Phycologia (2005) Volume 44 (2), 139–145 Published 26 April 2005

Dione and Minerva, two new genera from New Zealand circumscribed for
basal taxa in the Bangiales (Rhodophyta)
WENDY A. NELSON1*, TRACY J. FARR1 AND JUDY E.S. BROOM2
1 National Centre for Aquatic Biodiversity and Biosecurity, National Institute of Water and Atmospheric Research,
Private Bag 14-901, Wellington, New Zealand
2
Department of Biochemistry, University of Otago, P.O. Box 56, Dunedin, New Zealand

W.A. NELSON, T.J. FARR AND J.E.S. BROOM. 2005. Dione and Minerva, two new genera from New Zealand circumscribed
for basal taxa in the Bangiales (Rhodophyta). Phycologia 44: 139–145.

Two new monospecific genera of filamentous red algae from New Zealand are described, Dione arcuata and Minerva
aenigmata (Bangiales, Rhodophyta). Although these taxa conform to the morphological concept of the genus Bangia that
is in current use, analyses of nuclear small-subunit ribosomal RNA gene (nSSU) sequences have revealed that these taxa
are basal in the order Bangiales and differ significantly from each other and all other filamentous species in the order
Bangiales for which nSSU sequence data exist (Broom et al. 2004). Dione arcuata can be distinguished in the field by its
width, growth habit and the niche it occupies in the upper subtidal zone, whereas Minerva aenigmata is found on upper
intertidal rocks in areas of sand abrasion, and is indistinguishable morphologically from other filamentous Bangiales taxa.

INTRODUCTION indicate that at least three Porphyra-like genera should be

In the red algal order Bangiales, two morphologically defined
genera are recognized: those with filaments are placed in the
genus Bangia Lyngbye (Lyngbye 1819) and bladed taxa in
Porphyra C. Agardh (Agardh 1824). A critical turning point
in the understanding of relationships amongst taxa within the
Bangiales was the publication by Oliveira et al. (1995) of
phylogenetic analyses using nuclear small-subunit ribosomal
RNA gene (nSSU) sequences. This provided evidence that
there is not a simple sister-group relationship between Bangia
and a monophyletic Porphyra. Although making no formal
nomenclatural or taxonomic decisions, Oliveira et al. (1995)
explored the consequences of their findings. They observed
that synonymizing Porphyra with Bangia could create havoc
for commercial trade in nori seaweed and would result in a
monophyletic but molecularly divergent concept of Bangia.
The concept of the order Bangiales as a monophyletic lin-
eage is consistently supported by all recent phylogenetic treat-
ments (e.g. Ragan et al. 1994; Oliveira & Bhattacharya 2000;
Müller et al. 2001). The majority of studies on the Bangiales
have been centred in the North Pacific and North Atlantic
Oceans (e.g. Lindstrom & Cole 1992a, b; Miyata & Kikuchi
1997; Brodie et al. 1998; Müller et al. 1998, 2001; Klein et
al. 2003; Lindstrom & Fredericq 2003), with relatively few
authors including southern hemisphere taxa (e.g. Oliveira &
Coll 1975; Woolcott & King 1998; Broom et al. 1999; Nelson
et al. 2001a; Farr et al. 2003; Jones et al. 2004). As collec-
tions from a wider geographical region are examined, new
taxa discovered and data from molecular sequencing and phy-
logenetic analyses emerge, the need to examine generic con-
cepts in the Bangiales has become more urgent (Nelson et al.
2001b, 2003; Niwa et al. 2003; Broom et al. 2004). Lindstrom
& Fredericq (2003) consider that the data currently available
* Corresponding author (w.nelson@niwa.co.nz).
recognized.
The first species of Bangia was initially described as Con-
ferva atropurpurea Roth (Roth 1806) and was from a fresh-
water collection from Bremen, Germany. This was followed
by the description of a marine species, C. fuscopurpurea Dill-
wyn (Dillwyn 1802–1809). Although the distinction between
freshwater and marine species of Bangia [B. atropurpurea
(Roth) C. Agardh and B. fuscopurpurea (Dillwyn) Lyngbye,
respectively] has been maintained by some authors, freshwater
and marine populations of Bangia have been regarded as con-
specific by many workers from the time of Agardh (1883),
including the recent treatments by Kumano (2002) and Brodie
& Irvine (2003), although the latter refer the reader to Müller
et al. (2003). However, data from karyotype analyses as well
as from studies employing ribulose-1,5-bisphosphate carbox-
ylase/oxygenase large subunit gene (rbcL) and nSSU provide
evidence that marine and freshwater populations of Bangia
are distinct (Gargiulo et al. 1998; Müller et al. 1998, 2003).
Many species of Bangia have been described (e.g. 45
names listed in AlgaeBase species database; Guiry & Nic
Dhonncha 2002), but relatively few (c. eight species) are in
current usage. Of these, six species are highly restricted either
geographically or ecologically, as obligate epiphytes on spe-
cific hosts. Sheath & Cole (1984), examining Bangia in North
America, concluded there was no basis for continuing to rec-
ognize the seven segregate species described from marine
specimens. They concluded that the Pacific and Atlantic Ban-
gia differed and, based on taxonomic priority, assigned the
names B. vermicularis Harvey to Pacific populations and B.
atropurpurea to Atlantic Bangia. Sheath & Cole (1984) sug-
gested that the large variability in the size of Pacific popula-
tions of Bangia may be due to the ‘great age of this basin,
allowing more time to diversify’. After examining karyology
of Bangia populations in North America, Müller et al. (2003)
concluded that although there ‘appear to be distinct entities’
139
140 Phycologia, Vol. 44 (2), 2005
Fig. 1. Collection sites for Minerva aenigmata (triangles) and Dione arcuata (circle); Wellington (2) 5 Scorching Bay and Seatoun. Sites for
D. arcuata are c. 4 km apart.
along the North American coastlines, marine populations of
Bangia should be assigned to B. fuscopurpurea until their
complex relationships can be resolved. They referred fresh-
water populations to B. atropurpurea.
The southern hemisphere has traditionally been regarded as
an area with low diversity of Bangiales. Two species of Ban-
gia were recorded for the New Zealand region by Harvey
(1855): B. ciliaris Carmichael ex Harvey [now Porphyros-
tromium ciliare (Carmichael ex Harvey) Wynne] and the en-
demic B. lanuginosa Hooker f. & Harvey [subsequently trans-
ferred to Erythrotrichia lanuginosa (Harvey) De Toni; type
specimen collected by Colenso, housed in TCD (herbarium
abbreviations follow Holmgren et al. (1990)]. The latter spe-
cies has not been collected since. Laing (1928) provided the
first record of B. fuscopurpurea in New Zealand, recording it
from three specific sites around the South Island – Pelorus
Sound (in the Marlborough Sounds), New Brighton (Christ-
church) and Dunedin – although also referring to its growth
elsewhere. Laing (1928) reported that there were a number of
forms of this species, which he regarded as a ‘widely distrib-
uted aggregate species’. Levring (1955), in a revision of the
New Zealand Bangiales, referred to a single collection of B.
fuscopurpurea from Lyttelton (Christchurch). Adams (1994)
recorded this genus as uncommon in New Zealand, placing
all marine records under the name B. atropurpurea. There
have been no collections of Bangia from freshwater habitats
in New Zealand.
Recent studies of Bangiales in New Zealand have revealed
a high level of previously unsuspected cryptic diversity in this
order, with more than 33 Porphyra species reported by Nelson
et al. (2001b) and more than 10 Bangia reported by Farr et
al. (2001). A survey of Bangia in the New Zealand region
(Broom et al. 2004) has revealed a very high level of molec-
ular diversity, which is not reflected in the conservative mor-
phology of these organisms. These discoveries have been
made in the course of a research programme that includes
extensive field collections and observations, culture studies,
molecular sequencing and phylogenetic analyses.
Two entities have been collected in New Zealand that differ
markedly from each other and from all other filamentous ban-
gialean entities we have found. Sequence analyses have re-
vealed that they are basal to all other taxa so far discovered
in the Bangiales (entities BTS and BKE in Broom et al. 2004).
In this article we describe two new genera and species for
these basal bangialean taxa.
MATERIAL AND METHODS
Samples of Bangia were collected from intertidal rocky coast-
lines throughout the New Zealand region from 1998 to 2002
(Fig. 1). Samples were sorted and subsampled for culture ini-
tiation (culture conditions follow Knight & Nelson 1999),
dried in silica gel for later DNA extraction and molecular
sequencing analyses and herbarium specimens were retained
as vouchers (deposited in WELT).
Deoxyribonucleic acid extractions and polymerase chain-
reaction amplification followed the methods provided by
Broom et al. (2004). Subsamples of collections were also pre-
served in seawater–formalin. Permanent slides of preserved
material were measured using an Axiovert 200 microscope,
an Axiocam HR digital camera and Axiovision image soft-
ware v.4.1 (all Zeiss, Jena, Germany). Measurements were
made on either fresh collections (filament length) or on pre-
served samples (all other measurements). Features of the two
taxa described here are summarized in Table 1.
RESULTS
For all samples of Minerva W.A. Nelson and Dione W.A. Nel-
son on which the taxa described in this article are based, ap-
 Nelson et al.: New genera in Bangiales 141

Table 1. Comparison of Minerva aenigmata and Dione arcuata.

Minerva aenigmata Dione arcuata

Habitat high intertidal zone on rock upper subtidal zone on rock
Distribution throughout New Zealand (northern North Island to Kaikoura region, east coast of South Island
southern South Island)
Seasonality differs with location; associated with beach erosion early summer
(February–September)
Basal region uniseriate filament; rhizoids extending down within biseriate filament in rhizoidal area, uniseriate above;
common cell wall from lowermost 9–25(–43) cells rhizoids extending down within common outer cell
wall from lowermost 10–25 (paired) cells
Sterile filament width 17–40 mm 47–80 mm
Mature male filaments
(width) 30–55(–63) mm 115–150 mm
Mature sporangial fila-
ments (width) (42–)45–68(–77) mm 120–210 mm
Monoecious or dioecious both monoecious and dioecious filaments; male and both monoecious and dioecious filaments; male and
female regions intermixed on single filament, or female regions intermixed on single filament, or
occurring separately occurring separately
Archeospores present; produced in upper part of filament present; produced in upper part of filament
Conchocelis (in culture) common infrequent
Number of collections
(sites) 15 (12) 5 (2)

proximately 500 base pairs (bp) of sequence data were ob-
tained covering the V9 region of the nSSU (Neefs et al. 1993),
which has been shown to be an effective proxy for overall
nSSU variation in Porphyra and Bangia (Broom et al. 1999).
Near-complete nSSU sequences (c. 1750 bp) were obtained
for five collections of Minerva and two collections of Dione.
Within each taxon, all nSSU V9 sequences obtained were
identical, and the near-complete nSSU sequences within each
taxon were also identical to one another.
Taxonomic treatment
Minerva aenigmata W.A. Nelson, gen. et sp. nov.
Figs 2, 4–10
Filamenta simplicia flaccida ad 1.5–3.0(–10) cm alta, substratis sax-
osis in zona supera interaestuali, per rhizoidea a cellulis inferioribus
filamenti intra parietem communem exteriorem cellulosum exten-
dentia affixa. Filamenta uniseriata in parte infera, supra multiseriata,
13–23 mm lata in area rhizoideorum, 17–40 mm in parte sterili, in
parte fertili supera ampliantia ad 30–55(–63) mm (mascula), (42–)
45–68(–77) mm (feminea). Alga monoecia, spermatangiis et zygo-
tosporangiis praesumptis in filamentis aliquis intermixtis et ut vi-
detur singulatim in filamentis aliis. Spermatangia pallentia cellulis
omnibus; zygotosporangia praesumpta fusce pigmentifera. Archeos-
porae in parte superiore filamenti efferentes. Post germinationem
sporae filamenti ‘conchocelis’ procreant vel directe in filamenta
crescunt. Data de sequentiis 5 nSSU GenBank AY465355 (5 BTS,
Broom et al. 2004).
The filaments are simple, flaccid, 1.5–3(–10) cm long (Fig. 2), af-
fixed to rocky substrate in the upper intertidal zone (Fig. 3) by
rhizoids extending from the lower cells of the filament within a
common outer cell wall (Fig. 4). Filaments are uniseriate in their
lower part (Fig. 5), multiseriate above; they are 13–23 mm wide in
the rhizoid area, 17–40 mm wide in the sterile region, widening to
30–55(–63) mm (male) (Fig. 6) or (42–)45–68(–77) mm (female)
(Fig. 7) in the fertile upper filament. The alga is monoecious; sper-
matangia and presumed zygotosporangia are intermixed in some
filaments (Fig. 8) but apparently singly in others. The spermatangia
are pale, whereas presumed zygotosporangia are darkly pigmented.
Archeospores are produced in the upper part of the filament (Fig.

Fig. 2. Holotype of Minerva aenigmata (WELT A25775). Scale bar 5 1 cm.

Fig. 3. Puheke, North Island; type locality of Minerva aenigmata.
142 Phycologia, Vol. 44 (2), 2005

Figs 4–10. Minerva aenigmata. Scale bars 5 100 mm (Figs 4–9 all to the same scale).
Fig. 4. Basal region of filament showing rhizoids extending from lower cells of the filament within common wall (WELT A27031).
Fig. 5. Sterile filaments, initially with square cells (right) becoming shorter and broader (left filament) (WELT A27032).
Fig. 6. Male region of filament with spermatia being released (upper part of filament) (WELT A27033).
Fig. 7. Female region of filament, showing presumed zygotosporangia (WELT A23128).
Fig. 8. Filament with both male (small arrow) and female (large arrow) regions intermixed (WELT A23128).
Fig. 9. Archeospores ready to be released from upper part of filament (WELT A23129).
Fig. 10. Conchocelis filaments germinated from spores from field collected samples (WELT A27032).

9). Filament spores germinate to produce conchocelis (Fig. 10) or
develop directly into filaments.
HOLOTYPE (DESIGNATED HERE): Puheke, Great Exhibition Bay, North
Island, New Zealand; Nelson & Farr, 8 May 2001, WELT A25775
(Fig. 2).
SEQUENCE DATA: The nSSU GenBank AY465355 (5 BTS, Broom
et al. 2004)
ETYMOLOGY: Minerva – goddess of wisdom, hearth and weaving, in
reference to the basal position this taxon occupies (providing wis-
dom, knowledge) and the woven mat-like appearance on intertidal
rocks; aenigmata – mystery, obscure, riddle, in reference to the
puzzle presented by these morphologically simple filaments that dif-
fer so markedly in nSSU sequence from other taxa.
HABITAT:Growing on rock in the upper intertidal zone on exposed
to moderately sheltered open coasts.
DISTRIBUTION: Widespread around the New Zealand mainland from
the northern North Island to the southern South Island (Fig. 1).
Collections of this species have been made from throughout the
year at different locations.
REPRESENTATIVE SPECIMENS EXAMINED: Maketu, Bay of Plenty, Farr
& Williams, 5 May 2000, WELT A23131; Seatoun, Wellington,
Farr & Nelson, 12 February 2001, WELT A23128; Scorching Bay,
Wellington, Farr, 29 October 2001, WELT A23132; Manukau
Bluff, Canterbury, Neill & Jones, 20 September 2001, WELT
A23129; Cobden, Westland, Farr & Nelson, 11 March 2000, WELT
A26694, and 22 August 2000, WELT A23130; Warrington, Otago,
Broom & Jones, 5 September 2001, WELT A23127.
OBSERVATIONS: Minerva aenigmata is widespread in the New Zea-
land mainland region, occurring from northern North Island sites
(Puheke) through to the southern South Island near Dunedin (War-
rington) (Fig. 1), and was identified as ‘BTS’ in Broom et al.
(2004). This species occupies a habitat reported as typical for spe-
cies of marine Bangia, namely growing in the upper intertidal zone
on rock, often in areas which are scoured by sand or experience
frequent sand burial. This species has not been collected from
wooden or concrete surfaces. The seasonality of this species is hard
to determine from the field records to date. Collections have been
made from different sites in all months of the year. However, at
specific sites this species appears to have a strongly seasonal ap-
pearance that seems to be correlated with the seasonality of sand
deposition and erosion. In the Wellington region (central New Zea-
land) this species shows a very strong summer annual distribution,
appearing for c. 6 weeks each year and then completely disappear-
ing for the remainder of the year. The period when the species is
present coincides with local erosion of sand and uncovering of small
boulders and reef areas. After rocks become uncovered through
scouring this species appears very rapidly, forming thick felts on
bare rock surfaces within the upper intertidal zone. On the west and
east coasts of the South Island, records of this species are from
winter to spring, whereas in the northern North Island the collec-
tions have been made in late autumn.
Dione arcuata W.A. Nelson, gen. et sp. nov.
Figs 11, 13–17
Filamenta lata grossa curvata simplicia ad 1.5 cm longa, substratis
saxosis in zona umbrosa supera subaestuali per rhizoidea ex cellulis
binis inferioribus filamenti intra parietem communem exteriorem
cellulosum extendentia affixa. Filamenta uniseriata 47–60 mm lata
proxime super aream rhizoideorum, ampliantia ad 80 mm lata et
series multas facientia. Areae mares femineae filamentorum fertil-
ium aut intermixtae aut singulatim dispositae. Regiones spermatan-
giales 115–150 mm latae; zygotosporangia praesumpta 120–210 mm
lata. Archeosporae in partibus superioribus filamenti efferentes.
Sporae e filamentis pro parte maxima in filamenta nova directe cres-
cunt, ‘conchocelis’ formis raris. Data de sequentiis 5 nSSU
GenBank AY465354 (5 BKE, Broom et al. 2004).
Filaments are broad, coarse, curved and unbranched, up to 1.5 cm
long (Fig. 11); they are attached to rocky substrate in shaded parts
of the upper subtidal zone (Fig. 12) by rhizoids extending from
paired lower cells of the filament within a common outer cell wall
(Fig. 13). The filaments are uniseriate and 47–60 mm wide imme-
diately above the rhizoidal area, but widening and becoming mul-
tiseriate and up to 80 mm broad distally (Fig. 14). Fertile filaments
have male and female regions either mixed or singly. Spermatangial
regions are 115–150 mm wide (Fig. 15), whereas presumed zygo-
tosporangial regions are 120–210 mm wide (Fig. 16). Archeospores
are produced in the upper part of the filament. Spores of most fil-
 Nelson et al.: New genera in Bangiales 143

Fig. 11. Holotype of Dione arcuata (WELT A23126). Scale bar 5 1 cm.
Fig. 12. Rakautara, mouth of stream with rocks at extreme low water. One of two locations where Dione arcuata has been collected.

aments develop directly into new filaments; the conchocelis phase DISTRIBUTION: Highly restricted geographic distribution; to date
is rare (Fig. 17). found from only two locations 4.2 km apart on the east coast of the
South Island north of the Kaikoura Peninsula. This species has only
HOLOTYPE (DESIGNATED HERE):
Ohau Stream, Kaikoura, South Island,
been found in summer (November and December).
New Zealand, 5 November 2002, Neill & Farr, WELT A23126
(Fig. 11). REPRESENTATIVE SPECIMENS EXAMINED: Rakautara Stream, Kaikoura,
17 December 2000, Nelson, Farr & Williams, WELT A26702
SEQUENCE DATA: The nSSU GenBank AY465354 (5 BKE, Broom
(GenBank AY184345).
et al. 2004).
OBSERVATIONS: Dione arcuata is unusual amongst New Zealand
ETYMOLOGY: Dione – another name for Venus, the goddess who
Bangiales in that it has only been found at extreme low water level,
arose from the sea attended by cupids and dolphins – ‘The parent
and thus is predominantly subtidal, only being exposed at extreme
ocean work’d with heaving throes, and dripping wet the fair Dione
low water, spring tides. This species was identified as ‘BKE’ in
rose’ Anon. – in reference to the shore position occupied by Dione
Broom et al. (2004). To date only two populations of this species
on the edge of the subtidal zone; arcuata – curved, shaped like a
have been found, situated c. 4 km apart on the east coast of the
bow, in reference to the curved shape of the filaments.
South Island, north of the Kaikoura Peninsula (Fig. 1). At both sites
HABITAT:On rock in the low intertidal to upper subtidal on exposed a stream flows into the sea across cobble or boulder shores. The
open coast, most abundant on shaded faces. populations of D. arcuata have been found on north- and east-facing

Figs 13–17. Dione arcuata. Scale bars 5 100 mm (Figs 13–16 all to same scale).
Fig. 13. Basal region of filament showing rhizoids extending within a common cell wall from paired cells of the lower filament. Filament
becomes uniseriate again immediately above rhizoidal region (WELT A23126).
Fig. 14. Sterile filament showing broad uniseriate (right) and multiseriate filaments (left) (WELT A27030).
Fig. 15. Male region of filament showing release (upper) (WELT A23126).
Fig. 16. Female region of filament, presumed zygotosporangia (WELT A23126).
Fig. 17. Conchocelis filament germinated from spores from field collected material (WELT A26702).
144 Phycologia, Vol. 44 (2), 2005
boulders. The only records we have to date are from late spring and
early summer (November and December). Attempts to locate this
species at other times and in other locations have not been suc-
cessful. The curved growth form and large size of these filaments
are distinguishing characters that can be seen in the field.
DISCUSSION
Our research on southern hemisphere members of the Ban-
giales has revealed unanticipated diversity in the order, and
has shown that the simple morphology of these organisms
obscures significant levels of genetic diversity (Farr et al.
2003; Broom et al. 2004; Jones et al. 2004). The species D.
arcuata can be readily distinguished morphologically from all
other filamentous members of the Bangiales and it occupies a
unique habitat amongst the New Zealand members of the or-
der. The morphology of M. aenigmata, however, does not as-
sist identification or enable it to be distinguished from other
filamentous Bangiales found in New Zealand. It is common,
widespread and is found on upper intertidal rock surfaces that
frequently experience sand scour, the predominant habitat of
filamentous Bangiales in the New Zealand region.
Nelson et al. (2003) advocated the use of molecular se-
quence data of vouchered or type material in the description
of new taxa in the orders Bangiales and Erythropeltidales,
given the simplicity in morphology in these orders and their
well recognized combination of phenotypic plasticity and con-
vergent morphological characters. Although current generic
concepts of Porphyra and Bangia are untenable, as yet there
is no clear morphological or anatomical character set that will
allow the separation of genera consistent with phylogenetic
relationships: there is no immediate prospect of resolution to
questions about the circumscription of genera in this order
except with the use of molecular sequence data. The value of
sequence data in the identification and delineation of taxa in
the Bangiales is clearly established (Nelson et al. 2001a; Klein
et al. 2003; Lindstrom & Fredericq 2003).
Sequences of the nSSU gene from these two taxa differ
significantly from one another and from those of all other
Porphyra and Bangia sequences obtained to date. Alignment
of nSSU exon sequences from these two taxa with those of
other bangialean taxa poses some difficulties because of the
amount of sequence variation in the variable regions of the
gene. This confounds attempts to determine a ‘best’ align-
ment. Nevertheless it is apparent that more than 90 nucleotide
substitutions separate the nSSU exon sequences of these taxa
from one another and from all other bangialean nSSU exon
sequences in GenBank to date (Broom et al. 2004). The large
sequence divergences seen for these two taxa are in sharp
contrast with sequence differences observed for other pairs of
bangialean taxa. For example, nSSU exon sequences from the
Japanese taxa P. yezoensis Ueda (GenBank accession
AB013177) and P. tenera Kjellman (AB013175) differ from
one another by two nucleotide substitutions and one 2 bp in-
del; those from the saltmarsh taxon P. rediviva J.W. Stiller &
J.R. Waaland (AF175544) and the marine taxon P. purpurea
(Roth) C. Agardh (L26201) differ by only three nucleotide
substitutions; whereas the nSSU exon sequence of the foliose
P. yezoensis Ueda (AB013177) differs from that of the fila-
mentous Bangia sp. Newfoundland, Canada (AF169338) by
21 nucleotide substitutions and three indels of 2, 1 and 4 bp
in length.
The circumscription of two new genera in this order is the
beginning of a revision of the taxonomy of this order. Further
resolution of the relationships and taxonomy of the Bangiales
will require the inclusion of taxa from throughout the distri-
butional range of these organisms and, as observed by Lind-
strom & Fredericq (2003), involve the efforts of phycologists
working in different parts of the world. Bangia and Porphyra
as now defined have a worldwide distribution from the poles
to the tropics, and a solely regional focus in phylogenetic anal-
yses will result in flawed outcomes. Freshwater Bangia from
Europe and from North America form a strongly supported
clade in all molecular analyses carried out to date (e.g. Müller
et al. 2003; Broom et al. 2004): it is this entity that has pri-
ority in the genus Bangia. Broom et al. (2004) have illustrated
that other species currently classified as Bangia from marine
habitats fall within a number of distinct clades, some of which
also include species of Porphyra, and the phylogenetic rela-
tionships of a further number of taxa of ‘Bangia’ remain un-
resolved.
The scale of diversity found in New Zealand is exceptional
and places this region as a centre of diversity for the order
Bangiales. It is not clear whether similar diversity will be
found when equivalent molecular sequencing methodologies
are applied to floras in other areas. A much closer examination
of the Bangiales is required, however, before an adequate un-
derstanding of the diversity and phylogeography of these or-
ganisms will be achieved.
ACKNOWLEDGEMENTS
We thank our colleagues Kate Neill and Wyn Jones for assis-
tance with field collections. This research was funded by the
New Zealand Foundation for Research, Science and Technol-
ogy MNZX0202.
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Received 20 March 2004; accepted 10 October 2004
Communicating editor: R. Sheath (Guest Associate Editor)