Content uploaded by Michael Pimenov
Author content
All content in this area was uploaded by Michael Pimenov
Content may be subject to copyright.
MICHAEL G. PIMENOV, EUGENE V. KLJUYKOV & TATIANA A. OSTROUMOVA
Critical taxonomic analysis of Dichoropetalum, Johrenia, Zeravschania
and related genera of Umbelliferae-Apioideae-Peucedaneae
Abstract
Pimenov, M. G., Kljuykov, E. V. & Ostroumova, T. A.: Critical taxonomic analysis of Dichoropeta-
lum, Johrenia, Zeravschania and related genera of Umbelliferae-Apioideae-Peucedaneae. – Will-
denowia 37: 465-502. – ISSN 0511-9618; © 2007 BGBM Berlin-Dahlem.
doi:10.3372/wi.37.37208 (available via http://dx.doi.org/)
During critical investigation of the Umbelliferae in the Near East, N Africa and Europe some addi-
tional species were found to be putative relatives of Johrenia and the Peucedanum segregate
Johreniopsis. A comparative multivariate analysis of 41 species, based on 32 morphological diag-
nostic characters, revealed a revised taxonomic grouping of the species. The forgotten generic
name Dichoropetalum, typified by D. alpinum, is restored to accommodate a group of 26 SW Asian
and Mediterranean species, including species of the former genera Johreniopsis and Holandrea
(now regarded as sections), as well as some Peucedanum and Johrenia species. We recognize six
sections in Dichoropetalum, of which three are described as new. Two species of Johreniopsis,
poorly known before, are transferred to Zeravschania.Johrenia, on the contrary, is regarded as a
genus with five species only. Johrenia westii, which requires further investigation, belongs most
probably to the genus Ferulago. 28 new nomenclatural combinations are validated and lectotypes
are designated for several names.
Key words: Apiaceae, Holandrea, Johreniopsis, Peucedanum, Mediterranean region, SW Asia, tax-
onomy, morphology, carpology, multivariate analysis.
Introduction
The critical revision of Peucedanum L. s. ampl. and related genera (tribe Peucedaneae) remains
one of the most complicated taxonomic problems in the subfamily Apioideae of the family
Umbelliferae. Despite many new data on Peucedaneae accumulated during the last decades, both
morphological and molecular, the delimitation of the genera and other supraspecific taxa is yet ur-
gently to be revised. The current inappropriate supraspecific classification of the Peucedaneae is
partly the result of gradual (clinal) character variation (like in the Ligusticum alliance), partly of
the incomplete knowledge of its members, some of which are very rare and therefore poorly in-
vestigated.
Willdenowia 37 – 2007 465
The revision of the entire Peucedaneae is a project of the more distant future since it calls for
a complete comparative study of the numerous species spread across Europe, Asia, Africa and
even Oceania. However, we believe that there is an alternative way, and that Peucedanum
s. ampl. can be divided into natural groups by successive splitting of the most divergent taxa.
In the present contribution the traditional Peucedanum sect. Palimbioidea Boiss. is critically
revised in comparison with, on the one hand, putatively related taxa mainly from the Near East,
and, on the other hand, Peucedanum s.str. (i.e. P. sect. Peucedanum). Among the former are the
genera Holandrea Reduron & al. and Johreniopsis Pimenov, described on the basis of species ear-
lier included in this section.
Research history
Some Peucedanum segregates were separated by the first author in “Flora Iranica” (Pimenov
1987). One of them is Johreniopsis, the species of which were formerly placed in P. sect. Pa-
limbioidea. The name Johreniopsis was chosen due to habitual similarity with Johrenia. During
our further investigation of the Umbelliferae from Iran, Turkey and adjacent countries of the Near
East, the close affinity of some Turkish species to Johreniopsis became more and more evident.
Later, the new genus Holandrea (Reduron & al. 1997) was segregated with three European and
Caucasian species also formerly included in P. sect. Palimbioidea. The earlier generic names
Chabraea Raf. and Schlosseria Vuk., used for some species of Holandrea, were shown to be ille-
gitimate. Carpological descriptions of two European Holandrea species (under their traditional
names in Peucedanum) were given by Kowal & Wojterska (1973). When describing recently a
new species from Anatolia, P. isauricum, Parolly & Nordt (2004) made a comparative analysis of
related Turkish species, also including some fruit characters. Fruits of other closely related spe-
cies, however, were described rather scantily.
Evidently there are some major differences between the two species groups traditionally
treated as Peucedanum sect. Peucedanum and sect. Palimbioidea, in morphology, anatomy, che-
mical constituents, chromosome numbers and DNA sequences. In the published molecular
cladograms the species of Peucedanum s.str. (P. officinale, P. morisonii, P. coriaceum, P. galli-
cum) form a clade separate from the Holandrea clade containing H. carvifolia, H. pschavica, H.
schottii and P. achaicum (Downie & al. 2000, Spalik & al. 2004). According to Valiejo-Roman &
al. (2006) Johreniopsis scoparia also belongs to the Holandrea clade, just as Zeravschania Koro-
vin is rather distant, being more similar to Demavendia Pimenov and Haussknechtia Boiss., and
the genus Leutea Pimenov, separated from Peucedanum (Pimenov 1987), is closer to Ferula L.
than to Peucedanum s.str.
Chromosome numbers in many Peucedaneae are uniform, with prevalence of x = 11. All spe-
cies of Peucedanum sect. Palimbioidea, Johrenia, Zeravschania and Johreniopsis studied up to
now are diploid with 2n= 22 (Pimenov & al. 2002). This number was found in P. sect. Peu-
cedanum too, but in this section most species are hexaploids or tetraploids (P. officinale, P.
ruthenicum, P. morisonii, P. longifolium; the last taxon, however, being variable in chromosome
numbers).
Peucedanum sect. Palimbioidea and sect. Peucedanum differ also in their secondary metabo-
lites. All species of P. sect. Peucedanum contain linear furocoumarins, their specific component
is peucedanin (3-methoxy-2-isopropyl-7H-furobenzopyran-7-on). These compounds are not iso-
lated from species of P. sect. Palimbioidea (Carbonnier & al. 1978, Hadacek 1989). On the con-
trary, in P. carvifolia and P. schottii polyacetylene compounds were found (Hadacek 1989) that
are absent in the species of P. sect. Peucedanum.
Fruits of all Peucedanum s.l. species, as well as of some related taxa, are similar in their mor-
phology, adopted for wind dispersal. They have more or less developed winged marginal ribs and
subindivisible dorsal ribs. This similarity was a basis for a broad concept of Peucedanum, but we
believe that this fruit morphological similarly is homoplasic, a view that is supported by some
minor but essential differences in the inner mericarp structure (Kowal & Wojterska 1973).
466 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Among other morphological characters, the degree of development of the calyx teeth is diag-
nostic: they are present in Peucedanum s.str. and absent in all other taxa regarded.
Difference in leaf morphology is considerable. Leaf laminas in Johrenia, Johreniopsis and
Peucedanum sect. Palimbioidea are pinnate, but in Peucedanum s.str. they are ternate-multisect
with geniculate rhachis and segments, which are not situated in one plane. Basal leaf segments
have long petiolules in P. sect. Peucedanum, but are sessile or subsessile in P. sect. Palimbioidea.
Petioles are in cross section usually round with central vascular bundles in sect. Peucedanum, but
are falcate with a groove on the adaxial side without central vascular bundles in sect. Palim-
bioidea. All these differences, minor and sometimes overlapping as they may be, indicate consid-
erable divergence of the two groups, sufficient to exclude P. sect. Palimbioidea from Peucedanum.
The species included in Peucedanum sect. Palimbioidea have changed since its description by
Boissier (1872). Originally the section included the “oriental” species P. chabraei, P. pshawicum
(both currently placed in Holandrea, Reduron &. al. 1997), P. alpinum (currently placed in the re-
stored genus Ormosolenia, Pimenov 1992) and P. palimbioides, whereas P. chrysanthum, P. vitti-
jugum, P. meyeri, P. depauperatum and P. scoparia (plus P. spreitzenhoferi, Boissier 1888) were
attributed to the yellow-flowered group of section “Eupeucedana” (= P. sect. Peucedanum). Thel-
lung (1926) adopted P. sect. Palimbioidea within subg. Peucedanum, comprising the species P.
carvifolia and P. schottii, Shishkin (1951) adopted it with the species P. schottii, P. podolicum, P.
pschawicum and P. palimbioides. Frey (1989) followed mainly Thellung, whose classification he
regarded as the most practical; in Frey’s partial revision of P. sect. Peucedanum and Palimbioidea
the latter includes eight species (P. oligophyllum, P. aequiradium, P. carvifolia, P. schottii, P. gra-
minifolium, P. paucifolium, P. stridii, P. lavrentiadis).
Johrenia was described by Candolle (1829) based on J. dichotoma DC. Later Dichorope-
talum Fenzl, based on D. alpinum Fenzl, was described (Fenzl 1842) but soon after sunk by its
author into the synonymy of Johrenia (Fenzl 1843) and then forgotten. The basis for the modern
Johrenia taxonomy was laid by Boissier (1844, 1872), who placed 10 species (one of them in
question) in Johrenia, and divided them into two informal groups. The first (“vittae jugales ex-
teriores intra marginem spongiosum occultatae”) contains six species (J. selinoides, J. dichoto-
ma, J. fungosa, J. graeca, J. alpina and J. berytea). The second group was again subdivided into
two subgroups, each monotypic, with the species J. candollei and J. platycarpa, respectively.
Drude (1897-98) maintained Boissier’s classification of the genus, while erroneously adding the
Siberian Phlojodicarpus Turcz. to it. Bornmüller (1930) compiled a synopsis of the Johrenia
species and included descriptions of some new species, among which J. polyscias is adopted in
today’s floras. Tamamschjan (1951) demonstrated that Boissier’s main groups differ in fruit ana-
tomical features and recognized these groups in the rank of sections. She also showed that some
species were mistaken for Johrenia and actually belong to Ledebouriella H. Wolff, the present
Saposhnikovia Schischk. Currently, in Johrenia 36 validly published names or combinations are
known (Mozaffarian 2003, IPNI 2006).
Material and methods
43 species of Johrenia, Johreniopsis and Peucedanum from SW Asia, the S European and N Af-
rican Mediterranean and adjacent parts of Europe, as well as of the related genus Zeravschania
were included in our comparative morphological study (Table 1). Besides the species previously
included in P. sect. Palimbioidea (cf. Thellung 1926, Shishkin 1951, Frey 1988) we added to our
set (1) the N African P. munbyi Boiss., which is similar in habit to P. carvifolia and was de-
scribed (Boissier 1854) as closely related to P. chabraei, and (2) the Greek endemic P. achaicum
Halacsy, implicitly regarded as a P. carvifolia relative (Tutin 1968). The names of species, ini-
tially adopted in the present study, are presented in the left column of Table 1, their essential syn-
onyms, as used in literature, in the second column. The plant material, including types for most
species, was studied in ANK, B, BEI, BM, E, G, JE, K, L, LE, LIV, MANCH, MW, P, TARI, W
and WU (herbarium abbreviations according to Holmgren & Holmgren 1998-).
Willdenowia 37 – 2007 467
468 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Table 1. A priori and a posteriori accepted names of the Umbelliferae species included in the present study.
a priori accepted name basionym and essential synonyms a posteriori accepted name
1. Holandrea carvifolia (Vill.)
Reduron & al.
Peucedanum carvifolia Vill. Dichoropetalum carvifolia (Vill.) Pimenov & Kljuykov
2. H. pschawica (Rupr.) Reduron & al. Peucedanum pschawicum Rupr. Dichoropetalum pschawicum (Rupr.) Pimenov & Kljuykov
3. H. schottii (DC.) Reduron & al. Peucedanum schottii DC., P. petraeum W. D. J. Koch Dichoropetalum schottii (DC.) Pimenov & Kljuykov
4. Johrenia alpina (Fenzl) Fenzl Dichoropetalum alpinum Fenzl Dichoropetalum alpinum Fenzl
5. J. aromatica Rech.f. Dichoropetalum aromaticum (Rech. f.) Pimenov & Kljuykov
6. J. aurea Boiss. Dichoropetalum aureum (Boiss.) Pimenov & Kjuykov
7. J. berytea Boiss. Dichoropetalum depauperatum (Boiss.) Pimenov &
Kljuykov
8. J. dichotoma DC. Peucedanum haradjianii Rech. f. Johrenia dichotoma DC.
9. J. distans (Griseb.) Halacsy Caroselinum distans Griseb., Johrenia graeca Boiss. Johrenia distans (Griseb.) Halacsy
10. J. golestanica Rech. f. Dichoropetalum golestanicum (Rech. f.) Pimenov &
Kljuykov
11. J. paucijuga (DC.) Bornm. Ferula paucijuga DC. Dichoropetalum paucijugum (DC.) Pimenov & Kljuykov
12. J. platycarpa Boiss. Dichoropetalum platycarpum (Boiss.) Pimenov & Kljuykov
13. J. polyscias Bornm. Johrenia polyscias Bornm.
14. J. porteri Boiss. ?
15. J. ramosissima Mozaff. Dichoropetalum ramosissimum (Mozaff.) Pimenov &
Kljuykov
16. J. selinoides Boiss. Johrenia selinoides Boiss.
17. J. tortuosa (Fisch. & C. A. Mey.)
D. F. Chamb.
Eriosynaphe tortuosa Fisch.&C.A.Mey.,Johrenia
fungosa Boiss.
Johrenia tortuosa (Fisch.&C.A.Mey.)D.F.Chamb.
18. J. westii Post Ferulago westii (Post) Pimenov & Kljuykov
19. Johreniopsis oligactis (Rech. f. & Riedl)
Pimenov
Peucedanum oligactis Rech. f. & Riedl Zeravschania pauciradiata (Boiss.) Pimenov
20. J. scoparia (Boiss.) Pimenov Johrenia scoparia Boiss., Peucedanum scoparium
(Boiss.) Boiss.
Dichoropetalum scoparium (Boiss.) Pimenov & Kljuykov
21. J. seseloides (C.A.Mey.)Pimenov Ferula seseloides C. A. Mey., Peucedanum paucifolium
Ledeb., P. meyeri (Boiss.) Boiss.
Dichoropetalum seseloides (C.A.Mey.)Pimenov&
Kljuykov
22. J. stricticaulis (Rech. f.) Pimenov Peucedanum stricticaule Rech. f. Zeravschania stricticaulis (Rech. f.) Pimenov & Kljuykov
23. Peucedanum achaicum Halacsy Dichoropetalum achaicum (Halacsy) Pimenov & Kljuykov
24. P. alpigenum Boiss. Dichoropetalum alpigenum (Boiss.) Pimenov & Kljuykov
Willdenowia 37 – 2007 469
25. P. chryseum (Boiss.)D.F.Chamb. Anethum chryseum Boiss., Peucedanum chrysanthum
Boiss.
Dichoropetalum chryseum (Boiss.) Pimenov & Kljuykov
26. P. depauperatum Boiss. Dichoropetalum depauperatum (Boiss.) Pimenov &
Kljuykov
27. P. graminifolium Boiss. Dichoropetalum graminifolium (Boiss.) Pimenov &
Kljuykov
28. P. isauricum Parolly & Nordt Dichoropetalum isauricum (Parolly & Nordt) Pimenov &
Kljuykov
29. P. junceum (Boiss.) Mouterde Johrenia juncea Boiss., Peucedanum spreitzenhoferi
Dingler
Dichoropetalum junceum (Boiss.) Pimenov & Kljuykov
30. P. knappii Bornm. Peucedanum chenur Mozaff. Zeravschania knappii (Bornm.) Pimenov & Kljuykov
31. P. lavrentiadis Strid & Papan. Dichoropetalum lavrentiadis (Strid & Papan.)
Pimenov & Kljuykov
32. P. minutifolium (Janka) Velen. Peucedanum vittijugum subsp. minutifolium (Janka)
Kuzmanov & Andrejev
Dichoropetalum minutifolium (Janka) Pimenov & Kljuykov
33. P. munbyi Boiss. Dichoropetalum munbyi (Boiss.) Pimenov & Kljuykov
34. P. oligophyllum (Griseb.) Vandas Seseli oligophyllum Griseb. Dichoropetalum oligophyllum (Griseb.) Pimenov &
Kljuykov
35. P. palimbioides Boiss. Malabaila carvifolia Boiss. Dichoropetalum palimbioides (Boiss.) Pimenov & Kljuykov
36. P. stridii Hartvig Dichoropetalum stridii (Hartvig) Pimenov & Kljuykov
37. P. vittijugum Boiss. Dichoropetalum vittijugum (Boiss.) Pimenov & Kljuykov
38. Zeravschania aucheri (Boiss.) Pimenov Peucedanum aucheri Boiss. Zeravschania aucheri (Boiss.) Pimenov
39. Z. ferulifolia (Gilli) Pimenov Peucedanum ferulifolium Gilli Zeravschania ferulifolia (Gilli) Pimenov
40. Z. membranacea (Boiss.) Pimenov Peucedanum membranaceum Boiss. Zeravschania membranacea (Boiss.) Pimenov
41. Z. pauciradiata (Boiss.) Pimenov Peucedanum pauciradiatum Boiss. Zeravschania pauciradiata (Boiss.) Pimenov
42. Z. regeliana Korovin Zeravschania regeliana Korovin
43. Z. scabrifolia Pimenov Zeravschania scabrifolia Pimenov
The phenetic analyses include cluster analysis (UPGMA) and principal components analysis
(PCA) conducted by means of the NTSYSpc package for Windows (Rohlf 1994). The resulted
rectangular data matrix is shown in Table 2. Some data are missing, mainly for a few species
poorly represented in the available collections. For a principal components analysis (of charac-
ters and species) these missing character states were replaced by means for each character.
Characters. – The species were investigated in 33 selected morphological characters. The selec-
tion includes previously known diagnostic characters, additional carpological and other charac-
ters used in our previous revisions of complex taxonomic groups in the Umbelliferae (Kljuykov
& al. 2004). Among the characters 11 are vegetative and 22 reproductive, including 12 fruit char-
acters. The fruit structure of the species included in the present investigation is illustrated in Fig.
5-10.
The following characters and character states (with their coding) were recognized:
1. Life-form: monocarpics with unbranching rootstock = 0; polycarpics with branching root-
stock = 1.
2. Stem base: densely covered by remains of leaf petioles = 0; without remains of leaf petioles = 1.
3. Stem height (in cm): 7-40 = 0; 41-100 = 0.5; more than 100 = 1.
4. Stem ribs: stems rounded, without prominent ribs, or finely striate = 0; with slightly devel-
oped ribs = 0.5; with well developed ribs = 1.
5. Stem indumentum: none, glabrous = 0; pubescent = 1.
6. Leaf shape (in outline): leaves linear to lanceolate = 0; ovate to triangular = 1.
7. Leaf sheaths: long, linear = 0; short, linear = 0.5; short, triangular = 1.
8. Basal primary segments (attachment): sessile = 0; with short petiolules (up to 5 mm) = 0.5;
with long petiolules (more than 5 mm) = 1.
9. Terminal leaf lobes: filiform = 0; linear = 0.25; lanceolate = 0.5; ovate, toothed at the margin = 1.
10. Terminal leaf lobes: numerous, densely clustered = 0; not clustered = 1.
11. Upper cauline leaves: long = 0; short = 1.
12. Bracts: present = 0; absent = 1.
13. Min. number of rays per umbel: less than 5 = 0; more than 5 = 1.
14. Max. number of rays per umbel: less than 10 = 0; more than 10 = 1.
15. Umbel rays: very unequal = 0; slightly unequal = 0.5; almost equal = 1.
16. Bractlets (texture): completely herbaceous = 0; with narrow white margins = 1.
17. Bractlets (form in outline): subulate to linear = 0; linear-lanceolate = 0.5; lanceolate to
broadly lanceolate = 1.
18. Bractlets (relative to umbellules): longer than umbellule = 0; shorter than umbellule = 1.
19. Calyx teeth: well developed = 0; very short = 0.5; absent = 1.
20. Petals (colour): white = 0; whitish or yellowish = 0.5; yellow = 1.
21. Stylopodia: flat = 0; short-conical = 0.5; conical = 1.
22. Mericarps (in cross section): dorsally convex = 0; dorsally compressed = 1.
23. Dorsal mericarp ribs: indistinguishable (submerged into spongy pericarp tissue) = 0; fili-
form = 0.5; keeled or slightly inflated = 0.75; strongly spongily inflated = 1.
24. Marginal mericarp ribs: not developed = 0; shortly winged, swollen/inflated = 0.5; shortly
winged, triangular = 0.75; winged, thin = 1.
25. Mericarps (on commissural side): with destroyed parenchyma near carpophore = 0; com-
missural parenchyma not destroyed = 1.
26. Parenchyma cells with lignified pitted walls on dorsal mericarp side: developed = 0; absent
or sometimes only under vascular bundles = 1.
27. Marginal ribs inflated, composed of large mesocarp cells with lignified pitted walls: yes =
0; no = 1.
28. Hypendocarp (inner lignified layer of mesocarp): absent = 0; feebly marked (fragmentary)
= 0.5; strongly expressed, consisting of prosenchyma lignified cells = 1.
470 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Willdenowia 37 – 2007 471
Table 2. Data matrix for the 33 selected morphological characters of the 43 investigated species. For details see text.
29. Secretory ducts (vittae) in mericarp furrows (presence/absence): absent or inconstant, not
visible in each furrow = 0; always developed, more or less broad = 1.
30. Secretory ducts (vittae) in mericarp furrows (number): solitary = 0; several in each furrow =
0.75; cyclic, small = 1.
31. Commissural secretory ducts (vittae): absent = 0; two in each mericarp = 0.5; more than two = 1.
32. Rib secretory ducts: absent = 0; small = 0.5; large, exceeding vallecular vittae if present = 1.
33. Mericarp vascular bundles: compact = 0; consisting of several groups of vascular elements = 1.
Results
Characters. – Principal components analysis represents in plot form correlations between the
distribution of character states within the set of taxa. Projection of character points onto a plot of
1 and 2 principal components (Fig. 1) shows different similarity among characters; the majority
of them are low-correlated, although there are a few groups of highly correlated characters (13
and 30; 26 and 29; 15, 16 and 27; 6 and 28). Overall information from characters within each
such group is lower than the sum of information from independent characters. This character cor-
relation influences the species coordinates in the components analysis, and is not taken into con-
sideration in the cluster analysis.
Taxa excluded. – Two species are to be excluded from further discussion of the Johrenia-Johrenio-
psis-Zeravschania taxonomic complex. These are Johrenia porteri Boiss. and J. westii Post.
Johrenia porteri is a poorly known species. We have studied two sheets in G, including one
to be treated most probably as the holotype (an isotype is kept in BEI). This type material does
not allow to include the species in the comparative analysis. The collection at G is represented by
a part of the inflorescence with compact umbels and immature fruits. A picture of radical pinnate
leaves is attached to the type sheet, but its origin is unclear. In B we found an additional sheet
(“Kurd Dagh, 11.4.1893”), collected and determined by G. Post, with a rosette of radical leaves,
472 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Fig. 1. Character loadings in three-factor space showing character correlation (PCA).
Willdenowia 37 – 2007 473
Fig. 2. UPGMA phenogram of relationships among species investigated.
474 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Fig. 3. OTUs loadings in three-factor space (PCA).
Fig. 4. Two-factor plot (components 1 and 2) showing OTUs relationships in reduced character space.
similar to the above mentioned picture. Similar and more complete material of this gathering ex-
ists in BEI. Collected after the publication of the species, this material is not relevant for
typification and may even represent another species. Bornmüller (1930) proposed to attribute
this gathering to Pimpinella saxifraga, an attribution we doubt, but another Pimpinella species,
P. corymbosa, may be a good match. In G is another sheet (“Ravine frais de Froulok- Bassit (Al-
neto-Platanetum). 23.7.1955, Pabot”), which is in some characters similar to and was treated by
Mouterde (1970) as J. porteri. It has almost mature fruits, with slightly inflated marginal ribs,
one of the diagnostic characters of Johrenia. A more careful investigation of Pabot’s gatherings
showed, however, some differences from the type material of J. porteri. For instance, they have
well developed filiform deflexed bractlets exceeding the umbellule and the umbellules are not
compact but have considerably longer rays and peduncles. The radical leaves, although pinnate,
differ in outline and have longer teeth. It seems possible that this collection is identical or very
close to the newly described Peucedanum longibracteolatum Parolly & Nordt (2005). Therefore,
at present we cannot attribute J. porteri to Johrenia or to any other genus regarded in this analy-
sis. One possible solutions is an attribution of the generative shoots, kept in BEI and G, to Peuce-
danum junceum (see below under Dichoropetalum junceum), and the vegetative rosettes collected
later (B and BEI), to Pimpinella corymbosa, both species being distributed in the locus classicus
of J. porteri. If so, it would be no surprise that there are no more collections of the species. Nev-
ertheless, search for similar plants and collecting of more complete material at the type and
nearby localities is desirable.
Johrenia westii clearly differs from all investigated species in its fruit secretory system, con-
sisting of numerous comparatively small vittae, arranged cyclically. Among platyspermous Um-
belliferae of the region a similar fruit structure is observed only in the species of Ferulago. Well
developed bracts of J. westii are another character, unusual in the present set of species, and pe-
culiar for Ferulago. The vascular system of the mericarps of J. westii is rather unusual for
Apioideae, consisting of several separate groups of vessels; a similar structure has been observed
in Ferulago. The species has, however, umbels with 2-3 rays, whereas all local Ferulago species
have more umbel rays.
In Table 2 some species completely match other species (Johrenia berytea is apparently con-
specific with Peucedanum depauperatum and Johreniopsis oligactis with Zeravschania paucira-
diata). After the exclusion of Johrenia porteri, J. westii, J. berytea and Johreniopsis oligactis,
the final matrix of 41 species and 33 characters was processed.
Main groups of taxa. – Cluster analysis of the data matrix made in the UPGMA modification
(Fig. 2) allows to outline some main groups in our material. The relationships were further eluci-
dated by PCA. The first three factors were excluded on the basis of the character correlation ma-
trix. The character factor loadings were used to produce projections of OTUs in the three-factor
(Fig. 3) and two-factor space. All in all, three two-factor plots were obtained, including the one
reproduced in Fig. 4.
In both analyses, three distinct main groups were revealed, one consisting of several subgroups:
1.1. Holandrea carvifolia, H. schottii, H. pshawica, Peucedanum alpigenum, P. oligophyllum,
P. achaicum, P. graminifolium, P. isauricum, P. munbyi;
1.2. P. lavrentiadis, P. stridii;
1.3. Johrenia alpina, J. berytea, Peucedanum depauperatum, P. junceum, Johrenia aurea;
1.4. Johrenia golestanica, Johreniopsis scoparia, Johrenia platycarpa, J. ramosissima;
1.5. Johrenia aromatica, J. paucijuga;
1.6. Johreniopsis seseloides, Peucedanum vittijugum, P. chryseum, P. minutifolium, P. palim-
bioides;
2. Johrenia dichotoma, J. selinoides, J. polyscias, J. tortuosa, J. distans;
3. Johreniopsis oligactis, Zeravschania pauciradiata, Z. membranacea, Z. regeliana, Johreni-
opsis stricticaulis, Zeravschania ferulifolia, Z. scabrifolia, Z. aucheri, Peucedanum knappii.
Willdenowia 37 – 2007 475
Discussion
In the plot of the first three principal components (Fig. 3) and in the biplot of components 1 and 2
(Fig. 4) three groups are clearly distinguished.
Group 2, corresponding to Johrenia s.str., is clearly separated also in the biplot of compo-
nents 2 and 3 (not shown).
Group 3, containing the type of Zeravschania, Z. regeliana, should be interpreted as slightly
enlarged Zeravschania. It is clearly separated from the remaining species also in the biplot of
components 1 and 3 (not shown).
Group 1 (with subgroups 1.1-1.6) is a polymorphic taxon of the same rank as groups 2 and 3,
for which generic rank seems most appropriate. It contains the species that provide the types of
five generic names: (1) Chabraea Raf. 1840 (type: C. carvifolia (Vill.) Raf. ≡Peucedanum car-
vifolia Vill. ≡Holandrea carvifolia (Vill.) Reduron & al.), non Adans. 1763, (2) Dichoropetalum
Fenzl 1842 (type: D. alpinum Fenzl ≡Johrenia alpina (Fenzl) Fenzl), (3) Schlosseria Vuk. 1857
(type: S. chabraei Schloss. & Vuk. ≡Peucedanum carvifolia ≡Holandrea carvifolia), non Steud.
1841, (4) Johreniopsis Pimenov 1987 (type: J. seseloides (C. A. Mey.) Pimenov), and (5) Holan-
drea Reduron & al. 1997 (type: H. carvifolia (Vill.) Reduron & al.). Among these names, Cha-
braea Raf. and Schlosseria Vuk. are illegitimate as later homonyms (Reduron & al. 1997).
Priority has Dichoropetalum Fenzl, based on D. alpinum, which in our classification belongs to
group 1.3. The generic names Johreniopsis and Holandrea provide later synonyms. Dichoro-
petalum can be clearly divided into six subgroups, which we treat here as sections.
Taxonomy
KeytothegeneraOrmosolenia, Leutea, Peucedanum s.str., Zeravschania, Johrenia and Di-
choropetalum
1. Plants dwarf with low (up to 20 cm tall), creeping, leafless stems; blades of basal leaves en-
tire, round in outline, 3-lobed or deeply dissected (more rarely pinnate with 1-2 pairs of
leaflet); bracteoles 1-3; mericarps with numerous cyclic vittae ..... Ormosolenia
– Plants not dwarf, with erect, taller and leafy stems (if stems are low and leafless, basal
leaves bipinnate or biternate); leaf blades 2-4-dissected; bracteoles more than 3, usually nu-
merous; mericarp with 1-3 (rare up to 5) vallecular vittate ............ 2
2. Ultimate leaf segments cylindric, rigid, petiole-like; leaves almost all radical, stem leaves
size-reduced; umbel rays thickened in fruits; marginal mericarp ribs narrow . . Leutea
– Ultimate leaf segments flat, clearly differing from petioles; umbel rays not thickened; mar-
ginal mericarp ribs broader; if ribs narrow (Zeravschania), leaf segments flat, dentate . . 3
3. Calyx teeth present; petioles round, often with central vascular bundles; leaf lamina ternate-
multisect with segments not situated in one plane; primary segments long-petiolulate; ulti-
matesegmentslong(2-9cm),linear,entire .............. Peucedanum
– Calyx teeth inconspicuous or (in Zeravschania ferulifolia and Z. stricticaulis) very short;
petioles falcate in transection with a groove on the adaxial side, always without central bun-
dles; leaf lamina pinnate, plane; primary segments sessile or petiolulate; ultimate segments
short (<1 cm ), dentate or lobate (in Z. knappii linear, entire, 1-2 cm long) ..... 4
4. Bracts 3 or more; basal primary segments with long petiolules (more than 5 mm); mericarps
(incrosssection)dorsallyconvex ..................Zeravschania
– Bracts absent or, more rarely, 1-2; basal primary segments sessile or with short petiolules
(upto5mm);mericarps(incrosssection)dorsallycompressed .......... 5
5. Leaf sheaths long, linear; dorsal mericarp ribs indistinguishable (submerged into spongy
pericarp tissue); pericarp on commissural side thick, with two portions of non-lignified or
destroyed parenchyma near carpophore.................. Johrenia
– Leaf sheaths short, linear or triangular; dorsal mericarp ribs filiform, keeled, slightly or
strongly inflated; pericarp on commissural side thin, without destroyed parenchyma ..
............................... Dichoropetalum
476 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
1. Dichoropetalum Fenzl, Pugill. Pl. Nov. Syr.: 17. 1842.
Typus: Dichoropetalum alpinum Fenzl
Plantae polycarpicae vel monocarpicae, caulibus basi plerumque residuis foliorum emortuorum
dense tectis, glabris, rarius pubescentibus. Folia vaginis brevibus, linearibus vel triangulatibus,
laminis ambitu triangularibus vel lanceolatis, segmentis primariis basalibus sessilibus vel breve-
petiolulatis, lobis terminalibus linearibus, lanceolatis vel ovatis; folia caulina superiora integra,
longa vel brevia. Umbellae plerumque radiis valde inaequelongis, bracteis nullis. Bracteolae her-
baceae, subulatae, lineares vel lanceolatae, vulgo radiolis breviorae. Dentes calycini obsoleti. Pe-
tala alba vel flava. Stylopodia plana vel conica. Mericarpia dorso compressa, jugis dorsalibus
filiformibus, carinatis vel inflatis, marginalibus brevealatis, vix inflatis vel tenuibus alatis, cellulis
parenchymaticis e latere commissurali non destructis, e latere dorsali elignescentibus vel mem-
branis lignescentibus, porosis, hypendicarpiis evolutis, rarius obsoletis, vittis vallecularibus nullis
vel paucis, commissuralibus nullis, binis vel paucis, canaliculis jugalibus nullis, vel solitariis.
Endospermium ventre planum.
KeytothesectionsofDichoropetalum
1. Stemssimple;petalswhite.................. 1.2.D. sect. Stridia
– Stemsbranchingorleafy;petalswhite,pinkishoryellow............. 2
2. Leaf blades (in outline) linear to lanceolate ................... 3
– Leaf blades ovate to triangular ........................ 5
3. Numbersofraysperumbellessthan5 ............ 1.4.D. sect. Scoparia
– Numberofraysperumbelmorethan6..................... 4
4. Dorsal mericarp ribs filiform or keeled; marginal mericarp ribs winged, thin; parenchyma
cells with lignified pitted walls on dorsal mericarp side absent; furrow and commissural
ductspresent ..................... 1.6.D. sect. Johreniopsis
– Dorsal and marginal mericarp ribs inflated; all mesocarp of parenchyma cells with lignified
pitted walls; furrow and commissural ducts absent ..... 1.5.D. sect. Parajohrenia
5. Upper cauline leaves with long entire blade; petals white or pinkish (but yellow in D.
achaicum) ....................... 1.1.D. sect. Holandrea
– Upper cauline leaves with short entire blade; petals yellow . 1.3. D. sect. Dichoropetalum
1.1. Dichoropetalum sect. Holandrea (Reduron & al.) Pimenov & Kjuykov, comb. & stat. nov.
≡Holandrea Reduron & al. in J. Bot. Soc. Bot. France 1: 93. 1997.
Type: Dichoropetalum carvifolia (Vill.) Pimenov & Kljuykov
Plantae polycarpicae vel monocarpicae, caulibus foliosis, vulgo ramificantibus; laminis foliorum
ambitu oblongis, ovatis vel triangulatis, superioribus integris, longis. Petala alba, rarius flava vel
purpurea. Umbellae 3-15-radiatae. Cellulae mesocarpii parenchymaticae e latere dorsali elignes-
centes. Vittae valleculares solitariae vel paucae.
1. Bractlets longer than umbellule rays .............. (3)D. pshawicum
– Bractletsshorterthanumbellulerays ..................... 2
2. Umbelrays3-5 .............................. 3
– Umbelraysmorethan6 .......................... 4
3. Terminallobesoflowercaulineleaveslinear............. (9)D. munbyi
– Terminal lobes of lower cauline leaves ovate, dentate at the margins . . (8) D. isauricum
4. Mericarps with solitary vallecular vittae .................... 5
– Mericarp with 2-3 vallecular vittae ...................... 6
5. Styles 2-2.5 mm long, 2-3 times longer than stylopods ......... (2)D. schotii
– Styles up to 1 mm long, equal to or shorter than stylopods .... (7)D. graminifolium
6. Terminallobesofstemleavesverynarrow,upto1mmwideandupto8cmlong ....
.............................. (5)D. oligophyllum
Willdenowia 37 – 2007 477
– Terminallobesofstemleaves2-4mmwide,upto4cmlong ........... 7
7. Umbelraysscabrid ...................... (1)D. carvifolia
– Umbelraysglabrous...................... (6)D. achaicum
(1) Dichoropetalum carvifolia (Vill.) Pimenov & Kljuykov, comb. nov. Selinum carvifolia
Chabraei ex Crantz, Inst. Rei Herb. 2: 126. 1766 [& Strip. Austr. Fasc. 3: 62, t. 3, fig. 2. 1767],
nom. illeg., non L. (1762) Peucedanum carvifolia Vill., Prosp. Hist. Pl. Dauphiné: 25. 1779 [&
Hist. Pl. Dauphiné 2: 638. 1787] Selinum chabraei Jacq., Fl. Austriac. 5: 72.1778 [& in
Murray, Syst. Veg., ed. 14: 279. 1784] ≡Imperatoria chabraei (Jacq.) Spreng., Pl. Umb. Prodr.:
17. 1813 ≡Oreoselinum chabraei (Jacq.) Bess., Cat. Hort. Cremen: 94. 1818 ≡Peucedanum
chabraei (Jacq.) Rchb. in Moessler, Handb. Gewächse, ed. 2, 1: 448. 1827 ≡Palimbia chabraei
(Jacq.) DC., Prodr. 4: 176. 1830 ≡Pteroselinum chabraei (Jacq.) Rchb., Fl. Germ. Exc. 2: 453.
1832 ≡Caroselinum chabraei (Jacq.) Griseb., Spic. Fl. Rumel. Bithyn. 1: 374. 1843 ≡Schlos-
seria chabraei (Jacq.) Schloss. & Vuk., Fl. Croat.: 474. 1869 ≡Holandrea carvifolia (Vill.)
Reduron & al. in J. Bot. Soc. Bot. France 1: 93. 1997. – Lectotype (designated by Frey 1989:
280): [icon] Selinum carvifolia Chabraei ex Crantz, Stirp. Austr. Fasc. 3, t. 3, fig. 2. 1767.
=Ligusticum decussatum Moench, Method. Pl.: 81. 1794, p.p.
=Selinum podolicum Bess., Prim. Fl. Galic. 2: 392. 1809 ≡Oreoselinum podolicum (Bess.) M.
Bieb., Fl. Taur.-Cauc. 3: 210. 1819 ≡Peucedanum podolicum (Bess.) Eichw., Naturhist. Skizze:
155. 1830 ≡Peucedanum chabraei var. podolicum (Bess.) DC., Prodr. 4: 176. 1830.
=Selinum chabraei var. aestivale Holandre, Fl. Moselle, ed. 1: 146. 1829 ≡Peucedanum
chabraei var. aestivale (Holandre) Rouy & Camus, Fl. France 7: 386. 1901.
=Selinum chabraei var. autumnale Holandre, Fl. Moselle, ed. 1: 146. 1829 ≡Peucedanum
chabraei var. autumnale (Holandre) Rouy & Camus, Fl. France 7: 386. 1901.
=Pastinaca selinoides Vis. in Flora 12 (Ergänzungsbl. 1): 10. 1829 ≡Peucedanum chabraei var.
selinoides (Vis.) Vis., Fl. Dalmat.: 51.1850 ≡Peucedanum selinoides (Vis.) Fritsch., Excurs. Fl.
Oesterr.: 379. 1922, non DC. 1830.
=Peucedanum heterophyllum Vis. in Cat. Sem. Horti Patavini 3: 4. 1836 ≡Schlosseria hetero-
phylla (Vis.) Vuk. in Oesterr. Bot. Z. 7: 350. 1857 ≡Palimbia chabraei var. heterophylla (Vis.)
Schur, Enum. Pl. Transsilv.: 265. 1866.
=Palimbia decussata Schur, Sert. Fl. Transsilv.: 30. 1853.
=Peucedanum euphimiae Kotov in Bot. Zhurn. (Kiev) 1(2): 278. 1940.
Fruit structure.–SeeFig.5A.
Distribution. – Austria, Belgium, Bosnia & Herzegovina, Bulgaria, Czech Rep., Croatia, France,
Germany, Hungary, Italy, Luxembourg, Moldavia, the Netherlands, Romania, Russia, Serbia,
Slovakia, Spain, Switzerland, Ukraine.
Ref. – Under Peucedanum carvifolia: Lange (1880: 44), Calestani (1905: 232), Burnat (1906: 208,
in nota), Thellung (1926: 1375, fig. 2518), Hayek (1927: 1034), Beck-Mannagetta (1927: 470),
Degen (1937: 506), Rohlena (1942: 225), Stojanov & Stefanov (1948: 856), Dostal (1950: 1059),
Janchen ( 1956: 432), Chrtek & Hendrich (1962: 137), Soó (1966: 481), Tutin (1968: 362), Holub
(1972: 40), Rompaey & Devlosalle (1972: map 644), Nikoli6(1973: 282), Guinochet & Vilmorin
(1975: 475), Kuzmanov & Andreev (1982: 228, t. 46), Pignatti (1982: 233), Molero & Rovira
(1985: 537), Frey (1989: 280, fig. 37), Bruck (1988: 69), Fabri (1993: 312), Grulich (1997: 469, t.
94, fig. 2), Lukac& Regula-Revilacqua (1997: 112), Lauber & Wagner (1998: 772), Wisskirchen &
Haeupler (1998: 360), Guillen & Lainz (2003: 351, fig. 113). – Under Imperatoria chabraei:
Sprengel (1818: 64). – Under Oreoselinum chabraei: Marchall von Bieberstein (1819: 209), Besser
(1822: 12). – Under Peucedanum chabraei: Gaudin (1828: 330, comb. superfl.), Ledebour (1844:
308, comb. superfl.), Visiani (1850: 51), Todor (1958: 604), Fournier (1961: 688). – Under
Palimbia chabraei: Schur (1866: 265). – Under Selinum podolicum: Hoffmann (1816: 155). – Un-
der Peucedanum podolicum: Shishkin (1951: 195). – Under Peucedanum chabraei var. podolicum:
478 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Willdenowia 37 – 2007 479
Fig. 5. Schematic transects of mericarps – A: Dichoropetalum carvifolia; origin: Moravia orient.-merid., Un
Brod, Nivnice, 1933, Podpera 1246 (LE); B: D. pschawicum; origin: Russia, N Ossetia, valley of Bad river,
1.6.1977, Pimenov &al.168 (MW); C: D. oligophyllum; origin: Greece, Makedonia orient., prov. Senres,
23.7.1979, Greuter (E); D: D. achaicum; origin: Bouraïkos, au-dessous de village de Zachloron (Achaïe,
Gréce), 8.7.1896, Saint Lager (G); E: D. graminifolium; origin: Turkey, A4 Kastamonu, 20.9.1996, Vural 7059
(GAZI); F: D. munbyi; origin: Algerie, dep. d’Alger, terrains argieux humices pres de l’Oued Smar, 15.10.1953,
Dubois (W). – Abbreviations: dc = commissural secretory ducts, df = secretory ducts in furrows, dr = secretory
duct in distal part of rib, es = endosperm, ex = exocarp, p = parenchyma cells without pits, pp = parenchyma
cells with lignified pitted walls, vb = vascular bundles. – Scale bars = 1 mm.
Todor (1958: 606), Fabri (1993: 313, in nota). – Under Peucedanum chabraei var. aestivale: Todor
(1958: 606). – Under Peucedanum chabraei var. autumnale: Todor (1958: 606), Fabri (1993: in
nota). – Under Peucedanum carvifolia var. selinoides: Degen (1937: 506).
(2) Dichoropetalum schottii (Bess.) Pimenov & Kljuykov, comb. nov. Peucedanum schottii
Bess. in Candolle, Prodr. 4: 178. 1830 Holandrea schottii (Bess.) Reduron & al. in J. Bot. Soc.
Bot. France 1: 93. 1997. – Lectotype (designated by Hartvig 1986): “Cult. Crem. Besser”
(G-DC!; isotype (Vinogradova 2003): LE!).
=Peucedanum petraeum W. D. J. Koch, Syn. Fl. Germ., ed. 1: 304. 1837 Peucedanum schottii
var. petraeum W. D. J. Koch, Syn. Fl. Germ., ed. 2: 334. 1843 Peucedanum schottii subvar.
petraeum (W. D. J. Koch) Burnat, Fl. Alp. Marit. 4: 206. 1906. – Holotype: Italia “In rupibus et
locis asperis saxosis (zwischen Triest u. Fiume an Felsen der neuen Straße), Noe” (W; isotype: G).
Distribution. – Albania, Bosnia & Herzegovina, Croatia, France, Greece, Italy, Serbia.
Ref. – Under Peucedanum schottii: Hy (1901:105), Calestani (1905: 231), Burnat (1906: 205),
Thellung (1926: 1377: fig. 2519), Hayek (1927: 1034), Beck-Managetta (1927: 462), Degen
(1937: 507), Fournier (1961: 689), Tutin (1968: 362), Nikoli6(1973: 284, t. 54, fig. 2), Guinochet
& Vilmorin (1975: 474), Pignatti (1982: 233), Demiri (1983: 348), Molero & Rovira (1985: 537),
Hartvig (1986: 718), Frey (1989: 281, fig. 37), Bolòs & Vigo (1990: 472), Qosja (1992: 402, fig.
749), Lukac& Regula-Revilacqua (1997: 112), Guillen & Lainz (2003: 353). – Under Peuce-
danum schottii var. petraeum: Rohlena (1942: 225).
Note. – Burnat (1906) and Thellung (1926) gave Oenanthe karsthia Hacq. as a synonym of this
species. Reduron & al. (1997) noted, however, that the fruit description of O. karsthia does not
corresponds to the Peucedanum fruit type. J.-P. Reduron kindly informed us that herbarium mate-
rial of this species is probably lost. The species description by Besser of Peucedanum schottii was
published in Candolle’s “Prodromus” and the origin “In Vohlynia & Podolia” apparently refers to
a plant cultivated in the botanical garden at Kiew, because P. schottii is not distributed in the
Ukraine and adjacent countries, where only P. carvifolia occurs.
(3) Dichoropetalum pschawicum (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum pscha-
wicum Boiss., Fl. Orient. 2: 1020. 1872 ≡Holandrea pschawica (Boiss.) Reduron & al. in J. Bot.
Soc. Bot. France 1: 93. 1997. – Holotype: Georgia “Caucasus, Pschawia supra lacum Tana inter
1580-1200 hec, 23.9.1860, Ruprecht” (G-BOIS!; isotype: LE!)
–P. chabraei auct. non Rchb. : Boissier, Fl. Orient. 2: 1020.1872.
–P. carvifolia auct. non Vill. : Grossheim, Fl. Kavkaza 3: 182. 1932.
Fruit structure.–SeeFig.5B.
Distribution. – Russia (N Caucasus: Krasnodar Terr., Adygea, Stavropol Terr., Kabardino-Bal-
karia, N Ossetia), Georgia.
Ref. – Under Peucedanum pschawicum: Lipsky (1899: 326), Grossheim (1932: 180), Shishkin
(1951: 196), Tamamschjan (1967: 116, fig. 125), Mandenova (1984: 268), Menitsky (1991: 1761),
Ivanov (2001: 101).
(4) Dichoropetalum alpigenum (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum alpi-
genum Boiss., Fl. Orient. 2: 1020. 1872. – Holotype: Turkey “In alpibus Ponti Lazici loco non
indicato, K. Koch” (G-BOIS!).
Distribution. – Turkey: N (Pontic) Anatolia. Known only from the type gathering.
Note. – This species was regarded by Chamberlain (1972) and Frey (1989) as a synonym of Peu-
cedanum carvifolia. The large disjunction between Lazistan and the European area of Dicho-
ropetalum carvifolia casts doubt on their conspecifity, especially as Boissier (1872) showed at
480 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
least three differences between these species. Although the type material is scanty, some of these
differences can be confirmed. As no similar species are known in adjacent territories (the nearest is
D. pshawicum from the Main Caucasian Ridge), specific distinction of D. alpigenum is very prob-
able. Further search for the species seems to be necessary.
(5) Dichoropetalum oligophyllum (Griseb.) Pimenov & Kljuykov, comb. nov. ≡Seseli oligo-
phyllum Griseb., Spic. Fl. Rumel. Bithyn. 1: 359. 1843 ≡Peucedanum oligophyllum (Griseb.)
Vandas in Magyar Bot. Lapok 4: 110.1905. – Holotype: Makedonia “In pratis prope Mandani,
Mt Kobelitza in Scardi Macedoniae, 4000', Grisebach 936” (GOET).
=Peucedanum aequiradium Velen. in Sitzungsber. Königl. Böhm. Ges. Wiss., Math. Naturwiss.
Cl. 1889: 36. 1890 ≡Peucedanum oligophyllum subsp. aequiradium (Velen.) Achtarov in Izv.
Bulg. Bot. Druz. 9: 65. 1943 ≡Peucedanum carvifolia subsp. aequiradium (Vel.) Stoj. & Steph.,
Fl. Bulg.: 835. 1925. – Type: Bulgaria “In graminosis alpinis siccis praesertim inter Vaccinia in
cacumine montis Rujen montium Osogovska Planina, 1887, Velenovsky”(PRM).
Fruit structure.–SeeFig.5C.
Distribution. – Albania, Bulgaria, Greece, F.Y.R. Macedonia, Serbia.
Ref. – Under Seseli oligophyllum: Boissier (1872: 965), Demiri (1983: 3480). – Under Peuce-
danum oligophyllum: Hayek (1927: 1033), Stojanov & Stefanov (1948: 856), Tutin (1968: 362),
Strid & Papanicolaou (1981: 74), Kuzmanov & Andreev (1982: 229), Hartvig (1986: 717), Frey
(1989: 279, fig. 37), Qosja (1992 : 401, fig. 748). – Under Peucedanum aequiradium: Hayek
(1927: 1033), Nikoli6(1973: 282), Hartvig (1986: 717). – Under Peucedanum oligophyllum
subsp. aequiradium: Tutin (1968: 362, comb. superfl.), Strid & Papanicolaou (1981: 74)
(6) Dichoropetalum achaicum (Halacsy) Pimenov & Kljuykov, comb. nov. ≡Peucedanum achai-
cum Halacsy, Consp. Fl. Graec. Suppl. 1: 42. 1908. – Type: Greece “In rupibus vallis fluminis
Voreikos infra Zachloru Achaiae [Bouraïkos, au-dessous de village de Zachloron (Achaïe,
Gréce)], 8.7.1896, Saint Lager” (G!).
Fruit structure.–SeeFig.5D.
Distibution. – Greece.
Ref. – Under Peucedanum achaicum: Hayek (1927: 1033), Tutin (1968: 362).
(7) Dichoropetalum graminifolium (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum
graminifolium Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 314. 1844. – Holotype: Turkey “Olympe
de Bithynie, Brousse [Bursa], 8.1830, Aucher-Eloy 3758” (G!; isotypes: K!, P!).
Fruit structure.–SeeFig.5E.
Distribution. – Turkey: N (Pontic) Anatolia: Bursa, Kastamonu, Samsun; Central Anatolia:
Sivas.
Ref. – Under Peucedanum graminifolium: Tchihachev (1860: 435), Boissier (1872: 1017), Cham-
berlain (1972: 475), Frey (1989: 282, map 37), Heller & Heyn (1993: 41), Vural & Adigüzel
(1996: 60, fig. 1), Parolly & Nordt (2004: 141).
(8) Dichoropetalum isauricum (Parolly & Nordt) Pimenov & Kljuykov, comb. nov. ≡Peuce-
danum isauricum Parolly & Nordt in Willdenowia 34: 135. 2004. – Holotype: [Turkey], C4 An-
talya, NE Demirtas (Richtung Pass) [towards pass], 840 m, lichter Waldhang, Waldsaum [open
forest slope, forest fringe], Exp. W., 7.9.2001, Ulrich 1/12 (STU).
Fruit structure. – See Parolly & Nordt (2004).
Distribution. – Turkey (S Anatolia: Antalya).
Willdenowia 37 – 2007 481
(9) Dichoropetalum munbyi (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum munbyi
Boiss., Diagn. Pl. Orient., ser. 2, 2: 89. 1854. – Lectotype (designated here): Algeria “Ad vias
circa pagum la Senia prov. Oran, Balansa 598” (G!; isotype: K!); paralectotype: Oran, La Senia,
in humidis salinis, 5.8.1850, Munby (K!).
Fruit structure.–SeeFig.5F.
Distribution. – Algeria, Morocco.
Ref. – Under Peucedanum munbyi: Quézel & Santa (1963: 671).
1.2. Dichoropetalum sect. Stridia Pimenov & Kljuykov, sect. nov.
Type: Dichoropetalum lavrentiadis (Strid & Papan.) Pimenov & Kljuykov
Plantae polycarpicae, caulibus eramosis, aphyllis. Folia radicalia ambitu ovata vel triangulata.
Umbellae 6-13-radiatae. Petala alba. Cellulae mesocarpii paranchymaticae e latere dorsali elig-
nescentes. Vittae valleculares paucae vel nullae.
1. Stems densely covered at the base by a fibrous collar of leaf remains; mericarps with soli-
tary vallecular vittae .................... (10)D. lavrentiadis
– Stems at the base without a collar of leaf remains; mature mericarps without vallecular
vittae ............................. (11)D. stridii
(10) Dichoropetalum lavrentiadis (Strid & Papan.) Pimenov & Kljuykov, comb. nov. ≡Peuce-
danum lavrentiadis Strid & Papan. in Bot. Not. 133: 524. 1980. – Holotype: “Greece, Nom. Flo-
rinis/Pellis, Mt Kajmakcalan (Voras Oros), summit area, 2450-2520 m, a few hundred metres
from the Jugoslavian border, 17.8.1979, Strid & Papanicolaou 16599” (C; isotypes: ATH, B, G!,
LD).
Distribution. – Greece, Serbia, Albania.
Ref. – Under Peucedanum lavrentiadis: Hartvig (1986: 720, fig. 45, A-E), Frey (1989: 284, fig. 27).
(11) Dichoropetalum stridii (Hartvig) Pimenov & Kljuykov, comb. nov. ≡Peucedanum stridii
Hartvig in Strid, Mt. Fl. Greece 1: 720, fig. 45 F. ≡Peucedanum lavrentiadis subsp. multicaulis
Strid & Papan. in Bot. Not. 133: 524. 1980. – Holotype: “Greece, Nomos Ioannina, Ep. Konitsa,
Mt. Smolikas, SE side, place named Gangarantza, 2200-2400 m, on peridotite and serpentine,
25.7.1971, Stamatiadou 13511 bis” (ATH; isotypes: C, G!).
Fruit structure. – See Fig. 6A.
Distribution. – Greece.
Ref. – Under Peucedanum stridii: Frey (1989: 283, fig. 26).
1.3. Dichoropetalum sect. Dichoropetalum
Plantae polycarpicae, caulibus eramosis vel ramosis. Laminae foliorum ambitu ovatae vel trian-
gulatae, superiores integrae, breves. Umbellae 3-9-radiatae. Petala flava. Cellulae mesocarpii
parenchymaticae e latere dorsali elignescentes. Vittae valleculares solitariae.
1. Stemslow,upto20cmhigh;basalprimaryleafsegmentssessile ......... 2
– Stems25-100cmhigh;basalprimaryleafsegmentswithshortpetiolules ...... 3
2. Terminal leaf lobes lanceolate, dentate, not crowded; stems erect or spreading, commissural
vittae in mericarps present ................... (12)D. alpinum
– Terminal leaf lobes linear, crowded; stems prostrate; commissural vittae in mericarps ab-
sent ............................. (15)D. aureum
482 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
3. Leaves long-persistent; stems up to 100 cm high; fruits elliptic in outline; dorsal mericarp
ribs shortly keeled, marginal ribs shortly winged, triangular in cross section .....
............................. (13)D. depauperatum
– Leaves withered at flowering; stems 120-200 cm high; fruits obovate in outline; dorsal
mericarpribsfiliform,marginalribsnarrowlywinged........ (14)D. junceum
Willdenowia 37 – 2007 483
Fig. 6. Schematic transects of mericarps – A: Dichoropetalum stridii; origin: Greece, Nomos Ioannina, Ep.
Konitsa, Mt Smolikas, SE side, place named Gangarantza, 2200-2400 m, on peridotite and serpentine,
25.7.1971, Stamatiadou 13511bis (G); B: D. alpinum; Turkey, C6 Maras, 1600-2000 m, step, 10.8.1991,
Aytaç &Duman 4145 (GAZI); C: D. depauperatum; origin: Syria, Akher Dagh, 6000, Haradjan 1572 (LE);
D: D. junceum; origin: Israel, Jerusalem, 12.10.1951, Grizi (W); E: D. aureum; origin: Turkey, Anti-Taurus,
à 12 lieux à l’ESE de Cesarée (Cappadoce), 6.8.1856, Balansa 1007 (P). – Scale bars = 1 mm; for the abbrevi-
ations see caption of Fig. 5.
(12) Dichoropetalum alpinum Fenzl, Pug. Pl. Nov. Syr.: 57. 1842 ≡Johrenia alpina (Fenzl) Fenzl
in Russeger, Reisen 1: 961, t. 17. 1843 . – Holotype: Turkey “in alpibus Tauri occidentalis [In
monte Tauro], 8.1836, Kotschy 213” (G-BOIS!; isotypes: BM, E!, G!, K!, L!, LE!, M!, P!, US,
W!, WAG).
=?Peucedanum depauperatum var. alpinum Boiss., Fl. Orient. 2: 1019. 1872. – Lectotype (des-
ignated here): Israel/Lebanon/Syria “In summo Hermone [Hermon, regio alpino], 5.7.1846,
Boissier” (G-BOIS!); paralectotypes: Lebanon “locis super nivalibus derelictis in jugo versus
Baalbeck, 7500', 30.7.1855, Kotschy 357” (G-BOIS!, W!); Lebanon “in cacuminibus inter Ya-
mouny & Dimam,18.7.1864, Blanche 3221” (G-BOIS!).
Fruit structure. – See Fig. 6B. Type sheets of the species contain a packet with fruits, but these
fruits do not belong to the type plants, which are at anthesis. These fruits are the most probably
from Johrenia tortuosa, growing in the same region. True D. alpinum fruits were investigated on
the basis of another gatherings of the species.
Distribution. – Turkey (S Anatolia: Içel, Kahramanmara=), Lebanon.
Ref. – Under Johrenia alpina: Tchihachev (1860: 440), Bornmüller (1930: 47), Hiroe (1958: 186;
1979: 232, p.p.), Heller & Heyn (1993: 32).
(13) Dichoropetalum depauperatum (Boiss. & Balansa ex Boiss.) Pimenov & Kljuykov, comb.
nov. ≡Peucedanum depauperatum Boiss. & Balansa ex Boiss., Diagn. Pl. Orient., ser. 2, 5: 98.
1856. – Holotype: Turkey “Région montagneux superieure de Taurus, au-dessus de Boulgar-
maden, 15.9.1855, Balansa 617” (G-BOIS!; isotypes: G, GH, K!, P!, W!).
=Johrenia berytea Boiss. & Hausskn. ex Boiss., Fl. Orient. 2: 1012. 1872. – Holotype: Turkey
“In rupestribus montis Berytdagh Cataoniae 7000'-9000' [In rupestr. Berytdagh 7-9000'],
16.8.1865, Haussknecht 1158” (G-BOIS!; isotype: JE!).
Fruit structure.–SeeFig.6C.
Distribution. – Lebanon, Syria, Turkey (Central Anatolia: Afyon, Isparta, Kayseri, Nid8e; S Ana-
tolia: Antalya, Içel, Adana, Kahramanmara=, Hatay; E Anatolia: Malatya).
Ref. – Under Peucedanum depauperatum: Tchihachev (1860: 435), Post (1896: 363, fig. 190),
Handel-Mazzetti (1913: 52), Post (1932: 554, fig. 350), Thiébaut (1936: 167), Rechinger (1957:
315), Hiroe (1958: 179; 1979: 1562, p.p), Mouterde (1970: 648, t. 311, fig. 1), Chamberlain (1972:
436), Heller & Heyn (1993: 41), Fragman & al. (1999: 44). – Under Johrenia berytea: Bornmüller
(1930: 48), Chamberlain (1972: 437), Heller & Heyn (1993: 33).
(14) Dichoropetalum junceum (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Johrenia juncea
Boiss., Diagn. Pl. Orient., ser. 1, 10: 33. 1849 ≡Peucedanum junceum (Boiss.) Mouterde in Fl.
Djebel Druze: 161. 1953, non Spreng. 1820 ≡Peucedanum mouterdei M. Hiroe in Umbell.
World: 1574. 1979. – Holotype: Lebanon “Ad margines agrorum sterilium in Antilibano inter
Rascheya et radices Hermonis [circa Rascheya], 7.1846, Boissier” (G-BOIS!; isotype: G!).
=Peucedanum spreitzenhoferi Dingler in Flora 66: 210. 1883. – Syntypes: Lebanon “In campis
aridis lapidosis Antilibani in Wadi Faluj supra Jubb Jenin Coelosyriae, 3000', Dingler”(B,de
-
stroyed); Israel “Loco Nikephori dicto Palestinae, Spreitzenhofer” (B, destroyed).
=?Johrenia porteri Post ex Boiss., Fl. Orient., Suppl.: 266. 1888, quoad typum. – Holotype:
Turkey “In Syriae borealis collibus siccis ad Kapukham ditionis Marasch [Kapu bham dagh],
17.9.1887, Post” (G!; isotypes: BEI! G-BOIS!).
Fruit structure.–SeeFig.6D.
Distribution. – Israel, Jordan, Lebanon, Syria, Turkey (SW Anatolia: Hatay, Kahramanmara=).
Ref. – Under Johrenia juncea: Boissier (1872: 1013), Post (1896: 363; 1932: 553), Bornmüller (1930:
46), Thiébaut (1936: 166). – Under Peucedanum junceum: Mouterde (1970: 648, t. 311, fig. 2), Cham-
berlain (1972: 480), Heller & Heyn (1993: 41), Fragman & al. (1999: 44). – Under Peucedanum
484 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
spreitzenhoferi: Post (1932: 554), Thiébaut (1936: 167), Grünberg-Fertig & Zohary (1970: 303),
Zohary (1972: 440, t. 639), Boulos (1977: 99), Al-Eisawi (1982: 178), Parolly & Nordt (2004: 141).
(15) Dichoropetalum aureum (Boiss. & Balansa ex Boiss.) Pimenov & Kljuykov, comb. nov. ≡
Johrenia aurea Boiss. & Balansa ex Boiss., Diagn. Pl. Orient., ser. 2, 6: 81. 1859. – Holotype:
Turkey “In parte superiori monto Aslan dach Antitauri in Cappadocia [Région alpine superieure
de l’Aslan-Dach l’un pas pies de l’Anti-Taurus a 12 lieux à l’ESE de Cesarée (Cappadoce)],
6.8.1856, Balansa 1007” (G-BOIS!; isotypes: G!, MANCH! P!, W!).
Fruit structure.–SeeFig.6E.
Distribution. – Turkey (Central Anatolia: Kayseri; S Anatolia: Içel), Lebanon, Israel.
Ref. – Under Johrenia aurea: Tchihachev (1860: 441), Boissier (1872: 1013), Post (1896: 363),
Bornmüller (1930: 48), Post (1932: 553), Thiébaut (1936: 165), Mouterde (1970: 651, t. 314,
fig. 1), Chamberlain (1972: 437), Heller & Heyn (1993: 32).
1.4. Dichoropetalum sect. Scoparia Pimenov & Kljuykov, sect. nov.
Type: Dichoropetalum scoparium (Boiss.) Pimenov & Kljuykov
Plantae monocarpicae vel polycarpicae, caulibus ramosis, foliosis. Laminae foliorum ambitu
lanceolatae, superiores longae vel breves. Umbellae 3-9-radiatae. Petala flava. Cellulae meso-
carpii omnes parenchymaticae lignescentes vel e latere dorsali elignescentes. Vittae valleculares
nullae, solitariae vel paucae.
1. Monocarpicplants;stemsglabrous;terminalleaflobeslanceolatetoovate ..... 2
– Polycarpic plants; stems pubescent; terminal leaf lobes linear ...........3
2. Terminal leaf lobes lanceolate, without teeth or lobes or notches; stylopods conical; mar-
ginal mericarp ribs narrow, winged, vallecular vittae distinct, solitary; stems branched from
themiddlepart ...................... (16)D. golestanicum
– Terminal leaf lobes ovate, dentate; stylopods short-conical; marginal mericarp ribs shortly
winged, slightly swollen, vallecular vittae absent; stems strongly branched from the base .
............................. (19)D. ramossimum
3. Petioles and leaf blades soft; stems virgate, thin, slightly branched; upper cauline leaves with
long blades; dorsal mericarp ribs thin, filiform, marginal ribs narrow, winged, mericarp with
distinctvallecularvittae,secretoryductsinribsthinorabsent .... (17)D. scoparium
– Petioles and leaf blades rigid; stems multi-branched dichotomously; upper cauline leaves
with short blades; all mericarp ribs inflated, mericarp usually with inconspicuous vallecular
vittae,secretoryductsinribslarge .............. (18)D. platycarpum
(16) Dichoropetalum golestanicum (Rech. f.) Pimenov & Kljuykov, comb. nov. ≡Johrenia
golestanica Rech. f., Fl. Iran. 162: 376, t. 304. 1987. – Holotype: Iran “Persia, Gorgan: Jangal-e
Golestan prope Tang-e Rah, 420-500 m, 13.-14.10.1976, Terme & Matine 35109-E” (W!).
Fruit structure.–SeeFig.7A.
Distribution. – Turkmenistan, Iran (N: Golestan).
Ref. – Under Johrenia golestanica: Heller & Heyn (1993: 33), Jalili & Jamzad (1999: 677).
(17) Dichoropetalum scoparium (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Johrenia scoparia
Boiss., Diagn. Pl. Orient., ser. 1, 6: 61. 1846 ≡Peucedanum scoparium (Boiss.) Boiss., Diagn. Pl.
Orient., ser. 2, 2: 90. 1856 ≡Johreniopsis scoparia (Boiss.) Pimenov in Rechinger, Fl. Iran. 162:
455, t. 411. 1987 – Holotype: Iran “[Fars] In argillosis acidis alpis Kuh-Daena [Kuh-e-Dinar],
17.7.1842, Kotschy 676” (G-BOIS!; isotypes; BM!, C, E!, K!, LE!, M!, P!, US, W!).
Fruit structure.–SeeFig.7B.
Willdenowia 37 – 2007 485
486 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Fig. 7. Schematic transects of mericarps – A: Dichoropetalum golestanicum; origin: Iran, Golestan, Gorgan,
Jangal-e, Tang-e Gol, 600-700 m, 13.-14.10.1976, Termé &Matiné 35110 (E); B: D. scoparium; origin: Iran,
prov. Chaharmahal-e Bakhtiari, Sabz Kuh, Kuh-e Darreh, 2300-2750 m, 19.8.1986, Mozaffarian 58181
(TARI); C: D. platycarpum; origin: Iran, Khorassan, N Nishapour, Mirabadriva, 1750 m, 10.7.1995, Faghi-
haia &al.26066 (MD); D: D. ramosissimum; origin: Iran, Mazandaran, Kelardascht, Rudbarak, 1700 m, Mo-
zaffarian 72066 (TARI); E: D. aromaticum; origin: Turkey, Van, Çatak, 25.7.1954, Davis &Polunin 23240
(ANK); F: D. paucijugum; origin: Azerbaidzhan, Nakhichevan, Shakhbuz distr., Karababa, Kjukjuchai river,
12.6.1977, Pimenov &al.488 (MW). – Scale bars = 1 mm; for the abbreviations see caption of Fig. 5.
Distribution. – Iran (W: Azarbayjan, Kordestan, Kermanshah, Hamadan, Lorestan; Central:
Arak, Chaharmahal va Bakhteyari; S: Boyerahmad va Kohgiluye, Fars), Iraq.
Ref. – Under Johrenia scoparia: Boissier (1872: 1019). – Under Johreniopsis scoparia: Townsend
(1964: 74), Heller & Heyn (1993: 43), Jalili & Jamzad (1999: 678).
(18) Dichoropetalum platycarpum (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Johrenia platy-
carpa Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 308. 1844. – Holotype: Iran “In monte Elbourz,
Aucher-Eloy 4582” (G-BOIS!; isotypes: G!, P!, K!).
Fruit structure.–SeeFig.7C.
Distribution. – Turkmenistan, Afghanistan, Iran (Central: Semnan, Tehran; E: Khorassan; S:
Fars, Kerman).
Ref. – Under Johrenia platycarpa: Boissier (1844: 80; 1872: 1012), Bornmüller (1930: 49),
Mozaffarian (1983: 317), Rechinger (1987: 377), Pimenov & Kljuykov (1992: 112, fig. 1), Heller
& Heyn (1993: 33).
(19) Dichoropetalum ramosissimum (Mozaff.) Pimenov & Kljuykov, comb. nov. ≡Johrenia
ramosissima Mozaff. in Bot. Zhurn. (St. Petersburg) 88(4): 104, fig. 1. 2003. – Holotype: Iran
“Mazandaran: Kelardascht, Rudbarak, 1700 m, Mozaffarian 72066” (TARI!).
Fruit structure. – See Fig. 7D.
Distribution. – Iran (N: Mazandaran).
1.5. Dichoropetalum sect. Parajohrenia Pimenov & Kljuykov, sect. nov.
Type: Dichoropetalum paucijugum (DC.) Pimenov & Kljuykov
Plantae monocarpicae, caulibus ramosis. Laminae foliorum ambitu lanceolatae, superiores inte-
grae, breves. Umbellae 7-13-radiatae. Petala flava. Cellulae mesocarpii parenchymaticae omnes
membranis lignescentibus. Vittae valleculares commissuralesque obsoletae.
1. Plants with strong smell; stems almost round in cross section; bractlets linear; stylopods
short-conical; lateral umbels not overtopping the central one . . . (20) D. aromaticum
– Plants without strong smell; stems considerably ribbed; bractlets lanceolate; stylopods flat;
lateral umbels clearly overtopping the central one ........ (21)D. paucijugum
The position of this group of two species in Dichoropetalum remains unstable. Some small
changes in the coding of the character states can produce a grouping, in which sect. Parajohrenia
is joined with Johrenia, as its most distant fragment. Such treatment might be nearer to traditional
taxonomy of the two species, but contradicts their character set.
(20) Dichoropetalum aromaticum (Rech. f.) Pimenov & Kljuykov, comb. nov. ≡Johrenia aro-
matica Rech. f., Fl. Iran. 162: 376, t. 303. 1987. – Holotype: Iran “Kurdistan, Sanandaj, 35º03'N,
46º57'E, 1200-1400 m, low rolling mountains of dark slaty rock with open herbaceous vegeta-
tion, 21.6.1963, Jacobs 6973” (W!).
Fruit structure.–SeeFig.7E.
Distribution. – Iran (W: W Azarbayjan, Kordestan, Kermanshah), Turkey (E Anatolia: Van), Iraq.
(21) Dichoropetalum paucijugum (DC.) Pimenov & Kljuykov, comb. nov. ≡Ferula paucijuga
DC., Prodr. 4: 171. 1830 ≡Johrenia candollei Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 306. 1844
≡Johrenia paucijuga (DC.) Bornm. in Russk. Bot. Zhurn. 1-2: 9. 1910. – Holotype: Iran “in radi-
ces montium prope Badalan prope Khvoy, 7.6.1828, Szovits” (G-DC!).
Willdenowia 37 – 2007 487
=Johrenia persica Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 306. 1844. – Holotype: [Iran], Aucher-
Eloy 3664 (G!; isotype: P!)
=Seseli leucocoleum Stapf & Wettst. in Denkschr. Kaiserl. Akad. Wiss., Math.-Nat. Kl. 51, 2:
318. 1886. – Holotype: Iran “In locis rupestribus et vinetis prope Jalpan, 25.5.1882, Pichler”
(WU!).
Fruit structure.–SeeFig.7F.
Distribution. – Iran (W: W Azarbayjan, E Azarbayjan, Zanjan, Kordestan, Hamadan, Lorestan;
Central: Arak, Chaharmahal va Bakhteyari; S: Fars), Azerbaidzhan, Armenia.
Ref. – Under Johrenia paucijuga: Bornmüller (1911: 238), Grossheim (1927: 20, comb. superfl.),
Bornmüller (1930: 50), Grossheim (1932: 175), Shishkin (1951: 49, t. 5, fig.3), Karjagin (1955:
476), Tamamschjan (1967:104, map 115), Takhtajan & Fedorov (1972: 193), Mandenova (1973:
385, t. 175), Mozaffarian (1983: 122, 316), Rechinger (1987: 375, t. 302), Heller & Heyn (1993:
33).
1.6. Dichoropetalum sect. Johreniopsis (Pimenov) Pimenov & Kljuykov, comb.&stat.nov.≡
Johreniopsis Pimenov in Rechinger, Fl. Iran. 162: 454. 1987.
Type: Dichoropetalum seseloides (C. A. Mey.) Pimenov & Kljuykov
Plantae monocarpicae, caulibus ramosis foliosisque. Laminae foliorum ambitu lanceolatae, supe-
riores breves. Umbellae 6-14-radiatae. Petala flava. Cellulae mesocarpii parenchymaticae e latere
dorsali elignescentibus. Vittae valleculares solitariae vel paucae.
1. Stems covered by short hairs; mericarps with several vallecular vittae ........ 2
– Stems glabrous, mericarps with solitary vallecular vittae ............. 3
2. Umbel rays strongly unequal; bractlets subulate to linear; stylopods flat; dorsal mericarp
ribsfiliform ....................... (25)D. minutifolium
– Umbel rays ± equal; bractlets linear-lanceolate; stylopods short-conical; dorsal mericarp
ribsshortlykeeled..................... (26)D. palimbioides
3. Stems clearly ribbed ..................... (24)D. chryseum
– Stems round in cross section ........................ 4
4. Bractlets 2-3; terminal leaf lobes lanceolate; stylopods flat ..... (23)D. viittijugum
– Bractlets 3-7; terminal leaf lobes linear; stylopods conical ..... (22)D. seseloides
(22) Dichoropetalum seseloides (C. A. Mey.) Pimenov & Kljuykov, comb. nov. ≡Ferula sese-
loides C. A. Mey., Verz. Pfl. Casp. Meer.: 126. 1831 ≡Johrenia meyeri Boiss. in Ann. Sci. Nat.,
Bot., ser. 3, 1: 307. 1844 ≡Peucedanum meyeri (Boiss.) Boiss., Fl. Orient. 2: 1018. 1872 ≡Joh-
reniopsis seseloides (C. A. Mey.) Pimenov in Rechinger, Fl. Iran. 162: 455, t. 412. 1987. –
Holotype: Azerbaidzhan “in monte Beschbarmak, 21.7.1830, C. A. Meyer” (LE!, photo: W).
=Peucedanum conrathii Freyn in Bull. Herb. Boissier 3: 305. 1895. – Type: Georgia “Somchetia:
in collibus apricis circa Achtala, 7.1888, Conrath”.
Fruit structure.–SeeFig.8A.
Distribution. – Russia (N Caucasus: Daghestan), Iran (W: Kordestan; N: Golestan; Central: Sem-
nan, Chaharmahal va Bakhteyari), Azerbaidzhan, Georgia, Armenia, Turkey (N (Pontic) Anato-
lia: Coruh; E Anatolia: Kars, Bitlis).
Ref. – Under Peucedanum paucifolium: Shishkin (1951: 199), Karjagin (1955: 489), Tamamschjan
(1967: 115, map 124), Mandenova (1984: 267), Frey (1989: 282, map 37). – Under Peucedanum
meyeri: Lipsky (1899: 326), Grossheim (1932: 193), Chamberlain (1972: 476), Mandenova (1973:
405), Mozaffarian (1983: 124, t. 322). – Under Peucedanum conrathii: Grossheim (1932: 181). –
Under Johreniopsis seseloides: Heller & Heyn (1993: 43).
488 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
(23) Dichoropetalum vittijugum (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum vitti-
jugum Boiss., Fl. Orient. 2: 1018. 1872. – Lectotype (designated here): Greece “In regione infe-
riori montis Dirphyis Delphi Eubeae supra pagum Steni, 31.7.1858, Heldreich 1844” (G-BOIS!).
– Paralectotypes: Greece “In dumosis, radices Taygeti prope Gola inter frutices, 8.6.1846,
Sartori 1844” (G-BOIS!); “In monte Malevo Laconiae prope Hajos-Joannis, Orphanides 3194”
(G-BOIS!) “in mt. Chelmos prope Chalkinika, 18.-30.6.1852, Orphanides 2074” (G-BOIS!,
W[photo]); “prope Carpenisi Eurytaeniae ad rad. m. Veluchi, 9.8.1847, Samaritani”.
Fruit structure.–SeeFig.8B.
Willdenowia 37 – 2007 489
Fig. 8. Schematic transects of mericarps – A: Dichoropetalum seseloides; origin: N Azerbaidzhan, Besch-
barmak Mt, 10.8.1978, Pimenov &al.953 (MW); B: D. vittijugum; origin: Greece, Makedonia, Kastoria,
18.8.1988, Gardner &Gardner 4354 (E); C: D. chryseum; origin: Turkey, Antalya, between Kargi chai &
Belioter, 1000 m, 26.8.1947, Davis 14233 (E); D: D. minutifolium; origin: Bulgaria, 8.1902, Stribrny (E); E:
D. palimbioides; origin: Turkey, A7, Gümüshane, 10.8.1968, Baytop 14315 (E). – Scale bars = 1 mm; for the
abbreviations see caption of Fig. 5.
Distribution. – Albania, Bulgaria, Greece, Serbia, F.Y.R. Macedonia.
Ref. – Under Peucedanum vittijugum: Halacsy (1901: 640), Calestani (1905: 230), Hayek (1927:
1034), Rechinger (1943: 413), Stojanov & Stefanov (1948: 858), Tutin (1968: 362), Kuzmanov &
Andreev (1982: 226, t. 47), Demiri (1983: 348), Hartvig (1986: 718), Qosja (1992: 403, fig. 750).
(24) Dichoropetalum chryseum (Boiss. & Heldr. ex Boiss.) Pimenov & Kljuykov, comb. nov. ≡
Anethum chryseum Boiss. & Heldr. ex Boiss., Diagn. Pl. Orient., ser. 1, 10: 32. 1849 ≡Peuce-
danum chryseum (Boiss. & Heldr. ex Boiss.) D. F. Chamb. in Davis, Fl. Turkey 4: 475. 1972. –
Holotype: Turkey “In humilioribus Tauri Pamphylici [Pamphylia ad radices Tauri] inter Marla &
Adalia alt. 1000', 7.1845, Heldreich” (G-BOIS!; isotype: K!)
=Peucedanum chrysanthum Boiss., Diagn. Pl. Orient., ser. 2, 6: 86. 1859. – Lectotype (desig-
nated here): Turkey “In vineis prope Ouchak Phrygiae [Ouchak (Phrygie), vers 910 m d’alt.,
vignes], 8.1857, Balansa 1245” (G-BOIS!; isolectotypes: BM!, GH, JE!, K!, LE!, P!, US, W-photo).
Fruit structure.–SeeFig.8C.
Distribution. – Turkey (N (Pontic) Anatolia: Samsun; Central Anatolia: Kirikkale, Amasya,
Kütahya, U=ak, Denizli, Burdur, Isparta, Konya, Kir=ehir; W Anatolia: Manisa, Aydin, Mu8la; S
Anatolia: Antalya, Içel).
Ref. – Under Anethum chryseum: Tchihachev (1860: 436), Boissier (1872: 1026), Zohary (1972:
434). – Under Peucedanum chryseum: Heller & Heyn (1993: 41). – Under Peucedanum chry-
santhum: Tchihachev (1860: 435), Boissier (1872: 1018), Tamamschjan (1967: 115, map 124).
(25) Dichoropetalum minutifolium (Janka) Pimenov & Kljuykov, comb. nov. ≡Bunium minu-
tifolium Janka in Oesterr. Bot. Z. 22: 177. 1872 ≡Peucedanum minutifolium (Janka) Velen., Fl.
Bulg. Suppl.: 122. 1898 ≡Peucedanum vittijugum var. minutifolium (Janka) Kuzmanov & An-
drejev, Fl. Nar. Rep. Bblgarija 8: 228. 1982. – Type: Bulgaria “In steppis collinis mari nigro
vicinis inter Aitos & Burgas Thraciae orientalis, 2.7.1871, Janka”(BP).
=Peucedanum thracicum Velen. in Sitzungsber. Boehm. Ges. Wiss., Math. Naturwiss. Cl. 1892 :
379. 1893. – Type: “In collibus graminosis supra Stanimaka, 7.1892, Janka, Velenovsky &
Stribrny”.
Fruit structure.–SeeFig.8D.
Distribution.–Serbia,Croatia,Bulgaria.
Ref. – Under Peucedanum minutifolium: Hayek (1927: 1035), Nikoli6(1973: 285), Lukac&
Regula-Revilacqua (1997: 112).
(26) Dichoropetalum palimbioides (Boiss.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum
palimbioides Boiss., Fl. Orient. 2: 1021. 1872. – Holotype: Turkey “In Ponto Lazico ad Arda-
nutsch, K. Koch” (G-BOIS!).
=Malabaila carvifolia Boiss. & Balansa ex Boiss., Diagn. Pl. Orient., ser. 2, 6: 85. 1859 ≡
Pastinaca carvifolia (Boiss. & Balansa ex Boiss.) Koso-Pol. in Bull. Soc. Imp. Naturalistes
Moscou, ser. 2, 29: 112. 1916 ≡Peucedanum acuminatum M. Hiroe, Umbell. World: 1570. 1979. –
Holotype: Turkey “In vineis partis inferioris montis Alidagh prope Caesaream [Kayseri]
Cappadociae, 1300', 7.8.1866, Balansa 1016” (G!; isotypes: BM!, JE!, K!, US, W!).
=Peucedanum tomentellum Freyn & Sint. in Oesterr. Bot. Z. 44: 101. 1894. – Lectotype (desig-
nated here): Turkey “Paphlagonia, wilayet Kastamonu, Tossia: in collibus ad Karvak-Tschesme,
4.8.1892, Sintenis 4880” (B!; isolectotypes: BM!, G!, JE!, K!, LE!, P!, W!).
=Peucedanum fallax Freyn & Sint. in Oesterr. Bot. Z. 44: 102. 1894. – Lectotype (designated
here): Turkey “Paphlagonia, Tossia, in vineis montis Giaurdagh, 7.8.1892, Sintenis 4945”(B!;
isolectotypes: G!, JE!,K!, LE!, P!, W!).
490 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Fruit structure.–SeeFig.8E.
Distribution. – Turkey (N (Pontic) Anatolia: Kastamonu, Sinop, Çoruh; Central Anatolia: An-
kara, Kirikkale, Çankiri, Amasya, Tokat, Sivas, Konya, Nev=ehir, Kayseri; S Anatolia: Adana,
Kahramanmara=;EAnatolia:Gümü=hane, Bayburt, Erzurum, Malatya, Tunceli).
Ref. – Under Peucedanum palimbioides: Lipsky (1899: 326), Grossheim (1932: 182), Tamam-
schjan (1967: 116, map 125), Chamberlain (1972: 478), Heller & Heyn (1993: 42). – Under
Malabaila carvifolia: Tchihachev (1860: 460, “curvifolia”).
2. Johrenia DC. in Coll. Mém. 5 (Mém. Fam. Ombellifères): 54. 1829 [& Prodr. 4: 196. 1830]
Type: Johrenia dichotoma DC.
=Caroselinum Griseb., Spic. Fl. Rumel. Bithyn. 1: 374. 1843. – Type: Caroselinum distans
Griseb.
Plantae monocarpicae, caulibus basi residuis foliorum emortuorum obsoletis, glabris. Folia vaginis
longis, laminis ambitu ovatis vel triangulatibus, segmentis primariis basalibus sessilibus vel bre-
vepetiolulatis vel longepetiolulatis, lobis terminalibus linearibus vel lanceolatis; folia caulina su-
periora integra, brevia. Umbellae plerumque radiis valde inaequelongis, bracteis nullis. Bracteolae
herbaceae, subulatae, lineares vel lanceolatae, radiolis breviora. Dentes calycini obsoleti. Petala
flava. Stylopodia plana vel breveconica. Mericarpia dorso compressa, jugis inconspicuis, in
textum mesocarpii immersis, cellulis mesocarpii parenchymaticis lignescentibus, porosis, e latere
commissurali destructis circa carpophorum, hypendocarpiis evolutis, vittis vallecularibus nullis vel
paucis, maxime minutis, commissuralibus nullis, canaliculis jugalibus solitariis, frequenter mag-
nis. Endospermium ventre planum.
1. Basal primary leaf segments sessile; vallecular mericarp vittae distinct, several; umbels
3-5-rayed............................ (5)J. distans
– Basal primary leaf segments petiolulate; vallecular mericarp vittae absent or very thin;
umbels5-20-rayed ............................. 2
2. Umbels 15-20-rayed; stylopods flat ................ (2)J. selinoides
– Umbels 5-13-rayed; stylopods short-conical ...................3
3. Central part of dorsal mericarp surface in mature fruits green, clearly differing from white
spongy marginal part ..................... (1)J. dichotoma
– Central part of dorsal mericarp surface in mature fruits white, not differing in colour from
spongy marginal part ............................ 4
4. Stemsbranchingalmostfromthebase;mericarpsslightlycompresseddorsally.....
................................(3)J. polyscias
– Stems branching from the middle part; mericarp convex on dorsal surface . (4) J. tortuosa
(1) Johrenia dichotoma DC., Prodr. 4: 196. 1830. – Holotype: Lebanon “In Oriente ad Libanum,
de Labillardiere” (G-DC!).
=Peucedanum haradjianii Rech. f. in Ark. Bot., ser. 2, 5(1): 315, fig. 35. 1957. – Holotype: Tur-
key “Cassius: in monte Cassio, 4-5000', 6.1909, Haradjian 3924” (G; isotype: W!).
Distribution. – Turkey (Central Anatolia: Aksaray, Ankara, Isparta, Kayseri, Konya, Yozgat; W
Anatolia: Mu8la; S Anatolia: Adana, Antalya, Gaziantep, Hatay, Içel; E Anatolia: Erzincan,
Elazig), Greece (E Aegean Isls.), Iraq, Israel, Lebanon, Syria.
subsp. sintenisii Bornm. in Repert. Spec. Nov. Regni Veg. 28: 43. 1930.
Holotype: Turkey “Mardin, Balekri, 16.7.1888, Sintenis 1330” (JE!; isotypes: E! K! LD).
Distribution. – Turkey (S Anatolia: Içel, Kahramanmara=).
Fruit structure.–SeeFig.9A.
Willdenowia 37 – 2007 491
Ref. – Tchihachev (1860: 440), Boissier (1872: 1010), Post (1896: 551, fig. 188), Handel-Mazzetti
(1913: 51), Bornmüller (1930: 43, 1938: 285), Post (1932: 551, fig. 348), Mouterde (1970: 650, t.
313, fig. 1), Chamberlain (1972: 435), Heller & Heyn (1993: 33), Fragman & al. (1999: 35), Snogerup
& al. (2001: 149). – Under Johrenia dichotoma subsp. sintenisii: Chamberlain (1972: 435).
(2) Johrenia selinoides Boiss. & Balansa ex Boiss., Diagn. Pl. Orient., ser. 2, 5: 99. 1856. –
Holotype: Turkey “In faucibus Pylarum Ciliciarum (Gulek Bogasi) [Défile des Portus Cilicien-
nes, à 10 lieux au nord de Parscus], 9.1855, Balansa 616” (G-BOIS!; isotype: P!).
=Eriosynaphe kotschyana Fenzl ex Tchih., As. Min., Bot. 1: 433. 1860. – Lectotype (designated
here): Turkey “Tauro Cilicico, in Tauri alpes ‘Bulgar Dagh’, ad inclytas ‘Gullek Boghas’ in
regione montana 3800 ped., 1853, Kotschy” (W!).
Fruit structure.–SeeFig.9B.
Distribution. – Turkey (Central Anatolia: Konya; S Anatolia: Adana, Antalya, Hatay, Içel).
Ref. – Tchihachev (1860: 441), Boissier (1872: 1010), Post (1896: 362), Bornmüller (1930: 46),
Post (1932: 551), Thiébaut (1936: 166), Chamberlain (1972: 434, map 57, “silenoides”), Heller &
Heyn (1993: 33).
(3) Johrenia polyscias Bornm. in Repert. Spec. Nov. Regni Veg. 28: 44. 1930. – Holotype: Tur-
key “Agri Ancyritani ad meridiem versus oppidi, prope Cankai, in valle Kavakli-dere, c. 800 m,
16.7.1929” [“Galatia: ditionis oppidi Angora (Ancyra), in valle Kawakli-dere, c. 900 m, 16.7.
1929, Bornmüller 14139]” (B!; isotypes: E!, G!, K!, P!, W!).
Fruit structure.–SeeFig.9C.
Distribution. – Turkey (Central Anatolia: Ankara, Amasya, Tokat).
Ref. – Krause (1937: 114), Chamberlain (1972: 436, fig. 8), Heller & Heyn (1993: 33).
(4) Johrenia tortuosa (Fisch. & C. A. Mey.) D. F. Chamb. in Davis, Fl. Turkey 4: 436. 1972 ≡
Eriosynaphe tortuosa Fisch. & C. A. Mey., Ind. Sem. Horti Petrop. 5: 35. 1838. – Holotype: Tur-
key “In Anatolia [in Natolia] [? circa Amasya & Tokat], Wiedemann 35” ( LE!; isotypes: BM, P!)
=Johrenia fungosa Boiss., Diagn. Pl. Orient., ser. 1, 10: 33. 1849. – Holotype: Turkey “In mon-
tibus Lydiae [Tmolus supra Philadilphiam], 6.1842, Boissier” (G-BOIS!; isotypes: JE!, LIV!).
=Johrenia engleri Dingler in Flora 66: 212. 1883. – Holotype: Turkey “In lapidosis et rupes-
tribus montanis inter virgulta ad oppidum Biledschik Bithyniae, 7.1873, Dingler”(B).
–Johrenia dichotoma auct. non DC.: Boissier in Ann. Sci. Nat., Bot., ser. 3, 1: 304. 1844.
Fruit structure.–SeeFig.9D.
Distribution. – Turkey (N (Pontic) Anatolia: Bursa, Samsun; Central Anatolia: Ankara, Çorum,
Amasya, Tokat, Eski=ehir, Afyon, Burdur, Isparta, Konya, Ni8de, Yozgat, Kayseri; W Anatolia:
Izmir, Manisa; S Anatolia: Antalya, Içel, Adana), Syria.
Ref. – Heller & Heyn (1993: 33). – Under Johrenia fungosa: Tchihachev (1860: 441), Boissier
(1872: 1011), Post (1896: 362), Bornmüller (1930: 41), Post (1932: 552), Thiébaut (1936: 166),
Mouterde (1970: 650, t. 313, fig. 2). – Under Johrenia engleri: Boissier (1888: 266), Bornmüller
(1930: 45).
(5) Johreniadistans (Griseb.) Halacsy, Consp. Fl. Graec. 1: 634. 1901 ≡Caroselinum distans
Griseb., Spic. Fl. Rumel. Bithyn. 1: 374. 1843. – Type: Macedonia “In peninsula Hajion-Oros;
raro inter frutices sempervirentes pr. Hajianna in litore (substr. marmor)”.
=Johrenia graeca Boiss. & Spruner ex Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 305. 1844. –
Holotype: Greece “In parte orientali montis Hymetti Atticae versus prom. Sunium [im östlichen
Hymettus in Griechenland], Spruner” (G-BOIS!; isotypes: G!, W!).
492 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Willdenowia 37 – 2007 493
Fig. 9. Schematic transects of mericarps – A: Johrenia dichotoma; origin: Turkey, C5 Içel, Toros Da8lari,
near Çamliyayla, 37°09'N, 34°36'N, 18.8.96, Pimenov &al.T96-72 (MW); B: J. selinoides; origin: Turkey,
Antalya, distr. Alanya, Han Bo8azi forest, near Geyik Da., 1600 m, 30.8.1947, Davis 14713 (E); C: J.
polyscias; origin: Turkey, Portus Galaticus, 26.6.1890, Bornmüller 905 (K); D: J. tortuosa; origin: Turkey,
Tmolus & Sypulus, 8.1842, Boissier 908 (K); E: J. distans; origin: Greece, Mt Hymetus, Heldreich 910 (E). –
Scale bars = 1 mm; for the abbreviations see caption of Fig. 5.
=Johrenia thessala Bornm. in Repert. Spec. Nov. Regni Veg. 28: 37. 1930. – Holotype: Greece
“Thessala, Kabampaka, Hagios Stephanos, in fauce Karawa, 27.5.1896, Bornmüller 528”(LD).
Fruit structure.–SeeFig.9E.
Distribution. – Greece, F.Y.R. Macedonia.
Ref. – Calestani (1905: 224), Hayek (1927: 1026), Rechinger (1943: 410), Tutin (1968: 358).
3. Zeravschania Korovin in Bot. Mater. Gerb. Inst. Bot. Zool. Akad. Nauk Uzbeksk. SSR 12: 28.
Typus: Zeravschania regeliana Korovin
Plantae polycarpicae, caulibus rotundis, basi plerumque residuis foliorum emortuorum dense
tectis, glabris, rarius pubescentibus. Folia vaginis brevibus, laminis ambitu ovatis vel triangu-
laribus, segmentis primariis basalibus longepetiolulatis, lobis terminalibus linearibus, lanceolatis
vel ovatis, margine dentatis; folia caulina superiora integra, brevia. Umbellae radiis 3-15, plerum-
que subaequelongis, bracteis evolutis, integris. Bracteolae lanceolatae vel latelanceolatae, albo-
marginatae, vulgo radiolis breviorae. Dentes calycini obsoleti rarius breves. Petala alba vel flava.
Stylopodia breveconica, rarius conica. Mericarpia dorso convexa, jugis dorsalibus filiformibus,
marginatis brevetriangulatis, cellulis mesocarpii parenchymaticis a latere dorsali elignescentibus,
a latere commissurali non destructis, hypendocarpiis evolutis, vittis vallecularibus solitariis, com-
missuralibus binis. Endospermium ventre planum.
1. Lowerpartofstemcoveredbyshorthairs ............ (5)Z. stricticaulis
– Stemscompletelyglabrousinlowerpart.................... 2
2. Petalswhite................................ 3
– Petalsyelloworgreenishyellow ....................... 7
3. Terminalleaveslobeslanceolate ....................... 4
– Terminalleaflobesovate,dentate....................... 5
4. Stems hollow; leaves long-persistent, glabrous; mericarps ribs with secretory ducts ...
...............................(3)Z. regeliana
– Stems solid; leaves quickly withering, covered by short rigid hairs; mericarp ribs without
secretoryducts........................ (4)Z. scabrifolia
5. Calyx teeth broadly triangular, obtuse; stylopods conical ...... (6)Z. ferulifolia
– Calyx teeth inconspicuous; stylopods short-conical or dish-shaped ......... 6
6. High montane plants with thick branched caudex; stylopods dish-shaped .......
.............................. (7)Z. minjanensis
– Plants of low altitudes with taproots; stylopods short-conical ...... (8)Z. aucheri
7. Terminalleaflobeslanceolate,short .............. (1)Z. pauciradiata
– Terminal leaf lobes long, narrowly linear to filiform ............... 8
8. Leavesrigid;bracteoleswhite-membraneousatthemargins,lanceolatetoovate ....
............................. (2)Z. membranacea
– Leavessoft;bracteolesherbaceous,linear.............. (9)Z. knappii
(1) Zeravschania pauciradiata (Tamamsch.) Pimenov in Rechinger, Fl. Iran. 162: 459, t. 417.
1987 ≡Peucedanum pauciradiatum Tamamsch. in Trudy Bot. Inst. Akad. Nauk SSSR, Ser. 1 Fl.
Sist. Vyssh. Rast. 3: 225. 1936. – Holotype: Azerbaidzhan “prov. Nakhiczevan, c. Ordubad, M.
Schagal, 24.6.1920, Schelkovnikov &Kara-Murza” (ERE!).
=Peucedanum albostriatum Karjagin in Izv. Azerbaidzhansk. Fil. Akad. Nauk SSSR 5: 39.
1940. – Holotype: Armenia “distr. Megri, pr. custodiam Karschevan, ad fl. Arax, in schistosis
lapidosis siccis, 22.6.1934, Karjagin” (BAK).
=Peucedanum oligactis Rech. f. & Riedl in Anz. Oesterr. Akad. Wiss., Math-Naturwiss. Kl. 98:
251. 1961 ≡Johreniopsis oligactis (Rech. f. & Riedl) Pimenov in Rechinger, Fl. Iran. 162: 456, t.
413. 1987. – Holotype: Iran “prov. Kermanshah, inter Kermanshah & Sanandaj 94 km NNW
Taq-i Bustan, substr. Tonschiefer, 28.8.1957, Rechinger 14692”(W!)
494 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Fruit structure.–SeeFig.10A.
Distribution. – Iran (W: E Azarbayjan, Kordestan, Kermanshah), Azerbaidzhan, Armenia.
Ref. – Pimenov (1988: 79), Heller & Heyn (1993: 43). – Under Peucedanum pauciradiatum:
Shishkin (1951: 200), Karjagin (1955: 489), Tamamschjan (1967: 115, fig. 125), Mandenova
(1973: 405, t. 183). – Under Johreniopsis oligactis: Heller & Heyn (1993: 43), Jalili & Jamzad
(1999: 678).
(2) Zeravschania membranacea (Boiss.) Pimenov in Rechinger, Fl. Iran. 162: 458, t. 415. 1987
≡Peucedanum membranaceum Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 315. 1844. – Holotype:
Iran “prope Elamout [Persia, Alamut], Aucher-Eloy 4566” (P!; isotypes: G-BOIS!, K!, LE!, W!,
US).
=Cachrys leptorhabdos Bornm. in Repert. Spec. Nov. Regni Veg. 51: 103. 1942. – Holotype:
Iran “Dschekab inter Sultanabad & Kaschan, 7.1903, Strauss” (B!).
Fruit structure.–SeeFig.10B.
Distribution. – Iran (W: W Azarbayjan, E: Azarbayjan, Zanjan, Hamadan; N: Mazandaran; Cen-
tral: Arak, Esfahan; S: Fars).
Ref. – Pimenov (1988: 79), Heller & Heyn (1993: 43), Jalili & Jamzad (1999: 696). – Under Peuce-
danum membranaceum: Mozaffarian (1983: 124). – Under Cachrys leptorhabdos: Leute (1987:
211, in nota).
(3) Zeravschania regeliana Korovin in Bot. Mater. Gerb. Inst. Bot. Zool. Akad. Nauk Uzbeksk.
SSR 12: 28. 1948. – Holotype: Tadzhikistan “Vall. fl. Zeravschan, inter pag. Kschtut & [lacum]
Kuli-Kalon, 1882, Regel” ( LE!).
Fruit structure. – See Pimenov (1988)
Distribution. – Tadzhikistan, Uzbekistan.
Ref. – Shishkin (1950: 412), Korovin (1959: 351), Pimenov (1983: 273; 1988: 78), Korovin & al.
(1984: 152, t. 10, fig.3-5).
(4) Zeravschania scabrifolia Pimenov in Vvedensky, Consp. Fl. As. Med. 7: 375, 274. 1983. –
Holotype: Tadzhikistan “jugum Petri Primi, pars occidentalis, declivum septentrionale prope
trajectum Kamchirak, 2700 m, 18.8.1975, Pimenov 1127” (MW!; isotype: LE!).
Distribution. – Tadzhikistan.
Ref. – Korovin & al. (1984: 152), Pimenov (1988: 78).
(5) Zeravschania stricticaulis (Rech. f.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum stric-
ticaule Rech. f. in Repert. Spec. Nov. Regni Veg. 50: 258. 1941 ≡Johreniopsis stricticaulis
(Rech. f.) Pimenov in Rechinger, Fl. Iran. 162: 456, t. 414. 1987. – Holotype: Iran “Prov. Kho-
rassan, Montes Kopet-Dagh inter Quchan & Loftabad, in jugo Allah Akbar, 1800 m, 15.7.1937,
Rechinger 1749” (W!; isotypes: BM!, US).
Fruit structure.–SeeFig.10C.
Distribution. – Iran (E: Khorassan).
Ref. – Heller & Heyn (1993: 43), Jalili & Jamzad (1999: 678).
Note. – Akhani (in shed. herb. W) came to the same conclusion about the taxonomic attribution of
Peucedanum stricticaule to Zeravschania, instead of Johreniopsis.
Willdenowia 37 – 2007 495
496 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Fig. 10. Schematic transects of mericarps – A: Zeravschania pauciradiata; origin: Iran, Arak, Toureh Besri,
Kuh-e Aladagh, 25.8.1985, Mozaffarian 64194 (TARI); B: Z. membranacea; origin: Iran, E Azerbaijan,
c. 34 km S Mianeh, 1 km after Kahriz village, 1293 m, 37°13'N, 47°43'E, 2.9.2000, Akhani &Salimian 14214
(herb. Akhani); C: Z. stricticaulis; origin: Iran, Khorassan, Hazar Masdjid, inter Ardak et Tolgor, 1200-
1600 m, 7.-10.6.1948, Rechinger 4986 b (E); D: Z. ferulifolia; origin: Afghanistan, Kabul: Sher Darwasa,
1900 m, 19.6.1962, Hedge &Wendelbo 4312 (E); E: Z. aucheri; origin: Iran, Esfahan, Semirom Koh-e,
9.8.1978, Assadi &Mozaffarian 31662 (TARI). – Scale bars = 1 mm; for abbreviations see caption of Fig. 5.
(6) Zeravschania ferulifolia (Gilli) Pimenov in Rechinger, Fl. Iran. 162: 460, t. 418, 419. 1987 ≡
Peucedanum ferulifolium Gilli in Repert. Spec. Nov. Regni Veg. 61, 3: 204. 1959. – Holotype:
Afghanistan “bei Kabul, nach N exponierte Schlucht am Nordhang eines Berges am rechten
Kabulusur in den Tangi Ghari unweit des Wasserfalls Maipar, 1070 m, 22.6.1951, Gilli 2037”
(W!).
=Scaligeria paniculata Nasir in Biologia (Lahore) 9: 39. 1963. – Holotype: Pakistan “Baluchis-
tan, Koshkai, Dick Peddie 22”.
=Ferula kandaharica Rech. f., Fl. Iran. 162: 424, t. 383. 1987. – Holotype: Afghanistan “Kan-
dahar, in monte 15 km SW Kandahar, 1100 m, 22.5.1967, Freitag 811” (W; isotype: E!).
Fruit structure.–SeeFig.10D.
Distribution. – Afghanistan, Iran (E: Khorassan), Pakistan.
Ref. – Pimenov (1988: 78, p.p.). – Under Peucedanum ferulifolium: Nasir (1972: 137, fig. 41A-C).
– Under Scaligeria paniculata: Rechinger (1987: 233, in nota).
(7) Zeravschania minjanensis (Rech. f.) Rech. f., Fl. Iran. 162: 461, t. 420. 1987 ≡Eleuthe-
rospermum minjanense Rech. f. in Kongel. Danske Vidensk. Selsk. Skr. 13(4) [Symb. Afghan.
5]: 88. 1963. – Holotype: Afghanistan “Minjan, in jugo Minjan, 3600 m, 26.7.1937, Koelz
12718”(US).
Distribution. – Afghanistan.
(8) Zeravschania aucheri (Boiss.) Pimenov in Rechinger, Fl. Iran. 162: 459, t. 416. 1987 ≡Peu-
cedanum aucheri Boiss. in Ann. Sci. Nat., Bot., ser. 3, 1: 315. 1844. – Lectotype (designated by
Pimenov 1987): Iran “In Persiae borealis monte Dalmkou, Aucher-Eloy 4630” (G-BOIS!; isolec-
totypes: K!, W!).
=Peucedanum kotschyi Boiss., Diagn. Pl. Orient., ser. 1, 6: 63. 1846. – Holotype: Iran “Persia
australis, Kotschy 888” (G-BOIS!).
Fruit structure.–SeeFig.10E.
Distribution. – Iran (W: W Azarbayjan, E Azarbayjan, Hamadan, Lorestan; N: Mazandaran;
Central: Semnan, Tehran, Arak, Ghom, Esfahan, Chaharmahal va Bakhteyari; S: Boyerahmad va
Kohgiluye, Hormozgan).
Ref. – Pimenov (1988: 79), Heller & Heyn (1993: 43), Jalili & Jamzad (1999: 695). – Under
Peucedanum aucheri: Boissier (1872: 1022), Burkill (1909: 36), Nasir (1972: 139), Mozaffarian
(1983: 123, t. 319).
(9) Zeravschania knappii (Bornm.) Pimenov & Kljuykov, comb. nov. ≡Peucedanum knappii
Bornm. in Verh. Zool.-Bot. Ges. Wien 60: 121. 1910. – Holotype: Iran “Azerbeidjan, Qotursu
[Kotursu], distr. Afshar, Takht-i Soleiman, in pascuis montanis, 18.8.1884, Knapp” (B!).
=Peucedanum chenur Mozaff. in Bot. Zhurn. (St. Petersburg) 88(4): 116, fig. 5. 2003. –
Holotype: Iran “Kurdistan: Sanandaj, kuhhalye Chehelcheshme, Saral region, 2000-2350 m,
Mozaffarian 74795” (TARI!).
Fruit structure. – See Mozaffarian (2003, sub Peucedanum chenur).
Distribution. – Iran (W: W Azarbayjan, Kordestan).
Ref. – Under Peucedanum knappii: Rechinger (1987: 444), Heller & Heyn (1993: 41), Jalili &
Jamzad (1999: 684).
Species excludendae
Johrenia longifolia (Fisch. ex Spreng.) Calest. ≡Eriosynaphe longifolia (Fisch. ex Spreng.) DC.
Johrenia lycaonica Bornm. = Onopordon hispidum (Friv.) Griseb.
Willdenowia 37 – 2007 497
Johrenia nudiuscula (Turcz.) Palib. = Ferulopsis hystrix (Bunge ex Ledeb.) Pimenov
Johrenia pichleri Boiss. = Xanthoselinum alsaticum (L.) Schur
Johrenia platypoda Aitch. & Hemsl. ≡Galagania platypoda (Aitch. & Hemsl.) M. G. Vassilieva
& Pimenov
Johrenia popovii (Korovin) Korovin = Ferula nuratavica Pimenov
Johrenia racemoso-umbellata Gilli ≡Ferula racemoso-umbellata (Gilli) Rech. f.
Johrenia seseloides (Hoffm.) Koso-Pol. ≡Ledebouriella seseloides (Hoffm.) H. Wolff
Johrenia sieversii Koso-Pol. = Ledebouriella seseloides (Hoffm.) H. Wolff
Johrenia tordylium Fenzl ≡Coriandrum tordylium (Fenzl) Post
Johrenia villosa (Turcz. ex Fisch. & C. A. Mey.) Kudo ≡Phlojodicarpus villosus (Turcz. ex
Fisch. & C. A. Mey.) Ledeb.
Johrenia westii Post, Pl. Post. 3: 9. 1892 ≡Ferulago westii (Post) Pimenov & Kljuykov, comb.
nov. – Lectotype (designated here): Syria “In planitiae ad radices montis Halimat-Jabu
(Antilibani) in Wadi Kenniyyeh-er-Ras [plain at head of Wadi Kenuyyat-er-Vas], 27.7.1891,
Post 121” (BEI!; isolectotypes: B!, G!, K!)
Zeravschania pastinacifolia (Boiss. & Hausskn. ex Boiss.) Salimian & Akhani ≡Demavendia
pastinacifolia (Boiss. & Hausskn. ex Boiss.) Pimenov
Acknowledgements
This investigation was supported by grants from the Russian Foundation for Fundamental Inves-
tigations (RFFI). The curators of following herbaria are gratefully acknowledged for the loan of
specimens or scanning pictures and study facilities to examine collections: ANK, B, BEI, E, G,
K, LE, MW, P, TARI, W, WU. We express our gratitude to the referees Dr Gerald Parolly and Mr
Jean-Pierre Reduron for their constructive comments on a previous version of the paper.
References
Al-Eisawi, D. M. 1982: List of Jordan vascular plants. – Mitt. Bot. Staatssamml. München 18:
79-182.
Beck-Mannagetta, G. 1927: Flora Bosne, Herzegovine i oblasti Novogo Pasara. – Beograd &
Sarajevo.
Besser, W. 1822: Enumeratio plantarum hucus in Volhynia, Podolia, gub. Kiioviensi, Bessarabia
cis-tyraica et circa Odessam collectarum, simul cum observationibus in primitias florae Gali-
ciae austriacae. – Wilna.
Boissier, E. 1844: Plantae aucherianae II. – Ann. Sci. Nat., Bot., ser. 3, 1: 297-349
— 1856: Diagnoses plantarum orientalium, ser. 2. – Lipsiae.
— 1872: Flora orientalis 2. – Genève & Basel.
— (ed. Buser, R.) 1888: Flora orientalis supplementum. – Genève, etc.
Bolòs, O. de & Vigo, J. 1990: Flora dels Països Catalans 2. – Barcelona.
Bornmüller, J. 1911: Iter Persico-Turcicum 1892-93. Beiträge zur Flora von Persien, Babylo-
nien, Assyrien und Arabien. – Beih. Bot. Centralbl., Abt. 2, 28: 89-171.
— 1930: Kritische Bemerkungen über einige orientalische Arten der Gattung Johrenia nebst
Beschreibung zweier neuer Typen. – Repert. Spec. Nov. Regni Veg. 28(8-10): 33-53.
— 1938: Iter persico-turcicum 1892-93. Beiträge zur Flora von Persien, Babylonien, Assyrien,
Arabien (Fortsetzung II). – Beih. Bot. Centralbl, Abt. B, 58: 252-302.
Boulos, L. 1977: Studies on the flora of Jordan 5. On the flora of El Jafr-Bayir Desert. – Can-
dollea 32: 99-110.
Bruck, M. 1988: Untersuchungen über die Morphologie und Anatomie von Peucedanum carvi-
folia Vill. (Apiaceae). – Bull. Soc. Naturalistes Luxemburg 88: 69-80.
Burkill, I. H. 1909: A working list of the flowering plants of Baloutchestan. – Calcutta.
Burnat, E. 1906: Flore des Alpes Maritimes 4. – Genève, etc.
498 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Calestani, V. 1905: Contributo alla sistematica delle Ombrellifere d’Europa. – Webbia 1: 89-280.
Candolle, A. P. de 1829: Collection de mémoires 5. Mémoire sur la famille d’Ombellifères. –
Paris & Strasbourg.
Carbonnier, J., Fatianoff, O. & Molho, D. 1978: Phytochemie comparée des taxon rattachés à la
tribu des Peucedaneae (Umbelliferae-Apioideae). – Pp. 387-513 in: Cauwet-Marc, A.-M. &
Carbonnier, J. (ed.), Les Ombellifères, contributions pluridisciplinaires à la systématique.
Actes du 2e Symposium International sur les Ombellifères, Centre Universitaire de Per-
pignan, 18-21 mai 1977. – Perpignan.
Chamberlain, D. F. 1972: Peucedanum L. – Pp. 473-481 in: Davis, P. H. (ed.), Flora of Turkey
and the East Aegean Islands 4. – Edinburgh.
Chrtek, J. & Hendrich, R. 1962: Zu einigen Fragen der Art Peucedanum carvifolia. – Acta Univ.
Carol. Biol. 1962: 137-151.
Degen, A. von 1937: Flora velebitica 2. – Budapest.
Demiri, M. 1983: Flora ekskursioniste e Shoqiperise. – Tiranë.
Dostal, J. 1950: Kvetena CSR. – Praha.
Downie, S. P., Watson, M. F., Spalik, K. & Katz-Downie, D. S. 2000: Molecular systematics of
Old World Apioideae (Apiaceae): relationships among some members of tribe Peucedaneae
sensu lato, the placement of several island-endemic species, and resolution with the apioid
superclade. – Canad. J. Bot. 78: 506-528. [CrossRef]
Drude, O. 1897-98: Umbelliferae. – Pp. 63-272 in: Engler, A. & Prantl, K. (ed.), Die natürlichen
Pflanzenfamilien 3(8). –Leipzig.
Fabri, R. 1993: Flore générale de Belgique. Spermatophytes 5(2). –Meise.
Fenzl, E. 1842: Pugillus plantarum novarum Syriae et Tauri occidentalis primus. – Wien.
— 1843: Abbildungen und Beschreibungen neuer und selthener Thiere und Pflanzen in Syria
und im westlichen Taurus gesammelt von T. Kotschy. – Stuttgart.
Fournier, P. 1961: Les quatre flores de France. – Paris.
Fragman, O., Plitmann, U., Heller, D. & Shmida, A. 1999: Checklist and ecological data-base of
the flora of Israel and its surroundings. – Jerusalem.
Frey, R. 1989: Taxonomische Revision der Gattung Peucedanum: Sektion Peucedanum &Sek-
tion Palimbioidea (Umbelliferae). – Candollea 44: 257-327.
Gaudin, J. F. A. P. 1828: Flora helvetica sive historia stirpium hucusque cognitarum in Helvetia
2. – Zürich.
Grossheim, A. A. 1927: Nekotorye novye dlja Kavkaza vidy tzvetkovych rastenyi [Some spe-
cies of flowering plants, new for Caucasus]. – Vestn. Tiflissk. Bot. Sada, ser. 2, 3-4: 15-25.
— 1932: Flora Kavkaza 3. – Tiflis.
Gruenberg-Fertig, I. & Zohary, M. 1970: Nomenclature remarks on some plants of Palestina. –
Israel J. Bot 19: 293-304.
Grulich, V. 1997: Peucedanum L. – Pp. 406-418 in: Slavik, B. (ed.), Kvetena Ceské Republiky
5. – Prague.
Guillén, A. & Lainz, M. 2003: Peucedanum L. – Pp. 346-361 in: Castroviejo, S. (ed. gen.), Flora
iberica 10. –Madrid.
Guinochet, M. & Vilmorin, R. de 1975: Flore de France 2. –Paris.
Hadacek, F. 1989: Vergleichende phytochemische Untersuchungen in der Gattung Peucedanum
(Apiaceae-Apioideae).–Stapfia18: 1-186.
Halacsy, E. 1901: Conspectus florae graecae 1(3). –Leipzig.
Handel-Mazzetti, H. F. 1913: Pteridophyta und Anthophyta aus Mesopotamien und Kurdistan
sowie Syrien und Prinkipo. – Ann. K. K. Naturhist. Hofmus. Wien 27: 41-92.
Hartvig, P. 1986: Umbelliferae. – Pp. 655-735 in: Strid, A. & Tan, K. (ed.), Mountain flora of
Greece 1. – Cambridge, etc.
Hayek, A. 1927: Prodromus Florae peninsulae balcanicae 1(7/8). – Repert. Spec. Nov. Regni
Veg. Beih. 30(1). – Berlin.
Heller, D. & Heyn, C. C. 1993: Conspectus florae orientalis. An annotated catalogue of the flora
of the Middle East 7. – Jerusalem.
Willdenowia 37 – 2007 499
Hiroe, M. 1958: Umbelliferae of Asia (excluding Japan) 1. – Kyoto.
— 1979: Umbelliferae of the world. – Tokyo.
Hoffmann, G. F. 1816: Plantarum Umbelliferarum genera, ed. 2. – Mosquae.
Holmgren, P. K. & Holmgren, N. H. 1998- (continuously updated): Index herbariorum. – Pub-
lished on the internet http://sciweb.nybg.org/science2/IndexHerbariorum.asp.
Holub, J. 1972: Peucedanum carvifolia. – Pp. 40-41 in: Sojak J. (ed.), Plantae cechoslovacae
exsicatae, cent. 3. – Sborn. Nar. Mus. Praze, Rada B, Priv. Vedy 27.
Hy, l’Abbe 1901: Sur le Peucedanum schottii Besser. – Bull. Soc. Bot. France 48: 105-107.
Ivanov, A. 2001: Konspect flory Stavropolija [Conspectus of Stavropol province flora], ed. 2. –
Stavropol.
Jalili, A. & Jamzad, Z. 1999: Red Data Book of Iran. A preliminary survey of endemic, rare &
endangered plant species in Iran. – Tehran.
Janchen, E. 1956: Catalogus florae Austriae 1. –Wien.
Karjagin, I. I. 1955: Flora azerbaidzhana 6. – Baku.
Kljuykov, E. V., Liu, M., Ostroumova, T. A., Pimenov, M. G., Tilney, P. M. & Wyk, B.-E. van
2004: Towards a standardised terminology for taxonomically important morphological char-
acters in the Umbelliferae. – South African J. Bot. 70: 488-496.
Korovin, E. P. 1959: Umbelliferae. – Pp. 257-470 in: Vvedensky, A. I. (ed.), Flora uzbekistana
4. – Tashkent.
— , Pimenov, M. G. & Kinzikaeva, G. K. 1984: Umbelliferae. – Pp. 10-214 in: Ovczinnikov, P.
N. (ed.), Flora Tadzhikskoi SSR 7. – Moscow & Leningrad.
Kowal, T. & Wojterska, H. 1973: Morfologiczne i anatomiczne cechy diagnostyczne owoców
wybranych gatunkóv rodzaju Peucedanum L. – Prace Komis. Biol. 35(7): 457-491.
Krause, K. 1937: Ankaranin floru 2. – Ankara.
Kuzmanov, B. & Andreev, N. 1982: Peucedanum L. – Pp. 220-240 in: Jordanov, D. (ed.), Flora
na Narodna Republika Bblgarija 8. –Sofia.
Lange, J. 1880: Umbelliferae – Pp. 1-101 in: Willkomm, M. & Lange J., Prodromus florae
hispanicae 3. – Stuttgart.
Lauber, K. & Wagner, G. 1998: Flora helvetica, ed. 2. – Bern, etc.
Ledebour, C. F. 1844: Flora rossica 2. – Stuttgart.
Leute, G.-H. 1987: Alococarpum. – Pp. 210-211 in: Rechinger, K. H. (ed.), Flora iranica 162. –
Graz.
Lipsky, V. I. 1899: Flora Kavkaza. – Trudy Tiflissk. Bot. Sada 4.
Luca6, G. & Regula-Belivacqua, L. 1997: Apiaceae (Umbelliferae) – Pp. 104-116 in: Nicoli6,T.
(ed.), Index Florae croaticae 2. –Zagreb.
Mandenova, I. P. 1973: Umbelliferae. – Pp. 251-427 in: Takhatadjan, A. L. (ed.), Flora Armenii
6. – Erevan.
— 1984: Umbelliferae. – Pp. 128-326 in: Ketzchoveli, N. N. (ed.), Flora Georgiae 9. – Tbilisi.
Marchall von Bieberstein, F. 1819: Flora taurico-caucasica 3. – Charkov.
Menitsky, J. L. 1991: Konspect vidov semeistva Apiaceae (Umbelliferae) flory Kavkaza [Syn-
opsis of the species of the family Apiaceae (Umbelliferae) from the Caucasus]. – Bot. Zhurn.
(St. Petersburg) 76: 1749-1764.
Molero, J. & Rovira, A. M. 1985: Peucedanum schottii Besser ex DC. novedad para la flora
iberica. – Anales Jard. Bot. Madrid 42: 537.
Mouterde, P. 1970: Nouvelle flora du Liban et de la Syrie 2. – Beyrouth.
Mozaffarian, V. 1983: The family of Umbelliferae in Iran. Keys and distribution. – Tehran.
— 2003: New species and new records of Iranian Umbelliferae. – Bot. Zhurn. (St. Petersburg)
88(4): 104-124.
Nasir, E. 1972: Umbelliferae. – In: Nasir, E. & Ali, S. I. (ed.), Flora of West Pakistan 20. –
Rawalpindi.
Nikoli6, V. 1973: Apiaceae. – Pp. 183-349 in: Josifovi6,M.(ed.),FloraSRSerbije5. – Beograd.
500 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
Parolly, G. & Nordt, B. 2004: Peucedanum isauricum (Apiaceae), a striking new species from S
Anatolia with notes on the related P. graminifolium and P. spreitzenhoferi. – Willdenowia
34: 135-144. [CrossRef]
— & — 2005: A further new Peucedanum species (Apiaceae) from Taurus Mts, Turkey. –
Willdenowia 35: 97-105. [CrossRef]
Pignatti, S. 1982: Flora d’Italia. 2. – Bologna.
Pimenov, M. G. 1983: Umbelliferae. – Pp. 167-322 in: Vvedensky, A. I. (ed.), Conspectus flo-
rae Asiae mediae 7. – Tashkent.
— 1987: Leutea, Johreniopsis. – Pp. 445-450, 454-457 in: Rechinger, K. H. (ed.), Flora iranica
162. – Graz.
— 1988: Rod Zeravschania Korov.: vidy i taksonomicheskoe polozhenie v semeistve Umbelli-
ferae [The genus Zeravschania Korov.: the species and the taxonomic position in the family
Umbelliferae]. – Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 93(4): 75-80.
— 1992: Ormosolenia restored. – Edinburgh J. Bot. 49: 219-223.
— & Kljuykov, E. V. 1992: Johrenia platycarpa (Umbelliferae) – novyi vid dlja flory S.S.S.R.
[Johrenia platycarpa (Umbelliferae), a new species for the flora of the USSR]. – Bot. Zhurn.
(St. Petersburg) 77(7): 111-113.
— , Vasil’eva, M. G., Leonov, M. V. & Daushkevich, J. V. 2003: Karyotaxonomical analysis in the
Umbelliferae.–Enfield.
Post, G. E. 1896: Flora of Syria, Palestina and Sinai. – Beyrut.
— (rev. by J. E. Dinsmore) 1932: Flora of Syria, Palestina and Sinai, ed. 2, 1. –Beyrut.
Qosja, K. (ed.) 1992: Flora e Shqipërisë 2. Rosaceae - Umbelliferae. – Tiranë.
Quézel, P. & Santa, S. 1963: Nouvelle flora d’Algérie et des régions désertiques méridionales 2.
–Paris.
Rechinger, K. H. 1943: Flora Aegaea. Flora der Inseln und Halbinseln des Ägäischen Meeres. –
Wien.
— 1957: Zur Flora von Syrien, Lebanon und den angrenzenden türkischen Gebieten (Reliquiae
Samuelssonianae VI). – Ark. Bot. 5(1).
— (ed.) 1987: Flora iranica 162. – Graz.
Reduron, J.-P., Charpin, A. & Pimenov, M. 1997: Contribution à la nomenclature générique des
Apiaceae (Ombellifères). – J. Bot. Soc. Bot. France 1: 91-104.
Rohlena, J. 1942: Conspectus florae montenegrinae. – Preslia 20-21.
Rohlf, F. J. (ed.): NTSYS-pc. Numerical taxonomy and multivariate analysis system. Version
1.8. – New York.
Rompaey, E. van & Delvasalle, L. 1972: Atlas de la flore belge et luxembourgeoise: pteridophytes
et spermatophytes. – Bruxelles.
Shishkin, B. K. 1951: Peucedanum. – Pp. 168-203 in: Shishkin, B. K. & Bobrov, E. G. (ed.),
Flora SSSR 17. – Leningrad & Moskva.
Schur, F. 1866: Umbelliferae. – Pp. 244-276 in: Enumeratio plantarum Transsilvaniae exhibens.
– Vindobonnae.
Snogerup, S., Snogerup, B., Phitos, D. & Kamari, G. 2001: The flora of Chios island (Greece). –
Bot. Chron. (Patras) 14: 5-197.
Soó, R. 1966: A magyar flora és vegétacio Rendszertani-növényföldrajzi kézikönyve 2. – Budapest.
Spalik, K., Reduron, J.-P. & Downie, S. R. 2004: The phylogenetic position of Peucedanum sensu
lato and allied genera and their placement in tribe Selineae (Apiaceae, subfamily Apioideae).
– Pl. Syst. Evol. 243: 189-210. [CrossRef]
Sprengel, C. 1818: Species Umbelliferarum minus cognitae. – Halae.
Stojanov, N. A. & Stefanov, B. 1948: Flora na Bulgar’i’a, ed. 3. – Sofia.
Strid, A. & Papanicolaou, K. 1981: Floristic notes from the mountains of Northern Greece. Ma-
terials for the mountain flora of Greece 7. – Nordic J. Bot. 1: 66-82.
Takhtajan, A. L. & Fedorov, A. A. 1972: Flora erevana. – Leningrad.
Tamamschjan, S. G. 1951: De generibus Phlojodicarpus Turcz. et Stenocoelium Ldb. notae criti-
cae. – Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 14: 265-276.
Willdenowia 37 – 2007 501
— 1967: Umbelliferae. – Pp. 5-137 in: Grossheim, A. A., Flora Kavkaza, ed. 2, 7. – Leningrad.
Tchihachev, P. A. 1860: Asie Mineure. Description physique, statistique et archéologique de cette
contrée. Troisième partie. Botanique. – Paris.
Thellung, A. 1925-26: Umbelliferae. – Pp. 926-1537 in: Hegi G. (ed.), Illustrierte Flora von Mittel-
Europa 5(2). – München.
Thiébaut, J. 1936: Flora libano-syrienne 1. – Mém. Inst. Égypte 31.
Todor, I. 1958: Umbelliferae. – Pp. 326-652 in: Sbvulescu, T. (ed.), Flora Republicii Populare
Romîne 6. – Bukurestu.
Townsend, C. C. 1964: Notes on the Umbelliferae of Iraq II. – Kew Bull. 19: 69-75.
Tutin, T. G. 1968: Umbelliferae. – Pp. 315-375 in: Tutin, T. G., Heywood, V. H., Burges, N. A.,
Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. (ed.), Flora europaea 2. –
Cambridge, etc.
Visiani, R. 1850: Flora dalmatica sive enumeratio stirpium vascularium quas haetenus in Dal-
matia lectas et sibi observatas descriptis digessit rariorumque iconibus 3(1). – Lipsiae.
Vural, M. & Adigüzel, N. 1996: Notes on Turkish flora 1. Peucedanum graminifolium and Olym-
posciadium caespitosum (Apiaceae/Umbelliferae).–Ot3(1): 59-64.
Valiejo-Roman, K. M., Terentieva, E. I., Samigullin, T. H., Pimenov, M. G., Ghahremani-Nejad,
F. & Mozaffarian, V. 2006: Molecular data (nrITS-sequencing) reveal relationships among
Iranian endemic taxa of the Umbelliferae. – Feddes Repert. 117: 367-388. [CrossRef]
Wisskirchen, R. & Haeupler, H. 1998: Standardliste der Farn- und Blütenpflanzen Deutschlands.
– Stuttgart.
Zohary, M. 1972: Umbelliferae. – Pp. 378-453 in: Feinbrun-Dothan, N. (ed.), Flora Palaestina 2.
– Jerusalem.
Address of the authors:
Michael G. Pimenov, Eugene V. Kljuykov & Tatiana A. Ostroumova, Botanical Garden, Moscow
State University, Moscow 119992, Russia; e-mail: mgpimenov@mail.ru
502 Pimenov & al.: Dichoropetalum, Johrenia, Zeravschania and related genera
