A new treatment of Pilosella for the Euro-Mediterranean flora

Abstract

Bräutigam, S. & Greuter, W.: A new treatment of Pilosella for the Euro-Mediterranean flora [Notulae ad floram euro-mediterraneam pertinentes 24]. — Willdenowia 37:123–137 — ISSN 0511-9618; © 2007 BGBM Berlin-Dahlem. doi:10.3372/wi.37.37106 (available via http://dx.doi.org/) Recognising Pilosella as a genus distinct from Hieracium is justified both from a phylogenetic point of view (a more broadly defined Hieracium, to be monophyletic, would have to include at least two further genera along with Pilosella: Andryala and Hispidella) and for practical considerations. In Hieracium, almost all taxa are apomicts that rarely hybridise, and whenever they do, give rise to new, stable apomictic lines that are customarily given taxonomic recognition as species or subspecies. In Pilosella hybridisation is frequent, gene flow between populations (however defined) is considerable, and the recognition of microtaxa as if they were apomictic lines is unpractical. The classification here proposed rests on a framework of twenty accepted “basic” species (some with subspecies) or species aggregates. Hybrid progenies in which 2-3(-4) of these species or aggregates are believed to have participated are treated as 122 “collective species”, one per known or postulated parental combination. Each of these comprises one recognised species, or sometimes more than one when an included morphotype is stable over a significant, coherent area, or when the offspring of a particular subspecies or microspecies combination deserves recognition. A synopsis of the proposed classification is presented, and required new names and combinations are validated.

Full text

Notulae ad floram euro-mediterraneam pertinentes No. 24
SIEGFRIED BRÄUTIGAM & WERNER GREUTER
A new treatment of Pilosella for the Euro-Mediterranean flora
Abstract
Bräutigam, S. & Greuter, W.: A new treatment of Pilosella for the Euro-Mediterranean flora [Notulae
ad floram euro-mediterraneam pertinentes 24]. – Willdenowia 37: 123-137 – ISSN 0511-9618; © 2007
BGBM Berlin-Dahlem.
doi:10.3372/wi.37.37106 (available via http://dx.doi.org/)
Recognising Pilosella as a genus distinct from Hieracium is justified both from a phylogenetic point of
view (a more broadly defined Hieracium, to be monophyletic, would have to include at least two fur-
ther genera along with Pilosella: Andryala and Hispidella) and for practical considerations.In
Hieracium, almost all taxa are apomicts that rarely hybridise, and whenever they do, give rise to new,
stable apomictic lines that are customarily given taxonomic recognition as species or subspecies. In
Pilosella hybridisation is frequent, gene flow between populations(however defined) is considerable,
and the recognition of microtaxa as if they were apomictic lines is unpractical. The classification here
proposed rests on a framework of twenty accepted “basic” species (some with subspecies) or species
aggregates. Hybrid progenies in which 2-3(-4) of these species or aggregates are believed to have par-
ticipated are treated as 122 “collective species”, one per known or postulated parental combination.
Each of these comprises one recognised species, or sometimes more than one when an included
morphotype is stable over a significant, coherent area, or when the offspring of a particular subspecies
or microspecies combination deserves recognition. A synopsis of the proposed classification is pre-
sented, and required new names and combinations are validated.
Keywords: Hieracium, Europe, Mediterranean, classification, nomenclature, collective species.
Introduction
Up to the present day, Pilosella Vaill. is considered by many authors as a subgenus of Hiera-
cium L. Schultz & Schultz (1862) are among the first to accept it as a genus of its own (as did
19th century Hieracium specialists Fries, Arvet-Touvet and Norrlin at some stage of their life –
but not permanently). During the last decades generic recognition of Pilosella has gained in-
creased but not universal support. Several authors (including the present ones) have so far hesi-
tated to follow suit, the unassessed nomenclatural consequences of a change being a major
reason for their reluctance. Also, the question of how to deal with the American Hieracium subg.
Chionoracium Sch. Bip. (or as a genus, Stenotheca Monnier) under a narrow generic concept had
not yet been definitely answered.
Willdenowia 37 – 2007 123

On morphological grounds Pilosella is clearly distinct by cypsela features from Hieracium
sensu stricto. A number of further characteristics, when combined, support separation but are not
expressed in all species (e.g., the presence of stolons or of red veins on the abaxial ligule sur-
face). Past workers have experienced no difficulty in defining both groups and sharing the spe-
cies among them. In spite of abundant hybridisation within either group, no hybrids between
representatives of Pilosella and Hieracium are known. Nuclear DNA content (1C value)is about
twice as high in Hieracium as in Pilosella (Bräutigam & Bräutigam 1999, Vladimirov & Greil-
huber 2003).
The sequence analysis of both nuclear (ITS) and chloroplast DNA (trnT-trnL intergenic spacer;
matK gene) added considerably to our understanding of phylogenetic relationships (Fehrer & al.
2007a). The results consistently support the monophyly of a clade comprising Hieracium sensu
lato, Andryala L. and Hispidella Lam. The chloroplast DNA data, especially within Pilosella,
show no evident correlation with morphology. The observed patterns point to basal and/or inter-
group hybridisation events resulting in chloroplast “capture” followed by differential line sorting.
Chloroplast DNA data are thus indicative of the complex evolutionary mechanisms and reticulate
connections prevailing in the group since its very origin, rather than of morphologically expressed
phylogenetic affinities. The nuclear (ITS) data, however, yield a neat classification well compati-
ble with morphological data. Apart from the single, isolated species Hieracium intybaceum Jacq.,
which appears in a basal position with respect to the whole complex, it shows a subdivision into
three main clades: Hieracium (incl. subg. Chionoracium), Pilosella with its sister Hispidella, and
Andryala.
Phylogenetically it would be possible to define a natural genus Hieracium in a broad sense,
to include Pilosella, Andryala, and Hispidella. However, from a practical point of view a nar-
rower genus concept, with those four as distinct genera, is much preferable.
While apomixis is frequent or predominant in Old World Hieracium and Pilosella alike, it is
of a different nature: diplospory in the former, apospory in the latter.Concomitantly the patterns
of diversification differ. Whereas Hieracium, except for some sexual diploids, consists of
phenetically invariable apomictic lines, most Pilosella populations retain a varying but signifi-
cant degree of sexuality. The study of natural populations, the observed patterns of variation and
experimental crosses concur to demonstrate that hybridisation, involving all species, is a fre-
quent, recurrent phenomenon in Pilosella. There are all kinds of taxa in the latter genus, ranging
from fully sexual, both diploid and polyploid species, to almost totally apomictic ones occurring
as widespread, morphologically stable clones (Fehrer & al. 2007b). As a consequence, classifica-
tion in Pilosella must follow different criteria than those currently applied in Hieracium, where
the trend is toward recognition of very narrowly defined but constant, hybridogenous
agamospecies.
Classificationin[Hieracium subg.] Pilosella underwent extreme fluctuation over time. In his
seminal revision Fries (1862) recognised 42 species in all, to which Schultz & Schultz (1862)
added a dozen of their own plus an equal number of interspecific hybrids. When Nägeli & Peter
(1885) introduced their new taxonomic concept to hieraciology it resulted in a fourfold number
of species (164) plus countless subspecies. The next monographer, Zahn (1923), recognised 182
species – a relatively slight increase – but (to accommodate the countless microtaxa that had
meanwhile been described and named, especially by Nordic botanists) inflated subspecies num-
bers. As an example, in that work there are 624 numbered subspecies in Hieracium pilosella L.
(Pilosella officinarum) alone. Yet Zahn himself was critical of the meaning of such extreme pul-
verisation: Touton (1921: 71) quotes him as follows [our translation]: “The study of the species
pilosella in detail leads to nothing, because not a single fixed subspecies exists, but everything
flows together”; to which Touton added: “The species pilosella shares this fate with many other
Hieracium species”.
At the opposite extreme, a synthetic approach has been proposed recently by Tyler (2001).
Dealing only with the Nordic taxa, he reduced their species number to 8, of which only 3 were
further subdivided, plus 12 biparental nothospecies. He justified his new, “unprejudiced” classi-
124 Bräutigam & Greuter: New treatment of Pilosella for the Euro-Mediterranean flora

fication by stating “that all existing systems are inappropriate in the light of the complicated, and
largely unknown, reproductive biology of these taxa”. While his approach has undoubted merit,
it is not useful in the context of our Euro-Mediterranean inventory, for at least two obvious rea-
sons. First, extending it southward to cover other areas has yet to be attempted and would pre-
dictably result in conflicts of classification, as what is well defined and stable in one area need
not be so elsewhere; and second, adopting broadly defined units for organising and storing infor-
mation inevitably results in loss of valuable chorological and morphological data.
The system we have chosen for our broad, Euro-Mediterranean context is neither novel nor
unique. It builds upon good, modern precedent that has been successfully applied on a more re-
stricted geographical scale (see, in particular, Šljakov 1989 for European Russia and Mateo Sanz
2006 for Spain). In the first place, we recognise a relatively small number (20) of “basic” species,
or in some cases, small groups of species (termed “aggregates”). This part of our classification
follows the usual standards adopted for the Euro+Med database, as explained and implemented in
Med-Checklist (Greuter & al. 1984-89), meaning that “basic” species and their segregates may be
further subdivided into subspecies, or may include insufficiently known taxa of doubtful taxo-
nomic value (introduced by the word “also”, in the following Synopsis). We have, however, made
spare use of subspecies, defining them broadly – often in the sense of the greges of Zahn (1923).
Plants that do not fit into one of the basic species or aggregates are interpreted as hybrid off-
spring, and binomials are used to designate each postulated “parental” combination. These
hybridogenous taxa (or taxon swarms) we do not treat as nothospecies (as which they would not
be covered by either Euro+Med or Med-Checklist, but simply ignored) but term them “collective
species” (as which they can be included). Thus, to each different combination of postulated pa-
rental basic species (or aggregates) corresponds one collective species, provided that (a) it oc-
curs in nature and (b) it can be distinguished with some confidence from other possible
combinations. (The latter is not always the case: for example, crosses of Pilosella officinarum of-
ten cannot be distinguished from the corresponding crosses of P. peleteriana.) Our 122 collec-
tive species are of very unequal nature and value: they may comprise newly formed, primary
hybrids only, or correspond to stable hybridogenous species, or often they include both (Schuh-
werk & Fischer 2003, Fehrer & al. 2005). As a rule, each collective species consists of a single
taxonomic unit treated at species rank and designated by a binomial. However, on occasions we
found it practical to recognise more than one specific unit within a collective species. Such seg-
regates are useful when a particular morphotype is stable over a significant, coherent area, or
when the offspring of different subspecies or segregates of the same basic species differ recog-
nisably among themselves. The number of such segregates we kept deliberately low; more can be
added easily if and when the need is felt. In no case did we recognise subspecies within collec-
tive species.
Synopsis of Euro-Mediterranean Pilosella taxa
A. Basic species and aggregates (with subordinate species and subspecies)
1. Pilosella alpicola (Steud. & Hochst.) F. W. Schultz & Sch. Bip.; also:
Pilosella bonaquae (Buttler & W. Lippert) S. Bräut. & Greuter;
2. Pilosella argyrocoma (Fr.)F.W.Schultz&Sch.Bip.;
3. Pilosella aurantiaca (L.) F. W. Schultz & Sch. Bip.:
subsp. aurantiaca,
subsp. auropurpurea (Nägeli & Peter) S. Bräut. & Greuter,
subsp. decolorans (Fr.) T. Tyler;
4. Pilosella breviscapa (DC.) Soják;
5. Pilosella caespitosa aggr.:
Pilosella caespitosa (Dumort.) P. D. Sell & C. West; also:
subsp. colliniformis (Peter) P. D. Sell & C. West,
Pilosella onegensis Norrl.;
Willdenowia 37 – 2007 125

6. Pilosella castellana (Boiss. & Reut.) F. W. Schultz & Sch. Bip.;
7. Pilosella cymosa (L.) F. W. Schultz & Sch. Bip.:
subsp. cymosa,
subsp. sabina (Sebast. & Mauri) H. P. Fuchs,
subsp. vaillantii (Tausch) S. Bräut. & Greuter;
8. Pilosella echioides aggr.:
Pilosella brzovecensis (Horvat & Paw5³.) S. Bräut. & Greuter,
Pilosella caucasica (Nägeli & Peter) Sennikov,
Pilosella echioides (Lumn.) F. W. Schultz & Sch. Bip.:
subsp. echioides,
subsp. proceriformis (Nägeli & Peter) S. Bräut. & Greuter,
Pilosella procera (Fr.)F.W.Schultz&Sch.Bip.;
9. Pilosella galiciana (Pau) M. Laínz;
10. Pilosella glacialis (Reyn.)F.W.Schultz&Sch.Bip.;
11. Pilosella hoppeana (Schult.) F. W. Schultz & Sch. Bip.:
subsp. cilicica (Nägeli & Peter) P. D. Sell & C. West,
subsp. hoppeana,
subsp. macrantha (Ten.) S. Bräut. & Greuter;
12. Pilosella lactucella (Wallr.) P. D. Sell & C. West:
subsp. lactucella,
subsp. nana (Scheele) M. Laínz;
13. Pilosella macrotricha (Boiss.)F.W.Schultz&Sch.Bip.;
14. Pilosella officinarum Vaill.; also:
subsp. velutina (Hegetschw.) H. P. Fuchs
15. Pilosella peleteriana (Mérat) F. W. Schultz & Sch. Bip.:
subsp. peleteriana,
subsp. subpeleteriana (Nägeli & Peter) P. D. Sell;
16. Pilosella piloselloides aggr.:
Pilosella pavichii (Heuff.) Arv.-Touv.,
Pilosella piloselloides (Vill.) Soják:
subsp. bauhinii (Schult.) S. Bräut. & Greuter,
subsp. floccosa (Nägeli & Peter) S. Bräut. & Greuter,
subsp. magyarica (Peter) S. Bräut. & Greuter,
subsp. piloselloides,
subsp. praealta (Gochnat) S. Bräut. & Greuter,
subsp. rubrobauhinii (Schelk. & Zahn) S. Bräut. & Greuter;
17. Pilosella pseudopilosella (Ten.) Soják;
18. Pilosella saussureoides Arv.-Touv.;
19. Pilosella vahlii (Froel.) F. W. Schultz & Sch. Bip.;
20. Pilosella verruculata (Link) Soják:
subsp. akinfiewii (Woronow & Zahn) S. Bräut. & Greuter,
subsp. verruculata.
B. Intermediate, “collective” species (with their constituent species)
1. Pilosella alpicola <> cymosa: Pilosella petraea coll.:
Pilosella petraea F. W. Schultz & Sch. Bip.;
2. Pilosella alpicola <> hoppeana: Pilosella merxmuelleriana coll.:
Pilosella merxmuelleriana (S. Bräut.) S. Bräut. & Greuter;
3. Pilosella alpicola <> officinarum: Pilosella annae-vetterae coll.:
Pilosella annae-vetterae (Zahn) Soják
4. Pilosella argyrocoma <> pseudopilosella: Pilosella subulatissima coll.:
Pilosella subulatissima (Zahn) Mateo;
126 Bräutigam & Greuter: New treatment of Pilosella for the Euro-Mediterranean flora

5. Pilosella argyrocoma <> saussureoides: Pilosella nevadensis coll.:
Pilosella nevadensis (Arv.-Touv.) Mateo & Greuter;
6. Pilosella argyrocoma <> vahlii: Pilosella aranii coll.:
Pilosella aranii Mateo;
7. Pilosella aurantiaca <> caespitosa: Pilosella fuscoatra coll.:
Pilosella fuscoatra (Nägeli & Peter) Soják;
8. Pilosella aurantiaca <> caespitosa <> cymosa: Pilosella norrliniiformis coll.:
Pilosella norrliniiformis (Pohle & Zahn) Soják;
9. Pilosella aurantiaca <> caespitosa <> cymosa <> lactucella: Pilosella dimorphoides coll.:
Pilosella dimorphoides Norrl.;
10. Pilosella aurantiaca <> caespitosa <> lactucella: Pilosella subdecolorans coll.:
Pilosella subdecolorans (Norrl.) S. Bräut. & Greuter;
11. Pilosella aurantiaca <> cymosa: Pilosella guthnickiana coll.:
Pilosella guthnickiana (Hegetschw.) Soják;
12. Pilosella aurantiaca <> cymosa <> lactucella: Pilosella plaicensis coll.:
Pilosella plaicensis (Wo5.) Soják;
13. Pilosella aurantiaca <> cymosa <> officinarum: Pilosella biflora coll.:
Pilosella biflora (Arv.-Touv.) Arv.-Touv.;
14. Pilosella aurantiaca <> cymosa <> piloselloides: Pilosella macutensis coll.:
Pilosella macutensis (K. Malý & Zahn) Soják;
15. Pilosella aurantiaca <> echioides <> piloselloides: Pilosella muscelii coll.:
Pilosella muscelii (Prodán) S. Bräut. & Greuter;
16. Pilosella aurantiaca <> glacialis: Pilosella aurantella coll.:
Pilosella aurantella (Nägeli & Peter) Soják;
17. Pilosella aurantiaca <> glacialis <> hoppeana: Pilosella notha coll.:
Pilosella notha (Huter) S. Bräut. & Greuter;
18. Pilosella aurantiaca <> glacialis <> hoppeana <> lactucella: Pilosella amaurocephala coll.:
Pilosella amaurocephala (Peter) Soják;
19. Pilosella aurantiaca <> hoppeana: Pilosella substoloniflora coll.:
Pilosella substoloniflora (Peter) Soják;
20. Pilosella aurantiaca <> hoppeana <> lactucella: Pilosella eminens coll.:
Pilosella eminens (Peter) Soják;
21. Pilosella aurantiaca <> lactucella: Pilosella fusca coll.:
Pilosella blyttiana (Fr.)F.W.Schultz&Sch.Bip.,
Pilosella fusca (Vill.) Arv.-Touv.;
22. Pilosella aurantiaca <> lactucella <> officinarum: Pilosella peteriana coll.:
Pilosella peteriana (Käser) Holub;
23. Pilosella aurantiaca <> lactucella <> officinarum <> piloselloides: Pilosella moechiadia coll.:
Pilosella moechiadia (Peter) S. Bräut. & Greuter;
24. Pilosella aurantiaca <> lactucella <> piloselloides: Pilosella hyperborea coll.:
Pilosella hyperborea (Fr.)F.W.Schultz&Sch.Bip.;
25. Pilosella aurantiaca <> officinarum: Pilosella stoloniflora coll.:
Pilosella rubra (Peter) Soják,
Pilosella stoloniflora (Waldst. & Kit.) F. W. Schultz & Sch. Bip.;
26. Pilosella aurantiaca <> officinarum <> piloselloides: Pilosella trigenes coll.:
Pilosella trigenes (Peter) Soják;
27. Pilosella aurantiaca <> peleteriana: Pilosella bryhnii coll.:
Pilosella bryhnii (Omang) Soják;
28. Pilosella aurantiaca <> piloselloides: Pilosella calomastix coll.:
Pilosella calomastix (Peter) Soják,
Pilosella derubella (Gottschl. & Schuhw.) S. Bräut. & Greuter;
29. Pilosella breviscapa <> lactucella: Pilosella alturgelliana coll.:
Pilosella alturgelliana Mateo;
Willdenowia 37 – 2007 127

30. Pilosella caespitosa <> cymosa: Pilosella glomerata coll.:
Pilosella glomerata (Froel.) Fr.;
31. Pilosella caespitosa <> cymosa <> lactucella: Pilosella dubia coll.:
Pilosella dubia (L.) F. W. Schultz & Sch. Bip.,
Pilosella scandinavica (Dahlst.) Schljakov,
Pilosella tubulascens Norrl.;
32. Pilosella caespitosa <> cymosa lactucella <> officinarum: Pilosella polioderma coll.:
Pilosella polioderma (Dahlst.) Soják;
33. Pilosella caespitosa <> cymosa <> officinarum: Pilosella macranthela coll.:
Pilosella macranthela (Nägeli & Peter) Soják;
34. Pilosella caespitosa <> echioides <> lactucella <> piloselloides: Pilosella wolgensis coll.:
Pilosella wolgensis (Zahn) Soják;
35. Pilosella caespitosa <> echioides: Pilosella solacolui coll.:
Pilosella solacolui S. Bräut. & Greuter;
36. Pilosella caespitosa <> hoppeana: Pilosella levieri coll.:
Pilosella levieri (Peter) Soják [hoppeana <> onegensis];
37. Pilosella caespitosa <> hoppeana <> officinarum: Pilosella abakurae coll.:
Pilosella abakurae (Schelk. & Zahn) Soják;
38. Pilosella caespitosa <> hoppeana <> officinarum <> piloselloides: Pilosella aneimena coll.:
Pilosella aneimena (Nägeli & Peter) Soják;
39. Pilosella caespitosa <> lactucella: Pilosella floribunda coll.:
Pilosella cochlearis Norrl.,
Pilosella floribunda (Wimm. & Grab.) Fr.;
40. Pilosella caespitosa <> lactucella <> officinarum: Pilosella iserana coll.:
Pilosella iserana (R. Uechtr.) Soják,
Pilosella piloselliflora (Nägeli & Peter) Soják;
41. Pilosella caespitosa <> officinarum: Pilosella flagellaris coll.:
Pilosella flagellaris (Willd.) Arv.-Touv.,
Pilosella macrostolona (Gus. Schneid.) Soják,
Pilosella prussica (Nägeli & Peter) Soják;
42. Pilosella caespitosa <> officinarum <> piloselloides: Pilosella melinomelas coll.:
Pilosella leptoclados (Peter) Soják,
Pilosella melinomelas (Peter) Holub;
43. Pilosella caespitosa <> peleteriana: Pilosella chaetocephala coll.:
Pilosella chaetocephala (H. Hofm.) J. Holub;
44. Pilosella caespitosa <> peleteriana <> piloselloides: Pilosella dichotoma coll.:
Pilosella dichotoma (Lindeb.) Soják;
45. Pilosella caespitosa <> piloselloides: Pilosella polymastix coll.:
Pilosella erythrochrista (Nägeli & Peter) S. Bräut. & Greuter,
Pilosella polymastix (Peter) Holub;
46. Pilosella caespitosa <> pseudopilosella: Pilosella pawlowskiella coll.:
Pilosella pawlowskiella (Merxm.) Holub;
47. Pilosella cymosa <> echioides: Pilosella setigera coll.:
Pilosella setigera Fr.;
48. Pilosella cymosa <> echioides <> officinarum: Pilosella crassiseta coll.:
Pilosella crassiseta (Peter) Soják;
Pilosella cinereiformis (R. Meissn. & Zahn) S. Bräut. & Greuter
49. Pilosella cymosa <> echioides <> officinarum <> piloselloides: Pilosella setifolia coll.:
Pilosella setifolia (Touton) S. Bräut. & Greuter;
50. Pilosella cymosa <> echioides <> piloselloides: Pilosella megatricha coll.:
Pilosella megatricha (Borbás) Soják,
Pilosella schneidii (Schack & Zahn) S. Bräut. & Greuter;
128 Bräutigam & Greuter: New treatment of Pilosella for the Euro-Mediterranean flora

51. Pilosella cymosa <> glacialis: Pilosella laggeri coll.:
Pilosella laggeri (Rchb.f.)F.W.Schultz&Sch.Bip.;
52. Pilosella cymosa <> glacialis <> lactucella: Pilosella tendina coll.:
Pilosella tendina (Nägeli & Peter) Soják;
53. Pilosella cymosa <> glacialis <> lactucella <> officinarum: Pilosella tinctilingua coll.:
Pilosella tinctilingua (Zahn) Soják;
54. Pilosella cymosa <> glacialis <> officinarum: Pilosella pseudotrichodes coll.:
Pilosella pseudotrichodes (Zahn) Soják;
55. Pilosella cymosa <> hoppeana: Pilosella halacsyi coll.:
Pilosella halacsyi (Halácsy) Soják;
56. Pilosella cymosa <> lactucella: Pilosella corymbulifera coll.:
Pilosella corymbulifera (Arv.-Touv.) Arv.-Touv.;
57. Pilosella cymosa <> lactucella <> officinarum: Pilosella stenosoma coll.:
Pilosella stenosoma (Nägeli & Peter) Soják;
58. Pilosella cymosa <> lactucella <> piloselloides: Pilosella pseudosulphurea coll.:
Pilosella pseudosulphurea (Touton) Soják;
59. Pilosella cymosa <> officinarum: Pilosella kalksburgensis coll.:
Pilosella cymiflora (Nägeli & Peter) S. Bräut. & Greuter,
Pilosella kalksburgensis (Wiesb.) Soják;
60. Pilosella cymosa <> officinarum <> peleteriana: Pilosella fuernrohrii coll.:
Pilosella fuernrohrii (Vollm.) S. Bräut. & Greuter;
61. Pilosella cymosa <> officinarum <> piloselloides: Pilosella fallacina coll.:
Pilosella anchusoides Arv.-Touv.,
Pilosella fallacina (F.W.Schultz)F.W.Schultz,
Pilosella pilosellina (F.W.Schultz)Soják;
62. Pilosella cymosa <> peleteriana:Pilosella hybrida coll.:
Pilosella hybrida (Vill.) F. W. Schultz & Sch. Bip.;
63. Pilosella cymosa <> piloselloides: Pilosella ziziana coll.:
Pilosella bodewigiana (Zahn) Soják [cymosa <> pavichii],
Pilosella densiflora (Tausch) Soják,
Pilosella ziziana (Tausch) F. W. Schultz & Sch. Bip.
64. Pilosella cymosa <> piloselloides <> pseudopilosella: Pilosella litardiereana coll.:
Pilosella litardiereana (Zahn) Soják;
65. Pilosella echioides <> hoppeana: Pilosella balansae coll.:
Pilosella balansae (Boiss.) S. Bräut. & Greuter [hoppeana <> procera],
Pilosella erythrodonta (Zahn) S. Bräut. & Greuter,
Pilosella sterrochaetia (Nägeli & Peter) S. Bräut. & Greuter;
66. Pilosella echioides <> hoppeana <> officinarum: Pilosella grossheimii coll.:
Pilosella grossheimii (Zahn) Co=kunç. & Beyazo8lu;
67. Pilosella echioides <> hoppeana <> officinarum <> piloselloides: Pilosella chaetophyton coll.:
Pilosella chaetophyton (Zahn) S. Bräut. & Greuter;
68. Pilosella echioides <> hoppeana <> piloselloides: Pilosella budensis coll.:
Pilosella budensis (Borbás) Soják;
69. Pilosella echioides <> lactucella: Pilosella tephroglauca coll.:
Pilosella tephroglauca (Nägeli & Peter) Soják;
70. Pilosella echioides <> lactucella <> officinarum:Pilosella tephrophyton coll.:
Pilosella tephrophyton (Oborný & Zahn) Soják;
71. Pilosella echioides <> lactucella <> piloselloides: Pilosella occidentalis coll.:
Pilosella occidentalis (Nyár.) Soják;
72. Pilosella echioides <> officinarum: Pilosella bifurca coll.:
Pilosella bifurca (M.Bieb.)F.W.Schultz&Sch.Bip.,
Pilosella rothiana (Wallr.) F. W. Schultz & Sch. Bip.,
Pilosella schelkownikowii (Zahn) Soják [caucasica <> officinarum];
Willdenowia 37 – 2007 129

73. Pilosella echioides <> officinarum <> piloselloides: Pilosella heterodoxa coll.:
Pilosella euchaetia (Nägeli & Peter) Soják,
Pilosella heterodoxa (Tausch) Soják,
Pilosella heterodoxiformis (Zahn) S. Bräut. & Greuter;
74. Pilosella echioides <> piloselloides: Pilosella auriculoides coll.:
Pilosella auriculoides (Láng) Arv.-Touv.,
Pilosella calodon (Peter) Soják,
Pilosella procerigena (Litv. & Zahn) Sennikov [piloselloides <> procera],
Pilosella samokovensis (T. Georgiev & Zahn) S. Bräut. & Greuter [echioides <> pavichii];
75. Pilosella echioides <> piloselloides <> verruculata: Pilosella pannoniciformis coll.:
Pilosella pannoniciformis (Litv. & Zahn) Soják;
76. Pilosella echioides <> verruculata: Pilosella maschukensis coll.:
Pilosella maschukensis (Litv. & Zahn) Soják,
Pilosella woronowiana (Zahn) Soják [procera <> verruculata];
77. Pilosella galiciana <> pseudopilosella: Pilosella pseudogalliciana coll.:
Pilosella pseudogalliciana Mateo;
78. Pilosella galiciana <> vahlii: Pilosella unamunoi coll.:
Pilosella unamunoi (C. Vicioso) Mateo;
79. Pilosella glacialis <> hoppeana <> lactucella <> piloselloides: Pilosella salernicola coll.:
Pilosella salernicola (J. Vetter & Zahn) Soják;
80. Pilosella glacialis hoppeana: Pilosella acutifolia coll.:
Pilosella acutifolia (Vill.) Arv.-Touv.,
Pilosella pachypila (Peter) Soják,
Pilosella permutata (Nägeli & Peter) Soják;
81. Pilosella glacialis <> hoppeana <> lactucella: Pilosella lathraea coll.:
Pilosella brachycoma (Nägeli & Peter) H. P. Fuchs,
Pilosella lathraea (Peter) Soják;
82. Pilosella glacialis <> hoppeana <> officinarum: Pilosella basifurca coll.:
Pilosella basifurca (Peter) Soják;
83. Pilosella glacialis <> lactucella: Pilosella smithii coll.:
Pilosella smithii Arv.-Touv.;
84. Pilosella glacialis <> lactucella <> peleteriana: Pilosella aletschensis coll.:
Pilosella aletschensis (Zahn) Soják
85. Pilosella glacialis <> lactucella <> saussureoides: Pilosella triplex coll.:
Pilosella triplex (Peter) Soják;
86. Pilosella glacialis <> officinarum: Pilosella hypoleuca coll.:
Pilosella hypoleuca Arv.-Touv.;
87. Pilosella glacialis <> peleteriana: Pilosella subrubens coll.:
Pilosella subrubens Arv.-Touv.;
88. Pilosella glacialis <> piloselloides: Pilosella frigidaria coll.:
Pilosella frigidaria (Nägeli & Peter) Soják;
89. Pilosella glacialis <> pseudopilosella <> saussureoides: Pilosella faurei coll.:
Pilosella faurei Arv.-Touv.
90. Pilosella hoppeana <> lactucella: Pilosella viridifolia coll.:
Pilosella viridifolia (Peter) Holub;
91. Pilosella hoppeana <> lactucella <> vahlii:Pilosella noguerensis coll.:
Pilosella noguerensis Mateo;
92. Pilosella hoppeana <> officinarum: Pilosella hypeurya coll.:
Pilosella hypeurya (Peter) Soják;
93. Pilosella hoppeana <> officinarum <> piloselloides: Pilosella hypeurygenes coll.:
Pilosella hypeurygenes (A. W. Hill) S. Bräut. & Greuter;
94. Pilosella hoppeana <> officinarum <> pseudopilosella: Pilosella heteromelana coll.:
Pilosella heteromelana (Zahn) Mateo;
130 Bräutigam & Greuter: New treatment of Pilosella for the Euro-Mediterranean flora

95. Pilosella hoppeana <> officinarum <> verruculata: Pilosella sedelmeyeriana coll.:
Pilosella sedelmeyeriana (Zahn) S. Bräut. & Greuter;
96. Pilosella hoppeana <> peleteriana: Pilosella billyana coll.:
Pilosella billyana (de Retz) Mateo;
97. Pilosella hoppeana <> piloselloides: Pilosella ruprechtii coll.:
Pilosella arnoserioides (Nägeli & Peter) Soják,
Pilosella biglana (Bornm. & Zahn) S. Bräut. & Greuter [hoppeana <> pavichii],
Pilosella ruprechtii (Boiss.) Dostál;
98. Pilosella hoppeana <> pseudopilosella: Pilosella byzantina coll.:
Pilosella byzantina (Boiss.)P.D.Sell&C.West;
99. Pilosella hoppeana <> saussureoides: Pilosella eglandulosa coll.:
Pilosella eglandulosa (Sudre) Mateo;
100. Pilosella lactucella <> officinarum: Pilosella schultesii coll.:
Pilosella schultesii (F.W.Schultz)F.W.Schultz&Sch.Bip.,
Pilosella soleiroliana (Arv.-Touv. & Briq.) S. Bräut. & Greuter;
101. Pilosella lactucella <> officinarum <> piloselloides: Pilosella paragoga coll.:
Pilosella paragoga (Nägeli & Peter) Soják;
102. Pilosella lactucella <> peleteriana: Pilosella auriculiformis coll.:
Pilosella auriculiformis (Fr.)F.W.Schultz&Sch.Bip.;
103. Pilosella lactucella <> peleteriana <> piloselloides: Pilosella paragogiformis coll.:
Pilosella paragogiformis (Käser) Soják;
104. Pilosella lactucella <> piloselloides: Pilosella sulphurea coll.:
Pilosella koernickeana (Nägeli & Peter) Soják,
Pilosella sulphurea (Döll) F. W. Schultz & Sch. Bip.;
105. Pilosella lactucella <> pseudopilosella: Pilosella panticosae coll.:
Pilosella panticosae Mateo;
106. Pilosella lactucella <> pseudopilosella <> vahlii: Pilosella gudarica coll.:
Pilosella gudarica Mateo;
107. Pilosella lactucella <> saussureoides: Pilosella tardiuscula coll.:
Pilosella tardiuscula (Peter) Soják;
108. Pilosella lactucella <> saussureoides <> vahlii: Pilosella tremedalis coll.:
Pilosella tremedalis Mateo;
109. Pilosella lactucella <> vahlii: Pilosella pseudovahlii coll.:
Pilosella pseudovahlii (de Retz) Mateo;
110. Pilosella officinarum <> peleteriana: Pilosella longisquama coll.:
Pilosella longisquama (Peter) Holub;
111. Pilosella officinarum <> piloselloides: Pilosella brachiata coll.:
Pilosella arida (Freyn) Soják,
Pilosella brachiata (DC.)F.W.Schultz&Sch.Bip.,
Pilosella florentoides (Arv.-Touv.) P. D. Sell & C. West,
Pilosella leptophyton (Nägeli & Peter) S. Bräut. & Greuter,
Pilosella pavichiodes S. Bräut. & Greuter [officinarum <> pavichii],
Pilosella visianii F. W. Schultz & Sch. Bip.;
112. Pilosella officinarum <> pseudopilosella: Pilosella pintodasilvae coll.:
Pilosella pintodasilvae (de Retz) Mateo;
113. Pilosella officinarum <> saussureoides: Pilosella subtardans coll.:
Pilosella subtardans (Nägeli & Peter) Soják;
114. Pilosella officinarum <> verruculata: Pilosella kozlowskyana coll.:
Pilosella kozlowskyana (Zahn) Soják;
115. Pilosella peleteriana <> piloselloides: Pilosella promeces coll.:
Pilosella anobrachia (Arv.-Touv. & Gaut.) S. Bräut. & Greuter,
Pilosella mayeri (Vollm.) Soják,
Pilosella promeces (Peter) Holub;
Willdenowia 37 – 2007 131

116. Pilosella peleteriana <> pseudopilosella: Pilosella vansoestii coll.:
Pilosella vansoestii (de Retz) Mateo;
117. Pilosella peleteriana <> saussureoides: Pilosella portae coll.:
Pilosella portae (T. Durand & B. D. Jackson) Mateo & Greuter;
118. Pilosella piloselloides <> pseudopilosella: Pilosella fulviseta coll.:
Pilosella fulviseta (Bertol.) Soják;
119. Pilosella piloselloides <> pseudopilosella <> saussureoides: Pilosella tephrodes coll.:
Pilosella tephrodes (Nägeli & Peter) S. Bräut. & Greuter;
120. Pilosella piloselloides <> verruculata: Pilosella sintenisii coll.:
Pilosella sintenisii (Freyn) Soják;
121. Pilosella pseudopilosella <> saussureoides: Pilosella tardans coll.:
Pilosella tardans (Peter) Soják;
122. Pilosella saussureoides <> vahlii: Pilosella caballeroi coll.:
Pilosella caballeroi (Mateo) Mateo;
Appendix: Nomenclatural validations and comments
Pilosella anobrachia (Arv.-Touv. & Gaut.) S. Bräut. & Greuter, comb. nov. ≡Hieracium ano-
brachion Arv.-Touv. & Gaut., Hieracioth. 10: [in schedis] Gall. No. 547. 1900.
Pilosella aurantiaca subsp. auropurpurea (Nägeli & Peter) S. Bräut. & Greuter, comb. nov.
Hieracium aurantiacum subsp. auropurpureum Nägeli & Peter, Hierac. Mitt.-Eur. 1: 295. 1885.
Pilosella balansae (Boiss.) S. Bräut. & Greuter, comb. nov. Hieracium balansae Boiss., Diagn.
Pl. Orient., ser. 2, 6: 119. 1859.
Pilosella biglana (Bornm. & Zahn) S. Bräut. & Greuter, comb. nov. Hieracium biglanum
Bornm. & Zahn in Repert. Spec. Nov. Regni Veg. 16: 181. 1919.
Pilosella bonaquae (Buttler & W. Lippert) S. Bräut. & Greuter, comb. nov. Hieracium bo-
naquae Buttler & W. Lippert in Strid, Mount. Fl. Greece 2: 608. 1991.
Pilosella brzovecensis (Horvat & Paw5.) S. Bräut. & Greuter, comb. nov. Hieracium brzo-
vecense Horvat&Paw5. in Acta Soc. Bot. Poloniae 32: 486. 1963.
Pilosella chaetophyton (Zahn) S. Bräut. & Greuter, comb. nov. Hieracium chaetophyton Zahn
in Vestn. Tiflissk. Bot. Sada, ser. 2, 3-4: 32. 1927.
Pilosella cinereiformis (R. Meissn. & Zahn) S. Bräut. & Greuter, comb. nov. Hieracium
cinereiforme R. Meissn. & Zahn in Engler, Pflanzenr. 82: 1390. 1923.
The name Hieracium cinereiforme first appeared in a letter that Zahn addressed to
Touton and was published by the latter (in Jahrb. Nassauischen Vereins Naturk. 73:
72, 1921). However, conditions for valid publication were not met there, as no de-
scriptive elements referring to the species itself are included (see also comments
under Pilosella hypeurygenes, below). Zahn’s co-author “Meißner” could be iden-
tified as Richard Meißner, of Bernburg (c. 1855-1915), of whom an obituary notice
can be found in Zobel (Verz. Anhalt Phan. 4: III-IV. 1920).
Pilosella collina (Gochnat) Soják in Preslia 43: 185. 1971 Hieracium collinum Gochnat, Tent.
Med.-Bot. Cichorac.: 17. 1808. – Lectotype (designated here): The illustration of Hieracium Col-
linum in Gochnat, Tent. Med.-Bot. Cichorac.: t. I. 1808. – Epitype (designated here): “Hieracium
collinum Gochn. [manu Copin de Miribel]”; “Hieracium angustifolium Mt. Cenis” [manu
Bonjean?]; “hier. angustifol.M.Cenisesth. collinum Gochn.” [manu Villars], [Bonjean?]in
herb. Villars (GRM 1837.27543, photo!) [= Pilosella cymosa (L.) F. W. Schultz & Sch. Bip.].
Gochnat’s Hieracium collinum is a long standing source of confusion and debate.
Nägeli & Peter (Hierac. Mitt.-Eur. 1: 298. 1885) used it for the species that previous
132 Bräutigam & Greuter: New treatment of Pilosella for the Euro-Mediterranean flora

authors, and again Zahn (in Engler, Pflanzenr. 82: 1268. 1923), knew al H. pratense
Tausch 1828 and is now generally designated as H. caespitosum Dumort. 1827.
Zahn (l.c.: 1388, and elsewhere in his writings) included it in H. fallax Willd. 1809,
a name that for priority reasons it should have displaced. The name H. collinum is
not currently used in any sense. Gochnat’s original illustration shows fair similarity
with H. caespitosum, H. fallax and H. cymosum L. 1753 but does not coincide com-
pletely with any of them, so that it cannot be interpreted with full certainty.
The question of possible type material is still open (pers. comm. by Günter
Gottschlich, Tübingen). There are no specimens of either Gochnat or his mentor
Villars at Strasbourg (STR). One specimen labelled Hieracium collinum Gochnat is
present in Villars’ herbarium in Grenoble, of which Vincent Poncet kindly sent us a
photograph and which, as was confirmed by Gottschlich, belongs to H. cymosum.
However, it originates from Mt Cenis, so it can hardly be original material for H.
collinum, said to have been found near Marlenheim, Alsatia.
In order to obviate the need to either take up the name Pilosella collina (Goch-
nat) Soják in an unfamiliar sense or propose its formal rejection, we designate the
original illustration (being the single secure original element available) as the
lectotype of Hieracium collinum and, in the same time, designate an epitype that
belongs to H. cymosum, a name much older than H. collinum.
Pilosella cymiflora (Nägeli & Peter) S. Bräut. & Greuter, comb. nov. Hieracium cymiflorum
Nägeli & Peter, Hierac. Mitt.-Eur. 1: 424. 1885.
Pilosella cymosa subsp. vaillantii (Tausch) S. Bräut. & Greuter, comb. nov. Hieracium vaillan-
tii Tausch in Flora 11, Ergänzungsbl. 1: 57. 1828 Hieracium cymosum subsp. vaillantii (Tausch)
Nyman, Consp. Fl. Eur.: 452. 1879 Pilosella vaillantii (Tausch) Soják in Preslia 43: 184. 1971.
Pilosella derubella (Gottschl. & Schuhw.) S. Bräut. & Greuter, comb. nov. Hieracium deru-
bellum Gottschl. & Schuhw. in Ber. Bayer. Bot. Ges. 69-70: 148. 2000.
Pilosella echioides subsp. proceriformis (Nägeli & Peter) S. Bräut. & Greuter, comb. nov.
Hieracium echioides subsp. proceriforme Nägeli & Peter, Hierac. Mitt.-Eur. 1: 487. 1885 Hie-
racium proceriforme (Nägeli & Peter) Zahn in Engler, Pflanzenr. 82: 1369. 1923 Pilosella pro-
ceriformis (Nägeli & Peter) Soják in Folia Geobot. Phytotax. 6: 218. 1971.
Pilosella erythrochrista (Nägeli & Peter) S. Bräut. & Greuter, comb. nov. Hieracium erythro-
christum (Nägeli & Peter) Üksip in Komarov, Fl. SSSR 30: 520. 1960 Hieracium arvicola
subsp. erythrochristum Nägeli & Peter, Hierac. Mitt.-Eur. 1: 668. 1885.
This name must be used for the species currently known as Hieracium arvicola
Nägeli & Peter (Hierac. Mitt.-Eur. 1: 666. 1885) or Pilosella arvicola Soják (in Pres-
lia 43: 186. 1971), two illegitimate names: both were applied, upon publication, to a
species that included the type of the earlier and legitimate H. assimilatum (Norrl.)
Norrl. (in Mela, Lyhyk. Kasvioppi Kasvio, ed. 2: 2: 212, after Aug. 1884) = Pilosella
septentrionalis subsp. assimilata Norrl. (in Acta Soc. Fauna Fl. Fenn. 2(4): 147,
Feb.-June 1884).
Nägeli & Peter (l.c.: 672) treated Hieracium assimilatum as a subspecies of H.
arvicola. Thanks to the kindness of Alexander Sennikov, Helsinki, we could study
excellent digital images of original specimens of, or H. assimilatum, which we found
to be indistinguishable from Pilosella floribunda. This leaves the name H. arvicola
(typified by the type of Nägeli & Peter’s subsp. arvicola – see ICBN Art. 7.5) as the
oldest binomial that belongs to the present species. Nevertheless, being illegitimate,
it is unavailable for use, same as the homotypic P. arvicola.
The next possible candidate for naming the present species appeared to be
Hieracium curvulum Norrl. (in Mela, Suomen Koulukasvio, ed. 4: 526. 1899), treated
Willdenowia 37 – 2007 133

by Zahn (1923: 1476) as a subspecies of H. arvicola. Sennikov again kindly provided
type photographs, which show a plant that we consider to belong to Pilosella pilo-
selloides subsp. praealta.
Pilosella erythrodonta (Zahn) S. Bräut. & Greuter, comb. nov. Hieracium erythrodontum Zahn
in Vandas, Reliq. Formán.: 347. 1909 [et in Engler, Pflanzenr. 82: 1376. 1923].
We were at first disinclined to accept the basionym as validly published in 1909, for
exactly the same reasons as mentioned below under Pilosella hypeurygenes.When
proposing the new species Hieracium erythrodontum, Zahn described in detail the
equally new H. erythrodontum subsp. philippopolitanum andalsoprovideddescrip
-
tive matter for subsp. erythrodontum, but apparently none for the species as such. On
second thoughts, we accept that the short paragraph intercalated between the subspe-
cies descriptions, while not explicitly referring to the species as a whole, can be con-
sidered as an implicit species diagnosis. As Zahn mentions differences from two
other Hieracium species, he must logically have had the whole new species in mind.
Pilosella fuernrohrii (Vollm.) S. Bräut. & Greuter, comb. nov. Hieracium fuernrohrii Vollm.
in Denkschr. Königl. Bot. Ges. Regensburg 9: 72. 1905.
Pilosella heterodoxiformis (Zahn) S. Bräut. & Greuter, comb. nov. Hieracium heterodoxi-
forme Zahn in Engler, Pflanzenr. 82: 1514. 1923.
The name Hieracium heterodoxiforme Zahn first appears in a paper by Touton (in
Jahrb. Nassauischen Vereins Naturk. 74: 38. 1922), where it is not, however, validly
published for lack of a description of the species. Only the three constituent subspe-
cies are described, but their names are not validly published either (ICBN, Art. 43.1).
The situation is thus the same as for H. hypeurygenes, mentioned below. Subse-
quently, when he validly published H. heterodoxiforme, Zahn (l.c. 1923) adopted a
different circumscription for the species, restricting it to one of the three original sub-
species, for which he maintained the former designation subsp. nassovicum. As that
subspecies is coextensive with the species, it included the type of the species name.
This contravenes the present nomenclatural rules (ICBN, Art. 26.1), and yet again, H.
hypeurygenes subsp. nassovicum fell short of valid publication. It shares this fate
with the binomial H. nassovicum, proposed by Gottschlich (in Feddes Repert. 108:
105. 1997) as a new combination but for which there is no basionym.
Pilosella hoppeana subsp. macrantha (Ten.) S. Bräut. & Greuter, comb. nov. Hieracium pilo-
sella var. macranthum Ten., Fl. Napol. 4: 114. 1830 Hieracium macranthum (Ten.) Ten., Fl.
Napol. 5: 190. 1836 Pilosella macrantha (Ten.) F. W. Schultz & Sch. Bip. in Flora 45: 422. 1862
Hieracium pilosella subsp. macranthum (Ten.) Arcang., Comp. Fl. Ital.: 435. 1882 Hieracium
hoppeanum subsp. macranthum (Ten.) Nägeli & Peter, Hierac. Mitt.-Eur. 1: 125. 1885.
Pilosella hypeurygenes (A. W. Hill) [Co=kunç. & Beyazo8lu in Edinburgh J. Bot. 59: 320. 2002,
comb. inval., ex] S. Bräut. & Greuter, comb. nov. Hieracium hypeurygenes [ZahninVestn.
Tiflissk. Bot. Sada, ser. 2, 3-4: 32. 1927, nom. inval., ex] A. W. Hill, Index Kew., Suppl. 8: 114.
1933.
One of the tricky nomenclatural messes alas typical for Zahn’s Hieracium work.
When he proposed the new species H. hypeurygenes, Zahn did not describe it, but in-
stead he described the three new subspecies he recognized in it. As he did not de-
scribe the taxon as Art. 41.3(a) of the ICBN mandates (none of the subspecies is, of
course, co-extensive with the species), neither the species name nor (ICBN, Art.
43.1) any of the subspecies names were validly published. When in 1933 the species
name was listed as accepted in the Index Kewensis the situation was different, in so
far as Art. 41.3(b) permits validation by reference to a previously published descrip-
tion of ataxon. The difference between the and ais subtle but logically justified, and
no doubt intentional. One must thus accept Hieracium hypeurygenes A. W. Hill as be-
134 Bräutigam & Greuter: New treatment of Pilosella for the Euro-Mediterranean flora

ing validated by reference to Zahn’s 1927 description of H. hypeurygenes subsp.
hypeurygenes. Coîkunçelebi & Beyazo8lu, when proposing the combination Pilo-
sella hypeurygenes with reference to Zahn, did essentially the same as Hill had done
– but they were publishing in 2002. Reference to a Latin description was not then suf-
ficient to validate the name of a new taxon: indication of a type specimen by use of
the word “type” or an equivalent, and of the herbarium of deposit, would have been
necessary to obtain the benefit of Art. 33.7(d) and validate the intended combination.
Pilosella leptophyton (Nägeli & Peter) S. Bräut. & Greuter, comb. nov. Hieracium leptophy-
ton Nägeli & Peter, Hierac. Mitt.-Eur. 1: 642, 827. 1885.
Pilosella merxmuelleriana (S. Bräut.) S. Bräut. & Greuter, comb. nov. Hieracium merxmuel-
lerianum S. Bräut. in Bot. Jahrb. Syst. 107: 5. 1985.
Pilosella moechiadia (Peter) S. Bräut. & Greuter, comb. nov. Hieracium moechiadium Peter
in Bot. Jahrb. Syst. 5: 491. 1884 Hieracium cineraria subsp. moechiadium (Peter) Nägeli & Pe-
ter, Hierac. Mitt.-Eur. 1: 665. 1885
Pilosella muscelii (Prodán) S. Bräut. & Greuter, comb. nov. Hieracium muscelii Prodán in
Bul. ñti. Acad. Republ. Populare Romîne, Sect. Biol. ñti. Agric., Ser. Bot. 9: 311. 1957.
Pilosella nevadensis (Arv.-Touv.) Mateo & Greuter, comb. nov. Hieracium pilosella var. neva-
dense Arv.-Touv., Hier. Gall. Hisp. Cat.: 7. 1913 ≡Hieracium nevadense (Arv.-Touv.) Prain, In-
dex Kew., Suppl. 5: 130. 1921.
Pilosella notha (Huter) [Soják in Cas. Nár. Muz., Odd. Prír. 150: 139. 1982, comb. inval., ex] S.
Bräut. & Greuter, comb. nov. Hieracium nothum Huter, Fl. Gefäss-Pfl. Höhlenstein: 30. 1872.
Soják, in 1982, proposed the same combination but gave an earlier reference (Huter
in Österr. Bot. Z. 20: 338. 1870) for the inteded basionym. In that place, Hieracium
nothum appears as a nomen nudum, merely with an indication of the locality and sup-
posed parentage.
Pilosella pavichiodes S. Bräut. & Greuter, sp. nov. – Type: Romania, reg. Hunedoara, in quercetis
penes vicum Petreîti ad Sebeî,Borza (CL). – Latin decription: see Borza in Feddes Repert. Spec.
Nov. Regni Veg. 61: 92-92. 1958, sub Hieracium pavichiodes, nom. inval. [sine design. typi].
Pilosella piloselloides subsp. bauhinii (Schult.) S. Bräut. & Greuter, comb. nov. Hieracium
bauhinii Schult. in Besser, Prim. Fl. Galiciae Austriac. 2: 149. 1809 (post 16 Mar.); et Schult.,
Observ. Bot.: 164. 1809 (sero) Pilosella bauhinii (Schult.) Arv.-Touv. in Bull. Soc. Dauphin.
Echange Pl.: 280. 1880 Hieracium magyaricum subsp. bauhinii (Schult.) Nägeli & Peter,
Hierac. Mitt.-Eur. 1: 592. 1885.
Pilosella piloselloides subsp. floccosa (Nägeli & Peter) S. Bräut. & Greuter, comb. nov. Hie-
racium florentinum subsp. floccosum Nägeli & Peter, Hierac. Mitt.-Eur. 1: 550. 1885.
Pilosella piloselloides subsp. magyarica (Peter) S. Bräut. & Greuter, comb. nov. Hieracium
magyaricum Peter in Bot. Jahrb. Syst. 5: 217. 1884 Hieracium bauhinii subsp. magyaricum
(Peter) Zahn in Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 40: 703. 1906.
Pilosella piloselloides subsp. praealta (Gochnat) S. Bräut. & Greuter, comb. nov. Hieracium
praealtum Gochnat, Tent. Med.-Bot. Cichorac.: 17. 1808 Pilosella praealta (Gochnat) F. W.
Schultz & Sch. Bip. in Flora 45: 429. 1862 Hieracium florentinum subsp. praealtum (Gochnat)
Nägeli & Peter, Hierac. Mitt.-Eur. 1: 539. 1885 Hieracium piloselloides subsp. praealtum
(Gochnat) Zahn in Hegi, Ill. Fl. Mitt.-Eur. 6: 1230. 1929.
Pilosella piloselloides subsp. rubrobauhinii (Schelk. & Zahn) S. Bräut. & Greuter, comb. nov.
Hieracium bauhinii subsp. rubrobauhinii Schelk. & Zahn in Vestn. Tiflissk. Bot. Sada 29: 4.
1913 Hieracium rubrobauhinii (Schelk. & Zahn) Üksip in Komarov, Fl. SSSR 30: 460. 1960
Pilosella rubrobauhinii (Schelk. & Zahn) Sennikov in Bot. Zurn. 83(3): 69. 1998.
Willdenowia 37 – 2007 135

Pilosella portae (T. Durand & B. D. Jackson) Mateo & Greuter, comb. nov. Hieracium silvati-
cum Porta, Veg. Itin. Iber.: 45. 1892; et in Atti Imp. Regia Accad. Agiati Rovereto, ser. 2, 9: 45?.
1892 [non Hieracium sylvaticum (L.) Gouan 1773] Hieracium portae [Willk., Suppl. Prodr. Fl.
Hispan.: 118. 1893, nom. prov., ex] T. Durand & B. D. Jackson, Index Kew., Suppl. 1: 210. 1902.
Pilosella samokovensis (T. Georgiev & Zahn) S. Bräut. & Greuter, comb. nov. Hieracium
samokovense T. Georgiev & Zahn in Izv. Bulg. Bot. Druz. 5: 94. 1932.
Pilosella schneidii (Schack & Zahn) S. Bräut. & Greuter, comb. nov. Hieracium schneidii
Schack & Zahn in Behr, Herb. Hierac.: [in schedis] No. 910. 1938.
Pilosella sedelmeyeriana (Zahn) S. Bräut. & Greuter, comb. nov. Hieracium sedelmeyerianum
Zahn in Vestn. Tiflissk. Bot. Sada, ser. 2, 3-4: 28. 1927.
Pilosella setifolia (Touton) S. Bräut. & Greuter, comb. nov. Hieracium setifolium Touton in
Jahrb. Nassauischen Vereins Naturk. 74: 41. 1922.
Pilosella solacolui [Holub in Folia Geobot. Phytotax. 12: 306. 1977, basion. inval., ex] S. Bräut.
&Greuter,sp. nov. – Type: Romania, “(Crangu M.) R.S. [Râmnicu-Sarat]”, 1911, Brândza(CL
07071). – Latin Description: see Nyárády in Sbvulescu, Fl. Rep. Pop. Române 10: 723. 1965, sub
Hieracium solacolui Prodán (Fl. Român., ed. 2: 1144. 1939, nom. inval. [sine diagn. lat.] ex
Nyár., l.c., nom. inval. [sine design. typi]).
We are indebted to Irina Goia, Cluj, for providing us with a whole series of detailed
digital images of the type specimen.
Pilosella soleiroliana (Arv.-Touv. & Briq.) S. Bräut. & Greuter, comb. nov. Hieracium solei-
rolianum Arv.-Touv. & Briq. in Annuaire Conserv. Jard. Bot. Genève 5: 108. 1901 Hieracium
schultesii subsp. soleirolianum (Arv.-Touv. & Briq.) Zahn in Engler, Pflanzenr. 82: 1217. 1923.
Pilosella sterrochaetia (Nägeli & Peter) S. Bräut. & Greuter, comb. nov. Hieracium sterro-
chaetium Nägeli & Peter, Hierac. Mitt.-Eur. 1: 493, 817. 1885.
Pilosella subdecolorans (Norrl.) S. Bräut. & Greuter, comb. nov. Hieracium subdecolorans
Norrl. in Acta Soc. Fauna Fl. Fenn. 3(4): 69. 1888.
Pilosella tephrodes (Nägeli & Peter) S. Bräut. & Greuter, comb. nov. Hieracium tephrodes
Nägeli & Peter, Hierac. Mitt.-Eur. 1: 641, 827. 1885.
Pilosella verruculata subsp. akinfiewii (Woronow & Zahn) S. Bräut. & Greuter, comb. nov.
Hieracium incanum subsp. akinfiewii Woronow & Zahn in Vestn. Tiflissk. Bot. Sada 12: 13.
1908 Pilosella akinfiewii (Woronow & Zahn) Sennikov in Bot. Zurn. 83(3): 62. 1998.
Pilosella walteri-langii (Gottschl.) S. Bräut. & Greuter, comb. nov. Hieracium walteri-langii
Gottschl. in Mitt. Pollichia Jahrb. Natur Landschaft 77: 183. 1990.
Acknowledgements
We want to thank those who, in various ways, assisted by advice and by answering our questions
and requests. They include Birgit Gemeinholzer, Irina Goia, Günter Gottschlich, Heinz Helt-
mann, Gonzalo Mateo, Vincent Poncet, Franz Schuhwerk and Alexander Sennikov.
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Addresses of the authors:
Dr Siegfried Bräutigam, Staatliches Museum für Naturkunde Görlitz, Postfach 300 154, D-02806
Görlitz; e-mail: Siegfried.Braeutigam@smng.smwk.sachsen.de
Prof. Dr Werner Greuter, Botanischer Garten und Botanisches Museum Berlin-Dahlem, Freie
Universität Berlin, Königin-Luise-Str. 6-8, D-14195 Berlin; e-mail: w.greuter@bgbm.org
Willdenowia 37 – 2007 137