Parsimony Analysis and Cladistic Reclassification of the Relhania Generic Group (Asteraceae-Gnaphalieae)

12 Pages·1991·9.1 MB·

by  Arne A Anderberg

AND
PARSIMONY
ANALYSIS
Arne Anderberg^ and Kare Bremer
A,
CLADISTIG
THE
RECLASSIFICATION OF
RELHANIA GROUP
GENERIC
(ASTERACEAE-
GNAPHALIEAE)
Abstract
Parsimony analysis of the Relhania group of genera (Asteraceae-Gnaphalieae-Relhaniinae) yields a revised
hypothesis of their phylogeny. The genera Leysera L. and Oedera L. are demonstrated to have their closest relatives
A
Relhania which shown
within L'Her., to be paraphyletic as hitherto circumscribed. revised generic classification
is
proposed. Relhania and Leysera are redefined more narrowly. Rhynchopsidium DC. and the monotypic Nestlera
is
Sprengel are rees'ablished. Oedera amended to include some former Relhania species. The new genus Comhorhiza
is
&
Anderb. Bremer, with 2 species, described, and 14 new combinations Oedera and Comborhiza are made.
in
is
Because parsimony analysis produces better phylogenetic hypotheses than limited selections of alleged synapomorphies,
common
and because the latter approach hitherto has been more in generic classification, argued that, with
it is
improved phylogenetic hypotheses derived from parsimony analysis, generic reclassifications will become necessary
also in other groups.
In this paper we present a parsimony analysis knowledge of the phylogeny oi the tribe Gnaphalie-
and propose a cladistic reclassification of a group ae as a whole (Anderberg, 1991). Furthermore,
1976-
of South African Asteraceae genera belonging to another genus, not considered in Bremer's
the tribe Gnaphalieae. The genera were revised 1978 revisions, has recently been added to the
&
during the 1970s and were then the subjects of group (Anderberg Kallersjo, 1988).
some of the cladistic approaches in botany Bremer (1976a, b, 1978a, b) revised the tax-
first
onomy
(Bremer, 1976a, 1978a, At that time, and nomenclature of the genera Relhania^
b, b).
uniquely derived, qualitative characters that could Rosenia, Leysera^ Oreoleysera, and Antlthrixia.
be interpreted as synapomorphies, unequivocally In trying to circumscribe correctly these genera
indicating moncphyletic groups were sought. Thus, as monophyletic groups, Bremer relied on partic-
selected feature^s, mainly of the pappus structure ular characters as synapomorphies defining them,
crowning the were used as putative synapo- Thus, Relhania^ with 29 species, was distinguished
fruits,
morphies defining the genera. Today, cladistics by pappus of more or less connate scales and
is its
methodologically much more sophisticated and no Bremer also pointed out the hypothet-
bristles.
is
also considered a standard technique for analyzing ical nature of the generic delimitation:
phylogenies (Hull, 1989). Application of the par-
simony criterion (Farris, 1983) in computerized Admittedly the loss of pappus bristles might have
programs has made possible to use even large occurred several times. However, since there is no
it
Relhania
evidence that should be polyphyletic in
its
and homoplastic data sets including kinds of
all
we must
present circumscription, believe for the
I
information, minimizing a priori assumptions. Par-
time being rely on this chi as uniquely derived
simony analysis of the Relhania group of genera and keeping the genus together (Bremer 1976a:
9).
yields a revised hypothesis of their phylogeny, and
Relhania was amended by Bremer
hence include
a basis for a cladistic reclassification. to
not only the theoretical and methodological most of the species of the genus Nestlera, which
It is
background to the study of the phylogeny that has prior to Bremer's work was used to house all the
changed during the past 15 years, but also the species with epaleate receptacles. Bremer showed
Department cf Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, S- 04 05 Stockholm,
^ 1
Sweden.
^Department of Systematic Botany, University of Uppsala, P.O. Box 541, S-75I 21 Uppsala, Sweden.
1061-1072.
Ann. Missouri Hot. Card. 1991.
78:
1062 Annals of the
Garden
Missouri Botanical
made
that epaleate receptacles were not uniquely derived group of the rest of the Relhania group an
from
and could not be used to separate Nestlcra analysis of llie entire /^(?//iart/a group at the species
much
Relhania. Thus, Bremer (1976b) placed of level conceivable. Based on the characters pre-
and transferred some species sented Tables and we have performed a
A^e5//cra in /?('//m/H*a in 2,
1
Rosenia. The genus comprises four species parshnony analysis of the recognized species of the
to latter
1-4
The
and has a pappus of scales as well as scabrid group, using Antithrixia as outgroup. analysis
and taxonomic
implications are presented below.
bristles. its
B (1978a) genus
Later, revised the related
METHODS
pappus and mostly
Lcyscra. has a of scales five
It
plumose and was assumed be monophy-
bristles to
DATA
due to a synapomorphy in the very long pe-
letic
Bremer commented on pappus The was based mainly on the characters
duncles. Again, the analysis
characters used distinguish the three genera: used by Bremer (1976a, b, 1978a, b) in his re-
to
The character has been amended with
visions.
list
In Leysera the disc-floret pappus consists of scales
by specimens
and sometimes fewer) Rosenia further characters investigating the
five (or bristles, in
it
1-2 Museum
consists of scales and generally bristles, and in housed the Swedish of Natural History
in
Relhania there a disc-floret pappus of scales only.
is The Table Only two
characters are
listed in 1.
(S).
In three genera there are only scales and no
all
have been Oede-
species not available for study, viz.
These
pappus. differences
bristles in the ray-floret
Oedcra Most
ra muirii and laevis. of the char-
are rather slight and the derived conditions might
conceivably have been reached more than once but acters of Oedera muirii were possible to extract
genera
there is no information Indicating that the from the protologue of this species (Smith, 1927),
(Bremer
should he polyphyletic or paraphyletic.
but Oedera laevis has been omitted. However,
1978a:
371).
judging from De Candolle's (1838) and Harvey's
Bremer (1978b) removed one former Ley-
also
(1865) descriptions, seems to be a typical rep-
it
pappus and
sera species with several bristles de-
Oedera.
resentative of
monotypic genus Oreoleyscra.
scribed as the
it
Another numotypic germs, Antithrixia^ was noted
ANAI YSIS
Relhania, Rosenia, Leysera,
as a close relative of
and Oreoleysera. The data matrix Table 2 was computed with
in
&
Recently, Anderberg Kallersjo (1988) dis- a parsimony program (Hennig86, version writ-
1 .5)
between Oedera and some
cussed the similarity ten by and obtained from Farris (1988), and
S.
J.
A
species of Relhania. transfer of Oedera, which using a standard PC-AT. The following options
comprised from Anthemidcac
six species, the to were used: multiple hennig (mhennig*, construct-
the Relhania generic group was proposed. In a ing cladograms by adding the taxa
al initial
review of the tribe Gnaphalieae, Anderberg (1991) of the character matrix several
in diflferent se-
recognized the subtribe Relhaniinae comprising two quences, retaining the shortest cladograin of each)
major clades, viz. the Metalasia group, with 14 with a subsequent branch-breaker command (bb*,
genera, and the Relhania group, with the genera generating the multiple, equally parsimonious
all
Relhania, Rosenia, Leysera, Oreoleysera, An- The
cladograms can be cladograms
that found).
and Oedera. Anderberg considered Oe-
tithrixia, were rooted with a hypothetical ancestor corre-
c/cra to be the sister group of /?rMaA?/a. However, sponding the outgroup, Antithrixia. The mul-
to
no synapomorphy could be found to define Red- tistate characters 13, 16, 29, and 45 were coded
hania if Oedera was kept separate. Infraspecific as additive (cc + because the character states were
)
relationships between the two genera were not in- hypothesized represent gradual transformation
to a
Relhania
vestigated, but the possibiHty that could The and on
characters 21, 23, 43, 46,
series. 8,
—
be paraphyletic was noted. the other hand, were coded as nonadditive (cc
),
Furthermore, Anderberg showed that the mono- since no such hypothesis could be formulated in
typic genus Antithrixia the most plesiomorphic
is these cases,
Relhania The
representative of the group. other
synapomorphy pappus
five genera share a in their
Results
and Metalasia
absent Antithrixia the
scales, in
The 200
group. (Bremer, (1978b) erroneously stated that analysis resulted in equally parsimo-
Orf'o/rj.srrc/ lacks pa[)pus scales, whicli, albeit very nious cladograms, each 123 steps long, with a
One
narrow, do Oreoleysera). consistency index of 0.45. of these clado-
exist hi (ci)
presence of a monotypic genus as the sister grams shown in Figure 1. The strict consensus
Is
'I'lie
&
Volume Number 4 Anderberg Bremer 1063
78,
and
1991 Analysis Reclassification of
Relhania
Table 1. Characters used in the analysis of the Rel- Table 1. Continued.
^^=^=^==^^^^^^=^^=^^^^^=^^^^^^=
hania-groxxp. Plesiomorphic states are coded and apo-
morphic states and 3 with ^n/tfAriJcia as outgroup. 29. Floret tubes eglandular or with few scattered
1, 2, (0)
See Methods for coding of multistate characters. glands, (1) floret tubes regularly glandular, (2) floret
===^=^=^=z:^^zz^:^^:z=:z=^=zzz=^=^^^^^=^^
tubes with subulatc-triangular hairs.
1. (0) Perennial half-shrubs or shrubs, annual herbs. 30. (0) Anther appendage acute to obtuse, (1) anther
(1)
Subterranean woody rhizome appendage truncate.
2. (0) absent, (1) subter-
ranean woody rhizome 31. Anther branched, anther tails un-
present. (0) tails (1)
Stem not subdichotomously branched, stem branched.
3. (0) (1)
subdichotomously branched. 32. Style branches in disc-florets apically penicillate,
(0)
branches
branches unarmed, branches subspinescent or truncate, (1) style in disc-florets dorsally
4. (0) (1)
and rounded-obtuse.
apically penicillate,
spinescent.
Cypselas cypselas
5. Brachybliists present, brachyblasts absent. 33. (0) oblong-elliptic, (1) linear.
(0) (1)
6. (0) Leaves decussate, (1) leaves alternate. 34. (0) Cypselas of ray-florets terete or angular, (1) cyp-
Leaves spreading, leaves squarrose-recurved. selas of ray-florets sharply triquetrous.
7. (0) (1)
Leaves linear-oblong, leaves obovate-spathu- 35. (0) Cypselas of disc-florets terete or angular, (1) cyp-
8. (0) (1)
late, (2) leaves widely cordate to orbicular. selas of disc-florets flattened.
9. (0) Leaf margin entire, smooth, (1) leaf margin ap- 36. (0) Cypselas glabrous, (1) cypselas at least in ray-
pubescent.
parently denticulate to sparsely serrate. florets
10. Leaves blunt, leaves pungent. 37. (0) Cypselas of pubescent ray-florets pilose, (1) cyp-
(0) (1)
11. Leaves mid-nerved, leaves with at least three selas of pubescent ray-florets villose.
(0) (1)
mam 38. Cypsela trichomes straight, cypsela trichomes
nerves. (0) (1)
12. Leaves adaxially more densely pubescent than apically coiled.
(0)
39. Cypselas generally eglandular, cypselas reg-
abaxially, leaves adaxially glabrous. (0) (1)
(1)
very glandular.
13. Leaves glandular-hairy with stalked glands, ularly
(0) (1)
40. Cypsela epidermis smooth, cypsela epidermis
leaves glandular-punctate, leaves glandular-punc- (0) (1)
(2)
sunken with acute papillae.
with glands
tate, in pits.
Leafmargin margin 41. Cypselas with 5 vascular bundles (occasionally
14. not involute, leaf involute. (0)
(0) (1)
2-3
cypselas with vascular bundles.
15. Capitula or on short peduncles (generally 10), (1)
(0) sessile
pappus
25 mm), capitula on long peduncles (generally 42. (0) Pappus bristles in ray -florets present, (1)
(1)
25 mm). bristles in ray-florets absent.
Pappus numerous, pap-
16. (0) Capitula solitary, (1) capitula paired to cymose- 43. (0) bristles in disc-florets (1)
pappus
corymbose or congested, capitula many in dense pus bristles in disc-florets five, (2) bristles in
(2)
secondary hfsads. disc-florets one to four, (3) pappus, bristles in disc-
17. Synflorescense not surrounded by a leafy invo- florets absent.
(0)
synflorescense surrounded by a leafy 44. (0) Pappus bristles apically scabrid to barbellate, (1)
lucre, in-
(1)
pappus plumose.
bristles apically
volucre.
Pappus pappus connate,
18. (0) Involucral bracts with spreading limb, (1) invo- 45. (0) scales free, (1) scales
pappus
tubular.
lucral bracts with straight limb. (2)
= -
some 46. Basic chromosome number x 7 (2n 14, 28,
19. (0) At least involucral bracts spathulate, (1) (0)
= =
chromosome number x 5 (2n
basic 10),
involucral bracts not spathulate. 56), (1)
= =
chromosome number x 4
20. Receptacle without long squamae, receptacle (2) basic (2ri 8, 16).
(0) (1)
squamae.
with long
21. Receptacle naked, receptacle paleate, with
(0) (1)
paleae abaxially to the florets, (2) receptacle deeply Figure 2 was computed with the nelsen
^ree in
alveolate.
option and shows the clades consistently present
22. Paleae deciduous, paleae persistent.
(0) (1)
111 the equally parsimonious cladograms. Tn Fig-
in
all
23. (0) Paleae entire, (1) /paleae apicafly serrate to la- ures i andj 2o .tuhe .termmali t. axa are namedj as m
1
,^,
. . , . two
cmiate, paleae with lateral teeth. ^
(2) r^
\ ^ ^ tr oi^Uow-me i i- eenen•c i -r -
n ^*™ able z, the earlier classincation,
nA /n\ a u* a 1 ^ ^
24. rn>ay-flore^ts monomorphic, \ ray-floret^s dimor- '
(0) (1)
whereas the generic reclassification proposed below
1
names
indicated with in capitals. In the following
25. Ray-floret lamina usually 4-veined, (1) ray-floret is
(0)
Relhania Leysera and Oedera
lamina frequently with up to 10 veins. discussion, s.L, s.L,
whereas
26. Ray-floret tube cylindrical, ray-floret tube refer to the earlier circumscriptions,
(0) (1) s.s.
somewhat Relhania Leysera and Oedera
triquetrous. s.L refer
s.s., s.s.,
27. (0) Disc-florets perfect, (1) disc -florets functionally ^^ ^^^ proposed new delimitations
male, undivided.
style among 200
Variation the equally parsimoniously
somewhat
funnel-shaped,
28. (0) Disc-flor«?t corolla in-
100
cladograms restricted to alternative topol-
is
distinctly divided in tube and limb, (1) disc-floret
ogies within the screened right part of Figures
1
and
corolla distinctly divided in tube limb.
and 2 {Oedera s.L) as well as to two alternative
1064 Annals of the
Garden
Missouri Botanical
Table Data matrix. The characters are Table The taxon names follow the earlier classification.
2. listed in 1.
Inapplicable, unknown, or variable character states are coded -.
number
Character
0000000001 1222222222233333333334444444
1 1 1 1 1 1 1 1
1234567890123456789012345678901234567890123456
000000000000000000000—0000000000000-0000000—
Antithrixia flavicoma
1010000001000100--0000010100001000000000-
montana 00001
Oreoleysera
010011000001001000000~-00-0110101000— 100-
0001
Leysera longipes
0000-1000000001000-10—00001211010010001111102
Leysera gnaphalodes
1000110000000010001 10--00001211010010001111102
Leysera
tenella
10--00001211010010001111102
1000110000000010011
Leysera
leyseroides
001100000000000000001000000000000001000000201-
Rosenia glandulosa
R 00110000000000000000-000000000000001000000201-
osenia spinescens
000000000000000000000—0000000-000010000002010
Rosenia
opposltifoUa
00-00000000-00000000-0000000000000010000002010
Rosenia humilis
000010000000110000000—0000100100000—00013-1-
Relhania
dieterlenii
000011000000110000001000000100100000—00013-2-
Relhania
frutieosa
000011010000110000001000100100100000—00013-2-
Relhania
rotundifolia
0000110100001100000000000011001000011000013-10
Relhania
spathulifolia
000011010000110100000000000100100001100001301-
Relhania corymbosa
0000110100001101000000000011001000011000013-10
Relhania
tricephala
0000110100001100000000000011001000011000013-1-
Relhania relhanioides
000011-0000110010000100000011010000100-0013-10
Relhania
genistijolia
0000110000011001000010000001101000010000013-10
Relhania
viscosa
0000111000011001000010000001101010010000013-10
Relhania squarrosa
000011100001100100001000000110100000—10113-1-
Relhania
conferta
000010000101100100001000000110100000—00013-10
Relhania
steyniae
000011100000100100002—00011101000011000113-10
Relhania
garnotii
000011000001200101001000000110111000—0013-1-
Relhania multlpunetata
0000110000012001011010000001101000010000013-10
Relhania
resinifera
0000110000012001011010000001101-10010000013010
Relhania
sedifolia
0000110000012000000010000001001000010000013-1-
Relhania
foveolata
0000000000012001011010000001101000010000013-1-
Relhania nordenstamii
0000100000011000000010000001101000010000013-10
Relhania
unifiora
000010000001100100000—0000110100000—10013-1-
Relhania
silicieola
100011000000000000101020-001201000010100113-11
Relhania pumila
1000110000000000001010200001201000010100113-11
Relhania
sessilijlora
100011000000000000100—0-001211010010000013-21
Relhania
biennis
0100010000010000000000000001101000010000113-10
Relhania luberosa
001011000101100000000—00001101110010000113-1-
Relhania acerosa
0000100001001000000011000001001100010000113-10
Relhania decussata
Relhania pungens 00001 10001 10000000001 100-100200101 100001 13-10
1
000011-00111000000001100-100200101110000113-10
Relhania calycina
000011-00111-000000011001100200-01110000113-10
Relhania speciosa
000011101101000210001011100120100000—00013-1-
Oedera rapensis
000011121111000210001011100110100000—00013-1-
Oedera
hirta
000011121111000210001011100120100000—00013-1-
Oedera imbricata
000011101111000210001011100120100000—00013-10
Oedera intermedia
000010101111000210001011100100100000—00-13-1-
Oedera
niuirii
sister groups to Relhdnia tricephala, either R. Several analyses were also performed with slight
+
corymbosa or the species pair relhanioides modifications of the data matrix, omitting some
/?.
R. spathulifolia, within the screened part of characters or including others not accepted the
left in
The
Figures and 2 {Relhania majority of final matrix (ultimately rejected for various rea-
s.s.).
1
the clades in the cladogram in Figure ire present sons, for example, because of unreliable informa-
1
some
in the other equally parsimonious cladograms, tion in taxa or because of strong correlation
all
as shown by the strict consensus tree in Figure 2. with other included characters such as presence of
. L » '
Volume Number 4 Anderberg & Bremer 1065
78,
1991 and
Analysis Reclassification of
Relhania
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& 1067
Volume Number 4 Anderberg Bremer
78,
and
1991 Analysis Reclassification of
Relhania
glands on various organs). These analyses showed, of 10 Relhania species. This unfortunate, but
is
hand we want main-
as might be expected, that the clades supported no other alternative at if to
is
by single parallelisms are weak and easily broken tain monophyletic genera. Relhania iiseli is poorly
monophyletic group.
up even with changes the data matrix. defined but constitutes a
slight in
still
By we
However, in analyses, Rosenia and Oreoleysera accepting this second classification, gain a
all
came cladogram phylogenetic hypothesis with
successively out at the base of the detailed, predictive
with the remaining genera as one monophyletic more explanatory power, because more taxonomic
To
group. Furthermore, Leysera and Oedera were statements are made. reduce this information
all
and conceal
always firmly nested within a paraphyletic Rel- of phylogenetic interrelationships in
it
h ania a lumping such as outlined above would hardly
s
.1.
improve progress.
scientific
Consequently, we have resurrected Nestlera in
Discussion
original monotypic sense, reestablished the old
its
The consistency index 0.45 and the frequent ditypic genus Rhynchopsidium, described another
num-
occurrence of parallelisms and reversals on the ditypic genus Comhorhiza, and transferred a
Oedera
cladogram Figure show that there consid- ber of the former Relhania species to (see
in
1 is
erable homoplasy in the data, as is usually found Figs. 1, 2).
The Relhania 13 mostly with
in similarly sii^ed analyses of other groups. is restricted lo species,
impact of homoplastic characters on the phylo- hairy and glandular-punctate (not glandular-hairy)
The
and most and genus comprises
genetic hypothesis important, the par- leaves solitary capitula.
is
Bremer
simonious way which these characters can be two distinct groups, recognized also by
in
more
R.
distributed on a cladogram virtually impossible (1976a), as well as a isolated species,
is
to detect without computerized programs. As a garnoUi. The two groups ai'e R. acerosa to R,
scientific theory, the phylogeny reconstructed by speciosa, with five species (including the distinct
parsimony analysis superior to any phylogeny subgroup comprising R. pungens, R. calycina,
is
obtained from a limited selection of characters only, and R. speciosa)^ and R. dieterlenii to R. spa-
since the former takes into account all available thulifolia, with seven species (see Figs. 1, 2).
information. Parsimony analysis increasingly em- Leysera restricted to the three typical species
is is
ployed in taxonomic revisions, and offers possi- and thus the most well-defined of all the genera,
it is
for improved, phylogenetically more infor- with a whole set of synapomorphies, e.g., plumose
bilities
mative classifications, where the named taxa reflect pappus bristles (see Fig. 1). The former Leysera
evolutionary units rather than artificially composed longipes and Relhania tuberosa are grouped as
groups defined only by a few characters sister species in the new genus Comhorhiza. They
synapomorphy
In case, the analysis clearly demonstrates have a unique in their thick sub-
this
paraphyletic with iej^sera and terranean rhizomes. Although different in several
that /?e//ia;im
s.I. is
among
Oedera excluded. Delimitation the three characters, the two species are habitually similar,
genera has to be altered in order to meet the Comhorhiza virgata (= Relhania tuberosa) hav-
demand monophyletic To lump genera pedunculoid stems with reduced leaves beneath
taxa. ing
for all
into a single genus, Leysera sensu latissimo {Ley- the capitula, approaching the condition in C. Ion-
The
sera being the oldest name), would result in a very gipes. genus has adaxially glabrous, dorsally
heterogeneous taxon, defined only by characters glandular-hairy leaves, not glandular-punctate as
Oedera
Relhania and
such as absence of brachyblasts, alternate leaves, in s.s. s.l.
unbranched anther ray- The monotypic genus Nestlera and the ditypic
glandular floret tube, tails,
groups
without pappus, and corolla divided in tube genus /?/ij/ic/io/?5tc/mm are successive sister
florets
and limb. The characters are weak and can, in a to Leysera s.s. (see Figs. 1, 2). Except for the
wider context, also be shown to be parallelisms (secondarily) perennial L. gnaphalodes, they are
chromosome
by investigation of a larger number of taxa annuals or biennials with reduced base
(e.g.,
=
of the Gnaphalieae). A better alternative to split numbers, x 5 in Rhynchopsidium and Nestlera^
is
=
Relhania and Leysera and recognize smaller and x 4 in Leysera. (Humphries et al. (1978)
=
monophyletic genera that are more morphologi- reported 2/i 14 for L. leyseroides, a count that
homogeneous. This would mean the separa- must be doubted in the light of Blanca's (1983)
cally
tion of the apomorphic Leysera s.s., Rhynchop- detailed investigation of the karyology of the spe-
sidium, Comhorhiza, and Oedera from edich other, cies; Blanca showed that L. leyseroides is a tet-
=
=
The Oedera 2n
and from Relhania. of raploid with 16.)
definition
is
amended 12 former
hereby weakened due to the transfer to this genus Oedera, finally, is to include
1068 Annals the
of
Garden
Missouri Botanical
ReUiania species, and as such (Jedera cnar- apomorphies, the usual approach during
s.l. is cladistic
acterized by the generally glabrous, glandular- the middle-1970s. Thus, Bremer (1976a, 1978a)
punctate leaves, and the cymose-corymbose to con- defined genera on some pappus features, at the
same
gested capitula. Tnfraspecific relationships within time pointing out the possibility that the sup-
Oedera are uncertain. The analysis produced posed synapomorphies could be Parsimony
still false.
100 Oedera
equally parsimonious topologies within analysis of the characters has indeed produced
all
We
Nevertheless, must be concluded that Oedera a revised picture of the phylogeny. have
s.l. re-
it
s.s. firmly nested within the larger monophyletic circumscribed genera according to the best phy-
is
group that we have circumscribed as Oedera logenelic hypothesis available, as expressed the
s.l. in
Hence, Oedera s.s. cannot be maintained at the cladograms produced by the analysis. Reclassifi-
gcneric level, since would leave an unresolved cation of genera like Relhania formerly de-
it s.l.,
paraphyletic residue of former Relhania species. fined by unique, alleged synapomorphies, which
in
parsimony analysis of larger groups are shown to
become many
be necessary
false, is likely to in
Conclusion
other generic groups
as well.
The
former generic classification of the Rel- The present classification of the Relhania group
hania group was based on selected, alleged syn- summarized the following key:
in
is
la. Plants compact, cushion-forming
Oreoleysera
lb. Plants not cushion-forming.
Pappus
2a. of capillary bristles, scales absent Antithrixia
Pappus
2b. at least in ray-florets with conspicuous scales, bristles present or absent.
3a. Pappus bristles distinctly plumose Leysera
Pappus
3b. bristles scabrid-barbellate or absent.
Annual
4a. or biennfal herbs.
5a. Cypselas densely hairy with long, apically coiled hairs Rhynchopsidium
5b. Cypselas almost glabrous, Nestlera
4b. Perennial half-shrubs, shrublets or shrubs.
1-4
6a. Disc-flurets with pappus Rosenia
bristles
6b. Disc-florets without pappus
bristles.
7a. Plants glabrous (if hairy then with capitula congested in secondary heads), capitula
cymose-corymbose
generally to clustered in secondary heads solitary then with
(if
±
glabrous, distinctly one-nerved leaves) Oedera
±
7b. Plants hairy or glandular-hairy glabrous then with 3-9-nerved
(if distinctly leaves),
corymbose
capitula generally solitary (if then with hairy leaves).
Leaves
8a. with long glandular hairs Comborhiza
Leaves
8b. without long glandular hairs Relhania
DC,
Antithrixia Prodr. 6: 277. 1838. TYPE: A, connate, barbellate, capillary bristles in one to two
DC, Chromosome number unknown.
Jlaviconia rows.
Monotypic: A. Jlavicoma DC. Reference: Bre-
Shrublet. Leaves opposite or crowded on brach-
mer, 1978b.
concave and
yblasts, sessile, linear, adaxially to-
mentose, glandular-hairy, apically mucronate;
margin entire. Capitula solitary, terminal, het- Rosenia Thunb., Nov, Gen. 161 1800. TYPE:
/?.
.
erogamous, radiate. Involucral bracts with spathu- glandulosa Thunb.
laminas. Receptacle Female
late epaleate.
flat,
Synonym:
Polychaetia
Less.
fiorets yellow with purple bands dorsally,
pistillate,
radiate. In one row, fewer than the disc-florets. Shrublets. Leaves decussate or rarely alternate.
and pappus
C>/>.sr/a.s as in disc-florets. Disc-Jlorets straight, adaxially concave, tomentose, glandular-
perfect. Corolla yellow. Anthers ecalcarate with hairy; margin entire. Capitula heterogamous, ra-
short tails; endothecial tissue polarized; apical ap- diate, solitary or few together. Involucral bracts
pendage
acute. Style branches truncate with obtuse with spathulate laminas. Receptacle
often pa-
flat,
sweeping-hairs apically. Cypselas rodlike, with ten leate. Female florets yellow with purple bands dor-
X
{2 5) vascular bundles, glabrous or with a few sally, radiate, pistillate, in one row, fewer than the
scattered, elongated twin Pappus of Cypselas and pappus
hairs. slightly disc-florets. as in disc-florets.
Volume Number 4 Anderberg & Bremer 1069
78,
1991 Analysis and Reclassification of
Relhania
Disc-florets perfect. Corolla yellow. Anthers ecal- ecalcarate, with short tails; endolhecial tissue po-
carate with short tails; endothecial tissue polarized; larized; apical appendage acute. Style branches
±
apical appendage acute. Style branches truncate truncate with obtuse sweeping-hairs apically. Cyp-
with obtuse sweeping-hairs apically. Cypselas rod- selas narrowly elliptic with three vascular bundles,
like with five ascular bundles, sparsely hairy with densely hairy with elongated, apically coiled twin
^
± ±
elongated twin hairs. Pappus a crown of free hairs. Pappus a crown of free scales. Chro-
=
and mosome number 2n
scales, 1--4, barbellate, capillary bristles with 10.
Chromosome number 2n Two pumilum DC. (=
flattened apical portion. species: R. (L. Rel-
f.)
=
14, 28, 56. hania pumila (L. Thunb.), R. sessiliflorum (L.
f.)
Your R. glandulosa^\\\xr^.^ R. humilis DC. (= Relhania Thunb.).
s^ecie^: sessiflora (L.
f.) f.)
(Less.) Bremei", R. oppositifolia (DC.) Bremer, R. Reference: Bremer 1976a (spp. 21-22).
spinescens Reference: Bremer, 1976b.
D(!^.
Nestlera Sprengel, Kennt. Gewachse
Anleit.
2(2):
568. 1818. TYPE: (Jacq.) Sprengel.
A'. &/'r;z/i/s
Oreoleysera
Bremer, Bot. Not. 131: 450. 1978.
montana
Bremer.
TYPE: 0. (Bolus)
Synonyms: Columcllea nom. Stephanopap-
Jacq.
illeg.,
pus
Less.
Compact woody
forming dense
perennial,
tufts.
Leaves
Leaves alternate, narrowly oblong, Biennial herb. alternate or occasionally
sessile, flat-
tened, grayish tomentose on both surfaces, eglan- opposite, straight, adaxially concave, tomentose,
dular; margin entire. Capitula solitary, terminal glandular-hairy; glands with multicellular heads;
on long leafless peduncles, heterogamous, radiate. margin entire. Capitula heterogamous, radiate,
Involucral bracts with lanceolate laminas. Recep- solitary or few together. Involucral bracts with
tacle epaleate. Female yellow with pur- lanceolate laminas. Receptacle often paleate.
flat, florets flat,
Female
pie bands dorsally, pistillate, radiate, in one row. florets yellow with purple bands dorsally,
fewer than the disc-florets. Cypselas and pappus radiate, pistillate, in one row, fewer than the disc-
Cypselas and pappus
as in disc-florets. Disc-florets perfect. Corolla yel- florets. as in disc-florets. Disc-
low, glandular hairy. Anthers ecalcarate with short florets perfect. Corolla yellow, often with robust
endothecial tissue polarized; apical appendage muhicellular trichomes. Anthers ecalcarate, with
tails;
acute. Style branches truncate with obtuse sweep- short tails; endothecial tissue polarized; apical ap-
±
pendage
Cypselas truncate. Style-branches truncate with
ing hairs apically. rodlike with five vas-
cular bundles, moderately hairy with elongated twin obtuse sweeping-hairs apically. Cypselas rodlike
hairs. Pappus of free, barbellate, capillary bristles with five vascular bundles, sparsely hairy with elon-
in one row with an outer row of very narrow scales. gated twin hairs. Pappus a tubular crown of connate
Chromosome number unknown. scales. Chromosome number 2n 10.
Monotypic: 0. montana (Bolus) Bremer. Ref- Monotypic: N. biennis (Jacq.) Sprengel (= Rel-
erence: Bremer, 1978b. hania biennis (Jacq.) Bremer). Reference: Bremer,
1976a
(sp. 23).
Rhynchopsidium DC, Mem.
Geneve
Soc. Phys.
Leysera Amoen.
1249, 1763;
L. Sp. PI. ed. 2, 2:
283. 1836. type: R.
sessiliflorum (L.
7: f.)
Acad. 104, 1763; Gen. 431,
6: PI. ed. 6:
DC,
gnaphalodes
1764.
TYPE: L. (L.) L.
Synonym: Rhynchocarpus nom.
Less.,
illeg.
Synonyms: Asteropterus Adans., Callicornia Burm.
(Vaill.)
Longchampia
Leptophytus
Annual herbs. Leaves alternate or sometimes f., Callisia L., Cass.,
Willd., Pseudocrupina Velen., Leyseria Necker,
concave, tomentose,
opposite, straight, adaxially
Leyssera auct., ortogr. var.
margin Capitula heterog-
glandular-hairy; entire.
amous, radiate, solitary, sessile or pedunculate. Annual herbs or a perennial half-shrub or shrub-
Recep- Leaves
Involucral bracts with lanceolate laminas. alternate or occasionally opposite, ses-
let.
tacle flat-convex, paleate; paleae lanceolate with sile, linear, straight, mucronate, adaxially concave,
two lateral teeth, Fema/e^ore^5 yellow with purple tomentose, glandular-hairy; margin entire. Capit-
bands dorsally, radiate, pistillate, in one row, fewer ula heterogamous, radiate, solitary, terminal on
than the disc-florets. Cypselas and pappus as in long leafless peduncles, Involucral bracts with lan-
disc-florets. Di5<:-y?ore/5 perfect. Corolla yellow, of- ceolate laminas. Receptacle flat, marginally with
Female
ten with robust multicellular trichomes. Anthers prominent squamae. florets yellow with
,
1070 Annals the
of
Garden
Missouri Botanical
&
purple bands dorsally, radiate or niiniradiate, pis- C. virgata (N. E. Br.) Anderb. Bremer.
tillate, in one row, fewer tlian the disc-florets; tube Eponyniy: The name is deduced from the Greek
Cypsclas words combos, meaning and
often with robust, conical tricliomcs. as knot, tuber, rhi-
Pappus meaning
in disc-florets. of scales only. Disc-Jlorets za, root.
perfect. Corolla yellow, often with robust multi-
Fruticulus vel suffrutex. Caules rhizoinate vel tubere
Anthers ecalcarate, with short
cellular Iriclioines.
liypogaeo, crasso, lignoso exorientes. Folia alterna vel in
appendage
endothecial tissue polarized; apical
tails;
brachyblastis congesta, concava, inferne
sessilia, stricta,
truncate. Style branches truncate with obtuse
glandulis longistipitatis praedita, superne glabra, margine
sweeping-hairs apirally. Cypsclas rodlike, with three integra. Capituln heterogama, radiata, solitaria, ad apices
vascular bundles, glabrous or with scattered, elon- cauliuni paucifoliatorum vel pedunculorum longissimo-
runi. Bracteae involucri pluriseriatae lamina scariosa,
gated twin hairs. Pappus of free, apically plumose,
spathulata infuscata. Receptaculum planum vel convex-
capillary bristles in one row with an outer row of
um,
epaleatum. Flosculi radii sed dorse interdum
flavi
=
Chromosome number 2n
obtuse scales. 8, 16.
pur[)ureo-fasciati, pistillati, uniseriati, flosculos disci pauci-
Three gmiphalodes Cj/>5e/aeeaedem Pappus ex squam-
species: L. (L.) L., L. ley- ores. in flosculis disci.
seroides (Desf.) Maire, L. tenella DC. Reference: is angustis scariosis constans. Flosculi disci perfecti. Co-
rolla flava. Antherae ecalcaratae, breviter caudatae,
Bremer, 1978a.
appendicibus apicalibus acutis; endothecio polarato. Styli
rami Cypselae
apice truncati, pilis obtusis apicaliter. an-
Relhania L'Her., nom. cons.. Sert. And. 22, 6^^^^ ellipticae, tri- vel quinquevenosae, pilosae vel fere
1:
Pappus
glabrae. ex setis liberis, capillaribus, barbellatis,
1789. TYPE: R, Bremer.
friiticosa (L.)
squamis tantum squam-
uniseriatis et angustis scariosis vel
scariosis plus minusve connatis solum constans.
Synonyms: Osmites Laprirousia Thunb., nom. is
L.,
illeg.
Shrubs, shrublets, or half-shrubs. Leaves alter- Shrublet or suffrutex. Stems arising from sub-
nale or decussate, sessile, straight, adaxially terranean, thick woody tubers or rhizomes. Leaves
concave, tomentose, margin Capltula crowded on
entire. het- alternate or brachyblasts, sessile,
erogamous, radiate, solitary or sometimes cymose- straight, adaxially concave, glabrous above,
corymbose. Involucral bracts with spathulate lam- glandular-hairy below, margin Capltula
entire.
inas. Receptacle flat to convex or rarely deeply hoterogamous, radiate, solitary, on pedunculoid few-
alveolate, often paleate. Female Jlorcls yellow with leaved stems or on long leafless peduncles. Invo-
purple bands dorsally, radiate, pislillale, in one row, lucral bracts with spathulate laminas. Receptacle
fewer than the disc-florets. Cypsclas and pappus or to convex, epaleate. Female florets
flat flat
as In disc-florets. Disc-Jlorets perfect or function- yellow with purple bands dorsally, radiate,
pistillate,
ally male. Corolla yellow, sometimes with robust in one row, fewer than the disc-florets. Cypsclas
multicellular trichomas. Anthers ecalcarate, with as in disc-florets. Pappus of narrow scales. Disc-
short tails; endothecial tissue polarized; apical ap- florets perfect. Corolla yellow. Anthers ecalcarate,
pendage acute. Style branches truncate with obtuse with short endothecial tissue polarized; apical
tails;
sweeping-hairs apically (rarely obtuse with sweep- appendage acute. Style branches truncate with ob-
Cypsclas
ing-hairs dorsally). rodlike with 5 or 3 tuse sweeping-hairs Cypsclas narrowly
apically.
vascular bundles, glabrous or sparsely hairy with elliptic with five or three vascular bundles, glabrous
elongated twin hairs. Pap[)ns a crown of free to or sparsely hairy with elongated twin hairs. Pappus
connate scales without bristles, occasionally with of 5 barbellate capillary brisdes and narrow scales,
1
or 2 barbellate bristles in odd florets. Chromosome or of ± connate scales only. Chromosome number
=
number
2n 2n
It. 14.
T
wo
thirteen species: R. acerosa (DC.) Bremer, R. species:
calycina (L. L'Her., R. corymbosa (Bolus) Bre-
f.)
&
comb
longipes (Bremer) Anderb. Bremer,
mer, R. decussata L'Her., R. dieterlenii C.
{E. Phil-
R nov. Basionym: L^eysera longipes Bremer, Bot.
Bremer, fruticosa Bremer, R. garnotii
lips) (L.)
.
pungens Not. 131: 381. 1978.
(Less.) Bremer, R, L'Her., relhan-
/?.
&
ioides (Schltr.) Bremer, R. rotundifolia C. virgata (N. E. Br.) Anderb. Bremer, comb,
Les:
spathulijolia Bremer, R, spcciosa (DC.) nov. Basionym: Nestlera virgata N. E. Br.,
iirv
I U.1 V
.
Kew
1895: 25 (= Relhania tubcrosa
R. tricephala (DC.) Bremer. Reference: Bremer, Bull.
Bremer).
1976a 1-7, 25-29).
(spp. 13,
References: Bremer, 1976a 1978a
(sp. 24), (sp.
&
Comborhiza
Anderb. Bremer, gen. nov. TYPE:
4).