA). Hierbei wurde die
Monophylie des Subgenus bestaÈ tigt, aber die meisten traditionell benutzten Sektio-
nen waren entweder para- oder polyphyletisch. Eine neue Analyse mit erweitertem
Material (10 Arten und 150 Akzessionen zusaÈ tzlich) bestaÈ tigte die genannten Re-
sultate, insbesondere dass in verschiedenen monophyletischen Gruppen mehrfache
schnelle Radiationen auftraten. Schlieûlich wurden ungefaÈ hr 40 gut getrennte, mo-
lekulare Gruppen erkannt, diskutiert und taxonomisch eingeordnet. Ihr phylogen-
etischer Anschluss wurde aber oft nicht aufgeloÈst. Die Auftrennung von einigen
umfangreichen, ¹klassischen`` Sektionen war unumgaÈnglich, weshalb die Sektionen
Asteroprason R. M. FRITSCH und Procerallium R. M. FRITSCH sowie die Subsektionen
Humilicognata R. M. FRITSCH,Diffusoumbellata R. M. FRITSCH,Keratoprason R. M.
FRITSCH und Pharmakoprason R. M. FRITSCH neu beschrieben, und die Subsektionen
Decipientia (Omelczuk) R. M. FRITSCH und Stellata (F. O. KHASS. & R. M. FRITSCH)
R. M. FRITSCH als Sektionen neu kombiniert wurden. In diesem Zusammenhang
146
wurden vier Unterarten zu Arten erhoben, und Allium jaubertii R. M. FRITSCH an-
stelle eines illegitimen Namens geschaffen.
Im neu gefassten Konspekt von A. subg. Melanocrommyum werden 160 Arten
und Unterarten akzeptiert, weitere 53 als unklare Namen oder Synonyme. Sie wer-
den 20 Sektionen und 22 Untersektionen zugeordnet, wobei die Sektionen Acantho-
prason WENDELBO und Melanocrommyum WEBB &BERTHEL. nur in informelle Grup-
pen gegliedert sind. In diesem Zusammenhang wurden Lektotypen von 32 Arten
bestimmt bzw. validiert, Neotypen fuÈr A. isfairamicum O. FEDTSCH., A. suworowii
REGEL und A. viridiflorum POBED., sowie Epitypen fuÈr A. bakhtiaricum REGEL und
A. derderianum REGEL bestimmt.
Content
1. Introduction ..................................................................................... 147
2. Materials and methods ....................................................................... 151
2.1. Molecular investigations .............................................................. 151
2.2. Accessions (in numerical sequence) ................................................ 151
3. Results and discussion of molecular and morphological data...................... 167
3.1. Outgroups ................................................................................. 167
3.2. Basal grade ............................................................................... 167
3.3. Core clade, cluster 1.................................................................... 168
3.4. Core clade, cluster 2.................................................................... 192
3.5. Core clade, cluster 3.................................................................... 185
3.6. Core clade, cluster 4.................................................................... 187
3.7. Core clade, cluster 5.................................................................... 188
3.8. Core clade, cluster 6.................................................................... 190
3.9. Core clade, cluster 7.................................................................... 192
3.10. sect. Regeloprason ...................................................................... 200
3.11. sect. Thaumasioprason, not included in this study ............................ 201
3.12. Ornamental cultivars and hybrids.................................................. 201
4. Typifications .................................................................................... 202
5. Conspectus speciorum generis Allium L. subgeneris Melanocrommyum
(WEBB &BERTHEL.) ROUY ..................................................................... 209
5.1. Conspectus................................................................................ 209
5.2 Index of specific and infraspecific names applied in the ªConspectusº.. 214
6. Acknowledgements ............................................................................ 216
7. References........................................................................................ 217
1. Introduction
Allium is a very variable and taxonomically difficult genus naturally
distributed over the northern hemisphere. The most recent classification
proposal for this genus, based on morphological characters and consider-
ing also molecular data, accepted about 780 species belonging to fifteen
subgenera and 56 sections (FRIESEN & al. 2006). Taxonomic inventorying of
this genus focused recently mainly on the Mediterranean area, Southwest
and parts of Central Asia and resulted in the descriptions of more than 30
new species and subspecies raising this number to more than 800 species in
2009.
147
148
Melanocrommyum is the second largest Allium subgenus comprising
about 140 accepted species and subspecies in 2005 (FRIESEN & al. 2006).
Very recently about 15 taxa were newly described (DENIZ &SUMBUL 2004;
SEREGIN 2007; FRITSCH & al. 2007; FRITSCH &ABBASI 2009) and several hi-
therto undescribed ones are currently under investigation. Thus, at least
160 taxa belong to A. subg. Melanocrommyum. Characteristics for mem-
bers of this subgenus are true tunicated bulbs, annual roots, mostly broad
and flat leaves with subterranean (scarcely visible above earth) sheath
parts, strong and most often strictly upright scapes of varying length, and
large, fasciculate to globular inflorescences composed of many moderately
small to large, and often star-like, flowers. The flowers of several species
emit a sweet or in another way noticeable odor.
Despite many members of subg. Melanocrommyum do not own the
specific taste and smell of garlic and common onion, several species are
intensely used by the native population of Central Asia and Iran as vege-
tables and medicinal plants (KEUSGEN & al. 2006, ABBASI & al. 2008). In
general they contain less cysteine sulphoxides, the main pharmacologically
active substances, than garlic and common onion (KEUSGEN & al. 2008).
Surprisingly, species possessing remarkable antibiotic activity and partly a
much higher radical scavanger activity than garlic contain newly dis-
covered dithiodipyrroles (JEDELSKA
Â& al. 2008) and sulphur-pyridins
(KUSTERER &KEUSGEN 2009). This substances are also present in many well
known ornamental species (FRITSCH & al. 2008).
Former Allium classifications, used up to the 1950
th
, commonly in-
cluded the members of subg. Melanocrommyum in the bulbous sect. Mo-
lium G. DON EX KOCH because of certain morphological similarities. Spe-
cific features and structures of bulb and inflorescence development pre-
sent solely in Mediterranean members of sect. Molium bothered WENDELBO
1969 to accept this section in the strict sense at subgeneric level, and to
separate the remaining species as subgenus Melanocrommyum which he
even nominated as the ªthe most advanced of the subgeneraº without
presenting any argument. Additional studies detected more principal dif-
ferences between these subgenera which concern chromosome base num-
bers, the principles of leaf, scape, and flower anatomy, the course of annual
life cycle as perhaps the most important ones. Currently sect. Molium in
the strict sense is affiliated to subg. Amerallium TRAUB which has nomen-
clatural priority.
Earlier the long-rhizomatous growth and presence of slender, many-
scaled bulbs were postulated as ancient evolutionary level in the genus
Allium. Then the absence of rhizomes and presence of few-scaled, true
bulbs characterizing subg. Melanocrommyum as well as subg. Allium
should be the advanced level (HANELT & al. 1992). However, different mo-
lecular studies presented evidence that Allium species belong to three
evolutionary lineages, and rhizomatous as well as bulbous groups are pre-
149
sent in all of them (FRITSCH &FRIESEN 2002, FRIESEN & al. 2006). The basal
split in the phylogenetic dendrograms separates the bulbous subgenera
Microscordum (MAXIM.) N. FRIESEN, Nectaroscordum (LINDL.) ASCHERS.&
GRAEBN., and Amerallium (the latter comprising also rhizomatous sections)
from all other taxa of the genus. Subg. Melanocrommyum is the largest
group of a second evolutionary lineage jointly with the mono- or oligo-
typic, bulbous subgenera Caloscordum (HERB.) R. M. FRITSCH,Porphyr-
oprason (EKBERG)R.M.FRITSCH,Vvedenskya (KAMELIN)R.M.FRITSCH, and
the small rhizomatous subg. Anguinum (G. DON ex KOCH)N.FRIESEN. The
rhizomatous subgenera Butomissa (SALISB.) N. FRIESEN,Rhizirideum (G.
DON ex KOCH)WENDELBO, Polyprason RADIC
Â, Reticulatobulbosa (KAMELIN)
N. FRIESEN, and Cepa RADIC
Âas well as the bulbous subgenus Allium belong
together to the third evolutionary lineage (FRIESEN & al. 2006). Thus subg.
Melanocrommyum is neither a ªmost advancedº group nor is it closer re-
lated to the bulbous sections of subg. Amerallium and to subg. Allium.
A first subdivision of subgenus Melanocrommyum was already pro-
posed by Wendelbo 1966 when he described sect. Regeloprason WENDELBO.
Later (WENDELBO 1969) he accepted also the sections Melanocrommyum
WEBB &BERTHEL., Kaloprason K. KOCH, Acanthoprason WENDELBO, Mega-
loprason WENDELBO,and Thaumasioprason WENDELBO in a paper focusing
on Mediterranean and Southwest Asian species only. The many related
species occurring in Central Asia were later transferred to subg. Melano-
crommyum when KAMELIN 1973 proposed another classification. This au-
thor did not accept sect. Megaloprason and treated Kaloprason, Acantho-
prason, and Regeloprason at subsectional level only, described sect. Verti-
cillata KAMELIN and sect. Vvedenskya KAMELIN as new sections, and
transferred also the sections Porphyroprason,Microscordum, and Briseis
(SALISB.) STEARN to subg. Melanocrommyum. Later the Gatersleben Allium
working group (HANELT & al. 1992) presented a broad array of morpholo-
gical, anatomical, karyological, and chemical arguments to exclude again
the sections Microscordum,Vvedenskya, and Briseis, to add the new sec-
tions Acmopetala R. M. FRITSCH,Compactoprason R. M. FRITSCH (both
segregates of sect. Megaloprason sensu lato), and Miniprason R. M.
FRITSCH, and to transfer sect. Pseudoprason WENDELBO to subg. Melano-
crommyum (FRITSCH 1992). Finally the sections Acaule R. M. FRITSCH,Ar-
oidea F. O. KHASS. & R. M. FRITSCH, Brevicaule R. M. FRITSCH,and Popovia
F. O. KHASS. & R. M. FRITSCH were newly created by KHASSANOV &FRITSCH
1994 separating some morphologically rather isolated species of subg.
Melanocrommyum. A classification proposal of SEISUMS 1994 introduced
also a new section Triptera and four new subsections in subg. Melano-
crommyum, but these groups were never validly described.
Beginning in the 1990ies, the application of different molecular mar-
ker systems confirmed subg. Melanocrommyum as clade well separated
from other Allium subgenera and to consist of several subgroups. However,
different analyses displayed incongruent and often contradictory relations
of the rather few taxa studied. Some results could best be explained by
reticulate evolution (MES & al. 1997, 1999). Therefore detailed investiga-
tions analyzing sequences of the nuclear rDNA internal transcribed spacer
region (ITS) and the plastid DNA trnL-trnF region of about 430 accessions
representing more than 100 species were undertaken (GURUSHIDZE & al.
2008, GURUSHIDZE & al. in press). These results confirmed the monophyly
of A. subg. Melanocrommyum and indicated multiple radiations within the
group, but could not detect evidence for frequent introgressions. The in-
vestigated species form a basal grade and a core clade in the phylogenetic
tree. 7 different clusters within the core clade occur as monophyletic units
with all applied (phenetic, cladistic, and model-based) algorithms. The
relationships among these clusters were not resolved, but the clusters were
mostly well-supported in all analyses. Also some groups within these
cluster got statistical support enabling characterization of their phyloge-
netic positions. Altogether two groups were recognized in the grade, and 10
more groups in the clade. Five of these groups were additionally char-
acterized by different specific anatomical features of the septal nectaries
while the remaining groups shared another (unspecific) type of nectary.
Other (often small) groups were well separated with high support values
but their positions remained unresolved in this phylogenetic tree. The
phylogenetic analyses showed, as a main result, that most of the tradi-
tional taxonomic sections in subg. Melanocrommyum are either para- or
polyphyletic, and favor the circumscription of smaller sections. The oc-
currence of cryptic species in subg. Melanocrommyum was documented for
the first time in Allium.
The principal structure of this phylogenetic tree was confirmed by se-
quence analyses of the trnL-trnF region of chloroplast DNA (GURUSHIDZE
& al. in press). A statistical parsimony network of 80 chloroplast haplo-
types resulted in six main lineages which showed much congruence with
subunits of the ITS-based phylogenetic tree, and rather few species posi-
tions were more or less incongruent with the ITS data. Also the number of
inferred (missing in the data set) haplotypes was rather low.
Newly collected material enabled incorporation of more accessions
and more species to the material analyzed by GURUSHIDZE & al. 2008 in
order to prove splitting of traditional sections and especially the position
of small segregate groups and their statistical support. Below we will pre-
sent the final results of our investigation of subg. Melanocrommyum, in-
cluding detailed discussion of taxonomic changes only shortly mentioned
by GURUSHIDZE & al. 2008, and their nomenclatorial validations.
Most important is the conclusion, that taxa traditionally affiliated to
one section are not necessarily closely related but may belong to geneti-
cally only distantly related subgroups.
150
2. Materials and Methods
2.1. Molecular Investigations
For the present investigation we included ITS sequences of 578 accessions (acc.)
representing about 112 Melanocrommyum species and subspecies, 5 hybrid orna-
mental cultivars, and 8 outgroup species, from which 395 sequences were taken from
our previous publication, while the remaining 183 acc. were sequenced for this study.
All sequences were submitted to the EMBL nucleotide database. For detailed de-
scription of DNA extraction, PCR, and sequencing methods see GURUSHIDZE & al.
2008.
The data were analyzed using distance (Neighbor-Joining) and Bayesian algo-
rithms. For the present study we have not performed Maximum Parsimony (MP)
analyses due to following reasons. First, we showed that the Bayesian and MP results
were highly compatible for analyzing relationships among ITS sequences within
subgenus Melanocrommyum (GURUSHIDZE & al. 2008), and second, for such a large
data-matrix MP would be computationally extremely demanding. The model of se-
quence evolution was tested in MODELTEST 3.7 (POSADA &CRANDALL 1998), GTR+G
model was chosen as the best fitted model by the Akaike information criterion. This
model was used for distance calculations in Neighbor-Joining (NJ) analyses with
PAUP* (SWOFFORD 2002). Bayesian analyses (BI) was performed using MrBAYES 3.1
(RONQUIST &HUELSENBECK 2003). Eight chains were run for 10 million generations
under the chosen model of sequence evolution, sampling a tree every 1000 genera-
tions. The posterior probabilities were calculated after discarding the initial 25%
(non-stationary) of the resulting trees. The Bayesian tree with branch lengths and
posterior probabilities is shown schematically on Figure 1 and in detail on the Fig-
ures 2 A - 2 F.
2.2. Accessions (in Numerical Sequence)
The scheme of the list is as follows: a c c e s s i o n n u m b e r: species AUTHOR(s);
taxonomic affiliation (subgenus or section); origin (source, coordinates or collection
site; herbarium and voucher no. or photo)
0348: Allium oreophilum C. A. MEY.; subg. Porphyroprason; Graz Botanic Gar-
den Univ., Austria; (GAT). ± 0465: Allium macleanii BAKER;Compactoprason; Bota-
nical Garden Vrije Univ. Amsterdam, Netherlands; (GAT). ± 0515: Allium multi-
bulbosum JACQ.; Melanocrommyum; Leipzig Botanical Garden, Germany; (GAT). ±
0531: Allium stipitatum REGEL;Megaloprason; Dushanbe Botanical Garden, Tajiki-
stan; (GAT). ± 0612: Allium stipitatum REGEL;Megaloprason; Budapest Botanical
Garden, Hungary; (GAT). ± 0616: Allium backhousianum REGEL;Acmopetala; Bota-
nical Garden Univ. Budapest, Hungary; (GAT). ± 0779: Allium karataviense REGEL;
Miniprason; Jena Botanical Garden, Germany; (GAT). ± 0799: Allium atropurpureum
WALDST.&KIT.; Melanocrommyum; Jena Botanical Garden, Germany; (GAT). ± 0975:
Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan; Hissar mountain range,
Varzob valley; (GAT). ± 1017: Allium atropurpureum WALDST.&KIT.; Melano-
crommyum; private collection Dr. WANDELT, Quedlinburg; (GAT). ± 1033: Allium jes-
dianum BOISS.&BUHSE subsp. angustitepalum (WENDELBO)F.O.KHASS. & R. M.
FRITSCH;Megaloprason; Afghanistan, received from GoÈteborg Botanical Garden;
(GAT). ± 1044: Allium stipitatum REGEL;Megaloprason; Afghanistan, received from
GoÈteborg Botanicalal Garden; (GAT). ± 1082: Allium jesdianum BOISS.&BUHSE
151
subsp. angustitepalum (WENDELBO)F.O.KHASS. & R. M. FRITSCH;Megaloprason; Af-
ghanistan, received from GoÈ teborg Botanical Garden; (GAT). ± 1083: Allium jesdia-
num BOISS.&BUHSE subsp. angustitepalum (WENDELBO)F.O.KHASS. & R. M.
FRITSCH;Megaloprason; Afghanistan, received from GoÈ teborg Botanical Garden;
(GAT). ± 1090I: Allium stipitatum REGEL;Megaloprason; Iran; 344343N, 0483637E;
(IRAN 43978). ± 1119I: Allium jesdianum BOISS.&BUHSE subsp. jesdianum;Mega-
loprason; Iran; 360337N, 0465148E; (IRAN 43974). ± 1122I: Allium koelzii (WEN-
DELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran; 350050N, 0465421E; (IRAN). ±
1123I: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran; near
Sanandaj; (IRAN). ± 112G: Allium pseudowinklerianum R. M. FRITSCH &F.O.
KHASS.; Regeloprason; Kyrgyzstan; Fergan mountain range (isotype); (GAT). ± 1138I:
Allium akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason; Iran; prov.
Zanjan; (IRAN). ± 113G: Allium sochense R. M. FRITSCH &U.TURAKULOV;Re-
geloprason; Kyrgyzstan; Alai mountain range (isotype); (GAT). ± 114G: Allium der-
derianum REGEL;Acanthoprason; Iran; prov. Tehran, Karaj valley; (GAT). ± 1150:
Allium cyrilli TEN.; Melanocrommyum; Leipzig Botanical Garden; (GAT). ± 1164I:
Allium materculae BORDZ.subsp. graveolens R. M. FRITSCH;Acanthoprason; Iran;
335900N, 0500700E; (IRAN). ± 116G: Allium tulipifolium LEDEB.;Melanocrommyum;
Kazakhstan; steppe W Zelinograd; (GAT). ± 1178: Allium stipitatum REGEL;Mega-
loprason; Botanical Garden Univ. Strasbourg, France; (GAT). ± 117G: Allium aus-
troiranicum R. M. FRITSCH;Acanthoprason; Iran; 3222N, 05028E; (GAT). ± 11929:
Allium sp.2; Acanthoprason; Iran; prov. Kurdestan, Zanjan to Bijar, pass S Khur-
Khureh; (TARI 11929). ± 119G: Allium hexaceras VVED.; Acaule; Uzbekistan; Hissar
mountain range; (GAT). ± 1222: Allium jesdianum BOISS.&BUHSE subsp. angustite-
palum (WENDELBO)F.O.KHASS. & R. M. FRITSCH;Megaloprason; Moscow Main Bo-
tanical Garden; (GAT). ± 1224I: Allium stipitatum REGEL s. lat.; Megaloprason; Iran;
325523N, 0503330E; (IRAN). ± 1241I: Allium sp.;Acanthoprason; Iran; 302349N,
0515431E; (IRAN). ± 1300: Allium victorialis L.; subg. Anguinum; Russia; Altai;
(GAT). ± 1311: Allium stipitatum REGEL;Megaloprason; Tajikistan; 3849N, 06848E
(near type location); (GAT). ± 1315: Allium stipitatum REGEL;Megaloprason; Dush-
anbe Botanical Garden, Tajikistan; (GAT). ± 1323: Allium sarawschanicum REGEL;
Megaloprason; Tajikistan; 3849N, 06848E; (GAT). ± 1326: Allium sarawschanicum
REGEL;Megaloprason; Tajikistan; 3847N, 06850E; (GAT). ± 1327: Allium rosenorum
R. M. FRITSCH;Megaloprason; Tajikistan; 3858N, 06844E; (GAT). ± 1343: Allium sti-
pitatum REGEL;Megaloprason; Tajikistan; 3841N, 06852E; (GAT). ± 1345: Allium ro-
senorum R. M. FRITSCH;Megaloprason; Tajikistan; N Dushanbe; (GAT). ± 1384: Al-
lium lipskyanum VVED.; Regeloprason; Dushanbe Botanical Garden, Tajikistan;
(GAT). ± 1388: Allium cristophii TRAUTV.; Kaloprason; Botanic Garden Univ. Buda-
pest, Hungary; (GAT). ± 13H: Allium aff. isakulii R. M. FRITSCH &F.O.KHASS. subsp.
isakulii;Regeloprason; Uzbekistan; Fergan depression; (GAT). ± 14164: Allium der-
derianum REGEL;Acanthoprason; Iran; Taleqan; (TARI 14164). ± 14176: Allium ma-
terculae BORDZ.subsp. graveolens R. M. FRITSCH;Acanthoprason; Iran; prov. Tehran,
68 km from Tehran to Qom; (TARI 14176). ± 1502: Allium backhousianum REGEL;
Acmopetala; Khorog, Pamir Botanical Garden, Tajikistan; (GAT). ± 1623: Allium sti-
pitatum REGEL;Megaloprason; Tajikistan; Nurek; (GAT). ± 1625: Allium darwasicum
REGEL;Regeloprason; Tajikistan; Hissar mountain range; (GAT). ± 1626: Allium sar-
awschanicum REGEL;Megaloprason; Tajikistan; Javros, near Dushanbe; (GAT). ±
1631: Allium hollandicum R. M. FRITSCH;Megaloprason; private collection Dr.
152
FRITSCH, Gatersleben (type population); (GAT). ± 1650: Allium multibulbosum Jacq.;
Melanocrommyum; Leipzig Botanical Garden; (GAT). ± 16611: Allium nigrum L.;
Melanocrommyum; Greece, Crete Island; 351403N, 0240555E; (OSBU 16611). ±
16666: Allium nigrum L.; Melanocrommyum; Greece, Crete Island; 351741N,
0242524E; (OSBU 16666). ± 16744: Allium bisotunense R. M. FRITSCH;Melano-
crommyum; Iran; prov. Kermanshah, Kuh-e Parrou; (TARI 16744). ± 16799: Allium
aff. austroiranicum R. M. FRITSCH;Acanthoprason; Iran; prov. Kermanshah, Kuh-e
Bozab, macrowave station; (TARI 16799). ± 16833: Allium breviscapum STAPF;Acan-
thoprason; Iran; prov. Hamedan, Alvan Kuh, near Gandjnameh; (TARI 16833). ±
16867: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran; prov. Kurdestan,
Ghorveh towards Sanandaj; (TUH 16867). ± 16905: Allium saralicum R. M. FRITSCH;
Melanocrommyum; Iran; prov. Kurdestan, Dasht-e Zaghe on road from Hamadan to
Sanandaj c. 40 km E Sanandaj; (TARI 16905). ± 17362: Allium aff. ubipetrense R. M.
FRITSCH;Acanthoprason; Iran; prov. Azarbeijan, Tabriz to Ghar-e Chaman, before
this place; (TARI 17362). ± 1801: Allium hollandicum R. M. FRITSCH `Purple Sensa-
tion'; Megaloprason; Gesellschaft der Staudenfreunde, Germany; (GAT). ± 1869: Al-
lium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan; 3845N, 06919E (type loca-
tion); (GAT). ± 1880: Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan;
3847N, 06849E; (GAT). ± 1886: Allium rosenorum R. M. FRITSCH;Megaloprason;Ta-
jikistan; 3847N, 06848E; (GAT). ± 1894: Allium rosenorum R. M. FRITSCH;Mega-
loprason; Tajikistan; 3850N, 06849E; (GAT). ± 1897: Allium stipitatum REGEL;Mega-
loprason; Tajikistan; 3849N, 06846E; (GAT). ± 1900: Allium rosenorum R. M. FRITSCH;
Megaloprason; Tajikistan; 3849N, 06846E; (GAT). ± 1909: Allium sarawschanicum
REGEL;Megaloprason; Tajikistan; 3842N, 06853E; (GAT). ± 1911: Allium macleanii
BAKER;Compactoprason; Tajikistan, vegetable market in Dushanbe; (GAT). ± 1920:
Allium cristophii TRAUTV.;Kaloprason; Manchester Botanic Garden; (GAT). ± 19593:
Allium aff. akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason; Iran; prov.
Azarbeijan, W Rezaiyeh, hills W Silvana village; (TARI 19593). ± 19746: Allium sp.10;
Acanthoprason; Iran; (location not available); (TARI 19746). ± 19854: Allium aff.
hollandicum R. M. FRITSCH;Megaloprason; Iran; prov. Azarbeijan, SW of Rezaiyeh,
Silvana valley; (TARI 19854). ± 19932: Allium sp.2; Acanthoprason; Iran; prov.
Azarbeijan, 10 km on the road from Mianeh to Zanjan; (TARI 19932). ± 1I: Allium
robustum KAR.&KIR.; Melanocrommyum; Kazakhstan; 4719N, 08133E; (GAT). ± 1o:
Allium orientale BOISS. s. lat.; Melanocrommyum; Israel; Lehavim; (HUJ). ± 2: Allium
orientale BOISS. s. lat.; Melanocrommyum; Israel; Tel Krayot; (HUJ). ± 20228: Allium
aff. egorovae M. V. AGAB.&OGAN.; Acanthoprason; Iran; prov. Azarbeijan, Arasbaran
protected region; (TARI 20228). ± 2121: Allium stipitatum REGEL;Megaloprason;
Marburg Botanical Garden, Germany; (GAT). ± 2155H: Allium aff. hollandicum R. M.
FRITSCH;Megaloprason; Iran; prov. Azarbeijan, inter Rezaiyeh & Oshnaviyeh; (TARI
2155). ± 2162: Allium protensum WENDELBO;Kaloprason; Rostock Botanical Garden,
Germany; (GAT). ± 2162S: Allium noeÈanum REUT.exREGEL;Acanthoprason; Iran;
prov. Azarbeijan, in jugo Khan inter Baneh & Saqqez; (TARI 2162). ± 2164H: Allium
aff. hollandicum R. M. FRITSCH;Megaloprason; Iran; prov. Azarbeijan, Dasht-e Bel;
(TARI 2164). ± 2182: Allium verticillatum REGEL;Verticillata; Tajikistan; 3759N,
06834E; (GAT, photos). ± 2194: Allium atropurpureum WALDST.&KIT.; Melano-
crommyum; Bulgaria; Cape Emine; (GAT). ± 2239: Allium stipitatum REGEL;Mega-
loprason;VaÂcraÂtoÂt Botanical Garden, Hungary; (GAT). ± 2256: Allium rosenorum R.
M. FRITSCH;Megaloprason; Tajikistan; 3845N, 06918E (type location); (GAT). ± 2257:
153
Allium stipitatum REGEL;Megaloprason; Tajikistan; 3845N, 06918E; (GAT). ± 2262:
Allium nevskianum VVED.exWENDELBO;Kaloprason; Tajikistan; 3849N, 06850E;
(GAT). ± 2264: Allium stipitatum REGEL;Megaloprason; Tajikistan; 3903N, 06845E;
(GAT). ± 2266: Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan; 3902N,
06847E; (GAT). ± 2269: Allium nevskianum VVED.exWENDELBO;Kaloprason; Tajiki-
stan; 3842N, 06853E; (GAT). ± 2270: Allium rosenorum R. M. FRITSCH;Megaloprason;
Tajikistan; 384201N, 0685311E; (GAT). ± 2271: Allium rosenorum R. M. FRITSCH;
Megaloprason; Tajikistan; Kharangon valley N Dushanbe; (GAT). ± 2276: Allium
sarawschanicum REGEL;Megaloprason; Tajikistan; 3759N, 06834E; (GAT). ± 2379:
Allium neriniflorum (HERB.) BAKER; subg. Caloscordum; Mongolia; SE Sumber, So-
mon Chalchgol; (GAT). ± 2399: Allium nevskianum VVED.exWENDELBO;Kaloprason;
Tajikistan, received from Botanical Garden Tallin; (GAT). ± 2413: Allium macleanii
BAKER;Compactoprason; Chorog Botanical Garden, Tajikistan; (GAT). ± 2415: Allium
macleanii BAKER;Compactoprason; Chorog Botanical Garden, Tajikistan; (GAT). ±
2432: Allium sp.10; Acanthoprason; Iran; prov. Kurdestan, 15 km NE Banah; (TARI
2432). ± 2445: Allium stipitatum REGEL;Megaloprason; Cluj-Napoca Botanical Gar-
den, Romania; (GAT). ± 24865: Allium aff. egorovae M. V. AGAB.&OGAN.; Acantho-
prason; Iran; prov. Azarbeijan, Arasbaran protected area, between Kharil and Ma-
kiki; (TARI 24865). ± 2497H: Allium scotostemon WENDELBO;Thaumasioprason; Iran;
(data are missing); (TUH 2497). ± 2517: Allium aroides POPOV &VVED.; Aroidea; Uz-
bekistan, Tashkent Botanical Garden; (GAT). ± 2517A: Allium ellisii HOOK. f.; Kalo-
prason; Iran; (data missing); (TUH 2517). ± 2521: Allium severtzovioides R. M.
FRITSCH;Acmopetala; Uzbekistan; 4139N, 06947E; (GAT). ± 25222: Allium sp.3;
Acanthoprason; Iran; prov. Azarbeijan, Rezaiyeh, near Turkish border; (TARI 25222).
± 2530: Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan; 3905N, 06923E;
(GAT). ± 2537: Allium darwasicum REGEL;Regeloprason; Tajikistan; 3821N, 07003E;
(GAT). ± 2541: Allium rosenbachianum REGEL subsp. kwakense R. M. FRITSCH;
Megaloprason; Tajikistan; 3827N, 07011E; (GAT). ± 2549: Allium stipitatum REGEL;
Megaloprason; Tajikistan; 3830N, 07003E; (GAT). ± 2552: Allium rosenorum R. M.
FRITSCH;Megaloprason; Tajikistan; 3903N, 06854E; (GAT). ± 2557: Allium alex-
eianum REGEL;Kaloprason; Tajikistan; 3932N, 06838E; (GAT). ± 2566: Allium rose-
norum R. M. FRITSCH;Megaloprason; Tajikistan; 3912N, 06733E; (GAT). ± 2582: Al-
lium tricoccum SOL.; subg. Anguinum; USA; Whistler's Woods, Southern Cook
county, Illinois; (GAT, photos). ± 2615: Allium hollandicum R. M. FRITSCH; `Purple
Sensation'; Megaloprason; Botanic Garden Vrije Univ. Amsterdam; (GAT). ± 2616:
Allium multibulbosum JACQ.; Melanocrommyum; Botanical Garden Vrije Univ. Am-
sterdam; (GAT). ± 2618: Allium stipitatum REGEL;Megaloprason; Botanical Garden
Vrije Univ. Amsterdam, Netherlands; (GAT). ± 2657: Allium stipitatum REGEL;
Megaloprason; Dresden Botanical Garden, Germany; (GAT). ± 2673: Allium victor-
ialis L.; subg. Anguinum; Georgia; Caukasus, Teberda, Dombai; (GAT, photos). ±
2680: Allium backhousianum REGEL;Acmopetala; Kyrgyzstan; Alai mountain range;
(GAT). ± 2681: Allium backhousianum REGEL;Acmopetala; Kyrgyzstan; Alai moun-
tain range; (GAT). ± 2709: Allium decipiens FISCH.exSCHULT.&SCHULT. f. subsp.
quercetorum SEREGIN;Melanocrommyum; Ukraina; Crimea, Mt. Ayu-Dag; (GAT,
photos). ± 2735: Allium ramosum L.; subg. Butomissa; Alma-Ata Botanical Garden,
Kazakhstan; (GAT). ± 27559: Allium derderianum REGEL;Acanthoprason; Iran; prov.
Tehran, between Tehran and Karadj, above Kalak village; (TARI 27559). ± 27810:
Allium akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason; Iran; prov.
154
Azarbeijan, Assalem to Khalkhal after pass near Khalkhal; (TARI 27810). ± 2793:
Allium motor KAMELIN;Acmopetala; Uzbekistan, received from Dr. LEVICHEV, Lenin-
grad, Russia; (GAT). ± 2794: Allium stipitatum REGEL;Megaloprason; Uzbekistan;
village Samsarak near Tashkent; (GAT). ± 27959: Allium akaka S. G. GMELIN ex
SCHULT.&SCHULT. f.; Acanthoprason; Iran; prov. Azarbeijan, pass c. 20 km S Ahar on
road to Tabriz; (TARI 27959). ± 2797: Allium neriniflorum (HERB.) BAKER; subg. Ca-
loscordum; Novosibirsk Centr. Siber. Botanic Garden, Russia; (GAT). ± 2800: Allium
hollandicum R. M. FRITSCH;Megaloprason; from Bundesgartenschau (German Fed-
eral Garden Show) Frankfurt/Main; (GAT). ± 2846: Allium macleanii BAKER;Com-
pactoprason; Afghanistan, received from GoÈ teborg Botanical Garden; (GAT, photos).
± 2935: Allium hissaricum VVED.; Regeloprason; Tajikistan; 3801N, 06828E; (GAT). ±
2938: Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan; 3804N, 06830E;
(GAT). ± 2946: Allium sarawschanicum REGEL;Megaloprason; Tajikistan; 3841N,
06900E; (GAT). ± 2947: Allium hissaricum VVED.; Regeloprason; Tajikistan; 3841N,
06900E (type location); (GAT). ± 2952: Allium stipitatum REGEL;Megaloprason; Taji-
kistan; 4038N, 06935E; (GAT). ± 29524: Allium pseudobodeanum R. M. FRITSCH &
MATI N;Acanthoprason; Iran; prov. Tehran, Firuzkuh, Gaduk; (TUH 29524). ± 2966:
Allium tulipifolium LEDEB.;Melanocrommyum; Kazakhstan; 4356N, 07708E; (GAT).
± 2975: Allium fetisowii REGEL;Acmopetala; Kazakhstan; 4321N, 07651E (type loca-
tion); (GAT). ± 2976: Allium altissimum REGEL;Megaloprason; Kazakhstan; 4321N,
07500E; (GAT). ± 2977: Allium aflatunense B. FEDTSCH.; Megaloprason; Collection of
Dr. KAMENETZKAJA, Kazakhstan; (GAT). ± 2989: Allium karataviense REGEL;Mini-
prason; Uzbekistan; 4110N, 07008E; (GAT). ± 2992: Allium zergericum F. O. KHASS.&
R. M. FRITSCH;Acmopetala; Uzbekistan; 4055N, 07330E (type location); (GAT). ± 2E:
Allium chrysantherum BOISS.&REUT.; Melanocrommyum; Iran; prov. Zanjan, Bijar;
(IRAN 326/2). ± 3024: Allium schubertii ZUCC.;Kaloprason; private collection Dr.
WIERING, Bergen, Netherland; (GAT). ± 3028: Allium stipitatum REGEL;Mega-
loprason; private collection Dr. WIERING, Bergen Netherland; (GAT). ± 30358: Allium
akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason; Iran; prov. Azarbeijan,
Maku to Khoy, mountains SW Kelisa-Kandi; (TARI 30358). ± 3087: Allium stipitatum
REGEL;Megaloprason; Botanical Garden Univ. Strasbourg, France; (GAT). ± 3115:
Allium stipitatum REGEL;Megaloprason; Tajikistan; 3844N, 06825E; (GAT). ± 3118:
Allium lipskyanum VVED.; Regeloprason; Tajikistan; 3844N, 06828E; (GAT). ± 3121:
Allium suworowii REGEL;Acmopetala; Tajikistan; 3822N, 06942E; (GAT). ± 3123:
Allium trautvetterianum REGEL;Compactoprason; Tajikistan; 3822N, 06942E (near
type location); (GAT, photos). ± 3129: Allium hissaricum VVED.; Regeloprason; Taji-
kistan; 3823N, 06943E; (GAT). ± 3133: Allium komarowii LIPSKY;Compactoprason;
Tajikistan; 3905N, 06821E; (GAT). ± 3144: Allium komarowii LIPSKY;Compactopra-
son; Uzbekistan; 3850N, 06707E; (GAT). ± 3219: Allium jesdianum BOISS.&BUHSE
subsp. angustitepalum (WENDELBO)F.O.KHASS. & R. M. FRITSCH;Megaloprason;
private collection R. DADD, Wokingham, Berkshire, Great Britan; (GAT). ± 323H:
Allium chelotum WENDELBO;Melanocrommyum; Iran; prov. Gorgan, Golestan forest;
(IRAN 323). ± 3246: Allium stipitatum REGEL `Mount Everest'; Megaloprason; Firm of
Peter Nijsen, Heemstede, Netherlands; (GAT). ± 324H: Allium chelotum WENDELBO;
Melanocrommyum; Iran; prov. Gorgan, Golestan forest; (IRAN 324). ± 32H: Allium
aff. hollandicum R. M. FRITSCH;Megaloprason; Iran; prov. Azarbeijan, inter Rezaiyeh
& Oshnaviyeh; (TARI). ± 3335: Allium protensum WENDELBO;Kaloprason; Uzbeki-
stan; Kashkadarya area, near Uradarya; (GAT). ± 3347: Allium stipitatum REGEL;
155
Megaloprason; Kyrgyzstan; 4020N, 07238E; (GAT). ± 3355: Allium motor KAMELIN;
Acmopetala; Uzbekistan; 4117N, 06951E (near type location); (GAT). ± 3356: Allium
motor KAMELIN;Acmopetala; Uzbekistan; 411711N, 0695101E; (GAT). ± 3357: Allium
motor KAMELIN;Acmopetala; Uzbekistan; 4117N, 06951E; (GAT). ± 3362: Allium
majus VVED.; Compactoprason; Uzbekistan; 3920N, 06710E; (GAT). ± 34075: Allium
sp.aff. akaka S. G. GMELIN;Acanthoprason; Iran; (location missing); (TUH 34075). ±
34120: Allium scotostemon WENDELBO;Thaumasioprason; Iran; 3543N, 5212E; (TUH
34120). ± 34268: Allium shelkovnikovii GROSSH.; Acanthoprason; Iran; prov. Azar-
beijan, near Tabriz; (TUH 34268). ± 34269: Allium shelkovnikovii GROSSH.; Acantho-
prason; Iran; prov. Azarbeijan, campus of Tabriz University; (TUH 34269). ± 34349:
Allium sp.9; Acanthoprason; Iran; prov. Qazvin, Alamut Mts.; (TARI 34349). ± 343H:
Allium sp.1ªchlorotepalumº;Acanthoprason; Iran; prov. Esfahan, Esfahan to
Daran; (TARI). ± 34902: Allium haemanthoides BOISS.&REUT.exREGEL s. str.;
Acanthoprason; Iran; prov. Kurdestan, Novsoud 65 km to Marivan; (GAT). ± 34903:
Allium elburzense WENDELBO;Acanthoprason; Iran; prov. Tehran, Abali to Tehran;
(GAT). ± 34908: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran; prov. Qazvin,
Razan 24 km to Avaj; (GAT). ± 34911: Allium elburzense WENDELBO;Acanthoprason;
Iran; prov. Mazandaran, Dizin; (GAT). ± 35079: Allium elburzense WENDELBO;Acan-
thoprason; Iran; prov. Tehran, Damavand towards Tar Lake; (TUH 35079). ± 3625:
Allium kujukense VVED.; subg. Vvedenskya; Kazakhstan; 4245N, 07056E; (GAT). ±
3652: Allium suworowii REGEL;Acmopetala; Kazakhstan; 4308N, 07438E; (GAT). ±
3655: Allium saposhnikovii NIKITINA;Acmopetala; Kazakhstan; 4253N, 07106E;
(GAT). ± 3657: Allium suworowii REGEL;Acmopetala; Kazakhstan; 4149N, 06924E;
(GAT). ± 3659: Allium stipitatum REGEL;Megaloprason; Uzbekistan; 3917N, 06654E;
(GAT). ± 3661: Allium gypsaceum POPOV &VVED.; Popovia; Uzbekistan; 3815N,
06651E; (GAT, photos). ± 3666: Allium jesdianum BOISS.&BUHSE subsp. angustite-
palum (WENDELBO)F.O.KHASS. & R. M. FRITSCH;Megaloprason; Uzbekistan; 3755N,
06647E; (GAT). ± 3670: Allium stipitatum REGEL;Megaloprason; Uzbekistan; 3756N,
06645E; (GAT). ± 3671: Allium jesdianum BOISS.&BUHSE subsp. angustitepalum
(WENDELBO)F.O.KHASS. & R. M. FRITSCH;Megaloprason; Uzbekistan; 3756N,
06645E; (GAT). ± 3672: Allium protensum WENDELBO;Kaloprason; Uzbekistan;
3756N, 06645E; (GAT). ± 3673: Allium sarawschanicum REGEL;Megaloprason; Uz-
bekistan; 3919N, 06656E (type location); (GAT). ± 3676: Allium karataviense REGEL;
Miniprason; Kazakhstan; 4339N, 06838E (near type location); (GAT). ± 3686: Allium
sewerzowii REGEL;Acmopetala; Kazakhstan; 4250N, 06952E; (GAT). ± 3693: Allium
fetisowii REGEL;Acmopetala; Kazakhstan; 4321N, 07500E; (GAT). ± 3699: Allium tu-
lipifolium LEDEB.;Melanocrommyum; Kazakhstan; 4445N, 07623E; (GAT). ± 3703:
Allium aroides POPOV &VVED.; Aroidea; Uzbekistan; 3917N, 06654E; (GAT). ± 3714:
Allium sergii VVED.; Brevicaule; Kazakhstan; 4339N, 06838E; (GAT, photos). ± 3723:
Allium vvedenskyanum PAVLOV;Acmopetala; Kazakhstan; 4321N, 07500E (near type
location); (GAT, photos). ± 3724: Allium iliense REGEL;Regeloprason; Kazakhstan;
4365N, 07705E (type location); (GAT, photos). ± 3738: Allium stipitatum REGEL;
Megaloprason; Kazakhstan; Kurdai pass area, Kurdai gorge; (GAT). ± 3783: Allium
jesdianum BOISS.&BUHSE subsp. angustitepalum (WENDELBO)F.O.KHASS. & R. M.
FRITSCH;Megaloprason; Hoog & Dix-Export, Heemstede, Netherlands; (GAT). ± 3787:
Allium chodsha-bakirganicum GAFFAROV &TURAKULOV;Regeloprason; Kyrgyzstan;
Northern slopes of Turkestan mountain range (type location); (GAT, photos). ± 3884:
Allium macleanii BAKER;Compactoprason; Mark MCDONOUGH, Pepperell, USA;
156
(GAT). ± 3924: Allium bakhtiaricum REGEL;Megaloprason; Iran; 3212N, 05032E;
(GAT). ± 3927: Allium assadii SEISUMS;Acmopetala; Iran; 3332N, 05104E; (GAT). ±
3932: Allium elburzense WENDELBO;Kaloprason; Iran; prov. Tehran, Karaj valley;
(GAT). ± 3936: Allium elburzense WENDELBO;Kaloprason; Iran; 3601N, 05109E;
(GAT). ± 3940: Allium aff. zagricum R. M. FRITSCH;Acanthoprason; Iran; 3713N,
04746E; (GAT). ± 3947: Allium cardiostemon FISCH. & C. A. MEY.; Pseudoprason; Iran;
3804N, 04459E; (GAT). ± 3948: Allium materculae BORDZ.; Acanthoprason; Iran;
3814N, 04450E; (GAT). ± 3951: Allium jesdianum BOISS.&BUHSE subsp. jesdianum;
Megaloprason; Iran; 3132N, 05413E (type location); (GAT). ± 3953: Allium jesdianum
BOISS.&BUHSE subsp. jesdianum;Megaloprason; Iran; 3132N, 05413E (type loca-
tion); (GAT). ± 3958: Allium stipitatum REGEL;Megaloprason; Iran; 3219N, 05030E;
(GAT). ± 3962: Allium stipitatum REGEL;Megaloprason; Iran; 3222N, 05028E; (GAT).
± 3964: Allium stipitatum REGEL;Megaloprason; Iran; 3226N, 05006E; (GAT). ± 3967:
Allium stipitatum REGEL;Megaloprason; Iran; 3221N, 05021E; (GAT). ± 3I: Allium
tulipifolium LEDEB.;Melanocrommyum; Kazakhstan; 4423N, 07649E; (GAT). ± 3o:
Allium orientale BOISS. s. lat.; Melanocrommyum; Israel; near Sartaba; (HUJ). ± 3s:
Allium saralicum R. M. FRITSCH;Melanocrommyum; Iran; prov. Kurdestan, Saral
area; (HCAT). ± 4: Allium jesdianum BOISS.&BUHSE;Megaloprason; Iran; prov. Kur-
destan, Saral area; (HCAT). ± 40711: Allium ellisii HOOK. f.; Acanthoprason; Iran;
prov. Semnan, c. 50 km N Semnan; (TARI 40711). ± 40909: Allium aff. scotostemon
WENDELBO;Acanthoprason; Iran; prov. Semnan, c. 20 km NW Shahrud; (TARI 40909).
± 4522H: Allium sp.5; Acanthoprason; Iran; prov. Kurdestan, cultivated in Sanandaj;
(Sanandaj 4522). ± 4I: Allium viridulum LEDEB.;Melanocrommyum; Kazakhstan;
4543N, 08016E; (GAT). ± 5: Allium saralicum R. M. FRITSCH;Melanocrommyum; Iran;
prov. Kurdestan, Chatan; (HCAT). ± 5013H: Allium austroiranicum R. M. FRITSCH;
Acanthoprason; Iran; prov. Kurdestan, Sanandaj to Maryvan, Mt. Shaneshin; (Sa-
nandaj 5013). ± 5027: Allium dasyphyllum VVED.; Acmopetala; Kyrgyzstan; 4253N,
07135E (type location); (GAT). ± 5040: Allium karataviense REGEL;Miniprason; Uz-
bekistan; 4136N, 06955E; (GAT). ± 5043: Allium severtzovioides R. M. FRITSCH;Ac-
mopetala; Kazakhstan; 4245N, 07056E; (GAT). ± 5047: Allium saposhnikovii NIKI-
TINA;Acmopetala; Kazakhstan; 4254N, 07108E; (GAT). ± 5052: Allium fetisowii RE-
GEL;Acmopetala; Kyrgyzstan; 4253N, 07135E; (GAT). ± 5057: Allium chychkanense
R. M. FRITSCH;Acmopetala; Kyrgyzstan; 4215N, 07300E; (GAT). ± 5060: Allium
chychkanense R. M. FRITSCH;Acmopetala; Kyrgyzstan; 4210N, 07252E; (GAT). ±
5066: Allium schachimardanicum VVED.; Acmopetala; Uzbekistan; 3958N, 07143E
(type location); (GAT). ± 5068: Allium taeniopetalum POPOV &VVED. subsp. tur-
akulovii R. M. FRITSCH &F.O.KHASS.; Acmopetala; Kyrgyzstan; 3959N, 06957E (type
location); (GAT). ± 5076: Allium lipskyanum VVED.; Regeloprason; Uzbekistan;
3844N, 06757E; (GAT). ± 5078: Allium nevskianum VVED.exWENDELBO;Kaloprason;
Uzbekistan; 3845N, 06758E; (GAT). ± 5079: Allium nevskianum VVED.exWENDELBO;
Kaloprason; Uzbekistan; 3840N, 06757E; (GAT). ± 5080: Allium stipitatum REGEL;
Megaloprason; Uzbekistan; 3838N, 06757E; (GAT). ± 5081: Allium rosenorum R. M.
FRITSCH;Megaloprason; Uzbekistan; 3838N, 06757E; (GAT). ± 5092: Allium ovalifo-
lium HAND.-MAZZ.; subg. Anguinum; China; Provinz Quinghai, 120 km NO
ÈXining;
(GAT, photos). ± 5132: Allium rosenorum R. M. FRITSCH `Purple King'; Megaloprason;
Hoog & Dix Export, Heemstede, Netherlands; (GAT). ± 5133: Allium rosenorum R. M.
FRITSCH;Megaloprason; Hoog & Dix-Export, Heemstede, Netherlands; (GAT). ± 5135:
Allium-hybrid, subg. Melanocrommyum `Mars'; Hoog & Dix Export, Heemstede
157
Netherland; (GAT). ± 5136: Allium-hybrid, subg. Melanocrommyum `Lucy Ball';
Hoog & Dix Export, Heemstede Netherland; (GAT). ± 5137: Allium rosenorum R. M.
FRITSCH `Michael H. Hoog'; Megaloprason; Hoog & Dix-Export, Heemstede, Nether-
lands; (GAT). ± 5244: Allium taeniopetalum POPOV &VVED. subsp. taeniopetalum;
Acmopetala; Uzbekistan; 4021N, 06602E; (GAT). ± 5249: Allium taeniopetalum POPOV
&VVED. subsp. taeniopetalum;Acmopetala; Uzbekistan; 4023N, 06603E; (GAT). ±
5262: Allium cristophii TRAUTV.; Kaloprason; Turkmenistan; 3757N, 05759E; (GAT,
photos). ± 5263: Allium stipitatum Regel; Megaloprason; Turkmenistan; 3757N,
05759E; (GAT). ± 5264: Allium isakulii R. M. Fritsch & F. O. Khass. subsp. balkhani-
cum R. M. FRITSCH &F.O.KHASS.; Regeloprason; Turkmenistan; 3942N, 05428E (type
location); (GAT). ± 5272: Allium helicophyllum VVED.; Kaloprason; Turkmenistan;
3843N, 05616E; (GAT). ± 5278: Allium isakulii R. M. FRITSCH &F.O.KHASS. subsp.
subkopetdagense R. M. FRITSCH &F.O.KHASS.; Regeloprason; Turkmenistan; 3829N,
05622E (type location); (GAT). ± 5296: Allium cyrilli TEN.; Melanocrommyum; Tur-
key; 4157N, 02711E; (GAT). ± 5321: Allium multibulbosum JACQ.; Melanocrommyum;
Turkey; 3834N, 02727E; (GAT). ± 5352: Allium orientale BOISS.; Melanocrommyum;
Turkey; 3659N, 03030E; (GAT). ± 5365: Allium orientale BOISS.; Melanocrommyum;
Turkey; 3634N, 03022E; (GAT). ± 5371: Allium orientale BOISS.; Melanocrommyum;
Turkey; 3653N, 03020E; (GAT). ± 5447: Allium motor KAMELIN;Acmopetala; Uzbeki-
stan; Chatkal mountain range; (GAT). ± 5449: Allium severtzovioides R. M. FRITSCH;
Acmopetala; Uzbekistan; Chatkal mountain range, near Chimgan; (GAT). ± 5451:
Allium nevskianum VVED.exWENDELBO;Kaloprason; Uzbekistan; SW Hissar moun-
tain range, pass Artusgar; (GAT). ± 5455: Allium macleanii BAKER;Compactoprason;
Hoog & Dix-Export, Heemstede, Netherlands; (GAT). ± 5475: Allium stipitatum RE-
GEL;Megaloprason; Hoog & Dix-Export, Heemstede, Netherlands; (GAT). ± 5476:
Allium-hybrid, subg. Melanocrommyum `Globemaster'; Hoog & Dix-Export, Heem-
stede Netherland; (GAT). ± 5477: Allium-hybrid, subg. Melanocrommyum `Gla-
diator'; Hoog & Dix-Export, Heemstede Netherland; (GAT). ± 5478: Allium rose-
norum R. M. FRITSCH `Purple King'; Megaloprason; Hoog & Dix-Export, Heemstede,
Netherlands; (GAT). ± 5479: Allium-hybrid, subg. Melanocrommyum `His Ex-
cellency'; Hoog & Dix-Export, Heemstede Netherland; (GAT). ± 5480: Allium stipi-
tatum REGEL `White Giant'; Megaloprason; Hoog & Dix-Export, Heemstede, Nether-
lands; (GAT). ± 5531: Allium aschersonianum W. B ARBEY;Melanocrommyum; Israel;
Jordan valley, near the border to Jordan; (GAT, photos). ± 5562: Allium aflatunense B.
FEDTSCH.; Megaloprason; Kyrgyzstan, received from Olomouc collection; (GAT). ±
5614: Allium rosenbachianum REGEL s. l.; Megaloprason; Dushanbe Botanical Gar-
den, Tajikistan; (GAT). ± 5632: Allium aflatunense B. FEDTSCH.; Megaloprason; Kyr-
gyzstan; Central Tien-Shan mountain range, valley Kokomeren; (GAT). ± 5669: Al-
lium gypsaceum POPOV &VVED.; Popovia; Uzbekistan; 3812N, 06639E; (GAT, photos).
± 5676: Allium alaicum VVED.; Acmopetala; Kyrgyzstan; 4031N, 07234E; (GAT). ±
5679: Allium pseudowinklerianum R. M. FRITSCH &F.O.KHASS.; Regeloprason; Kyr-
gyzstan; 4058N, 07333E; (GAT). ± 5680: Allium backhousianum REGEL;Acmopetala;
Kyrgyzstan; 4058N, 07333E; (GAT, photos). ± 5685: Allium fetisowii REGEL;Acmope-
tala; Kyrgyzstan; 4245N, 07433E; (GAT). ± 5686: Allium saposhnikovii NIKITINA;Ac-
mopetala; Kyrgyzstan; 4245N, 07433E (type location); (GAT). ± 5688: Allium fetisowii
REGEL;Acmopetala; Kyrgyzstan; 4235N, 07454E; (GAT). ± 5690: Allium karataviense
REGEL;Miniprason; Kyrgyzstan; 4230N, 07321E; (GAT). ± 5692: Allium arkitense R.
M. FRITSCH;Acmopetala; Kyrgyzstan; 4150N, 07158E (type location); (GAT). ± 5694:
158
Allium aflatunense B. FEDTSCH.; Megaloprason; Kyrgyzstan; 4150N, 07158E (near
type location); (GAT). ± 5695: Allium dodecadontum VVED.; Acmopetala; Kyrgyzstan;
4150N, 07158E (type location); (GAT). ± 5699: Allium viridiflorum POBED.; Verti-
cillata; Kyrgyzstan; 4150N, 07158E (type location); (GAT). ± 5702: Allium kar-
ataviense REGEL;Miniprason; Kyrgyzstan; 4150N, 07158E; (GAT). ± 5707: Allium
dodecadontum VVED.; Acmopetala; Kyrgyzstan; 4139N, 07202E; (GAT). ± 5713: Al-
lium karataviense REGEL;Miniprason; Tajikistan; 4017N, 06926E; (GAT). ± 5778: Al-
lium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmopetala; Uzbekistan; 4108N,
07022E; (GAT, photos). ± 5780: Allium costatovaginatum KAMELIN &LEVICHEV;Ac-
mopetala; Uzbekistan; 4108N, 07022E; (GAT, photos). ± 5783: Allium costatovagina-
tum KAMELIN &LEVICHEV;Acmopetala; Uzbekistan; 4108N, 07022E; (GAT, photos). ±
5786: Allium isakulii R. M. FRITSCH &F.O.KHASS. subsp. isakulii;Regeloprason;
Tajikistan; Mogoltau mountain range 4017N, 06927E (type location); (GAT). ± 5788:
Allium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmopetala; Uzbekistan; 6957N,
04138E; (GAT). ± 5789: Allium severtzovioides R. M. FRITSCH;Acmopetala; Uzbeki-
stan; 6957N, 04138E; (GAT). ± 5791: Allium suworowii REGEL;Acmopetala; Uzbeki-
stan; 4845N, 04000E (near type location); (GAT). ± 5793: Allium karataviense REGEL;
Miniprason; Uzbekistan; 4104N, 07034E; (GAT). ± 5803: Allium karataviense REGEL;
Miniprason; Tajikistan; 3951N, 07040E; (GAT, photos). ± 5808: Allium cupuliferum
REGEL subsp. cupuliferum;Regeloprason; Uzbekistan; Nuratau mountain range;
(GAT). ± 5837: Allium decipiens FISCH.exSCHULT.&SCHULT. f. subsp. quercetorum
SEREGIN;Melanocrommyum; Ukraina; Crimea, N Mt. Ayu-Dag; (GAT). ± 5844: Allium
sewerzowii REGEL;Acmopetala; Kazakhstan; Aksu-Zhabagly natural reserve; (GAT,
photos). ± 5873: Allium caspium (PALL.) M. BIEB. subsp. baissunense (LIPSKY)F.O.
KHASS. & R. M. FRITSCH;Kaloprason; Uzbekistan; 3812N, 06715E (type location);
(GAT, photos). ± 5878: Allium nevskianum VVED.exWENDELBO;Kaloprason; Uzbe-
kistan; 3835N, 06728E; (GAT). ± 5879: Allium severtzovioides R. M. FRITSCH;Acmo-
petala; Uzbekistan; 4140N, 06946E (type location); (GAT). ± 5885: Allium tashkenti-
cum F. O. Khass. & R. M. FRITSCH;Acmopetala; Uzbekistan; 4132N, 07002E; (GAT,
photos). ± 5888: Allium karataviense REGEL;Miniprason; Uzbekistan; 4112N, 07015E;
(GAT). ± 5907: Allium rosenorum R. M. FRITSCH;Megaloprason; Uzbekistan; 3850N,
06707E; (GAT). ± 5908: Allium komarowii LIPSKY;Compactoprason; Uzbekistan;
Hissar mountain range, near Kaltakul; (GAT). ± 5910: Allium taeniopetalum POPOV &
VVED. subsp. mogoltavicum (VVED.) R. M. FRITSCH &F.O.KHASS.; Acmopetala; Uz-
bekistan; 4121N, 06956E; (GAT). ± 5916: Allium stipitatum REGEL;Megaloprason;
Turkmenistan; 3758N, 05801E; (GAT). ± 5917: Allium regelii TRAUTV.; Regeloprason;
Turkmenistan; 3748N, 05822E (near type location); (GAT, photos). ± 5919: Allium
isakulii R. M. FRITSCH &F.O.KHASS . subsp. subkopetdagense R. M. FRITSCH &F.O.
KHASS.; Regeloprason; Turkmenistan; 3830N, 05705E; (GAT, photos). ± 5920: Allium
cristophii TRAUTV.; Kaloprason; Turkmenistan; 3821N, 05658E; (GAT). ± 5922: Allium
brachyscapum VVED.; Acanthoprason; Turkmenistan; 382030N, 0565630E; (GAT,
photos). ± 5927: Allium caspium (PALL.) M. BIEB. subsp. caspium;Kaloprason; Turk-
menistan; 3915N, 05518E; (GAT). ± 5931: Allium isakulii R. M. FRITSCH &F.O.KHASS.
subsp. balkhanicum R. M. FRITSCH &F.O.KHASS.; Regeloprason; Turkmenistan;
3938N, 05421E; (GAT, photos). ± 5936: Allium caspium (PALL.) M. BIEB. subsp. cas-
pium;Kaloprason; Turkmenistan; 400142N, 0525242E; (GAT, photos). ± 5937: Allium
regelii TRAUTV.; Regeloprason; Turkmenistan; 384136N, 0564442E; (GAT, photos). ±
5942: Allium giganteum REGEL;Compactoprason; Turkmenistan; 3757N, 05801E;
159
(GAT). ± 5967: Allium akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason;
Turkey; 3823N, 04323E; (GAT, photos). ± 5977: Allium cardiostemon FISCH. & C. A.
MEY.; Pseudoprason; Turkey; 3903N, 04345E; (GAT). ± 5978: Allium cardiostemon
FISCH. & C. A. MEY.; Pseudoprason; Turkey; 3916N, 04402E; (GAT). ± 5980: Allium
akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason; Turkey; 3930N, 04408E;
(GAT). ± 5I: Allium robustum KAR.&KIR.; Melanocrommyum; Kazakhstan; 4709N,
08206E; (GAT). ± 6: Allium aff. minutiflorum REGEL;Acanthoprason; Iran; prov.
Kurdestan, Chatan; (HCAT). ± 6081: Allium cardiostemon FISCH. & C. A. MEY.; Pseu-
doprason; Armenia; 4023N, 04416E; (GAT). ± 6086: Allium cardiostemon FISCH.&C.
A. MEY.; Pseudoprason; Armenia; 4006N, 04448E; (GAT, photos). ± 6086H: Allium
kingdonii STEARN; subg. Cyathophora; China; 2702N, 10014E; (MB 14122). ± 6090H:
Allium prattii C. H. WRIGHT; subg. Anguinum; China; 3038N, 09312E; (OSBU). ±
6102: Allium cristophii TRAUTV. s. lat.; Kaloprason; Iran; Alborz mountain range;
(GAT, photos). ± 6114: Allium bucharicum REGEL;Kaloprason; Tajikistan; 371745N,
0691528E (type location); (GAT, photos). ± 6119: Allium insufficiens VVED.; Acmope-
tala; Tajikistan; 372249N, 0691243E; (GAT). ± 6120: Allium rosenbachianum REGEL
subsp. rosenbachianum;Megaloprason; Tajikistan; 372159N, 0691242E; (GAT). ±
6122: Allium giganteum REGEL;Compactoprason; Tajikistan; 374358N, 0693945E;
(GAT). ± 6123: Allium rosenbachianum REGEL subsp. rosenbachianum;Mega-
loprason; Tajikistan; 374342N, 0693844E; (GAT, photos). ± 6126: Allium bucharicum
REGEL;Kaloprason; Tajikistan; 374531N, 0694000E; (GAT, photos). ± 6129: Allium
trautvetterianum REGEL;Compactoprason; Tajikistan; 374748N, 0701429E; (GAT,
photos). ± 6130: Allium trautvetterianum REGEL;Compactoprason; Tajikistan;
394752N, 0701334E; (GAT). ± 6131: Allium trautvetterianum REGEL;Compactopra-
son; Tajikistan; 374733N, 0701459E; (GAT, photos). ± 6132: Allium rosenbachianum
REGEL subsp. rosenbachianum;Megaloprason; Tajikistan; 374730N, 0701415E;
(GAT). ± 6133: Allium karataviense REGEL;Miniprason; Tajikistan; 374755N,
0701411E; (GAT). ± 6134: Allium darwasicum REGEL;Regeloprason; Tajikistan;
374755N, 0701411E; (GAT). ± 6137: Allium rosenbachianum REGEL subsp. ro-
senbachianum;Megaloprason; Tajikistan; 3759N, 07011E; (GAT). ± 6138: Allium
darwasicum REGEL;Regeloprason; Tajikistan; 375905N, 0701051E (near type loca-
tion); (GAT, photos). ± 6140: Allium rosenbachianum REGEL subsp. rosenbachianum;
Megaloprason; Tajikistan; 375858N, 0701028E; (GAT). ± 6141: Allium suworowii RE-
GEL;Acmopetala; Tajikistan; 375857N, 0701025E; (GAT). ± 6143: Allium darwasicum
REGEL;Regeloprason; Tajikistan; 380431N, 0701332E (type location); (GAT, photos). ±
6144: Allium darwasicum REGEL;Regeloprason; Tajikistan; 380426N, 0701346E (type
location); (GAT). ± 6145: Allium suworowii REGEL;Acmopetala; Tajikistan; 375548N,
0701538E; (GAT). ± 6147: Allium karataviense REGEL;Miniprason; Tajikistan;
380320N, 0702229E; (GAT). ± 6148: Allium giganteum REGEL;Compactoprason;Ta-
jikistan; Panj valley; (GAT). ± 6150: Allium macleanii BAKER;Compactoprason; Taji-
kistan; 382629N, 0705556E; (GAT). ± 6151: Allium schugnanicum VVED.; Mega-
loprason; Tajikistan; 372410N, 0712949E (type location); (GAT, photos). ± 6153: Al-
lium chitralicum F. T. W ANG &TANG;Brevicaule; Tajikistan; 371200N, 0713208E;
(GAT, photos). ± 6157: Allium hissaricum VVED.; Regeloprason; Tajikistan; 381729N,
0691504E; (GAT). ± 6158: Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan;
381730N, 0691432E; (GAT). ± 6159: Allium rosenbachianum REGEL subsp. kwakense
R. M. FRITSCH;Megaloprason; Tajikistan; 381728N, 0691502E; (GAT). ± 6161: Allium
suworowii REGEL;Acmopetala; Tajikistan; 384841N, 0684827E; (GAT, photos). ±
160
6162: Allium verticillatum REGEL;Verticillata; Tajikistan; 384840N, 0684845E;
(GAT). ± 6169: Allium taeniopetalum POPOV &VVED. subsp. mogoltavicum (VVED.) R.
M. FRITSCH &F.O.KHASS.; Acmopetala; Uzbekistan; 410730N, 0702135E; (GAT,
photos). ± 6171: Allium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmopetala;
Uzbekistan; 413206N, 0695329E; (GAT, photos). ± 6172: Allium motor Kamelin; Ac-
mopetala; Uzbekistan; 413100N, 0700200E; (GAT, photos). ± 6174: Allium severtzo-
vioides R. M. FRITSCH;Acmopetala; Uzbekistan; Chimgan massif, Aksai; (GAT). ±
6175: Allium karataviense REGEL;Miniprason; Uzbekistan; 414429N, 0700438E;
(GAT). ± 6176: Allium severtzovioides R. M. FRITSCH;Acmopetala; Uzbekistan;
414542N, 0700504E; (GAT). ± 6185: Allium gypsaceum POPOV &VVED.; Popovia; Uz-
bekistan; 380519N, 0672654E; (GAT, photos). ± 6187: Allium caspium (PALL.) M. BIEB.
subsp. baissunense (LIPSKY)F.O.KHASS. & R. M. FRITSCH;Kaloprason; Uzbekistan;
380533N, 0672651E (type location); (GAT, photos). ± 6188: Allium giganteum REGEL;
Compactoprason; Uzbekistan; 380533N, 0672651E; (GAT, photos). ± 6189: Allium ar-
oides POPOV &VVED.; Aroidea; Uzbekistan; 384946N, 0670715E; (GAT). ± 6191: Al-
lium rosenorum R. M. FRITSCH;Megaloprason; Uzbekistan; 384906N, 0670740E;
(GAT). ± 6192: Allium sarawschanicum REGEL;Megaloprason; Uzbekistan; 384900N,
0670753E; (GAT, photos). ± 6196: Allium cupuliferum REGEL subsp. cupuliferum;Re-
geloprason; Uzbekistan; 402154N, 0660057E (near type location); (GAT, photos). ±
6201: Allium verticillatum REGEL;Verticillata; Uzbekistan; 400438N, 0674336E;
(GAT, photos). ± 6205: Allium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmope-
tala; Uzbekistan; 414202N, 0695555E (near type location); (GAT, photos). ± 6208: Al-
lium motor KAMELIN;Acmopetala; Uzbekistan; 410702N, 0701418E; (GAT). ± 6236:
Allium chelotum WENDELBO;Melanocrommyum; Iran; 372136N, 0560043E; (GAT). ±
6240: Allium helicophyllum VVED.; Kaloprason; Iran; 371851N, 0560011E; (GAT). ±
6246: Allium cristophii TRAUTV.; Kaloprason; Iran; 372411N, 0561200E; (GAT). ±
6248: Allium sarawschanicum REGEL;Megaloprason; Iran; 372636N, 0560439E;
(GAT). ± 6249: Allium ellisii HOOK. f.; Kaloprason; Iran; 363558N, 0585559E; (GAT). ±
6254: Allium ellisii HOOK. f.; Kaloprason; Iran; 372604N, 0583526E; (GAT). ± 6255:
Allium ellisii HOOK. f.; Kaloprason; Iran; 372955N, 0583525E; (GAT). ± 6258: Allium
giganteum REGEL;Compactoprason; Iran; 373134N, 0583703E; (GAT). ± 6259: Allium
sarawschanicum REGEL;Megaloprason; Iran; 373134N, 0583703E; (GAT). ± 6261: Al-
lium jesdianum BOISS.&BUHSE subsp. jesdianum;Megaloprason; Iran; 361201N,
0590514E; (GAT). ± 6262: Allium kuhsorkhense R. M. FRITSCH &JOHARCHI;Acantho-
prason; Iran; 361201N, 0590514E; (GAT, photos). ± 6263: Allium giganteum REGEL;
Compactoprason; Iran; 361207N, 0590530E; (GAT, photos). ± 6278: Allium stipitatum
REGEL;Megaloprason; Tajikistan; 390625N, 0685112E (type location); (GAT). ± 6279:
Allium stipitatum REGEL;Megaloprason; Tajikistan; 390733N, 0685150E (near type
location); (GAT). ± 6282: Allium komarowii LIPSKY;Compactoprason; Tajikistan;
391056N, 0685449E; (GAT, photos). ± 6284: Allium alexeianum REGEL;Kaloprason;
Tajikistan; 390336N, 0682045E; (GAT). ± 6289: Allium komarowii LIPSKY;Compacto-
prason; Tajikistan; 390400N, 0682024E; (GAT). ± 6290: Allium rosenorum R. M.
FRITSCH;Megaloprason; Tajikistan; 390303N, 0681715E; (GAT). ± 6291: Allium alex-
eianum REGEL;Kaloprason; Tajikistan; 390320N, 0681648E; (GAT). ± 6292: Allium
winklerianum REGEL;Regeloprason; Tajikistan; 390322N, 0681604E; (GAT). ± 6296:
Allium komarowii LIPSKY;Compactoprason; Tajikistan; 390850N, 0681718E (near
type location); (GAT). ± 6298: Allium rosenorum R. M. FRITSCH;Megaloprason; Taji-
kistan; 391220N, 0674829E; (GAT). ± 6299: Allium komarowii LIPSKY;Compactopra-
161
son; Tajikistan; 391220N, 0674829E; (GAT, photos). ± 6300: Allium stipitatum REGEL;
Megaloprason; Tajikistan; 390641N, 0675050E; (GAT). ± 6303: Allium alexeianum
REGEL;Kaloprason; Tajikistan; 390612N, 0675035E; (GAT). ± 6304: Allium sar-
awschanicum REGEL;Megaloprason; Tajikistan; 390642N, 0675038E; (GAT). ± 6311:
Allium rosenorum R. M. FRITSCH;Megaloprason; Tajikistan; 391234N, 0674320E;
(GAT). ± 6313: Allium komarowii LIPSKY;Compactoprason; Tajikistan; Penjakent re-
gion, near Obiborik; (GAT, photos). ± 6314: Allium sarawschanicum REGEL;Mega-
loprason; Tajikistan; 391238N, 0674312E; (GAT). ± 6315: Allium winklerianum RE-
GEL;Regeloprason; Tajikistan; 390428N, 0685206E; (GAT, photos). ± 6340: Allium ro-
thii ZUCC.; Melanocrommyum; Jordan; 321901N, 0373528E; (LI). ± 6342: Allium rothii
ZUCC.; Melanocrommyum; Jordan; 320115N, 0362500E; (LI). ± 6345: Allium rothii
ZUCC.; Melanocrommyum; Jordan; 300142N, 0352739E; (LI). ± 6350: Allium cupuli-
ferum REGEL subsp. cupuliferum;Regeloprason; Uzbekistan; 403131N, 0664604E;
(GAT). ± 6354: Allium cupuliferum REGEL subsp. cupuliferum;Regeloprason; Uzbe-
kistan; Nuratau, north of village Uhum; (GAT). ± 6356: Allium cupuliferum REGEL
subsp. cupuliferum;Regeloprason; Uzbekistan; 403123N, 0664351E; (GAT). ± 6357:
Allium altissimum REGEL;Megaloprason; Uzbekistan; 403120N, 0664508E; (GAT). ±
6362: Allium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmopetala; Uzbekistan;
411456N, 0694914E; (GAT). ± 6364: Allium costatovaginatum KAMELIN &LEVICHEV;
Acmopetala; Uzbekistan; 411449N, 0694926E; (GAT). ± 6366: Allium karataviense
REGEL;Miniprason; Uzbekistan; 411448N, 0694933E; (GAT, photos). ± 6369: Allium
motor KAMELIN;Acmopetala; Uzbekistan; 411443N, 0695016E; (GAT). ± 6370: Allium
stipitatum REGEL;Megaloprason; Uzbekistan; 411433N, 0694916E; (GAT, photos). ±
6372: Allium taeniopetalum POPOV &VVED. subsp. mogoltavicum (VVED.) R. M.
FRITSCH &F.O.KHASS.; Acmopetala; Uzbekistan; 412138N, 0695532E; (GAT, photos).
± 6373: Allium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmopetala; Uzbekistan;
412127N, 0695529E; (GAT). ± 6377: Allium severtzovioides R. M. FRITSCH;Acmope-
tala; Uzbekistan; 414206N, 0700841E; (GAT). ± 6388: Allium elburzense WENDELBO;
Kaloprason; Iran; 360648N, 0511938E; (GAT). ± 6390: Allium derderianum REGEL;
Acanthoprason; Iran; 360904N, 0511840E; (GAT). ± 6392: Allium elburzense WEN-
DELBO;Kaloprason; Iran; 354522N, 0515723E (type location); (GAT). ± 6397: Allium
akaka S. G. GMELIN ex SCHULT.&SCHULT. f.; Acanthoprason; Iran; 373512N,
0483717E (epitype accession); (GAT). ± 6398: Allium akaka S. G. GMELIN ex SCHULT.
&S
CHULT. f.; Acanthoprason; Iran; 373422N, 0483440E; (GAT). ± 6402: Allium ma-
terculae BORDZ.; Acanthoprason; Iran; 381958N, 0454828E; (GAT). ± 6404: Allium
shelkovnikovii GROSSH.; Acanthoprason; Iran; 381940N, 0454745E; (GAT). ± 6405:
Allium shelkovnikovii GROSSH.; Acanthoprason; Iran; 380817N, 0455437E; (GAT). ±
6406: Allium materculae BORDZ.; Acanthoprason; Iran; 380817N, 0455437E; (GAT). ±
6425: Allium giganteum REGEL;Compactoprason; Tajikistan; 384925N, 0695414E;
(GAT). ± 6427: Allium macleanii BAKER;Compactoprason; Tajikistan; 384357N,
0703557E; (GAT). ± 6431: Allium darwasicum REGEL;Regeloprason; Tajikistan;
383734N, 0704007E; (GAT). ± 6434: Allium winklerianum REGEL;Regeloprason;Ta-
jikistan; 383813N, 0704246E (type location); (TAD). ± 6438: Allium giganteum REGEL;
Compactoprason; Tajikistan; 382556N, 0705524E; (GAT). ± 6445: Allium macleanii
BAKER;Compactoprason; Tajikistan; 383938N, 0720004E; (GAT). ± 6446: Allium gi-
ganteum REGEL;Compactoprason; Tajikistan; 382914N, 0714742E; (GAT). ± 6449:
Allium stipitatum REGEL;Megaloprason; Tajikistan; 383425N, 0705103E; (GAT). ±
6450: Allium intradarvazicum R. M. FRITSCH;Regeloprason; Tajikistan; 383724N,
162
0705149E; (GAT). ± 6452: Allium intradarvazicum R. M. FRITSCH;Regeloprason;Ta-
jikistan; 383802N, 0705206E (type location); (GAT). ± 6454: Allium intradarvazicum
R. M. FRITSCH;Regeloprason; Tajikistan; 382924N, 0705120E; (GAT). ± 6461: Allium
rosenbachianum REGEL subsp. kwakense R. M. FRITSCH;Megaloprason; Tajikistan;
384224N, 0703311E; (GAT). ± 6475: Allium ubipetrense R. M. FRITSCH;Acanthopra-
son; Iran; 354749N, 0492124E; (GAT). ± 6478: Allium breviscapum STAPF;Acantho-
prason; Iran; 344537N, 0482604E (type location); (GAT). ± 6479: Allium stipitatum
REGEL;Megaloprason; Iran; prov. Hamadan, Assad Abad district; (GAT). ± 6480: Al-
lium breviscapum STAPF;Acanthoprason; Iran; 344022N, 0482512E; (GAT). ± 6484:
Allium stipitatum REGEL;Megaloprason; Iran, vegetable market in Assad Abad;
(GAT). ± 6485: Allium stipitatum REGEL;Megaloprason; Iran, vegetable market in
Hamadan; (GAT). ± 64855: Allium aff. akaka S. G. GMELIN ex SCHULT.&SCHULT. f.;
Acanthoprason; Iran; prov. Zanjan, Mahneshan, Anguran, Belgheis Mts.; (TARI
64855). ± 6487: Allium hamedanense R. M. FRITSCH;Melanocrommyum; Iran;
344511N, 0483615E (type location); (GAT). ± 6489: Allium bisotunense R. M. FRITSCH;
Melanocrommyum; Iran; 342725N, 0473442E (type location); (GAT). ± 6490: Allium
noeÈanum REUT.exREGEL;Melanocrommyum; Iran; 341640N, 0464933E; (IRAN). ±
6491: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran;
341407N, 0464012E; (GAT). ± 6496: Allium jesdianum BOISS.&BUHSE subsp. re-
mediorum R. M. FRITSCH;Megaloprason; Iran, vegetable market in Kerend; (GAT). ±
6498: Allium jesdianum BOISS.&BUHSE subsp. remediorum R. M. FRITSCH;Mega-
loprason; Iran; 345807N, 0462744E (type location); (GAT). ± 6499: Allium koelzii
(WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran; 345814N, 0462755E; (GAT).
± 6500: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran;
345814N, 0462755E; (IRAN). ± 6501: Allium koelzii (WENDELBO)K.PERSS.&WEN-
DELBO;Pseudoprason; Iran; 342543N, 0472301E; (GAT). ± 6502: Allium stipitatum
REGEL;Megaloprason; Iran; 352152N, 0465210E; (GAT). ± 6503: Allium ubipetrense
R. M. FRITSCH;Acanthoprason; Iran; 352214N, 0465129E (type location); (IRAN
43975). ± 6506: Allium saralicum R. M. FRITSCH;Melanocrommyum; Iran; 354023N,
0470722E (type location); (GAT). ± 6507: Allium ubipetrense R. M. FRITSCH;Acan-
thoprason; Iran; 360057N, 0465552E; (GAT). ± 6508: Allium jesdianum BOISS.&
BUHSE subsp. jesdianum;Megaloprason; Iran; 360101N, 0485548E; (GAT). ± 6509:
Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran; 360336N,
0465147E; (IRAN). ± 6516: Allium derderianum REGEL;Acanthoprason; Iran;
360212N, 0511200E; (GAT). ± 6519: Allium elburzense WENDELBO;Kaloprason; Iran;
360646N, 0511936E; (GAT). ± 6523: Allium jesdianum BOISS.&BUHSE subsp. angu-
stitepalum (WENDELBO)F.O.KHASS. & R. M. FRITSCH;Megaloprason; Uzbekistan;
380122N, 0665014E; (GAT). ± 6525: Allium verticillatum REGEL;Verticillata; Uzbe-
kistan; 380220N, 0665121E; (GAT). ± 6526: Allium protensum WENDELBO;Kalopra-
son; Uzbekistan; 380240N, 0665128E; (GAT, photos). ± 6528: Allium alexeianum RE-
GEL;Kaloprason; Uzbekistan; 3938N, 06829E; (GAT). ± 6530: Allium komarowii
LIPSKY;Compactoprason; Uzbekistan; 393827N, 0683011E; (GAT). ± 6531: Allium
taeniopetalum POPOV &VVED.; Acmopetala; Uzbekistan; 394040N, 0682903E; (GAT).
± 6532: Allium suworowii REGEL;Acmopetala; Uzbekistan; 394536N, 0682441E;
(GAT). ± 6533: Allium karataviense REGEL;Miniprason; Uzbekistan; 414207N,
0700801E; (GAT). ± 6535: Allium tashkenticum F. O. KHASS. & R. M. FRITSCH;Acmo-
petala; Uzbekistan; 414204N, 0700816E; (GAT). ± 6553: Allium darwasicum REGEL;
Regeloprason; Tajikistan; 385325N, 0705542E; (GAT). ± 6557: Allium darwasicum
163
REGEL;Regeloprason; Tajikistan; 385233N, 0710538E; (GAT). ± 6565: Allium robus-
tum KAR.&KIR.; Melanocrommyum; Kazkhstan; 4747N, 08220E; (GAT). ± 6570: Al-
lium rosenbachianum REGEL subsp. kwakense R. M. FRITSCH;Megaloprason; Tajiki-
stan; 385422N, 0710135E; (GAT). ± 6597: Allium rosenbachianum REGEL subsp. kwa-
kense R. M. FRITSCH;Megaloprason; Tajikistan; 384228N, 0703303E; (GAT). ± 6612:
Allium materculae BORDZ. subsp. graveolens R. M. FRITSCH;Acanthoprason; Iran;
341721N, 0501144E; (GAT). ± 6613: Allium materculae BORDZ. subsp. graveolens R.
M. FRITSCH;Acanthoprason; Iran; 341544N, 0494532E; (GAT). ± 6616: Allium moder-
ense R. M. FRITSCH;Melanocrommyum; Iran; 340613N, 0493834E (type location);
(GAT). ± 6617: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran; 335215N,
0492649E; (GAT). ± 6620: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran;
335205N, 0492613E; (GAT). ± 6621: Allium ubipetrense R. M. FRITSCH;Acanthopra-
son; Iran; prov. Markazi, N Razvand massif; (IRAN). ± 6622: Allium ubipetrense R. M.
FRITSCH;Acanthoprason; Iran; 335210N, 0492628E; (GAT). ± 6625: Allium jesdianum
BOISS.&BUHSE subsp. jesdianum;Megaloprason; Iran; 335204N, 0492627E; (GAT). ±
6627: Allium materculae BORDZ.subsp. graveolens R. M. FRITSCH;Acanthoprason;
Iran; 335837N, 0500704E; (GAT). ± 6628: Allium materculae BORDZ.subsp. graveolens
R. M. FRITSCH;Acanthoprason; Iran; 335841N, 0500647E; (GAT). ± 6629: Allium
moderense R. M. FRITSCH;Melanocrommyum; Iran; 355825N, 0500642E; (IRAN). ±
6630: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran; 335305N, 0485812E;
(GAT). ± 6632: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran; 333325N,
0484950E; (GAT). ± 6633: Allium zagricum R. M. FRITSCH;Acanthoprason; Iran;
333324N, 0484948E; (GAT). ± 6635: Allium jesdianum BOISS.&BUHSE subsp. re-
mediorum R. M. FRITSCH;Megaloprason; Iran; prov. Lurestan, market in Khorram
Abad; (GAT). ± 6637: Allium jesdianum BOISS.&BUHSE subsp. remediorum R. M.
FRITSCH;Megaloprason; Iran; 335836N, 0482021E; (GAT). ± 6638: Allium jesdianum
BOISS.&BUHSE subsp. remediorum R. M. FRITSCH;Megaloprason; Iran; 335834N,
0482041E; (GAT). ± 6639: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseu-
doprason; Iran; 335828N, 0482052E; (GAT). ± 6640: Allium zagricum R. M. FRITSCH;
Acanthoprason; Iran; 335813N, 0482140E; (GAT). ± 6641: Allium zagricum R. M.
FRITSCH;Acanthoprason; Iran; 335813N, 0482140E; (GAT). ± 6643: Allium zagricum
R. M. FRITSCH;Acanthoprason; Iran; 333836N, 0482731E (type location); (GAT). ±
6644: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran;
333836N, 0482731E; (GAT). ± 6648: Allium haemanthoides BOISS.&REUT.exREGEL s.
str.; Acanthoprason; Iran; 350053N, 0462133E (type location); (GAT). ± 6651: Allium
keusgenii R. M. FRITSCH;Melanocrommyum; Iran; 343858N, 0473650E (type location);
(GAT). ± 6652: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason;
Iran; 343910N, 0473656E; (GAT). ± 6657: Allium derderianum REGEL;Acanthoprason;
Iran; 355200N, 0512330E; (GAT). ± 6658: Allium elburzense WENDELBO;Kaloprason;
Iran; 355144N, 0512307E; (IRAN). ± 6682: Allium nigrum L.; Melanocrommyum;
Greece; 351046N, 0243359E; (GAT, photos). ± 6692: Allium materculae BORDZ.subsp.
graveolens R. M. FRITSCH;Acanthoprason; Iran; 332906N, 0515758E; (IRAN). ± 6694:
Allium sp.1ªchlorotepalumº;Acanthoprason; Iran; 325512N, 0503350E; (GAT). ±
6697: Allium sp.1ªchlorotepalumº;Acanthoprason; Iran; 325523N, 0503330E (type
location); (GAT). ± 6699: Allium sp.;Acanthoprason; Iran; 325523N, 0503330E;
(IRAN). ± 6703: Allium cathodicarpum WENDELBO;Regeloprason; Iran; 302339N,
0531258E; (IRAN). ± 6704: Allium austroiranicum R. M. FRITSCH;Acanthoprason;
Iran; 294113N, 0520326E; (GAT). ± 6706: Allium jesdianum BOISS.&BUHSE;Mega-
164
loprason; Iran; 293730N, 0520038E; (IRAN). ± 6708: Allium austroiranicum R. M.
FRITSCH;Pseudoprason; Iran; 293712N, 0520031E; (GAT). ± 6709: Allium austroir-
anicum R. M. FRITSCH;Acanthoprason; Iran; 301806N, 0515755E; (GAT). ± 6710: Al-
lium austroiranicum R. M. FRITSCH;Acanthoprason; Iran; 301806N, 0515755E; (GAT).
± 6711: Allium stipitatum REGEL s. lat.; Megaloprason; Iran; 302349N, 0515431E;
(IRAN). ± 6712: Allium austroiranicum R. M. FRITSCH;Acanthoprason; Iran; 302349N,
0515431E; (GAT). ± 6716: Allium jesdianum BOISS.&BUHSE subsp. jesdianum;
Megaloprason; Iran; 313637N, 0540558E (type location); (GAT). ± 6722: Allium sco-
tostemon WENDELBO;Thaumasioprason; Iran; 354707N, 0522233E; (IRAN). ± 6734:
Allium austroiranicum R. M. FRITSCH;Acanthoprason; Iran; 325523N, 0503330E;
(GAT). ± 6736: Allium verticillatum REGEL;Verticillata; Uzbekistan; 300953N,
0665846E; (GAT, photos). ± 6737: Allium caspium (PALL.) M. BIEB. subsp. baissunense
(LIPSKY)F.O.KHASS. & R. M. FRITSCH;Kaloprason; Uzbekistan; 373207N, 0664119E;
(GAT, photos). ± 6738: Allium gypsaceum POPOV &VVED.; Popovia; Uzbekistan;
373517N, 0663258E; (GAT, photos). ± 6739: Allium verticillatum REGEL;Verticillata;
Uzbekistan; 380427N, 0681332E; (GAT, photos). ± 6740: Allium caspium (PALL.) M.
BIEB. subsp. baissunense (LIPSKY)F.O.KHASS. & R. M. FRITSCH;Kaloprason; Uzbe-
kistan; 380716N, 0661323E; (GAT, photos). ± 6741: Allium nevskianum VVED.ex
WENDELBO;Kaloprason; Uzbekistan; 383028N, 0673930E; (GAT, photos). ± 6742: Al-
lium suworowii REGEL;Acmopetala; Uzbekistan; 383208N, 0673353E; (GAT, photos).
± 6743: Allium nevskianum VVED.exWENDELBO;Kaloprason; Uzbekistan; 383157N,
0673444E; (GAT, photos). ± 6744: Allium giganteum Regel; Compactoprason; Uzbe-
kistan; 380446N, 0672536E; (GAT, photos). ± 6745: Allium protensum WENDELBO;
Kaloprason; Uzbekistan; 381510N, 0670140E; (GAT, photos). ± 6746: Allium gigan-
teum REGEL;Compactoprason; Uzbekistan; 380828N, 0680659E; (GAT, photos). ±
6747: Allium rosenorum R. M. FRITSCH;Megaloprason; Uzbekistan; 385045N,
0670706E; (GAT, photos). ± 6748: Allium komarowii LIPSKY;Compactoprason; Uzbe-
kistan; 305028N, 0670736E; (GAT, photos). ± 6749: Allium majus VVED.; Compacto-
prason; Uzbekistan; 391156N, 0671715E; (GAT, photos). ± 6750: Allium sp.;Acmope-
tala?; Uzbekistan; 410539N, 0704231E; (GAT, photos). ± 6913H: Allium sp. 3; Acan-
thoprason; Iran; prov. Azarbeijan, Piranshahr, Mt. Dalanpar; (Sanandaj 6913). ± 6I:
Allium tulipifolium LEDEB.; Melanocrommyum; Kazakhstan; 4556N, 08003E; (GAT).
± 70664: Allium aff. egorovae M. V. AGAB.&OGAN.; Acanthoprason; Iran; prov. Azar-
beijan, Arasbaran to Ahar 7 km to Barzid village; (TARI 70664). ± 7600H: Allium sp.4
ªmozaffarianiiº;Melanocrommyum; Iran; prov. Kurdestan, Sanandaj to Divandarreh,
near Abbas-Abad; (Sanandaj 7600). ± 7709H: Allium ubipetrense R. M. FRITSCH;
Acanthoprason; Iran; prov. Kurdestan, Saghez to Baneh, Piromaran village; (Sa-
nandaj 7709). ± 7880H: Allium sp.8; Acanthoprason; Iran; prov. Kurdestan, culti-
vated in Sanandaj; (Sanandaj 7880). ± 7I: Allium tulipifolium LEDEB.; Melano-
crommyum; Kazakhstan; 443400N, 0763800E; (GAT). ± 8196H: Allium austroir-
anicum R. M. FRITSCH;Acanthoprason; Iran; prov. Kurdestan, Mt. Koremyryam SW
Sanadaj; (Sanandaj 8196). ± 8464H: Allium sp.3; Acanthoprason; Iran; prov. Kurde-
stan, Maryvan; (Sanandaj 8464). ± 8465H: Allium sp.3; Acanthoprason; Iran; prov.
Kurdestan, Maryvan; (Sanandaj 8465). ± 8480H: Allium aff. egorovae M. V. AGAB.&
OGAN.; Acanthoprason; Iran; prov. E Azarbeijan, Arasbaran, Mt. Doghoron; (Sa-
nandaj 8480). ± 8684H: Allium sp.4ªmozaffarianiiº;Melanocrommyum; Iran; prov.
Kurdestan, Maryvan, Darband Dezli; (Sanandaj 8684). ± 8858H: Allium sp.5; Acan-
thoprason; Iran; prov. Kurdestan, Maryvan, Mt. Dalani; (Sanandaj 8858). ± 8860H:
165
Allium bisotunense R. M. FRITSCH;Melanocrommyum; Iran; prov. Kurdestan, culti-
vated in Sanandaj; (Sanandaj 8860). ± 8861H: Allium bisotunense R. M. FRITSCH;
Melanocrommyum; Iran; prov. Kurdestan, Kamiaran; (Sanandaj 8861). ± 8863H: Al-
lium sp.5; Acanthoprason; Iran; prov. Kurdestan, Maryvan, Mt. Dalani; (Sanandaj
8863). ± 8I: Allium tulipifolium LEDEB.; Melanocrommyum; Kazakhstan; 444500N,
0762100E; (GAT). ± 9: Allium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudo-
prason; Iran; prov. Kurdestan, Saral area; (HCAT). ± 9001I: Allium hooshidaryae
MASHAYEKHI,ZARRE &R.M.FRITSCH;Compactoprason; Iran; prov. Kurdestan (near
type location); (Sanandaj). ± 9002I: Allium aff. koelzii (WENDELBO)K.PERSS.&WEN-
DELBO;Pseudoprason; Iran; prov. Kurdestan; (Sanandaj). ± 9003I: Allium hae-
manthoides BOISS.&REUT.exREGEL s. str.; Acanthoprason; Iran; prov. Kurdestan;
(Herbarium Sanandaj). ± 9I: Allium protensum WENDELBO;Kaloprason; Kazakhstan;
444500N, 0762100E; (GAT). ± ISR09: Allium aff. nigrum L.; Melanocrommyum;Is-
rael; Mt. Meriron; (HUJ). ± JA009: Allium austroiranicum R. M. FRITSCH;Acantho-
prason; Iran; prov. Esfahan, 5 km E Daran; (GAT, photos). ± JA014: Allium austroir-
anicum R. M. FRITSCH;Acanthoprason; Iran; prov. Chaharmahal-Bakhtiari, Kuh-e
Kallar; (GAT, photos). ± JA015: Allium austroiranicum R. M. FRITSCH;Acanthopra-
son; Iran; prov. Chaharmahal-Bakhtiari, Kuh-e Kallar; (GAT, photos). ± JA022: Al-
lium koelzii (WENDELBO)K.PERSS.&WENDELBO;Pseudoprason; Iran; prov. Kerman-
shah, Noah Kuh near Kerend; (GAT, photos). ± JA026: Allium breviscapum STAPF;
Acanthoprason; Iran; prov. Hamedan, Kuh-e Alvand; (GAT, photos). ± JA029: Allium
aff. derderianum REGEL;Acanthoprason; Iran; prov. Qazvin, pass N Qazvin; (GAT,
photos). ± JA031: Allium aff. derderianum REGEL;Acanthoprason; Iran; prov. Qazvin,
second pass N Qazvin; (GAT, photos). ± JA039: Allium materculae BORDZ. subsp.
graveolens R. M. FRITSCH;Acanthoprason; Iran; 342721N, 0502574E; (GAT, photos). ±
JA047: Allium austroiranicum R. M. FRITSCH;Acanthoprason; Iran; 321432N,
0501312E; (GAT, photos). ± JA050: Allium austroiranicum R. M. FRITSCH;Acantho-
prason; Iran; 321675N, 0501339E; (GAT, photos). ± JA057: Allium ubipetrense R. M.
FRITSCH;Acanthoprason; Iran; prov. Kurdestan, 10 km W Sanandaj; (GAT, photos). ±
JA060: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran; 352322N, 0464340E;
(GAT, photos). ± JA066: Allium ubipetrense R. M. FRITSCH;Acanthoprason; Iran;
364447N, 0453213E; (GAT, photos). ± JA076: Allium aff. ubipetrense R. M. FRITSCH;
Acanthoprason; Iran; 364110N, 0484420E; (GAT, photos). ± JA079: Allium austroir-
anicum R. M. FRITSCH;Acanthoprason; Iran; prov. Esfahan, near Melmand; (GAT,
photos). ± JA082: Allium austroiranicum R. M. FRITSCH;Acanthoprason; Iran; prov.
Chaharmahal-Bakhtiari, pass Cheri; (GAT, photos). ± JA090: Allium jesdianum
BOISS.&BUHSE subsp. jesdianum;Megaloprason; Iran; 302747N, 0514103E; (GAT,
photos). ± JA097: Allium sp.1ªchlorotepalumº;Acanthoprason; Iran; 325507N,
0503357E; (GAT, photos). ± JA105: Allium ubipetrense R. M. FRITSCH;Acanthoprason;
Iran; 335246N, 0492719E; (GAT, photos). ± JA115: Allium stipitatum REGEL;Mega-
loprason; Iran; 331948N, 0495115E; (GAT, photos). ± OS: Allium schubertii ZUCC.;
Kaloprason; Botanical Garden OsnabruÈ ck, Germany; (OSBU). ± Un10: Allium sp.6
(aff. A. minutiflorum); Melanocrommyum ?; Iran; prov. Kurdestan, Saral area;
(HCAT). ± Un8: Allium sp.7 (aff. A. hooshidaryae); Compactoprason ?; Iran; prov.
Kurdestan, Saral area; (HCAT).
166
3. Results and Discussion of Molecular and Morphological Data
Addition of more species and accessions did not change the principal
genetic substructure of A. subg. Melanocrommyum shown by GURUSHIDZE
& al. 2008. The investigated species formed a basal grade characterized by
statistically well supported divisions, and a core clade (Fig. 1). The 7 dif-
ferent clusters in the core clade retained as monophyletic units in the ap-
plied (phenetic, cladistic, and model-based) algorithms. The phylogenetic
relationships among these clusters were not resolved, but the clusters were
more or less well-supported in all analyses. Also some groups within and
beside of these clusters got good statistical support despite their phyloge-
netic positions remained unclear. Altogether two groups (A, B) were re-
cognized in the grade, and 10 more groups (C ± L) in the clade. Thus,
clusters and main groups of the ITS-tree, as discussed by GURUSHIDZE & al.
2008, form a well applicable measure for the following presentation of ge-
netic relations.
3.1. Outgroups
Several species of the subgenera Anguinum, Butomissa, Caloscordum,
Porphyroprason, Reticulatobulbosa, and Vvedenskya were included in this
study as outgroup (Fig. 2 A). All accessions clustered only basally thus
confirming the monophyly of subg. Melanocrommyum. The position of
several species differs from the generic classification shown by FRIESEN &
al. 2006, which may well be caused by long branch attraction as already
discussed by GURUSHIDZE & al. 2008.
3.2. Basal Grade
3.2.1. sect. Longibidentata
Allium fetisowii and A. chychkanense from western Tianshan moun-
tain range were recently split off from sect. Acmopetala at sectional level
(FRITSCH 2009). They are morphologically rather similar (FRITSCH 2009), but
their molecular characters differ remarkably and show much diversity.
Analysis of additional material can perhaps clarify these relations.
These species are sister to all other clusters and groups (Fig. 1, Fig. 2
A) of subg. Melanocrommyum. Morphologically they are similar to some
members of sect. Acmopetala and were formerly included there (KHASSA-
NOV &FRITSCH 1994). According to our molecular data, sect. Acmopetala is
to be split in several parts positioned in different clusters of the core clade
(see chapters 3.5.4., 3.7.3 - 3.7.5, 3.8.1).
3.2.2. sect. Decipientia
Another species group, hitherto affiliated to sect. Melanocrommyum,
contains the basal A. decipiens from Crimea and Caucasus, A. viridulum
167
(which is exceptional in having granulous periclinal walls of seed coat
without verrucae, FRITSCH & al. 2006), A.chelotum from northern Iran, and
A. tulipifolium from South Siberia (Fig. 2 A). Here we should point to an
error in naming voucher specimens published in GURUSHIDZE & al. 2008
where A. viridulum and A. robustum were merged. It is perhaps a sign for a
phylogenetically old bifurcation that A. chelotum from northern Iran is
close to the geographically very distant A. tulipifolium alliance. Our mo-
lecular data do not support separation of the South Kazakh steppe plants
of A. tulipifolium as A. vakhtinae (nom. nud.) by SEISUMS 1994, ªholotypeº
in LE. Material of some possibly related species from Mongolia and wes-
tern China was not available for study.
The above discussed species are characterized by rather narrow (at
least 8 times longer than wide) and not undulate leaf blades, rose, lilac, or
carmine (not white) tepals, tuberculate (not glossy or nearly smooth) sur-
face of ovaries, and dominating S-like undulation of anticlinal walls of the
seed coat (FRITSCH & al. 2006). They occupy a taxonomic position far from
sect. Melanocrommyum (which constitutes the molecularly well char-
acterized ªgroup Cº, GURUSHIDZE & al. 2008, distantly positioned in the
core clade, see Fig. 1 and chapters 3.3.2. and 3.3.3.) and represent a well
separable section:
Allium sect. Decipientia(O
MELCZUK)R.M.FRITSCH,comb. & stat.
nov.
Basionym: Allium [sect. Melanocrommyum s. lat.] series Decipientia
OMELCZUK in Ukr. Bot. Zh. 19, 3: 71, 1962. Type: Allium decipiens FISCH.ex
SCHULT.&SCHULT.f.
The well-supported sistergroup relationship of A. robustum, another
Siberian taxon, to A. zergericum and the whole core clade would support
recognition as monotypic section, but we were not able yet to find out re-
liable morphological characters separating A. robustum from sect. Deci-
pientia.
Surprisingly, A. zergericum, which is sister to the large core clade,
belongs morphologically to the Alai-Fergan group of sect. Acmopetala (see
chapter 3.8.1). On the other hand, several characters of the seed coat
(FRITSCH & al. 2006) are much more similar to the species of sect. Long-
ibidentata than to the Alai-Fergan group. More material would be essen-
tial to verify this topology of A. robustum and A. zergericum.
3.3. Core Clade, Cluster 1
All other investigated species were positioned in the core clade. The
first cluster (Figs. 2 A, 2 B) consisted mainly of members of the sections
Melanocrommyum and Acanthoprason, both in the narrow sense, sect.
Porphyroprason, as well as a few species of sect. Regeloprason. Three main
groups and some taxa with unresolved relations were also included here.
168
3.3.1. sect. Pseudoprason in the strict sense
This well supported and rather homogeneous group occupies the most
basal position in cluster 1 (Fig. 2 A). It is divided in a small basal subgroup
(A. hooshidaryae as well as two undetermined taxa only known from single
herbarium specimens), plus A. koelzii. The first species shows some mor-
phological similarity to sect. Compactoprason (where it was affiliated hi-
therto) as well as to A. koelzii. All taxa positioned in this group stem from
the Northwest Iranian parts of Zagros mountain range. The peculiar
flower characters (FRITSCH & al. 2007) as well as the permanently green
ovaries are a good reason to hold A. koelzii separate at sectional level, but
additional material would be necessary to clarify the correct relations to
the other taxa. The singular separate position of acc. 9 may point to a
hybrid state of this herbarium specimen.
3.3.2. sect. Melanocrommyum in the narrow sense
Although we added 20 more accessions in comparison to GURUSHIDZE
& al. 2008, the species we investigated represent only a small part of up to
50 species affiliated to this section by different authors. Most basal and
sister to the other three groups and sect. Acanthoprason (Fig. 1, Fig. 2 A)
remained group ªCº of GURUSHIDZE & al. 2008 comprising at basal position
some East Mediterranean species, A. aschersonianum,A. schubertii (ex-
ceptional here as member of sect. Kaloprason which was also reported
from the Libyan Cyrenaica), and A. nigrum as neo-typified by SEISUMS
1998. These species own green coarse ovaries. Advanced are two subgroups
of closely related ªspecies-pairsº from Central to Southeast Europe and
the West Mediterranean characterized by smooth and at begin of anthesis
black ovaries. These are the pair A. multibulbosum (A. nigrum in the tra-
ditional sense) and A. atropurpureum, and as second pair A. orientale
(plants from Turkey) and A. cyrilli. All these species share one haplotype of
the trnL-trnF region of chloroplast DNA, and most taxa possess also ad-
ditional species-specific haplotypes (GURUSHIDZE & al. in press).
3.3.3. sect. Melanocrommyum in the wide sense
Other species traditionally affiliated to sect. Melanocrommyum group
together (Fig. 2 A) but their phylogenetic relations to sect. Acanthoprason
remained unresolved. They all share black ovaries. One small but well se-
parated subgroup comprises only A. bisotunense and A. keusgenii from
NW Iran, another subgroup A. noeÈanum, A. saralicum, and a hitherto un-
described species occuring also in NW Iran. A third subgroup comprises A.
ªorientaleº and A. ªnigrumº (from Israel, which do not belong to these
species in a strict sense), A. rothii (an East Mediterranean desert species),
and, more distantly related, A. chrysantherum sensu lato (to which another
169
170
Fig. 1. Schematic overview of the presentation of results in the chapter ª3. Results
and Discussionº
171
Fig. 2 A. Bayesian dendrogram, basal part: outgroups, basal grade, lower part of
cluster1 of core clade
172
Fig. 2 B. Bayesian dendrogram, second part: core clade, upper part of cluster 1 (sect. Acanthoprason).
173
Fig. 2 C. Bayesian dendrogram, third (submedian) part: core clade, clusters 2, 3, and 4.
174
Fig. 2 D. Bayesian dendrogram, fourth (upper median) part: core clade, clusters 5, 6, and lower part of cluster 7.
175
Fig. 2 E. Bayesian dendrogram, fifth (below upper) part: core clade, median part of cluster 7.
176
Fig. 2 F. Bayesian dendrogram, uppermost part: core clade, upper part of cluster 7.
not yet described species belongs) from Iran and Iraq, and the fourth sub-
group the rather inhomogeneous (but without geographic correlation) A.
cardiostemon from Armenia, Turkey, and Iran. Also the A. pseudo-
winklerianum alliance of sect. Regeloprason (see next chapter) belongs to
the fourth subgroup.
Our results strongly underline that several well separated taxa exist in
this area, and completely disagree with the taxonomic concept presented
by DE WILDE-DUYFJES 1976 for the A. orientale group. A careful morpho-
logical and taxonomic analysis of this group based on more accessions
from the eastern and south-eastern Mediterranean region seems urgently
necessary.
As long as the phylogenetic relationships of these different subgroups
are unresolved, all these species should remain members of sect. Melano-
crommyum. This holds also true for A. cardiostemon because our data ex-
clude the alternatively proposed affiliation to sect. Pseudoprason (GRE-
GORY & al. 1998). As far as investigated, the different trnL-trnF haplotypes
(GURUSHIDZE & al. in press) of all the species discussed in this chapter be-
long to one lineage confirming thus the affiliation to the same taxonomic
group.
3.3.4. Allium pseudowinklerianum Alliance of sect. Regeloprason
Only few species of sect. Regeloprason belong to the first cluster (Fig.
2 A). In the dendrogram of GURUSHIDZE & al. 2008 they formed jointly with
A. cardiostemon a well supported subgroup, and the separation became
even better in our dendrogram. Despite all species share pinkish-purple
flowers, their leaf characters differ strongly. A. pseudowinklerianum (Fer-
gan mountain range) has straight, broad, and smooth leaves, but A. so-
chense (Central Turkestan mountain range) has very narrow, curled leaves.
Both taxa are narrow endemics from Kyrgyzstan. The leaves of A. hissar-
icum (Central Tajikistan) are rough, linear, and moderately narrow. One
accession of this species was also included in this subgroup, but all other
accessions belong to cluster 7 (chapter 3.9.4., Fig. 2 D). Analyses of more
material would be essential to prove this position in cluster 1 as sister of A.
cardiostemon.
Rather small and stout plants developing more or less stinging tepals
after anthesis were traditionally affiliated to sect. Acanthoprason which is
typified by A. akaka. However, WENDELBO 1973 used this name in a very
broad sense subsuming also A. shelkovnikovii and A. haemanthoides
(WENDELBO 1971 accepted this name for a very variable taxon) as sub-
species under A. akaka. Because herbarium specimens named A. akaka
looked rather divers, detailed studies of living plants were undertaken
which showed that besides A. akaka, A. shelkovnikovii, and A. hae-
manthoides s. str., at least three more phenotypes of A. haemanthoides s.
177
lat. can be separated among Iranian accessions (FRITSCH 2008). They were
later described as new species A. zagricum,A. ubipetrense, and A. aus-
troiranicum (FRITSCH &ABBASI 2009). Occurrence of three different chlor-
oplast haplotypes of the trnL-trnF region (GURUSHIDZE & al. in press) in
this alliance underline the inhomogeneous structure of this group. Many
additional accessions collected in 2007 and 2008 could be added to the
analyses resulting in a better resolution and more detailed subdivision
than presented by GURUSHIDZE & al. 2008 as group D of cluster 1 (Fig. 1).
Our new molecular data (Fig. 2 B) strongly support recognition of several
taxa belonging to some statistically well supported groups. These groups
will be presented and discussed in the next chapters. Two accessions from
herbaria (7880H, 25222) were singled out between these groups. More ma-
terial of them would be essential to verify whether these accessions are
hybrids or represent new species. Because all the accessions forming this
cluster were collected in Iran and adjacent Turkey, we accept sect. Acan-
thoprason as solely Southwest Asian group. Our molecular data strongly
support the conclusion that this section has not only a centre of diversity in
this region but also undergoes an intense phase of differentiation and ra-
diation. Our data allow to predict that collection and molecular study of
much additional material of many morphotypes will probably improve the
circumscription and taxonomic position of many below discussed species,
but will also add more accessions occupying unclear positions. It remains
doubtful whether a stable infra-sectional classification of sect. Acantho-
prason can be constructed currently.
3.3.5. Allium austroiranicum Subgroup (sect. Acanthoprason)
The accessions clustering in this well separated but heterogeneous
group of sect. Acanthoprason refer to the second group of A. hae-
manthoides (according to FRITSCH 2008) distinguished by a very stout sta-
ture and very broad, shining, yellow-green leaves. Two Iranian species are
placed here, which differ by some flower characters and were therefore
affiliated to different taxonomic alliances in the conspectus (chapter 5).
Allium zagricum was collected in the province Lorestan, and A. austroir-
anicum from the provinces Fars and Chaharmahal-Bakhtiari is divided in
two geographical clusters. Three herbarium accessions (16799, 5013H,
8196H) of A. austroiranicum from Kurdestan were positioned in the A.
haemanthoides alliance (chapter 3.3.9.). These vouchers represent variable
populations typical for A. austroiranicum, but morphologically these
plants could alternatively represent segregating populations of a hybrid of
A. haemanthoides with one of the species having dark-colored filament
tips. Somewhat surprising was the insertion of one accession of A. sp.1
ªchlorotepalumº (343H). Other accessions of this still undescribed species
from Central Iran form a well supported subgroup close to the A. grave-
178
olens subgroup (see chapter 3.3.8.). One accession (6632) of A. upipetrense
(see chapter 3.3.6.) also included here may represent a hybrid plant having
homogenized ITS sequences.
3.3.6. Allium ubipetrense Alliance (sect. Acanthoprason)
This well supported group of sect. Acanthoprason includes mainly A.
ubipetrense from eastern parts of province Markazi accompanied by one
accession of A. graveolens and a still undescribed species. More surprising
was the insertion of A. ubipetrense accessions in other groups of sect.
Acanthoprason discussed in the chapters 3.3.5., 3.3.8., 3.3.9., and 3.3.11.
This widespread molecular affiliation is difficult to explain, because we
did not find yet any correlation of morphological character states with
these topologies, which could support recognition of several species. The
assumption that A. ubipetrense represents a morphotype which developed
parallel in several evolutionary lines seems us more probable than to ac-
cept many cases of recent hybridization events of A. ubipetrense, which
might have led to different homogenized sequences responsible for posi-
tions close to the respective parents. All these different accessions were
collected in Northwest Iran in the mountain ranges South of Lake Or-
umiyeh.
3.3.7. Allium derderianum Alliance (sect. Acanthoprason)
Only plants collected in Alborz mountain range constitute the A. der-
derianum subgroup (acc. 34075 could not be traced and re-determined
yet). This subgroup is homogeneous, despite morphologically divers ac-
cessions (FRITSCH 2008: 60 f.) were included. Also a morphologically still
more deviating, recently recognized taxon (A. sp.9) only known yet from
the area of Mt. Alamut, does not show molecular differences. WENDELBO
1971 included in A. derderianum also A. austroiranicum. These plants are
similar only at a first glance and occur in mountain ranges in Central Iran.
They are only distantly related to A. derderianum and belong to a sub-
group discussed in chapter 3.3.5.
Basal to this clade and rather well supported, the type accession of A.
moderense (6616) is positioned. A second accession of this species (6629)
occupies an unresolved position. These positions are unexpected because
this species belongs morphologically to the A. colchicifolium alliance of
sect. Melanocrommyum (FRITSCH &ABBASI 2009). The molecular position
may indicate that A. derderianum or another species of sect. Acanthopra-
son was involved in the evolution of A. moderense.
3.3.8. Allium graveolens Subgroup (sect. Acanthoprason)
This weakly supported Iranian subgroup is composed of rather differ-
ent taxa showing a weak geographic correlation. Most accessions belong to
179
the newly described A. materculae subsp. graveolens occurring in the
provinces Markazi and Esfahan (FRITSCH &ABBASI 2009). This subspecies
is morphologically very similar to the typical subspecies (see chapter
3.3.11) which has a more northern distribution. Despite this similarity, our
molecular data strongly favour to accept this taxon at species level.
A l l i u m graveolens (R. M. FRITSCH)R.M.FRITSCH,comb. & stat. nov.
Basionym: Allium materculae subsp. graveolens R.M. FRITSCH in
Rostaniha 9 Suppl. 2: 31, 2009.
Also three accessions of A. ubipetrense (discussed in chapter 3.3.6.)
from the pre-mountains along the borderlines of the provinces Zanjan and
Hamedan were located in this subgroup. Also the only accession (6487) of
A. hamedanense, a peculiar species only known from the mountains south
of Hamedan (FRITSCH &ABBASI 2009), another accession (11929) of A. sp.2
from the Northwest of province Kurdestan (discussed in chapter 3.3.11.),
and an aberrant, purple-flowering accession (64855) very similar to A.
akaka from the western edge of province Zanjan are included. Finally we
have to mention the well supported subclade of A. sp.1ªchlorotepalumº
occurring in the northwestern corner of province Esfahan. (One herbarium
voucher of this taxon included in the A. austroiranicum subgroup, chapter
3.3.5., is morphologically different and may represent a hybrid.) Allium
sp.1ªchlorotepalumº is morphologically characterized by initially black
and shining ovaries like the taxa discussed in chapter 3.3.3. On the other
hand, also A. hamedanense owns shining (but not black) ovaries, and the
ovaries of A. graveolens are faintly glossy. Therefore we can state that more
or less glossy ovaries characterize this subgroup.
3.3.9. Allium haemanthoides Alliance (sect. Acanthoprason)
Also this subgroup of group ªDº (Fig. 1) of Gurushidze & al. 2008 was
strongly enlarged by addition of many acessions and herbarium vouchers.
It became a well supported, mainly geographically characterized, and
morphologically and taxonomically heterogeneous subgroup. It comprises
all investigated accessions of A. haemanthoides in the strict sense, several
accessions of A. ubipetrense (discussed in chapter 3.3.6.), and three hi-
therto undescribed new taxa. Tiny plants with very short, nearly sub-
orbicular leaves and purplish tepals from the provinces Azarbeijan and
Kurdestan south of Lake Orumiyeh were preliminary named A. sp.3, but
plants of A. sp.5 from central parts of province Kurdestan are mostly lar-
ger and possess large, broadly lanceolate leaves and multiflorous, dense
inflorescences. They are candidates for being described as new taxa. On
the other hand, A. sp.10 (acc. 2432) is perhaps only a variant of A. hae-
manthoides having narrow leaves and dark-purple flowers. Investigation
of additional living material would be essential. Also three accessions of A.
180
austroiranicum from province Kurdestan were affiliated here. They were
already discussed in chapter 3.3.5.
3.3.10. Allium breviscapum Alliance (sect. Acanthoprason)
Four accessions of this local endemic from Alvand massif near Hame-
dan in northwestern Iran could be analyzed. They are the well supported
sister group to the A. haemanthoides alliance (discussed in the chapter
before) although A. breviscapum is morphologically more similar to A.
derderianum. This molecular position reflects perhaps an ancient hybrid
offspring of A. breviscapum with a taxon of the A. haemanthoides alliance
and possibly A. derderianum as parents.
3.3.11. sect. Acanthoprason in the strictest sense
The A. akaka alliance was contradictory treated by different authors.
Because only the geographical region (the Iranian province Ghilan) but not
the exact collection site of the type specimen is known, Iranian material
from that area fitting well the original description was studied (FRITSCH
2008) and designated as epitype (FRITSCH &ABBASI 2009). These epitypic
plants (acc. 6398) and more accessions from the Iranian provinces Ardabil,
West and East Azarbeijan form one well supported subgroup. Our material
allows to conclude that the distribution of A. akaka reaches the Tabriz
area in the West. Additional material of acc. 17362 would be necessary to
verify whether determination as A. aff. ubipetrense is correct. Two acces-
sion of A. aff. shelkovnikovii (34268, 34269) from Tabriz town area clus-
tered also in this subgroup, but two more accessions of this species (6404,
6405), collected about 20 km apart from Tabriz, clustered in the second
subgroup discussed below. This taxon differs morphologically well from A.
akaka s. str. (FRITSCH 2008), but not much from A. egorovae (see below),
and therefore affiliation of all accessions (also of acc. 70664 named A. aff.
egorovae) to the second subgroup was expected. We cannot explain why
they were included in the first subgroup.
Rather closely related is the well supported second subgroup com-
prising A. materculae s. str. (from North Iran and adjacent republics, see
chapter 3.3.8.), a species preliminary named A. shelkovnikovii (see discus-
sion in FRITSCH 2008) collected North of Tabriz, and plants from eastern
Turkey and West of Lake Orumiyeh hitherto also named A. akaka and
discussed below. Several herbarium specimens from the mountains near
the Araks valley (border to republic Azerbaijan) preliminarily named A.
egorovae and another still undescribed species (A. sp.2) known yet only
from fruiting herbarium specimens from the province Azarbeijan belong
also here. Beside A. materculae, these taxa differ from A. akaka s. str. by
narrow filaments with dark tips and the different forms of tepals. Inclu-
sion of acc. 3940, named with some doubt A. zagricum despite it was col-
181
lected in the province Zanjan and not in Lorestan, may indicate that this
accession also belongs to A. sp. 2. However, another accession preliminary
named in this way was included in the A. graveolens subgroup (chapter
3.3.8.).
The East Turkish ± Northwest Iranian ªA. akakaº plants are divers.
Turkish plants differ remarkably by narrow, bluish-pinkish tepals. Like
some Iranian plants they fit better the description and the geographic re-
gion of A. latifolium JAUB.&SPACH. Unfortunately, his name is a later
homonym and thus illegitimate. Therefore a new name is necessary:
Allium jau bertii R. M. FRITSCH,nom. novum =A. latifolium JAUB.&
SPACH, Ill. pl. orient. 2: t. 103, 1846, non A. latifolium W. Y OUNG, Cat. arbr.
Amer. 28, 1783, necque A. latifolium GILIB., Excerc. phyt. 2: 470, 1792.
Analysis of living plants in addition to the herbarium specimens
would be urgently necessary to characterize all these taxa more in detail
and to apply the names correctly.
3.4. Core Clade, Cluster 2
Three well separated groups recognized by GURUSHIDZE & al. 2008
were also present in our data, but as single units without further resolution
of phylogenetic connections (Fig. 2 C).
3.4.1. Allium suworowii Alliance
This most basal and well supported subgroup of cluster 2 in the den-
drogram of GURUSHIDZE & al. 2008 contains only one species morphologi-
cally completely different from the other species of cluster 2. A. suworowii
is widely distributed (from western and central Tian Shan mountain ran-
ges up to the western pre-mountains of western Hissar mountain range,
northern Afghanistan, and the western Pamir area of Tajikistan). There-
fore it is not surprising that our accessions are also genetically divers.
Perhaps addition of much more accessions would allow to find out whether
morphologically and geographically well circumscribed, infraspecific
groups exist.
A. suworowii was most often affiliated to sect. Megaloprason in the
widest sense. Only KHASSANOV &FRITSCH 1994 selected it as type species of
subsect. Spiralitunicata (sect. Acmopetala) including also A. fibriferum, A.
insufficiens, and A. vvedenskyanum. Our current molecular data support
closer relations of A. suworowii to sect. Megaloprason in the strictest sense
which were also confirmed by possessing identical chloroplast haplotypes
(GURUSHIDZE & al. in press). A. insufficiens and A. vvedenskyanum belong
to other groups (see chapters 3.6.1. and 3.7.3.), and A. fibriferum could not
be analyzed. Thus, subsect. Spiralitunicata contains probably only one or
two species and should be affiliated under sect. Megaloprason.
182
3.4.2. Allium regelii of sect. Regeloprason
Two accessions of this morphologically striking species from Turkme-
nistan, Southwest Afghanistan, and adjacent areas of Iran were available.
They clustered jointly with the above discussed A. brachyscapum (chapter
3.3.10) alliance but formed a well supported sub-clade (Fig. 2 C). This
points to a rather isolated position of A. regelii in sect. Regeloprason.
3.4.3. subsect. Humilicognata
This small but well supported clade occupies a middle position in the
second clade of GURUSHIDZE & al. 2008 (Fig. 2 C). It consists of two well
supported alliances: A. regelii (discussed below), and a group of three
species possessing tepals reflexed after anthesis and rather long, patent
filaments. Therefore WENDELBO 1971 affiliated the two species he knew to
sect. Megaloprason. The first species A. brachyscapum is a small endemic
from the alpine belt of Koppeh Dagh mountain range in Turkmenistan and
Iran. Much taller but otherwise similar plants known from semi-desert
plains and hills in North and Central Iran were separated some time ago at
species level as A. assadii (SEISUMS 2000). The third species A. scotostemon
differs remarkably by much narrower leaves, broader tepals, and purple
filaments. It is a narrow endemic of Central Alborz mountain range in
North Iran.
These taxa do not belong to sect. Acmopetala subsect. Stellata (pro-
posed by FRITSCH & al. 2002 for A. assadii) but to sect. Megaloprason in the
strictest sense (see chapter 3.6.1.) because A. brachyscapum and A. ro-
senbachianum share identical chloroplast haplotypes (GURUSHIDZE & al. in
press). They are affiliated to a new subsection:
Allium [sect. Megaloprason s. str.] s u b s e c t . Humilicognata R. M.
FRITSCH,subsect. nova
Type species: A. brachyscapum VVED.
D i a g n o s i s: Scapus brevis flexuosus folia plerumque excedens vel
subelongatus strictus. Inflorescentia plus minusque hemisphaerica, den-
siuscula. Perigonium stellatum; tepala basi libera, mox reflexa et plus
minusque spiraliter contorta, filamenta carnosa subulata. Ovarium stipi-
tatum, verruculosum.
Scapes either short and flexuous much shorter than leaves or some-
what longer and straight. Inflorescence +hemispherical, rather dense.
Flowers star-like, tepals free near base, later reflexed and spirally con-
torted. Filaments fleshy, subulate. Ovary stalked, with a rough surface.
3.4.4. sect. Asteroprason
The largest and most advanced group of the second cluster (in the
dendrogram of GURUSHIDZE & al. 2008, Fig. 1) comprised the well sepa-
183
184
rated alliances of the important ornamental A. cristophii,ofA. elburzense,
and the few investigated accessions of A. helicophyllum. The separation of
A. ellisii and A. kuhsorkhense was not resolved (Fig. 2 C).
All the species included here are morphologically well separated one
from another but share loose globular inflorescences and star-like patent
tepals. A. cristophii is naturally growing in the Turkmen and Iranian
Koppeh Dagh mountain range and in the easternmost parts of Alborz
mountain range. It shows remarkable infraspecific diversity (FRITSCH 2008)
which may explain the certain degree of inhomogeneity of its subgroup.
The taxonomic still unclear A. ellisii is a narrow endemic of Binalud and
eastern Koppeh Dagh mountain ranges, A. elburzense and the perhaps
conspecific A. pseudobodeamum occur in the central Elburz mountain
range, A. helicophyllum also in the Turkmen and Iranian Koppeh Dagh
mountain range, and A. kuhsorkhense in the lower mountain massifs south
of Mashhad in the province Khorassan.
All available molecular data (GURUSHIDZE & al. 2008, GURUSHIDZE &
al. in press) gave clear evidence that traditional affiliation of these species
neither to sect. Acanthoprason (WENDELBO 1971) nor to sect. Kaloprason
(GREGORY & al. 1998) is correct. They represent a new section:
Allium sect. Asteroprason R. M. FRITSCH,sect. nova
Type species: A. elburzense WENDELBO
D i a g n o s i s: Scapus subelongatus conicus flexuosus brevior quam fo-
lia. Inflorescentia semiglobosa usque subglobosa, laxa usque subcompacta.
Flores plus minusque plana stellata, tepala patentia vel demum recurvata
post anthesin longitudinaliter convoluta.
Scape conical and flexuous, somewhat elongated but shorter than leaf
blades. Inflorescence semiglobose to subglobose, loose to somewhat con-
tracted. Flowers more or less flat star-like, tepals patent or later somewhat
recurved, after anthesis longitudinally enrolled.
Allium subsect. Asteroprason R. M. FRITSCH,subsect. nova
Type species: A. elburzense WENDELBO
D i a g n o s i s: Statura plantae compacta, folia glabra, tepala post an-
thesin longitudinaliter convoluta non autem aculeata.
Plants of compact stature, leaves glabrous, tepals after anthesis long-
itudinally enrolled but not prickly.
Species of the typical subsection Asteroprason are small plants having
smooth leaves and longitudinally enrolled, crumpled, not prickly tepals in
the fruiting stage. Only gradually different are the larger stature and hairy
leaves of A. cristophii, but the tepals remain straight and surround later
the capsules as stiff prickles (subsect. Christophiana). Our material named
A. ellisii belongs also to the latter subsection, but other morphotypes fit-
ting also the description of A. ellisii could not be studied yet as living
plants. The smallest species is A. monophyllum, a rare taxon distributed in
the alpine parts of Koppe Dagh mountain range. It is morphologically most
185
similar to the species of sect. Asteroprason, but no material was available
to prove this affiliation by molecular data.
3.5. Core Clade, Cluster 3
Addition of more species and accessions led to a clear substructure of
this cluster (Fig. 2 C) compared with the weak relations reported by GUR-
USHIDZE & al. 2008. Additionally, the haplotypes of trnL-trnF sequences of
the investigated species of all included groups belong to the same lineage
(GURUSHIDZE & al. in press) which confirmed the close relationship of all
members of cluster 3.
3.5.1. sect. Verticillata
Only two species belong to this section, A. verticillatum from western
parts of Tajikistan and Uzbekistan, and A. viridiflorum, a narrow endemic
from the northwestern edge of Fergan depression in Kyrgyzstan. Both are
rather small and inconspicuous in the general appearance but share an
unique character: the leaf blades are longitudinally divided into several
narrow, thread-like tips appearing like single leaves. It was unexpected
that A. verticillatum belongs to two well separated and distant molecular
subgroups positioned basally in cluster 3: most basal the accessions from
West of 688longitude, and slightly more advanced the investigated acces-
sions from East of this longitude (Fig. 2 C). Both subgroups of A. verti-
cillatum show high genetic diversity. Their insertion in cluster 3 may re-
present long branch attraction, as discussed already by GURUSHIDZE & al.
2008. The unique structure of the seed testa cells (flat granular periclinal
and finely undulated anticlinal cell walls, FRITSCH & al. 2006) disagrees
completely to all other species of cluster 3.
The only investigated accession of A. viridiflorum occupies a well
supported position in cluster 6 (see chapter 3.8.1., Fig. 2 D) among mor-
phologically dissimilar species from the same geographic region. This po-
sition may perhaps point to an ancient hybridization event. Also the dif-
ferent ornamentation of seed testa cells (FRITSCH & al. 2006) could be
caused by hybridization. More accessions of both species would be essen-
tial to verify these positions in the ITS dendrogram.
3.5.2. Allium cupuliferum Alliance, sect. Regeloprason
Sister to A. taeniopetalum and the A. balkhanicum alliance is the or-
namental species A. cupuliferum (endemic in the Aktau and Nuratau
massifs up to the northern slopes of Turkestan mountain range in Uzbeki-
stan). Although it shares several morphological characters with the species
of the A. balkhanicum alliance, it is only distantly related (Fig. 2 C) and
constitutes a well separated subgroup. A rather large genetic distance be-
tween acc. 6196 from Aktau massif and the other accessions from Nuratau
massif (West Uzbekistan) may reflect long-standing geographic separation.
3.5.3. Allium balkhanicum Alliance, sect. Regeloprason
A. isakulii was described to comprise three geographically separated
subspecies (FRITSCH 2000). However, the molecular analyses of GURUSHIDZE
& al. 2008 did not confirm these relations: the only accession of the typical
subspecies was positioned jointly with the A. pseudowinklerianum alli-
ance in cluster 1, and the other subspecies were included in cluster 3 where
they formed one subgroup of group F. Our new data (Fig. 2 C) do not con-
firm these different positions. After re-sequencing, the only accession of
A. isakulii s. str., a narrow endemic of Nuratau and Mogoltau massifs in
Uzbekistan and Tajikistan, occupied a sister position to the only accession
of A. iliense s. str., distributed in the hilly semi-deserts nearby the Ili river
in South Kazakhstan. Both species are sister to subsp. balkhanicum. An-
other accession most similar to subsp. isakulii from a newly detected site
in Kurama mountain range in Uzbekistan became sister to sect. Stellata.
Large molecular distances and separate positions of subsp. isakulii, subsp.
balkhanicum (a narrow endemic of Great Balkhan massif in Turkmeni-
stan), subsp. subkopetdagense (from lower mountains of Turkmen Central
Kopetdag mountain range), and the Central Iranian A. cathodicarpum
underlined, that all these taxa should be accepted at species level:
Allium subkopetdagense (R. M. FRITSCH &F.O.KHASS.) R. M.
FRITSCH,comb. & stat. nov.
Basionym: Allium isakulii R. M. FRITSCH &F.O.KHASS. subsp. sub-
kopetdagense R. M. FRITSCH &F.O.KHASS.inO.A.ASHURMETOV & al.
(eds.), Plant Life S-W. Central Asia, Tashkent: 65, plate 1H, 2000.
Allium balkhanicum (R. M. FRITSCH &F.O.KHASS.) R. M. FRITSCH,
comb. & stat. nov.,
Basionym: Allium isakulii R. M. FRITSCH &F.O.KHASS. subsp. bal-
khanicum R. M. FRITSCH &F.O.KHASS.inO.A.ASHURMETOV & al. (eds.),
Plant Life S-W. Central Asia, Tashkent: 65, plate 1I, 2000.
3.5.4. sect. Stellata
The most advanced position in cluster 3 (Fig. 2 C) is occupied by three
subspecies of A. taeniopetalum. They share a specific anatomy of nectaries
(GURUSHIDZE & al. 2008) with the A. balkhanicum alliance (chapter 3.5.3.).
The phylogenetic relations of the subspecies were not resolved. The rather
large distances between them may reflect geographic diversity, but already
inside of subsp. taeniopetalum from Uzbekistan the diversity is remark-
able (acc. 5244 and 5249 were collected in the Aktau massif not far from
one another, and acc. 6531 in the western Turkestan mountain range).
Subspecies turakulovii occurs in the Eastern Turkestan (adjacent Kyrgyz-
stan) mountain range. It remains open whether the analysis of topo-typical
plants of subsp. mogoltavicum from Mogoltau massif (North Tajikistan, all
186
investigated accessions stem from Chatkal mountain range in Uzbekistan
200 km apart) could enhance the molecular differentiation.
Although the lax globular inflorescence of A. taeniopetalum is more
similar to sect. Megaloprason s. str., this species shares stellate tepals with
some species of sect Acmopetala s. lat. Unfortunately, the phylogenetic re-
lations to these sections remained as unresolved as shown by GURUSHIDZE
& al. 2008. However, A. taeniopetalum shares identical haplotypes of trnL-
trnF sequences (GURUSHIDZE & al. in press) with sect. Kaloprason, subsect.
Keratoprason (chapter 3.9.6.), and A. jesdianum (see chapter 3.9.9.) but not
with the sections Megaloprason and Acmopetala in the strict sense.
Therefore recognition at sectional level seems appropriate:
Allium sect. Stellata (F. O. KHASS. & R. M. FRITSCH)R.M.FRITSCH,
comb. & stat. nov.
Basionym:Allium [sect. Acmopetala] subsect. Stellata F. O. KHASS.&
R. M. FRITSCH, Linzer Biol. BeitraÈ ge 26: 976, 1994. Type species: A. taenio-
petalum VVED.
3.6. Core Clade, Cluster 4
This cluster was divided in three well-supported groups with un-
resolved phylogenetic relations (Fig. 2 C). Two groups contain taxa of the
A. rosenbachianum alliance (in the strict sense, not A. rosenbachianum of
gardens = A. rosenorum which belongs to cluster 7). Species of sect. Re-
geloprason constitute the third group.
3.6.1. sect. Megaloprason in the strictest sense
One group comprises several accessions of A. rosenbachianum s. str.,
as well as one accession each of A. chitralicum (which will be discussed in
chapter 3.9.7.) and A. schugnanicum. The latter species, a narrow endemic
from shady montane slopes of western Pamir mountain massifs, is mor-
phologically very close to A. rosenbachianum which occurs on shady slopes
and in submontane broad-leafed shrub and forest associations in southern
and southeastern Tajikistan. Analyses of more accessions would be neces-
sary in order to decide whether A. schugnanicum is not more than a mon-
tane variant of A. rosenbachianum.W
ENDELBO 1971 used the name A. ro-
senbachianum for A. jesdianum subsp. angustitepalum (see chapter 3.9.9.).
It is surprising that all analyzed accessions of A. rosenbachianum
subsp. kwakense (which shares the area of distribution with subsp. ro-
senbachianum) belong to another group which is further subdivided (Fig. 2
C). The clear molecular distance to A. rosenbachianum favor to recognize
this taxon at species level:
Allium kwakense (R. M. FRITSCH)R.M.FRITSCH,comb. & stat. nov.
Basionym: A. rosenbachianum REGEL subsp. kwakense R. M. FRITSCH
in Candollea 48: 419, 1993.
187
The only analyzed accession of A. insufficiens, distributed in south-
eastern Tajikistan and adjacent parts of Afghanistan, is sister to A. kwa-
kense. The long distance of A. insufficiens to all other species of cluster 4
implies recognition perhaps as monotypic subsection. Analysis of more
accessions would be essential to confirm this position in sect. Mega-
loprason s. str.
3.6.2. Allium intradarvazicum Alliance
A. intradarvazicum is a recently described species (FRITSCH 2009) only
known from the Khumbov valley in central Darvaz mountain range, Taji-
kistan. Surprisingly, it forms the third group of cluster 4 (Fig. 2 C) together
with several accessions of the morphologically rather dissimilar A. dar-
wasicum from southern Darvaz, Vakhsh, and Hissar mountain ranges.
Other accessions of the latter species were located in cluster 7, and possi-
ble reason for the splitting will be discussed in more detail in chapter 3.9.4.
The position of A. intradarvazicum and A. darwasicum in cluster 4 was
unexpected because the morphologically most similar species of sect. Re-
geloprason,A. winklerianum and A. hissaricum, were also included in
cluster 7 group J (Fig. 2 D) of the ITS-based dendrogram of GURUSHIDZE &
al. 2008.
3.7. Core Clade, Cluster 5
Addition of many accessions did not much improve the resolution of
the phylogenetic tree presented by GURUSHIDZE & al. 2008. The phyloge-
netic relationships of the well supported five larger groups remained still
unresolved (Fig. 2 D). As far as analyzed by GURUSHIDZE & al. in press, the
trnL-trnF sequences of all members of the clusters 5 and 6 belong to one
lineage which underlines a rather high degree of general relationship.
3.7.1. sect. Miniprason
A. karataviense, the only species of this section, occupies a wide nat-
ural area of distribution from Kazakh Syrdarya Karatau mountain range
and Chu-Ili-mountains in the North up to the Panj valley along the border
between Tajikistan and Afghanistan (and perhaps up to Hindu Kush range
in eastern Afghanistan and the Kugitang massif near the northeastern
edge of Turkmenistan) in the South. It was somewhat surprising that both
accessions from Panj valley (6133, 6147) are well supported segregates
(Fig. 2 D) despite they do not differ morphologically. On the other hand,
red flowering plants from Chatkal mountain range (accs. 2989, 5040, 5888,
6366) are part of the homogeneous main group (completely homogeneous
also in the NJ-dendrogram, not shown), but material of the newly de-
scribed subsp. henrikii RUKSANS were not available for study. As far as
known hitherto, A. karataviense is the only species of subg. Melano-
188
crommyum having 2n = 18 chromosomes. All other species have 2n = 16 or
multiples of it (FRITSCH &ASTANOVA 1998).
3.7.2. sect. Brevicaule
Only single accessions of two (out of three) species could be analyzed.
A. sergii (a rare endemic species from central Syrdarya Karatau in Ka-
zakhstan) occupies a phylogenetically unresolved and very distant position
in cluster 5 (Fig. 2 D). It might be another case of long branch attraction
already discussed by GURUSHIDZE & al. 2008 for A. verticillatum.
The second species A. chitralicum occurs in the higher mountains of
East Afghanistan, Pakistan, and Tajikistan. The only accession in-
vestigated (from Tajikistan) was positioned in cluster 4 among A. ro-
senbachianum (sect. Megaloprason s. str., see chapter 3.6.1., Fig. 2 C). Thus
both species were placed beside morphologically dissimilar species from
the same geographical regions. More material, also of the third species A.
eugenii, a narrow endemic of Great Balkhan massif in Turkmenistan,
seems essential to resolve this discrepancy. The current data would also be
compatible with the assumption that sect. Brevicaule is perhaps only an
artificial aggregate of morphologically relatively similar but only very
distantly related species.
3.7.3. Allium dasyphyllum Alliance
Another well supported group with unresolved relations (Fig. 2 D)
consists of the only analyzed accessions of two narrow endemics from the
northern limit of western Tianshan mountain range, A. dasyphyllum (from
the Kyrgyz Talassian Alatau) and A. vvedenskyanum (from the Kazakh
Chu-Ili-mountains). Formerly they were affiliated to different subsections
of sect. Acmopetala discussed in chapters 3.5.4. and 3.8.1. These species
are morphologically well separated from another as well as from A.
tschimganicum and the other species included in cluster 5. Analysis of
more accessions seems essential prior to recognition as separate taxonomic
subgroup.
3.7.4 subsect. Pharmakoprason
A. tschimganicum (= A. motor, see chapter 3.13. below), a narrow Uz-
bek endemic from the Chatkal mountain range south and east of Tashkent,
forms a homogeneous and well separated alliance in the ITS dendrogram
(Fig. 2 D). It differs from the morphologically most similar species A. se-
vertzovioides and other species discussed in the next chapter also by an-
other nectary anatomy (GURUSHIDZE & al. 2008). Additionally, only this
species is used as wild collected vegetable having remarkable medicinal
properties (KEUSGEN & al. 2006) and contains sulphurpyrrols (GURUSHIDZE
189
2008). Therefore it should be accepted as monotypic new healthy onion
subsection:
Allium [sect. Acmopetala] subsect. Pharmakoprason R. M.
FRITSCH,subsect. nova
Type species: A. tschimganicum O. FEDTSCH., quoad lectotypum (= A.
motor KAMELIN &LEVICHEV)
D i a g n o s i s: Scapus valde elongatus proportione tenuis, strictus, folia
plerumque excedens. Folium vaginatum extimum tenue sine colore. In-
florescentia sphaerica, densa. Perigonium stellatum; tepala basi libera,
mox reflexa et plus minusque contorta. Filamenta interiora dilata basi
utrinque triangulato-dentata. Ovarium verruculosum.
Scape very long and rather slender, straight, much longer than the leaves.
Outer sheath leaf thin, colorless. Inflorescence spherical, dense. Flowers star-
like, tepals basally free, later reflexed and +contorted. Bases of inner
filaments widened, with a triangular tooth at both sides. Ovary rough.
3.7.5. Western Tianshan Geographical Clade of sect. Acmopetala
This group is characterized by a specific type of nectary anatomy and
was named ªgroup Hº by GURUSHIDZE & al. 2008. It consists of two well
supported subgroups but with unresolved phylogenetic relations. All spe-
cies included here are endemics of different parts of the western Tianshan
mountain range in Uzbekistan, Kyrgyzstan, and Kazakhstan. The smaller
subgroup (Fig. 2 D) contains only one molecularly inhomogeneous species,
A. costatovaginatum. It possesses a hard, ribbed, and long-lasting basal
sheath leaf and was hitherto affiliated to subsect. Durovaginata like A.
severtzovioides. The latter species forms the second subgroup together
with A. tashkenticum and A. sewerzowii, belonging to subsect. Inornatae,
plus A. saposhnikovii, the only species of subsect. Albidiflora. It remained
unclear whether the accession 6364 of A. saposhnikovii from northern
slopes is a hybrid or deserves taxonomic recognition, whereas plants from
southern slopes (5780, 5783) share all molecular character with the other
species. Also A. tashkenticum is morphologically and molecularly some-
what divers. This union of several morphologically differing species in one
molecularly rather homogeneous subgroup was not expected. It is difficult
to predict whether the addition of more accessions of A. sewerzowii and
other missing taxa (A. rudolfii,A. tokaliense) could result in a better re-
solution in separate taxa. Alternatively, subsections Inornatae and Albidi-
flora could be abandoned and all species inserted in subsect. Dur-
ovaginata.
3.8. Core Clade, Cluster 6
This small group is the sister to the species-rich cluster 7 of GURUSH-
IDZE & al. 2008 and consists mainly of monotypic, well supported groups
190
having mostly badly resolved phylogenetic relations (Fig. 2 D). Most spe-
cies included share a general morphological similarity and belong to sect.
Acmopetala. Their trnL-trnF haplotypes belong to the same lineage as
those of cluster 5 (GURUSHIDZE & al. in press). However, all studied mem-
bers of cluster 6 share a specific type of nectary anatomy (GURUSHIDZE &
al. 2008) which underlines a close relationship.
3.8.1. Alai-Fergan Geographical Clade of sect. Acmopetala
This small cluster comprises five species occurring in a rather re-
stricted geographic area in Central Asia which belongs partly to Kyrgyz-
stan, partly to Uzbekistan. Most basal are A. schachimardanicum (a nar-
row endemic of Alai mountain range South of Fergana town) and A. do-
decadontum (endemic of the central Chatkal mountain range and the Sar-
ychilek area). Hitherto both species were included in subsect. Long-
ibidentata, but our molecular data deny that affiliation (see chapter 3.2.1.).
Addition of more accessions would be essential to verify their current
phylogenetic position and to decide whether their preliminary taxonomic
affiliation to subsect. Acmopetala discussed below is a good solution.
Also the only investigated accession of A. viridiflorum, sect. Verti-
cillata (see chapter 3.5.1.) was positioned in cluster 6 but shows unresolved
relations.
The advanced position in cluster 6 is occupied by two subgroups. One
of them comprises A. aflatunense in the strictest sense and A. arkitense,
two morphologically strongly differing endemics of Sarychilek area, plus
A. alaicum, a rare endemic from the northern slopes of Alai mountain
range. Most surprising was the inclusion of A. aflatunense which shows
only small differences of quantitative leaf, scape, and flower characters to
A. stipitatum variants without haired leaves positioned in cluster 7 (see
chapter 3.9.11.). Further detailed morphological analyses can perhaps
trace additional reliable differences to the A. stipitatum alliance. No less
surprising was the position of A. zergericum (endemic of the Fergan
mountain range) as sister to the core clade (discussed in chapter 3.2.2.)
because this taxon is morphologically close to A. arkitense and A. alaicum
(FRITSCH & al. 2002).
The second subgroup comprises only A. backhousianum, an endemic
of the northern slopes of Alai mountain range. It differs morphologically
remarkably from all other species belonging to cluster 6. This taxonomic
distance is also expressed by the molecular position as subgroup of its own
(Fig. 2 D). On the other hand, molecular and anatomical data underline a
far-reaching phylogenetic homogeneity of cluster 6. Therefore the A. afla-
tunense morphotype is probably another instance for the assumption dis-
cussed by GURUSHIDZE & al. 2008 in the cases of A. stipitatum and A. ro-
senorum (see chapter 3.9.11.), that independent evolutionary lineages seem
191
to have converged on similar phenotypes. Then, A. aflatunense s. str.
should be affiliated to subsect. Acmopetala, but other similar phenotypes
remain under A. stipitatum, sect. Procerallium.
The above discussed four groups (chapters 3.7.3 ± 3.7.5., 3.8.1) re-
present the majority of species hitherto affiliated to sect. Acmopetala after
splitting off several species as new sections Longibidentata (chapter 3.2.1.)
and Stellata (chapter 3.5.4.), and transferring two species to sect. Mega-
loprason (subsect. Spiralitunicata, chapter 3.4.1.). The species remaining
in sect. Acmopetala share trnL-trnF haplotypes belonging exclusively to
lineage II (GURUSHIDZE & al. in press), and most of them share nectary
types of either group H or group I (GURUSHIDZE & al. 2008). Also the seed
coat structure studied by FRITSCH & al. 2006 does not contradict this nar-
rower circumscription of sect. Acmopetala, though the rather great varia-
tion did not allow to separate similar subgroups as above presented. Only
the sections Longibidentata and Stellata were separable by specific seed
coat character states. Thus, we can accept section Acmopetala in the strict
sense as a natural group, with the subsections Acmopetala (in a wide
sense), Durovaginata (without A. tschimganicum), Pharmakoprason, and
possibly Inornatae and Albidiflora. The phylogenetically closest relative is
sect. Miniprason, member of cluster 5, but the relations to other sections
were not resolved.
3.9. Core Clade, Cluster 7
This cluster is large and most complicated containing several groups
having badly or not resolved phylogenetic relations. GURUSHIDZE & al. 2008
accepted three well supported main groups termed ªJº, ªKº and ªLº
(Figs. 1, 2 D, 2 E, 2 F), and did not discuss the smaller ones. Addition of
much more living accession and herbarium vouchers did not much change
the general structure but remarkably enlarged several groups. Our data
allow to recognize altogether at least 11 groups.
3.9.1. Allium lipskyanum Alliance, sect. Regeloprason
Also in cluster 7, a few species of sect. Regeloprason were included in
two different subgroups. One well supported, small subgroup contains A.
lipskyanum, an endemic of Southwest and Central Hissar mountain range
in Tajikistan and Uzbekistan, and A. chodsha-bakirganicum, known only
from the type location in central Turkestan mountain range, western Kyr-
gyzstan. Both show a moderate morphological similarity, which was ex-
pressed by the long distances after the basal bifurcation. Addition of more
material would be essential to verify whether they actually belong to one
clade. The other subgroup was located inside of sect. Compactoprason and
will be discussed in chapter 3.9.4.
192
3.9.2. Allium komarowii Subgroup (sect. Compactoprason)
Only A. komarowii, naturally distributed in the northwestern Turke-
stan up to Southwest Hissar mountain ranges in Tajikistan and Uzbeki-
stan, is included in this homogeneous subgroup (Fig. 2 D) which is iden-
tical with the morphologically well separated, monotypic subsect. Komar-
owiana of sect. Compactoprason. The infra-sectional subdivision is dis-
cussed in the chapter below.
3.9.3. sect. Compactoprason
Beside of three species of sect. Regeloprason (see next chapter), this
group comprises only species of subsect. Erectopetala (Fig. 2 D). In a basal
position is A. giganteum, distributed in an area stretching from Central
Koppeh Dagh in Turkmenistan and Iran via northern Afghanistan to South
Tajikistan and even the Panj valley in western Pamir region. It is divided into
a basal subgroup comprising accessions from all parts (beside Afghanistan)
of the above mentioned area of distribution. Somewhat advanced is another,
rather well supported subgroup composed of Tajik (acc. 6746 stems from
Uzbekistan near the Tajik border) accessions only. The members of these
subgroups do not differ morphologically, and the small geographic and mo-
lecular differences do not justify their recognition at infraspecific level.
Rather homogeneous and phylogenetically more derived than A. gi-
ganteum is our material of A. macleanii (known from Central Hissar, Pa-
mir, and Alai mountain ranges via Northwest Afghanistan to East Afghan
Hindu Kush mountain range), if we assume that the accessions 0465 and
3884 (received from botanical collections) may represent hybrids and oc-
cupy therefore isolated positions. However, material from Afghanistan
would be essential to verify these positions in this cluster.
It was unexpected that the members of the morphologically only
moderately divers sect. Compactoprason were positioned in different mo-
lecular subclusters. The only species of subsect. Spiralopetala,A. majus
(a rare species from southwestern Hissar mountain range), is morphologi-
cally rather similar to A. giganteum but was placed among the derived
sub-clade of sect. Kaloprason (chapter 3.9.8.) close to taxa occurring in the
same geographic region. Also A. trautvetterianum, morphologically still
more similar to A. giganteum, is sister to A. sarawschanicum and will be
discussed below in chapter 3.9.6. In terms of molecular characters, sect.
Compactoprason is heterogeneous. On the other hand, inclusion of all
members of this section in only three subgroups of cluster 7 underline a not
too low degree of relatedness. Thus sect. Compactoprason is also no artifi-
cial group. However, at the moment we do not know enough character
differences in order to propose a more natural subdivision. Also the seed
coat characters of these species are rather similar (FRITSCH & al. 2006) and
do not present clues for another subdivision of this section.
193
194
3.9.4. sect. Regeloprason subsect. Odoratae
Three montane to subalpine species of this subsection were placed
between A. giganteum and A. macleanii, sect. Compactoprason discussed
above. The type species of subsect. Odoratae,A. darwasicum, is widely
distributed from central and southern parts of Darvaz mountain range up
to Central and south-western Hissar range in Tajikistan. This species
forms one moderately homogeneous sub-clade being the sister to both the
other species. A. hissaricum from Central Tajikistan forms another homo-
geneous, well separated sub-clade, and A. winklerianum (occurring in the
Central Darvaz up to Central Hissar mountain ranges) is part of the most
derived but well supported sub-clade jointly with A. macleanii. Recently
A. winklerianum was reported by XU&KAMELIN 2000 to occur also in the
upper Ili valley of Chinese Tian Shan mountain range, but material was
not available to us.
These are problematical results, not only concerning the nearly iden-
tical ITS-sequences of members of two really dissimilar sections. Here only
three accessions from the type location (most southern end of Darvaz
mountain range) of A. darwasicum were included. Other analyzed acces-
sions from this and other locations are clustering with A. intradarvazicum
among species of sect. Megaloprason (see chapter 3.6.2.). Genome-wide
screening by RAPD analysis of GURUSHIDZE & al. 2008 provided evidence
that the investigated accessions of A. darwasicum belong to two morpho-
logically indistinguishable different evolutionary lineages affiliated to the
clusters 4 and 7. Clear reasons for this case of cryptic species were not
found. It seems extremely unlikely that several introgressions from A. in-
tradarvazicum and subsequent homogenization of the rDNA loci towards
the introgressed type could be responsible for the occurrence of A. darwa-
sicum accessions in cluster 4.
3.9.5. sect. Aroidea
This monotypic section is morphologically well characterized and
molecularly well separated, but the phylogenetic relationships are not re-
solved (Fig. 2 E). The only species A. aroides is a rare endemic in the wes-
tern part of Uzbek Saravshan mountain range with an outpost in the
northeastern corner of Turkmenistan. The seed coat has flat periclinal
walls with evenly granulose ornamentation, and nearly straight anticlinal
walls (FRITSCH & al. 2006) which could be explained either as ancient or as
advanced (extremely reduced) sculptures.
3.9.6. Allium sarawschanicum ± A. trautvetterianum Clade
The union of these morphologically deviating species having also dif-
ferent areas of distribution in a molecularly well supported clade (Fig. 2 E)
is really surprising.
195
A. sarawschanicum shares many morphological characters and ecolo-
gical preferences with A. rosenbachianum s. str., the type species of sect.
Megaloprason (see chapter 3.6.1.). The natural area of distribution of A.
sarawschanicum stretches from the central and western parts of Sarav-
shan and Hissar mountain ranges in Uzbekistan and Tajikistan via
Northwest Afghanistan to the central and eastern parts of Koppeh Dagh
mountain range in Turkmenistan and Iran. But the well supported se-
paration in two sub-clades does not reflect geographic diversity, because
accessions from Hissar mountain range, composing the somewhat sepa-
rated sub-clade, are also present in the basal clade. Our molecular data do
also not support to split plants from Turkmenistan as A. pseudozer-
avschanicum as proposed by SEISUMS 1992, 2000.
A singular character of A. sarawschanicum is the presence of six ra-
dial horn-like to rib-like outgrowths on the top of the ovary. This character
is rare in subg. Melanocrommyum and not known from any other member
of sect. Megaloprason s. lat. The molecular data underline a separate po-
sition, but we hesitate to describe a new section as long as the phylogenetic
difference to the sections Procerallium, Acmopetala, and Compactoprason
remains unresolved. Currently recognition of a new monotypic subsection
of sect. Megaloprason s. str. seems us the best solution:
Allium [sect. Megaloprason s. str.] s u b s e c t . Keratoprason R. M.
FRITSCH,subsect. nova
Type species: A. sarawschanicum REGEL
Scapus valde elongatus, strictus, levis, folia excedens. Folia in numero
2±3 (vel unum), sublate lanceolata, stricta apice nutantia. Inflorescentia
sphaerica, densa. Perigonium stellatum patelliforme, tepala lanceolata
acuta. Filamenta basi dilatata connata interiora utrinque indistincte uni-
dentata supra subulata. Ovarium breve hexasulcatum pyriforme apice
toris acutis radialibus vel cornibus brevibus 6 provisum.
Scape very long, straight, smooth. Leaves shorter than scape, up to 3 in
number, +widely lanceolate, straight with down hanging tips. In-
florescence globose and dense. Flowers star-like bowl-shaped, tepals lan-
ceolate and acute, filaments basally widened and united, inner filaments
basally with one tooth at every side, above subulate. Ovary shortly and 6-
furrowed pear-shaped, with 6 sharp horn-like or radial outgrowths.
A. trautvetterianum is only distributed in a small area in southeast
Tajikistan roughly along longitude 708E and latitude from 37.58to 38.58N.
It shares only character states very common in subg. Melanocrommyum
with A. sarawschanicum. All other characters and character states differ
remarkably and show a high degree of similarity to A. giganteum.Mor-
phologically, A. trautvetterianum belongs certainly to sect. Compactopra-
son in the strict sense, and we do not hesitate to leave it there. Difficult to
explain is the molecular diversity (Fig. 2 E) because most of the in-
vestigated accessions were collected in the narrow vicinity of one village.
3.9.7. sections Popovia and Acaule
Section Popovia comprises only A. gypsaceum and is also morpholo-
gically well characterized, well separated by molecular characters, and
occupies sister position to sect. Kaloprason s. str. (Fig. 2 E). This position
may explain that the seed coat of A. gypsaceum shows a variable undula-
tion of anticlinal cell walls like most members of sect. Kaloprason (FRITSCH
& al. 2006).
The only analyzed accession of A. hexaceras, the only species of sect.
Acaule, is well separated as sister of the sections Popovia and Kaloprason,
but the relationship to the other taxonomic groups of cluster 7 remained
unresolved. This rare species is only known from two separate places in the
subalpine belt of the Hissar mountain range in Tajikistan and Uzbekistan.
Analysis of more material seems essential in order to confirm this position.
3.9.8. sect. Kaloprason in the strict sense
Our molecular data support recognition of this well separated but
heterogeneous section in a rather narrow sense without the A. cristophii
and A. elburzense alliances forming a well separated clade of cluster 2 (see
chapter 3.4.4., Fig. 2 C). The type species A. caspium (widely distributed in
sandy deserts from Pakistan and Iran to South Russia and Kazakhstan)
occupies a basal position (Fig. 2 E) jointly with A. bucharicum, a narrow
endemic of southern Tajikistan. More advanced is the position of A. alex-
eianum, whereas A. nevskianum (vicarious species with A. alexeianum
from central and southwestern Hissar, western Saravshan, and western
Turkestan mountain ranges), A. protensum (clay and stony deserts from
Afghanistan to the eastern Karakum desert and Lake Balkhash area in
Kazakhstan), and A. baissunense, an ecologically specialized, narrow en-
demic of South Tajikistan and South Uzbekistan, occupies basal, inter-
mediate, and derived positions. The latter species is morphologically very
similar to A. caspium and was accepted as subspecies by KHASSANOV &
FRITSCH 1994, but our molecular data would equally support recognition at
species level. The next relative of this section is A. gypsaceum, the only
member of sect. Popovia. We already mentioned in chapter 3.9.3. that we
did not find yet obvious reasons for the inclusion of A. majus, belonging to
sect. Compactoprason, among members of sect. Kaloprason.
One main sub-cluster of cluster 7 comprises the important orna-
mentals A. stipitatum,A. altissimum,A. jesdianum,A. rosenorum, and A.
hollandicum (= A. aflatunense of Dutch bulb trade). A close genetic re-
lationship of these morphologically similar species traditionally affiliated
to sect. Megaloprason s. lat. was expected.
The well supported group ªKº (GURUSHIDZE & al. 2008) contained four
well separated and closely related subgroups. Many additional accessions
blurred this picture: Now the Central Asian subclade is divided in the
196
badly resolved A. stipitatum alliance and the well separated A. rosenorum
alliance (Figs. 2 E, 2 F); sister is the complicated Iranian A. stipitatum ± A.
remediorum alliance, and the well resolved Afghan-Iranian A. jesdianum
alliance jointly with a second group of A. rosenorum from Central Asia.
3.9.9. Allium jesdianum Alliance (sect. Procerallium)
This group is composed of two somewhat heterogeneous clades. One
contains plants from the type location of A. jesdianum from Central Iran,
one accession (6261) from Binalud mountain range in NE Iran probably
belonging to the recently described A. oriento-iranicum (NESHATI & al.
2009), and all available accessions of A. jesdianum subsp. angustitepalum
of Afghan origin from botanic gardens, from bulb trade, and own collec-
tions in South Uzbekistan which are somewhat heterogeneous. These data
strongly support the close relationship of A. angustitepalum WENDELBO
and A. jesdianum ascertained by KHASSANOV &FRITSCH 1994. One acces-
sion of A. stipitatum (1044) positioned here was initially collected in Af-
ghanistan and may represent a hybrid with subsp. angustitepalum.
The second, well supported sub-clade contains only accessions of A.
rosenorum from western Zaravshan and Hissar mountain ranges. All this
material was collected West of 688longitude in Tajikistan and Uzbekistan.
Other morphologically indistinguishable accessions collected East of this
longitude belong to another subgroup presented in chapter 3.9.11. The
taxonomic problems of this separation is discussed below.
Difficult to explain is the unresolved inclusion of A. stipitatum acces-
sion 3347 (from the northern premountains of Alai mountain range in
Kyrgyzstan) in the A. jesdianum alliance because there is neither a geo-
graphic nor a taxonomic relation. The determination was correct, but re-
collection and re-sequencing of material was impossible hitherto.
3.9.10 NW Iranian Geographic Clade
As already discussed by GURUSHIDZE & al. 2008, this phylogenetically
not resolved and heterogeneous group is formed by all investigated acces-
sions of A. stipitatum from Iran (= A. hirtifolium, Zagros mountain range
in NW Iran north of 308N latitude), the only available accession of A.
bakhtiaricum (from Bakhtiar Mountains), and the ornamental A. hollan-
dicum (ªA. aflatunenseº of bulb traders) as derived sub-clade. Beside ad-
ditional accessions of the above mentioned taxa, also other taxa from
Northwest Iran were included in this clade: the recently described A. jes-
dianum subsp. remediorum (FRITSCH &ABBASI 2009), some more accessions
named A. jesdianum, and also a taxon most similar to A. hollandicum but
separated as rather well supported extra subgroup. These data enable us
now to set the discussion of some taxonomic problems on a stronger fun-
dament of facts.
197
First we can only repeat that the Northwest Iranian accessions of A.
stipitatum are morphologically not separable from the most common mor-
photype of A. stipitatum in Central Asia. We accept them as geographically
separated variants of the generally rather divers A. stipitatum alliance (see
next chapter). The borderline separating these variants is apparently si-
tuated south of the North Iranian mountain ranges Alborz and Koppeh
Dagh. However, a closer relation to A. hollandicum must be denied now
because that species constitutes a well separated subgroup which includes
also one accession of A. jesdianum from Kurdestan. This ornamental stems
certainly from Iran, but is apparently closer related and was probably se-
lected from a taxon preliminarily affiliated to A. jesdianum as discussed
below. It was really surprising that morphologically very similar plants
from the mountains West of Lake Orumiyeh did not cluster jointly with A.
hollandicum but as rather well supported extra subgroup. This position
supports to describe them as a new taxon.
Problematic remains the taxonomic treatment of the accessions named
A. jesdianum subsp. jesdianum and subsp. remediorum. Several accessions
from the provinces Markazi, Lurestan, Kermanshah, and Kurdestan be-
long to a slightly separated subgroup related to A. hollandicum, but the
only two investigated accessions from province Shiraz (6706, JA090) were
inserted beside A. stipitatum. Morphologically, the accessions 1119, 6508,
and 6625 do not differ from typical A. jesdianum and might represent a
parallel case as described for A. rosenorum in the chapters 3.9.9. and
3.9.11. The above mentioned plants from province Shiraz are slender in the
general appearance. Careful morphological comparison will be essential to
decide whether the name A. kazerouni PARSA can be applied to them. The
third group of accessions fit well the characters of subsp. remediorum.
3.9.11. Allium stipitatum and A. rosenorum Alliances
All wild A. rosenorum accessions from the area east of the 688long-
itude in Tajikistan (southern slopes of central Hissar mountain range,
there is also the type location) belong to one well supported clade. They are
closely allied to conspecific plants from botanic gardens and bulb trade,
without any subdivision. These molecular data strongly underline a clear
separation of A. rosenorum from A. jesdianum as proposed by KHASSANOV
&FRITSCH 1994.
The second alliance got low support but otherwise is very homo-
geneous, comprising accessions collected in South Kazakhstan, Uzbeki-
stan, Tajikistan, and in the Turkmen Ashgabat area, as well as accessions
from botanic gardens and bulbs sold in shops. Most accessions belong to A.
stipitatum (moderately tall plants with more or less hairy leaves, some-
times only with teeth along the leaf margins, verified by plants from the
type location), two accessions to A. altissimum (less tall plants with nar-
198
rower hairless leaves and a lower number of darker flowers, but wider te-
pals), and some more accessions to a third morphotype traditionally named
A. aflatunense and characterized by taller and later flowering plants with
hairless and rather wide leaves. These facts strongly support the already
earlier discussed assumption (FRITSCH 1993) that A. stipitatum is a highly
variable species and possess morphotypes more similar to A. aflatunense
(but plants from the type location of A. aflatunense were affiliated to sect.
Acmopetala, see chapter 3.8.1.) and other ones fitting the description of A.
altissimum. Unfortunately, we were not able to analyze plants from the
type location of A. altissimum in southeastern Tajikistan anywhere near
Baljuan. It must remain undecided whether that taxon is a good species or
just a variant of A. stipitatum.
One prominent taxonomic problem became visible among the species
of the above discussed group ªKº: several morphologically inseparable
taxa were affiliated to different molecular subgroups which are also geo-
graphically characterized. If we regard these molecular differences as im-
portant argument for separation of taxa at species or at least at subspecies
level, we should separate the ªwesternº accessions of A. rosenorum as
taxon close to A. jesdianum and the accessions of A. jesdianum from
Northwest Iran as taxon close to A. stipitatum, must rise A. jesdianum
subsp. remediorum to species level, and must accept A. hirtifolium and A.
stipitatum as separate species. However, these taxa can only be determined
knowing their geographic origin. In the parallel case discussed in chapter
3.3.6., new names should be given to the accessions of A. ubipetrense in-
serted among the groups dicussed in the chapters 3.3.5., 3.3.8., 3.3.9., and
3.3.11. It remains very doubtful whether an extremely detailed morpholo-
gical comparison of the above discussed taxa may resolve this problem.
GURUSHIDZE & al. 2008 assumed in A. rosenorum and A. stipitatum, that
independent evolutionary lineages seem to have converged on similar
phenotypes. An alternative offspring by hybridization seems not probable,
because only a single accession of A. rosenorum (and no accession of A.
stipitatum) possessed a deviating chloroplast haplotype (GURUSHIDZE & al.
in press). Thus only singular cases of introgression with chloroplast cap-
ture occur.
Irrespective of the above discussed problems, the taxa of the chapters
3.9.9.±3.9.11. are closely related and should represent a formal section.
Because the traditionally applied sectional name Megaloprason is nomen-
claturally typified by A. rosenbachianum which represents a well sepa-
rated clade (chapter 3.6.1.), a new section is created here to which the
subsections Costatae (type species: A. jesdianum) and Elatae (type species:
A. stipitatum) should be transferred.
Allium [subg. Melanocrommyum] sect. Procerallium R. M.
FRITSCH,sect. nova
199
Type species: A. stipitatum REGEL
Scapus valde elongatus proportione tenuis, strictus, levis vel costatus
(in sicco semper costatus), folia minimum aequantia plerumque excedens.
Folia recurvata. Inflorescentia plerumque sphaerica, densa. Perigonium
stellatum; tepala etiam basi libera, saepe mox reflexa et plus minusque
contorta. Ovarium plerumque manifeste stipitatum, verruculosum.
Scape very long and slender, straight, smooth or ribbed (in dry state
always ribbed), mostly much longer than (rarely equal to) the recurved
leaves. Inflorescence commonly globose and dense. Flowers star-like, te-
pals free near base, soon reflexed and +contorted. Ovary commonly dis-
tincly stalked, with rough surface.
3.10. sect. Regeloprason
This is one of the moderately small sections comprising about 18 spe-
cies and subspecies which were affiliated to the subsections Regeloprason
(c. 10 species and subspecies) and Odoratae (c. 8 species). All species pos-
sess narrowly campanulate to funnel-shaped flowers with connate basal
parts of tepals and filaments. The dimensions of nearly all plant parts are
moderate, the filaments are always remarkably shorter than the tepals,
only white, yellow, and different kinds of pinkish-purple flowers occur.
Only length, width, number, and structure of leaves and density of in-
florescences are rather variable. Therefore sect. Regeloprason is generally
regarded as one of the best circumscribed infra-subgeneric groups of subg.
Melanocrommyum.
It was very surprising to see that molecular data draw a contradictory
picture. Members of sect. Regeloprason were scattered over the ITS-based
phylogenetic tree reported by GURUSHIDZE & al. 2008, and also the haplo-
types of the trnL-trnF region of chloroplast DNA belong to well-separated
lineages (GURUSHIDZE & al. in press). 17 species and subspecies were in-
vestigated and split in eight separate groups belonging to clusters 1, 2, 3, 4,
and 7 (Fig. 1). It must be concluded that only members of one group are
closely related, because the genetic distances to all other taxa of this sec-
tion are large. This fact can perhaps best be explained by accepting the
flower form of sect. Regeloprason as ancient structure, which did not much
change, contrary to other morphological and genetic characters, during the
fast diversification (rapid radiation) which happened during evolution of
the current subg. Melanocrommyum. The detection of cryptic species only
in a member of this section, A. darwasicum, (discussed in chapter 3.9.4.)
may also be seen as confirmation of a conservative morphology. Probably
this section is phylogenetically old. We may imagine that basal group(s)
did not survive, and several descendants developed independently during
later phases of evolution but retained the specific flower characters. Also
the very variable characters of seed coats in this section (FRITSCH & al.
200
2006: missing verrucae in A. regelii, variable amplitudes and wave lengths
of anticlinal wall undulation among the other species) may point to dif-
ferent directions of evolution.
Our results show a strong infra-subsectional diversity of subsect. Re-
geloprason and a quite separate position of its type species A. regelii.
Therefore another subsection will be split off here:
Allium [sect. Regeloprason] subsect. Diffusoumbellata R. M. FRITSCH,
subsect. nov.
Type species: A. cupuliferum REGEL
Folia oblonga usque anguste ovata, crassa, patentia usque solum ver-
sus adpressa. Inflorescentia iam ante anthesin rara fasciculata, post an-
thesin laxissima plus minusque sphaerica pedicellis inaequilongis.
Leaves oblong to narrowly ovate, thick, patent or adpressed to the soil.
Inflorescence initially loosely fasciculate, after anthesis very loose and
more or less globose with pedicels of unequal length.
This subsection comprises the alliances of A. cupuliferum and A. isa-
kulii located in cluster 3.
The separate position in the clusters 1 and 4 favors A. pseudo-
winklerianum, A. intradarvazicum, and A. sochense as candidates for one
more subsection, but they share a slender general stature, narrow and
strict leaves, and compact inflorescences with sweet-smelling flowers with
typical members of subsect. Odoratae.
3.11. sect. Thaumasioprason, not Included in this Study
This oligotypic section contains three Afghan endemics A. mirum
(type species), A. caroli-henrici, and A. cucullatum. Another recently de-
scribed species A. khozratense (FRITSCH 2009), a narrow endemic from
Khozratishoh mountain range in Tajikistan, was also preliminarily af-
filiated to this section. These species are morphologically rather well se-
parated from members of all other sections and should be respected also
furthermore. Material of them was not available for molecular analyses
yet.
3.12. Ornamental Cultivars and Hybrids
About 15 ornamental cultivars were initially included in this study,
and 12 probes showed single or homogenized ITS copies and occupied
clear positions in the dendrograms. `Purple Sensation' (accs. 1801, 2615) is
heterogeneous but does not really differ from other accessions of A. hol-
landicum. Our molecular data do not support the thesis that his cultivar is
a hybrid. Similarly, `Michael H. Hoog' and `Purple King' (accs. 5132, 5137,
5478) belong clearly to A. rosenorum, and `Mount Everest' and `White
Giant' (accs. 3246, 5480) to Central Asian A. stipitatum. These positions
completely confirm the results of FRIESEN & al. 1997. Also `Mars' and
201
`Gladiator' (accs. 5135, 5477) were positioned inside of Central Asian A.
stipitatum, despite they are morphologically deviating. `Gladiator' is
probably a hybrid of A. hollandicum, the only parent found by FRIESEN &
al. 1997, with A. stipitatum as second parent because it shows double
peaks clearly uniting the stipitatum and hollandicum copies (15 wobble
positions). `Mars' is most probably a mutant of A. stipitatum only (also
FRIESEN & al. 1997 found no other genome). `Lucy Ball' and `His Ex-
cellency' (accs. 5136, 5479) occupy an unresolved position in the clade
containing Iranian accessions of A. stipitatum, A. jesdianum s. lat., and A.
hollandicum. `Lucy Ball' also may be a hybrid with a homogenized ITS
sequence between any of these species (FRIESEN & al. 1997 accepted A.
hollandicum as one parent) and any taxon from another clade, while the
accession no. 5479 showed additionally the ITS copy of A. macleanii,
pointing that A. macleanii is another parent of 'His Excellency'. Also
`Globemaster' (acc. 5476) in sect. Asteroprason is placed next to A. cristo-
phii, which fits well as one parent, but the position is indifferent concern-
ing A. macleanii (both recognized by FRIESEN & al. 1997) as a second par-
ent. However, in the sequence of this accession several nucleotide positions
showed double peaks, which correspond well to the sequence of A. ma-
cleanii. Thus our data confirmed both parents of this cultivar found by
FRIESEN & al. 1997. Our data also showed that the A. macleanii like se-
quence was partially homogenized towards the A. cristophii type.
4. Typifications
Several species affiliated in the conspectus (next chapter) were treated
in differing sense by several authors. In some cases the type material was
interpreted in different way, or consists of morphologically differing
plants. As far as possible and practical, lectotypes or neotypes are desig-
nated below. Also the holotype sheets of rather many Central Asian species
bear several plants often differing one from another. Then also single plant
should be designated as lectotypes. This selection requires careful com-
parison of all available material, including living specimens, which could
not be done in the frame of the current work. Such nomenclatural deci-
sions must remain one of the assignments for further regional revisions or
a general taxonomic revision of subg. Melanocrommyum.
Lectotype of A. albopilosum WRIGHT (designated here): [no la-
bel, lettering on the sheet] ªAllium albopilosum C. H. WRIGHT in Gard.
Chron. XXXIV p. 34, with fig. 138±02. ELLIS. Hort. Kew. 9 June 1903. Fig-
ured for Bot. Mag. 7982º (K, barcoded K000464557)
Material sent by VA N TUBERGEN to Kew was mentioned in the original
description. Three sheets containing parts of such plants are present in K.
The inflorescence depicted in Botanical Magazine (also mentioned in the
202
original description) is designated herewith as lectotype. It is visible that
the inflorescence was divided in two (somewhat unequal) halves. The ac-
companying leaf may belong to the type plant, but evidence is missing.
Lectotype of A. atriphoenicum BORNM. (designated here): ªAllium
atrophoeniceum (sic!) BORNM. m. (ex aff. A. cardiostemon), Erzurum, am
Khash-Khash-Dagh, 1934 VII. leg. KOTZSCHº (JE)
According to the text of description, type specimens of A. atriphoeni-
cum were given to ªHerb. Haussknecht, Weimar, und Herb. d. Technischen
Hochschule, Dresdenº. The specimen in JE is extant containing one plant
and the draft of a description in BORNMU
ÈLLER's handwriting. It is desig-
nated here as lectotype.
Epitype of A. bakhtiaricum REGEL (designated here): [Iran,
prov.] ªChaharmahal-Bakhtiari, 5 km S of Farsan, Deh Cheshmeh Pirghar,
17.5.1994 leg. R. FRITSCHº (IRAN 286)
The holotype specimen of A. bakhtiaricum (Iran: Montys Bakhtinis,
leg. BODE init. Maji 1840, LE) is in a bad state and shows only one (for-
merly two) inflorescences and parts of scapes. The problematic inter-
pretation of this name was discussed in detail by FRITSCH 1996 who also
gave an emendated description based on plants newly collected in the
Bakhtiar mountains. Objections raised by SEISUMS 2000 against this inter-
pretation where answered by FRITSCH & al. 2002. In order to warrant a
stabile application of the name, a voucher of the plants discussed by
FRITSCH 1996 is designated here as epitype.
Lectotype of A. breviscapum STAPF (designated here): [Iran:]
ªAuf steinigen AbhaÈ ngen bei Gentschnahme, 19 (?) Maiº, second label:
ªAllium breviscapum STAPF, In declivibus saxosis ad Ganjname, 19./5. 82,
leg. Th. PICHLERº (WU)
The type sheet at WU contains two plants. The left plant was marked
by an asterisk and designated as lectotype by WENDELBO (label dated
1967), but this selection was apparently never published. It shall be vali-
dated herewith.
Lectotype of A. dasyphyllum VVED. (designated here): [Ka-
zakhstan:] ªProv. Syr-Darja, distr. Aulie-ata, Montes Alexandri in tesquis
subalpinis saxosis prope Utsch-bulak; 8. VII. 1924, leg. MOKEEVA et PO-
POVº, [Herbarium Florae Asiae Mediae 57. Allium dasyphyllum VVED. sp.
n.] (the very right plant, TASH)
Apparently a rather high number of syntypes were distributed as the
above mentioned exsiccata in 1925. One plant of the specimen stored at
TASH is herewith designated as lectotype. This sheet belongs probably to
203
the material used by VVEDENSKY himself and transferred to TASH in the
early 1990ies.
Epitype of A. derderianum REGEL (designated here): ªIran, prov.
Tehran, Karaj, Chalus road, Asara, slopes N of the road. 23. 5. 2006 leg.
ABBASI, M., FRITSCH, R., KEUSGEN, M.º (IRAN 44046)
REGEL 1875 mentioned material of three collectors in the protologue,
out of which ªPersia borealis, leg. DERDERIANº (LE) was designated as
lectotype by AGABABIAN &OGANESIAN 2000. However, this was not really
helpful, because the collection site of DERDERIANS plants is not known, and
A. derderianum is a rather variable taxon (FRITSCH 2008). Therefore a spe-
cimen with known collection site is designated as epitype which shows
narrowly linear-triangulate tepals about 10 mm long and 1 mm wide
(FRITSCH 2008: 61, fig. 8A) as originally described.
Lectotype of A. dilutum STAP F (designated here): ªIter persicum
Dr. J. E. POLAK 1882. Allium dilutum STAPF. Persia borealis. Inter Kaswin
et Zerschk, 5. 5. leg. Th. PICHLERº (WU)
The type sheet at WU contains two plants. The right plant and the
bulb in the centre were marked by a cross and designated as lectotype by
WENDELBO (label dated 1981), but this selection was apparently never
published. It shall be validated here. WENDELBO intended perhaps to cor-
rect the earlier statement (WENDELBO 1966: 49) ªA. dilutum STAPF of which
I have had the type on loan from Kew . . .º.
Lectotype of A. eginense FREYN (designated here): ªP. SINTENIS:
Iter orientale 1890. No. 2436 Allium chrysantherum B. & REUT. Armenia
turcica. Szanduk, in campis. 17. V. det. J. FREYNº (the left-hand, narrow-
leafed plant, WU)
Two different numbers of exsiccata collected by SINTENIS were men-
tioned in the protologue. Several of these syntypic sheets are extant in
several herbaria, but no voucher annotated by FREYN could be traced. One
plant of the voucher in WU is designated here as lectotype which fits per-
haps best the original description.
Lectotype of A. ellisii HOOK. f. (designated here): ªAllium ex af-
finitate A. karataviensis REGEL Bot. Mag. 6451 from Hon. Ch. ELLIS, Ha-
slemere, June, 1900. Collected in Persia by Ney ELIASº, second label ªAl-
lium sp. n. A. karataviense Allium Ellisii, HOOK. f. June 6. 1900 Type of
Bot. Mag. t. 7875º (K, barcoded K000464378)
This specimen delivers all the data mentioned in the protologue and
bears also the determination by HOOKER.WENDELBO added the remark ªTo
me this seems to be a coarse specimen of A. bodeanum RGL.º in 1966. A
204
second sheet at K (K000464377) containing flowers, capsules and an in-
florescence may also represent original material: ª. . . April, 1897. Sent for
name by Honbl. C. ELLIS.º.
Lectotype of A. giganteum REGEL (designated here): ªEx horto
bot. Petropolitano 82.6 Allium giganteum RGL.º, second label ªlectotypus
A. SEISUMS 1992º (two inflorescences on the left side, LE)
In the protologue, beside a wrong offspring (ªStammt aus dem Hima-
laya º) only cultivated material was mentioned (ª es wurden uns von Hrn.
Frank MILES in Bingham, Nottinghamshire, Zwiebeln mitgetheilt.º). Later
(REGEL 1887: 362) he cited as material ªIn Turcomaniae angustiis (RADDE),
prope Merw (A. DONOWAN)º. Because herbarium vouchers labelled in this
way could not be traced at LE, we must conclude that these were also liv-
ing plants. Also the ªTypusº cited by WENDELBO 1971: 88: ªPlanta quae-
dam culta, probabiliter e bulbis in Persia inter Mashhad et Chacha, O'DO-
NOVAN, LEª was not among the authentic specimens in LE (alternatively it
might be the rephrased second reference of 1887). But a sheet containing
two inflorescences of cultivated plants collected in 1882 may well re-
present that material sent by MILES and used by REGEL when describing A.
giganteum in 1883. Also the label was written by REGEL. Herewith the hi-
therto unpublished designation of SEISUMS shall be confirmed and vali-
dated.
Lectotype of A. hirtifolium BOISS. (designated here): ªIn Persia
prope Ispahan, leg. AUCHER No. 5389º (G, iso-lectotype K)
BOISSIER cited two herbarium vouchers in the protologue, and those
cited above was named as type by WENDELBO 1971: 85. WENDELBO's selec-
tion was a good choice and shall also formally be validated here as lecto-
typification.
Neotype of A. isfairamicum O. F EDTSCH. (designated here):
[Kirgizstan:]ªAlaijskij khrebet: Langar (Ucz-Kurganskij), 28. VI. 1904, leg.
B. A. FEDTSCH.º, second label ªneotypus A. SEISUMS 1992º. (LE)
The first, very short description by O. FEDTSCHENKO did not mention
offspring of the plants, and the second description by B. FEDTSCHENKO in
1907 (ªA. isphairamicumº) described the source of the material as ªMontes
Alaici: In decliviis inter Ucz-Kurgan et Karaul, VI. 1904, (B. FEDTSCHENKO
!!)º. However, herbarium vouchers from this site could not be traced nei-
ther in LE nor in FEDTSCHENKO's herbarium at TASH. Therefore it seems
the best solution to validate the hitherto unpublished designation of SEI-
SUMS. Because a clear indication is missing that this sheet was used when
A. isfairamicum was described, only a neotype can be designated.
205
Lectotypus of A. jenischianum REGEL (designated here): ªPer-
sia Jenischº (LE)
WENDELBO 1966: 49 analyzed both syntypes mentioned in the proto-
logue and decided ªOne sheet is labelled 'Persia Jenisch' and this must be
considered the type sheetº. Though I was not able to see the type when I
visited LE, the decision of Wendelbo was correct and shall be validated
herewith.
Lectotype of A. kazerouni PARSA (designated here): ªAllium
Saporis STAPF cult. hort Vindob. 1886 Bulb. in jugo Kotael Henan prope
Kasrun legit STAPFº, second label ªAllium Kazerouni PARSA in Kew Bull.
1949, 34. Determinavit 22.IV.47 G. PARSAº (the very right plant, K, bar-
coded K000464369)
The type sheet contains beside some more labels and notes a plant
with 3 leaves laid in the press when still in buds, 3 inflorescences, and one
inflorescence with 2 fragmented leaves designated as lectotype. Bulbs are
missing.
Lectotype of A. kharputense FREYN &SINT. (designated here):
ªP. SINTENIS: Iter orientale 1889. No. 711 Allium kharputense FREYN &SINT.
n. sp. Armenia turcica, Kharput, in campis ad Miadun, 8. V.º (the right
plant, WU)
As in the case of A. eginense, two different numbers of exsiccata col-
lected by SINTENIS and distributed to several herbaria were mentioned in
the protologue, but no annotated sheet could be traced. The designation
follows WENDELBO 1971: 79, who mentioned the LD specimen of no. 711 as
ªTypus (isotypus)º.
Lectotype of A. olivieri BOISS. (designated here): [Iraq:] ªMeso-
potamia inter Mossul et Baghdad, leg. OLIVIERº (G, not seen)
BOISSIER mentioned two voucher specimens in the protologue. WEN-
DELBO 1971: 78 named only one of them as ªTypusº. The second syntype
was apparently not traced because it was also not mentioned under the
material seen. Therefore WENDELBO's selection is followed here to desig-
nate the lectotype.
Lectotype of A. saposhnikovii NIKITINA (designated here):
[Kirgizstan:] ªKirgizskij khrebet, predgor'ya nad Chon-Arykam lesopo-
sadki Akademii (Premountains of Kirgiz mountain range above Chon-
aryk, forest plantings of academy) 1 iyunya [June] 1956 leg. E. V. NIKITINAº,
second label ªLecto-Typus 26.5.1997 Reinhard M. FRITSCHº (FRU)
In the protologue, ªPredgoryÂa khrebta Kirgizskij Ala-Too. Gora Pas-
pel'dyk. Stepnye fitotsenozy. 1. VI. 1956, E. V. NIKITINA (in Herb. Ac. Sci.
206
Kirghiz. Frunze)º was described as type. However, a voucher bearing ex-
actly this label could not be found, but several vouchers with a rephrased
description of the collection site with identical date and collector are pre-
sent in LE and FRU. It is very probable that they represent syntypes. The
lectotype chosen in 1997 shall be validated here.
Neotype of A. suworowii REGEL (designated here): ªEx horto
bot. Petropolitano 82.5 Allium Suworowi RGL. Fl. turkest.º, second label
ªLectotypus 1992 A. SEISUMSº (LE)
No herbarium specimen from the type location ªIn deserto kirghisico
prope pagum Uralsk (A. REGEL)º could be traced in LE. Probably also this
taxon was described from cultivated living plants collected by A. REGEL in
Turkestan. Also in TASH no voucher from this place (a former post station
about 90 km S Tashkent) was found. SEISUMS proposed a voucher of a plant
cultivated in St. Petersburg and labelled in E. REGEL's handwriting as lec-
totype specimen. Though it contains a leaf part and an inflorescence with a
part of the scape only, a better voucher is not known. Because this material
was collected after the publication of A. suworowii, it can only be desig-
nated as neotype.
Lectotype of A. trilophostemon BORNM. (designated here):
ªAllium ± Cilicia: in m. Tauro et Antitauro leg. DIECK 1906 cult. in ZoÈschen
1907 VI. 10 com. pl. viv. (small plant in the left corner above, B, barcoded B
10 0268786)
Four plants annotated by BORNMU
ÈLLER are still extant in B. A large
plant labelled ªAllium trilophostemon BORNM. sp. n. Fedde Repert. 1912 ?
cult. in horto meo (Weimar) pl. viv. a 1906 leg. DIECK 1911. VI. leg. J.
BORNMU
ÈLLERº (probably collected after submission of the description), and
the lectotype specimen were mounted at one sheet. A second sheet bears
one non-flowering and one flowering plant collected 19. 06. 1912 (after
publication of the description at 31.12.1911) together with an analytic fig-
ure of the flower parts.
Lectotype of A. tschimganicum O. F EDTSCH. (designated here):
ªAllium tschimganicum B. F. Ol'gino, cult. 1902 g. Semena iz Chimgana
poseyali IX. 1897, tsvety v 1j raz v 1901 g. (Seeds from Chimgan, sown
Sept. 1897, flowered for the first time in 1901)º with a second label ªlec-
totypus A. SEISUMS 1992º (LE)
=A. motor KAMELIN &LEVICHEV
Olga FEDTSCHENKO described this taxon in Progress. Sadov. Ogorodn.
No. 36: 332, 1906 (in Russian) as a tall plant with narrower leaves and
violet flowers. A longer description (as far as known written by VVE-
DENSKY) in the Russian book of M. G. POPOV (ed.), Key Pl. Envir. Tashkent:
207
65, 1923, mentioned linear, subobtuse, c. 4 mm long tepals and 5±21 mm
wide leaves having cartilagineously dentate margins. These characters fit
rather well the later described taxon A. costatovaginatum also occurring in
the Chimgan area. Obviously VVEDENSKY interpreted A. tschimganicum
without having seen the authentic specimens (which were labelled by him
ªPotius A. Severtzovii RGL.º only at 23.I.1924). Later VVEDENSKY treated A.
tschimganicum as synonym of A. fetisowii and A. sewerzowii, but revived
A. tschimganicum in VVEDENSKY &KOVALEVSKAYA 1971: 53, 84 as taxon
separated by pink tepals from the pinkish-violet flowering A. ªsevertzoviiº
sensu VVED.
Only several years ago herbarium specimens of A. tschimganicum
collected in FEDTSCHENKO's garden prior to the description of this species
were traced in LE. These plants are 70 ± 100 cm high, have +lanceolate,
acute tepals 5±7 mm long, inner filaments with small side teeth, rough
ovaries, and leaves 30 ± 50 cm long and up to 30 mm wide. These characters
fit neither to A. fetisowii nor to A. sewerzowii s. str. nor to A. costatovagi-
natum, but could well belong to A. motor or to A. severtzovioides. Violet
flowers as mentioned in the original description fit better to the purplish-
pink tepals of A. motor which bears regularely side-teeth at the base of
inner filaments, whereas A. severtzovioides owns clear pink flowers and
only sometimes side-teeth. Leaf blades and scape characters of these two
species are similar, but A. severtzovioides bears a long, coarse, and rather
hard sheath leaf covering above soil the basal part of the plants mostly till
anthesis. A. motor bears only a small, soft, and quickly decomposing
sheath leaf. Unfortunately, no authentic plant bears bulbs, and no remains
of sheath leaves are visible at all. Certainly the outstanding and carefully
working botanists Olga and Boris FEDTSCHENKO would have recognized the
conspicuous sheath leaf if it was present, and would have taken care that it
remains at the specimens laid into the press. Thus, there are three argu-
ments to regard A. tschimganicum as conspecific with A. motor.
The authentic plants were mounted at 3 sheets. The sheet containing
one large plant without bulb in anthesis shall be validated here as lecto-
type.
Neotype of Allium viridiflorum POBED. (designated here):
ªSemennoe vosproisvedenie tipa, dostavlenno v oranzhereyu N 24, s Fer-
ganskogo khr., ur. Arkit, Dzhalalabadskoj obl. O. I. NEUSTRUEVOJ v 1945 g.
(Reproduction from seed of the type, delivered to the glasshouse no. 24
from Fergan Mts., place Arkit, district Dshalalabad, by O. I. NEUSTRUEVA
in 1945), 15. VIII. 1952 leg. E. G. POBEDIMOVAº (the left upper plant, LE)
It was clearly said in the Russian text of the original description and
was also indicated in the protologue ªTypus: Distr. Dshelalabad. In decli-
viis montium prope Arkit, culta in horto Instituta botanici Academiae
Scientarum URSS, leg. O. NEUSTRUEVA 1945º that this species was de-
208
scribed from a living plant (valid according to art. 37.1 ICBN). However, a
voucher specimen of this cultivated plant could not be traced in LE, and
only the above mentioned specimen referred to NEUSTRUEVA or POBEDI-
MOVA. Because that voucher of A. viridiflorum was not available when this
species was described, only a neotype could be designated.
Lectotype of A. vvedenskyanum PAVLOV (designated here):
[Kazakhstan:] Allium Vvedenskyanum N. PAVL . sp. n. Kaz. SSR Almaa-
tinsk. obl., sukhaya step bliz st. Otar [region Alma-Ata, dry steppe near
station Otar] 2. VI. 1940, No. 583 Leg. + Det. N. V. PAVLOV (MW)
Probably syntypes of this species were distributed to several herbaria,
but a specimen in AA could not be seen. The specimen housed in MW is in
a good state, and the label bears the determination in PAVLOV's hand-
writing. It is designated here as lectotype.
5. Conspectus speciorum generis Allium L. subgeneris Melano-
crommyum (WEBB &BERTHEL.) ROUY
5.1. Conspectus
1. sect. Longibidentata (R.M.F
RITSCH)R.M.FRITSCH
Allium fetisowii REGEL, type species (= A. simile Regel), Allium chych-
kanense R. M. Fritsch
2. sect. Decipientia (O MELCZUK)R.M.FRITSCH
Allium decipiens FISCH.exSCHULT.&SCHULT. f., type species, subsp.
decipiens,Allium decipiens subsp. quercetorum SEREGIN, Allium che-
lotum WENDELBO,Allium grande LIPSKY,Allium roborowskianum RE-
GEL,Allium robustum KAR.&KIR.,Allium sinkiangense F. T. W ANG &Y.
C. TANG,Allium tulipifolium LEDEB., Allium viridulum LEDEB.
3. sect. Regeloprason WENDELBO
3.1 s u b s e c t . Regeloprason (WENDELBO)KAMELIN s. str. Allium
regelii TRAUTV., type species of section and subsection Re-
geloprason (= A. yatei AITCH.&BAKER), Allium victoris VVED.
3.2 s u b s e c t . Diffusoumbellata R. M. F RITSCH Allium cupuli-
ferum REGEL, type species, subsp. cupuliferum,Allium cupuli-
ferum subsp. nuratavicum R. M. FRITSCH &BESHKO,Allium bal-
khanicum (R. M. FRITSCH &F.O.KHASS.) R. M. FRITSCH,Allium
cathodicarpum WENDELBO,Allium iliense REGEL s. str., Allium
isakulii R. M. FRITSCH &F.O.KHASS.(=A. cupuliferum sensu KA-
MELIN subsp. nuratense KAMELIN), Allium subkopetdagense (R. M.
FRITSCH &F.O.KHASS.) R. M. FRITSCH
3.3 s u b s e c t . Odoratae R. M. F RITSCH Allium darwasicum REGEL,
type species, Allium chodsha-bakirganicum GAFFAROV &TUR-
AKULOV
,Allium hissaricum VVED., Allium intradarvazicum R. M.
FRITSCH,Allium lipskyanum VVED., Allium pseudowinklerianum
209
210
R. M. FRITSCH &F.O.KHASS., Allium sochense R. M. FRITSCH &U.
TURAKULOV,Allium winklerianum REGEL s. str.
4. sect. Melanocrommyum WEBB &BERTHEL.s. str.
4.1 A l l ium nigrum alliance Allium nigrum L., nom. cons., type
species of subgenus, section, and subsection Melanocrommyum
[= A. afrum (ZUCCAGNI)KUNTH,A. magicum L., nom. rej., A.
bauerianum BAKER], Allium struzlianum OGAN.
4.2 Allium asclepiadeum alliance Allium asclepiadeum BORNM.,
Allium chrysantherum BOISS.&REUT.(=A. reflexum BOISS.&
REUT. non F. DIETR.), Allium eginense FREYN,Allium kharputense
FREYN &SINT.,Allium nemrutdaghense KIT TAN &SORGER,Allium
olivieri BOISS., Allium saralicum R. M. FRITSCH,Allium shatakiense
RECH.f.,Allium stenopetalum BOISS.&KOTSCHY ex REGEL,Allium
urmiense KAMELIN &SEISUMS
4.3 Allium bisotunense alliance Allium bisotunense R. M.
FRITSCH,Allium keusgenii R. M. FRITSCH
4.4 Allium cardiostemon alliance Allium cardiostemon FISCH.&
C. A. MEY.(=A. atriphoeniceum BORNM.,A. nabelekii KAMELIN &
SEISUMS,A. trilophostemon BORNM.), Allium mariae BORDZ.,Al-
lium woronowii MISCZ.exGROSSH.(=A. leonidis GROSSH.)
4.5 Allium colchicifolium alliance Allium colchicifolium
BOISS., Allium haussknechtii NA
ÂBE
ÏLEK,Allium libani BOISS., Al-
lium moderense R. M. FRITSCH,Allium straussii BORNM.
4.6 Allium multibulbosum alliance Allium atropurpureum
WALDST.&KIT.,Allium cyrilli TEN., (= A. auctum OMELCZUK,A.
fragrans CIRILLO ex TEN.), Allium elmaliense DENIZ &SU
ÈMBU
ÈL, Al-
lium multibulbosum JACQ.(= A. nigrum auct. non L., A. mon-
spessulanum GOUAN,A. speciosum CIRILLO), ? Allium rhetoreanum
NA
ÂBE
ÏLEK
4.7 Allium noeÈanum alliance Allium karamanoglui KOYUNCU &
KOLLMANN,Allium noeÈanum REUT.exREGEL (= A. dilutum STAPF,
A. jenischianum REGEL)
4.8 Allium orientale alliance Allium aschersonianum BARBEY
(incl.subsp. ambiguum BE
ÁG.&VACC.), Allium crameri ASCHERS.&
BOISS., Allium dumetorum FEINBR.&SZELUB.,Allium fedtschenkoi
NA
ÂBE
ÏLEK,Allium lachnophyllum PAINE,Allium lycaonicum SIEHE,
Allium orientale BOISS., Allium tel-avivense EIG,?Allium tuber-
genii FREYN
4.9 Allium rothii alliance Allium rothii ZUCC.,Allium vinicolor
WENDELBO
5. sect. Acanthoprason WENDELBO
5.1 Allium akaka alliance Allium akaka S. G. GMELIN ex SCHULT.
&S
CHULT. f., type species of sect. Acanthoprason
211
5.2 Allium austroiranicum ALLIANCE Allium austroiranicum R.
M. FRITSCH,Allium jaubertii R. M. FRITSCH (= A. latifolium JAUB.&
SPACH non W. YOUNG necque GILIB.)
5.3 Allium derderianum alliance Allium breviscapum STAPF,
Allium derderianum REGEL,Allium egorovae M. V. AGAB.&OGAN.,
?Allium ramazanicum PARSA,Allium shelkovnikovii GROSSH., Al-
lium vasilevskajae OGAN.
5.4 Allium haemanthoides alliance Allium haemanthoides
BOISS.&REUT.exREGEL s. str., Allium zagricum R. M. FRITSCH
5.5 Allium materculae alliance Allium materculae BORDZ.,Al-
lium graveolens (R. M. FRITSCH)R.M.FRITSCH
5.6 Allium minutiflorum alliance Allium hamedanense R. M.
FRITSCH,Allium minutiflorum REGEL
5.7 Allium ubipetrense alliance Allium ubipetrense R. M.
FRITSCH (= A. haemanthoides var. lanceolatum BOISS.)
6. sect. Pseudoprason (WENDELBO)K. PERSS.&WENDELBO
Allium koelzii (WENDELBO)K.PERSS.&WENDELBO, type species, Allium
hooshidaryae MASHAYEKHI,ZARRE &R.M.FRITSCH
7. sect. Asteroprason R. M. FRITSCH
7.1 s u b s e c t . Asteroprason R. M. FRITSCH Allium elburzense
WENDELBO, type species of section and subsect. Asteroprason,Al-
lium helicophyllum VVED., Allium monophyllum VVED., Allium
kuhsorkhense R. M. FRITSCH &JOHARCHI,Allium pseudobodeanum
R. M. FRITSCH &MAT IN
7.2 s u b s e c t . Christophiana TSCHOLOK.Allium cristophii
TRAUTV., nom. & orth. cons., type species (= A. albopilosum WRIGHT,
A. bodeanum REGEL, nom. rej., A. walteri REGEL), Allium ellisii
HOOK.f.
8. sect. Stellata (F.O.K
HASS.&R. M. FRITSCH)R.M.FRITSCH
Allium taeniopetalum POPOV &VVED., type species, subsp. taeniopeta-
lum, Allium taeniopetalum subsp. mogoltavicum (VVED.) R. M. FRITSCH
&F.O.KHASS.(=A. baschkyzylsaicum KRASSOVSK.), Allium taeniope-
talum subsp. turakulovii R. M. FRITSCH &F.O.KHASS.
9. sect. Megaloprason WENDELBO s. str.
9.1 s u b s e c t . Megaloprason R. M. F RITSCH Allium rosenbachia-
num REGEL s. str., type species of section and subsection Mega-
loprason, Allium insufficiens VVED., Allium kwakense (R. M.
FRITSCH)R.M.FRITSCH,Allium schugnanicum VVED.
9.2 s u b s e c t . Humilicognata R. M. F RITSCH Allium brachysca-
pum VVED., type species, Allium assadii SEISUMS (= A. brachysca-
pum sensu WENDELBO), Allium scotostemon WENDELBO
9.3 s u b s e c t . Keratoprason R. M. F RITSCH Allium sarawschani-
cum Regel, type species, (= A. pseudozeravschanicum POPOV &
VVED.exB.FEDTSCH.&POPOV)
9.4 s u b s e c t . Spiralitunicata R. M. F RITSCH Allium suworowii
REGEL, type species, Allium fibriferum WENDELBO
10. s e c t . Miniprason R. M. F RITSCH
Allium karataviense REGEL (= A. cabulicum BAKER,A. singulifolium
RECH. f.; ? incl. subsp. henrikii RUKSANS)
11. sect. Acmopetala R. M. FRITSCH
11.1 s u b s e c t . Acmopetala R. M. F RITSCH Allium backhousianum
REGEL, type species of section and subsection Acmopetala (= A.
gulczense O. FEDTSCH.), Allium aflatunense B. FEDTSCH. non hort.,
Allium alaicum VVED., Allium arkitense R. M. FRITSCH,Allium be-
keczalicum LAZKOV,Allium calocephalum WENDELBO,Allium da-
syphyllum VVED., ? Allium kurdaicum BAJTENOV,Allium pangasi-
cum TURAKULOV,Allium schachimardanicum VVED., Allium vve-
denskyanum PAVLOV,Allium zergericum F. O. KHASS.&R.M.
FRITSCH
11.2 s u b s e c t . Albidiflora R. M. F RITSCH Allium saposhnikovii NI-
KITINA (= A. collis-magni KAMELIN s. str.)
11.3 s u b s e c t . Durovaginata R. M. F RITSCH Allium costatovagi-
natum KAMELIN &LEVICHEV, type species, (= A. rudolfii TUR-
AKULOV), Allium dodecadontum Vved., Allium severtzovioides R.
M. FRITSCH (= A. sewerzowii auct. non REGEL), Allium tokaliense
KAMELIN &LEVICHEV
11.4 s u b s e c t . Inornatae R. M. F RITSCH Allium sewerzowii REGEL s.
str., type species, Allium tashkenticum F. O. KHASS. & R. M.
FRITSCH,(=A. collis-magni auct. non KAMELIN)
11.5 s u b s e c t . Pharmakoprason R. M. F RITSCH Allium tschimga-
nicum O. FEDTSCH.s.str.(=A. motor KAMELIN &LEVICHEV)
12. s e c t . Verticillata KAMELIN
Allium verticillatum REGEL, type species, Allium viridiflorum POBED.
13. s e c t . Compactoprason R. M. F RITSCH
13.1 s u b s e c t . Erectopetala F. O. K HASS.Allium giganteum REGEL,
type species of sect. Compactoprason and subsect. Erectopetala (=
A. procerum TRAUTV.exREGEL), ? Allium isfairamicum O.
FEDTSCH.,Allium macleanii BAKER (= A. elatum REGEL,A. lucens
NIKITINA, nom. invalid.), Allium trautvetterianum REGEL
13.2 s u b s e c t . Komaroviana F. O. K HASS. & R. M. F RITSCH Allium
komarowii LIPSKY
13.3 s u b s e c t . Spiralopetala F. O. K HASS. & R. M. F RITSCH Allium
majus VVED.
14. s e c t . Procerallium R. M. F RITSCH
14.1 s u b s e c t . Elatae R. M. F RITSCH Allium stipitatum REGEL,type
species of sect. Procerallium and subsect. Elatae (= A. hirtifolium
BOISS.), Allium altissimum REGEL,Allium botschantzevii KAMELIN
212
14.2 s u b s e c t . Costatae R. M. F RITSCH Allium jesdianum BOISS.&
BUHSE, type species, subsp. jesdianum,Allium jesdianum subsp.
angustitepalum (WENDELBO)F.O.KHASS. & R. M. FRITSCH (= A.
ecornutum F. O. KHASS.&MALTZEV), Allium jesdianum subsp.re-
mediorum R. M. FRITSCH, Allium bakhtiaricum REGEL,Allium hol-
landicum R. M. FRITSCH,(=A. aflatunense hort. non B. FEDTSCH.), ?
Allium kazerouni PARSA,Allium rosenorum R. M. FRITSCH (= A.
rosenbachianum auct. non REGEL)
15. s e c t . Aroidea F. O. K HASS. & R. M. F RITSCH
Allium aroides POPOV &VVED.
16. s e c t . Acaule R. M. F RITSCH
Allium hexaceras VVED.
17. s e c t . Popovia F. O. K HASS. & R. M. F RITSCH
Allium gypsaceum POPOV &VVED.
18. s e c t . Thaumasioprason WENDELBO
Allium mirum WENDELBO, type species, Allium caroli-henrici WEN-
DELBO,Allium cucullatum WENDELBO,Allium khozratense R. M.
FRITSCH
19. s e c t . Kaloprason K. K OCH
19.1 s u b s e c t . Kaloprason (K. K OCH)KAMELIN s. str. Allium cas-
pium (PALL.) M. BIEB., type species of section and subsection Ka-
loprason, subsp. caspium (= A. brahuicum BOISS.), Allium caspium
subsp. baissunense (LIPSKY)F.O.KHASS. & R. M. FRITSCH (= A.
rhodanthum VVED.), Allium bucharicum REGEL
19.2 s u b s e c t . Ligulifolia R. M. F RITSCH Allium alexeianum REGEL
s.str., type species, Allium hindukuschense KAMELIN &SEISUMS,
Allium nevskianum VVED.exWENDELBO,Allium protensum WEN-
DELBO (= A. schubertii auct. non ZUCC.)
19.3 s u b s e c t . Schubertia KAMELIN Allium schubertii ZUCC.s. str.
20. s e c t . Brevicaule R. M. F RITSCH
Allium sergii VVED., type species, Allium chitralicum F. T. W ANG &TANG
s. str. (= A. pauli VVED., A. badakhshanicum WENDELBO), Allium eugenii
VVED.
21.Species incertae sedis
Allium triste KUNTH &BOUCHE
Â(type not known, perhaps a member of
sect. Kaloprason or sect. Asteroprason), Allium chitralicum sensu
WENDELBO 1971 (no authentic material seen, probably a member of sect.
Procerallium), Allium lallemantii REGEL &RACH.(type not known,
perhaps a member of sect. Decipientia), Allium loratum BAKER (type
plants are in buds only, perhaps a member of sect. Compactoprasum or
sect. Decipientia)
213
214
5.2. Index of Specific and Infraspecific Names Applied in the
ªConspectusº
Scientific name Position
Allium aflatunense B. F
EDTSCH
. non hort. 11.1
Allium aflatunense hort. non B. F
EDTSCH
. 14.2
Allium afrum (Z
UCCAGNI
)K
UNTH
4.1
Allium akaka S. G. G
MELIN
ex S
CHULT
.&
S
CHULT
.f.
5.1
Allium alaicum V
VED
. 11.1
Allium albopilosum W
RIGHT
7.2
Allium alexeianum R
EGEL
s.str. 19.2
Allium altissimum R
EGEL
14.1
Allium arkitense R. M. F
RITSCH
11.1
Allium aroides P
OPOV
&V
VED
.15
Allium aschersonianum B
ARBEY
4.8
Allium asclepiadeum B
ORNM
.4.2
Allium assadii S
EISUMS
9.2
Allium atriphoeniceum B
ORNM
.4.4
Allium atropurpureum W
ALDST
.&K
IT
. 4.6
Allium auctum O
MELCZUK
4.6
Allium austroiranicum R. M. F
RITSCH
5.2
Allium backhousianum R
EGEL
11.1
Allium badakhshanicum W
ENDELBO
20
Allium bakhtiaricum R
EGEL
14.2
Allium balkhanicum (R. M. F
RITSCH
&
F. O. K
HASS
.) R. M. F
RITSCH
3.2
Allium baschkyzylsaicum K
RASSOVSK
.8
Allium bauerianum B
AKER
4.1
Allium bekeczalicum L
AZKOV
11.1
Allium bisotunense R. M. F
RITSCH
4.3
Allium bodeanum R
EGEL
, nom. rej. 7.2
Allium botschantzevii K
AMELIN
14.1
Allium brachyscapum sensu W
ENDELBO
9.2
Allium brachyscapum V
VED
. 9.2
Allium brahuicum B
OISS
. 19.1
Allium breviscapum S
TAPF
5.3
Allium bucharicum R
EGEL
19.1
Allium cabulicum B
AKER
10
Allium calocephalum W
ENDELBO
11.1
Allium cardiostemon F
ISCH
. & C. A. M
EY
. 4.4
Allium caroli-henrici W
ENDELBO
18
Allium caspium (P
ALL
.) M. B
IEB
.subsp.
caspium
19.1
Allium caspium subsp. baissunense (L
IPSKY
)
F. O. K
HASS
. & R. M. F
RITSCH
19.1
Allium cathodicarpum W
ENDELBO
3.2
Allium chelotum W
ENDELBO
2
Scientific name Position
Allium chitralicum F. T. W
ANG
&T
ANG
s. str. 20
Allium chitralicum sensu W
ENDELBO
21
Allium chodsha-bakirganicum G
AFFAROV
&
T
URAKULOV
3.3
Allium chrysantherum B
OISS
.&R
EUT
. 4.2
Allium chychkanense R. M. F
RITSCH
1
Allium colchicifolium B
OISS
. 4.5
Allium collis-magni auct. non K
AMELIN
11.4
Allium collis-magni K
AMELIN
s. str. 11.2
Allium costatovaginatum K
AMELIN
&
L
EVICHEV
11.3
Allium crameri A
SCHERS
.&B
OISS
. 4.8
Allium cristophii T
RAUTV
. 7.2
Allium cucullatum W
ENDELBO
18
Allium cupuliferum R
EGEL
subsp.
cupuliferum
3.2
Allium cupuliferum sensu K
AMELIN
subsp.
nuratense K
AMELIN
3.2
Allium cupuliferum subsp. nuratavicum
R. M. F
RITSCH
&B
ESHKO
3.2
Allium cyrilli T
EN
. 4.6
Allium darwasicum R
EGEL
3.3
Allium dasyphyllum V
VED.
11.1
Allium decipiens F
ISCH
.exS
CHULT
.&S
CHULT
.
f. subsp. decipiens
2
Allium decipiens subsp. quercetorum Seregin 2
Allium derderianum R
EGEL
5.3
Allium dilutum S
TAPF
4.7
Allium dodecadontum V
VED
. 11.3
Allium dumetorum F
EINBR
.&S
ZELUB
.4.8
Allium ecornutum F. O. K
HASS
.&M
ALTZEV
14.2
Allium eginense F
REYN
4.2
Allium egorovae M. V. A
GAB
.&O
GAN
.5.3
Allium elatum R
EGEL
13.1
Allium elburzense W
ENDELBO
7.1
Allium ellisii H
OOK
. f. 7.2
Allium elmaliense D
ENIZ
&S
U
ÈMBU
ÈL
4.6
Allium eugenii V
VED
.20
Allium fedtschenkoi N
A
ÂBE
ÏLEK
4.8
Allium fetisowii R
EGEL
1
Allium fibriferum W
ENDELBO
9.4
Allium fragrans C
IRILLO
ex T
EN
. 4.6
Allium giganteum R
EGEL
13.1
Allium grande L
IPSKY
2
215
Scientific name Position
Allium graveolens (R. M. F
RITSCH
)
R. M. F
RITSCH
5.5
Allium gulczense O. F
EDTSCH
.11.1
Allium gypsaceum P
OPOV
&V
VED
.17
Allium haemanthoides B
OISS
.&R
EUT
.ex
R
EGEL
s. str.
5.4
Allium haemanthoides var. lanceolatum B
OISS
. 5.7
Allium hamedanense R. M. F
RITSCH
5.6
Allium haussknechtii N
A
ÂBE
ÏLEK
4.5
Allium helicophyllum V
VED
. 7.1
Allium hexaceras V
VED
.16
Allium hindukuschense K
AMELIN
&S
EISUMS
19.2
Allium hirtifolium B
OISS
. 14.1
Allium hissaricum V
VED
. 3.3
Allium hollandicum R. M. F
RITSCH
14.2
Allium hooshidaryae M
ASHAYEKHI
,Z
ARRE
&
R. M. F
RITSCH
6
Allium iliense R
EGEL
s. str. 3.2
Allium insufficiens V
VED
. 9.1
Allium intradarvazicum R. M. F
RITSCH
3.3
Allium isakulii R. M. F
RITSCH
&F.O.K
HASS
. 3.2
Allium isfairamicum O. F
EDTSCH
.13.1
Allium jaubertii R. M. F
RITSCH
5.2
Allium jenischianum R
EGEL
4.7
Allium jesdianum B
OISS
.&B
UHSE
subsp.
jesdianum
14.2
Allium jesdianum subsp. angustitepalum
(W
ENDELBO
)F.O.K
HASS
. & R. M. F
RITSCH
14.2
Allium jesdianum subsp. remediorum
R. M. F
RITSCH
14.2
Allium karamanoglui K
OYUNCU
&K
OLLMANN
4.7
Allium karataviense R
EGEL
10
Allium karataviense R
EGEL
subsp. henrikii R
UKSANS
10
Allium kazerouni P
ARSA
14.2
Allium keusgenii R. M. F
RITSCH
4.3
Allium kharputense F
REYN
&S
INT
. 4.2
Allium khozratense R. M. F
RITSCH
18
Allium koelzii (W
ENDELBO
)K.P
ERSS
.&
W
ENDELBO
6
Allium komarowii L
IPSKY
13.2
Allium kuhsorkhense R. M. F
RITSCH
&
J
OHARCHI
7.1
Allium kurdaicum B
AJTENOV
11.1
Allium kwakense (R. M. F
RITSCH
)
R. M. F
RITSCH
9.1
Scientific name Position
Allium lachnophyllum P
AINE
4.8
Allium lallemantii R
EGEL
&R
ACH
.21
Allium latifolium J
AUB
.&S
PACH
non
W. Y
OUNG
necque G
ILIB
.)
5.2
Allium leonidis G
ROSSH
. 4.4
Allium libani B
OISS
. 4.5
Allium lipskyanum V
VED
. 3.3
Allium loratum B
AKER
21
Allium lucens N
IKITINA
13.1
Allium lycaonicum S
IEHE
4.8
Allium macleanii B
AKER
13.1
Allium magicum L., nom. rej. 4.1
Allium majus V
VED
. 13.3
Allium mariae B
ORDZ
.4.4
Allium materculae B
ORDZ
.5.5
Allium minutiflorum R
EGEL
5.6
Allium mirum W
ENDELBO
18
Allium moderense R. M. F
RITSCH
4.5
Allium monophyllum V
VED
. 7.1
Allium monspessulanum G
OUAN
4.6
Allium motor K
AMELIN
&L
EVICHEV
11.5
Allium multibulbosum J
ACQ
.4.6
Allium nabelekii K
AMELIN
&S
EISUMS
4.4
Allium nemrutdaghense K
IT
T
AN
&S
ORGER
4.2
Allium nevskianum V
VED
.exW
ENDELBO
19.2
Allium nigrum auct. non L. 4.6
Allium nigrum L. 4.1
Allium noeÈanum R
EUT
.exR
EGEL
4.7
Allium olivieri B
OISS
. 4.2
Allium orientale B
OISS
. 4.8
Allium pangasicum T
URAKULOV
11.1
Allium pauli V
VED
.20
Allium procerum T
RAUTV
.exR
EGEL
13.1
Allium protensum W
ENDELBO
19.2
Allium pseudobodeanum R. M. F
RITSCH
&
M
ATI N
7.1
Allium pseudowinklerianum R. M. F
RITSCH
&
F. O. K
HASS
.
3.3
Allium pseudozeravschanicum P
OPOV
&
V
VED
.exB.F
EDTSCH
.&P
OPOV
9.3
Allium ramazanicum P
ARSA
5.3
Allium reflexum B
OISS
.&R
EUT
. non F. D
IETR
. 4.2
Allium regelii T
RAUTV
. 3.1
Allium rhetoreanum N
A
ÂBE
ÏLEK
4.6
Allium rhodanthum V
VED
. 19.1
Allium roborowskianum R
EGEL
2
216
Scientific name Position
Allium robustum K
AR
.&K
IR
.2
Allium rosenbachianum auct. non R
EGEL
14.2
Allium rosenbachianum R
EGEL
s. str. 9.1
Allium rosenorum R. M. F
RITSCH
14.2
Allium rothii Z
UCC
.4.9
Allium rudolfii T
URAKULOV
11.3
Allium saposhnikovii N
IKITINA
11.2
Allium saralicum R. M. F
RITSCH
4.2
Allium sarawschanicum R
EGEL
9.3
Allium schachimardanicum V
VED
. 11.1
Allium schubertii auct. non Z
UCC
.19.2
Allium schubertii Z
UCC
.s. str. 19.3
Allium schugnanicum V
VED
. 9.1
Allium scotostemon W
ENDELBO
9.2
Allium sergii V
VED
.20
Allium severtzovioides R. M. F
RITSCH
11.3
Allium sewerzowii auct. non R
EGEL
11.3
Allium sewerzowii R
EGEL
s. str. 11.4
Allium shatakiense R
ECH
.f.4.2
Allium shelkovnikovii G
ROSSH
. 5.3
Allium simile R
EGEL
1
Allium singulifolium R
ECH
.f. 10
Allium sinkiangense F. T. W
ANG
&Y.C.T
ANG
2
Allium sochense R. M. F
RITSCH
&
U. T
URAKULOV
3.3
Allium speciosum C
IRILLO
4.6
Allium stenopetalum B
OISS
.&K
OTSCHY
ex
R
EGEL
4.2
Allium stipitatum R
EGEL
14.1
Allium straussii B
ORNM
.4.5
Allium struzlianum O
GAN
. 4.1
Allium subkopetdagense (R. M. F
RITSCH
&
F. O. K
HASS
.) R. M. F
RITSCH
3.2
Scientific name Position
Allium suworowii R
EGEL
9.4
Allium taeniopetalum P
OPOV
&V
VED
.subsp.
taeniopetalum
8
Allium taeniopetalum subsp. mogoltavicum
(V
VED
.) R. M. F
RITSCH
&F.O.K
HASS
.
8
Allium taeniopetalum subsp. turakulovii
R. M. F
RITSCH
&F.O.K
HASS
.
8
Allium tashkenticum F. O. K
HASS
. & R. M.
F
RITSCH
11.4
Allium tel-avivense E
IG
4.8
Allium tokaliense K
AMELIN
&L
EVICHEV
11.3
Allium trautvetterianum R
EGEL
13.1
Allium trilophostemon B
ORNM
.4.4
Allium triste K
UNTH
&B
OUCHE
Â
21
Allium tschimganicum O. F
EDTSCH
. s. str. 11.5
Allium tubergenii F
REYN
4.8
Allium tulipifolium L
EDEB
.2
Allium ubipetrense R. M. F
RITSCH
5.7
Allium urmiense K
AMELIN
&S
EISUMS
4.2
Allium vasilevskajae O
GAN
.5.3
Allium verticillatum R
EGEL
12
Allium victoris V
VED
. 3.1
Allium vinicolor W
ENDELBO
4.9
Allium viridiflorum P
OBED
.12
Allium viridulum L
EDEB
.2
Allium vvedenskyanum P
AVLOV
11.1
Allium walteri R
EGEL
7.2
Allium winklerianum R
EGEL
s. str. 3.3
Allium woronowii M
ISCZ
.exG
ROSSH
. 4.4
Allium yatei A
ITCH
.&B
AKER
3.1
Allium zagricum R. M. F
RITSCH
5.4
Allium zergericum F. O. K
HASS
.
&R.M.F
RITSCH
11.1
6. Acknowledgements
These investigations were mainly based on the taxonomic Allium reference col-
lection in the IPK Gatersleben. This rich collection was gathered over a period of
more than 25 years of field-work in many countries of Central and Southwest Asia,
Asia Minor, and Israel. We would like to thank all organizers and local scientists
giving support during these research missions and working facilities in the herbaria,
especially the directors and staff of the Botanical Institutes in Dushanbe (Tajikistan),
Tashkent (Uzbekistan), Almaty (Kazakhstan), and in Tbilisi (Georgia), as well as
Neriman O
ÈZHATAY (Istanbul), Rina KAMENETZKY (Jerusalem), Mehrdad ABBASI, Sha-
hin ZARRE and Ali A. MAASSOUMI (Tehran), and Michael KEUSGEN (Marburg). RMF is
especially indebted to Tamara F. KOCHKAREVA (formerly Dushanbe), Furkat O.
KHASSANOV (Tashkent), Isakul TURAKULOV (Khojent) who introduced him into the
Allium flora of Central Asia, and to many local botanists in the Iranian provinces
visited which supported field-work as well as supplied working facilities during re-
vision of herbarium specimens. Substantial support and taxonomic consultation
concerning type specimens in LE by Igor LEVICHEV is also gratefully acknowledged.
We would also like to express our gratitude to the staff of the Taxonomy department
of IPK, to Petra OSWALD, Brunhilde WEDEMEIER, and Birgit WOHLBIER for technical
work in the lab, to Saeideh MASHAYEKHI for DNA extraction and sequence analysis of
several Iranian accessions, and to Christina KOCH, Klaus PISTRICK, and Volkmar
HECKER for maintaining the Allium collection. Funding for a part of the collecting
activities by VolkswagenStiftung (Hannover, Germany) under the general funding
theme ¹Zwischen Europa und Orient ± Mittelasien/Kaukasus im Fokus der Wis-
senschaftª is gratefully acknowledged.
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