New Zealand Journal of Botany, 1993, Vol.

31: 43-58 43
O028-825X/93/3101-O043 $2.50/0 © The Royal Society of New Zealand 1993

Systematics of New Zealand Inuleae (Compositae-Asteraceae)—3

Numerical phenetic analysis of leaf anatomy and flavonoids

ILSE BREITWIESER* INTRODUCTION

JOSEPHINE M. WARD This study uses analysis of chemical and anatomical
Department of Plant and Microbial Sciences leaf data to elucidate taxonomic relationships in the
University of Canterbury New Zealand Inuleae*. A representative selection
Private Bag 4800 of species of all indigenous genera except Craspedia
Christchurch, New Zealand is analysed as well as critical species from Tasmania
and a single species of Anaphalis from the
*Present address: Botanischer Garten und Botanisches Himalayas, included for comparison with the New
Museum Berlin-Dahlem, Konigen-Luise-Strasse 6- Zealand species recently transferred by Webb (1987)
8, D-1000 Berlin 33, Federal Republic of Germany. to this genus. Taxa examined are listed in Table 1.
Of the nine genera of the Inuleae which are
indigenous to New Zealand, Craspedia is part of the
Abstract Forty-five species of Inuleae Angianthus complex centred in Australia and refer-
(Compositae), predominantly from New Zealand able to Bentham's (1873a) subtribe Angianthinae.
and Tasmania, were subjected to numerical phenetic The remaining eight genera all fall within Bentham's
analysis using 49 leaf anatomy and 38 leaf flavonoid subtribe Gnaphaliinae, and are in the groups of
characters. Results indicate that on the basis of genera clustered around Anaphalis, Gnaphalium,
overall similarity in this suite of characters: (1) and Helichrysum (Merxmiiller et al. 1977)^.
Leucogenes is a good genus; (2) an undescribed
taxon of Inuleae is a distinct monotypic genus; (3) The classification of New Zealand species of
New Zealand species currently placed in Anaphalis, Gnaphaliinae has been strongly affected by
Cassinia, and Ewartia are generically distinct but European ideas, especially those of Bentham and
are referred to the wrong genera; (4) similarity Hooker. In the nineteenth century, most herbaceous
between species of Ewartia and Gnaphalium New Zealand species were referred to one of the two
indicates that the boundary between these genera large, widespread genera Gnaphalium and
needs to be reassessed; (5) both Helichrysum and Helichrysum, distinguished by the ratio of female
Raoulia are heterogeneous and need revision; (6) in
New Zealand, Pseudognaphalium is distinct from
other gnaphalioid genera.
*Anderberg (1989) regards the tribe Inuleae sensu lato as
paraphyletic. He has divided it into three tribes, with all
Keywords Compositae; Asteraceae; Inuleae; New Zealand genera falling into the largest of these, the
Gnaphaliinae; Anaphalis; Cassinia; Ewartia; Gnaphalieae. However, paraphyly of the Inuleae is not yet
Gnaphalium; Haastia; Helichrysum; Leucogenes; sufficiently established to warrant abandoning the traditional
Pseudognaphalium; Pterygopappus; Raoulia; leaf concept of this tribe (Bremer et al. 1992). We do not accept
the revised subtribes proposed for the Gnaphalieae by
anatomy; flavonoids; taxonomy; systematics; Anderberg (1991), which separate closely related
numerical phenetics; New Zealand; Tasmania (hybridising as well as phenetically similar; Ward 1981,
1993b) New Zealand genera at the subtribal level. Generic
name changes proposed by Anderberg (1991) are noted
but not used in this paper. They will be discussed and
recommendations will be made regarding their use in a
forthcoming revision of New Zealand Inuleae.

subtribal name Gnaphaliinae is used throughout this

B92012 paper in the sense of Bentham, not the much broader sense
Received 31 March 1992; accepted 3 February 1993 of Merxmiiller et al. (1977).
44 New Zealand Journal of Botany, 1993, Vol. 31

Table 1 Species included in this paper.

Anaphalis A. triplinervis Himalayas

A. keriensis New Zealand
A. rupestris New Zealand
A. subrigida New Zealand
A. trinervis New Zealand
Cassinia C. aculeata Tasmania
C. longifolia Tasmania
C. fulvida New Zealand
C. leptophylla New Zealand
Ewartia E. catipes Tasmania
E. meredithiae Tasmania
E. planchonn Tasmania
E. sinclairii New Zealand
Gnaphalium sect. G. involucratum New Zealand
Euchiton G. mackayi New Zealand
G. nitidulum New Zealand
G. traversii New Zealand
Helichrysum sect. H. backhousei Tasmania
Ozothamnus H. obcordatum Tasmania
H. lanceolatum New Zealand
H. coralloides New Zealand
H. depressant New Zealand
H. dimorphum New Zealand
H. intermedium New Zealand
H. parvifolium New Zealand
sect. H. bellidioides New Zealand
Xerochlaena H. fllicaule New Zealand
Leucogenes L. grandiceps New Zealand
L. leontopodium New Zealand
Pseudognaphalium subg. Laphangium P. luteoalbum New Zealand
Raoulia subg. Raoulia R. hookeri New Zealand
R. tenuicaulis New Zealand
R. glabra New Zealand
R. cinerea New Zealand
R. species 1 New Zealand
subg. Mistura R. petriensis New Zealand
subg. Psychrophyton R. grandiflora New Zealand
R. hectorii New Zealand
R. bryoides New Zealand
R. eximia New Zealand
R. species 2 New Zealand
Inuleae genus 1 undescribed species New Zealand
Haastia H. pulvinaris New Zealand
H. sinclairii New Zealand
Pterygopappus P. lawrencei Tasmania
Solid lines show present taxonomic limits. Dotted lines show limits suggested by this study.
Breitwieser & Ward—N.Z. Inuleae 45

to hermaphrodite florets in the capitulum. The effects Ewartiothamnus. The integrity of Ewartia in
of this have been to group the New Zealand taxa Australia has also been questioned (Ward 1981,
into widely distributed genera with which they have 1993b; Anderberg 1991).
little in common and to obscure differences among Gnaphalium sensu lato has traditionally been a
the New Zealand taxa themselves. Classification has vast, heterogeneous, widespread genus characterised
also been hampered by the overlapping distributions by its herbaceous habit, usually tomentose leaves,
of critical character states among New Zealand taxa. and capitula with predominantly female florets. The
A given species may possess diagnostic character New Zealand species in Allan (1961) were shown
states of several genera, with its generic position by Drury (1970) to comprise three distinct groups.
changing as different characters are emphasised. The "achyroclinoid group", with divided stereomes,
Of the genera of Gnaphaliinae indigenous to New had one indigenous species, G. luteoalbum,
Zealand, Anaphalis is characterised by conspicuous subsequently transferred to Pseudognaphalium by
white involucral bracts, dorsal sweeping hairs all Hilliard & Burtt (1981). The "anaphalioid group"
along the style arms, and subdioecy. It is a mainly had four indigenous species, transferred by Webb
Asian genus with four species, all endemic, in New (1987) to Anaptialis as outlined above. The
Zealand. The New Zealand species were classified "gnaphalioid group" remained in Gnaphalium, with
by Bentham (1873b) as Gnaphalium section all New Zealand species falling within sect.
Anaphalioides, connecting Gnaphalium to Euchiton, which has its greatest species con-
Anaphalis and Helichrysum. Separate generic status centration in this country and for which separate
has been proposed or suggested for them generic status has recently been proposed
(Kirpichnikov & Kuprijanova 1950; Drury 1970; (Anderberg 1991). It is characterised by a stolon-
Merxmiiller et al. 1977; Anderberg 1991), but Webb iferous habit, achenes with paired papillae (Drury
(1987) transferred them to Anaphalis, pending more 1970), pappus hairs with basal cilia, and capitula in
detailed investigation of their relationships. Several terminal and sometimes axillary involucrate clusters,
authors (Drury 1970, 1971; Ward 1981, 1993b; or solitary. Of the 14 indigenous species, two, G.
Webb 1987) have noted the similarity between these mackayi and G. nitidulum, differ in their non-
species and the endemic Helichrysum bellidioides, stoloniferous mat habit and achenes with imbricate
which was not formerly classified with them in rather than paired papillae. Bentham (1873b) noted
Gnaphalium because it lacked the preponderance of that G. nitidulum is scarcely distinguishable from
female florets characteristic of that genus. Ewartia (then Raoulia) planchonii of Tasmania.
About half the New Zealand species are endemic,
Cassinia is an Australasian genus of some 21
the remainder being shared with Australia and one,
species, characterised by a shrub habit and
G. involucratum, also with Taiwan, Java, and the
homogamous capitula which have white involucral
Philippines (Webb 1988).
bracts and are clustered in secondary inflorescences.
It is distinguished from Helichrysum sect. Helichrysum, like Gnaphalium, has been a very
Ozothamnus by having receptacular paleae among large widespread genus, to which a heterogeneous
the florets. The New Zealand species (five in Allan assortment of New Zealand species with pre-
1961, but reduced to one by Webb 1988) are all dominantly hermaphrodite florets has been referred.
endemic. Sometimes they lack receptacular paleae. Of these, two have been transferred to Leucogenes
It has been suggested (Hooker 1864; Hilliard & Burtt (Beauverd 1910), one to Raoulia (Beauverd 1912),
1973; Breitwieser 1990, in press) that they are closer and one to Ewartia (Cheeseman 1925). There remain
to Australian species of Helichrysum sect. two species in the herbaceous perennial section
Ozothamnus than to Cassinia. Xerochlaena and seven in the mainly woody section
Ewartia has four Australian species which have Ozothamnus, which is recognised at generic level
a mat habit, terminal capitula which are solitary or by Anderberg (1991). Of the seven species in sect.
a few in a cluster, subdioecy, and dorsal sweeping Ozothamnus, one, Helichrysum lanceolatum, is a
hairs all along the style arms. The single New shrub with broad, spreading leaves and capitula in
Zealand species, which is endemic, and which was fasciculate clusters. It is quite unlike any other
transferred from Helichrysum by Cheeseman (1925), gnaphalioid New Zealand taxon. A further five are
has none of these features and appears to be palpably whipcord shrubs with small, imbricate, appressed
misplaced in Ewartia (Ward 1981, 1993b). leaves and solitary, sessile capitula. The habit of the
Anderberg (1991) has proposed generic status for remaining species, Helichrysum dimorphum, is
the New Zealand species, naming the new genus remarkable. Not only is the plant scandent (the only
46
example of this habit in the New Zealand Inuleae)
but the shoot apex produces alternating and quite
different leaf forms, one scale-like, and appressed
to the stem, the other broad, and strongly deflexed
basally. H. dimorphum grows with and is inter-
mediate in morphology between H. depressum and
H.filicaule. It is now established (Allan 1961) that
it is not a direct hybrid of these two species as
suggested by Wall (1920), but the possibility of
hybrid origin remains.
Leucogenes was erected by Beauverd (1910) for
the two species (both endemic to New Zealand) of
Helichrysum sect. Leontopodioides, which are Anaphalis triplinervis which was taken from
characterised by capitula like those of Helichrysum
sect. Ozothamnus but arranged, like those of
Leontopodium, in a dense terminal cyme surrounded
by a ring of conspicuous bracts.
Raoulia is a large, endemic New Zealand genus
of small-leaved mat or cushion plants with solitary,
sessile, terminal capitula (for an account see Ward
1981, 1982a). It has long been regarded as
heterogeneous, with the subgenera Raoulia and
Psychrophyton sometimes given generic status
(Beauverd 1910; Anderberg 1991), but more usually
regarded as sections (Bentham 1873b; Kirk 1899;
Cheeseman 1906) or subgenera (Beauverd 1912;
Cheeseman 1925; Allan 1961).
Inuleae genus 1 (CANU 35556) is an unpublished
monotypic genus (Ward unpubl.) described by Allan
(1961, p. 714) under Leucogenes incertae sedis. He
compared it with Haastia sinclairii and Leucogenes
and suggested a possible hybrid origin. It differs
from both these genera in possessing, in addition to
terminal capitula, lateral capitula in the axils of the
upper foliage leaves.
The other genera included in this study are
Haastia and Pterygopappus. Haastia was originally
placed with the "gnaphalioid Compositae" (Hooker
1864) but included in the tribe Astereae by Kirk
(1899), Cheeseman (1906, 1925), and Allan (1961)
because it lacks the tailed anthers of the Inuleae.
Merxmliller et al. (1977), without presenting
reasons, transferred the type species (H. pulvinaris)
to the Inuleae and commented that the other two
species "seem to represent quite another genus".
Pterygopappus is a monotypic Tasmanian endemic
genus, a small-leaved mat plant with solitary,
terminal capitula and florets with a few broad,
flattened, serrulate pappus hairs. It was included in
their "Gnaphalium group" by Merxmiiller et al.
(1977), who suggested a position close to Haastia
pulvinaris, whereas Anderberg (1991) associated it
with Raoulia subg. Psychrophyton (as Psychrophyton).
New Zealand Journal of Botany, 1993, Vol. 31
The present study applies techniques of numerical
phenetic analysis to chemical and anatomical leaf
characters in New Zealand Gnaphaliinae. This is one
of a series of studies including taxonomic, bio-
systematic, phenetic, and cladistic approaches and
leading to a full taxonomic revision of this group.
MATERIAL AND METHODS
All material was collected in the field between
January and March of 1987-89, except material of
cultivation. Details of voucher specimens are given
in Appendix 2. Names follow Allan (1961) or Webb
(1987, 1988) for New Zealand species and Curtis
(1963) for Tasmanian species. The genus of Inuleae
and the two species of Raoulia for which published
names do not exist are designated by numbers within
their ranks and referenced to a CANU herbarium
number. They are Inuleae genus 1 (CANU 35556),
Raoulia species 1 (an alpine mat plant with blackish
leaves, CANU 34359), and Raoulia species 2 (a
small cushion plant of Central Otago mountains,
CANU 35559)*.
For the numerical analyses, three data matrices
of 45 taxa with, respectively, 49 leaf anatomy
characters, 38 flavonoid characters, or all 87
characters, were used. (Flavonoid distribution is
shown in Table 3. Anatomy characters are listed in
Appendix 1.)
For leaf anatomy studies, leaves were fixed in 3%
glutaraldehyde and embedded in 1B-A (Polysciences
Inc. embedding kit) as described in Breitwieser
(1990, in press). Transverse sections of 4 iim
thickness were stained in aqueous methylene blue/
azure blue. Detailed comments on the leaf anatomy
characters and descriptions of the leaf anatomy of
each taxon examined are given in Breitwieser (1990,
in press).
For leaf flavonoid analysis, two-dimensional
paper chromatograms of crude leaf extracts were
prepared following the techniques of Mabry et al.
(1970) and Markham (1982). Two-dimensional
paper chromatograms (2D-PCs) having spots with
low mobility in the solvent system TBA/15% HOAc
were rerun in 50% HOAc. The identity of the spots
*In Ward (1981,1982a, b) and Breitwieser (1990, in press),
Raoulia species 1 and 2 are referred to, respectively, as
Raoulia sp. "M" and Raoulia sp. "L". In Breitwieser (1990,
in press), Inuleae genus 1 is referred to as genus "Z".
Breitwieser & Ward—N.Z. Inuleae 47

was determined by their Rf values, by their colour Adams 1974; Harborne 1975; Harborne & Turner
under UV (366 nm) before and after fuming with 1984; Breitwieser 1990). For the data matrix, the
ammonia, by spraying with a 1% solution of flavonoid characters were coded for presence and
Naturstoffreagenz-A in methanol, and by co- absence. Where the presence of a spot varied within
chromatography, as well as, in some specimens, by one taxon, or the identity of one spot could not be
UV absorption spectroscopy and by hydrolysis. determined, the state was entered as "no
These procedures give results considered sufficiently comparison".
reliable for use in preliminary systematic com- Similarities between the species were calculated
parisons. Problems in the use of phenolic spot using Gower's general coefficient of similarity for
patterns have been discussed fully elsewhere (e.g., qualitative and quantitative characters (Gower
1971). The similarity values were clustered by
UPGMA (the unweighted pair group method using
Table 2 Rf values and fluorescence characteristics of arithmetic averages; Sokal & Michener 1958), which
leaf flavonoids in 45 species of Inuleae. generally gives the least amount of distortion of a
similarity matrix (Rohlf 1970; Sneath & Sokal 1973;
Colour Rf Rf value Rf value Ward 1981). The program used for the numerical
under value (xlOO) (xlOO) analyses was "Gower", by C. M. Frampton, G. A.
Spot Colour UV (xlOO) in 15% in 50% Findlay, and J. M. Ward, Christchurch.
no. under UV +NH3 inTBA HOAc HOAc

1 purple green 47 45
2 purple green 37 39 RESULTS
3 purple green 38 48
4 purple green 45 59 Flavonoids
5 purple green 33 61 Examination of the individual 2D-PCs under UV
6 purple green 40 20 light revealed the presence of 65 different UV-
7 purple green 52 16
8 yellow yellow 60 2
fluorescent or absorbing spots. These were, with the
9 purple yellow 68 2 exception of the 14 bright yellow fluorescent spots,
10 purple purple 77 2 probably phenolic or flavonoid in nature (Markham
11 purple yellow 20 20 1982). On the 2D-PCs, all spots relevant to the
12 purple yellow 37 30 present study typically appeared dark purple or
13 purple green 16 47 yellow. They are distinguished by their colour
14 purple green 12 65 changes in NH3 vapour and by their Rf values in
15 purple purple 27 51 TBA, 15% aqueous acetic acid, and 50% aqueous
16 purple yellow 15 33
acetic acid (Table 2). Other blue fluorescent spots
17 purple green 85 2
18 purple purple 90 10 (a-k) were encountered in almost all taxa. They are
19 yellow yellow 90 2 probably simpler phenolics such as cinnamic acids
20 yellow yellow 80 2 and their glycosides. Their 2D-PC pattern is often
21 purple green 23 >10 6 inconsistent and for this reason no attempt has been
22 purple green 37 >10 15 made in this study to use these components in a
23 yellow yellow 27 >10 42 taxonomic interpretation. The only compound
24 yellow yellow 55 >10 33 appearing as a red spot, identified as a cyanidin-O-
25 purple yellow 41 >10 23
glucoside, was excluded for the same reason. Figure
26 purple purple 73 >10 37
27 purple purple 92 42 1 shows the master composite chromatogram of all
28 purple yellow 51 >10 27 compounds derived from all individual chromato-
29 purple green 55 >10 42 grams, while Fig. 2 shows the master composite
30 purple green 61 >10 37 chromatogram of flavonoids which were used in the
31 yellow yellow 41 >10 23 analyses. The chromatograms characterised by spots
32 yellow yellow 59 30 with low mobility in standard solvent systems were
33 purple yellow 63 >10 43 rerun in 50% acetic acid to obtain better resolution.
34 yellow yellow 65 >10 52
35 purple yellow 69 >10 50 Only the spots with low mobility can be seen here,
36 purple purple 68 2 as shown in the master chromatogram (Fig. 3), since
37 yellow yellow 37 2 all the other spots ran with the solvent front.
38 purple purple 84 25 The semi-quantitative distribution of the leaf
Table 3 Semi-quantitative distribution of leaf flavonoids in 45 species of Inuleae. Key—Relative concentration (as judged visually): +++ = high, ++ = moderate, -p..
+ = low, ± = trace, ? = inconsistent °°

Spot
Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Anaphalis keriensis +++ +++ , + ± ±

A. rupestris ++++++ . + ± ± + + .
A. subrigida ± +++ . ± ++ .
A trinervis + +++ . ±
A. triplinervis + ± + + ± . ±
Cassinia aculeata + . . ++ +++ +++ . ++ ++ . .
C.fiilvida ± ± . . ? + . . . . ++++++++++++
C. leptophylla ± ± . . ? + . . . . + + + +
C. longifolia + +++ ++ . + + . .
Ewartia catipes +++ . . ++ ± + + . . + . . +
E. meredithiae +++ + ++ ± . . + + . .. ±
E. planchonii + + + + . . + + + . + +
E. sinclairii . + . . . +++ ± ±
Gnaphalium involucratum + + + + . . + . . + ± . . . .
G. mackayi + + . . ± + . ±
G. nitidulum ++ + . . +
G. traversii + + + ?
Haastia pulvinaris +++ . ± +
H. sinclairii + + +. ? . . . . +++± +
Helichrysum backhousei ± . . . ± + +
H. bellidioides +++ ± . +++ +++ ++
H. coralloides ? ? . . . ? ++
H. depressum +++ +++ +++ ? ^
H. dimorphum +++ ^
H. filicaule + + + . + + . + + fsj
H. intermedium +++ ++ + £J
H. lanceolatum ?± ^
H. obcordatum ± . . . +++ ± . . . . + + + + +++ +++ J5
H. parvifolium ± ± + +++ +++ l_^
Leucogenes grandiceps . + + + + +++ +++ . ++ ++ ± . ± ± + O
L. leontopodium . + ? ? . . . +++ +++ + ± + . +++ . ± . . ++ + +++ ± + + + + . . . g
Pseudognaphalium luteoalbum ± . . ± ± . . ± a
Pterygopappus lawrencei + ++ . . ? +++ + + ++ ^
l f
Raoulia bryoides ± . . + + ~>
R. cinerea . ± ± . . ± ± W
R. eximia ++.++++ + Q
R. glabra + . . . . ? +++ +++ . + . + . +++ p
R . grandiflora ++ + + . . + + + . +++ . + . ± . +++ . . . . f~
R. hectorii . ± . + + + ? . . . + + + g
R . hookeri . ± ± ± ± + + + . . + . . + + + . . ± ± . + . . . . <§,
R . species 1 . + + i"*1
R. species 2 + + . . ± + . . . + + + <^
R. petriensis . ± . ± ++++++ . . ? ± . ++ O_
R. tenuicauUs . ± ± ± ± ± +++ . . + . . +++ . . ± ± . + . . . . '
Genus 1 ± + ± ± + . . + . . + . . ? . . ? . . . . *-
Breitwieser & Ward—N.Z. Inuleae 49

3U
I
&
1—

CM

1 TBA - ^
Fig. 1 Master composite chromatogram of all compounds in the leaves of 45 species of Inuleae. Spot colour key:
continuous outline - purple or yellow spot; broken outline - blue spot; dotted outline - bright yellow fluorescent spot;
hatched spot - red spot. Numbering and lettering: all flavonoids are numbered (see Table 2, 3). Lettering refers to the
following: a - k, other phenolic compounds; R, rutin marker.

flavonoids shows that all of the 45 species examined
are distinguished by their flavonoid pattern (Table
3), although in some the differences are minor or of
a quantitative nature only. Limited investigation of
infraspecific flavonoid variation indicated it is either
absent or minor and of a quantitative rather than a
qualitative nature. Relative intensities of spots refer
only to the pattern of spots for the taxon under
consideration and are not comparable between taxa.
Three major groups of spots can be distinguished
(Fig. 4). These are the group of flavonol-3-
glycosides in the centre of the 2D-PC (spots I), the
spots with little mobility in standard solvent systems
near the origin (spots II), and the spots with little or
no mobility in 15% aq. acetic acid, but very high
mobility in TBA (spots III). Spots III may have been
surface flavonoids since they could not be detected
on the chromatograms if the leaves were shaken in
chloroform before extraction in methanol. Species
with only spots I are the New Zealand Anaphalis
spp., Helichrysum bellidioid.es and H. filicaule,
Haastia pulvinaris and H. sinclairii, Raoulia eximia
and Raoulia species 1, and the Tasmanian
Pterygopappus lawrencei. The New Zealand species
of Cassinia and the Tasmanian species of
Helichrysum studied have spots III in common. The
species which are characterised by spots II only are
Anaphalis triplinervis, Pseudognaphalium
luteoalbum, Raoulia bryoides, R. grandiflora,
Raoulia species 2, all Tasmanian Ewartia spp., and
the New Zealand Gnaphalium species studied except
G. traversii. Helichrysum lanceolatum is the only
50 New Zealand Journal of Botany, 1993, Vol. 31

•
o
%
u
<
in
i—
CM

1 TBA - •
Fig. 2 Master composite chromatogram of flavonoids (TBA/15% HO Ac) in the leaves of 45 species of Inuleae. Spot
colour (before/after spraying with NH3): hatched spot - purple/green; dotted spot - purple/yellow; double-hatched spot
- purple/purple; blank spot - yellow/yellow. Numbering and lettering: all flavonoids are numbered (see Table 2, 3). R,
rutin marker.

species without spots belonging to any of these three Flavonoid phenogram

major spot groups. The species not mentioned have
spots I as well as spots II. The most immediately apparent feature of the five
Leucogenes leontopodium has the highest number main clusters in Fig. 5 is their lack of conformity
of flavonoid compounds (15), followed by L. with either current (Allan 1961; Webb 1988) or
grandiceps (12), Raoulia tenuicaulis (11),/?. hookeri suggested (Ward 1981,1993a, b) taxonomic arrange-
(10), R. grandiflora (9), and Inuleae genus 1 (7). By ments. They do, however, contain some taxo-
contrast, Helichrysum lanceolatum and H. nomically interesting groups. Thus, the uppermost
dimorphum are characterised by one spot only. large cluster (D) includes representatives of no less
than seven of the total 11 genera, but it groups: the
whipcord species of Helichrysum (A); the New
Leaf anatomy Zealand species of Anaphalis and the two her-
The characters used for the numerical analysis are baceous species of Helichrysum (B); and the two
shown in Appendix 1. Details of comparative leaf species of Haastia (C).
anatomy in New Zealand and related Tasmanian The second large cluster (E) shows little structure
Inuleae is given in Breitwieser (1990, in press). or taxonomic correlation, although it does contain
Breitwieser & Ward—N.Z. Inuleae 51

;
I_o
"55
u
<
5?
o
CO

1 TBA - •

Fig. 3 Master composite chromatogram of flavonoids (TB A/50% HOAc) in the leaves of 45 species of Inuleae. Spot
colour (before/after spraying with NH3): hatched spot - purple/green; dotted spot - purple/yellow; double-hatched spot
- purple/purple; blank spot - yellow/yellow. Numbering and lettering: all flavonoids are numbered (see Table 2, 3).

all three of the Australian species of Ewartia and
three of the four species of Gnaphalium. Clusters F
and H are mixtures with almost as many genera as
species, but G groups the New Zealand species of
Cassinia with the Australian species of Helichrysum
sect. Ozothamnus rather than Australian Cassinia.
Leaf anatomy phenogram
Figure 6 shows a clear hierarchical structure, good
correlation with existing taxonomy, and a large
number of isolated single species. Cluster A
associates the Australian Ewartia catipes with most
of Raoulia, and is linked to Leucogenes which
constitutes cluster B. Cluster C comprises three of
the four species of Gnaphalium (but not the same
three which are grouped in Fig. 5), while D contains
all of Cassinia, all of the non-whipcord species of
Helichrysum except H. lanceolatum, and the other
two Australian species of Ewartia. Haastia makes
up cluster E, while F is the taxonomically unlikely
pairing of Gnaphalium involucratum with Anaphalis
triplinervis from the Himalayas. The whipcord forms
of Helichrysum constitute cluster G, which is quite
closely linked to the only other species in this
analysis which has inverse-dorsiventral leaves,
Raoulia petriensis. Cluster H, comprising the four
New Zealand species of Anaphalis, is both tighter
and more isolated than any of those above.
The isolated species in Fig. 6 are generally those
which also occupy isolated taxonomic positions.
52
•p
3
u
CM
1 TBA - •
Fig. 4 Spot groups in the chromatograms of 45 species of Inuleae.
Pterygopappus lawrencei and Inuleae genus 1 are
both monotypic genera, Raoulia petriensis is alone
in its subgenus, and Pseudognaphalium luteoalbum
is the only species of its genus in this analysis.
Ewartia sinclairii is distinct from the rest of its
genus, as are Raoulia cinerea and Raoulia species 1
(Ward 1981, 1993a, b; Breitwieser 1990, in press).
The isolation of Helichrysum lanceolatum in the
phenogram is an expression of its very distinctive
leaf anatomy (Breitwieser 1990, in press) and may
also reflect an incorrect taxonomic position.
Combined phenogram
Figure 7 shows associations based on all the
characters from both the leaf anatomy and the leaf
flavonoid analyses. Although this phenogram is not
as well structured as Fig. 6, it shows many of the
same features. The New Zealand species of
New Zealand Journal of Botany, 1993, Vol. 31
Solvent front
TS III
Anaphalis again form a very distinct group (G) as
do the whipcord species of Helichrysum (A), again
loosely associated with Raoulia petriensis. The
remaining species of Raoulia are not closely linked,
although, with the exception of the two aberrant
species R. cinerea and Raoulia species 1, they form
a loose cluster (B) which also includes Inuleae genus
1 and a subcluster (C) of the two alpine mat species
of Gnaphalium with the Australian species of
Ewartia.
The New Zealand species of Cassinia are again
allied with the Australian species of Helichrysum
sect. Ozothamnus and more distantly with the
Australian species of Cassinia (D). Again the
Himalayan species of Anaphalis associates with
Gnaphalium involucratum (E), and the two species
of Haas tia are paired (F). The final cluster (H) again
links the herbaceous species of Helichrysum with
Breitwieser & Ward—N.Z. Inuleae 53
similarity

Fig. 5 Phenogram of similarity among 45 species of Fig. 6 Phenogram of similarity among 45 species of
Inuleae, based on leaf flavonoid data. Taxa are abbreviated Inuleae, based on leaf anatomy data. Abbreviations are the
to the first three letters of the generic and specific names. same as for Fig. 5.

the broadleaved form of the scandent Helichrysum herbaceous species of Helichrysum (in the flavonoid
dimorphum, but with the addition of the aberrant analysis only). They are never associated with the
Raoulia species 1 and one of the stoloniferous Himalayan Anaphalis triplinervis. Further com-
species of Gnaphalium. parison with Anaphalis may confirm that the New
The isolated taxa in Fig. 7 are the same as those Zealand species should be recognised as the separate
in Fig. 6 except for the inclusion of Raoulia species genus Anaphalioides Kirp., with the possible
1 in a cluster and the isolation of another two species inclusion of Helichrysum bellidioides and H.
of Raoulia, R. glabra and & grandiflora, as well as filicaule.
the separation of the two species of Leucogenes both The four species Cassinia fulvida and C.
from each other and from all other taxa. leptophylla of New Zealand and Helichrysum (sect.
Ozothamnus) backhousei and H. obcordatum of
Australia are consistently associated*, and are the
only species with "group III" flavonoids. It is clear
DISCUSSION that the New Zealand species of Cassinia are closer
The genera of Gnaphaliinae occurring in New
Zealand are discussed here in terms of similarities
in leaf flavonoids and anatomy.
*The position of//, obcordatum in Fig. 6 is slightly aberrant
The New Zealand species of Anaphalis are linked due to the average clustering technique (see discussion in
together in all three analyses, but the only other taxa Ward 1993b), its highest similarity being in fact to C.
with which they are associated are the two leptophylla.
54 New Zealand Journal of Botany, 1993, Vol. 31

examined in possessing spot 16. The two non-

stoloniferous species, G. mackayi and G. nitidulum,
do not form a group separate from the other two
species.
A close relationship between Gnaphalium and
Australian Ewartia is suggested by their common
1 1c possession of spots of group II and lack of spots of
groups I and III. The exception is G. traversii with
' H
consistent spots present only in group I.
sP2 = 1 L Within Helichrysum the two species of the
exi
1 herbaceous sect. Xerochlaena, H. bellidioides and
1 1 H. filicaule, have leaves which are alike in both
flavonoids and anatomy. With regard to flavonoids,
their association with the New Zealand species of
Anaphalis suggests both groups may best be referred
to a single genus. Not only do these six species lack
any spots of groups II or III, but H. bellidioides and
A. keriensis are identical apart from quantitative
differences. However, such an amalgamation is not
supported by their leaf anatomy, in which H.
bellidioides and H. filicaule are quite unlike
Anaphalis and closest to two other species of
Helichrysum.
The whipcord species of Helichrysum sect.
Ozothamnus (H. coralloides, H. depressum, H.
intermedium, and H. parvifolium), together with the
whipcord form of//, dimorphum, comprise a clearly
Fig. 7 Phenogram of similarity among 45 species of delimited group in all three phenograms. The leaf
Inuleae, based on leaf flavonoid and leaf anatomy data. of the whipcord form of H. dimorphum is ana-
Abbreviations are the same as for Fig. 5. tomically very similar to that of//, depressum, while
the broadleaved form is most similar to H.
obcordatum and H. filicaule. However, a hybrid
origin of H. dimorphum from H. depressum and H.
to at least these Australian species of Helichrysum filicaule is not supported by the flavonoid analysis.
sect. Ozothamnus than they are to the Australian Far from showing the flavonoids of both these
species of Cassinia. species, as might be expected in a hybrid, H.
Within Ewartia no species are consistently dimorphum is virtually devoid of any flavonoids.
associated. The isolation of the only New Zealand The sole remaining New Zealand species of
species, E. sinclairii, is not surprising since it is Helichrysum sect. Ozothamnus, H. lanceolatum, is
known to be an anomaly in this genus (see above). the only taxon in this study which lacks spots in any
However, the diversity among the three Australian of the three groups. It possesses only spot 12, which
species indicates that a reappraisal of their position it shares with Cassinia, the Australian species of
in relation to other Australasian taxa is required. The Helichrysum sect. Ozothamnus, and Pterygopappus.
association of E. planchonii with Gnaphalium and In leaf anatomy, H. lanceolatum is distinct from all
particularly the identical flavonoid spots in this other taxa. Its affinities remain a mystery.
species and G. mackayi support the view that the The two Australian species of Helichrysum sect.
boundary between these two genera needs revision. Ozothamnus are closer to Cassinia than to
The leaves of the four species of Gnaphalium sect. Helichrysum of New Zealand, which itself seems to
Euchiton are not all similar in either their flavonoids, span several distinct genera.
in which G. traversii is quite separate with not one The most remarkable feature of Leucogenes is its
spot in common with any of the other species, or very large number of flavonoid spots nearly all of
their anatomy, in which G. involucratum differs which belong to groups I and II. L. leontopodium is
markedly. G. nitidulum is unique among all taxa particularly distinctive, with 17 spots, two of which
Breitwieser & Ward—N.Z. Inuleae
are unique to this species. Although the two species
differ in many of their spots, causing their separation
in Fig. 5, they share more spots (at least 7) than
almost any other taxa, being equalled only by L.
leontopodium and Raoulia grandiflora (7) and by
Cassinia fulvida and C. leptophylla (7). In leaf
anatomy, Leucogenes constitutes a distinct entity
with links to Raoulia.
Pseudognaphalium is distinct in both leaf
anatomy and flavonoids. [The distant link with
Ewartia sinclairii in Fig. 5 is an aberration caused
by the average clustering technique (see Ward
1993b). The two taxa have only two spots in
common.] All its flavonoids belong to spot group
II, a characteristic which it shares with most species
of Gnaphalium and Ewartia as well as with
Anaphalis triplinervis and three species of Raoulia.
In addition, it is notable for its large quantity of bright
yellow, fluorescent spots.
Although Raoulia has long been regarded as
heterogeneous, most species are similar in their leaf
anatomy. Exceptions are R. cinerea, R. petriensis,
and Raoulia species 1, each isolated not only from
the rest of Raoulia but from all other taxa as well.
Otherwise, the species of subg. Raoulia form a group
and the species of subg. Psychrophyton are quite
unified with the exception of ^?. grandiflora. Ewartia
catipes, formerly included in Raoulia (Hooker 1860),
strongly resembles these species in leaf anatomy.
Raoulia is much more diverse in its flavonoids,
the only associations being two pairs of species (R.
hookeri with R. tenuicaulis in subg. Raoulia and R.
bryoides with Raoulia species 2 in subg.
Psychrophyton), and a grouping of two of the
anatomically isolated species, ^?. cinerea and
Raoulia species 1, with Gnaphalium traversii. Of
note is the presence of spot 13 combined with the
absence of spot 21, found in all three species of the
pulvinate group (R. bryoides, R. eximia, and Raoulia
species 2) but elsewhere only in Pterygopappus.
Raoulia grandiflora shares all nine of its spots with
one or both species of Leucogenes, with spot 30
restricted to these three species, thus providing a
further clue in the puzzle of the relationships among
the species of Leucogenes and Raoulia subg.
Psychrophyton (Ward 1981, 1993b).
The spread of the species of Raoulia in Fig. 7,
based on all leaf anatomy and flavonoid characters,
is similar to that found by Ward (1993b). In both
studies, R. cinerea and R. petriensis are isolated,
Raoulia species 1 is separated from the rest of the
genus, and R. eximia, R. bryoides, Raoulia species
2, and R. hectorii are grouped together, as are R.
55
hookeri and R. tenuicaulis. The relationships of R.
glabra and R. grandiflora remain somewhat
equivocal. The former again shows similarities to,
but also substantial differences from, Raoulia subg.
Raoulia, while the latter shows affinity both to
Raoulia subg. Psychrophyton and to Leucogenes.
The undescribed Inuleae genus 1 is distinct in leaf
anatomy, while in flavonoids it is similar to Raoulia
hookeri and R. tenuicaulis. A large number of spots
confined to groups I and II is found only in these
three species and in R. petriensis and Leucogenes
grandiceps. Inuleae genus 1 has been likened to
Haastia sinclairii and to L. grandiceps (Allan 1961),
but it is quite unlike these taxa in leaf anatomy and
unlike H. sinclairii in its flavonoids. Overall (Fig.
7) its affinities seem to be with Raoulia.
The correct tribal position of Haastia is not
resolved here because, while it is similar to other
genera of the Inuleae, we have no data for
comparison with genera of the tribe Astereae to
which Haastia has traditionally been referred.
However, the presence of secretory canals in the
leaves of H. pulvinaris (Breitwieser 1990, in press)
militates against its inclusion in the subtribe
Gnaphaliineae where this feature is, as far as we
know, unrecorded. Haastia pulvinaris and H.
sinclairii are clearly more similar to each other than
to any other taxa included in this study, in spite of
the suggestion by Merxmiiller et al. (1977) that they
are not congeneric. As well as being paired in all
three phenograms, they alone possess the flavonoid
spot 15.
The endemic, monotypic Tasmanian genus
Pterygopappus is also distinct in both leaf anatomy
and flavonoids. Apart from one spot in group I, its
spots all lie outside the three main groups, with spot
38 being unique. No support is found for a taxonomic
position near Haastia pulvinaris, as was tentatively
suggested by Merxmiiller et al. (1977).
In conclusion, results of this study confirm that
generic boundaries in New Zealand Inuleae need
revision. Leucogenes seems a satisfactory genus and
in both Pseudognaphalium and Inuleae genus 1 the
single New Zealand species is quite distinct from
other taxa examined. However, Haastia appears
misplaced in the Gnaphaliinae. In Anaphalis,
Cassinia, and Ewartia the New Zealand species may
be in the wrong genus, and the lack of unity in
Gnaphalium and its links to Australian species of
Ewartia indicate that it too would repay further
study. Helichrysum and Raoulia are heterogeneous
and their need of revision (now in progress) is
evident.
56
ACKNOWLEDGMENTS
This paper represents part of a dissertation submitted by
the first author in partial fulfilment of the requirements of
a Ph.D. degree at the University of Canterbury, New
Zealand, and made possible by financial support in the form
of a scholarship from the Konrad-Adenauer-Stiftung,
Institut fur Begabtenfiirderung, Germany, and a grant
provided by the Miss E. L. Hellaby Trust, New Zealand.
We wish to thank John Lovis and David Lloyd for criticising
the manuscript, John Walker and John McCallum for advice
on chemical methods, and Rainer Vogt for help with
fieldwork and preparation of the figures.
REFERENCES
Adams, R. P. 1974: II. On "Numerical chemotaxonomy"
revisited. Taxon 23: 336-338.
Allan, H. H. 1961: Flora of New Zealand. Vol. I.
Wellington, Government Printer.
Anderberg, A. A. 1989: Phytogeny and reclassification of
the tribe Inuleae (Asteraceae). Canadian journal
of botany 67: 2277-2296.
Anderberg, A. A. 1991: Taxonomy and phytogeny of the
tribe Gnaphalieae (Asteraceae). Opera Botanica
104: 1-195.
Beauverd, G. 1910: Contribution a l'etude des Composees.
IV: Recherches sur la tribu des Gnaphaliees.
Bulletin de la Societe Botanique de Geneve sir.
2, 2: 207-252.
Beauverd, G. 1912: Contribution a l'etude des Composees.
VI: Nouveaux Leontopodium et Raoulia. Bulletin
de la Societe Botanique de Geneve ser. 2, 4: 12-
55.
Bentham, G. 1873a: Notes on the classification, history
and geographical distribution of Compositae.
Botanical journal of the Linnean Society 13: 335-
577.
Bentham, G. 1873b: Compositae. In: Bentham, G.;
Hooker, J. D. Genera Plantarum Vol. II. London,
Reeve.
Breitwieser, I. 1990: Leaf anatomy and chemotaxonomy
in Gnaphaliinae (Inuleae-Compositae).
Unpublished Ph.D. thesis, University of
Canterbury, New Zealand.
Breitwieser, I. in press: Comparative leaf anatomy of
New Zealand and Tasmanian Inuleae
(Compositae). Botanical journal of the Linnean
Society.
Bremer, K.; Jansen, R. K.; Karis, P. O.; Kim, K. J. 1992:
Current research on Asteraceae phytogeny and
classification. Compositae newsletter20/21: 1-3.
Cheeseman, T. F. 1906: Manual of the New Zealand
flora. Wellington, Government Printer.
New Zealand Journal of Botany, 1993, Vol. 31
Cheeseman, T. F. 1925: Manual of the New Zealand
flora. 2nd ed. Wellington, Government Printer.
Curtis, W. M. 1963: The student's flora of Tasmania.
Vol. II. Tasmania, Government Printer.
Drury, D. G. 1970: A fresh approach to the classification
of the genus Gnaphalium with particular reference
to the species present in New Zealand (Inuleae-
Compositae). New Zealand journal of botany 8:
222-248.
Drury, D. G. 1971: The American spicate cudweeds
adventive to New Zealand: (Gnaphalium section
Gamoc/iaeta-Compositae). New Zealand journal
ofbotany 9: 157-185.
Gower, J. C. 1971: A general coefficient of similarity and
some of its properties. Biometrics 27: 857-871.
Harborne, J. B. 1975: Biochemical systematics of the
flavonoids. In: Harborne, J. B.; Mabry, T. J.;
Mabry, H. ed. The flavonoids. London, Chapman
& Hall.
Harborne, J. B.; Turner, B. L. 1984: Plant
chemosystematics. New York, Academic Press.
Hilliard, O. M.; Burtt, B. L. 1973: Notes on some plants
of southern Africa chiefly from Natal: 3. Notes
from the Royal Botanic Garden Edinburgh 32:
303-387.
Hilliard, O. M.; Burtt, B. L. 1981: Some generic concepts
in Compositae-Gnaphaliinae. Botanical journal
of the Linnean Society 82: 181-232.
Hooker, J. D. 1860: The botany of the antarctic voyage of
H.M. discovery ships Erebus and Terror, in the
years 1839-1843, under the command of Captain
Sir James Clark Ross. Vol. III. Flora Tasmaniae
pt 1. London, Reeve.
Hooker, J. D. 1864: Handbook of the New Zealand flora.
London, Reeve.
Kirk, T. 1899: The students' flora of New Zealand.
Wellington, Government Printer.
Kirpichnikov, M. E.; Kuprijanova, L. A. 1950:
Morphological-geographical and palynological
contributions to the understanding of the genera
of the subtribe Gnaphaliinae. Acta Instituti
Botanici Academiae Scientiarum URSS, series 1,
9: 7-37.
Mabry, T. J.; Markham, K. R.; Thomas, M. B. 1970: The
systematic identification of flavonoids. Berlin,
Springer Verlag.
Markham, K. R. 1982: Techniques of flavonoid
identification. London, Academic Press.
Merxmuller, H.; Leins, P.; Roessler, H. 1977: Inuleae—
systematic review. In: Heywood, V. H.; Harborne,
J. B.; Turner, B. L. ed. The biology and chemistry
of the Compositae. London, Academic Press.
Rohlf, F. J. 1970: Adaptive hierarchical clustering
schemes. Systematic zoology 19: 58-82.
Breitwieser & Ward—N.Z. Inuleae
Sneath, P. H. A.; Sokal, R. R. 1973: Numerical taxonomy.
San Francisco, W. H. Freeman.
Sokal, R. R.; Michener, C. D. 1958: A statistical method
of evaluating systematic relationships. University
of Kansas science bulletin 38: 1409-1438.
Wall, A. 1920: Helichrysum dimorphum Cockayne—a
hybrid? Transactions of the New Zealand Institute
52: 106-107.
Ward, J. M. 1981: Numerical phenetics and the
classification of Raoulia (Gnaphaliinae-
Compositae). Unpublished Ph.D. thesis,
University of Canterbury, New Zealand.
Ward, J. M. 1982a: A key, synopsis and concordance for
Raoulia. Mauri Ora 10: 11-19.
Ward, J. M. 1982b: A field guide to Raoulia. Canterbury
Botanical Society journal 16: 33-41.
Appendix 1 Characters used in the numerical
phenetic analysis of leaf anatomy.
1. Cuticle thickness, adaxial cf. abaxial thickness: (a)
thicker (b) thinner (c) equal. 2. Cuticle thickness, margin:
(a) thickened (b) not thickened. 3. Cuticle thickness,
midrib: (a) thickened (b) not thickened. 4. Epidermis,
adaxial cf. abaxial thickness: (a) thicker (b) thinner (c)
equal. 5. Epidermis, average cell height of adaxial
epidermis: mm. 6. Epidermis, average cell height of
abaxial epidermis: mm. 7. Epidermis, cell shape: (a) wider
than high (b) higher than wide. 8. Epidermis, adaxial
epidermal cell size: (a) uniform (b) not uniform. 9.
Epidermis, abaxial epidermal cell size: (a) uniform (b) not
uniform. 10. Epidermis, cell size in abaxial midrib: (a)
higher than the other abaxial epidermal cells (b) same size
as the other abaxial epidermal cells. 11. Epidermis, cell
modifications in adaxial midrib: (a) none (b) 4 narrower
cells (c) more than 4 narrower cells. 12. Epidermis,
occasional interstitial cells: (a) absent (b) present. 13.
Epidermis, one taller cell in adaxial midrib: (a) absent (b)
present. 14. Stomata, arrangement: (a) only adaxial (b)
more on adaxial side (c) equal numbers on both sides (d)
more on abaxial side (e) only abaxial. 15. Stomata,
position: (a) guard cells level with unspecialised epidermal
cells (b) guard cells raised (c) guard cells and one adjoining
cell raised (d) guard cells and two or more adjoining cells
raised. 16. Stomata, size of substomatal chambers: (a) less
than half the leaf width (b) half the leaf width (c) more
than half the leaf width. 17. Mesophyll, spongy
chlorenchyma: (a) present (b) absent. 18. Mesophyll, shape
of spongy chlorenchyma cells: (a) different to palisade
cells (b) similar to palisade cells. 19. Mesophyll, large
central clear cells: (a) present (b) absent. 20. Mesophyll,
arrangement: (a) almost homogeneous (b) not
homogeneous. 21. Mesophyll, if not homogeneous: (a)
57
Ward, J. M. 1993a: Systematics of New Zealand Inuleae
(Compositae-Asteraceae)—1. A numerical
phenetic study of the species of Raoulia. New
Zealand journal of botany 31: 21-28.
Ward, J. M. 1993b: Systematics of New Zealand Inuleae
(Compositae-Asteraceae)—2. A numerical
phenetic study of Raoulia in relation to allied
genera. New Zealand journal of botany 31: 29-
42.
Webb, C. J. 1987: In: Connor, H. E.; Edgar, E. Name
changes in the indigenous New Zealand flora,
1960-1986 and nomina nova IV, 1983-1986. New
Zealand journal of botany 25: 115-170.
Webb, C. J. 1988: Asteraceae: Inuleae. In: Webb, C. J.;
Sykes, W. R.; Garnock-Jones, P. J. Flora of New
Zealand. Vol. IV. Christchurch, New Zealand,
Botany Division, Department of Scientific and
Industrial Research.
poorly differentiated (b) well differentiated. 22.
Mesophyll, if poorly differentiated: (a) oval/round/oval
cells distinguishable (b) oval/round cells distinguishable.
23. Mesophyll, arrangement of palisade chlorenchyma:
(a) only abaxial (b) not only abaxial. 24. Mesophyll,
arrangement of palisade chlorenchyma: (a) equally on both
sides (b) not equally on both sides. 25. Mesophyll, small
to medium central clear cells: (a) present (b) absent. 26.
Mesophyll, small round cells next to abaxial epidermis:
(a) present (b) absent. 27. Mesophyll, palisade cell shape:
(a) oblong (b) oval. 28. Mesophyll, palisade cells next to
epidermis: length/width ratio. 29. Mesophyll, length of
palisade cells next to epidermis: mm. 30. Mesophyll,
arrangement: (a) dorsiventral (b) equifacial. 31.
Mesophyll, thickness: palisade/spongy chlorenchyma
ratio. 32. Mesophyll, arrangement at the leaf margin: (a)
at one side only (b) around the peripheries (c) absent. 33.
Midrib, protrusion: mm. 34. Midvein, bundle-sheath:
number of cell layers. 35. Midrib, palisade cells in midrib:
(a) absent (b) present. 36. Midrib, abaxial collenchyma:
(a) present (b) absent. 37. Midvein, sclerenchyma caps:
(a) present (b) absent. 38. Midvein, sclerenchyma caps if
present: (a) adaxial (b) abaxial (c) adaxial and abaxial.
39. Midrib, shape on the non-protruding side: (a) straight
(b) not straight. 40. Midrib, sclerenchymatous cells in
mesophyll: (a) present (b) absent. 41. Midvein, bundle-
sheath: (a) sclerenchymatous (b) not sclerenchymatous.
42. Lateral ribs, protrusion: (a) present (b) absent. 43.
Lateral ribs, palisade cells: (a) absent (b) present. 44.
Lateral ribs, bundle-sheath extension: (a) present (b)
absent. 45. Lateral ribs, abaxial collenchyma: (a) present
(b) absent. 46. Leaf, outline: (a) normal (b) crenulate (c)
scale-like (d) needle-like. 47. Secretory canals: (a) present
(b) absent. 48. Spaces between epidermis and mesophyll:
(a) present (b) absent. 49. Sclerenchymatous cells in
mesophyll: (a) present (b) absent.
58 New Zealand Journal of Botany, 1993, Vol. 31

Appendix 2 Material examined. (Hook, f.) Benth. et Hook, f.: /. Breitwieser & R.W. Vogt
776 (A); /. Breitwieser & R. W. Vogt 765 (A); /. Breitwieser
The voucher specimens of material examined are listed & R.W. Vogt 637 (F); /. Breitwieser & R.W. Vogt 619 (A
below. Numbers are lodged at CANU. (A) = leaf anatomy, & F). H. dimorphum Ckn.: /. Breitwieser & R.W. Vogt
(F) = flavonoids, (A & F) = leaf anatomy and flavonoids. 820 (A); /. Breitwieser & R.W. Vogt 770 (A & F). H.
intermedium Simpson: /. Breitwieser & R.W. Vogt 803
Anaphalis keriensis (Cunn.) C. Webb: /. Breitwieser & (A); /. Breitwieser & R.W. Vogt 542 (F); /. Breitwieser &
R.W. Vogt 827 (A & F); /. Breitwieser & R.W. Vogt 887 R.W. Vogt 825 (A); /. Breitwieser & R.W. Vogt 817 (A).
(A). A. rupestris C. Webb: Ward 88364 (A & F). A. H. filicaule Hook, f.: /. Breitwieser & R.W. Vogt 769 (A
subrigida (Colenso) C. Webb: /. Breitwieser & R. W. Vogt & F); /. Breitwieser & R.W. Vogt 805 (A); /. Breitwieser
861 (F); /. Breitwieser & R.W. Vogt 862 (A & F); /. & R.W. Vogt 890 (F); Ward 84012 (A). H. lanceolatum
Breitwieser & R.W. Vogt 898 (A). A. trinervis (Forst. f.) (Buchanan) Kirk: /. Breitwieser & R.W. Vogt 809 (A); /.
F. Muell.: /. Breitwieser & R.W. Vogt 544 (A & F); /. Breitwieser & R.W. Vogt 781 (A & F). H. obcordatum
Breitwieser & R.W. Vogt 597 (A); /. Breitwieser & R.W. (DC.) F. Muell. ex Benth.: /. Breitwieser & R.W. Vogt
Vogt 653 (F). A. triplinervis (Sims) S.B. Clarke: /. 675 (A); /. Breitwieser & R.W. Vogt 730 (A & F). H.
Breitwieser & R.W. Vogt 836 (A & F). Cassinia aculeata parvifolium Yeo: /. Breitwieser & R.W. Vogt 789 (A &
R. Br.: /. Breitwieser & R. W. Vogt 673 (A); /. Breitwieser F). Leucogenes grandiceps (Hook, f.) Beauverd: /.
& R.W. Vogt 756 (A); /. Breitwieser & R.W. Vogt 710 Breitwieser & R.W. Vogt 613 (A & F); /. Breitwieser &
(F); /. Breitwieser & R. W. Vogt 660 (A). C. fulvida Hook, R. W. Vogt 760 (A). L. leqntopodium (Hook, f.) Beauverd:
f.: /. Breitwieser & R.W. Vogt 826 (A); /. Breitwieser & /. Breitwieser & R.W. Vogt 847 (A & F).
R.W. Vogt 828 (A & F); /. Breitwieser & R.W. Vogt 584 Pseudognaphalium luteoalbum (L.) Hilliard et B.L. Burtt:
(A & F). C. leptophylla (Forst. f.) R. Br.: /. Breitwieser & /. Breitwieser & R. W. Vogt 891 (A); /. Breitwieser & R. W.
R. W. Vogt 845 (A); /. Breitwieser & R. W. Vogt 834 (F); /.Vogt 611 (F); /. Breitwieser & R.W. Vogt 600 (A & F).
Breitwieser & R.W. Vogt 784 (A). C. longifolia R. Br.: /. Pterygopappus lawrencei Hook, f.: /. Breitwieser & R.W.
Breitwieser & R.W. Vogt 725 (A & F). Ewartia catipes Vogt 705 (A); /. Breitwieser & R.W. Vogt 677 (F); /.
(DC.) Beauverd: /. Breitwieser & R.W. Vogt 724 (A & Breitwieser & R. W. Vogt 682 (A). Raoulia bryoides Hook,
F). E. meredithiae (F. Muell.) Beauverd: /. Breitwieser & f.: /. Breitwieser & R.W. Vogt 793 (A & F); /. Breitwieser
R. W. Vogt 697 (A); /. Breitwieser & R. W. Vogt 670 (A && R. W. Vogt 800 (A). R. cinerea Petrie: /. Breitwieser &
F). E. planchonii (Hook. F.): /. Breitwieser & R.W. Vogt R.W. Vogt 922 (A & F); Ward 88265 (A & F). R. eximia
738 (A & F); /. Breitwieser & R.W. Vogt 708 (A). E. Hook, f.: /. Breitwieser & R.W. Vogt 831 (A); /.
sinclairii (Hook, f.) Cheesem.: /. Breitwieser & R.W. Vogt Breitwieser & R.W. Vogt 772 (A & F). R. glabra Hook.
786 (A & F); /. Breitwieser & R.W. Vogt 792 (A). f.: /. Breitwieser & R. W. Vogt 546 (A & F); /. Breitwieser
Gnaphalium involucratum Forst. f: /. Breitwieser & R. W. & R.W. Vogt 808 (A); /. Breitwieser & R.W. Vogt 883
Vogt 824 (A & F). G. mackayi (Buchanan) Ckn: /. (F); /. Breitwieser & R.W. Vogt 766 (A). R. grandiflora
Breitwieser & R.W. Vogt 572 (F); /. Breitwieser & R.W. Hook, f.: /. Breitwieser & R. W. Vogt 586 (F); /. Breitwieser
Vogt 810 (A); /. Breitwieser & R.W. Vogt 833 (A). G. & R. W. Vogt 571 (F); /. Breitwieser & R. W. Vogt 799 (A);
nitidulum Hook, f.: /. Breitwieser & R.W. Vogt 638 (A); /. Breitwieser & R. W. Vogt 830 (A); /. Breitwieser & R. W.
/. Breitwieser & R.W. Vogt 811 (A). G. traversii Hook, f.: Vogt 737 (A). R. hectorii Hook, f.: /. Breitwieser & R. W.
/. Breitwieser & R. W. Vogt 801 (A); /. Breitwieser & R. W. Vogt 581 (A); /. Breitwieser & R.W. Vogt 614 (F); /.
Vogt 779 (A); /. Breitwieser & R. W. Vogt 589 (F). Haastia Breitwieser & R.W. Vogt 566 (A). R. hookeri Allan: /.
pulvinaris Hook, f.: /. Breitwieser & R. W. Vogt 905 (A & Breitwieser & R.W. Vogt 821 (A); /. Breitwieser & R.W.
F); /. Breitwieser & R.W. Vogt 909 (A & F). H. sinclairii Vogt 768 (A); /. Breitwieser & R.W. Vogt 654 (F); /.
Hook, f.: /. Breitwieser & R.W. Vogt 906 (A & F). Breitwieser & R.W. Vogt 610 (F); /. Breitwieser & R.W.
Helichrysum backhousei (Hook, f.) F. Muell. ex Benth.: Vogt 629 (F); /. Breitwieser & R.W. Vogt 806 (A). R.
/. Breitwieser & R. W. Vogt 743 (A); /. Breitwieser & R. W.species 2: /. Breitwieser & R.W. Vogt 580 (A & F). R.
Vogt 669 (A); /. Breitwieser & R.W. Vogt 723 (A); /. species 1: /. Breitwieser & R.W. Vogt 829 (A); /.
Breitwieser & R.W. Vogt 688 (F); /. Breitwieser & R.W. Breitwieser & R.W. Vogt 771 (A & F). R. petriensis Kirk:
Vogt 698 (A & F). H. bellidioides (Forst. f.) Willd.: /. /. Breitwieser & R.W. Vogt 612 (A & F). R. tenuicaulis
Breitwieser & R.W. Vogt 547 (A & F); /. Breitwieser & Hook, f.: /. Breitwieser & R. W. Vogt 592 (F); /. Breitwieser
R. W. Vogt 548 (F); /. Breitwieser & R. W. Vogt 577 (F); /. & R. W. Vogt 607 (F); /. Breitwieser & R. W. Vogt 802 (A);
Breitwieser & R.W. Vogt 587 (A & F); /. Breitwieser & /. Breitwieser & R. W. Vogt 822 (A); /. Breitwieser & R. W.
R.W. Vogt 813 (A); /. Breitwieser & R.W. Vogt 631 (A). Vogt 761 (A); /. Breitwieser & R.W. Vogt 609 (A & F).
H. coralloides (Hook, f.) Benth. et Hook, f.: /. Breitwieser Genus " 1 " : Ward 89097 (A); /. Breitwieser & R.W. Vogt
& R. W. Vogt 787(A & F); Ward88158 (A). H. depressum 795 (A & F); /. Breitwieser & R.W. Vogt 907 (F).