Generic and Sectional Limits in Senecio (Compositae): I.

Progress Report
Author(s): C. Jeffrey, P. Halliday, M. Wilmot-Dear and S. W. Jones
Reviewed work(s):
Source: Kew Bulletin, Vol. 32, No. 1 (1977), pp. 47-67
Published by: Springer on behalf of Royal Botanic Gardens, Kew
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 Generic and sectional limits in Senecio
(Compositae):
I. Progress Report
C. JEFFREY, P. HALLIDAY, M. WILMOT-DEAR* & S. W. JONES+

Summary. Proposalsfor a revisionof genericand sectionallimits in Senecioand allied genera

are outlined and, on the basis of the study of 182 speciesof Seneciosensu lato, a schemeis
put forwardto serve as a basisfor the incorporationand integrationof furtherdata. Taxo-
nomic implicationsare brieflydiscussed.

INTRODUCTION

The very variable generic and sectional concepts employed by different

authors in Senecioand allied genera have made it impossible to give definitive
answers to enquiries requesting the correct names of many species and/or
specimens submitted for identification. This unsatisfactory situation, and
the need to adopt meaningful generic concepts for an account of the
Senecioneae for the 'Flora of Tropical East Africa', prompted the initiation of
the present study. Much of the inconsistency in treatments hitherto accorded
to the group has resulted from authors confining their studies to plants of
particular geographical areas; classifications applicable on a world-wide
basis have not been produced. The sheer size of the group (an estimated
3,000 species) poses problems if a holistic approach be attempted. To enable
a study to be completed in a reasonable time some choice, both of species
and of characters to be investigated, must be made and in the choice of
characters, ease and readiness of observation has to be a paramount con-
sideration. Such an eclectic approach has obvious dangers, but it is used in
this study, not to produce a definitive classification, but to provide a uniform
criterion throughout the group, against which all available evidence might
be evaluated and consistent systematic conclusions therefore drawn. It
envisages the following stages of study by which, it is hoped, the often opposed
requirements of practicality and inclusiveness may be reconciled.

I. Easily observed characters exhibited by the surfaces of the cypselas,

the pappus, the style-arm apices, the anther-collars, the anther-bases, the
ray florets and the abaxial surfaces of the leaves are selected for examination.
Data thus obtained are supplemented, when considered necessary, by study
of the vegetative habit, leaf shape, venation and attachment, flower colour,
inflorescence type, involucre type, the endothecial (exothecial) cell thicken-
ings, the pollen grain surface, the pollen grain wall stratification, the
configuration of the stigmatic surfaces and, when available, by chromosome
numbers, obtained from the compilations of Fedorov (1969), Moore (1973,
1974) and occasionally other sources.

Acceptedfor publication5 December1975.

* The Herbarium, Royal BotanicGardens,Kew.
$ Departmentof Botany,BritishMuseum (NaturalHistory),London.
47

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48 KEW BULLETIN VOL. 32(I)

A preliminary selection of species is made, of which the above

2.
characters are studied, partly on the basis of existing taxonomic treatments,
partly on the basis of some fifteen years' working knowledge of the group. It
is hoped in this way to select 'modal' species around which other species
might reliably be clustered on the basis of rapid examination and/or data
already available in the literature. In the latter process, further 'modal'
species may be selected for detailed examination should it be found neces-
sary.
3. All other available information, obtained from the literature, is
integrated with the results obtained from the above two stages, to produce a
classification, with taxonomic concepts consistent on a world-wide basis.
4. A critical revision of the nomenclature at subtribal, generic, sub-
generic, sectional, subsectional and serial ranks is carried out.
This paper summarizes the results obtained from study of the first 182
of the initially chosen 'modal' species (stages I and 2). Only species currently
classified within Senecioin the Kew herbarium have been considered at this
early stage. While this restriction of choice is to a large extent purely
arbitrary, it does ensure that variation in Seneciois considered and, hopefully,
understood before representatives of the currently most generally accepted
segregate genera are examined and attempts made to integrate them into
the taxonomic system.

MATERIAL AND METHODS

All material was taken from herbarium specimens selected (by C. J.) from
the Herbarium of the Royal Botanic Gardens, Kew (K). For scanning
electron microscopy (by M. W.-D.), dry cypselas and unacetolyzed pollen
grains were mounted directly on to stubs using double-sided sellotape, gold-
coated, and photographed at approximately 30, X 100oo,X 300, x1ooo,
x 3000 (cypselas) and x 2-3000 and x 6-7000 (pollen grains) on a Jeol
JSM I scanning electron microscope. For light microscopy, florets and pieces
of leaf were softened by boiling, dissected and mounted in lactic acid,
examined and drawn (by P. H. and S. W. J.) under Nikon SMZ-2 binocular
dissecting and Watson Service monocular compound microscopes. For
transmission electron micrography of sections of pollen grains, we are greatly
indebted to ProfessorJohn J. Skvarla, University of Oklahoma. A list of the
species and specimens studied is deposited in the library of the Royal
Botanic Gardens, Kew. Determinations are by the senior author.

RESULTS

The results are presented in two parts: an account and discussion of

variation in the characters studied; and a grouping of species based on
analysis of this variation.

I. Characters Studied

I. Cypselasurfaces
Cypselas are glabrous or bear duplex hairs of varying distribution and
density. The hairs vary in length/breadth ratio, shape of the apex, and

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GENERIC AND SECTIONAL LIMITS IN SENECIO 49
prominence of the wall thickenings. The superficial cells vary in length/
breadth ratio, prominence of the transverse and longitudinal walls, and in
the sculpture of the surface-smooth, striate or papillose; 'smooth' is
employed here to cover all states apart from the latter two.
2. Pollengrainsurfaces
The pollen grains are uniformly tricolporate and echinate and vary
only in the number, density and length/breadth ratio of the spines, and in
the degree of prominence of the colpi.
3. Pappus
The pappus may be uniform, composed of only one type of hair, or
dimorphic, composed of two distinct hair types. Pappus-hairs of uniform
pappi are either tapered, with divergent teeth, or less often clavate with
subconnivent teeth. The dimorphic pappus has, in addition to numerous
tapered hairs, a number of fluked hairs (Drury, 1967), with retrorsely
directed apical teeth, or rarely, tapered hairs with merely reduced teeth.
Pappus-hairs themselves vary in colour, texture, stoutness, number of cells
in cross-section (that at about five cells below the apex is quoted here), cell
length/breadth ratio, and angle of projection and prominence of the teeth.
Pappus-hairs may be present in all florets or absent from the ray-florets.
4. Style-armapices
The style-arm apices may be truncate, convex, rounded or rarely
rhombic and bear papillae which vary in length, shape and distribution.
Usually they fringe the apices, run down the outer surfaces of the style-arms
and rapidly become less prominent as they recede from the apices. They are
termed decurrent when this tendency to run down is very marked and
accompanied by only a very gradual decrease in length. Sometimes a median
fascicle of longer, more or less fused papillae is present on the apex of each
style-arm, in addition to the fringing apical papillae.
5. Anther-collars
The anther-collars vary in shape from cylindrical to balusterform
(Drury, 1967), in length, number of cells, cell shape and in degree of
differentiation of enlarged, thin-walled basal cells.
6. Anther-bases
The anther-bases are always sterile and vary from very short to
exceeding the length of the anther-collar; they are obtuse, acute or, occasion-
ally, lobed.
7. Ray-florets
Ray-florets may be present, present but reduced, or absent, and vary in
venation and in the presence or absence of glandular hairs on the corolla-
tube. Such hairs may be uniseriate and/or biseriate, clavate or capitate, and
vary in length and number of cells. Ray-florets are termed 'hairy' if both
biseriate and uniseriate hairs are present. Non-glandular hairs are occasion-
ally also present.
8. Leaf-hairs
Leaves are sometimes glabrous but usually bear glandular hairs,
similar to, and varying like, those on the ray-florets; non-glandular hairs
are often also present and are of two main types-uniform, with all cells
I--D

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50 KEW BULLETIN VOL. 32(I)

similar (apart from sometimes an apically setiferous or mucilaginous cell)

throughout their length; or non-uniform, with short basal cells and an apical
elongated cell or cells. The former vary in length, cell shape and number,
the latter in the number of cells in each part, the degree of differentiation
between the parts, and the orientation of the cross-walls in the apical part;
this orientation forms a continuous series from transverse, through oblique,
to oblique and laterally extended; in the latter case, the hairs are incipiently,
asymmetrically or symmetrically T-shaped.
9. Habit
Plants may be annual herbs, perennial subscapose or uniformly leafy
(described as 'leafy') herbs, subshrubs, shrubs or small trees; some are
epiphytes, some scandent, some succulent, some variously otherwise
specialized in life-form.
10o. Leaf shape,venationand attachment
Leaves may be petiolate, pseudopetiolate (with petioloid base) or
sessile, pinnately (including parallelopinnately), pinnatopalmately or
palmately veined, entire, toothed or variously pinnately or palmately
divided, often expanded, auricled, amplexicaul or sheathing at the base,
occasionally peltate, sometimes succulent, ericoid or reduced to scales;
petioles may be concavo-convex or more or less terete in cross-section;
occasionally they have a persistent thickened base.
I I. Involucre
The bracts of the involucre are usually free, rarely more or less united,
and usually provided with a few outer accessory bracts (calyculate); rarely
these are lacking (ecalyculate).
I2. Flowercolour
Radiate heads are usually homochromous and yellow, rarely orange-
yellow, purplish, reddish or white, less often heterochromous with white
rays and yellow disc. Discoid heads vary, white, yellow, brown, red or purple
in various shades.

13. Inflorescence
Capitula may be (rarely) solitary or in terminal, terminal and upper
axillary or sometimes axillary cymose corymbs, compound corymbs,
compound paniculoid thyrses, or racemiform thyrses.
14. Endothecialcells
Endothecial cells vary from short and thickwalled, with thickenings
confined to the poles of the cells (polar of Drury, 1967), to elongated, thin-
walled, with thickenings along the whole length of the radial walls (radial
of Drury, 1967).

15. Pollengrain walls

Pollen grain walls vary in thickness and structure, particularly in
development and length of the columellae and in the thickness and degree
of elaboration of the foot layer of the ectoexine.
16. Stigmaticsurfaces
The stigmatic surfaces vary from widely separated throughout their
length, through variously approximated or contiguous along their whole

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GENERIC AND SECTIONAL LIMITS IN SENECIO 51

length to completely confluent and covering the whole inner surfaces of the
style-arms.
DISCUSSION

The most important facts to emerge from this preliminary stage of the
study are that intermediate states are always found between the various
character-states of the above characters and that no one character-state is
consistently correlated with any character-state of any other character. As
a result, it is impossible to define groups of species other than on the basis of
syndromes of rather loosely associated character-states and it is also impos-
sible meaningfully to allocate character-states to classes, such as might be
necessary for their scoring for use in numerical analyses. This continuity of
character-states and lack of correlation between states of different characters
is observed, it should be emphasized, in all characters, including those, such
as length and distribution of style-arm papillae, length of anther-bases,
shape and degree of cellular differentiation of the anther-collars, type of
leaf-hairs, distribution of endothecial cell thickenings, and configuration of
the stigmatic surfaces, that at one time or another have been considered by
various authors to be of use in defining genera or groups of genera. The
importance to be accorded to any character-state in the construction of taxa
can, and must be, assessed only by its consideration against the background
of other character-states with which it is associated in any particular species,
and on that basis is to be assessed for that particular species only. Any
attempt at overall weighting of a character-state-or, conversely, at non-
weighting of all character-states-throughout the group as a whole would
ignore both the eclectic nature of the study and the existence of different
character-state syndromes in different species and would therefore be unlikely
to result in a meaningful classification.

2. Groupings of the Species

As a result of study of the above characters, the species were grouped as
follows:
three major series (A, B and C)
16 groups (I-XVI)
62 clusters (x-62)
of one or more 'modal' species. Furthermore, within the very large group IX,
the clusters are grouped into five subgroups (a-e) of which a, the largest, is
subdivided into six minor series (i-vi).
A. Anther-collars balusterform, with enlarged basal cells, rarely sub-
balusterform with indistinctly differentiated basal cells; stigmatic surfaces
usually narrow and widely separated, less often broad, contiguous or con-
fluent; endothecial cell thickenings radial, very rarely polar, cells usually
elongated; basic chromosome numbers 20, 10, 23, 30, 45, 50, 5, 52 (16, 18,
19, 22, 46, 60, 69, 80, >8o also recorded).
I. Frutescent; leaves pinnately veined, glabrous; inflorescence terminal
corymboid cymes; capitula radiate, dull yellow, rays 5-veined, glabrous;
cypselas glabrous, cells rather short, smooth; pappus-hairs uniform, stout,
tapered, 3-4 cells across below the apex; style-arm papillae rather long, a

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52 KEW BULLETIN VOL. 32(I)

median fascicle of longer papillae present; anther-bases very short, obtuse;

anther-collars cylindrical, cells uniform.
Australasian species, forming SenecioGroup III of Drury (1973); it does
not fit into any of his three assemblages and its isolated position is emphasized
by the present study.
A count, n = 19, is recorded for this species (Drury, personal communica-
tion).
I. S. insularis Benth.
II. Herbaceous, perennial, sometimes subscapose, or variously woody,
often scandent; leaves pinnately to palmately veined; inflorescences
compound terminal and upper axillary thyrses or terminal and axillary
corymbiform thyrses or cymes, often dense; capitula radiate or discoid, rays
yellow, disc yellow or whitish; rays various, o-4-veined, glabrous or with
uniseriate or biseriate hairs; cypselas glabrous or with short, obtuse hairs,
cells elongated, smooth or striate; pappus uniform, hairs moderately stout,
tapered; style-arm papillae medium to long, a median fascicle often present;
anther-collars balusterform, the basal cells often little differentiated; anther-
bases long, acute, obtuse or lobed, the length of the anther-collars.
Eastern Asian, Indian, Madagascan ---I and tropical African species; some of
the Asian and all the Afro-Madagascan species are climbers of distinctive
habit, corresponding to the genus CisampelopsisMiq. and resembling
MikaniopsisMilne-Redh. in this respect. S. wallichii, S. acuminatus,S. alatus
and S. corymbosus are referred by Hooker (1881) to Seneciosect. Synotis,and
also by Koyama (1969), who divides them amongst ser. Erectae (S.
ser. Synotis(S. wallichii,S. alatus) and ser. Scandentes
acuminatus,S. densiflorus),
(S. corymbosus).S. mikanioidesis referred to Seneciosect. Scandentesby Chater &
Walters (1976) and Harvey (1865) and (with S. rectiramus)by Muschler
(1909). S. leucopappus represents 'Group XIII' of Humbert (1963), S.
mikanioidesis referred to Seneciosect. Delairia by Cabrera (1957).
A count giving n = Io is recorded for S. mikanioides.
2. S. wallichiiDC.; S. acuminatus DC.; S. alatusDC.; S. densiflorus DC.; S.
corymbosus DC. (S. araneosusDC.); S. rectiramusBak.; S. leucopappus DC.; S.
mikanioidesWalp.
III. Herbaceous or softly woody, somewhat pachycaul, subsucculent,
erect or sarmentose; leaves pinnately or pinnatopalmately veined; in-
florescences dense terminal or terminal and axillary compound corymbiform
cymes; capitula discoid, yellow or pale yellow; cypselas glabrous or with
scattered hairs, cells rather short, smooth or striate; pappus uniform, hairs
moderately stout to moderately slender, tapered or subclavate; style-arm
papillae medium to long, a median fascicle sometimes present; anther-
collars subcylindrical, elongated, basal cells sometimes little differentiated;
anther-bases short, obtuse to acute, up to I the length of anther-collars.
Tropical African species, all referred to Seneciosect. Scandentesby Muschler
(1909); they approach in overall morphology certain other species referred
at present to the genus Crassocephalum Moench.
3. S. stuhlmanniiKlatt. Cypselas hairy; pappus-hairs subclavate, moder-
ately stout; style-arms with median fascicle of papillae.
4. S. syringifoliusO. Hoffm.; S. gigas Vatke. Cypselas glabrous; pappus-
hairs moderately slender, tapered; style-arms without median fascicle of
papillae.

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GENERIC AND SECTIONAL LIMITS IN SENECIO 53
IV. Herbaceous, perennial or annual, subsucculent; leaves pinnately
veined, often spathulate; inflorescences terminal corymbose cymes, or
capitula solitary; capitula radiate or discoid, rays and disc yellow; rays 4-
veined, with biseriate clavate hairs; cypselas with short, obtuse, spirally
thickened hairs, cells moderately elongated to short, smooth; pappus
uniform, hairs slender to very slender, tapered; style-arm papillae medium
to very long; anther-collars balusterform; anther-bases short, acute.
Tropical African species, representing Seneciosect. Spathulifoliiof Muschler
(1909). In overall morphology and ecalyculate involucres, resemble species
referred to the genera Emilia Cass. and OthonnaL.
Counts giving n = 5 are recorded for both S. hockiiand S. abyssinicus;this
number is known within the Senecioneaeotherwise only for some annual
Emilia species.
5. S. hockiiDe Wild. & Muschl.; S. abyssinicus A. Rich.

V. Herbaceous, perennial; leaves pinnately veined; inflorescences

terminal and upper axillary corymbose cymes; capitula radiate, rays and
disc yellow; rays 7-veined, glabrous; cypselas glabrous, cells elongated;
pappus uniform, hairs rather stout, tapered; style-arm papillae very long,
median fascicle present; anther-collars balusterform, basal cells little
differentiated; anther-bases short, obtuse.
Brazilian species; represents Seneciosect. Dichroaof Cabrera
(1957).
6. S. organensisCasar.

VI. Woody, arborescent, frutescent or suffrutescent, often scandent;

leaves pinnately veined; inflorescences terminal, terminal and upper
axillary or axillary thyrses, often copious; capitula radiate or discoid, rays
and disc yellow; rays 4-7-veined, glabrous or with uniseriate or biseriate
hairs; cypselas glabrous, cells elongated, smooth or striate; pappus uniform,
hairs moderately stout to moderately slender, tapered; style-arm papillae
long, acute to obtuse; anther-collars subcylindrical to balusterform, basal
cells little to distinctly differentiated; anther-bases long, acute, 2-I1 the
length of the anther-collars, rarely short, obtuse.
Tropical Central and South American species; they represent Senecio
sect. Streptothamniof Greenman (1902) (S. parasiticus) and Cabrera (1957)
(S. desiderabilis),Senecio sect. Myriocephalusof Cabrera (1957) (S. pellu-
cidinervius,S. brachycodon,
S. glaziovii) and Seneciosect. Cacaliastrum
of Cabrera
(1957) (S. grandis).
A count giving n = 45-50 is recorded for S. desiderabilis.
7. S. parasiticusHemsl.; S. desiderabilisVell.; S. pellucidinerviusBak.; S.
brachycodonBak. Suffrutices, often scandent; leaves all exauriculate; capitula
radiate; anther-bases long, acute.
8. S. grandis Gardn. Frutices, erect; upper leaves sessile, auriculate;
capitula discoid; anther-bases long, acute.
9g.S. glaziovii Bak. Trees; leaves all petiolate, exauriculate; capitula
discoid; anther-bases short, obtuse.

VII. Suffrutescent, scandent; leaves pinnately veined; inflorescences

terminal corymbose cymes; capitula radiate, rays orange-red, disc orange-
yellow; rays about 4-veined, with uniseriate capitate hairs; cypselas with
long, finely striate subobtuse hairs, cells short, undulate-papillose; pappus

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54 KEW BULLETIN VOL. 32(I)

uniform, hairs moderately stout, tapered; style-arm papillae rather long,

inserted on expanded rhombic apex of style-arms; anther-collars baluster-
form, basal cells little differentiated; anther-tails very short, obtuse.
Tropical Central and South American species; represents Seneciosect.
Convolvuloideiof Greenman (1902), later segregated as a distinct genus
Pseudogynoxys (Greenm.) Cabrera (1950).
A count giving n = 45 is recorded for S. berlandieri.
xo. S. berlandieri(DC.) Hemsl.
VIII. Herbaceous, somewhat suffrutescent or pachycaul; leaves pinnately
veined; inflorescences copious terminal corymbose compound cymes;
capitula radiate; rays 5-veined, glabrous; cypselas glabrous, cells short,
undulate-papillose; pappus uniform, hairs moderately stout, tapered; style-
arm papillae long, acute, median fascicle present; anther-collars baluster-
form, elongated, basal cells little differentiated; anther-bases very short,
obtuse.
Tropical Central American species, representing Seneciosect. Multinervii
of Greenman (I902).
A count of n = c. 50 is recorded for a specimen referred to the affinity
of S. multivenius.
1I. S. multiveniusBenth.
IX. Herbaceous, perennial or annual, often subscapose, suffrutescent or
frutescent, sometimes succulent, sometimes scandent, rarely pachycaul
arborescent; leaves pinnately, pinnatopalmately or palmately veined;
inflorescences usually terminal corymbose cymes, sometimes thyrsoid or
capitula solitary, terminal; capitula radiate, subradiate or discoid, rays
yellow, orange-yellow, purplish or rarely white, disc yellow, orange-red,
purplish or brownish; rays 3-Io-veined, glabrous or with uniseriate or
biseriate hairs; cypselas glabrous or with usually spirally thickened obtuse
hairs; cells short to long, smooth or striate, often papillose; pappus uniform
or dimorphic, hairs stout to very slender, tapered or tapered and fluked;
style-arm papillae medium, median fascicle usually absent; anther-collars
nearly always balusterform with clearly differentiated basal cells; anther-
bases short, rarely as long as the anther-collars, obtuse to acute.
a. Suffrutices or perennial herbs, usually leafy, leaves more or less
pinnately divided or lobed; rays and disc usually yellow; cypselas with
usually smooth cells; pappus uniform, stout to slender, tapered, flexuous,
white, present in ray-florets.
i. Perennial rhizomatous usually leafy herbs or dwarf suffrutescent herbs
with yellow or orange-yellow radiate capitula; rays 4(-8)-veined, glabrous
or with usually uniseriate hairs, cypselas with smooth cells.
Eurasian species, variously referred as follows:
to Cacalia: S. arachnanthus by Hu (1966); to Ligulariasubgen. Dolichorhiza;
S. renifoliusby Pojarkova (1961); to Seneciosect. Reniformes:S. renifoliusby
Boissier (1875); to Seneciosect. Amplectentes: S. pseudoarnicaby Greenman
(1902); to Seneciosect. Quadridentati:S. taraxacifoliusby Schischkin (1961) and
Boissier (1875); to Seneciosect. Oliganthi:S. othonnaeby Schischkin (1961)
and Boissier (1875); to Seneciosect. Jacobaea:S. morrisonensis
by Kitamura
(1942) and Koyama (1969), S. erucifoliusand S.jacobaea by Chater & Walters
(1976), Boissier (1875) and Schischkin (ser. Erucifolii) (1961), S. bicolor and

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GENERIC AND SECTIONAL LIMITS IN SENECIO 55
S. trapezuntinusby Boissier (1875), and S. abrotanifoliusand S. adonidifoliusby
Chater & Walters (1976); to Seneciosect. Incani:S. pseudoarnica (ser. Arnicoidei)
by Schischkin (1961), S. bicoloralso by Greenman
and S. bicolor(ser. Cinerariae)
(1902) and with S. leucophyllus, S. incanus,S. ambiguusand S. gnaphalodes,by
Chater & Walters (1976); to Seneciosect. Carniolici:S. incanusby Schischkin
(I96I); to Seneciosect. Velutini:S. gnaphalodesby Boissier (1875); to Senecio
sect. Coriacei:S. auriculaby Muschler (1909); to Seneciosect. Crocoserides: S.
pseudoarnica by Kitamura (1942) and Koyama (1969); and to Senecio sect.
Doria: S. othonnaeand S. auriculaby Chater & Walters (1976).
Counts giving the following numbers are recorded: n = 18, 20 for S.
pseudoarnica; n = 20 for S. morrisonensis;n = 20 for S. erucifolius; n = 20,
6o, - 8o for S. incanus; n = 20 for S. bicolor; n = 16, 20, 40 for S. jacobaea;
n = 20 for S. leucophyllus;n = 20, 30, 50 for S. abrotanifolius and n = 20 for
S. adonidifolius.
Seneciorenifoliusstands apart from the other species in its palmately veined,
reniform leaves, but is Senecio-like,not Ligularia-like, in endothecial cell
thickenings, style-arm papillae and pappus-hairs. The position of S.
trapezuntinusis uncertain, and it might be grouped with IX e 46.
12. S. renifolius(C. A. Mey.) Sch. Bip; S. taraxacifolius(M. Bieb.) DC.;
S. pseudoarnicaLess. Perennial rhizomatous herbs; leaf-hairs uniform or
subuniform; cypselas glabrous, cells elongated; rays 4-8-veined, glabrous or
with one-celled hairs; pappus-hairs stout to very stout, 2-4 cells across below
apex; anther-collars sub-balusterform, basal cells poorly differentiated.
13. S. arachnanthusFranch.; S. faberi Hemsl.; S. morrisonensis Hayata.
Perennial rhizomatous herbs; leaf-hairs non-uniform or glandular; cypselas
glabrous, cells elongated; rays 4-veined, glabrous or with uniseriate hairs;
pappus-hairs stout, 2 cells across below apex; anther-collars balusterform.
14. S. othonnaeM. Beib.; S. erucifoliusL. Perennial herbs; leaf-hairs
uniform; cypselas densely hairy, hairs long, slightly spirally thickened,
apices often emarginate, cells short to elongated; rays 4-veined, hairy;
anther-collars balusterform.
x5. S. incanusL.; S. ambiguus(Biv.) DC.; S. gnaphalodesSieb.; S. bicolor
(Willd.) Tod. (S. cinerariaDC.). Perennial herbs or dwarf suffrutices; leaf-
hairs non-uniform; cypselas glabrous or hairy, cells short to moderately
elongated; rays 4-veined, glabrous; pappus-hairs stout to moderately stout,
2-3 cells across below apex; anther-collars balusterform.
A6. S. trapezuntinusBoiss.; S. jacobaeaL.; S. leucophyllusDC.; S. auricula
Bourg. Perennial sometimes subscapose herbs; leaf-hairs non-uniform,
glandular, or absent; cypselas hairy, cells short to moderately elongated;
pappus-hairs rather slender, 2 cells across below apex; anther-collars
balusterform.
17. S. abrotanifoliusL.; S. adonidifoliusLoisel. Perennial herbs with 2-3-
pinnatisect glabrous leaves; cypselas glabrous, cells elongated; pappus-hairs
rather slender, 2 cells across below apex; anther-collars balusterform.

ii. Suffrutices, suffrutescent or perennial herbs, rarely succulent; leaves

sometimes reduced; rays and disc yellow, rays 2-5-veined, sometimes
reduced, often absent, glabrous or with biseriate or uniseriate hairs; cypselas
with smooth, usually elongated, cells; anther-collars balusterform.
African species, referred as follows: to Seneciosect. Pinifolii: S. pinifolius
by Harvey (1865) and Muschler (1909); Seneciosect. Aphylli: S. junceusby

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56 KEW BULLETIN VOL. 32(I)

Harvey (1865); Seneciosect. Leptolobi:S. achilleifoliusand S. bipinnatusby

Harvey (1865); Seneciosect. Kleinioidei: S. junceus by Muschler (1909); Senecio
sect. Emilianthoidei:S. bipinnatusand S. achilleifoliusby Muschler (1909);
Seneciosect. Microlobiby Harvey (1865) and Seneciosect. Griseiby Muschler
(1909): S. oliganthus;Seneciosect. Bethencourtia:S. palmensisby Hoffman
(1890); Seneciosect. Rigidi by Harvey (1865) and Seneciosect. Stenophylliby
Muschler (1909): S. pubigerusand S. rigidus; to Seneciosect. Tuberosiby
Muschler (1909): S. tuberosus;and to Seneciosect. Montani by Muschler
(190o9): S. steudelii.
z8. S. pinifolius (L.) Lam.; S. junceus(Less.) Harv.; S. achilleifoliusDC.;
S. bipinnatus(Thunb.) Less. Suffrutescent, or leafless succulent herbs; leaves
glabrous; cypselas glabrous, cells elongated; rays 4-5-veined, glabrous or
absent; anther-bases acute, 1 to 1 the length of anther-collars.
xg. S. oliganthusDC. Suffrutices; leaf-hairsnon-uniform; cypselas glabrous,
cells elongated; rays absent; anther-bases equalling the anther-collars, acute.
20. S. palmensis (Nees) DC. Shrubby; leaves glabrous; cypselas hairy,
cells moderately elongated, walls prominent; rays reduced, 2-3-veined,
glabrous; anther-bases equalling the anther-collars, acute.
21. S. pubigerusL.; S. rigidusL.; S. diphyllusDe Wild. & Muschl. Suffrutes-
cent or perennial, sometimes tuberous herbs; leaves hairy, hairs non-
uniform, with stout tapered base and short to long narrower apical cell;
cypselas densely hairy, hairs finely thickened; cells short, walls prominent;
rays 4-veined, glabrous or with biseriate hairs, or absent; anther-bases
short, up to 3 the length of anther-collar, acute.
22. S. tuberosusA. Rich. Perennial tuberous herbs; leaves glabrous;
cypselas hairy, hairs long, finely thickened, obtuse to emarginate at apex,
cells elongated, walls prominent; rays absent; anther-bases short, obtuse.
23. S. steudeliiA. Rich. Perennial leafy herbs; leaves hairy, hairs non-
uniform, with 1-4-celled base and elongated apical cell; cypselas glabrous,
cells short, smooth, with prominent walls; rays 4-veined, with small uni-
seriate hairs; anther-bases short, subacute.

iii. Suffrutices, perennial or biennial herbs, sometimes subscapose; leaves

lobed or entire, glabrous or with non-uniform hairs; rays and disc yellow,
rays 4-veined, venation usually open, glabrous or usually with short bi-
seriate or uniseriate hairs; cypselas glabrous or hairy, cells moderately
elongated to fairly short, smooth; pappus moderately stout, 2-celled below
apex; anther-collars balusterform; anther-bases short to medium, acute or
obtuse.
Indian species; S. lavandulifoliusis referred to Seneciosect. Pseudojacobaea
by
Hooker (1881), the others by him to Seneciosect. Jacobaea.
A count giving n = 20 is recorded for S. neelgherryanus.
24. S. obtusatusDC.; S. lavandulifolius DC.; S. griffithiiHook. f. & Thoms.;
S. neelgherryanusDC.

iv. Suffrutices or leafy perennial herbs; leaves lobed to entire, hairs non-
uniform or uniform; rays and disc yellow, rays 4-5-veined, with short
biseriate or uniseriate hairs; cypselas hairy, cells short, smooth or sub-
papillose; pappus-hairs slender, 2-celled below apex; anther-collars baluster-
form; anther-bases short, acute.

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GENERIC AND SECTIONAL LIMITS IN SENECIO 57
North American species, referred by Greenman (1902) as follows: to
Seneciosect. Suffruticosi:S. douglasii,S. flaccidus;to Seneciosect. Eremophili:S.
eremophilus;to Seneciosect. Incani:S. palmeri.
Counts giving n = 10, 2o are recorded for S. douglasii.
25. S. douglasiiDC.; S. eremophilus
Richards.; S. palmeriGray; S. flaccidus
Less.

v. Dwarf suffrutices or perennial rhizomatous herbs, rarely subscapose;

capitula radiate, subradiate or discoid, disc yellow, rarely orange-red or
brownish, rays yellow or orange-yellow, rarely white, 3-6-veined, with
usually short uniseriate glandular hairs, rarely also with eglandular hairs;
cypselas glabrous or hairy, cells elongated to short, smooth or rarely papillose;
pappus-hairs moderately stout, 2-celled below apex, uniform or rarely
subdimorphic (some hairs with reduced teeth); anther-collars balusterform,
often long; anther bases rather short, up to 1 (1) the length of anther-collar,
obtuse to acute; style-arm papillae medium, rarely with a median fascicle.
South American species, referred by Cabrera (I948, 1957) to the following
sections of Senecio:Haplostichia:S. arnicoides;Corymbocephalus: S. glabratus
(subsect. Glabrati),S. conyzifolius(subsect. Viscosi); Columbaria: S. calocephalus;
Wernerioides: S. wernerioides; Leucanthemifolium:S. werdermannii; Metazanthus:
S. subdiscoideus; Chenopodioides: S. amphibolus;Andina:S. santelicis;Suffrutecius:
S. tricuspidatus(subsect. Caespitosi),S. darwinii(subsect. Subincani); Oreophyton:
S. subulatus; Graveolens:S. graveolens;Xerosenecio:S. patagonicus(subsect.
Filaginoidei); Otites:S. otites; Websteria:S. websteri;Paranaia:S. paranensis;and
Acanthifolium: S. acanthifolius.
Counts giving the following numbers are recorded: n = 20 for S.
amphibolus; n = 52 for S. calocephalus; n = 20 for S. websteri. The n = 52
count suggests a possible affinity with the genus Werneria.
Senecioacanthifolius
stands apart from the other species in its white rays and
median fascicle of style-arm papillae. S. conyzifoliusalso is somewhat
anomalous in its brownish corollas and filiform female outer florets.
26. S. arnicoidesHook. & Arn. (S. plantagineusColla); S. glabratusHook. &
Arn.; S. calocephalus Poepp.; S. wernerioidesWedd.; S. werdermannii Greenm.;
S. subdiscoideus Wedd. (S. purpuratusPhil.); S. amphibolus Wedd.; S. santelicis
Phil.; S. tricuspidatusHook. & Arn. (S. longipesHook. f.); S. subulatusHook.
& Arn.; S. graveolensWedd. Dwarf suffrutices, sometimes rather succulent,
or perennial sometimes subscapose herbs; capitula sometimes solitary; disc
yellow or rarely orange-yellow; rays yellow or rarely orange-red, 3-6-veined,
reduced and evascular, or absent, glabrous or with short capitate hairs;
cypselas glabrous or hairy, cells smooth; anther-bases short.
27. S. patagonicusHook. & Arn.; S. darwinii Hook. & Arn. Perennial
suffrutescent herbs; leaf-hairs non-uniform; capitula sometimes solitary;
rays and disc yellow, rays short, 4-veined, with short capitate hairs, or absent;
cypselas glabrous, cells strongly papillose; anther-bases short, obtuse.
28. S. otites DC. Perennial rhizomatous herbs; leaf-hairs uniform; rays
and disc bright yellow; rays 4-veined, with uniseriate hairs; cypselas hairy,
with moderately elongated smooth cells; anther-bases about ? the length of
anther-collar, acute.
29ag.S. websteriHook. f. Perennial herbs; leaves palmatopinnately veined;
leaf-hairs non-uniform; rays 3-veined, with uniseriate capitate and biseriate

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58 KEW BULLETIN VOL. 32(I)
clavate hairs; cypselas glabrous, cells elongated, smooth; anther-bases about
* the length of anther-collar, acute.
30. S. paranensisMalme. Perennial herbs; leaf-hairs non-uniform; rays
and disc yellow; rays 5-veined, with capitate and eglandular hairs; cypselas
glabrous, cells elongated, smooth; anther-bases I the length of anther-collar,
subacute.
3x. S. conyzifoliusBak. Perennial herbs; leaves sessile, auriculate, hairs
uniform; capitula disciform, brownish, marginal florets tubular, without
rays, with uniseriate and eglandular hairs; cypselas glabrous, cells
moderately elongated, smooth; anther-bases very short, obtuse.
32. S. acanthifoliusHombr. & Jacq. Perennial rhizomatous herbs; rays
4-veined, with uniseriate hairs, white; disc yellow; cypselas glabrous, cells
elongated, smooth; style-arm papillae medium, obtuse, a median fascicle
of much longer acute papillae also present; anther-bases very short, obtuse.
vi. Suffrutices or suffrutescent herbs, often succulent, or perennial
herbs, rarely annual succulent herbs; leaves pinnately or parallelo-
pinnately veined, leaves glabrous or with non-uniform hairs; rays and disc
yellow, rays 4-veined, with biseriate hairs or glabrous, or rays absent;
cypselas hairy, cells striate, subpapillose, rarely smooth; anther-bases up to
1 the length of anther-collar, acute; style-arm papillae various, rarely with
a median fascicle.
African, Madagascan and Australasian species, referred as follows: to
Seneciosect. Rigidi: S. arniciflorusby Harvey (1865); to Seneciosect. Grisei:
S. arniciflorusby Muschler (19o09); to Seneciosect. Kleinioidei and sect.
Orientalesby Muschler (1909) and to his groups 16 and 17 by Humbert
(1963) respectively: S. melastomifoliusand S. hildebrandtii.S. quadridentatus
represents the 'erechthitoid Senecios' of Belcher (1956).
Counts giving n = 2o are recorded for S. hildebrandtii and S. gregorii.
Seneciogregoriistands apart from the other species in its annual habit,
ecalyculate involucres with fused bracts, median fascicle of stigmatic papillae,
and in features of the pappus and cypsela surface. S. hypoleucus also is some-
what anomalous in its smooth cypsela cells.
33. S. arniciflorusDC., S. melastomifolius Bak.; S. hildebrandtiiBak.; S.
gregoriiF. Muell.; S. quadridentatus Labill.; S. hypoleucusBenth.
b. Perennial or annual leafy herbs; rays and disc yellow; cypselas with
usually striate cells; pappus uniform, stout to slender, tapered, rigid, often
reddish, absent in ray-florets.
Indian species, referred to Seneciosect. Madaractisby Hooker (1881) and
(S. saxatilis) by Koyama (1969).
34- S. saxatilis DC. Perennial herbs; leaf-hairs uniform; rays 4-veined,
glabrous; cypselas glabrous, cells moderately elongated, smooth.
35. S. edgeworthiiHook. f.; S. grahamiiHook. f. Perennial or annual herbs;
leaf-hairs non-uniform; rays 4-veined, with biseriate hairs; cypselas hairy,
hairs obtuse or apiculate, cells moderately elongate, striate.
c. Perennial sometimes subscapose or often annual herbs, less often
suffrutescent, rarely scandent, usually leafy, leaves usually pinnately
divided and lobed, often sessile and auriculate; rays and disc yellow or
purplish; cypselas with papillose, often striate, cells, usually hairy; pappus-
hairs stout to very slender, tapered, flexuous, uniform or dimorphic (some
hairs persistent, fluked), white, present in ray-florets.

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GENERIC AND SECTIONAL LIMITS IN SENECIO 59
Mostly southern hemisphere species, a few in eastern Asia and some,
mostly annuals, in Eurasia and North America, variously referred to the
following sections of Senecio:S. scandensto Synotisby Hooker (188I1),Kitamura
(1942) and Koyama (1969) (ser. Scandentes);S. dumetorum to Adamantinaby
Cabrera (1957); S. pohlii to Corymbocephalus subsect. Simplices and S.
brasiliensisto subsect. Brasilensesby Cabrera (1957); S. linifoliusto Leptophylli
by Chater & Walters (1976) and to Fruticulosiby Muschler (19o09); S.
inaequidens,S. erubescens and S. cinerascens respectively to Leptophylli,Sinuosi
and Rigidi by Harvey (1865) and to Stenophylli,Viscosiand Griseiby Muschler
(1909); to sect. Senecio(syn. Annui, Obaejaceae):S. subdentatus by Schischkin
(1961) (ser. Subdentati), S. viscosus by Greenman (1902), Chater & Walters
(1976), Schischkin (1961) (ser. Viscosi)and Boissier (1875), S. gallicus by
Chater & Walters (1976) and Muschler (I909), S. californicusby Greenman
(1902), S. vulgaris by Greenman (1902), Cabrera (1948, 1957), Chater &
Walters (1976), Harvey (1865), Schischkin (1961) (ser. Vulgares), Boissier
(1875) and Muschler (I909), S. vernalis by Chater & Walters (1976),
Schischkin (I961) (ser. Vernales)and Boissier (1875), S. sylvaticus by
Greenman (1902), Cabrera (I948), Chater & Walters (1976), Schischkin
(1961) (ser. Sylvatici), Boissier (1875) and Muschler (o1909), S. desfontaineiby
Hu (1968), and S. littoreusby Harvey (1865) and Muschler (1909); S. littoreus
is referred to his 'Group 5' by Humbert (1963), S. erechtitoidesto his 'Group
Io' and by Muschler (I909) to sect. Montani; to sect. Jacobaea are referred S.
and S. ramosusby Hooker (1881) and S. delphiniifolius
desfontainei by Muschler
is referred to sect. Delphiniifoliusby Chater & Walters
(1909); S. delphiniifolius
(1976).
Counts giving the following numbers are recorded: n = Io for S. scandens;
n = 20 for S. linifolius; n = 30 for S. subdentatus; n = 20 for S. viscosus;
n = Io for S. gallicus; n = 19, 20 for S. vulgaris; n = 20 for S. vernalis, S.
sylvaticus,S. desfontaineiand S. delphiniifolius.
36. S. scandensD. Don; S. stauntoniiDC. Perennial erect or scandent herbs;
leaf-hairs uniform; rays and disc yellow; rays 4-6-veined, with uniseriate
and sometimes biseriate hairs; cypselas with moderately long papillose cells;
pappus uniform, cells short to moderately elongated.
37. S. dumetorumGardn.; S. pohlii Bak.; S. brasiliensis(Spreng.) Cass.
Perennial or biennial sometimes subscapose herbs; leaf-hairs uniform or non-
uniform; rays and disc yellow, rays 5-7-veined, hairy, or rays absent and
corollas magenta; cypselas with moderately long, almost smooth papillose
or strongly striate subpapillose cells; pappus uniform or dimorphic.
38. S. linifolius L. Suffrutices; leaf-hairs uniform; rays and disc yellow,
rays 5-veined, with biseriate hairs; cypselas with smooth papillose cells;
pappus uniform.
39. S. erubescens Ait.; S. inaequidens DC.; S. cinerascensAit. Suffrutices or
perennial to annual herbs, sometimes subscapose; leaf-hairs non-uniform;
rays and disc yellow, rays 4-veined, with short biseriate hairs, or rays absent
and corollas purplish; cypselas with moderately elongated to short, smooth
or striate papillose cells; pappus uniform or dimorphic.
4o. S. subdentatusLedeb.; S. viscosusL.; S. gallicus Chaix; S. californicus
DC.; S. vulgarisL.; S. vernalisWaldst. & Kit.; S. sylvaticusL.; S. desfontainei
Druce; S. littoreusThunb.; S. erechtitoides Bak.; S. lautusWilld.; S. ramosus
DC. Annual herbs; leaf-hairs uniform; rays and disc yellow, rays usually
4-veined, glabrous or with short biseriate hairs, or rays absent; cypselas

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60 KEW BULLETIN VOL. 32(1)
with long hairs and moderately elongated to short, smooth or striate, papil-
lose cells; pappus uniform, subdimorphic or dimorphic.
S. delphiniifolius
Vahl. Annual herbs; leaf-hairs uniform; rays and disc
4x.
pale yellow, rays 4-veined, hairy; cypselas with short, subglobose hairs;
pappus uniform.
d. Perennial erect sometimes subsucculent or succulent scandent suffrut-
ices; leaves palmately or palmatopinnately veined, usually reniform,
sometimes peltate; rays and disc yellow or purplish; cypselas with elongated
to short, striate, sometimes papillose cells, glabrous or hairy; pappus uniform,
tapered, sometimes dimorphic (with fluked hairs) white, flexuous, present
in ray-florets.
Macaronesian, tropical and south African species. S. malvifoliusand S.
heritierihave generally been referred by most authors to Seneciosect. Pericallis,
including the former by Chater & Walters (1976). S. auriculatissimus by
Muschler (1909) and S. angulatus by Muschler (1909), Harvey (1865) and
Chater & Walters (1976) have been referred to Seneciosect. Scandentes,S.
oxyriifoliusto Seneciosect. Paucifolii by Harvey (1865) and Seneciosect.
Cinerariiphylli by Muschler (1909).
Counts giving n = 30 are recorded for S. heritieriand S. oxyriifoliusand
n = 30, 46 for S. angulatus.
42. S. malvifolius(L'Herit.) DC.; S. heritieriDC. Erect herbs; leaf-hairs
non-uniform; rays and disc purplish; rays 4-7-veined; pappus uniform.
43. S. auriculatissimusBritten; S. oxyriifoliusDC.; S. angulatusLinn. f.
Erect or scandent; leaf-hairs uniform, short, or absent; rays and disc yellow;
rays 4-veined, or absent; pappus uniform or dimorphic.
e. Perennial, often subscapose, less often suffrutescent or arborescent
pachycaul herbs; leaves pinnately veined, often undivided and toothed, the
upper often sessile, auriculate or amplexicaul; rays and disc yellow; cypselas
with elongated smooth cells, often glabrous; pappus uniform, hairs tapered,
white, present in ray-florets.
Madagascan, African, Eurasian, North, Central and South American
species, variously referred to the following sections of Senecio-Plantaginei:
S. lasiorhizusby Harvey (1865); Polyrhizi:S. lasiohizusby Muschler (1909);
Imbricati:S. karaguensisby Muschler (i9o9); Arborei:S.johnstoniiby Muschler
(1o909); Emilianthei: S. panduriformis and S. bupleuroidesby Muschler (19o9);
Rigidi: S. panduriformis
by Harvey (1865) ; Paucifolii:S. bupleuroides
by Harvey
(1865); Velutini:S. hypochionaeus
and S. ciliciusby Boissier (1875); Crocoserides:
S. paludosusby Boissier (1875) and Schischkin (1961) (ser. Paludosi), S.
orientalis by Boissier (1875) and Schischkin (1961) (ser. Subalpini), and S.
racemosusby Boissier (1875) and Schischkin (i961) (ser. Macrophylli); Doria:
S. paludosus,S. nemorensis,
S. doronicum,
S. doriaand S. resedifolius
by Chater &
Walters (1976) and S. doriiformis
by Boissier (1875); Jacobaea:S. nemorensis
by
Kitamura (1942) and S. chysanthemoides by Hooker (1881) and Muschler
(1909); Oliganthi: S. nemorensis by Boissier (1875); Pseudooliganthi: S.
nemorensisby Schischkin (1961); Montani: S. renardii by Schischkin (1961);
Hualtata:S. pulcherand S. hualtataby Cabrera (I957); Aurei:S. toluccanus, S.
aureus and S. resedifoliusby Greenman (1902); Scaposi: S. resedifoliusby
Schischkin (1961); Columbiani: S. triangularis by Greenman (19o02); Lobati:
S. eurycephalus
and S. neomexicanus
by Greenman (1902); Mulgediifolii: S.

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GENERIC AND SECTIONAL LIMITS IN SENECIO 61
decorusby Greenman (1902); Amplectentes:S. pudicus by Greenman (1902);
and Tomentosi:S. bellidifoliusand S. tomentosus
by Greenman (1902).
Counts giving the following numbers are recorded: n = 10, 40 for S.
johnstonii; n = 20 for S. bupleuroides;n = 20 for S. hypochionaeus,S. racemosus;
n = 20, 30 for S. orientalis; n = 20, 40 for S. doronicum; n = 20 for S.
nemorensisand S. chrysanthemoides;
n = 20 for S. hualtataand S. toluccanus;
n = 22 for S. aureus; n = 23, 40, 46, 69 for S. resedifolius; n = 23 for S.
eurycephalus;n = 20 for S. bellidifolius; n = 20, 23 for S. neomexicanusand S.
tomentosus;n = 10, 20, 40 for S. triangularis; n = 20 for S. decorus.
The position of S. bupleuroides is uncertain; it could as well be placed, on
the present data, with IX a i x8. Likewise, S. renardiiand S. chrysanthemoides
might be grouped with IX a i x6.
44. S. lasiorhizusDC. (S. coronatusHarv.); S. karaguensis0. Hoffm.; S.
johnstonii Oliv. Perennial sometimes pachycaul tree-like herbs; leaf-hairs
non-uniform; cypselas glabrous or hairy; ray-florets 5-9-veined, glabrous
or hairy, sometimes with eglandular hairs.
45. S. gossypinusBak.; S. denisii H. Humb.; S. panduriformisHilliard (S.
pandurifoliusHarv.); S. bupleuroidesDC. Perennial suffrutescent or leafy
herbs; leaf-hairs non-uniform or absent; cypselas glabrous; ray-florets 4-6-
veined, glabrous or with uniseriate hairs, or rays absent.
46. S. hypochionaeus Boiss.; S. paludosusL.; S. racemosus(M. Bieb.) DC.; S.
orientalisWilld. (S. pseudoorientalis Schischk.); S. doronicumL.; S. doriaL.; S.
ciliciusBoiss.; S. nemorensis L.; S. doriiformisDC.; S. chrysanthemoides DC.; S.
renardiiWinkl. Perennial leafy or subscapose herbs; leaf-hairs uniform or
non-uniform; cypselas glabrous or hairy; ray-florets 3-Io-veined, with
uniseriate and/or biseriate hairs or glabrous.
47. S. pulcherHook. & Arn.; S. hualtataDC.; S. toluccanusDC.; S. aureus
L.; S. resedifoliusLess.; S. eurycephalus Torr. & Gray; S. bellidifoliusH.B.K.;
S. neomexicanus Gray; S. tomentosus Michx.; S. triangularisHook.; S. decorus
Greenm.; S. pudicusGreene (S. cernuusGray). Perennial often subscapose
herbs; leaf-hairs non-uniform, rarely uniform or absent; cypselas glabrous or
hairy; ray-florets 4-7-veined, glabrous or with sub-biseriate hairs, or rays
absent.

B. Anther-collars cylindrical, without enlarged basal cells; stigmatic

surfaces broad, confluent; endothecial cell thickenings polar or radial, cells
short; involucres ecalyculate; basic chromosome number 24 (13, 25 also
recorded).
X. Perennial rhizomatous or biennial herbs; leaves pinnately veined,
toothed; leaf-hairs non-uniform or uniform; inflorescence terminal, often
subumbelliform, corymbose cyme; capitula radiate, rays yellow, orange-
yellow, red or dark violet, disc yellow or purplish-yellow; rays 4-7-veined,
glabrous; cypselas glabrous or with long, acute, spirally-thickened hairs,
with short, smooth or striate cells; pappus-hairs stout, uniform, 2-3 cells
across below apex; style-arm papillae short; anther-bases short, up to 2

the length of anther-collar.

Holarctic species referred as follows: to Senecio sect. Tephroseris:S.
papposusand S. rivularisby Chater & Walters (1976) and Schischkin (1961)
(ser. Papposi and ser. Alpestresrespectively), S. pierotii and S. Jfammeusby
Kitamura (1942) and Koyama (1969), and S.Jfammeus by Schischkin (1961)

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62 KEW BULLETIN VOL. 32(I)
(ser. Porphyranthi);to Seneciosect. Eriopappus:S. congestusby Chater &
Walters (1976) and Schischkin (1961); to Senecio sect. Cinerarioidei: S.
congestusby Greenman (i9o2).
Counts giving the following numbers are reported: n = 24 for S. pierotii;
n = 13, 24 for S. flammeus; n = 24, 25 for S. congestus.
48. S. papposus(Reichenb.) Less.; S. rivularis(Waldst. & Kit.) DC. (S.
crispatusDC.); S. pierotiiMiq.; S. flammeusTurcz.; S. congestus(R.Br.) DC.
(S. palustris(L.) DC., S. arcticusRupr.).
XI. Perennial rhizomatous herbs; leaves pinnately veined and lobed;
leaf-hairs non-uniform; inflorescence terminal, subumbelliform corymbose
cyme; capitula radiate, rays 3-4-veined, with short biseriate hairs; rays and
disc pale yellow; cypselas glabrous, cells elongated, obscurely striate,
papillose; pappus-hairs stout, 3-celled across below apex, uniform; style-arm
papilllae short; anther-bases short, acute.
Eastern Asian species; referred to Seneciosect. Nemosenecioby Kitamura
(1942) and Koyama (1969).
Counts of n = I0o,20, 24 are recorded for S. nikoensis;according to Koyama
(1969), the counts of Io and 20oare erroneous.
49. S. nikoensisMiq.
C. Anther-collars cylindrical, without enlarged basal cells, rarely sub-
balusterform with indistinctly differentiated basal cells; stigmatic surfaces
usually broad, contiguous or confluent, rarely narrow and widely separated;
endothecial cell thickenings polar or radial, cells short; basic chromosome
numbers 30, 26 (27 also recorded).

XII. Perennial herbs; leaves palmately or pinnatopalmately veined;

inflorescences terminal corymbose cymes, paniculoid or racemiform
thyrses; capitula radiate, rays and disc yellow, or discoid, whitish or reddish,
rarely yellow; rays 4-5-veined, glabrous, or absent; cypselas glabrous, cells
elongated, smooth or striate; pappus-hairs tapered, uniform, stout, 2-4 cells
across below apex; stigmatic papillae medium to long; anther-bases short to
long, up to I4 the length of anther-collar; anther-collars cylindrical or sub-
balusterform, basal cells uniform or slightly enlarged.
Eastern Asian species, variously referred as follows; S. kramerito Cacalia
by Hu (1966) and Syneilesis by Kitamura (1942), Koyama (1969) and
Robinson & Brettell (1973a); C. peltifolia to Cacalia by Hu (1966); S.
makineanusto Miricalia by Kitamura (1942), Koyama (1969) and Robinson
& Brettell (1973a); S. zuccarinii to Cacalia by Hu (1966), Kitamura (1942)
(sect. Cacalia ser. Candidae), Koyama (1969) (sect. Delphiniifolia) and to
Koyamacaliaby Robinson & Brettell (I973a); C. kiusiana to Cacalia by
Kitamura (1969) (sect. Cacalia ser. Rufescentes) and Koyama (1969) (sect.
Delphiniifolia) and to Koyamacaliaby Robinson & Brettell (1973a); S.
monanthusto Cacaliaby Hu (1966), Kitamura (1942) and Koyama (1969)
(sect. Cacalia ser. Monanthae);S. cyclotusto Cacalia by Hu (1966) and to
Koyamacaliaby Robinson & Brettell (1973a); S. sagittatusto Cacaliaby Hu
(1966), Pojarkova (1961) (ser. Hastatae), Kitamura (1942) (sect. Cacalia ser.
Caudatae)and Koyama (I969) (sect. Cacalia)and to Koyamacalia by Robinson
& Brettell (1973a); S. tanguticusto Ligulariaby Hu (1966), to Cacaliasect.
Vaginalesby Koyama (1969) and to Sinacaliaby Robinson & Brettell (1973a);

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GENERIC AND SECTIONAL LIMITS IN SENECIO 63
S. przewalskiito Ligulariaby Hu (1966) ; and L. sibiricato Seneciosect. Ligularia
by Hooker (1881), to Ligularia by Chater (1976), Boissier (1875), Pojarkova
(1961) and Kitamura (1942) (subgen. Ligularia ser. Racemosae).
Counts giving the following numbers are recorded: n = 26 for S. krameri,
C. peltifolia, S. zuccarinii and S. kiusiana; n = 26, 27 for S. makineanus;n = 30
for S. sagittatus,S. tanguticus,S. przewalskiiand L. sibirica.
The species of this group correspond to the genera CacaliaL. (in the sense
of Asiatic authors, syn. KoyamacaliaRobins. & Brettell, including its segre-
gates SyneilesisMaxim. and Miricalia Kitamura) and LigulariaCass. (incl.
SinacaliaRobins. & Brettell), but also include some never referred other than
to Senecio.
50. S. krameriFr. & Say. (Syneilesispalmata (Thunb.) Maxim.); Cacalia
peltifolia Mak.; S. makineanus Yatabe; S. zuccariniiMaxim. (Cacaliadelphinii-
jolia Sieb. & Zucc.); Cacalia kiusianaMak.; S. monanthusDiels; S. cyclotus
Bur. & Franch.; S. sagittatusSch. Bip. (Cacaliahastata L.) Inflorescences
thyrsoid; capitula discoid, whitish or reddish, rarely yellow, usually
nodding; cypsela cells smooth; style-arm papillae long, apically fasciculate,
tending to be decurrent; leaf-hairs uniform or non-uniform; endothecial
cells thick-walled, thickenings polar.
51. S. maximowicziiC. Winkl.; S. tanguticusMaxim. (S. henryiHemsl.);
S. przewalskii Maxim.; Ligularia sibirica (L.) Cass. (S. ligularia Hook. f.).
Inflorescences usually thyrsoid; capitula radiate, rays and disc yellow;
cypsela cells smooth; style-arm papillae medium to long, tending to be
long-decurrent on to the style-arms and style; leaf-hairs uniform or absent;
endothecial cells short, thick-walled, thickenings polar.
52. S. winklerianusHand.-Mazz. (S. acerifolius C. Winkl.); S. trinervis
Chang. Inflorescences corymbiform; capitula radiate, rays and disc yellow;
cypsela cells striate; style-arm papillae short, obtuse, marginal; leaf-hairs
uniform or non-uniform; endothecial cells short, thick-walled, thickenings
polar.

XIII. Arborescent, woody; leaves pinnately veined; inflorescences dense

terminal corymbose cymes; capitula discoid, purplish; cypselas hairy, hairs
long, finely spirally thickened, subacute, cells elongated, smooth; pappus-
hairs tapered, uniform, stout, 2 cells across below apex; style-arm papillae
medium, obtuse; anther-bases about I the length of anther-collar; anther-
collars cylindrical, basal cells uniform.
Eastern Asian species, corresponding to the genus Dendrocacalia Nakai to
which it is referred by Kitamura (1942) and Koyama (1969).
Nakai.
53. S. crepidifolius

XIV. Herbaceous, perennial; leaves pinnately veined; inflorescences

terminal, corymbiform; capitula radiate, yellow; cypselas hairy, hairs long,
finely spirally thickened, acute-acuminate, cells elongated, smooth; pappus-
hairs tapered, uniform, rather slender; style-arm papillae short, obtuse;
anther-bases acute, about j the length of anther-collar; anther-collars
cylindrical, basal cells uniform.
Australasian species; for S. lyallii, a count of n = 30 is recorded.
54. S. lyallii Hook. f.

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64 KEW BULLETIN VOL. 32(I)

XV. Fruticose or arborescent; leaves pinnately veined; inflorescences

terminal corymbose or terminal and upper axillary paniculoid compound
cymes; capitula radiate, white; cypselas glabrous or with short obtuse hairs,
cells elongate, smooth or striate; pappus-hairs clavate, uniform, rather
stout; style-arm papillae very short; anther-bases lobed or acute, I to
equalling the length of the anther-collar; anther-collars cylindrical, basal
cells uniform.
Australasian species; S. myrianthoshas striate, hairy cypselas, short ray
florets, acute anther-bases, and leaves with non-uniform (T-shaped) hairs;
it corresponds to Senecio Group I of Drury (1973) and to the genus
BrachyglottisJ. R. & G. Forst., to which it has been referred by him. S. kirkii
has smooth, glabrous cypselas, long ray florets, lobed anther-bases and
glabrous leaves; it corresponds to SenecioGroup 2 of Drury (1973), which,
with Group I, he includes in his Assemblage I.
A count giving n = 30 is recorded for S. kirkii.
55. S. myrianthosCheesem. (Brachyglottismyrianthos(Cheesem.) Drury);
S. kirkii Kirk.

XVI. Trees, shrubs, less often herbaceous; leaves pinnately or pinnato-

palmately veined; inflorescences terminal and upper axillary corymbiform
cymes or thyrses; capitula radiate, yellow; cypselas glabrous or with
spirally-thickened obtuse hairs, cells elongated, smooth or striate; pappus-
hairs tapered, subclavate or clavate, uniform, stout to slender, 2-3 cells
across below apex; style-arm papillae medium to short, obtuse; anther-bases
obtuse to acute, short or usually from J to equalling the length of the anther-
collar; anther-collars cylindrical or sub-balusterform, basal cells uniform or
slightly enlarged.
Australasian, Madagascan, South and Central American species. S.
hectorisand S. perdicioidesare included in his SenecioGroup VII by Drury
(I973), S. laxifoliusin his Group VIII; both are included in his Assemblage
III which, he states, is related to a group of herbaceous species including S.
lagopus.S. brunonisrepresents his Group V, which he also includes in his
Assemblage III. S. yegua and S. cymosusare referred by Cabrera (1948) to
Seneciosect. Myriocephalus, but are very distinct from the other species of that
section. S. hypargyreus and S. ripariusrepresent, respectively, SenecioGroups 2
and I of Humbert (1963). S. salignusis referred to Seneciosect. Fruticosiby
Greenman (1902) and is segregated as the monotypic genus Barkleyanthus by
Robinson & Brettell (I973b). To Seneciosect. Palmatinerviiare referred S.
petasitis, S. roldanaand S. hartwegiiby Greenman (1902), as is S. petasitisby
Chater & Walters (1976); S. barba-johannisis referred by Greenman (1902)
to Seneciosect. Fruticosi.By Robinson & Brettell (I973b), S. barba-johannis,
S.
petasitis, S. roldana and S. are
hartwegii referred to the genus Roldana,and S.
grandifolius to the genus Telanthophora.
Counts giving the following numbers are recorded: n = 30 for S.
perdicioides; n = 30 for S. salignus; n = 30 for S. petasitis; and n = 30, " 38
for S. grandifolius.
56. S. hectoris Buchan.; S. laxifolius Buchan.; S. lagopus Raoul; S.
perdicioidesHook. f. Trees, shrubs or herbs; leaves pinnately veined;
inflorescences corymbiform; cypselas glabrous, cells smooth; pappus-hairs
subclavate, tapered; ray-florets glabrous, rays 4-7-veined; style-arm

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GENERIC AND SECTIONAL LIMITS IN SENECIO 65
papillae very short; anther-collars cylindrical, cells uniform; anther-bases
acute, j-i the length of anther-collar.
57. S. brunonis(Hook. f.) J. H. Willis (S. centropappus F. Muell.). Arbores-
cent; leaves pinnately veined; inflorescencescorymbiform; cypselas glabrous,
cells smooth; pappus-hairs tapered, teeth elongated near apex; ray-florets
glabrous, rays 7-veined; style-arm papillae very short; anther-collars
cylindrical, cells uniform; anther-bases acute, about I the length of anther-
collar.
58. S. yegua (Colla) Cabr.; (S. denticulatus(Hook. & Arn.) DC.); S.
cymosusRemy. Arborescent or fruticose; leaves pinnately veined; inflores-
cences corymboid or thyrsoid; cypselas hairy, cells smooth or striate; pappus-
hairs tapered; ray-florets glabrous, rays 4-veined; style-arm papillae short;
anther-collars cylindrical, cells uniform; anther-bases subacute, J to * the
length of anther-collar.
59. S. hypargyreus DC.; Arborescent or fruticose; leaves pinnately veined;
inflorescences corymboid; cypselas hairy, cells striate; pappus-hairs tapered;
ray-florets 6-veined, with uniseriate clavate hairs, style-arm papillae short;
anther-collars sub-balusterform, cells uniform; anther-bases long, lobed,
equalling the anther-collars.
6o. S. riparius DC.; Shrubs; leaves pinnately veined; inflorescences
corymboid; cypselas glabrous, cells smooth; pappus-hairs clavate; ray-
florets 4-veined, glabrous; style-arm papillae very short; anther-collars sub-
balusterform, basal cells slightly enlarged; anther-bases acute, about ? the
length of anther-collar.
6x. S. salignusDC. (Barkleyanthus salicifolius(H. B. K.) Robins. & Brettell).
Shrubs; leaves pinnately veined; inflorescences thyrsoid; cypselas hairy,
cells strongly striate; pappus-hairs tapered; rays-florets4-6-veined, glabrous;
style-arm papillae rather short; anther-collars cylindrical, cells uniform;
anther-bases very short, subacute.
62. S. grandifoliusLess.; S. barba-johannis DC.; S. petasitis (Sims) DC.; S.
roldana DC.; S. hartwegii Benth. Shrubs or suffrutescent herbs; leaves
pinnately or pinnatopalmately veined, leaves ovate to orbicular, more or
less cordate; inflorescences corymboid, sometimes abruptly terminal;
cypselas glabrous or hairy, cells elongated, smooth or striate; pappus-hairs
subclavate or tapered; ray-florets 4-veined, glabrous or hairy; style-arm
papillae medium, often decurrent on to style-arms; anther-collars cylindrical,
cells uniform; anther-bases obtuse to acute, j to equalling the length of the
anther-collars.

CONCLUSIONS

The establishment of an internally consistent taxonomic scheme for

Senecioon a worldwide basis makes clear some hitherto overlooked anomalies.
It becomes obvious that the extent of variation is far too great to be accom-
modated within a single genus and indeed, that the variation-range amongst
species currently referred to Seneciooverlaps and even exceeds the combined
ranges exhibited by species currently referred to several other genera. The
existence of discontinuities in character-state syndromes, however, makes
taxonomic division of the complex possible, in spite of the absence of absolute
discontinuities in particular characters. It also becomes obvious that widely
different generic concepts are currently held, both in different parts of the
I-E

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66 KEW BULLETIN VOL. 32 (I)
world and in treatments of different groups of species within the complex.
Thus, while within group XII, no fewer than four distinct genera are
recognized by Japanese botanists, e.g. Kitamura (1942), Ohwi (1965) and
Koyama (1969), all members of groups I-VIII are currently included in
Senecio,notwithstanding that they differ from one another overall far more
than do the species included in group XII. Within group XVI, in
Australasia, Madagascar and South America, the species are commonly
accepted as belonging to Senecio,apart from a suggestion by Drury (1973)
that the Australasian species might be segregated as the genus Centropappus
Hook. f., whereas for the Central American species of the group, Robinson &
Brettell (I973b) accept no fewer than three distinct genera, Roldana,
Barkleyanthus and Telanthophora. Examples of such inconsistencies could be
repeated ad nauseam.
The evidence so far considered in the present study suggests that the name
Senecioshould be applied only to species of group IX, within which its type
species, S. vulgaris,falls, and which is, perhaps luckily from the nomenclatural
point of view, larger than all the other groups here recognized put together.
Serious consideration should be given to treating each of the sections I-
VIII and X-XVI, as distinct genera, either by establishing them as such or
by showing their species to be congeneric with other widely recognized
genera yet to be treated in this study. The three anomalous species of group
IX, S. renifolius,S. acanthifoliusand S. gregorii, likewise require further
exploration of their status and affinities. The grounds on which generic
distinctions withinany of these groups have been made should, on the other
hand, be the subject of critical re-investigation.
Above the level of the groups (tentative genera) here recognized, each of
the three major series might well be recognizable eventually at sub-tribal
level. In the other direction, within the still vast genus Senecio,as restricted
to group IX, it is clear there is need for a very thorough review of the
delimitation of taxa, of the allocation of species and of the nomenclature, at
sectional, subsectional and serial ranks. The widely different concepts of
such taxa held by different authorities, and their very various allocations
of species to them, have rendered them virtually meaningless.
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