New Zealand Journal of Botany. 1977, Vol. 15: 39-86.

39

A taxonomic revision of the New Zealand species of

Asplenium

P. J. BROWNSEY

Department of Botany, Victoria University of Wellington, Private Bag,

Wellington, New Zealand*

(Received 16 August 1976)

ABSTRACT

A taxonomic revision of the fern genus Asplenium in the New Zealand region has been
carried out. Fourteen species and four subspecies are recognised. One species, A. terrestre.
and three subspecies, A. obtusatum subsp. northlandicum, A. flaccidum subsp. haurakiensc,
and A. terrestre subsp. maritimum, are described for the first time. Typification, synonymy,
range of morphological variation, chromosome number, habitat, and distribution are described
for each taxon. A dichotomous key to the species is presented.
The origin and development of the New Zealand Asplenium flora is briefly discussed.
It has its closest affinities with that of temperate Australia from where most of the species
have probably been derived. However, some species such as A. trichomanes, A. polyodon,
and A. obtusatum have affinities with Northern Hemisphere, tropical, or South American
floras and have therefore ultimately reached New Zealand from much further afield.
The genus displays a higher mean level of ploidy (5.9) in this country than in any other
area for which records exist; of the 18 sexually reproducing cytotypes, 9 are tetraploid,
1 is hexaploid, and 8 are octoploid. Most of the tetraploid species have distributions extending
outside New Zealand and they are therefore considered ancestral. The higher polypoids are
almost all endemic and are thought to have originated within New Zealand from the tetraploid
taxa.

INTRODUCTION

Asplenium with over 700 species distributed
throughout the world is one of the largest genera
within the Filicopsida. The family Aspleniaceae is
believed to be of relatively recent origin and is still
in an active state of evolution with most of the
genera and many of the species rather poorly defined
(Lovis 1973, 1977). In recent years an intensive
cytogenetic and biosystematic investigation of the
European representatives of the family has resulted
in a much better understanding of the evolution and
taxonomy of this group of ferns. It is now known
that hybridisation and chromosomal doubling have
been of prime importance and that reticulate evolu-
tion through auto- and allopolyploidy has produced
several morphologically very similar but genetically
quite distinct species (see reviews by Vida 1972,
Lovis 1977).
Present address: National Museum, Private Bag,
Wellington, New Zealand.
To ascertain whether Asplenium has developed
in a comparable manner in other parts of the world,
and perhaps to provide some evidence for the geo-
graphic origin of this enormous genus, a similar
biosystematic investigation has been started on the
New Zealand representatives. It will be many years
before the evolution of these species is as clearly
documented as that of their European counterparts,
but the preliminary results of this investigation have
provided sufficient information for a much needed
taxonomic revision of the genus in New Zealand.
Many authors (e.g., Hooker 1855, Allan 1961) have
remarked on the plasticity and difficulties of defining
species of Asplenium in this country, but, apart from
an investigation by Knowlton (1957) which was
terminated by his premature death, and some
chromosome counts on individual species (Brownlic
1954, 1958, 1961), no detailed study of the genus
has been made in recent times. The present paper
presents new cytological and morphological evidence
which more clearly delimits the taxa.
40
MATERIALS AND METHODS
Plants were collected from throughout the New
Zealand region and grown in cultivation in Welling-
ton under approximately uniform environmental
conditions. Herbarium material was collected at the
same time and voucher specimens are deposited in
CHR, WELT, and WELTU.
Meiotic chromosome preparations were obtained
by fixing young sporangia and staining according
to the method described by Manton (1950, pp.
295-6). Mitotic preparations were obtained by the
method outlined in Roy & Manton (1965). Material
which has been examined cytologically is listed
immediately after the chromosome number given for
each taxon or cytotype.
Measurements of the exospore were made from
spores mounted in gum chloral (a sample of 50
spores was measured for each population). The range
of mean values for a specified number of populations
is given for each species and subspecies, the figures
in brackets indicating the extreme sizes of individual
spores. Rhizome scales have been drawn by camera
lucida from permanent preparations in Canada balsam
after dehydrating in a series of ethanols and clearing
in xylol.
Distribution maps covering the North, South, and
Stewart Islands have been prepared from herbarium
material examined in AK, AKU, CHR, WELT,
WELTU, the personal herbaria of Mr A. E. Esler
(Botany Division, c/o Plant Diseases Division, DSIR,
Auckland), Dr J. E. Braggins and Mr A. E. Wright
(University of Auckland), and my own personal
collections. Lists of material examined have not been
given, but the sheets in the above herbaria have
been appropriately annotated.
Also, the occurrence of the different species on
all the main islands in the New Zealand region has
been listed in the same abbreviated fashion as in
Allan (1961); i.e., K, Kermadec Islands; N, North
Island; S, South Island; Ch., Chatham Islands; St.,
Stewart Island; Sn., Snares Islands; A, Auckland
Islands; C, Campbell Islands; Ant., Antipodes Island.
In determining the taxonomic rank to be assigned
to particular taxa, precedents set in naming the Euro-
pean Asplenium flora have been followed. It is hoped
that some equivalence in the treatment of this cos-
mopolitan genus in different areas will thus be
achieved. Hence, the level of subspecies has been
applied to taxa which are either cytologically or
ecologically distinct but which can be clearly dis-
tinguished morphologically only by means of micro-
characters. The term variety is reserved for occas-
New Zealand Journal of Botany 15, 1977
ional variants, or distinct forms confined to limited
geographical areas. Species which show marked
polymorphy in many of their populations have not
been taxonomically subdivided, but their ranges of
variation are illustrated.
Wherever possible details of the type specimen
are given. For those species first described by
Forster, lectotypes have been designated because it
is not possible to identify the holotypes. Carolin
(1963) indicates, that for species in the "Prodromus",
specimens in BM and GOET have the greatest claim
for selection. These have been examined. Specimens
from BM have been chosen as the lectotypes because
they are of generally superior quality, those in GOET
frequently consisting only of fragments of a frond.
KEY TO THE NEW ZEALAND SPECIES
OF ASPLENIUM — see opposite page.
1. ASPLENIUM FLABELLIFOLIUM Cav. Descr.
PL: 257 (1802)—as A. flavelifolium* (Fig. 1)
A. flabellifolium var. ramosum Col. Trans. N.Z.
Inst. 20: 228 (1888).
•The spelling of the specific epithet in Cavanilles'
original description is flavelifolium but this was later
corrected to flabellifolium by Swartz (1806; pp. 88,
273). Eichler (1965) suggested that "The original
spelling . . . ought to be retained, as it is difficult
to justify an alteration of this kind which is neither
a correction of a typographic nor of an orthographic
error". However, the derivation of the original name
flavelifolium is obscure, unless it means yellow-
leaved. This seems unlikely because (i) the leaves
are not particularly yellow in this species, (ii) there
is no mention of yellow leaves in the original
description - indeed, they are stated as being bright
green, (iii) if Cavanilles had wanted to use an
epithet meaning yellow-leaved he would surely have
used flavifolium. Thus, the name flavelifolium is
either nonsensical or quite inappropriate, and it is
extremely improbable that Cavanilles intended to
produce such a meaningless epithet for his new
species. It seems much more likely that he meant
the name to refer to the very characteristic fan-
shaped leaves of this fern which he specifically
mentioned in the protologue. The name he intended
was almost certainly flabellifolium but in publishing
it as flavelifolium two kinds of errors may have
been perpetrated - a typographic error in which
Cavanilles' handwritten "b" was mistaken for a
"v", and an orthographic error in which one " 1 "
was used instead of two. Article 73 of the Inter-
national Code of Botanical Nomenclature sanctions
the correction of both types of error and the change
from flavelifolium to flabelifolium to flabellifolium
is therefore entirely within reason. Hence, I have
rejected Eichler's arguments for retaining the name
A. flavelifolium and followed the now widely
accepted practice first established by Swartz.
Brownsey—N.Z. species of Asplenium 41

KEY TO THE NEW ZEALAND SPECIES OF ASPLENltjM

/ Fronds simply pinnate 2

Fronds 2 — or more pinnate //
2 Fronds < 3 cm wide 3
Fronds > 3 cm wide 6
3 Stipe and rachis scaly; usually < 8 pairs pinnae 4
Stipe and rachis without scales; usually > 8 pairs pinnae 5
4 Fronds thick and fleshy; pinnae with scales on underside only; plant of exposed
sea cliffs 4. A. obtusatum
Fronds not thick and fleshy; pinnae often with scales on both surfaces; plant of
calcareous soils 7. A. lyallii
5 Stipe and rachis green; fronds lax and rooting at tip 1. A. flabellifolium
Stipe and rachis brown; fronds stiff and erect 2. A. trichomanes
6 Rachis dark chocolate brown throughout; pinnae irregularly and doubly
serrate 3. A. polyodon
Rachis green, at least on the upper surface (sometimes turning pale brown when
dried); pinnae entire or regularly serrate 7
7 Pinnae linear, > 10 times as long as broad 12. A. flaccidum
Pinnae < 10 times as long as broad 8
8 Pinnae prominently serrate; sori extending right to margin between serrations
6. A. scleroprium
Pinnae entire or minutely serrate; sori not reaching margin 9
9 Lowest pinnae usually lobed or with at least one distinct pinnule; sometimes with
scales on both surfaces; plant of calcareous soils 7. A. lyallii
Lowest pinnae not lobed or incised, lacking scales on upper surface 10
10 Rachis and foliar scales with very long hair-like apices (Fig. 9K-0); pinnae firm
but not thick and fleshy, glossy above, accuminate to ± acute 5. A. lucidum
Rachis and foliar scales ovate to narrowly ovate with short hair-like apices
(Fig. 9A-J); pinnae thick and fleshy, dull above, obtuse to ± acute
4. A. obtusatum
// Fronds bearing bulbils 9. A. bulbiferum
Fronds not bearing bulbils 12
12 Rhizome creeping; fronds thin, glossy above 8. A. lamprophyllum
Rhizome not creeping; fronds dull above 13
13 Sori remote from margin, 4-10 mm long 7. A. lyallii
Sori submarginal, or, if remote from margin, < 4 mm long 14
14 Fronds 3-4 pinnate at base; ultimate segments long and narrow, but broadened
in region of sorus; plant of Kermadecs 14. A. shuttleworthianum
Fronds 2-pinnate at base, or, if 3-4 pinnate, ultimate segments not broadened in
region of sorus 15
15 Fronds 3-4 pinnate at base; ultimate segments linear, mostly < 1 mm wide; pinnae
and pinnules crowded and overlapping; stipe stout 11. A. richardii
Fronds 2-3 pinnate at base; not with all above combination of characters 16
16 Fronds thin; sori 2-4 mm long 17
Fronds thick and leathery or fleshy; sori 2-10 mm long 18
(A. flaccidum agg.)
17 Fronds with < 15 pairs pinnae (if c. 15 pairs pinnae, pinnules with distinct but very
slender stalks) 10. A. hookerianum
Fronds with > 15 pairs pinnae (if c. 15 pairs pinnae, pinnules sessile or on very
short broad stalks) 9. A. bulbiferum
18 Spores* pale yellow, surface granular but with few or no prominent ridges
(Fig. 29 M-P) 19
Spores* dark brownish yellow, surface with many very prominent ridges
(Fig. 30 O-R) 20
19 Fronds limp and flaccid; plant usually epiphytic 12a. A. flaccidum subsp. flaccidum
Fronds firm and erect; plant of exposed sea cliffs, confined to east coast regions
north of lat. 38° 12b. A. flaccidum subsp. haurakiense
20 Fronds thick and fleshy; plant of exposed sea cliffs
13b. A. terrestre subsp. maritimum
Fronds leathery, but not fleshy; plant of forest floor or rocky inland areas
13a. A. terrestre subsp. terrestre

•Spores mounted in gum chloral or other suitable medium and viewed at c. X 200.
42 New Zealand Journal of Botany 15, 1977

Fig. 1 Fronds of Asplenium flabellifolium ( x 1). Octoploid cytotype (A-C), hexaploid cytotype (D-F). A.
NZ 191, Eastbourne, Wellington; B, NZ 834, Turakirae Head, Wellington; C, NZ 847, Seatoun, Wellington;
D, NZ 177, Kapiti Island; E, NZ 887, Rangitoto Island; F, NZ 789, Wharariki Beach, NW Nelson.

TYPE: In MA (not seen). 2n = 277-280 (exact number uncertain) (Fig. 31B).

DESCRIPTION: Rhizome short, erect, bearing dark
brown subulate scales with long filiform apices up
to 6 X 0.5 mm (Fig. 4M-S). Stipes 1-10 cm long,
green, slender, lacking scales. Laminae linear,
7-25 X 1-2.5 cm, light green, flaccid and often pros-
trate, pinnate. Raches green, weak, lacking scales,
frequently extending beyond the uppermost pinnae
and rooting at their apices. Pinnae 8-25 pairs,
gradually decreasing in size from base to apex, fan-
shaped, crenate-dentate, cuneate at base, lowest 5-15
X 5-20 mm. Sod radiating along veins, up to 6 mm
long. Spores (37)43-51(62) Mm long, (30)36-41(51)
fim wide (8 populations), perispore with numerous
crests and ridges (Fig. 29C-D).
CHROMOSOME NUMBER: 2n = 210 or 211 (Fig. 31A).
NZ 887, Rangitoto Island; NZ 216, Stronvar, Waira-
rapa; NZ 177, 178, Kapiti Island; NZ 789, Whara-
riki Beach, NW Nelson.
NZ 181, 186, Eastbourne, Wellington; NZ 314, Karori,
Wellington; NZ 834, Turakirae Head, Wellington;
NZ 847, Breaker Bay, Wellington; NZ 775, Port
Hills, Christchurch.
HABITAT AND DISTRIBUTION: N, S. A. flabellifolium
is found in scattered localities from the Bay of Islands
to Otago (Fig. 2), but it is absent from large areas
of the west coast of both islands and from the far
south. It grows from sea level to 1 250 m in dry rocky
ground, on lava and scoria fields, in grassy areas
below scrub, and sometimes in open beech forest; it
never occurs in deep forest. Its greater abundance in
open scrub and the margins of agricultural land,
particularly in Canterbury and the Wairarapa, sug-
gests that it may be a species which has considerably
extended its range with the advent of European
colonisation. Wherever it occurs, it appears as a pros-
trate plant and spreads rapidly by its rooting tips.
Brownsey—N.Z. species of Asplenium
Fig. 2 Distributions of
Asplenium flabellifolium and
A. polyodon.
A. flabellifolium
COMMENT: A. flabellifolium is a very distinctive
species confined to New Zealand and parts of southern
Australia and Tasmania. It was first investigated
cytologically by Brownlie (1958) who reported
n. = c.27O in plants from Lyttelton, indicating that
this was a sexual c.l6-ploid species (the base number
(x) in Asplenium is 36).
More recently, Lovis (1973) investigated a collec-
tion from Australia and showed that there were
c.212 univalents present at meiosis. Prothalli raised
from spores of this collection proved to be apogam-
ous and Lovis therefore pointed out that the taxon
was an approximately hexaploid apomict of the
Asplenium aethiopicum type*. Because plants of A.
flabellifolium from Australia and New Zealand are
very similar morphologically, this evidence clearly
*i.e., reduction of the chromosome number is circum-
vented by nuclear restitution after asynaptic meiosis
(Braithwaite 1964) and not by a failure of mitosis
in the formation of the spore mother cells, as in
most apomictic ferns (Manton 1950, Walker 1966).
43
A. polyodon
suggested the possibility that the material investigated
by Brownlie was not a sexual c. 16-ploid species but
an approximately octoploid apomict.
During the present investigation, plants of A.
flabellifolium from several localities have been
examined at meiosis and, in every case, a very dis-
tinctive diakinesis has been observed in which only
univalents are present (Fig. 31 A, B). This confirms
the fact that A. flabellifolium is an apomictic species
in New Zealand as well as Australia. Two separate
chromosome numbers have been recorded, 2n = 210~
211 and 2n = 277-280. The lower number clearly cor-
responds to that reported in Australian material, but,
although there is some slight doubt as to whether the
exact number is 210 or 211, it seems certain that it
is not as high as the 212 suggested by Lovis (1973).
The number of the higher cytotype has been deter-
mined with less accuracy, but it is sufficiently clear
to indicate that in reporting n = c.27O Brownlie (1958)
mistook the univalents in this apomictic species for
bivalents, thereby implying a level of plbidy twice
as high as is the case.
44
The two cytotypes of A. flahellifolium represent
approximately hexaploid and octoploid levels of
ploidy, but it is now certain that their chromosome
numbers do not correspond to exact multiples of
36. Thus, one would anticipate 216 and 288 chromo-
somes respectively, but the actual numbers of 210-
211 and 277-280 indicate that some aneuploidy has
occurred. One rather large chromosome is regularly
present in both cases and, as suggested by Lovis
(1973), some of the missing chromosomes may have
been absorbed by fusion.
No constant morphological distinction of any kind
has been observed between the two different cyto-
types and neither are they obviously restricted to
particular localities. Hence, they have not been
accorded any separate taxonomic status.
2. ASPLENIUM TRICHOMANES L Sp. PI.:
1080 (1753).
DESCRIPTION: Rhizome stout, erect, bearing dark
brown subulate scales up to 5 mm long. Stipes 1-10 cm
long, dark brown, shining, stiff, lacking scales except
at the very base. Laminae linear, 5-30 cm long, dark
green, sub-coriaceous, pinnate. Raches dark brown,
shining, stiff, lacking scales. Pinnae sub-sessile, 15-25
(or more) pairs, oblong to ± orbicular, crenate-
serrate to ± entire, 2-10 X 1-5 mm. Sori up to 3 mm
long.
CHROMOSOME NUMBER: TWO distinct chromosome
numbers have been reported for this species in New
Zealand, n = 72 and n = 108 (Brownlie 1954; Lovis
1956, 1958).
HABITAT AND DISTRIBUTION: N, S. A. trichomanes
displays a considerable range of morphological and
cytological variation over its virtually cosmopolitan
distribution. A world-wide biosystematic investigation
of this intricate group is in preparation (Lovis, unpub-
lished data). However, in New Zealand A. tricho-
manes is a very distinctive fern and is apparently
quite unrelated to any other native species with the
possible exception of A. flabellifolium. In a detailed
but unpublished study of A. trichomanes in New
Zealand, Lovis (1956) showed that two different
cytotypes were present which could only be dis-
tinguished morphologically by the structure of the
rhizome scales and by the size and markings of the
spores (mean values: 34-39 /nn long in the tetraploid
and 39-47 pm long in the hexaploid). The tetraploid.
which in Europe is the most common cytotype, is
rare here and apparently confined to local areas of
the North Island. The second cytotype, a hexaploid.
is the characteristic form in New Zealand. It is
widespread in the South Island from sea level to
New Zealand Journal of Botany 15. 1977
900 m, occurring on both limestone and greywacke
rock, but in the North Island it is confined to
scattered localities on calcareous soil. Outside New
Zealand it is known from only a single locality in
Australia (Gudgenby Creek, Australian Capital
Territory — Lovis, pers. comm.).
3. ASPLENIUM POLYODON Forst. f. Prodr.: 80
(1786) (Fig. 3)
Trichomanes adiantoides L. Sp. PI. 2: 1098
(1753)
Asplenium falcatum Lam. Encyc. 2: 306 (1786)
(torn, abort.
Tarachia polyodon (Forst. f.) Presl. Epim. Bot.:
77 (1851)
Asplenium adiantoides (L.) C. Chr. Ind. Fil.:
99 (1906) non Lam. 1786, non Raddi 1819.
non Raoul 1844
Asplenium adiantoides var. polyodon (Forst. f.)
C. Chr. Ind. Fil.: 126 (1906)
Asplenium forsterianum Col. Tasm. J. nat. Sci.
2: 171 (1846)
Asplenium falcatum var. caudatum (Forst. f.)
Allan, Fl. N.Z.: 71 (1961)
TYPE: Not located (not in BM or GOET).
DESCRIPTION: Rhizome stout, short creeping, densely
covered in red-brown, narrowly triangular scales up
to 10 X l m m . Stipes 10-30 cm long, dark brown,
still", densely covered in scales similar to but smaller
than those of the rhizome (Fig. 4A-E). Laminae
lanceolate, 25-50 (or more) X 10-20 cm, dark green
and glossy above, paler and dull below, frequently
pendulous, pinnate. Raches dark chocolate brown,
very scaly. Pinnae 25 (or more) pairs, narrowly
angular-ovate to ovate, sometimes with a large
rounded basal acroscopic lobe, acuminate, doubly
serrate, 5-10 x 1—2 cm, scaly and with prominent
veins on underside. Sori often slightly curved away
from the midrib, up to 2 mm long. Spores (26)30-37
(40) fim long, (15)18-21(25) /an wide (6 popula-
tions), perispore with a few very prominent ridges
(Fig. 30A-B).
CHROMOSOME NUMBER: 2n = 144. NZ 679, Spraggs
Bush, Waitakere Ranges, Auckland; NZ 377, Bay of
Many Coves, Marlborough Sounds.
Tetraploid chromosome numbers have also been
recorded for this species in Australia and Ceylon
(Manton & Sledge 1954), Malaya [Malaysia] (Manton
1954), and the Himalayas (Bir 1960).
HABITAT AND DISTRIBUTION: K. N, S, Ch., St. A.
polyodon occurs widely in forested areas from North
Cape to Stewart Island, though it is now absent
from most of the east coast of the South Island
Brovvnsey—N.Z. species of Asplenium 45

Fig. 3 Fronds of Asplenium polyodon ( x | ) . A. NZ 864. Carter's Bush. Wairarapa; B. NZ 928, Kapiti
Island; C. NZ 1069. Lighthouse Track. Stewart Island.
 Q R
Fig. 4 Camera lucida drawings of scales ( x 20), each taken from a different population. A-E, stipe scales
from Asplenium polyodon; F-L, stipe scales from A. lyallii (F & I from bipinnate plants, others from pinnate
specimens); M-S, rhizome scales from A. ftabellifolium (M, N, P from hexaploid cytotype; O, Q, R, S from
octoploid cytotype).
Brownsey—N.Z. species of Asplenium
where forest has been cleared (Fig. 2). It is found
principally as an epiphyte from sea level to c. 900 m,
but it also grows on fallen logs and even on the
ground. In the far south, A. polyodon grows
terrestrially in coastal forest and scrub, and often
develops into very large plants. A substantial
distance separates the plants in these localities from
those in Westland which are more typically epiphytic,
but whether this apparent disjunct distribution is real
needs further investigation. There is nothing to suggest
that the terrestrial plants are morphologically different
to the epiphytic ones apart from their larger size.
COMMENT: Until recently this fern has been widely
referred to as either A. adiantoides (L) C. Chr. (e.g.
Cheeseman 1925) or A. falcatum Lam. (e.g., Allan
1961, Crookes 1963). However, Morton (1967) has
shown that the correct name is A. polyodon Forst. f.
The confusion has arisen because the original descrip-
tion of this species (as Trichomanes adiantoides L.
by Linnaeus in "Species Plantarum'") was based on
two separate elements - one from Ceylon illustrated
by Burmann ("Thesaurus Zeylanicus", t.43, 1737),
and one from Africa illustrated by Plukenet ("Phyto-
geographia", t.123, 1692). These are now known
to be quite distinct species.
Lamarck was the first to realise this, but unfor-
tunately compounded the existing confusion by retain-
ing Linnaeus' specific epithet for the African plant
which he called Asplenium adiantoides Lam. How-
ever, it seems that the type of Trichomanes adian-
toides L. was actually the plant from Ceylon (Morton
1967) to which Lamarck erroneously gave the new
name Asplenium falcatum Lam. and cited the Bur-
mann illustration, but which he should have called
A. adiantoides (L.) Lam. The name A. falcatum
Lam. is therefore illegitimate because the type of the
Linnean species Trichomanes adiantoides was quoted
in synonymy.
Christensen tried to remedy this error by proposing
the new combination Asplenium adiantoides (L.) C.
Chr. (Ind. Fil.: 99, 1906) for the Ceylon material
(i.e., that which Lamarck had called A. falcatum).
However, this name is also illegitimate, being a later
homonym three times over (A. adiantoides Lam.
Encyc. 2: 306, 1786; A. adiantoides Raddi, Opusc.
Sci. Bol. 3: 291, 1819; A. adiantoides Raoul, Ann.
Sci. nat. Ser. Ill 2: 115, 1844).
The oldest acceptable name for this species com-
plex is therefore A. polyodon, first proposed by
Forster in his "Prodromus" but until recently quoted
in synonymy with either A. falcatum Lam. or A.
adiantoides (L.) C. Chr. The species extends through-
out much of the Indo-Malaysian, Australian, and
Pacific regions and is very variable. However, Skotts-
berg (1942) made a thorough investigation of the
47
taxon throughout its range of distribution and con-
cluded that plants from Mauritius, Timor, the
Phillipine Islands, New Zealand, Australia, and New
Caledonia were basically similar.
Within the New Zealand region, further nomen-
clatural confusion has been caused by the existence of
a rather narrow-pinnuled form of A. polyodon on
the Kermadec Islands. This material has sometimes
been recognised as A. caudatum Forst. f. (e.g.,
Cheeseman 1925) or taxonomically distinguished as a
new variety, A. falcatum var. caudatum (Forst. f.)
Allan (e.g., Allan 1961, Crookes 1963). However,
it is clear from examination of Forsters material in
BM and GOET that A. caudatum is a distinct species
which has deeply dissected, more or less parallel-
sided pinnae bearing sori only c. 4-5 mm long. There
is no evidence to suggest that Forster's A. caudatum
was collected in New Zealand and plants comparable
to it do not grow in this region today. Copeland
(1932) indicates that the type locality was probably
in the Society Islands and that this species has a
distribution extending from the New Hebrides to
Hawaii. The plants on the Kermadecs are therefore
probably best regarded as one extreme of the range
of variation in the polymorphic A. polyodon complex.
4. ASPLENIUM OBTUSATUM Forst. f. Prodr.:
80 (1786) (Figs 5, 6)
A. obliquum Forst. f. Prodr.: 80 (1786)
A. ohtusatum var. obliquum (Forst. f.) Hook. f.
Fl. Antarct. 1: 108 (1844)
A. apice-dentatum Homb. et Jacq. Voy. Pole
Sud: pi. 1A (1853)
A. lucidum var. obliquum (Forst. f.) Moore,
Ind. Fil.: 142 (1859)
TYPE: Lectotype in BM; New Zealand, coll. J. R.
and G. Forster.
DESCRIPTION: Rhizome stout, often forming a hard
woody mass above ground, bearing brown, shiny,
ovate, acuminate scales up to 15 X 4 mm. Stipes
2-30 cm long, dark brown at base, green above, stout,
covered in ovate or narrowly triangular scales with
acute or acuminate apices. Laminae ovate to lanceo-
late, 4-40 X 2-20 cm, dull green, very thick and
fleshy, pinnate. Raches green, stout, prominently
grooved and scaly. Pinnae 4-18 pairs, narrowly ovate
to oblong, obtuse to acute, crenate-serrate to ± entire,
cuneate at base, 10-100 X 7-25 mm, scaly on the
underside. Sori up to 10 mm long, not reaching to
lamina edge. Perispore of spores slightly winged but
with few crests or ridges (Fig. 29E-H).
 New Zealand Journal of Botany 15, 1977

Fig. 5 Fronds of Asptenium obtusatum subsp. obtusatum ( x J). A, NZ 1045, Curio Bay. SE Otago; B, NZ
1059, Maori Bay, Stewart Island; NZ 471, Cape Foulwind, Westport; D, NZ 1170, Ship Cove, S Westland;
E, NZ 1058, Maori Bay. Stewart Island; F, NZ 996, Tomahawk Beach, Dunedin.
Brownsey—N.Z. species of Asplenium
4a. A. OBTUSATUM Forst. f. subsp.
OBTUSATUM (Fig. 5)
DIAGNOSIS: Stipes 4-30cm long, covered in ovate
scales (Fig. 9A-E). Laminae usually ovate, 6-40
X 5-20 cm. Pinnae 5-15 pairs, 20-100 X 10-25 mm.
Spores (35)40-43(49) Mm long, (21)24-27(33) ^m
wide (5 populations).
CHROMOSOME NUMBER: n = 72 (Fig. 31C). NZ 977,
Cape Terawhiti, Wellington; NZ 472, Cape Foulwind,
Westport; NZ 465, Punakaiki; NZ 998, Tomahawk
Beach, Dunedin.
The same chromosome number has also been
recorded for this taxon from Stewart Island (Brownlie
1958) and Tristan da Cunha (Manton & Vida 1968).
HABITAT AND DISTRIBUTION: N, S, Ch., St., Sn., Ant,
A, C. (Fig. 7). A. obtusatum subsp. obtusatum is
commonest in the South Island where it occurs more
or less continuously around the west and south coasts
from Cape Farewell to Moeraki Point; there is also
an outlying population on Banks Peninsula. In
Nelson, the Marlborough Sounds, and on the east
coast as far south as Banks Peninsula it is apparently
replaced by A. lucidum (Fig. 7). In the Cook Strait
region there is a population on Brothers Island, but
the only recently confirmed record from the North
Island is at Cape Terawhiti*.
In the far south, A. obtusatum is abundant around
the coasts of the islands of Foveaux Strait and on
the sub-Antarctic islands.
In the northern part of its New Zealand range,
subsp. obtusatum is a rather small plant found grow-
ing in isolated populations only on the most exposed
coastal rocks. In the south, however, it grows
luxuriantly not only on exposed cliffs but also further
back in the shelter of the coastal vegetation where
it attains a height of 1 m or more and often forms
an important association with Hebe elliptica (Forst.
f.) Pennell and Blechnum durum (Moore) Chr.
Nowhere, however, does it extend beyond the
influence of salt spray.
COMMENT: The Forster material of A. obliquum
in BM and GOET has been examined. In none of
its characters does it exceed the range of variability
in A. obtusatum subsp. obtusatum and, hence, plants
referred to it by Forster and later workers should
be regarded as merely large forms of A. obtusatum.
Field work in localities from which such plants have
*In WELT there are old herbarium specimens from
Kapiti Island and from Somes Island in Wellington
Harbour.
49
been collected indicates that there is often a com-
plete range of forms within a few yards, from small
plants in the most exposed sites to much larger
plants sheltered by scrub vegetation. The differences
in morphology are entirely due to changes in the
environment, and, because the various forms are
regularly found together in close proximity, there
seems little justification for maintaining any taxo-
nomic distinction between them.
4b. A. OBTUSATUM Forst. f. subsp. NORTH-
LANDICUM Brownsey subsp. nov. (Fig. 6)
Planta plerumque parvior quam subsp. obtusatum.
Stipites 2-8 cm longi, paleis anguste triangularibus
vestiti (Fig. 9F-J). Laminae lanceolatae vel ovatae,
4-20 X 2-7 cm. Pinnae 4-18 jugatae, 10-40 X
7-12 mm. Sporae (39)45-52(60) /on longae, (25)29-
33(38) /ITO. latae (Fig. 29G-H). Numerus somaticus
chromosomatum 288; in meiosi paria 144 observantur
(Fig. 31E).
HOLOTYPUS: CHR; NZ 667, Karekare, Auckland
(NZMS 1 N41:000432). Growing in crevices on the
cliffs at the south end of the beach. Coll. P. J.
Brownsey, 19.3.1974. (CHR 308924)
DIAGNOSIS: Generally a smaller plant than subsp.
obtusatum. Stipes 2-8 cm long, covered in narrowly
triangular scales (Fig. 9F-J). Laminae lanceolate to
ovate, 4-20 X 2-7 cm. Pinnae in 4-18 pairs, 10-40
X 7-12mm. Spores (39)45-52(60) Mm long, (25)29-
33(38) /j.m wide (6 populations).
cm
10-
5-
0J
Fig. 6 Fronds of Asplenium obtusatum subsp.
northlandicum ( x 1). A, B, NZ 667, Karekare,
Auckland; C, NZ 603, Little Barrier Island.
50
CHROMOSOME NUMBER: n = 144 (Fig. 31E). N7. 667,
Karekare, Auckland.
HABITAT AND DISTRIBUTION: K, N. Apart from one
locality at Karekare, A. obtusatum subsp. north-
landicum is found only on the east coast of the
North Auckland peninsula extending from the Three
Kings to Slipper Island off Coromandel (Fig. 7). It
is separated from the tetraploid subspecies by virtually
all the remaining coastline of the North Island. It
grows only on coastal rocks or amongst scrub
exposed to salt spray, but it is a rather small plant
and is not as important a constituent of the coastal
vegetation as the southern subspecies. It is so far
not known outside New Zealand.
COMMENT: The disjunct distribution of A. obtusatum
in New Zealand is accompanied by a change in
the chromosome number, populations in the
south being tetraploid and those in the north
A. obtusatum
• subsp.
northlandicum
• subsp.
obtusatum
A. lucidum
New Zealand Journal of Botany IS, 1977
octoploid, but the gross morphology of the two
cytotypes is very similar. The tetraploid differs signi-
ficantly only in having stipe scales which are more
broadly ovate (cf. Fig. 9A-E, F-J) and spores which
are smaller than those of the octoploid (cf. Fig. 29E.
G). Despite their geographic separation, the two
cytotypes can only be distinguished morphologically
by these micro-characters and hence they have not
been accorded full species status.
A. obtusatum occurs commonly around the
southern coastlines of Australia and South America
as well as on many islands of the southern Atlantic
and Pacific Oceans. Material from Tristan da Cunha
is known to be tetraploid (Manton & Vida 1968)
and therefore presumably corresponds to subsp.
obtusatum. Collections from elsewhere have yet to
be cytologically examined, but, because of the dis-
covery of two distinct cytotypes in New Zealand, it
is very desirable that this species be more thoroughly
investigated on a world-wide basis.
Fig. 7 Distributions of
Asplenium obtusatum and
A. lucidum.
Brownsey—N.Z. species of Asplenium
5. ASPLENIUM LUCIDUM Forst. f. Prodr.: 80
(1786) (Fig. 8)
A. oblongifolium Col. Tasm. J. nat. Sci. 2: 171
(1846)
A. lucidum var. paucifolium Hook. f. Handbk.
N.Z. Fl.: 371 (1864)
A. iTurvillei Mett. Kuhn Linn. 36: 95 (1869)
A. obtusatum var. lucidum (Forst. f.) Hook, et
Baker Syn. Fil.: 207 (1873)
A. obtusatum var. integrifolium Szyszyl. in
Wawra, Itin. princ. S. Coburgi: 122 (1888).
TYPE: Lectotype in BM; coll. G. Forster (type
locality given in "Prodromus" is New Zealand).
DESCRIPTION: Rhizome stout, often forming a hard
woody mass above ground, bearing pale brown,
shiny, ovate, acuminate scales up to 30 X 7 mm.
Stipes 8-20 cm long, dark brown, stout, densely
covered in narrow scales with very long filiform
apices (Fig. 9K-O). Laminae oblong to elliptic,
18-100 X 10-35 cm, dark green and glossy above,
pinnate. Raches brown below, green above, stout,
slightly ridged, scaly. Pinnae 4-15 pairs, lanceolate
to narrowly oblong or ovate, acuminate, crenate-
serrate to ± entire, cuneate at base, 4-15 X 1-3 cm,
frequently covered in very small hair-like scales on
the underside. Sori up to 20 mm long, not reaching
lamina edge. Spores (27)33-38(44) fim long, (18)20-
24(26) /im wide (5 populations), perispore a com-
plex reticulate network (Fig. 29A-B).
CHROMOSOME NUMBER: n = 7 2 (Fig. 31D). NZ
Rangitoto Island; NZ 462, Greymouth.
Brownlie (1958) reported n = 72 in material of
this species from Queen Charlotte Sound.
HABITAT AND DISTRIBUTION: K, N, S, Ch. (Fig.
A. lucidum is found commonly in the North Island
where it occupies a wide variety of habitats from
coastal cliffs to deep forest and occurs both terres-
trially and epiphytically. It is abundant in coastal
regions but becomes less frequent towards the vol-
canic plateau. In the South Island it is reasonably
common in coastal areas of NW Nelson and in the
Marlborough Sounds, but is rare further south appar-
ently reaching no further than Greymouth on the
West Coast and Banks Peninsula in the east*. Its
absence from the central North Island and from the
far south is probably due to an inability to tolerate
cold.
*I have found many herbarium specimens originat-
ing from further south and labelled as A. lucidum,
but these have all proved to be A. obtusatum subsp.
obtusatum.
51
Cheeseman (1925) and Allan (1961) state that
A. lucidum has been recorded on Norfolk Island and
Lord Howe Island. However, collections from Lord
Howe Island have been variously identified as A.
lucidum, A. milnei Carr., and A. obtusatum (for
details, see Oliver 1917). Clearly, further investi-
gation is required before the existence of A. lucidum
outside the New Zealand region can be confirmed.
COMMENT: Taxonomists have previously relied on
the texture, colour, and shape of the pinnae to dis-
tinguish A. obtusatum from A. lucidum. However,
these characters vary greatly in different environ-
ments and considerable confusion has surrounded
the identity and distribution of the two taxa, even
to the extent that they have sometimes been reduced
to varieties of the one species. However, there is no
doubt that two quite separate species do exist and
that in most cases they can be satisfactorily distin-
guished. The principal source of confusion appears
to have been the very large plants of A. obtusatum
which occur in the far south and which superficially
resemble A. lucidum in their general habit and pinna
shape. However, A. lucidum is clearly distinct in that
it has thinner pinnae with acuminate apices and a
very glossy upper surface; the stipe scales are narrow
with extremely elongated hair-like apices (Fig.
9K-O); the spores have a very characteristic pattern
consisting of a complex reticulate network (Fig.
29A-B) and they are also slightly smaller (mean
values 33-38 X 20-24 /tm). A. obtusatum is charac-
terised by fleshy pinnae which have obtuse apices
885,
and are usually dull green; the stipe scales are
broadly ovate without long hair-like projections (Fig.
9A-E); the spores have a slightly winged and ridged
perispore (Fig. 29E-F), and they are somewhat larger
(mean values 40-43 X 24—27 /im in subsp. obtusatum)
7).
than those of A. lucidum. On this basis it is now
believed that A. lucidum is entirely replaced in the
far south by A. obtusatum subsp. obtusatum, and that
A. lucidum extends only as far as Greymouth and
Banks Peninsula in the South Island. On the southern
edge of its distribution, particularly on the West
Coast, plants of A. lucidum are unusually small
(Fig. 8A) and their stipe scales sometimes lack very
long hair-like apices. Nevertheless, their spore charac-
ters indicate that they belong to A. lucidum and not
to A. obtusatum.
The two species may also be confused in the far
north where both A. lucidum and A. obtusatum
subsp. northlandicum occupy similar habitats on sea
cliffs of the Hauraki Gulf area. The stipe scales are
not always a reliable taxonomic guide in this region
because those of A. obtusatum subsp. northlandicum
are narrower and have longer apices than those of
subsp. obtusatum (cf. Fig. 9F-J, A-E). Never-
theless, the important difference in chromosome
52 New Zealand Journal of Botany IS. 1977

D
Fig. 8 Fronds of Asplenium lucidum ( x i ) . A, NZ 809, Takaka Hill, Nelson; B, NZ 913, Kapiti Island;
C. NZ 1171, Buller Gorge, Nelson; D, NZ 532, Little Barrier Island.
Brownsey—N.Z. species of Aspleniutn

Fig. 9 Camera lucida drawings of stipe scales ( x 20), each from a different population. A-E, Asptenium
obtusatum subsp. obtusatum; F-J, A. obtusatum subsp. northlandicum; K-O, A. lucidum; P-R, A. scleroprium.
54
number between A. lucidum and A. obtusatum subsp.
northtandicum is manifested in the much larger spore
size of the latter (mean values 33-38 X 20-24 /j.m
in A. lucidum, 45-52 X 29-33 ,um in A. obtusatum
subsp. northlandicum), and the two taxa may be
reliably distinguished on this basis alone.
6. ASPLENIUM SCLEROPRIUM Homb. et Jacq.
Voy. Pole Sud: pi. ID (1853) (Fig. 10)
A. flaccidum var. aucklandicum Hook. f. Fl.
Antarct. 1: 109 (1844)
A. lucidum var. scleroprium Moore, Ind. Fil.:
142 (1859)
A. lucidum var. aucklandicum (Hook, f.) Allan,
Fl. N.Z.: 1012 (1961)
A. aucklandicum (Hook, f.) Crookes, N.Z. Ferns
ed. 6: 334 (1963)
TYPE: Not located.
DESCRIPTION: Rhizome short, stout, erect, bearing
red-brown, ovate, acuminate scales up to 25 x 4 mm.
Stipes 15-50 cm long, green on upper surface, brown
below, stout, covered in narrowly triangular scales
with long filiform apices (Fig. 9P-R). Laminae
lanceolate, 15-50 X 8-20 cm, dark green, thick and
leathery, pinnate. Raches green, stout, prominently
grooved and scaly. Pinnae 6-17 pairs, lanceolate,
acuminate, often broadened and lobed towards cuneate
base, deeply and regularly crenate-serrate, 5-10 X
1-2 cm, scaly on underside. Sori reaching the lamina
margin in sinuses, not more than 10 mm long. Spores
(38)48-53(61) Mm long, (24)28-33(42) pm wide (5
populations), perispore slightly winged, but with few
ridges (Fig. 29I-J).
CHROMOSOME NUMBER: n=144 (Fig. 32C).
1010, 1012, Bluff, Southland.
HABITAT AND DISTRIBUTION: S, St., Sn., A.
scleroprium is an endemic species known only from
a very few collections most of which have been
made on the Auckland Islands; others originate
from the Snares, from one locality on the mainland
at Bluff, and from Ernest, Big South Cape, and
Codfish Islands around Stewart Island. (The site of
an early collection at Half Moon Bay was not
relocated during recent field work on Stewart Island.)
The species is confined to coastal areas and often
grows alongside A. obtusatum on cliffs and in scrub
exposed to salt spray.
COMMENT: Accounts of the coastal vegetation on
islands in the far south (e.g., Fineran 1966a, b; 1969;
1973) frequently mention a very variable mixture of
plants which have usually been referred to the A.
New Zealand Journal of Botany 15, 1977
obtusatum complex. It is now clear that most of
these plants are forms of A. obtusatum subsp.
obtusatum varying according to their degree of
exposure on the cliff. However, there is no doubt
that a second species, A. scleroprium, also occurs in
this habitat. Previously, this taxon has been con-
sidered a variety of both A. lucidum and A. flaccidum,
and it has also been confused with A. obtusatum
subsp. obtusatum. However, by virtue of its octoploid
chromosome number, it is cytologically distinct from
all these taxa, and may be further distinguished by
its larger spore size. In gross morphology it has some
similarity to both A. flaccidum and A. terrestre (cf.
Figs 10, 21, 25) but its pinnae are generally broader
and less deeply divided, and they are characteristic-
ally glossy green on the upper surface. From A.
lucidum (Fig. 8) and the two subspecies of A.
obtusatum (Figs 5, 6) it may be distinguished by
its narrower more lanceolate pinnae which have
deeper and more regular serrations, and because the
sori extend to the edge of the pinna in the sinuses;
in A. lucidum and A. obtusatum the sori stop short
of the lamina margin.
7. ASPLENIUM LYALLII (Hook, f.) Moore Ind.
Fil.: 143 (1859) (Fig. 11)
A. lucidum var. lyallii Hook. f. Fl. N.Z. 2: 33
(1855)
A. anomodum Col. Trans. N.Z. Inst. 15: 309
(1883)
A. lucidum var. anomodum (Col.) Cheeseman,
Man. N.Z. Fl.: 991 (1906)
TYPE: Holotype in K; Otago, Middle Island, N.Z.,
coll. Lyall.
NZ
DESCRIPTION: Rhizome stout, ascending, bearing
brown subulate to narrowly triangular scales up to
25 X 2 mm. Stipes 3-20 cm long, pale brown at base
A. and on underside, green elsewhere, covered with
scales similar to but smaller than those of the
rhizome (Fig. 4F-L). Laminae lanceolate to elliptic,
4—40 X 2—20 cm, dark shiny green to dull grey-green,
pinnate to bipinnate. Raches normally green, occa-
sionally brown on the underside, often grooved,
scaly. Pinnae 2-16 pairs, ovate to narrowly oblong,
obtuse to acuminate, serrate to ± entire, stalked,
1-10 X 0.5-5 cm, often covered on both surfaces
with tiny scales. Lowermost pinnae normally at least
partially lobed or divided at the base, sometimes
completely pinnate; pinnules stalked, lanceolate to
elliptic, obtuse, serrate to entire, up to 3 X 1.5 cm.
Sori up to 10 mm long, not reaching lamina edge.
Spores (36)40-54(60) Mm long, (23)27-35(40) Mm
wide (13 populations), perispore prominently winged
and ridged (Fig. 29K-L).
Brownsey—N.Z. species of Asplenium
Fig. 10 Fronds of Asplen-
ium scleroprium ( x | ) .
A-C. NZ 1016. Bluff.
CHROMOSOME NUMBER: n=144 (Fig. 32D). NZ 268,
Manaia, Taranaki; NZ 744, Woodside Creek, Marl-
borough; NZ 277, Kaikoura Peninsula; NZ 854, Dan
Rogers Gully, Banks Peninsula; NZ 986, Castle Hill.
Canterbury.
These results agree with those of Brownlie (1958)
who reported n=144 in populations from Lyttelton
and Weka Pass, Canterbury. No evidence has been
found to confirm the finding by Knowlton (1957)
of 2n = c.l44 in populations of A. anomodum from
Puketautahi. Hawke's Bay and 2n = 216 in A. lyallii
from the Taieri River mouth, Otago.
55
HABITAT AND DISTRIBUTION: N, S, Ch., St. A. lyallii
is an endemic species largely confined to calcareous
soils and found at various sites along the drier east
coast and in limestone upland areas of the interior
(Fig. 12). In the North Island it occurs principally
in the mountain areas of Hawke's Bay and along
the Wairarapa coast, but there are also scattered
localities on the Egmont coast and at Te Kuiti, its
northernmost limit. In the South Island there are
more extensive populations in the limestone areas of
NW Nelson and Canterbury, and coastal populations
in Marlborough, SE Otago, Stewart Island, and parts
of Fiordland. It occurs from sea level to 1 400 m. On
 G
Fig. 11 Fronds of Asplenium lyallii ( x i ) . A, NZ 801, Takaka Hill, Nelson; B, NZ 786, Castle Hill, Canter-
bury; C, D, NZ 763, Mt Pleasant, Port Hills, Christchurch; E, NZ 730, Woodside Creek, Marlborough; F,
NZ 722, Woodside Creek, Marlborough; G, NZ 504. Manaia, Taranaki.
Brownsey—N.Z. species of Asplenium
A. lyallii
Fig. 12 Distribution of Asplenium lyallii.
the coast it grows in open forest or forest remnants,
particularly amongst the roots of Corynocarpus
laevigatus J. R. et G. Forst. At higher altitudes it is
frequently found in crevices or beneath overhangs
of limestone rock.
COMMENT: AS understood here, A. lyallii includes
all the forms which have previously been assigned
to the two taxa A. anomodum and A. lucidum var.
lyallii. The latter was first described by Hooker
(1855) who indicated that it differed from A.
lucidum only by possessing bipinnate fronds. How-
ever, Moore (1858-9) evidently attached greater
significance to this character and raised the taxon
to specific rank as A. lyallii.
A. anomodum is a later name coined by Colenso
(1883) to describe some apparently new and dis-
tinctive pinnate plants. Colenso presumably believed
that his A. anomodum was entirely different to A.
lyallii because, although he pointed out characters by
57
which A. anomodum could be distinguished from
both A. obtusatum and A. hookerianum, he found
it unnecessary to mention A. lyallii.
However, as more collections were made and the
known ranges of variability of A. anomodum and
A. lyallii increased, it became evident that they were
closely related and that the distinctions between
them were not so clear. Subsequent authors have
treated this situation in various ways. Cheeseman
1906) reduced both taxa to varieties of A. lucidum.
although in the second edition of his "Manual"
(Cheeseman 1925) he reinstated A. anomodum as a
full species distinct from A. lucidum var. lyallii in
the size, shape, and cutting of the frond. Allan
(1961) retained A. anomodum but rejected A. lyallii
altogether, suggesting that such forms might be
hybrids between A. lucidum and A. bulbiferum. On
the other hand, Crookes (1963) maintained all three
taxa, A. lucidum, A. lyallii, and A. anomodum, as
full species, though she admitted that the distinction
between the later two was somewhat doubtful.
During the present investigation, many wild popu-
lations have been examined and many plants brought
into cultivation. In the wild, both pinnate and bipin-
nate plants have been found growing together in
habitats ranging from coastal cliffs to inland lime-
stone mountains. In cultivation, a wide range of
these morphological forms have reverted to a sur-
prisingly uniform condition in which the fronds are
essentially pinnate with only the lowest pinnae par-
tially lobed or incised. Pinnate and bipinnate plants
show the same ranges of variation with respect to
spore pattern, spore size (mean values 40-54 X 28-35
itim in pinnate forms, 42-50 x 27-33 nm in bipin-
nate forms), and the shape of the stipe scales (Fig.
4F-L). All the poulations investigated have been
uniformly octoploid. Hence, it seems clear that plants
previously referred to the two species A. lyallii and
A. anomodum rea'ly belong to one extremely poly-
morphic taxon, and that the later name, A.
anomodum, should be rejected as a synonym of A.
lyallii.
Nevertheless, there is some suggestion that plants
of this complex growing in coastal areas, particularly
on non-calcareous soils, have dark shiny green fronds
which are larger, more divided, and with scales con-
fined to the lower surface, whereas plants of lime-
stone upland areas have du'l grey-green fronds which
are smaller, less divided, and have scales on both
surfaces. It is possible that these forms should be
treated respectively as var. lyallii and var. anomodum.
However, the association between frond form and
habitat is not strong, and many exceptions to this
tendency have been found making it difficult to
justify any taxonomic distinction between them.
58 New Zealand Journal of Botany 15, 1977

cm
r 2O

-10

- O

ifrj£\WF^

A
Fig. 13 Fronds of Asplenium lamprophyllum ( x I ) . A, NZ 560, Little Barrier Island; B, NZ 529. Smiths
Bush, Auckland.
Brownsey—N.Z. species of Asplenium
Hence, I prefer to regard A. lyallii as a single poly-
morphic species without any further formal sub-
division in rank.
8. ASPLENIUM LAMPROPHYLLUM Carse
Trans. N.Z. Inst. 56: 81 (1926) (Fig. 13)
TYPE: Lectotypc in CHR; lava flats, Mt. Wellington,
Auckland, coll. H. Carse, 30.5.1924, 27.10.1927.
DESCRIPTION : Rhizome creeping, up to 15 cm long,
pale green with a few scattered scales, stoloniferous.
Stipes 6-20 cm long, pale green above, brown below,
deeply grooved, covered in small, very dark, triangular
to ovate scales which have thick cell walls (Fig.
17F-J). Laminae lanceolate to elliptic, 15-60 X
7-20 cm, light green, glossy above, thin, bipinnate.
Raches green, scaly, prominently grooved. Pinnae
12-20 pairs, lanceolate to narrowly ovate, acuminate,
stalked, 3-10 X 0.5-2.5 cm, scaly on underside; lower
pinnae themselves pinnate, upper ones pinnatifld.
r*1?..
A. lamprophyllum
Fig. 14 Distributions of
A splenium lamprophyllum
and A. bulbijerum.
59
Pinnules sessile or shortly stalked, elliptic, often
deeply serrate, 10-20 X 5-15 mm. Sori 3-10 mm long,
nearer mid-vein than margin. Sporangia orange-
brown. Spores (30)38-42(50) Mm long, (20)24-29(35)
^m wide (4 populations), perispore very pale and not
prominently patterned (Fig. 30E-F).
CHROMOSOME NUMBER: n=72 (Fig. 32F). NZ 652,
Hikurangi, Whangarei; NZ 681, Smith's Bush, Auck-
land.
The same chromosome number was recorded by
Brownlie (1961) for material from New Plymouth.
HABITAT AND DISTRIBUTION: N. A. lamprophyllum
is an endemic species almost entirely confined to
regions north of latitude 38°N, with only very
scattered populations extending as far south as
Wanganui on the west coast (Fig. 14). It frequently
carpets extensive areas on rocky ground or clay banks
in shady forest, growing from its creeping rhizome
and spreading by stolons. It occurs from sea level
to 600 m.
u r
A. bulbiferum
60
COMMENT: A. lamprophyllum is sometimes con-
fused with A. bulbijerum but can easily be distin-
guished from this species by its creeping rhizome,
thick-walled stipe scales (cf. Fig. 17A-E. F-J),
glossy frond, absence of bulbils, long orange sori.
rather featureless spores (cf. Fig. 30E-F, K-L)
and the fact that it is never more than bipinnately
divided (cf. Figs 13, 15). It also frequently smells
of oil of wintergreen when crushed.
9. ASPLENIUM BULBIFERUM Forst. f. Prodr.:
80 (1786) (Figs 15. 16)
TYPE: Lectotype in BM; New Zealand, Dusky Bay.
coll. J. R. and G. Forster.
DESCRIPTION: Rhizome short, stout, erect, bearing
ovate scales up to 15 X 5 mm. Stipes 5-30 cm long,
brown on underside, green above, stout, covered in
small brown ovate scales. Laminae lanceolate to
elliptic, 15-120 X 7-30 cm, bi- to tripinnate, some-
times bearing bulbils. Raches green, scaly, promin-
ently grooved. Pinnae 15—30 (or more) pairs, ovate
to narrowly ovate, acuminate, shortly stalked, 3-20
X 1-5 cm, scaly on underside, basal pair pointing
downwards when fresh. Secondary pinnae sessile or
shortly stalked, very narrowly elliptic to ovate or
elliptic, obtuse, deeply serrate or sometimes almost
pinnate, decreasing in size from base to apex, basal
acroscopic pinnule often enlarged (up to 4 X 1 cm).
Ultimate pinnules narrowly oblong, ± entire to
crenate-serrate, up to 1 cm long. Sori numerous,
broad, submarginal, 2-4 mm long. Perispore of spores
irregularly ridged (Fig. 30K-N).
9a. A. BULBIFERUM Forst. f. subsp.
BULBIFERUM (Fig. 15)
DIAGNOSIS: Stipe scales ovate, lacking long apical
projections (Fig. 17A-E). Laminae pale green, usually
bulbiferous. Spores (33)41-48(55) /an long, (21)28-
32(37) fim wide (3 populations).
CHROMOSOME NUMBER: n = 72 (Fig. 32A). NZ 31,
Wilton Bush, Wellington; 2n=144, NZ 707, Bull
Stream, Akatarawa Range, Wellington.
Browniie (1954) also reported n = 72 in this
species.
9b. A. BULBIFERUM Forst. f. subsp.
GRACILLIMUM (Col.) Brownsey, comb. nov.
(Fig. 16)
BASIONYM: A. gracillimum Col. Trans. N.Z. lnst.
22: 453 (1890)
New Zealand Journal of Botany 15, 1977
TYPE: Holotype in WELT; Dannevirke. coll. W.
Colenso. (WELT P3348)
DIAGNOSIS: Very similar to subsp. bulbijerum but
differing in the following characters. Stipe scales with
apices drawn into long filiform projections (Fig.
17K-O). Laminae dark green, rarely bulbiferous.
Spores (33)41-48(55) i>m long, (21)28-32(37) ^m
wide (3 populations).
CHROMOSOME NUMBER: n=144 (Fig. 32B). NZ 661.
Karekare, Auckland; NZ 55, Lake Rotoaira, Tonga-
riro; NZ 1164, Carter's Bush. Gladstone, Wairarapa.
HABITAT AND DISTRIBUTION: N, S, St., Ch.. A. (Fig.
14). A. bulbijerum occurs commonly in wooded
areas from North Cape to Stewart Island, and is
absent only from the drier parts of South Canter-
bury and Central Otago where forest has been
removed. It is a characteristic component of the
ground-layer in most forests and grows from sea
level to 1 100 m. It has not been possible to prepare
separate distribution maps for the two subspecies of
A. bulbijerum because herbarium specimens can only
be distinguished by reference to micro-morphological
characters. However, it seems likely from personal
field observations that both subspecies are equally
widespread and probably occur sympatrically in
many areas.
COMMENT: The A. bulbijerum complex is wide-
spread and extremely polymorphic, and it has there-
fore attracted considerable attention from taxonomists
who have proposed many different names for its
various forms. It is now clear that, at least in New
Zealand, there are two different cytotypes in this
complex, and that some morphological distinction
is associated with the increase in chromosome num-
ber. Thus, the tetraploid form is usually bulbiferous.
has fronds which are often a pale green, stipe scales
which lack long filiform apices (cf. Fig. 17A-E,
K-O) and spores which are smaller (mean values
34-37 x 23-26 ^m). By contrast, the octoploid form
is rarely bulbiferous, has darker green fronds, more
attenuated stipe scales and larger spores (mean values
41-48 X 28-32 /xm). On this basis, the two cytotypes
have been referred to separate subspecies.
A. bulbijerum was first described by Forster, and
examination of his material in BM and GOET indi-
cates that plants collected by him were all bulbi-
ferous and had stipe sca'es and a spore size typical
of the tetraploid cytotype. The frond shape of his
specimens also corresponds to that of known tetra-
ploid plants. The name A. bulbijerum subsp. bulbi-
jerum is therefore retained for this cytotype.
At the octoploid level there is a much greater
range of frond dissection than amongst the tetraploid
Brownsey—N.Z. species of Asplenium 61

A
Fig. 15 Fronds of Asplenium bulbiferum subsp. bulbiferum ( X I ) . A, NZ 906. Rapid; B. NZ 284, Keith
George Park, Wellington; C, NZ 35, Wilton Bush, Wellington.

plants (cf. Figs 15. 16). At one extreme there
are forms with rather broad and only partially incised
secondary pinnae (e.g.. Fig. 16A) and, at the other,
forms which have their secondary pinnae completely
divided into rather narrow ultimate pinnules (e.g.,
Fig. 16D). However, it is clear from examination of
many herbarium specimens that these types have
many intermediates (e.g.. Fig. 16B, C) and that
there is a complete morphological continuum between
the two extremes. It is proposed, therefore, that
the octoploid cytotype should be treated as a single
polymorphic subspecies encompassing all these
varieties. However, individual populations rarely
include all of these forms and the more extreme
frond types are usually found in different localities.
It is possible therefore that some morphological
differentiation may be related to ecological factors,
but until it can be more thoroughly investigated in
relation to environmental conditions this subspecies
will not be further subdivided.
62 New Zealand Journal of Botany 15, 1977

Fig. 16 Fronds of Aspleniunt bulbiferum subsp. gracillimum ( x })• A, NZ 1071, Lighthouse Track, Stewart
Island; B, NZ 231, Stronvar, Wairarapa; C. NZ 656. Karekare, Auckland; D, NZ 71, Lake Rotoaira, Tongariro.
Browrtsey—N.Z. species of Asplenium 63

Fig. 17 Camera lucida drawings of stipe scales ( x 20), each taken from a different population. A-E, Asplenium
bulbiferum subsp. bulbiferum; F-J, A. lamprophyllum; K-O, A. bulbiferum subsp. gracillimum; P-S. A.
richardii; T-Z'. A. hookerianum (T-X, A. hookerianum, Y-Z', A. colensoi).
64
The most appropriate name for the octoploid
subspecies appears to be a new combination. A.
bulhiferum subsp. gracillimum (Col.) Brownsey.
Colenso's type specimen of A. gracillimum is very
similar to the frond illustrated in Fig. 16A and the
range of forms in his herbarium includes some similar
to Fig. 16B. The spore size and stipe scales of
Colenso"s material also fall within the range of varia-
tion exhibited by plants now known to be octoploid.
Frond forms similar to that in Fig. 16D have
been referred by some authors to A. bulbiferum var.
tripinnatum Hook. f. (e.g., Field 1890, plate XXII1.
fig. 6; Crookes 1963, p. 345). However, examination
of Hooker's type material indicates that his concept
of var. tripinnatum was quite different, the frond
being considerably more divided than anything illus-
trated in Fig. 16. It has very long narrow tertiary
pinnae which are up to 15 mm long and 4 mm wide,
and which are themselves divided into segments.
Furthermore, the sori are up to 5 mm long which is
significantly greater than for any other specimen of
A. bulbiferum I have examined. The possibility there-
fore exists that Hookers specimen may be a hybrid,
but because it is unfortunately immature this hypo-
thesis cannot be tested by reference to the spores,
and the taxon must remain of equivocal status.
Whatever its status may eventually prove to be,
there is no doubt that A. bulbiferum var. tripinnatum
has been widely misinterpreted by later workers, and
that forms such as Fig. 16D have been incorrectly
referred to it. Because these varieties have apparently
received no other taxonomic recognition, subsp.
gracillimum is interpreted here on a broader basis
than by Colenso and includes the entire range of
variation illustrated in Fig. 16. Hence, this taxon
is more clearly defined by reference to chromosome
number and spore size than it is by the degree of
frond dissection.
One other name which has been proposed
within the A, bulbiferum aggregate deserves further
mention. A. laxum R.Br. was described from Tas-
mania as a species very similar to A. bulbiferum
but differing in that it lacked bulbils. Examination
of the type specimen in BM, however, indicates that
the plant has aborted spores and is therefore very
probably a hybrid. Other collections made by Brown
in Australia and referred to A. laxum have good
spores, but clearly the name must now be applied
to the hybrid, whatever its parentage may be. The
representatives of the A. bulbiferum complex in
Australia need further investigation in the light of
the present revision of New Zealand material.
New Zealand Journal of Botany 15, 1977
10. ASPLENIUM HOOKERIANUM Col. Tasm. J.
nat. Sci. 2: 169 (1846) (Fig. 18)
A. colensoi Col. Tasm. J. nat. Ssi. 2: 170 (1846)
A. adiantoides Raoul Ann. Sci. nat. Ser. Ill 2:
115 (1844) non Lam. 1786
A. adiantoides Raoul var. minus Hook f. in Hook.
Ic. PI. 10: t.983 (1854)
A. adiantoides Raoul var. colensoi Hook. f. in
Hook. Ic. PI. 10: t.984 (1854)
A. adiantoides Raoul var. hookeriana Hook. f.
Fl. N.Z. 2: 35 (1855)
A. hookerianum var. colensoi Moore Ind. Fil.:
137 (1858)
A. richardii var. colensoi Hook. Sp. Fil. 3: 197
(1860)
A. ornatum Col. Trans. N.Z. Inst. 22: 452
(1890)
A. svmmetricum Col. Trans. N.Z. Inst. 31: 264
(1899)
TYPE: Holotype in WELT; between Matamata and
Maungatautari, coll. W. Colenso, January 1842.
DESCRIPTION: Rhizome short, erect, bearing numer-
ous brown, ovate, acuminate scales up to 10 X 2 mm.
Stipes 2-10 cm long, pale brown below, green above,
densely covered in small, subulate to narrowly ovate
scales with filiform apices (Fig. 17T-Z). Laminae
lanceolate to rhombic, 4—25 X 1-15 cm, dark green,
thin, normally bipinnate but often pinnate when
young and almost tripinnate in well-grown speci-
mens. Raches green, slender, and very scaly. Pinnae
5-15 pairs, very narrowly ovate to ovate, obtuse to
acuminate, long-stalked, 1-8 X 0.5-2 cm, basal pair
pointing upwards when fresh. Pinnules stalked, 3-10
pairs, linear to suborbicular, crenate to deeply
incised (or almost pinnate), 3-12 X 3-10 mm, tending
to lie at 90° to plane of frond in well-grown speci-
mens. Sori sub-marginal on narrow pinnules, remote
from margins on broad segments, 1-3 mm long.
Spores (25)31-37(42) /mi long, (19)23-27(33) ^m
wide (9 populations), perispore very prominently
winged and ridged (Fig. 30G-H).
CHROMOSOME NUMBER: n=72 (Fig. 33C). NZ 67.
Lake Rotopounamu, Tongariro; NZ 118, Okahukara
Forest, Tongariro; NZ 213, 277, Stronvar, Wairarapa;
NZ 912, Kapiti Island; NZ 848, Khandallah, Welling-
ton; NZ 318, Karori, Wellington; NZ 711, Linden.
Wellington; NZ 796, Takaka Hill, Nelson.
Brownlie (1954, 1958) recorded n = 72 for both
A. hookerianum and A. colensoi.
HABITAT AND DISTRIBUTION: N, S, St. A. hookerianum
is one of the most common and widespread species
of Asplenium in New Zealand, having been recorded
from Kaitaia to Stewart Island (Fig. 19). However,
there are no recent collections of the species from
further north than Kaipara Harbour and its present
Brownsey—N.Z. species of Asplenium 65
cm
15-1

IO-

5-

OJ

Fig. 18 Fronds of Asplenium hookerianum ( x 1). A, NZ 1145, Carters Bush, Wairarapa; B, NZ 933, Kapiti
Island; C, NZ 194, Stronvar, Wairarapa; D, NZ 306, Karori. Wellington; E. NZ 234. Stronvar, Wairarapa;
F, NZ 769, Sugarloaf. Port Hills, Christchurch; G. NZ 942, Whangiamoana, Wairarapa.
66
A. hookerianum
A. richardii
occurrence in the North Auckland peninsula needs
investigation. Tt grows from sea level to 1 250 m and
is found particularly on clay banks or rocky outcrops
in scrub and open forest, or on the ground in dis-
turbed forest remnants.
Outside New Zealand, this species has apparently
been recorded only once. Tn 1874, F. v. Mueller
collected a specimen (in MEL) from rocks at 3 000—
4 000 ft (900-1 200 m) on the Upper Hume River,
Australia. Willis (1962) states that this locality is
most likely to have been on the slopes of Mt
Kosciusko, on the New South Wales side of the
Murray River. Clearly this record needs confirmation,
but, even if the species is now extinct in Australia,
it suggests that A. hookerianum once had a much
more widespread distribution extending beyond New
Zealand.
COMMENT: AS understood here, A. hookerianum is
an extremely polymorphic species and one which
includes several previously described taxa. Of these.
New Zealand journal of Botany 15. 1977
Fig. 19 Distributions of
Asplenium hookerianum and
A. richardii.
the one which has received the most widespread
recognition is A. colensoi, a form with very narrow
pinnules. This name was first mentioned by Colenso
in a letter to W. J. Hooker (Hooker 1842), but not
validly published until 2 years later when descrip-
tions of both A. colensoi and A. hookerianum
appeared together (Colenso 1846). Moore (1858-
1859) later reduced A. colensoi to a variety of A.
hookerianum and this combination has been accepted
by most subsequent authors, although Crookes
(1963) retained both taxa as full species.
Colenso later described two more species, A.
ornatum and A. symmetricum. However, examination
of the type specimens (in AK and WELT respec-
tively) indicates that these are merely forms of his
earlier species A. hookerianum and A. colensoi, and
should therefore be rejected.
The real problem concerns the true status of A.
hookerianum and A. colensoi. In the wild, these two
forms are very frequently found together in the same
Brownsey—N.Z. species of Asplenium
populations and, indeed, it is rather unusual to find
extreme forms of either type growing alone. The
polymorphism in frond shape is particularly notice-
able amongst young plants, different individuals
producing either rather broad pinnules, or much
narrower pinnules. However, as they reach maturity,
both in the wild, and especially in cultivation, there
is a strong tendency for plants of both forms to
produce fronds of an intermediate shape, possessing
neither the very broad pinnules of A. hookerianum
nor the very narrow ones of A. coletuoi. Further-
more, the two taxa show the same rather narrow
range of variation with respect to spore pattern, spore
size (mean values 31-35 X 23-27 iim in A. hookeri-
anum, and 31-37 X 23-26 /im in A. colensoi) and
the shape of the stipe scales (cf. Fig. 17T-X, Y-Z').
Cytologically they are both uniformly tetraploid.
Hence there can be no justification for maintaining
two separate species. Because the two names were
published simultaneously neither has priority; accord-
ingly I have chosen to retain only A. hookerianum
because this is the more widely used and therefore
more appropriate name.
It is possible that fronds bearing very narrow
pinnules should continue to be recognised as a dis-
tinct variety of A. hookerianum. However, in practice
it is frequently very difficult to assign a particular
specimen to one or other taxa on this basis. Thus,
many plants have very narrow pinnules on their
youngest fronds, but gradually acquire broader pin-
nules as they get older (and vice versa). Further-
more, although there is a considerable range of
variation with respect to frond dissection, the
extremes of variability are often found together and
are not obviously correlated with particular habitats
or geographical locality. Recognition of var. colensoi
as a separate taxon is therefore difficult to justify
and it has been rejected.
A. hookerianum is usually a much smaller, more
delicate plant than A. bulbiferum, but confusion
between large plants of the former and small plants
of the latter sometimes arises. Large specimens of
A. hookerianum can be distinguished by the presence
of very delicate but nevertheless quite distinct stalks
to the pinnules. Small plants of A. bulbiferum
generally have sessile pinnules, or very short broad
stalks. A. bulbiferum also often has a narrow frond
(more than twice as long as broad) which is flattened
in one plane, and a basal pair of pinnae which point
downwards in fresh specimens. A. hookerianum has
fronds which are often less than twice as long as
broad (Fig. 18), pinnules tending to lie at 90° to
the plane of pinnae and raches, and a lower pair of
pinnae which point upwards.
67
11. ASPLENIUM RICHARDII (Hook, f.) Hook, f.
Fl. N.Z. 2: 35 (1855) (Fig. 20)
A. adiantoides Raoul var. richardii Hook. f. in
Hook. Ic. PI. 10: t.977 (1854)
A. raoulii var. richardii Mett. U. e. Farngatt.:
118 (1859)
TYPE: Holotype in P; New River, Southern Island,
N.Z., herb. A. Richard.
DESCRIPTION: Rhizome short, stout, erect, bearing
dark brown subulate scales up to 20 X 2 mm. Stipes
5—15 cm long, brown on underside, green above,
densely covered in subulate scales with filiform apices
(Fig. 17P-S). Laminae ovate to narrowly ovate,
10-25 X 4-12 cm, dark green, relatively thin, normally
tripinnate. Raches green, very scaly, slightly grooved.
Pinnae 10-15 crowded and overlapping pairs, ovate
to narrowly ovate, sub-acute, stalked, 2-8 X 1-4 cm.
Secondary pinnae stalked, ovate, 10-20 X 10-15 mm,
again pinnate or pinnatifid. Ultimate segments linear,
acute or sub-acute, up to 8 mm long. Pinnae and
pinnules not flattened in one plane but spreading in
three dimensions. Sori 2-4 mm long, submarginal.
Spores (38)44-49(55) pm long, (27)32-36(40) Mm
wide (6 populations), perispore prominently winged
and ridged (Fig. 30I-J).
CHROMOSOME NUMBER: n=144 (Fig. 32E). NZ 422,
Hooker Valley, Mt Cook.
This number was also recorded by Iirownlie
(1954).
HABITAT AND DISTRIBUTION: N, S. A. richardii is an
endemic species almost entirely confined to mountain
districts of the South Island, being found in the
North Island only on Mt Honokawa, Mt Ruapehu,
and parts of the Kaimanawa Range (Fig. 19). It
extends throughout the mountain regions of the South
Island, growing on rocky outcrops, in crevices,
amongst boulders, and on stream banks particularly
under beech forest. It is found on both limestone
and greywacke rock and occurs up to 1 400 m, but
is more frequent east of the Main Divide.
COMMENT: A. richardii can be confused with well-
grown specimens of A. hookerianum, a species to
which it is closely related but from which it is clearly
distinguished by its octoploid chromosome number
and its larger spores (cf. Fig. 30G-H, I-J).
A. richardii can also be confused with A. terrestre,
a species with which it often grows in close associa-
tion. The two taxa are both octoploid and have
very similar spores, but A. richardii is characterised
by its thinner frond with long narrow pinnules. (Fig.
20), its shorter sori (2-4mm), the dense covering
 New Zealand Journal of Botany 15. 1977

Fig. 20 Fronds of Asplcnium richardii (X i). A, D. NZ 419. Black Birch Stream. Mt Cook; B. NZ 401.
Glencoe Creek, Mt Cook; C, NZ 271, Cobb Valley, Nelson.

of scales on the stipe and rachis, and by its habit 12. ASPLEN1UM FLACCIDUM Forst. f. Prodr.:
of producing pinnae and pinnules in three dimensions. 80 (1786) (Figs 21, 23)
A. terrestre has a more leathery frond with broad
pinnules flattened into one plane (Fig. 25), longer Coenopteris flaccida Thunb. Nov. Act. Pet. 9:
sori (2-7 mm), and fewer scales on the stipe and 158, t.D, f.1-2 (1795)
C. novae-zelandiae Spreng. Schrad. J. Bot. 2: 269
rachis. (1799)
Fig. 21 Fronds of Asplenium fiaccidum subsp. fiaccidum ( x i ) . A. NZ 38. Wilton Bush, Wellington; B.
NZ 441. Haast, South Westland; C. NZ 387, Terenikau Valley. Tararuas; D. NZ 63, Lake Rotopounamu.
Tongariro: E. NZ 1004. Lake Fergus. Fiordland.
70
Darea fiaccida Willd. Sp. PI. 5: 245 (1810)
D. odontites Willd. Sp. PI. 5: 245 (1810)
A. heterophyllum A. Rich. Essai Fl. N.Z.: 74
(1832)
TYPE: Lectotype in BM; New Zealand, Dusky Bay,
coll. J. R. and G. Forster.
DESCRIPTION: Rhizome short, stout, erect, bearing
dark brown subulate or ovate scales up to 20 X 2 mm.
Stipes 5-20 cm (or more) long, brown on underside,
green above, flaccid or firm and erect, sparingly
covered in small subulate or ovate scales. Laminae
lanceolate to elliptic, 10-90 (or more) X 4-25 cm,
dull green, thick, leathery and either limp and
pendulous or stiff and erect, pinnate to bipinnate.
Raches green, sparingly scaly. Pinnae in 5-20 (or
more) pairs, linear to narrowly ovate, acuminate,
long stalked, 2-15 X 0.5-2 cm; degree of dissection
very variable, sometimes only divided into very short
obtuse segments, sometimes pinnate with linear acute
pinnules up to 15 X 2 mm. Sori submarginal, linear,
2-10 mm long.
12a. A. FLACCIDUM Forst. f. subsp.
FLACCIDUM (Fig. 21)
DIAGNOSIS: Rhizome bearing subulate scales. Stipes
5-20 cm (or more) long, flaccid, bearing small subu-
late scales with long filiform apices (Fig. 24A-E).
Laminae lanceolate to elliptic, 15-90 (or more) X
5-25 cm, leathery, limp and pendulous. Pinnae 5-20
(or more) pairs, linear, 5-15 X 0.5-2 cm. Pinnules
very variable in length, from oblong and obtuse to
linear and acute, up to 15 X 2 mm. Basal acroscopic
pinnule occasionally much longer than that next to it.
Sori 2-10mm long. Spores (31)36^44(50) /im long,
(19)23-27(33) pm wide (10 populations), perispore
with a prominent wing but lacking ridges (Fig.
29M-N). Plant usually epiphytic, but sometimes
terrestrial.
CHROMOSOME NUMBER: n = 72 (Fig. 33A). NZ 857,
Renata Ridge, Akatarawa Range, Wellington; NZ 32,
36, Wilton Bush, Wellington; NZ 840, Seatoun,
Wellington.
Brownlie (1954) reported n = 72 for A. flaccidum
sens. lat.
HABITAT AND DISTRIBUTION: K, N, S, Ch., St. (Fig.
22). A. flaccidum subsp. flaccidum is a common fern
throughout the forested regions of New Zealand
from North Cape to Stewart Island, being absent only
in the drier parts of Otago and Canterbury from
which the forest has been cleared. It grows principally
New Zealand Journal of Botany 15, 1977
as an epiphyte on various native trees, occurring
from sea level to the tree line, but is sometimes also
found on the ground. Outside New Zealand its distri-
bution includes parts of Australia and some of the
Pacific Islands.
12b. A. FLACCIDUM Forst. f. subsp.
HAURAKIENSE Brownsey, subsp. nov. (Fig. 23)
Rhizoma paleis ovatis cum parietibus cellularum
crassissimis vestitum. Stipes 5-12 cm longus, erectus,
firmus, paleis anguste vel late ovatis cum parietibus
cellularum crassissimis vestitus (Fig. 24F-I). Lamina
oblonga vel elliptica, 10-30 X 4-15 cm rigida, erecta.
Pinnae 5-12 jugatae, lineares vel anguste ovatae,
20-80 X 8-16 mm. Pinnulae oblongae obtusaeque vel
lineares acutaeque, usque ad 7 X 2 mm. Pinnula
basalis acroscopica multo longior quam pinnula
vicina, usque ad 4 cm longa, pinnatifida. Sori 2-7 mm
longi. Sporae (38)43-49(54) urn longae, (20)26-
31(38) pm latae; perisporae interdum alatae,
plerumque non cristatae. Numerus somaticus
chromosomatum 144; in meiosi paria 72 observantur.
Habitat in rupibus orae.
HOLOTYPUS: CHR; NZ 894, Oneroa Beach, Waiheke
Island, Auckland (NZMS 1 N42:528688). Growing
amongst coastal rocks. Coll. PJ. Brownsey, 1.9.1974.
(CHR 308926)
DIAGNOSIS: Rhizome bearing ovate scales with very
thick cell walls. Stipes 5-12 cm long, erect, firm,
bearing narrowly to broadly ovate scales with very
thick cell walls (Fig. 24F-I). Laminae oblong to
elliptic, 10-30 X 4-15 cm, stiff, erect. Pinnae 5-12
pairs, linear to narrowly ovate, 20-80 X 8-16 mm.
Pinnules oblong and obtuse to linear and acute, up
to 7 X 2 mm. Basal acroscopic pinnule frequently
much longer than that next to it, up to 4 cm long,
and itself pinnatifid. Sori 2-7 mm long. Spores
(38)43-49(54) Mm long, (20)26-31(38) Mm wide (10
populations); perispore sometimes winged, usually
lacking ridges (Fig. 29 O-P). Plant of coastal rocks.
CHROMOSOME NUMBER: n=72 (Fig. 33B). NZ 640,
Lion Rock, Bay of Islands; NZ 582, Shag Point,
Little Barrier Island; NZ 894, Oneroa, Waiheke
Island.
HABITAT AND DISTRIBUTION: N. A. flaccidum subsp.
haurakiense is found along the eastern side of the
North Auckland peninsula from the Three Kings
Islands to the Hauraki Gulf and Coromandel, extend-
ing locally into the Bay of Plenty (Fig. 22). It is so
far not known outside New Zealand. Plants of this
subspecies are almost always found growing terres-
trially and are restricted to coastal areas within the
immediate vicinity of salt spray, either on exposed
rock or amongst coastal scrub. The species is replaced
on Auckland's west coast by A. terrestre subsp.
maritimum (Fig. 26).
Brownsej—N.Z. species of Asplenium
Fig. 22 Distribution of
Asplenium ftaccidum.
A. flaccidum
subsp. flaccidum
COMMENT: The present investigation has shown that
the A. flaccidum aggregate comprises two quite
separate cytotypes, tetraploid and octoploid, and that
the associated morphological differences are sufficient
to warrant the recognition of two distinct species,
A. flaccidum and A. terrestre. Each of these can be
further subdivided into two subspecies distinguished
largely on the basis of ecological differentiation.
Material of A. flaccidum in BM and GOET
collected by Forster has been examined. The name is
retained here for plants which are usually epiphytic,
have long pendulous leathery fronds, and which are
never more than bipinnately divided (Fig. 21). They
have small subulate stipe scales (Fig. 24A-E), spores
which are prominently winged but rarely ridged
(Fig. 29M-N), and are tetraploid. There remains a
wide range of frond forms within this species, but
it is a more clearly circumscribed taxon than has
been interpreted by other authors and it corresponds
closely to material named by Forster.
A distinct ecological race of this species is des-
cribed as A. flaccidum subsp. haurakiense. This is
71
A. flaccidum
subsp.
haurakiense
similar to subsp. flaccidum in having many of the
same frond characters, an almost identical spore
pattern, and being tetraploid. It is distinguished
primarily by its coastal habitat and restricted geo-
graphical range, its firm erect habit, its larger spores
(cf. Fig. 29M-N. O-P), and by its very thick-
walled, ovate, stipe scales (cf. Fig. 24A-E, F-I).
13. ASPLENIUM TERRESTRE Brownsey, sp. nov.
(Figs 25, 27)
Rhizoma breve, erectum, paleis fuscis subulatis
usque ad 15 X 2 mm obtectum. Stipes 5—15 cm
longus, infra fuscus, supra viridis, paleis subulatis
vel anguste triangularibus cum apicibus longis
filiformibus vestitus (Fig. 24J-N, Q-V). Lamina
lanceolata vel elliptica, 10-30 X 5-20 cm, atroviridis,
coriacea aut crassissima carnosaque, cernua aut rigida
erectaque, bi- vel tri-pinnata. Rhachis viridis,
squamata, manifeste canaliculata. Pinnae 7-20
jugatae, inferiores ovatae, superiores peranguste
ovatae vel ellipticae, acutae vel acuminatae, 3-10 X
0.5-3 cm, paleis minutis infra vestitae. Pinnulae
superiores lineares acutae integrae sessilesque;
inferiores ovatae vel anguste ellipticae, pinnatifidae
 cm
20-i

10-

D
Fig. 23 Fronds of Asplenium flaccidum subsp. haurakiense ( x t ) . A, NZ 641, Lion Rock, Bay of Islands; B.
NZ 1132, Poor Knights Islands; C, D, NZ 578, Awaroa Stream, Little Barrier Island; E, NZ 890, Mataitai,
Waiheke Island.
Brownsey—N.Z. species of Asplenium

Fig. 24 Camera lucida drawings of stipe scales ( x 20), each taken from a different population. A-E, Asplenium
flaccidum subsp. flaccidum; F-I, A. flaccidum subsp. haurakiense; J-N, A. terrestre subsp. terrestre; O-P,
A. shuttleworthianum; Q-V, A. terrestre subsp. maritimum.
74
vel pinnatae, petiolatae, usque ad 30 X 10 mm. Sori
submarginales, 2-7 mm longi. Sporae (38)43-51(56)
/tm longae, (25)29-34(38) /ira latae, perisporae mani-
feste alatae cristataeque (Fig. 30 O-R). Numerus
somaticus chromosomatum 288; in meiosi paria 144
observantur.
HOLOTYPUS: CHR; NZ 96, Blowhard Bush, Napier-
Taihape Road (NZMS 1 N123:861491). Growing on
the ground beneath mixed forest. Coll. P. J. Brownsey,
30.12.1973. (CHR 308925)
DESCRIPTION: Rhizome short, erect, bearing dark
brown subulate scales up to 15 X 2 mm. Stipes 5-15
cm long, brown below, green above, covered in
subulate to narrowly triangular scales with long
filiform apices (Fig. 24J-N, Q-V). Laminae lanceolate
to elliptic, 10-30 X 5-20 cm, dark green, leathery or
very thick and fleshy, slightly drooping or stiff and
upright, bi- to tripinnate. Raches green, scaly,
prominently ridged. Pinnae 7-20 pairs, those at the
base ovate, those above very narrowly ovate or
elliptic, acute to acuminate, stalked, 3-10 X 0.5-3 cm,
covered on underside in tiny scales. Upper pinnules
linear, acute, entire and sessile; lower pinnules ovate
to narrowly elliptic and pinnatifid or pinnate, up to
30 X 10 mm. Sori submarginal, 2-7 mm long. Spores
(38)43-51(56) f«n long, (25)29-34(38) ^m wide (14
populations), perispore prominently winged and
ridged (Fig. 30 O-R). 288 chromosomes in the
sporophyte; 144 bivalents at meiosis.
HABITAT AND DISTRIBUTION: N, S. (Fig. 26). A.
terrestre is an endemic species with a distribution
confined to the North and South Islands and extend-
ing from Auckland to Bluff. It is a terrestrial plant
found growing on the forest floor, on bluffs and
rocky outcrops, or on sea cliffs.
13a. A. TERRESTRE Brownsey subsp. TERRESTRE
(Fig. 25)
DIAGNOSIS: Laminae lanceolate to elliptic, 10-30 X
5-15 cm, thick and leathery, usually slightly droop-
ing. Pinnae 8-20 pairs, those at the base ovate, those
above very narrowly ovate or elliptic, often with
long relatively undivided acuminate tips. Upper
pinnules linear and entire, lower pinnules narrowly
elliptic and pinnatifid to pinnate, up to 30 X 8 mm.
CHROMOSOME NUMBER: n=144 (Fig. 33D). NZ 121,
Okahukara Forest, Tongariro; NZ 96, Blowhard Bush,
Napier-Taihape Road; NZ 312, Karori, Wellington;
NZ 270, Woodside Creek, Marlborough.
HABITAT AND DISTRIBUTION: N, S. (Fig. 26). A.
terrestre subsp. terrestre is found principally in the
forests of North Canterbury and Marlborough, but
New Zealand Journal of Botany 15, 1977
extends locally to many parts of the South Island.
In the North Island it occurs mainly on the volcanic
plateau and in the mountains of the Hawke"s Bay
area, but also reaches Rotorua, Mt Egmont, and
the Rimutakas. It grows terrestrially in forested areas
but may also be found on bluffs and rocky outcrops
up to 1 100 m.
13b. A. TERRESTRE Brownsey subsp.
MARITIMUM Brownsey, subsp. nov. (Fig. 27)
Lamina pro ratione brevis lataque, elliptica, 10-25
X 7-20 cm, crassissima, carnosa, rigida, erecta.
Pinnae 7-15 jugatae, inferiores ovatae, superiores
anguste ovatae, acutae. Pinnulae superiores lineares
integraeque, inferiores ovatae vel ellipticae pinna-
tifidaeque, usque ad 20 X 10 mm. Numerus somaticus
chromosomatum 288; in meiosi paria 144 observantur.
HOLOTYPUS: CHR; NZ 12, Ohiro Bay, Wellington
(NZMS 1 Nl64:300142). Growing on greywacke
coastal cliffs. Coll. P. J. Brownsey. 15.12.1973.
(CHR 308927)
DESCRIPTION: Laminae relatively short and broad,
elliptic, 10-25 X 7-20 cm, very thick and fleshy, stiff
and upright. Pinnae 7-15 pairs, those at the base
ovate, those above narrowly ovate, acute. Upper
pinnules linear and entire, lower pinnules ovate to
elliptic and pinnatifid, up to 20 X 10 mm.
CHROMOSOME NUMBER: n=144 (Fig. 33E). NZ 669,
Karekare, Auckland; NZ 508, Manaia, Taranaki; NZ
12, Ohiro Bay, Wellington.
HABITAT AND DISTRIBUTION: N, S. (Fig. 26). A.
terrestre subsp. maritimum has a distribution centred
on Cook Strait where it occurs more or less con-
tinuously around the Wellington coast from Cape
Palliser to Pukeraa Bay and Kapiti. It is common in
the Marlborough Sounds, and extends down the
Marlborough coast as far as Banks Peninsula. Else-
where it is known only from Cape Farewell, the
Taranaki coast, and from the west coast of Auck-
land. It is found on coastal cliffs within the influence
of salt spray, either on exposed rocks where plants
are rather stunted, or amongst coastal scrub where
the plants grow much bigger.
COMMENT: A. terreslye is a species which has been
included within A. flaccidum by most previous authors
but which is clearly distinct from that taxon because
of its octoploid chromosome number, its more highly
divided frond (cf. Figs 21, 23, and 25, 27), and by
its more prominently ridged spore pattern (cf. Figs
29M-P, 30 O-R). It differs from the epiphytic
A. flaccidum subsp. flaccidum in growing terrestrially
and having a more or less erect frond and a larger
spore size. It can also be distinguished from A.
Brownsey—N.Z. species of Asplenium 75

c
Fig. 25 Fronds of Asplenium terrestre subsp. terrestre ( x 1). A, NZ 1042, Bluff Hill, Southland; B. NZ
771, Sugarloaf, Port Hills, Christchurch; C. NZ 724, Woodside Creek, Marlborough; D. NZ 403. Glencoe
Creek, Mt Cook; E, NZ 109, Okahukara Forest, Tongariro.
76
A. terrestre
subsp.
terrestre
A. terrestre
subsp.
maritimum
flaccidum subsp. haurakiense by the lack of a greatly
elongated basal acroscopic pinnule on each pinna
(cf. Figs 23. 27).
A. flaccidum subsp. flaccidum frequently occurs
together with A. terrestre subsp. terrestre in forested
areas; in parts of the Cook Strait region where forest
extends to the coast it also overlaps with subsp.
maritimum. In both these areas hybrids between A.
flaccidum and A. terrestre are reasonably common.
They will be described in more detail in another
publication, but it is sufficient to say here that the
hybrids produce only aborted spores and are com-
pletely sterile. The recognition of two separate species
is therefore fully justified not only on morphological
and cytological evidence but also on genetical
grounds.
The distinction between the two subspecies of
A. terrestre is primarily ecological, subsp. terrestre
being largely confined to inland areas, and subsp.
maritimum to coastal regions, but there is also some
degree of morphological differentiation. Thus, subsp.
maritimum has a much thicker and more fleshy frond,
New Zealand Journal of Botany 15, 1977
Fig. 26 Distribution of
Asplenium terrestre.
and pinnae which are generally broader than those
in subsp. terrestre (cf. Figs 25, 27).
It is possible that A. terrestre was first detected
as a distinct species by Armstrong (1882) when he
described A. canterburiense from various localities
in Canterbury. His description is very similar to that
given here for A. terrestre, but the only specimen
of his which I have been able to locate* consists of
a single frond which has aborted spores and is
clearly a hybrid, probably between A. terrestre and
A. richardii. This hybrid is extremely similar to A.
terrestre (Brownsey, in preparation) and probably
occurs reasonably frequently wherever the two
parents are found together. Armstrong's collections
from other localities cited in his original description
of A. canterburiense may have included plants of
the parent species as well as the hybrid, but until
such specimens are located the name A. canter-
buriense must obviously be retained for A. richardii
X terrestre.
"One sheet in the Armstrong Herbarium, Christchurch
Botanic Gardens, labelled Mt Arrowsmith, 4 000 It.
coll. J. F. Armstrong. March 1869.
Fig. 27 Fronds of Asplenium terrestre subsp. marilimum (X {). A, NZ 929. Kapiti Island; B, NZ 509.
Manaia, Taranaki; C, NZ 673, Karekare, Auckland; D. NZ 353. Titahi Bay. Wellington; E. NZ 838. Seatoun,
Wellington; F. NZ 837, Seatoun. Wellington.
78
Fronds of A. terrestre subsp. terrestre may some-
times be confused with those of A. bulbiferum
subsp. gracillimum (cf. Figs 16B, 25B). However,
A. terrestre subsp. terrestre can usually be
recognised by its thicker, more leathery frond, by
its longer sori (2-7 mm), and sometimes by the
presence of long, relatively undivided, acuminate tips
to the pinnae. A. bulbiferum subsp. gracillimum has
a thinner frond, shorter sori (2-4 mm), and usually
has pinnae which are divided throughout their length
into more or less distinct pinnules.
14. ASPLENIUM SHUTTLEWORTHIANUM
Kunze Farrnkr. 1: 26, t.14 (1840) (Fig. 28)
A. flaccidum var. shuttleworthianum Hook. f.
Handbk. N.Z. Fl.: 274 (1864)
TYPE: BM; Pitcairn Island, coll. Cuming 1374.
DESCRIPTION: Rhizome short, erect, bearing red-
brown, linear attenuate scales. Stipes up to 15 cm
long, green above, brown below, sparingly covered in
small linear scales with long filiform apices (Fig. 24
O-P). Laminae oblong to elliptic, 15-90 X 10-25 cm,
yellow-green, coriaceous, 3- to 4-pinnate. Raches
green, almost lacking scales, prominently grooved.
Pinnae ovate to narrowly ovate, acuminate, stalked,
A oJ
New Zealand Journal of Botany 15, 1977
up to 12 X 5 cm. Secondary pinnae linear to lanceo-
late, up to 15 X 2 mm, often pinnatifid. Ultimate
segments linear, subacute, slightly expanded in the
region of the sori. Sori near tips of ultimate seg-
ments, solitary, broad, submarginal, c.2mm long.
Spores (46)51-63(71) ^m long, (32)34-44(50) Mm
wide (3 populations), perispore prominently winged,
granular, but not conspicuously ridged (Fig. 30C-D).
CHROMOSOME NUMBER: n=c.l44. NZ 1112, Raoul
Island, Kermadecs.
HABITAT AND DISTRIBUTION: K. In the New Zea-
land region, A. shuttleworthianum is known only
from the Kermadecs where it apparently occupies
various habitats from coastal scrub to inland forest,
and where it grows both terrestrially and epiphytic-
ally. It also occurs on Pitcairn and other Pacific
islands.
COMMENT: This species is closely related to A.
terrestre but distinguished from it by a greater degree
of frond dissection (cf. Figs 25, 28), its very
large spores (cf. Fig. 30C-D, O-P), and by the
characteristic broadening of the ultimate pinnules in
the region of the sori. Because of its remote geo-
graphic distribution it has been very little studied.
cm
2 On
10-
Fig. 28 Fronds of
Asplenium shuttleworth-
B ianum ( x I ) . A, NZ 690,
Raoul Island; B, NZ 1112,
Raoul Island.
Brownsey—N.Z. species of Asplenium

••

K <

N o
Fig. 29 Exospore and perispore of spores of Asplenium species (all X 400).
A-B, A. lucidum, NZ 552, Little Barrier Island; C-D, A. flabellifolium, NZ 178.
Kapiti Island; E-F, A. obtusatum subsp. obtusatum, NZ 790, Wharariki Beach.
NW Nelson; G-H, A. obtusatum subsp. northlandicum, NZ 687, Slipper Island.
Bay of Plenty; I-J, A. scleroprium, CHR 195053. Ernest Island, off Stewart
Island; K-L, A. lyallii, NZ 810, Takaka Hill, Nelson; M-N, A. flaccidum
subsp. ftaccidum, NZ 378, Bay of Many Coves, Marlborough; O-P. A. flaccidum
subsp. haurakiense, NZ 890, Mataitai. Waiheke Island.

DISCUSSION relatives in the Northern Hemisphere. Nevertheless,

The New Zealand representatives of Asplenium
undoubtedly have their closest affinities with the
flora of temperate parts of Australia. Some species,
such as A. bulbiferum and A. flaccidum, form a
characteristic component of the forest vegetation in
both regions. Other species, however, have affinities
with more distant floras. A. obtusatum, for example,
is a common coastal plant in higher latitudes of the
entire Southern Hemisphere; A. polyodon is primarily
a plant of tropical regions, whereas A. trichomanes
is of cosmopolitan distribution and has its closest
it is clear that most of these species must have
ultimately arrived in New Zealand from Australia,
presumably as a result of spores being blown across
the Tasman Sea in the predominating westerly air-
stream.
A striking feature of Asplenium in New Zealand
is the degree of polymorphism shown by its different
species, a fact which has been remarked on by many
workers, and which is probably due to divergent
evolution in the relatively impoverished flora of this
oceanic archipelago. The number of species which
80 New Zealand Journal of Botany 15, 1977

s \ D
/ •

J '

Fig. 30 Exospore and perispore of spores of Asplenium species (all X 400).

A-B, A. polyodon, NZ 295, Keith George Memorial Park, Wellington; C-D,
A. shuttleworthianum, NZ 690, Raoul Island; E-F, A. lamprophyllum, NZ 560,
Little Barrier Island; G-H, A. hookerianum, NZ 796, Takaka Hill, Nelson;
I-J, A. richardii, NZ 426, Hooker Valley, Mt Cook; K-L, A. bulbiferum subsp.
bulbiferum, NZ 911, Kapiti Island; M-N, A. bulbiferum subsp. gracillimum, NZ
231, Stronvar, Wairarapa; O-P, A. terrestre subsp. terrestre, NZ 794, Takaka Hill.
Nelson; Q-R, A. terrestre subsp. maritimum, NZ 505, Manaia, Taranaki.

have successfully established themselves is rather Nevertheless, morphological diversification is not

small, but many of these have almost certainly been
able to spread into a greater variety of habitats than
in the continental and tropical floras from which
they have migrated, and evolution in response to a
variety of new environmental conditions has there-
fore taken place.
the only change to have resulted from the evolution
of these species. Hybridism has also played its part,
and the occurrence of intermediate forms has almost
certainly contributed to the inability of different
authors to consistently define the limits of the New
Zealand Asplenium species. Taxa whose distributions
Brownsey—N.Z. species of Asplenium 81

* i . •
/

Fig. 31 Permanent acetocarmine preparations of diakinesis (all X 1 000). A, Asplenium

flabellifolium, NZ 789, Wharariki Beach, NW Nelson, showing 210-211 univalents; B, A.
flabellifolium, NZ 834, Turakirae Head, Wellington, showing 277-280 univalents; C, A.
obtusatum subsp. obtusatum, NZ 472, Cape Foulwind, Westport, showing 72 bivalents; D,
A. lucidum, NZ 462, Greymouth, showing 72 bivalents; E, A. obtusatum subsp. northlandicum,
NZ 667, Karekare, Auckland, showing 144 bivalents.
82 New Zealand Journal of Botany 15, 1977

A B

V 7 **• ,

Fig. 32 Permanent acetocarmine preparations of diakinesis (all X 1 000). A, Asplenium

bulbiferum subsp. bulbiferum, NZ 707. Bull Stream, Akatarawas, showing 72 bivalents; B,
A. bulbiferum subsp. gracillimum, NZ 55, Lake Rotoaira. Tongariro, showing 144 bivalents;
C, A. scleroprium, NZ 1012, Bluff, showing 144 bivalents; D, A. lyallii, NZ 986, Castle Hill,
Canterbury, showing 144 bivalents; E, A. richardii, NZ 422, Hooker Valley, Mt Cook,
showing 144 bivalents; F. A. lamprophyllum, NZ 652. Hikurangi. Whangarei. showing 72
bivalents.
Brownsey—N.Z. species of Asplenium

0
% ^ -

E
Fig. 33 Permanent acetocarmine preparations of diakinesis (all X 1 000). A,
Asplenium flaccidum subsp. ftaccidum, NZ 36, Wilton Bush, Wellington, showing
72 bivalents; B, A. flaccidum subsp. haurakiense, NZ 640, Lion Rock, Bay of
Islands, showing 72 bivalents; C, A. hookerianum, NZ 796, Takaka Hill, Nelson,
showing 72 bivalents; D, A. terrestre subsp. terrestre, NZ 96, Blowhard Bush,
showing 144 bivalents; E, A. terrestre subsp. maritimum, NZ 669, Karekare,
Auckland, showing 144 bivalents.

overlap do occasionally hybridise, and, in disturbed
areas, particularly where forest has been opened up,
some species spread more prolifically than usual
and may give rise to several hybrid individuals.
However, unlike many of the polymorphic genera
of trees and shrubs in New Zealand, hybrids between
even closely related species of Asplenium are usually
sterile, and the idea that there are abundant hybrid
swarms exhibiting various degrees of introgression
is quite erroneous. Most hybrid individuals can be
immediately recognised by their aborted and shrivelled
spores and therefore provide a useful reference point
for denning the limits of their parent species.
A crucial role in the evolution of Asplenium
in New Zealand has been played by polyploidy and
84
the present investigation has revealed several new
and unsuspected cytotypes amongst the more com-
mon species. One such example is the A. flaccidum/
terrestre aggregate. Here, increase in chromosome
number has been accompanied by morphological
divergence and the establishment of a sterility barrier
between the two cytotypes. Clearly, the extreme poly-
morphism of A. flaccidum remarked on by earlier
authors was partly due to the inclusion within the
one taxon of two quite separate species which are
themselves now known to be fairly polymorphic. A
similar situation almost certainly exists amongst
other species aggregates for which more than one
chromosome number has now been determined.
It is very significant that all the newly discovered
cytotypes are at the octoploid level. Moreover, it is
remarkable that within the New Zealand Asplenium
flora there is not a single diploid species. The genus in
this country includes approximately equal numbers of
tetraploids (9) and octoploids (8) and displays a
higher mean level of ploidy (5.9) than in any other
area for which sufficient records exist (cf. Lovis
1973). The situation in New Zealand contrasts
strongly with that in Europe where over half the
Asplenium species are diploid and the mean level of
ploidy is only 2.9. In fact, there is a gradual increase
in the percentage of diploid species and a decrease
in the mean level of ploidy as one moves northwards
from New Zealand through the tropics and into the
Northern Hemisphere. This has led Lovis (1973) to
speculate that Asplenium may have originated in
Gondwanaland with "the main diversification occur-
ring after the genus had reached tropical latitudes,
only subsequently migrating into northern temperate
regions". Thus, Asplenium may now display its
highest level of ploidy in the areas where it has
existed longest. Lovis supports this hypothesis by
pointing out that, in contrast to such northern genera
as Dryopteris and Cystopteris which are deciduous
and eminently well adapted to the seasonal climate
of northern Europe, Asplenium is evergreen and
many species may be found undergoing meiosis
throughout the year.
Nevertheless, many of the higher polyploid sexual
species of Asplenium in New Zealand are endemic
and have almost certainly originated here, whereas
the tetraploid species are very largely of wider distri-
bution. Thus, there are three natural groups of closely
related species in New Zealand: (1) the A. flaccidum
aggregate with thick and rather leathery bipinnate
fronds, (2) the A. hookerianum/bulbiferum complex
with thin highly dissected fronds, and (3) the A.
lucidumjobtusatum complex with thick fleshy pinnate
fronds.
New Zealand Journal of Botany 15, 1977
Within the A. flaccidum aggregate there are just
two species*; of these, one (A. flaccidum sens, str.)
is tetraploid and widely distributed in Australia, New
Zealand, and Polynesia; the other (A. terrestre) is
octoploid and thought to be endemic to New Zea-
land. Botanists familiar with the floras of both
Australia and New Zealand have often remarked on
the polymorphism and wide ecological range of A.
flaccidum in this country compared to its more
uniform morphology and constant epiphytic habit
in Australia, suggesting that the terrestrial species
(A. terrestre) is absent from that region.
Within the A. hookerianum/bulbiferum group
there are two cytotypes of A. bulbiferum of which
the tetraploid occurs in both Australia and New
Zealand, but of which the octoploid is so far only
known from this country. Similarly, the tetraploid
species, A, hookerianum, is known from both coun-
tries, but its octoploid relative, A. richardii, is
endemic to New Zealand.
Within the A. lucidumjobtusatum aggregate, the
tetraploid cytotype of A. obtusatum is widely dis-
tributed in the Southern Hemisphere; A. lucidum is
also thought to occur on a few islands outside the
New Zealand region. However, the various octoploid
taxa, A. obtusatum subsp. northlandicum, A. sclero-
prium, and A. lyallii are all strictly endemic.
The remaining taxa are neither closely related to
each other nor to the three groups mentioned above.
A. trichomanes has two cytotypes of which the
tetraploid is cosmopolitan (though rare in New
Zealand) but the hexap'oid is almost confined to
this country. A. lamprophyllum is a tetraploid species
apparently restricted to the northern part of the
North Island but closely related to A. cuneatum
Lam. of widespread tropical distribution. A. polyodon
is also tetraploid and in New Zealand is at the
southernmost limit of its range which extends through
the Indo-Malaysian tropical region. The remaining
species, A. flabellifolium, is apomictic and therefore
not strictly comparable to the sexual taxa, but it has
hexaploid cytotypes in both New Zealand and Aus-
tralia whereas the octoploid is so far only known
from this country.
The evidence therefore strongly suggests that the
New Zealand Asplenium flora has two components:
an ancestral element of widely distributed, sexually-
reproducing, tetraploid species, and an endemic
element of higher polyploid taxa which have
originated within New Zealand from these tetraploid
'Excluding A. shuttleworthianum which occurs only
on the fringes of the New Zealand Botanical Region.
Brownsey—N.Z. species of Asplenium
parents. The derivation of the individual octoploids
will be discussed in a future publication when
evidence from naturally occurring hybrids can also
be considered. However, it seems likely that processes
of auto- and allopolyploidy may have been as impor-
tant in the evolution of the New Zealand Asplenium
flora as they are now known to have been in
Europe and North America.
ACKNOWLEDGMENTS
This work has been carried out during the tenure
of a Post-doctoral Research Fellowship supported by
Victoria University of Wellington. I am extremely
grateful to Professors H. D. Gordon and J. K. Heyes
for the opportunity to work in the Department of
Botany, and for financial assistance in carrying out
field work.
I thank the following who have collected material
for me during this investigation: Mr R. G. Bagnall,
Drs J. E. and R. E. Beever, Drs A. E. and B. D.
Bell, Mr B. D. Bell, Dr J. E. Braggins, Mr R. J.
Chinnock, Mr H. D. Christophers, Dr G. B. Cone,
Dr D. R. Given, Mr G. S. Hardy, Professor J. K.
Heyes, Miss M. McDonald, Mrs E. Miller, Dr F. B.
Sampson, Dr J. E. Sheridan, Mr C. J. Smuts-Kennedy,
Mr B. V. Sneddon, Mr G. Stephenson, Mr G. Walls,
and Mr A. E. Wright. I also thank my wife and
many friends who have assisted me with field work.
I am indebted to the curators of the following
herbaria who have sent valuable material on loan to
Wellington: AK, AKU, BM, CHR, GOET, K, P, W,
WELT. I particularly thank Miss B. H. Macmillan
(Botany Division, DSIR) for packing and dispatch-
ing the entire collection of Asplenium in CHR, Dr
J. E. Braggins, Mr A. E. Esler, and Mr A. E.
Wright for allowing me to examine their personal
herbaria, and Mr L. J. Metcalf of Christchurch
Botanic Gardens for sending me type specimens from
the Armstrong Herbarium.
Finally, special thanks go to Mr R. Lucas for
his care in maintaining a large collection of plants
in cultivation, to Mrs J. Benfield for help in the
preparation of the illustrations, to Dr W. A. Sledge
and Dr J. D. Lovis for their criticism of the manu-
script and valuable advice on matters of nomen-
clature, and to Dr R. Ross of the British Museum
for his opinion on the correct name for A. flabelli-
folium.
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