Botanical Journal of the Linnean Socieb (1988), 98: 27-36. With 3 figures
A classification of the genus Nothofagus
(Fagaceae)
WILLIAM R. PHILIPSON, F.L.S.
Department of Botany, University of Canterbury, .New zealand
AND
MELVA N. PHILIPSON
Botany Division, DSIR, Private Bag, Christchurch, .New zealand
Received February 1987, revised and accepted for publication December 1987
PHILIPSON, W. R. & PHILIPSON, M. N., 1988. A clesaificadon of the genus N o t b f a g w
(hg8ce8e). The genus Nothofagus is subdivided into two subgenera, two sections and three
subsections, employing characters of the pollen, cupule and vernation, together with the incidence
of leaf-fall. The subdivision into subgenera also agrees with present geographical distribution and
with characters of the secondary xylem. One subgeneric and one subsectional name are new.
Reasons are given for departures from previous classifications.
ADDITIONAL KEY WORDS:-Cladistics
~
biogeography.
CONTEN’I’S
Introduction . . . . . . . . .
A synopsis of the subgeneric taxa of Nothofagus .
Acknowledgements
. . . . . . .
References. . . . . . . . . .
. . . . . . . . . .
. . . . . . . . . .
. . . . . . . . . .
. . . . . . . . . .
27
33
35
35
INTRODUCTION
In a previous paper we reported the occurrence of several types of leaf
vernation in Nothofagus and drew attention to correlations between the different
types and other characters within the genus, including their geographical
distribution (Philipson & Philipson, 1979). The taxonomic significance of these
correlations was referred to, but at the time we did not wish to propose formal
amendments to the existing classification of the species. In view of some
favourable responses to the changes suggested in that paper (e.g. Wardle,
1984: 17; Tomlinson, 1984: 55) it is now considered appropriate to present our
ideas formally.
The principal difference between the classification proposed here and that of
van Steenis (1953, 1972), is that the primary division of the genus adopted is
0024-4074/88/090027
+ 10 $03.00/0
27
0 1988 The Linnean Society of London
28
W. R. PHILIPSON AND M. N. PHILIPSON
between the tropical species of New Guinea and New Caledonia, and the
temperate species of South America, Australia and New Zealand. van Steenis,
on the other hand, emphasized the distinction between the deciduous, plicate
species and the remainder of the genus. Our reason for rejecting his
arrangement is that the tropical species stand apart from the remainder of the
genus in the possession of brassii-type pollen, in the characters of the secondary
xylem (Dadswell & Ingle, 1954; Patel, 1986) and in their vernation. I n addition
they all bear a 2-partite cupule, although this character does occur in one of the
temperate species (Nothofagus pumilio). The species with deciduous leaves and
plicate vernation also form a natural group, but they do not appear to be so
distinct from the temperate evergreen species as we have seen the tropical
species to be. A number of characters are either shared by virtually all
temperate species whether deciduous or evergreen (e.g. the 4-partite cupule and
the short apertures of the pollen) or they vary similarly in both the deciduous
and evergreen groups (e.g. fusca and rnenciesii types of pollen). For these reasons
the category of subgenus has been introduced to accommodate the tropical
species whereas the distribution between the temperate deciduous and the
temperate evergreen species has been retained at the sectional level.
A second significant amendment to the former classification concerns our
treatment of N . solandri (Hook. fil.) Oerst. van Steenis emphasized the tripartite
cupule by placing this species (together with N . clzfortioides (Hook. fil.) Oerst.,
now considered synonymous) in a separate subsection, but we prefer to unite it
in a subsection with the two other New Zealand species with revolute vernation,
namely, N . fusca (Hook. fil.) Oerst. and N . truncata (Colenso) Cockayne. We
recognize two other subsections among the temperate evergreen species. Both
have vernation of the plane type but differ in their geography and in the
character of the appendages on their cupules. They also differ in their pollen
types, but as the distinction between the mentiesii and fusca types of pollen is
slight, this may be of little significance. Subsection Menriesiae comprises three
Australasian species with meneiesii-type pollen and distinctive glandular-tipped
processes on the cupule, while subsection Quadripartitae comprises three South
American species with fusca-type pollen and cupular lamellae with simple toothlike appendages. For nomenclatural reasons two of these subsections must bear
the inappropriate names Tripartite and Quadripartitae.
In view of their long isolation it might be expected that the two populations
of tropical species, in New Guinea and New Caledonia respectively, could be
distinguished by some morphological features which would justify their
recognition as distinct taxa within the subgenus Brassospora. An interesting
difference in phyllotaxis reported by van Steenis (1971) might be useful for this
purpose, but the evidence is inadequate at present. According to van Steenis the
phyllotaxis of the New Caledonian species is partly spiral and partly distichous,
whereas the New Guinean species have a strictly distichous phyllotaxis. van
Steenis himself records that seedlings of a New Guinean species ( N . rubra) have
spiral phyllotaxis on their primary axis. The four New Zealand species also have
primary seedling axes with spiral phyllotaxis, and their distichous lateral
branches occasionally revert to spiral phyllotaxis (Philipson, 1988). van Steenis
does not indicate how the two types of phyllotaxis reported for the New
Caledonian species are distributed nor their relative abundance, but it is evident
that the presence of two types of phyllotaxis is not confined to New Caledonia,
CLASSIFICATION OF NOTHOFAGUS
29
though spiral phyllotaxis may be more prevalent there than elsewhere. Since the
distinction between the New Caledonian and the New Guinean species may be
one of degree only, we have avoided the creation of sections in subgenus
Brassospora, though this may well become possible with more information.
Modifications to the van Steenis classification have been proposed informally
by other authors and these must now be considered. Melville (1973) published a
phylogenetic tree of the genus based “on all available characters”.
Unfortunately, he does not explain how these characters relate to his
dendrogram, but his subdivisions of the temperate evergreen species appear to
us less natural than those in the key published by van Steenis (1953) so that his
scheme cannot be regarded as an improvement. Melville’s diagram illustrating
the evolution of cupule and nutlet characters is more explicit. For example, he
derives the 2-partite cupule of the tropical species from the 4-partite condition
by way of the 3-partite cupule of N. solandri. This progression overlooks the
features of N. solandri (wood, vernation, pollen) which preclude its acceptance as
a stage in the evolution of the 2-partite group of species.
Humphries ( 1981, 1983) published a cladogram for species of Nothofagus
which corresponds closely with the proposals made by us in 1979 and now
formalized. The first dichotomy in the scheme proposed by Humphries
corresponds to our division of the genus into two subgenera, and the temperate
subgenus is again divided by a dichotomy into deciduous and evergreen
branches corresponding to our two sections.
It is in the evergreen clade that slight differences occur between our scheme
and that of Humphries. The branching points here are shown as a trichotomy
rather than two dichotomies. This difficulty could have been resolved using the
evidence of vernation. The three species with revolute vernation would have
split off from the remaining six temperate evergreen species, all of which have
plane vernation. These six species would, in turn, become resolved into two
groups of three species, each, distinguished by geography, pollen type and the
nature of the appendages of their cupules. However, in a later paper
incorporating additional data (Humphries, Cox & Nielsen, 1986) the phylogeny
of Nothofagus is related to those of a number of its parasites. The cladogram of
Nothofagus which showed maximal fit with the parasite groups gives a
satisfactory separation of the temperate evergreen species into a revolute group
fusca, truncata and solandri) and two, more closely associated, groups with plane
vernation (betuloides, nitida dombeyilmoorei, cunninghamii, menriesii).
Wardle (1984) modifies the scheme of Humphries by including the evidence
of vernation. His fig. 1.1 gives the clearest arrangement published at that time,
but it retains the principal division between deciduous and evergreen species.
Tanai (1986) studied venation of extant species of Nothofagus in order to
facilitate his examination of fossil leaves. He subdivides the venation pattern
into five groups to which he provides a key. The groups are congruent with the
infrageneric taxa proposed by us, and he states that the tropical species are
pronouncedly different from the temperate groups. I t is, therefore, surprising
that he proposes a phylogenetic relationship that shows the tropical species (our
subgenus Brassospora) as a sister group of the species included in our subsection
menriesii. This requires that the brassii type pollen of the living species to be
unrelated to the early fossil brassii pollen type. With this one exception, the
phylogenetic scheme of Tanai conforms closely to ours, though separating our
W. R. PHILIPSON AND M. N. PHILIPSON
30
section I
.--------------Nafhafagus
(temperate South
America, Tasmania)
subsection I Quadrlparflfae
(temperate South
America)
subsection 2 Menzies/ae
(Australia, New
Zealand l
subQenus I
Nofhofagus
section 2
Calusparassus
subsection 3 Triparfifae
(New Zealand)
L..,_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
subgenus 2
Brassosporo
(New Guinea,
New Caledonia)
Figure I. Diagram of relationships between the various categories of taxa within tht· genus
.Nothofagus.
subsections Q.uadripartitae and Men;:,iesii more widely than we consider
appropriate in view of their similar vernation.
The inter-relations of the various taxa within the genus Nothofagus are
summarized in Fig. I.
A character state matrix is given in Table 2 using the characters set out in
Table 1, and this information forms the basis of the cladograms in Figs 2 and 3.
The outgroup of Nothofagus is considered to be the genus Fagus as this is the only
other genus in the subfamily Fagoideae. Nothtifagus was raised to family status by
Kuprianova ( 1962), and several recent authors have supported this separation
from Fagus (Nixon, 1984;Jones, 1986; Romero, 1986; Zavada & Dilcher, 1986),
though it has been adopted by neither Takhtajan (1987) nor Cronquist (1981).
As the distinctiveness of Nothtifagus pollen has played a leading role in this
separation, the occurrence of very distinct pollen types within the genus
Trigonobalanus should be borne in mind (Forman, 1964). The retention of Fagus
and Nothtifagus in the same family is supported by their seedling morphology
TABLE
I.
2.
3.
4.
5.
6.
7.
8.
9.
Habit
Vernation
Vernation
Pollen
Pollen apperturcs
Cupule
J Infl.
Terminal <jl flower
Cupule lamellae
l. Characters used in the cladograms in Figs 2 and 3
Primitive
Advanced
Deciduous
Plicate
Plicate
3-Colpate
Long
4-Partite
Few-flowered
Present
Toothed, fringed or entire
Evergreen
Conduplicate
Plane/revolute
5-8-Colporate
Short
2-Partite
IX- Flowered
Absent
With glandular processes
CLASSIFICATION OF NOTHOFAGUS
31
TABLE
2. Character state matrix (see Table 1 for the characters)
Subgenus Nothofagus section Nothofagus
Section Calusparassus subsection Quadripartitae
Subsection Menziesiae
Subsection Tripartitne
Subgenus Brassospora
Fagus
1
2
3
4
5
6
7
8
9
0
1
1
1
1
0
0
0
-
1
1
1
1
1
0
0
1
1
0
0
0
0
l
l
l
0
I
O
l
1
1
1
0
1
0
0
0
1
0
1
1
0
0
0
1
0
0
0
1
0
0
which links these two genera (Philipson, 1988) in contrast to the seedlings of the
other two subfamilies of the Fagaceae (sensu lato). Fagus and Nothofagus have
been regarded as sister groups (e.g. Humphries, 1981; Humphries & Parenti,
1986) and this would remain valid even if their status were changed. Of the
subgeneric taxa within Nothofagus, section Nothofagus resembles Fagus most
closely as it shares with it the deciduous habit with plicate vernation and a 4partite cupule. Consequently these character states are regarded as primitive
within Nothofagus. In using pollen grain characteristics it is considered that the
distinction between brassii (long aperture) type grains and non-brassii (short
aperture) type (i.e. fusca and menriesii types combined) is more significant than
the difference between the fusca and menziesii types. Evolution of leaf vernation is
considered to have taken two distinct directions, (1) from plicate to the
conduplicate condition found in subgenus Brassospora, and (2) from plicate to
the plane condition in subsections Quadripartitae and Menciesiae leading to the
morphologically more extreme and geographically restricted revolute vernation
of subsection Tripartitae. I n considering the distinction of 4-partite and 2-partite
cupules which are so distinctive of the two subgenera, the occurrence of
section
No?hoiagus
subsection
OwdriprN?or
subsection
Manzirsiar
subsection
TriparN?ar
Figure 2. A cladogram of Nothofagus assuming that elongated pollen appertures have evolved in
parallel (see Table I for key to characters).
32
W. R. PHILIPSON AND M. N. PHILIPSON
Fagus
section
Nathofagus
su bsection
Ouadripartitae
su bsect ion
Menziesiae
subsect ion
Tripar titoc
subgenus
Brassospom
Figure 3. A cladograrn of Nothofagus assuming that the evergreen habit has evolved in parallel (see
Table 1 for key to characters).
2-partite cupules in Jv. pumilio and of 3-partite cupules in S. solandri is neglected,
as these conditions are considered to have been derived independently and must
be given no more weight than specific characters.
Both cladograms involve the parallel evolution of one character. In the first
(Fig. 2), it is assumed that the short apertures of the menziesii and fusca type
pollen grains have been acquired independently in sections Jvothofagus and
Calusparassus, while in the second (Fig. 3) it is assumed that the evergreen habit
has evolved independently in subgenus Brassospora and in section Calusparassus.
The latter is considered the most probable. It conforms best with the
classification in Fig. 1, which is derived on independent grounds, viz. the several
congruent characters from diverse parts of the plant which distinguish the
subgenus Brassospora. Further, changes in leaf-fall appear more probable than in
pollen morphology, since it is not uncommon to find deciduous and evergreen
members of the same genus. I t must also be borne in mind that, until more is
known about the evolution of Fagus, the alternative possibility that the ancestral
FaguslJvothofagus was evergreen cannot be dismissed. In that case, the evergreen
habit would not have evolved in parallel within Nothofagus. Instead the
deciduous habit would have arisen in parallel in Fagus and section Nothofagus. If
this should be the correct interpretation, the cladogram in Fig. 2 would be
favoured.
A comparison of Fig. 3 with the cladogram published by Humphries (1981:
fig. 15) reveals a very general similarity, but with four principal differences.
Firstly, N. solandri and N. clijfortioides were separated by Humphries on the basis
of their usually 3-partite cupule, whereas here they are united with 3.fusca and
N . truncata to form subsection Tr@artitae on the basis of their unique vernation.
It would be possible to recognize further subdivision of subsection Tripartitae on
cupule characters, but this is not done in this treatment which considers only the
major groups of the species of Nothofagus. Secondly, Humphries shows no
CLASSIFICATION OF NOTHOFAGUS
33
separation of the species here placed in subsections Quadripartitae and Tripartitae,
whereas in the present cladogram the Tripartitae are separated by their
distinctive vernation. Thirdly, subdivision within the South American species
and the tropical species which are shown in the cladogram of Humphries are
not adopted here. This is partly because the present classification does not take
subdivisions to this fine level and also because adequate synapomorphies have
not been established. Some of the groupings proposed by Humphries may not be
valid. For example, the reduced cupules of N. carrii and N. resinosu may have
been acquired independently, and the separation of N . perryi and N. nudu from
other New Guinean species, and of N . discoidea from the bulk of the New
Caledonian species is probably unwarranted. Fourthly, the synapomorphy used
by us to distinguish subgenus Nothofagus from subgenus Brassospora is the shorter
pollen apertures of the former. At the same point of his cladogram Humphries
uses the presence of tracheids in the xylem. As this is regarded by him as a
primitive condition, it is not a synapomorphy and cannot serve to establish the
unity of the species linked to this branch of his cladogram.
A SYNOPSIS OF THE SUBGENERIC TAXA OF NOTHOFAGUS
Nothofagus Blume, M u s . Bot. Lugd.-Bot., I : 307 (1850), nom. conserv.
Fagaster Spach, Hist. Nut. VLg. Phan., 11: 142 (1842), nom. re$.
Calucechinus Hombr. & Jacq. in Dumont d’urville, Voy. Pol. Sud. G’ Oc. Bot.
Atlas, Dicot. ( 1844), norn. rejic.
Calusparassus Hombr. & Jacq. in Dumont d’Unville, Voy. Pol. Sud. G’ Oc. Bot.
Atlas, Dicot. ( 1844), nom. rejic.
Lophozonia Turcz., Bull. Soc. Imp. N a t . Moscou, I : 396 (1858).
Trisyngyne Baill., Adansonia, 11: 136 (1873).
Subgenus 1. Nothofagus
Leaves mostly distichous, deciduous or evergreen; leaf vernation plicate, plane
or revolute; cupule 4-partite or rarely 3- or 2-partite; pollen of menziesii or fusca
tY Pe.
LECTOTYPE: N . antarctica (G. Forst.) Oerst. Sixteen species in southern Chile,
Argentina, Australia and New Zealand.
Section 1. Nothofagus
Calucechinus Hombr. & Jacq. in Dumont d’ Urville, Calucechinus Hombr. &
Jacq. in Dumont d’ Urville, Voy. Pol. Sud. 6’ Oc. Bot. Atlas, Dicot. (1844).
Lophozonia Turcz., Bull. Sot. Imp. &at. Moscou, I : 396 (1858).
Fagus section Eufagus A. DC. in DC. Prodr., 16(2): 117-123.
Fagus section Nothofagus Benth. & Hook., Gen. PI., 3: 410 (1880).
Nothofagus subgenus Lophozonia Krasser, Ann. Hofmus. Wien, 11: 162 ( 1896).
Nothofagus subgenus Molischia Krasser, Ann. Hofmus. Wien, 11: 162 (1896).
Nothofagus section Calucechinus (Hombr. & Jacq.) Krasser, Ann. Hofmus. Wien,
11: 163 (1896).
Nothofagus section Deciduae Steenis, Blumea, 7: 146 ( 1952).
Nothofagus section Plicatae Steenis, Blumea, 7: 306 (1952).
Nothofagus section Calucechinus subsection Antarcticae Steenis, J . A m . Arb., 34:
334 (1953).
Nothofagus section Calucechinus subsection Pumiliae Steenis, 3. A m . Arb., 34: 336
(1953).
34
W. R. PHILIPSON AND M. N. PHILIPSON
Leaves deciduous; leaf vernation plicate; cupule 4-partite (rarely 2-partite),
N . antarctica (G. Forst.) Oerst. Eight species, with seven species in
temperate South America and one in Tasmania:
N . alessandri Espinosa
N . gunnii (Hook. fil.) Oerst.
N . alpina (Poepp. & Endl.) Oerst.
N . obliqua (Mirb.) Oerst.
N. antarctica (Forst.) Oerst.
N . procera (Poepp. & Endl.) Oerst.
N . glauca (Phil.) Krasser
N . pumilio (Poepp. & Endl.) Krasser
TYPE:
Section 2. Calusparassus (Hombr. & Jacq.) Krasser, Ann. Hofmus. Wien, 11:
163 (1896).
Calusparassus'Hombr. & Jacq. in Dumont d' Urville, Voy. Pol. Sud. €8 Oc. Bot.
Atlas, Dicot. ( 1844).
Fagus subgenus Calusparassus Miq., Ann. Mus. Bot. Lugd-Bat., 1: 103 (1863).
Nothofagus section Sempervirentes Steenis, Blumea, 7: 146 ( 1952).
Nothofagus section Planae Steenis, Blumea, 7: 306 (1952).
Leaves evergreen; leaf vernation plane or revolute; cupule 4-partite (rarely 3partite).
LECTOTYPE: N . betuloides (Mirb.) Oerst. Nine species in Chile, Argentina,
Australia and New Zealand (see under subsections).
Subsection 1. Quadripartitae Steenis, 3. Am. Arb., 34: 337 (1953).
Leaf vernation plane; cupular lamellae with simple tooth-like appendages;
pollen fusca type.
N . betuloides (Mirb.) Oerst. Three species in temperate South
LECTOTYPE:
America:
N . betuloides (Mirb.) Oerst.
N. nitida (Phil.) Krasser
N . dombeyi (Mirb.) Oerst.
Subsection 2. Menxiesiae Philipson & Philipson, subsect. nov.
Nothofagus foliis sempervirentibus in gemma planis; cupulis processibus
glandulosis praeditis; polline mentiesii- typii.
Leaf vernation plane; cupule with glandular-tipped processes; pollen menziesii
t Y Pee
TYPE: N. menriesii (Hook. fil.) Oerst. Three species in Australia and New
Zealand:
N. cunninghamii (Hook. fil.) Oerst.
N . moorei (F. Muell.) Krasser
N. menziesii (Hook. fil.) Oerst.
Subsection 3. Tripartitae Steenis, 3. Am. Arb., 34: 338 (1953).
Leaf vernation revolute; cupule with transverse fringed or entire lamellae; pollen
fusca type.
TYPE:N . solandri (Hook. fil.) Oerst. Three species in New Zealand:
N . fusca (Hook. fil.) Oerst.
N. truncata (Colenso) Cockayne
N . solandri (Hook. fil.) Oerst.
Subgenus 2. Brassospora Philipson & Philipson, subgenus nov.
Trisyngyne Baill., Adansonia, 11: 136 (1873).
Nothofagus subsection Bipartitae Steenis, 3. Am. Arb., 34: 338 (1953).
Nothofagus foliis sempervirentibus in gemma conduplicatis; cupulis 2-partitis;
polline brassii-typi.
CLASSIFICATION O F N O T H O F A G U S
35
Leaves distichous or spirally arranged, evergreen; vernation conduplicate;
cupule 2-partite; pollen brassii type.
TYPE: N. brassii Steenis. About 18 species from New Guinea and New
Caledonia:
N. aequilateralis (Baum-Bod.) Steenis
N . grandis Steenis
N. balansae (Baill.) Steenis
N. nuda Steenis
N . baumanniae (Baum-Bod.) Steenis
N. perryi Steenis
N. brassii Steenis
N . pseudoresinosa Steenis
N. carrii Steenis
N . pullei Steenis
N . codonandra (Baill.) Steenis
N . resinosa Steenis
N . crenata Steenis
N. rubra Steenis
N. discoidea (Baum-Bod.) Steenis
N. starkenborghii Steenis
N. Jlaviramea Steenis
N . womersltyi Steenis
ACKNOWLEDGEMENTS
We wish to thank Professor David Lloyd of the University of Canterbury for
reading successive drafts and suggesting modifications; Dr Elizabeth Edgar and
Dr Philip Garnock-Jones of Botany Division, D.S.I.R., for assistance with
nomenclatural and cladistic problems; and Mr W. Simes of the Forest Research
Institute, Rangiora, for seed of Nothofagus.
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