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An overview of Nitella in Australia (Characeae, Charophyta)

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www.publish.csiro.au/journals/asb Australian Systematic Botany, 22, 193–218

An overview of Nitella (Characeae, Charophyceae)

in Australia

Michelle T. Casanova

Royal Botanic Gardens Melbourne, Birdwood Avenue, South Yarra, Vic. 3141, Australia and 273 Casanova Road,
Westmere, Vic. 3351, Australia. Email: amcnova@netconnect.com.au

Abstract. The genus Nitella Ag. in algal family Characeae is characterised by furcate (forked) branchlets, compressed
oospores (i.e. oval in cross-section), terminal antheridia and a 10-celled coronula on the oogonium. Species of Nitella are
submerged plants that grow in a variety of wetland and riverine habitats. Approximately 89 taxa of Nitella (species,
subspecies, varieties and forms) have been described on the basis of Australian collections, and published estimates of the
number of species range from 18 to >35. The lower value is based on the assumption that infra-specific variation is great,
species have a wide distribution, monoecy and dioecy are not indicative of speciation and the number of furcations and the
ratio of branchlet segment lengths vary for a species owing to the environment in which they grow. The higher value is based
on evidence that morphological characters are relatively constant for a species, that oospore variation is a good indication of
speciation and that monoecious and dioecious entities are not inter-fertile. An overview of Australian members of the genus is
presented here as a framework for further taxonomic work. Representatives of all three subgenera of Nitella occur in Australia,
with subgenus Nitella poorly represented, and subgenera Tieffallenia and Hyella equally speciose. The subgenera are defined
here in relation to the Australian taxa they contain. In the present treatment, section Migularia is transferred from subgenus
Tieffallenia to subgenus Hyella, and several species are transferred to subgenus Tieffallenia. Within subgenus Tieffallenia,
variation in vegetative and oospore morphology is useful for distinguishing among sections and species. However, although
members of subgenus Hyella display a similar range of variation in vegetative morphology, most of the species have similar,
reticulate, oospore ornamentation. Australia is home to a large number of endemic species of Nitella, many of which are
dioecious. The total number of species and the degree of endemism have been underestimated in earlier studies, and it is likely
that more than 50 species of Nitella will be recognised on the basis of Australian specimens. A key to the subgenera, and keys
to sections in the subgenera are provided.

Introduction of New England (Macdonald and Hotchkiss 1956; Chambers and

Members of family Characeae are macroalgae that occur in fresh,
brackish and saline non-marine waters worldwide. Australia has
five of the six genera that are currently recognised, including
Chara L., Nitella Agardh, Lamprothamnium J.Groves, Tolypella
(A.Braun) A.Braun and Lychnothamnus (Rupr.) Leonhardi.
Nitellopsis Hy appears to be restricted to the northern
hemisphere. The genus Nitella is characterised by furcate
(forked) branchlets (Fig. 1), compressed oospores (i.e. oval in
cross-section), terminal antheridia and a 10-celled coronula on the
oogonium (Wood 1965). Specimens of Nitella have been
collected from Australia since Robert Brown found Nitella
congesta (R.Br.) A.Braun in 1803 (Brown 1810). Early
naturalists sent specimens to Braun and Nordstedt in Europe
for determination. During the middle and late 1800s, 22 species
had been described for Australia (Nordstedt 1889) and by the
early 20th century, 33 species were recognised (Nordstedt 1918).
In the 1930s, Groves and Allen (1935) investigated the
Queensland members of Characeae, recognising 20 Nitella
species recorded previously and two additional species
(N. phauloteles and N. comptonii; see Table 1 for authorities).
In the 1950s, Beth Williams (nee Macdonald) worked on
charophyte taxonomy at Sydney University and the University
Williams 1959; Williams 1959). She clarified the circumscription
of N. cristata and N. diffusa and described N. reticulata and
N. microteles. Unfortunately, many specimens from Australian
herbaria gathered for examination by Williams were damaged or
destroyed in the Armidale fire in early 1958 (Wood and Williams
1967). In 1960, Richard Wood came to Australia on a Fulbright
Fellowship, collected widely, and used this collection as the basis
of his revision of Australian species, which resulted in recognition
of 15 species of Nitella for Australia; the remainder of described
species were relegated to subspecies, varieties or forms, or
amalgamated with northern hemisphere species (Wood 1972).
After Wood’s revision (Wood 1972), there was a period of
time when little charophyte research was undertaken in Australia.
However, towards the end of the 20th century, there was a flurry
of taxonomic activity. Van Raam (1995) revised the Characeae
of Tasmania, resulting in 18 Nitella species. He retained eight of
the Nitella species recognised by Wood (1972), reinstated six
previously described species, raised two subtaxa to species, made
three new varietal combinations and described two new species
(N. haagenii and N. monopodiata). N. partita and N. sonderi have
been reinstated after examination of type material and new
collections (Casanova et al. 2003; Casanova 2007a) and

 CSIRO 10 June 2009 10.1071/SB08039 1030-1887/09/030193

194 Australian Systematic Botany M. T. Casanova

(a) (b)

(c) (d )

Fig. 1. Branchlet whorls of Nitella (as viewed from above). (a) Heteroclemous whorl of a specimen of r022
Nitella congesta (R.Br.) A.Braun from Mount Monster, South Australia, with ~40 branchlets in three whorls
at each node (M.T. Casanova, 27.x.2007, MEL). (b) Homeoclemous whorl of a specimen p907 N. sonderi
A.Braun from Tang Tang Swamp, with 6 even, 1–2  furcate branchlets in the whorl (K. Ough CS1a,
28.iv.2006, MEL). (c) Homeoclemous whorl of a specimen of r089 Nitella sp. from Howden, Tasmania, with 8,
evenly 3  furcate, macrodactylous branchlets in a whorl (H. and A. Wapstra HAW/03a, 20.vi.2008, MEL).
(d) Homeoclemous whorl of a specimen of r090 Nitella sp. aff. furcata from Howden, Tasmania, with
8 brachydactylous branchlets in a whorl (H. and A. Wapstra HAW/03b, 20.vi.2008, MEL).

additional species have been described (N. woodii, Hotchkiss and
Imahori 1988a; N. ignescens, N. ungula, García 1996;
N. paludigena Casanova and Karol 2008). Hotchkiss and
Imahori (1988a) reported on the occurrence of N. imahorii,
N. imperialis and N. horikawae in New South Wales in 1968,
and Sakayama et al. (2005) reported on a collection of
N. morongii in Queensland. García and Chivas (2006) listed
37 species of Nitella for Australia, including records of 15 species
not seen by the authors, records of two New Zealand species and
two additional, undescribed taxa. Casanova et al. (2007) revised
the N. hookeri complex, raising N. arthroglochin to species.
Problems concerning the taxonomy of species in Chara and
Nitella have been summarised (Casanova 2005a, 2007a) and
García and Chivas (2006) gave a description of the distribution
and ecology of 62 species of Australian charophytes on the basis
of literature records for ~18 species and on their own collections.
These works are contributing to the recognition of the importance
of charophytes in the ecology of Australian wetlands, although
the taxonomy remains difficult because of Wood’s (1972)
erroneous revision, difficult and out-of-date keys, and a lack of
clarification about what actually defines a species (Casanova
2005a).
There have been few explicit descriptions of what constitutes a
species in the Characeae. The earliest species concept for
Australian Characeae was developed by Alexander Braun
(Braun 1843, 1849). He was the first person to examine in
detail several specimens from the region. Braun introduced
several subcategories in his treatments, although it is not
always clear whether they represent varieties, forms or merely
descriptive names (Wood 1950). From the descriptions given in
Braun and Nordstedt (1882) it appears that Braun’s subspecies
were assigned to entities that had the same overall appearance,
although they differed in the shape of the end-cell or construction
of dactyls, and that varieties differed in size (var. minor, var.
major), or compactness (var. fastigiata, var. penicillata), which
are characters that can vary as a consequence of growing
conditions (e.g. Henricson et al. 2006). Nordstedt finished
Braun’s life work (Braun and Nordstedt 1882) by following
Braun’s notes after Braun died in 1877 (Allen 1953).
Nordstedt appears to be the first to investigate oospore
An overview of Nitella in Australia
Table 1. List of species of Nitella (Ag.) A.Braun that have been published based on Australian specimens
In many cases Wood (1972) wrote that forms ‘might’ be designated, and gave only brief, comparative descriptions in English, with a
specimen cited as an example of each. The specimens cited are labelled ‘type’. In several cases, the names would be illegitimate if
the forms were raised to variety or species because the combination already exists
Classification by Wood (1972)
Subgenus Nitella
Section Rajia
Subgenus Nitella
Section Palia
Subgenus Tieffallenia
Section Tieffallenia
Subgenus Tieffallenia
Section Muelleria
Subgenus Tieffallenia
Section Vogania
Subgenus Tieffallenia
Section Gioallenia
Australian Systematic Botany 195
Taxon
N. acuminata A.Braun ex Wallm.
N. acuminata A.Braun ex Wallm. var. acuminata. f. belangeri (A.Braun) R.D.Wood
N. stuartii A.Braun
N. furcata (Roxb. ex Bruz.) Ag. em. R.D.Wood
N. furcata var. zollingeri (A.Braun) Zaneveld
N. oligospira T.F.Allen
N. australiensis (Nordstedt) van Raam
N. microcarpa A.Braun
N. polyglochin A.Braun
N. orientalis T.F.Allen
N. partita Nordstedt
N. tumida Nordstedt
N. pseudoflabellata A.Braun in A.Braun and Nordstedt
N. pseudoflabellata var. imperialis f. gelatinosa R.D.Wood
N. pseudoflabellata var. imperialis f. gracilis R.D.Wood
N. pseudoflabellata var. imperialis f. delicatula R.D.Wood
N. pseudoflabellata var. imperialis f. diffusa R.D.Wood
N. pseudoflabellata var. imperialis f. bancroftii R.D.Wood
N. pseudoflabellata var. imperialis f. australis R.D.Wood
N. pseudoflabellata var. imperialis f. flabellata R.D.Wood
N. pseudoflabellata f. australiana Nordstedt
N. imperialis (T.F.Allen) Sakayama
N. conformis Nordstedt
N. leptosoma Nordstedt
N. comptonii J.Groves
N. mucosa (Nordst.) J.Groves
N. confusa (R.D.Wood) J.C.van Raam
N. batrachosperma Thuillier acc. Reichenbach
N. haagenii J.C.van Raam
N. paludigena M.T.Casanova & K.G.Karol
N. monopodiata J.C.van Raam
N. sonderi A.Braun
N. phauloteles J.Groves
N. subtilissima A.Braun
N. subtilissima var. subtilissima f. diffusa R.D.Wood
N. subtilissima var. subtilissima f. dendroidea R.D.Wood
N. subtilissima var. micophylla f. tasmanica R.D.Wood
N. microphylla A.Braun
N. gloeostachys A.Braun
N. gloeostachys var. major A.Braun
N. ignescens A.García
N. verticillata (Fil. et G.O.Allen ex Fil) R.D.Wood
N. robertsonii A.Braun & Nordst.
N. aemula A.Braun
N. penicillata A.Braun ( = N. gunnii var. penicillata nom. illeg)
N. penicillata var. fastigiata A.Braun ( = N. gunnii var. fastigiata nom. illeg.)
196 Australian Systematic Botany M. T. Casanova

Table 1. (continued )
Classification by Wood (1972) Taxon
Subgenus Decandollea 
 N. lhotzkyi (A.Braun) A.Braun ( = N. conglobata var. lhotzkyi nom. illeg.)
Section Decandollea
N. lhotzkyi f. subtilissimoides R.D.Wood
N. lhotzkyi f. major R.D.Wood
N. biformis A.Braun ( = N. conglobata var. biformis nom. illeg.)
N. heterophylla (A.Braun) A.Braun
N. congesta (R.Brown) A.Braun
N. hyalina (DC.) Ag.
N. hyalina var. brachyactis A.Braun
N. ungula A.García
Subgenus Hyella 
 N. myriotricha A.Braun ex Kütz.
Section Hyella
N. leptostachys A.Braun em R.D.Wood
N. leonhardii R.D.Wood ( = N. interrupta Braun. (ined.) non-N. interrupta (Rupr.) Kütz.)
N. polycephala A.Braun
N. polygyra (A.Braun) ex Kütz.
N. tasmanica Muell. ex A.Braun
N. tasmanica subspecies gelatinfera var. gelatinifera f. major A.Braun & Nordst.
N. tasmanica ssp. gelatinfera var. gelatinifera f. minor A.Braun & Nordst.
N. tasmanica ssp. gelatinfera var. gelatinifera f. elongata R.D.Wood
N. tasmanica ssp. gelatinfera var. gelatinifera f. compacta R.D.Wood
N. tasmanica ssp. gelatinfera var. gelatinifera f. zonata R.D.Wood
N. tasmanica ssp. gelatinfera var. gelatinifera f. tenuis R.D.Wood
N. gelatinifera R.D.Wood ( = N. gelatinosa (A.Braun) A.Braun non-C. gelatinosa Roth)
N. gelatinifera var. opaca A.Braun
N. gelatinifera var. microcephala (A.Braun) J.C.van Raam
N. gelatinifera var. microcephala f. simplex R.D.Wood
N. gelatinifera var. podostachya (A.Braun) J.C.van Raam
N. gelatinifera var. cladostachya (A.Braun) J.C.van Raam
N. gelatinifera var. cryptostachya (A.Braun) R.D.Wood
N. gelatinifera var. genuina A.Braun & Nordst.
N. gelatinifera var. afoliata R.D.Wood
N. gelatinifera var. beaugleholii R.D.Wood
Subgenus Hyella 
 N. cristata A.Braun
Section Migularia
N. cristata var. cristata f. robusta R.D.Wood
N. cristata var. ambigua A.Braun
N. reticulata M.B.Williams
N. diffusa A.Braun
N. cristata var. diffusa f. laxa R.D.Wood
N. cristata var. diffusa f. obscura R.D.Wood
N. remota A.Braun & Nordst.
N. microteles M.B.Williams
N. woodii Hotchkiss & Imahori
N. arthroglochin (A.Braun) M.T.Casanova
N. tricellularis Nordstedt
N. hookeri A.Braun

features, and relied on them to distinguish different species
(Nordstedt 1888, 1889). Groves (1922, 1927) also examined
and described oospore features, and he delineated several new
species of Nitella; however, he died before he finished working on
the Queensland Characeae, and left his work to be published
posthumously by G. O. Allen (Groves and Allen 1935), so his
species concept for Australian Characeae (especially for Nitella)
remained unjustified. Zaneveld (1940) referred to both subspecies
and varieties for his treatment of the Malaysian Characeae and
Allen (1953, 1954) maintained the species concept developed by
his mentor, Groves, and was ‘less concerned’ with varieties than
Wood (1948) had been. In all these works, authors have grappled
with the fact that within the Characeae there is a large number of
similar entities that vary from each other in microscopic details. In
many cases, the authors saw few representatives of each, so the
conclusion that there were broad species that exhibited a large
degree of morphological variation was a reasonable one.
Chambers and Williams (1959) changed the approach to
charophyte taxonomy by introducing an experimental basis to
the concept of species. Williams (1959) did not describe new
varieties, and distinguished among species on the basis of
breeding compatibility, oospore features and branchlet
segment-length ratios. Her species were very narrowly defined
but the approach was well supported by the biological species
An overview of Nitella in Australia
concept (Proctor 1975). However, Wood (1972) revived Braun’s
broad species concept for Australian charophytes, and reduced
Williams’ species to varieties. He carried Braun’s broad species
concept even further and recognised not only subspecies and
varieties, but forms as well (Wood 1965, 1972).
Wood’s most lasting contribution to charophyte taxonomy
might be his development of a phylogenetic framework of
relationships among taxa on the basis of more than one
morphological character (Wood 1964). He divided the genus
Nitella into three subgenera (Nitella, Hyella and Tieffallenia),
and subgenera into sections. He gave a clear definition of
taxonomic levels in Characeae for the first time (Wood 1965),
on the basis of the untested premises of Braun, Nordstedt
and Zaneveld (Proctor 1980). Species were defined as having
‘intergrading sets of recombining characters’, the level
‘subspecies’ was used where additional infra-specific categories
were needed, varieties were thought to differ in having ‘one or a
restricted set of intergrading recombining characters and by
having some especially distinctive feature or features’ and forms
were thought to ‘each differ from other combinations of
characters, [forming] very constant populations, often apparently
a single genotype . . . largely clonal in nature’ (Wood 1965: p. 24).
Wood suggested that his ‘forms’ wereequivalent to ‘microspecies’,
i.e. narrowly defined species, and gave a ‘microspecies index’ in
which each of his ‘forms’ were listed as species with the relevant
authorities. In this way, many of the entities that did not conform to
Wood’s broad species concept were actually described by him as
species. From this, we can deduce that Wood thought species in
Characeae should bebroadly defined; however, ifyou weregoing to
define them more narrowly then you could use his ‘forms’ as
species. His reason for creating this unwieldy taxonomy
(e.g. Nitella pseudoflabellata ssp. imperialis var. imperialis
f. sonderi instead of N. sonderi; Casanova 2007a) was that
recognition of all the microspecies would create a ‘formidable’
number of species. He anticipated that forms would not be retained
or needed in the future (Wood 1964); however, history has not
shown this to be the case.
Proctor’s work on Chara improved the definition of species
in Characeae (summarised in Proctor 1975), and he found that
many of the taxa in the Characeae were ‘species complexes’ of
interrelated entities, some morphologically identical, some
capable of crossing to produce fertile offspring. Clues to
breeding incompatibility in Chara were different arrangements
of gametangia, different morphology and geographic isolation.
Recent advances in nucleotide sequencing, scanning electron
microscopy of oospores and further study of Nitella support
Proctor’s approach and indicate that Wood’s microspecies
concept actually delineates biological species more accurately
(Sakayama et al. 2002, 2004, 2005), and is more useful
in biodiversity studies and surveys (e.g. Casanova 2004)
than Wood’s broad species concept. However, revision of
Australian members of genus Nitella is not just the simple task
of raising forms to species level, because in some cases
(e.g. Casanova 2007a) several microspecies or forms have
been delineated for what can be shown to be a single
taxonomic entity (on the basis of morphology). Also the
amalgamation of dioecious and monoecious species without
preserving the dioecious taxon as a separate form or variety
means the entire group (as defined by Wood 1965) needs to be
Australian Systematic Botany 197
examined. This can be done only if the morphological
characters that indicate species-level differences in Nitella can
be defined. Additionally, when thorough surveys have been
undertaken for charophytes in Australia, new taxa have been
found in most catchments surveyed (Casanova 1993; García
1999; Casanova 2004, 2005b; Schult et al. 2006; Casanova
and Karol 2008).
The oospores of Nitella species provide a useful tool in
charophyte taxonomy. Nordstedt (1888, 1889) relied on them
to distinguish among species, and they were used by numerous
subsequent workers for species-level delineation (Allen 1892,
1894, 1896; Groves and Bullock-Webster 1920; Groves and
Allen 1935; Zaneveld 1940; Williams 1959; Haas 1994). The
differences among oospores of type specimens (John and
Moore 1987; Casanova and Karol 2008) and the correlation of
oospore characters and particular nucleotide sequences
(Sakayama et al. 2002, 2004, 2005, 2006) confirm that
oospores possess characters that are useful for distinguishing
among species. Surveys of oospore variation in species
throughout their geographic range (de Winton et al. 2007;
Casanova 2007a) have indicated that oospores are consistent
for species delineated on morphological grounds. It remains to be
seen whether entities with the same oospore features are the
same biological species (i.e. can interbreed to produce fertile
offspring). Until breeding studies are done, however, the
available evidence supports the use of oospore morphology as
an indicator of speciation in Nitella.
The aim of the present study was to provide an overview of the
genus Nitella in Australia, to explicitly state the morphological
criteria on which species can be determined and to build a
framework for further taxonomic study in this genus.
Materials and methods
More than 750 fresh, pressed and spirit-preserved specimens of
Nitella in Australian and overseas herbaria and several private
collections were examined for the present study, including the
type material of 38 taxa. Each specimen was allocated a number
(p### or r###) so that all data obtained and stored for that
specimen (i.e. measurements, photographs, chromosome
counts, scanning electron micrographs, nucleotide sequences)
could be related to the specimen. Initial examination involved
measurement (size, number) directly on the specimen
(e.g. number of branchlets, number of furcations, presence/
absence of heads and mucus) and under a microscope
(e.g. axis and branchlet diameter, number and morphology of
dactyls, arrangement of gametangia). Oospore features were
examined and measured by light microscopy and scanning
electron microscopy. The specimens referred to in the present
study are listed in Table 2.
Fresh specimens were obtained in field surveys (e.g. Casanova
2004) and from cultures of seed-bank material in a greenhouse
(following the methods of Casanova 2004). In the greenhouse,
plastic containers (185 mm  125 mm  50 mm) containing
~300 g of wetland soil were inundated to a depth of 10–14 cm
in large tanks and the charophytes that germinated from them
were examined. When oospores were present they were removed
from the specimens for examination with a scanning electron
microscope (SEM), after which plants were gathered and pressed,
198 Australian Systematic Botany M. T. Casanova

Table 2. List of species of Nitella (Ag.) A.Braun illustrated in the study

Species are listed alphabetically. Herbarium codes follow ‘Index Herbariorum’. ‘Reference number’ refers to the number allocated to that specimen in the author’s
notes and in the present paper

Species Locality and date Collector and collector no. Herbarium Reference no. Figures
Nitella. sp. Howden, Tas., 20.vi.2008 H. and A. Wapstra HAW/03a MEL r089 Fig. 1c
N. acuminata A.Braun ex Moochalabra Dam, WA, A.A. Mitchell 3703B PERTH p149 Figs 3, 4
Wallm. 27.vii.1994
N. arthroglochin (A.Braun) Corralinga Ck, Fleurieu E.N.S. Jackson 75 AD p604 Figs 81, 82
M.T.Casanova Peninsula, SA, 23.iii.1959
N. australiensis (Nordst.) Lake Dulverton Tas. M.T. Casanova, MEL p807 Figs 7, 8
J.C.van Raam 22.xi.2005,
N. comptonii J.Groves Enoggera Reservoir, Brisbane, R.D. Wood 60-11-21-23 NY p912 Figs 29, 30
Qld, 21.xi.1960
N. confusa (R.D.Wood) Cynthia Bay, Lake St Clair, R.D. Wood 61-1-2-12A AD r036 Figs 17, 18
J.C.van Raam Tas., 24.xii.1961
N. congesta (R.Br.) A.Braun Gahnia Lagoon 5, Brinkworth M.A. Brock MEL p255 Figs 43, 44
wetlands, SA, 24.ii.1997
N. congesta (R.Br.) A.Braun Mt Monster, SA, 27.x.2007 M.T. Casanova MEL r022 Fig. 1a
N. cristata A.Braun Wetland near Interlaken, Tas., M.T. Casanova MEL p815 Figs 77, 78
30.xi.2005
N. cristata var. ambigua Barleyfields Lagoon near J. Divola MEL p266 Figs 75, 76
A.Braun Uralla, NSW, 29.ii.1996
Nitella sp. aff. furcata Howden, Tas., 20.vi.2008 H and A Wapstra HAW/03b MEL r090 Fig. 1d
N. gelatinifera R.D.Wood Canning River, WA, xi.1841 L. Preiss MEL p059 Figs 61, 62
N. gelatinifera var. afoliata Rocky River, Flinders Chase R.D. Wood 60-12-27-1 AD p946 Figs 67, 68
(R.D.Wood) J.C.van Raam NP, Kangaroo I., SA,
27.xii.1960
N. gelatinifera var. Boggy waters, Mersey River, C. Stuart 559 MEL r054 Figs 65, 66
cryptostachya (A.Braun) Tas.
J.C.van Raam
N. gelatinifera var. Wimmera, Vic. P. Eckhart 83 MEL r057 Figs 63, 64
microcephala (A.Braun)
J.C.van Raam
N. heterophylla A.Braun Condinar Pool, Pilbara region, M. Lyons and S. Lyons 3000 PERTH p637 Figs 47, 48
WA, 5.ix.2003
N. hyalina Agardh East dam at Casanovas, near M.T. Casanova MEL p468 Figs 41, 42
Chatsworth, Vic.,
24.xi.2002
N. imperialis (T.F.Allen) Denmark, WA, 9.x.1960 R.D. Wood 60-10-9-5 NY p957 Figs 27, 28
Sakayama
Nitella sp. 4 aff. leonhardii Blue Mountains, NSW, J.L. Porter 297 MEL p821 Figs 51, 52
R.D.Wood 24.i.2006
Nitella sp. 5 aff. leonhardii Freshwater Lake, Grampians M.T. Casanova MEL p299 Figs 53, 54
R.D.Wood region, Vic., 3.i.12
N. leptostachys A.Braun Creek near Mt Barker, WA, M.T. Casanova MEL p464 Figs 49, 50
2.x.2002
N. lhotzkyi A.Braun Lake Alexandrina, SA, E. Wollaston 61-1-1-1 NY p992 Figs 45, 46
1.i.1961
N. microteles M.B.Williams Creek 3 miles east of M.B. Williams 108 NSW r113 Figs 73, 74
Nimmitabel, NSW,
7.ii.1955
N. monopodiata J.C.van Raam 5 miles north of Golden Valley R.D. Wood 61-1-31-2A AD r040 Figs 19, 20
PO, Tas., 31.i.1961
N. myriotricha A.Braun Sunnybank, Qld, 14.viii.1926 C.T. White 3221 BM p901 Figs 55, 56
Nitella sp. 6 aff. myriotricha Yamburram Range, NT, N. Walsh 3773 MEL p105 Figs 57, 58
A.Braun 15.v.1994
Nitella sp. 1 aff. orientalis Rudal River, Pilbara region, M. Lyons and D. Mickle 3057 PERTH p641 Figs 9, 10
T.F.Allen WA, 29.v.2004
Nitella sp. 2 aff. orientalis Rudal River, Pilbara region, M. Lyons and D. Mickle 3054 PERTH p647 Figs 11,12
T.F.Allen WA, 29.v.2004
N. paludigena M.T.Casanova Ettrick Rd, Darlots Ck, Vic., A.C. Beauglehole 23, MEL p897 Figs 21, 22
& K.G.Karol 20.xii.1960
An overview of Nitella in Australia Australian Systematic Botany 199

Table 2. (continued )
Species Locality and date Collector and collector no. Herbarium Reference no. Figures
N. partita Nordst. Pied Stilt Swamp, Nocoleche J.L. Porter MEL p271 Figs 13, 14
Nature Reserve, Paroo River
region, NSW, 1.vi.2001
N. penicillata A.Braun Macquarie River at Ross H. and A. Wapstra HAW/01 MEL r082 Figs 15, 16
Bridge, Tas., 5.vi.2008
N. phauloteles J.Groves Dawson River, Qld, R.D. Wood 60-11-22-9 NY p956 Figs 23, 24
22.xi.1960
Nitella sp. 3. aff. Nerang, Qld, 2.xii.1960 R.D. Wood 60-12-2-3 NY p959 Figs 31, 32
pseudoflabellata A.Braun
Nitella sp. 8. aff. Lake St Clair, Tas., 25.i.2004 T. Dugdale and S. Talman 9b MEL P530 Figs 35, 36
pseudoflabellata A.Braun
N. remota A.Braun ex A.Braun Bangham Forest Reserve, SA, R.J. Bates 26076 AD r032 Figs 79, 80
& Nordstedt Oct. 1991
N. reticulata M.B.Williams Dune pond, Maroubra Beach, M.B. Williams 341 NSW r112 Figs 71,72
NSW, 20. vii.1955
N. sonderi A.Braun Tang Tang Swamp, Vic., K. Ough 8.2d, MEL p934 Figs 25, 26
10.v.2006
N. sonderi A.Braun Tang Tang Swamp, Vic., K. Ough CS1a MEL p907 Fig. 1b
28.iv.2006
N. stuartii A.Braun Tang Tang Swamp, Vic., K. Ough TT1i MEL p939 Figs 5, 6
22.i.2002
N. subtilissima A.Braun Lanark, near Branxholme, M.T. Casanova MEL p230 Figs 37, 38
Vic., DATE
N. subtilissima var. Field ditch pool near Mt R.D. Wood 60-11-9-8 AD r039 Figs 39, 40
subtilissima f. diffusa Gambier SA, 9.xi.1960
R.D.Wood
N. tasmanica Muell. ex South Esk River, Tas., C. Stuart 218 MEL p312 Figs 59, 60
A.Braun 25.xii.1848
N. tasmanica subspecies Creek 2 miles SW of Snelling R.D. Wood 60-12-29-11 NY, AD p947 Figs 69, 70
gelatinifera var. [ton] Beach, Kangaroo
beaugleholii R.D.Wood Island, SA, 29.xii.1960
Nitella sp. 7 aff. tricellularis Lake St Clair, Tas., 25.i.2004 T. Dugdale and S. Talbot 9a MEL p524 Figs 83, 84
Nordstedt
N. verticillata (Fil. et Lake Edward, SA, 13.vi.2004 M.T. Casanova MEL p612 Figs 33, 34
G.O.Allen ex Fil.)
R.D.Wood

or preserved in 70% alcohol. If oospores were not present, the
plants were either returned to the culture to mature, or kept in jars
on a windowsill until they matured and oospores were freely
released. If antheridia were present, chromosome counts were
attempted following the methods outlined in Casanova (1997).
Where possible and appropriate, oospores were removed from
the herbarium specimens or obtained as above from live material.
Approximately half of the specimens examined had oospores.
They were prepared for SEM examination by cleaning with a
detergent solution, by methods modified from those of Crawford
et al. (2001). Sometimes the enveloping cells were removed by
hand with fine needles, and sometimes freely released oospores
from fresh or cultured plants were not cleaned at all. For type
specimens, very old material, or specimens with a single oospore
available for examination, the oospores were handled with great
care and with minimal manipulation. Oospores were removed
from the stubs after microscopy and were stored in alcohol with
the specimen for possible future examination.
The data were used to compare individuals and allocate them
to species on the basis of the keys of Braun and Nordstedt (1882),
Nordstedt (1918), Groves and Allen (1935), Williams (1959),
Wood (1965, 1972), van Raam (1995) and Casanova et al.
(2007). Specimens were compared with type material and
type descriptions, and extensive review of the literature and
historical specimens was undertaken to determine historical
species concepts and allocate species names. In general, it was
considered insufficient to reinstate a species on the basis of a
single individual, and at least three gatherings with similar
oospores was needed to confirm a taxon.
Results
Examination of a large number of specimens, including type
specimens and their oospore ornamentation, and reference to the
scant literature on culture experiments on species of Nitella
facilitated the development of a species concept alternative to
that of Wood (1965). The major criteria used in the present study
to distinguish among species of Nitella were oospore
morphology, sexual state, dactyl morphology and the pattern
of branchlet furcations.
Consistent differences exist among the oospores of species in
subgenera Nitella and Tieffallenia and these differences are
generally visible under the light microscope as well as the
200 Australian Systematic Botany M. T. Casanova

scanning electron microscope. In these two subgenera, oospore
ornamentation is a valuable indicator of speciation. Sexual
characters that coincided with differences among oospores are
monoecy and dioecy, the development of highly contracted,
mucus-covered fertile heads or spikes, antheridia size and the
number of chromosomes. Vegetative characters that coincided
with differences among oospores were the number of branchlets,
the number of furcations, the number of branchlet segments at
each furcation (Fig. 1), the number of dactyl cells and the shape of
the dactyl end-cell (Fig. 2). Branchlet segment ratios (i.e.
macrodactyly v. brachydactyly: Fig. 1c, d), branchlet and axis
width ratios and the evenness of branchlet whorls were also
useful. Within subgenus Hyella, where oospores were (almost
without exception) reticulately ornamented, the size and shape
of the oospore, depth of the reticulum, the number of meshes
across the fossa and the presence of a flange on the striae
were useful in distinguishing among species. However, these
features are easy to see on SEM images only. The value of
SEM images is that they provide, on a single examination,
detection of characters that aid determination. Vegetative
characters can be modified by environment (e.g. removed by
herbivores, or be absent because of immaturity or senescence),
and it is a combination of vegetative characters that allows
determination, rather than a single feature. The subgenera and
sections of Nitella (sensu Wood 1965) are described and species
in them are listed below.

(i ) (a) (b) (c)

(d )
(e) (f )

(h)
(g)

Fig. 2. Dactyls (terminal branchlet segments) of Nitella spp. in Australia. (a) Anarthrodactylous (single-celled) dactyls.
(b) Bicellulate dactyls in which the penultimate cell is narrowed towards the end, and the end-cell is confluent with the end of
the penultimate cell. (c) Bicellulate dactyls in which the end-cell is confluent with the end of the penultimate cell.
(d) Bicelluate dactyls in which the end-cells are mucronate (bristle-tipped). (e) Bicelluate dactyls in which the end-cells are
cuspidate or partite. (f) Variable dactyls with 2 or 3 cells. (g) Pluricelluate (3-celled) dactyls in which the end-cell is allantoid.
(h) Pluricellulate dactyls in which the terminal two cells are reduced to a 2-celled mucro. (i) Pluricelluate dactyls (7-celled) in
which the end-cell is reduced, conical and acute.
An overview of Nitella in Australia
Key to the subgenera of Nitella in Australia
1. Terminal branchlet segments consisting of single cells
(anarthrodactylous) ....................................................subgenus Nitella
Terminal branchlet segments consisting of 2 to many cells
(arthrodactylous) .................................................................................2
2. Terminal branchlet segments consisting of 2 (rarely 3) cells (bicellulate),
oospore ornamentation papillate, verrucate, granulate, spongy, fibrous,
scabrous or smooth ............................................ subgenus Tieffallenia
Terminal branchlet segments consisting of 3–7 cells (pluricellulate),
oospore ornamentation reticulate or flanges on the striae united into an
apical crest ................................................................. subgenus Hyella
Nitella subgenus Nitella
Subgenus Nitella is characterised by single-celled dactyls
(Fig. 2a). Karol et al. (2001) found in nucleotide-sequencing
studies that this subgenus is polyphyletic, and that sections Rajia
and Palia (the two sections that are represented in Australia)
belong to genetically distant branches of genus Nitella. Each of
these sections is represented by a single species in Australia.
Key to the sections of subgenus Nitella in Australia
1. Branchlets in a single whorl at axis nodes, oospore smooth.........................
............................................................section Rajia: Nitella acuminata
Branchlets in two or more whorls at the axis nodes, oospore with reticulate
ornamentation ........................................ section Palia: Nitella stuartii
Section Rajia
Nitella acuminata has elongate, single-celled dactyls (Fig. 2a)
and homeoclemous whorls. This species is rare in Australia and
few specimens exist in Australian herbaria. It is a tropical species,
and is most abundant and diverse in South-east Asia (Groves
1922; Zaneveld 1940; Han and Li 1994). Wood thought that the
few specimens he saw from Australia were possibly N. acuminata
f. belangeri, although insufficient material was available at that
time to confirm this determination (Wood 1972). García and
Chivas (2006) reported on three localities for N. acuminata from
Queensland; however, these authors do not give varietal or form
designation. The oospores of Australian specimens (Figs 3, 4) are
smooth, with five or six flanged striae. Specimens referred to
N. acuminata (not f. belangeri) by other authors (Leitch et al.
1990) have a diversity of oospore wall morphologies, including
scabrous and smooth. The description of the oospore of
N. acuminata f. belangeri in Wood (1965) is ‘flanged’ and
‘coarsely granulate’, although this determination cannot be
ruled out or confirmed on the basis of the single oospore that
was available for examination (p149). Collection and
examination of additional material is needed to confirm the
presence of N. acuminata in Australia.
Section Palia
Nitella stuartii is the only species in this section in Australia
(p939, Figs 5, 6). The vegetative morphology of this species is
characterised by single-celled dactyls and heteroclemous whorls
Australian Systematic Botany 201
(Fig. 1a as an example of heteroclemy). It occurs sparsely in
Tasmania (Wood 1972; van Raam 1995), in Victoria (Casanova
and Karol 2008), through the Northern Tablelands of New South
Wales (Casanova 1993), in south-eastern Queensland (Groves
and Allen 1935), and has been collected in New Zealand
(Nordstedt 1888; Wood and Mason 1977), India (Pal et al.
1962) and China (Han and Li 1994). There is some variation
in the morphology of oospores for this taxon (John and Moore
1987; Casanova 1991; Han and Li 1994); some Indian and
Australian specimens have oospores ornamented with a
beaded reticulum and some Australian specimens have
oospores with a simple reticulum (John and Moore 1987: figs
13, 14, 16, 17, 53). Some of the variation is due to differences in
the maturity of oospores selected for SEM (Casanova 1991).
Australian specimens are distinctively shrubby, and they are
generally found in the shallows of fresh, flowing water
(Casanova and Karol 2008) or in shallow, temporary,
freshwater wetlands (Casanova 1993).
Nitella subgenus Tieffallenia
Tieffallenia has terminal branchlet segments (dactyls) with short,
conical end-cells, much smaller than the preceding dactyl cell(s)
(Fig. 2b–d). Most species in this subgenus have two-celled
dactyls, but species with dactyls consisting of more than two
cells were included in this subgenus by Wood (1965) if the end-
cell was conical and reduced in comparison with the other dactyl
cells (Fig. 2f, h). Work on nucleotide sequences by Karol (2004)
indicated that such two- or three-celled species (in section
Migularia) should not be included in subgenus Tieffallenia,
rather they should be referred to subgenus Hyella. The
subgenus Tieffallenia is divided into several sections
containing Australian species: sections Tieffallenia, Gioallenia
and Decandollea, each with several species, and sections
Muelleria and Vogania, with one species each. All members
of this subgenus have homeoclemous branchlet whorls
(Fig. 1b–d), except members of section Decandollea, which
have heteroclemous whorls (Fig. 1a).
Key to sections in subgenus Tieffallenia
1. Branchlets in a single whorl at the nodes ................................................ 2
Branchlets in two or three whorls at the nodes, oospores with spongy,
fibrous wall and flange construction, or ‘string of beads’ (cystoseriate)
ornamentation ...................................................... section Decandollea
2. End-cells regularly cuspidate or 2–3–4 partite, oospores flanged, oospore
wall papillate.................................... section Muelleria: Nitella partita
End-cells on most branchlets coming to a single point, rarely
bifurcate..............................................................................................3
3. Dactyls highly inflated, almost spherical; oospores slightly flanged and
with ornamentation of irregular projections ........................................
.......................................................... section Vogania: Nitella tumida
Diameter of terminal branchlet segments similar to the other
segments .............................................................................................4
4. Dactyls much shorter than preceding segments
(brachydactylous)................................................... section Tieffallenia
Dactyls as long, or longer than preceding segments
(macrodactylous) .....................................................section Gioallenia
202 Australian Systematic Botany M. T. Casanova

3 4

5 6

7 8

9 10
An overview of Nitella in Australia Australian Systematic Botany 203

Section Tieffallenia Section Muelleria

The Nitella species in this section were defined by Wood (1965)
as large plants with mucronate end-cells (Fig. 2d) or
brachydactyly (Fig. 1d), with reproductive whorls rarely
contracted into heads and with no mucus. According to
Wood’s (1972) treatment, Australia has N. furcata (subsp.
furcata and orientalis) only in this subgenus. The separation
of this section on morphological criteria is questionable
because the characters that define it are ones that either vary in
relation to environment (plant size) or can vary during plant
development (brachydactyly). Additionally, the placement of
species in the same group on the basis of either reduced end-
cells or reduced dactyls is odd, because whole dactyls and end-
cells are not analogous structures on Australian members of
subgenus Tieffallenia. All Australian species that could be
members of this subgenus have bicellulate dactyls and a
single whorl of branchlets at the axis nodes (Fig. 1b–d:
homeoclemous). Cosmopolitan N. furcata was first mentioned
for Australia by Groves and Allen (1935), on the basis of a
specimen from the Endeavour River (Qld). They amalgamated
N. furcata with N. polyglochin (from India) and separated it
from N. oligospira f. australiensis, N. orientalis and
N. microcarpa on the basis of the distinctively elongate upper
row of cells in the coronula of the oogonium, and the
clustered oogonia. Wood (1972) went further and
amalgamated all of these brachydactylous taxa into N. furcata,
along with N. phauloteles, and stated that the available
specimens of N. furcata were almost invariably sterile or poor
material. Sakayama et al. (2002) found that N. furcata was
distinguishable from other taxa amalgamated with it in
Japan, on the basis of nucleotide sequences and oospore
ornamentation, as well as vegetative features. Specimens of
N. furcata in Japan (Sakayama et al. 2002: figs 4–6) have
oospores different from those of specimens corresponding
to Wood’s (1972) description of N. furcata in Australia
(p641, p647, Figs 7–12). Van Raam (1995) resurrected
N. australiensis ( = N. oligospira f. australiensis, p807,
Figs 7, 8) on the basis of Tasmanian material and
distinguished it from N. oligospira of North and South
America, and from N. furcata f. indica of Indonesia. Although
specimens that are similar to Wood’s broad concept of N. furcata
have been found in Australia, it is possible that N. furcata does
not occur in Australia; rather, there may be several endemic
species. The status of this section in subgenus Tieffallenia
requires further examination.
Nitella partita (p271, Figs 13, 14) is the only species in this
section, which is defined by the presence of cuspidate end-cells
(Fig. 2e). In Wood’s (1965) treatment, he recognised N. partita on
the basis of Nordstedt’s (1891) illustrations and description (the
type being lost). Later, when no further material of this species had
been seen, he amalgamated it with N. furcata as an unusual variant
(Wood 1972). Since then, new collections of N. partita have been
made (Casanova et al. 2003). As Wood (1972) noted, partite or
cuspidate cells can occur as an abnormality on several species of
charophytes (Han and Li 1994). In those cases, the occurrence
of such a cell is occasional, and normal cells are found on most of
the plant. In N. partita, end-cells vary in the number of points
(2–4 per cell), although partite end-cells occur on every dactyl
examined, and on every plant in the populations seen so far.
Further, specimens have been collected (rarely) from the field,
as well as grown under defined conditions in a greenhouse or tank
culture. It is likely that this species is ephemeral in its occurrence
and rarely collected. It is distributed in ephemeral flood-
plain wetlands of New South Wales and Queensland arid-
zone rivers (Casanova et al. 2003), as well as on Middle
Island in Western Australia (Casanova 2003). Its oospores are
distinctively flanged (Figs 13, 14). Records of N. partita in China
(Han and Li 1994) appear to be in error.
Section Vogania
Nitella tumida is the only species in this section, so the section is
characterised by the features of the species. There are three
records of N. tumida from inland (usually saline) water bodies
(Wood 1972; García and Chivas 2006). The axes are long and
thin, with whorls of inflated branchlets at their apices. An isotype
specimen exists in MEL; however, oospores of this species have
not been available for examination. Nordstedt (1891) described
the oospores as yellowish to bright brown, 315–330 mm long,
with seven or eight slightly flanged, prominent ridges. The
ornamentation is smooth, or with more or less irregular
projections 4–6 mm in diameter (Nordstedt 1891; Wood 1965).
Section Gioallenia
This section is characterised by species with conical end-cells,
the diameter of which is similar to the penultimate dactyl cell
and elongate dactyls (Fig. 2b, c). Wood (1965) retained
N. pseudoflabellata and N. subtilissima as Australian species
in section Gioallenia, and recognised a large amount of variation

Figs 3–10. Figs 3, 4. Oospore of a specimen of p149, Nitella acuminata A.Braun ex Wallm., from Moochalabra Dam, Western Australia (A.A. Mitchell 3703B,
27.vii.1994, PERTH). Fig. 3. Whole oospore (316 mm  225 mm), showing smooth ornamentation and 5 or 6 striae with low flanges. Scale bar = 100 mm.
Fig. 4. Detail of oospore wall, showing smooth surface and low flanges on the striae. Scale bar = 20 mm. A single oospore was available for examination, so it is
possible the specimen is immature. Figs 5, 6. Oospore of a specimen of p939, Nitella stuartii A.Braun, from Tang Tang Swamp, Victoria (K. Ough TT1i, 22.i.2002,
MEL). Fig. 5. Whole oospore (230 mm  195 mm), with reticulate ornamentation on the fossa wall and unornamented flanges on the striae. Scale bar = 100 mm.
Fig. 6. Detail of oospore wall, showing shallow reticulate ornamentation and the base of a flange. Scale bar = 10 mm. This is similar to the type material of N. stuartii
(BM, MEL). Figs 7, 8. Oospore of a specimen of p807, Nitella australiensis (Nordst.) J.C.van Raam, from Lake Dulverton, Tasmania, seed-bank culture
(M.T. Casanova, 22.xi.2005, MEL). Fig. 7. Whole oospore (314 mm  195 mm) with no discernable ornamentation, striae consisting of weak ridges.
Scale bar = 100 mm. Fig. 8. Detail of oospore wall showing shallowly rippled surface. Scale bar = 5 mm. Several freely released oospores were examined
for this specimen, and the specimen illustrated is representative of the material examined. Figs 9, 10. Oospore of a specimen of p641, Nitella sp. 1 aff. orientalis
T.F.Allen, from Rudal River, Pilbara region, Western Australia (M. Lyons and D. Mickle 3057, 29.v.2004, PERTH). Fig. 9. Whole oospore (205 mm  165 mm),
showing apparently smooth fossa wall and 5 or 6 flanged striae. Scale bar = 100 mm. Fig. 10. Detail of oospore wall showing finely granulate surface with delicate,
fimbriate flanges on the striae. Scale bar = 20 mm.
204 Australian Systematic Botany M. T. Casanova

11 12

13 14

15 16

17 18
An overview of Nitella in Australia Australian Systematic Botany 205

within those taxa by delineating four varieties and nine forms in
N. pseudoflabellata and three varieties and five forms in
N. subtilissima (Wood 1972). The section is enlarged here to
include four species from Tasmania (N. penicillata (r082, Figs 15,
16), N. confusa (r036, Figs 17, 18), N. haagenii, N. monopodiata
(r040, Figs 19, 20), one from New South Wales (N. ignescens),
N. paludigena (p897, Figs 21, 22) and N. aemula from South
Australia and Victoria, and N. phauloteles (p956, Figs 23, 24)
from Queensland (see section Tieffallenia above). N. sonderi
(p934, Figs 25, 26) is also included in this section. It appears that
monoecious N. imperialis (p957, Figs 27, 28, similar to the type of
N. imperialis from Japan), N. comptonii (p912, Figs 29, 30, from
the same locality as a specimen recorded by Groves and Allen
1935), N. pseudoflabellata (p956, Figs 31, 32) and dioecious
N. robertsonii (with smooth oospores, Braun and Nordstedt 1882;
Groves and Allen 1935) also occur in Australia. N. verticillata
(p612, Figs 33, 34) is transferred from section Muelleria (Wood
1965) to section Gioallenia here, on the basis of its bicellulate
dactyls. Good, fertile specimens of this species are found
infrequently (García and Chivas 2006) because of its small
stature (Wood 1972). The status of larger plants in New South
Wales that meet the description of N. verticillata (Hotchkiss and
Imahori 1988b) needs investigation. N. batrachosperma was
listed for Australia by Nordstedt (1918); however, the only
specimen with this determination in the National Herbarium of
Victoria (Lake Wellington, Mueller 2) has oospores consistent
with those of N. lhotzkyi. N. mucosa (described on the basis of
New Zealand material) was recorded for Australia by Nordstedt
(1918) although the Australian specimens examined so far are
different from the two taxa present in New Zealand (de Winton
et al. 2007).
A preliminary investigation (Casanova 2007b) of the oospore
variation among the entities remaining in N. pseudoflabellata
sensu Wood (1972) reveals that there are at least five different
oospores among them that cannot be allocated to previously
described taxa (e.g. N. sp. 8 aff. pseudoflabellata; Figs 35, 36).
It is likely that there are some new species in this section. The
presence of N. horikawae and N. imahorii (Hotchkiss and Imahori
1988a) has not been confirmed, and a comparison of the oospores
of the type of N. morongii (Casanova and Karol 2008) with those
illustrated by Sakayama et al. (2005) does not support the record
of N. morongii for Australia.
Similarly, N. subtilissima needs revision. Three specimens
collected in sequence by Drummond from the same place (Swan
River, Drummond 7, 9 and 10) were thought unlikely to be the
type material of three different species (Wood 1972), so they were
relegated to varieties of N. subtilissima. However, the
descriptions of the species on the basis of these specimens
differ significantly (e.g. in sexual state, presence of mucus and
contraction of the fertile branchlets). Specimens similar to the
description of N. subtilissima, with no mucus, contracted fertile
spikes, brachydactylous sterile branchlets (Fig. 1d) and oospores
with a granulate ornamentation (p230, Figs 37, 38) occur in
Victoria and New South Wales. The oospores of specimens
allocated to forms of N. subtilissima described by Wood
(1972) (e.g. var. subtilissima f. diffusa (r039, Figs 39, 40))
differ significantly. N. gloeostachys, with finely granulate
oospore ornamentation, was reinstated by van Raam (1995)
after examination of type material. On the basis of oospore
variation alone, the status of the subspecific taxa and the
species amalgamated with N. subtilissima by Wood needs
examination.
Section Decandollea
Decandollea contains species with bicellulate dactyls in which
the end-cell is confluent with the penultimate cell (Fig. 2c), and
with two or more whorls of branchlets at the nodes
(heteroclemous whorls, Fig. 1a). Monoecious N. hyalina
(p468, Figs 41, 42) is the well recognised representative of
this section and has a worldwide distribution. Despite a large
range in morphology in this species, all specimens so far
examined have fibrous oospore ornamentation. According to
Wood’s (1972) revision, Australia has the only additional
species in the section, all dioecious; N. congesta (p255,
Figs 43, 44), N. lhotzkyi (varieties lhotzkyi (p992, Figs 45, 46),
biformis and minor) and N. penicillata (here transferred to section
Gioallenia). Dioecious N. heterophylla, described on the basis of
a specimen from Western Australia, was retained by Wood (1972)
as a form of N. lhotzky; however, it should be reinstated as a
separate species because (after examination of ~30 specimens
from the Pilbara region and elsewhere in Western Australia) its
vegetative morphology and oospore (p637, Figs 47, 48) have
been found to be consistently different from those of N. lhotzkyi,
N. congesta and N. hyalina. Another species, N. ungula,
described by García (1996), was found to be related to these
taxa (Karol 2004), and García and Chivas (2006) mentioned N. cf.
lhotzkyi.
Wood’s placement of N. penicillata in this section is
somewhat inexplicable, as the original description of

Figs 11–18. Figs 11,12 Oospore of a specimen of p647, Nitella sp. 2 aff. orientalis T.F.Allen, from Rudal River, Pilbara region, Western Australia
(M. Lyons and D. Mickle 3054, 29.v.2004, PERTH). Fig. 11. Whole oospore (266 mm  235 mm), showing shallow reticulate ornamentation on the fossa wall and
6 striae with weak flanges. Scale bar = 100 mm. Fig. 12. Detail of oospore wall, showing shallow, incomplete, reticulate ornamentation and weak flanges.
Scale bar = 20 mm. Figs 13, 14. Oospore of a specimen of p271, Nitella partita Nordst., from Pied Stilt Swamp, Nocoleche Nature Reserve, Paroo River region,
New South Wales (J.L. Porter, 1.vi.2001, MEL). Fig.13. Whole oospore (233 mm  255 mm), showing granulate oospore wall and 5 or 6 striae with large, firm
flanges. Scale bar = 100 mm. Fig. 14. Detail of oospore wall, showing irregular grains on the surface of the fossa. Scale bar = 20 mm. Figs 15, 16. Oospore of a
specimen of r082, Nitella penicillata A.Braun, from Macquarie River at Ross Bridge, Tasmania (H. and A. Wapstra HAW/01, 5.vi.2008, MEL). Fig.15. Whole
oospore (250 mm  172 mm), showing papillate ornamentation and 6 or 7 striae with smooth flanges. Scale bar = 50 mm. Fig. 16. Detail of oospore wall,
showing papillate projections and low flanges on the striae. Scale bar = 20 mm. Nordstedt (1918) described this ornamentation as‘small, tooth-like (dentate)
scales’. Figs 17,18. Oospore of the isotype specimen of r036, Nitella confusa (R.D.Wood) J.C. van Raam, from Cynthia Bay, Lake St Clair, Tasmania (R.D. Wood
61-1-2-12A, 24.xii.1961, AD). Fig. 17. Whole oospore (358 mm  307 mm), showing smooth to rippled ornamentation and ~7 striae with low ridges. Scale
bar = 100 mm. Fig. 18. Detail of oospore wall, showing textured ornamentation and low ridges on the striae. Scale bar =20 mm. Wood’s (1972) description of the
oospore was ‘membrane imperfectly finely reticulate’. It is possible that this oospore is immature.
206 Australian Systematic Botany M. T. Casanova

19 20

21 22

23 24

25 26

Figs 19–26. Figs 19, 20. Oospore of a specimen of r040, Nitella monopodiata J. C.van Raam ( = N. subtilissima var. microphylla f. tasmanica R.D.Wood), from
Tasmania (R.D. Wood 61-1-31-2A, 31.i.1961, AD). Fig. 19. Whole oospore (300 mm  220 mm), showing granulate oospore ornamentation and ~9 striae with
smooth flanges. Scale bar = 100 mm. Fig. 20. Detail of oospore wall, showing evenly spaced, close-set granulate ornamentation and smooth flanges on the striae.
Scale bar = 20 mm. Figs 21, 22. Oospore of a specimen of p897, Nitella paludigena M.T.Casanova & K.G.Karol, from Ettrick Rd, Darlots Creek, Victoria (A.
C. Beauglehole 23, 20.xii.1960, MEL). Fig. 21. Whole oospore (230 mm  194 mm) from the dried specimen, with indistinct vermiferous ornamentation and thick
ridges on the 8 striae. Scale bar = 50 mm. Fig. 22. Detail of oospore wall near the base of the oospore. Ornamentation of sinuous and simple vermiferous papillae.
Scale bar = 10 mm. This is similar to the type material of N. paludigena (MEL). Figs 23, 24. Oospore of a specimen of p956, Nitella phauloteles J.Groves, from
Dawson River, Queensland (R.D. Wood 60-11-22-9, 22.xi.1960, NY). Fig. 23. Whole oospore (265 mm  210 mm), showing apparently smooth oospore
ornamentation with 7 flanged striae. Scale bar = 100 mm. Fig. 24. Detail of oospore wall, showing minutely granulate ornamenation that continues up the flanges.
Scale bar = 5 mm. Figs 25, 26. Oospore of a specimen of p934, Nitella sonderi A.Braun, from Tang Tang Swamp, Victoria, seed-bank culture (K. Ough 8.2d,
10.v.2006, MEL). Fig. 25. Whole oospore (240 mm  208 mm), showing papillate ornamentation on the fossa wall, and 8-flanged, ornamented striae. Scale
bar = 100 mm. Fig. 26. Detail of apex of the oospore, showing the long papillae, some of which are joined at the base, sometimes arranged in rows perpendicular to
the striae. Scale bar = 20 mm. This is similar to the type material of N. sonderi (MEL).
An overview of Nitella in Australia Australian Systematic Botany 207

27 28

29 30

31 32

33 34

Figs 27–34. Figs 27, 28. Oospore of a specimen of p957, Nitella imperialis (T.F.Allen) Sakayama, from Denmark, Western Australia (R.D. Wood 60-10-9-5, 9.
x.1960, NY). Fig. 27. Whole oospore (307 mm  270 mm), showing apparently smooth fossa wall and 7 or 8 low ridges on the striae. Scale bar = 50 mm.
Fig. 28. Detail of oospore wall, showing flattened granulate ornamentation. Scale bar = 10 mm. This is similar to the oospore of the type material of N. imperialis
(NY). Figs 29, 30. Oospore of a specimen of p912, Nitella comptonii J.Groves, from Enoggera Reservoir, Brisbane, Queensland (R.D. Wood 60-11-21-23,
21.xi.1960, NY). Fig. 29. Whole oospore (297 mm  202 mm), showing granulate ornamentation, with 7 or 8 ridges on the striae. Scale bar = 100 mm. Fig. 30.
Detail of oospore wall and striae, showing a band of papillae or grains along the centre of the fossa wall. Scale bar = 20 mm. This is similar to the oospore depicted in
Sakayama et al. (2006). Figs 31, 32. Oospore of a specimen of p959, Nitella sp. 3. aff. pseudoflabellata A.Braun, from Nerang, Queensland (R.D. Wood 60-12-2-3,
2.xii.1960, NY). Fig. 31. Whole oospore (285 mm  250 mm), showing felted, fibrous oospore wall and 8 or 9 striae. Scale bar = 100 mm. Fig. 32. Detail of oospore
wall, showing wall ornamentation of felted fibres with pores, from which both the oospore wall and the short flanges on the striae are constructed. Scale bar = 20 mm.
This differs from the description of the oospore of the type material of N. pseudoflabellata (Allen 1953). Figs 33, 34. Oospore of a specimen of p612, Nitella
verticillata (Fil. et G.O.Allen ex Fil.) R.D.Wood, from the seed bank of Lake Edward, South Australia (M.T. Casanova, 13.vi.2004, MEL). Fig. 33. Whole oospore
(~175 mm  150 mm) viewed from the apex, showing close-set granulate ornamentation and indistinct striae. Scale bar = 50 mm. Fig. 34. Detail of the base of the
oospore, showing the impression of the basal cells of the oogonium with tightly packed, rounded papillate ornamentation. Scale bar = 10 mm.
208 Australian Systematic Botany M. T. Casanova

35 36

37 38

39 40

41 42

Figs 35–42. Figs 35, 36. Oospore of a specimen of p530, Nitella sp. 8 aff. pseudoflabellata, from Lake St Clair, Tasmania (T. Dugdale and S. Talman 9b, 25.
i.2004, MEL). Fig. 35. Whole oospore (360 mm  290 mm), with a felted appearance and 8 or 9 striae with small flanges. Scale bar = 100 mm. Fig. 36. Detail of
oospore wall, showing fine, papillate-granulate ornamentation and small, shallowly ornamented flanges. Scale bar = 10 mm. Figs 37, 38. Oospore of a specimen of
p230, Nitella subtilissima A.Braun, from Lanark, near Branxholme, Victoria (M.T. Casanova, MEL). Fig. 37. Whole oospore (280 mm  251 mm), showing
apparently smooth ornamentation and 6 striae with fimbriate flanges. Scale bar = 100 mm. Fig. 38. Detail of oospore wall, showing finely granulate–papillate
ornamentation. Scale bar = 5 mm. This is similar to the oospore of the type of N. subtilissima var. microphylla depicted by Leitch et al. (1990). Figs 39, 40. Oospore
of a specimen of r039, Nitella subtilissima var. subtilissima f. diffusa R.D.Wood, from a Field ditch pool near Mt Gambier, South Australia (R.D. Wood 60-11-9-8,
9.xi.1960, AD). Fig. 39. Whole oospore (182 mm  146 mm), showing smooth oospore surface and ~5 striae with low ridges. Scale bar = 50 mm. Fig. 40. Detail of
oospore wall, with minutely granulate ornamentation and low ridges on the striae. Scale bar = 10 mm. Figs 41, 42. Oospore of a specimen of p468, Nitella hyalina
Agardh, from the East dam at Casanovas, near Chatsworth, Victoria (M.T. Casanova, 24.xi.2002, MEL). Fig. 41. Whole oospore (350 mm  280 mm), showing 7-
or 8-flanged striae and fibrous construction. Scale bar = 100 mm. Fig. 42. Detail of oospore wall, showing matted fibrous oospore wall and flanges.
Scale bar = 20 mm. This is similar to other material of N. hyalina depicted in John and Moore (1987).
An overview of Nitella in Australia Australian Systematic Botany 209

43 44

45 46

47 48

49 50

Figs 43–50. Figs 43, 44. Oospore of a specimen of p255, Nitella congesta (R.Br.) A.Braun, from Gahnia Lagoon 5, Brinkworth wetlands, South Australia
(M.A. Brock, 24.ii.1997, MEL). Fig. 43. Whole oospore (506 mm  354 mm), showing smooth oospore wall and 7 or 8 striae with smooth flanges and an apical
crest. Scale bar = 200 mm. Fig. 44. Detail of oospore wall, showing smooth surface with underlying spongy construction where a flange has been damaged during
cleaning. Scale bar = 20 mm. Figs 45, 46. Oospore of a specimen of p992, Nitella lhotzkyi A.Braun, from Lake Alexandrina, South Australia (E. Wollaston
61-1-1-1, 1.i.1961, NY). Fig. 45. Whole oospore (320 mm  266 mm), showing verrucate ornamentation and 7- or 8-flanged striae. Scale bar = 100 mm.
Fig. 46. Detail of oospore wall, showing ornamentation of scattered verrucae connected by low walls into an uneven reticulate pattern. Scale bar = 10 mm. This is
similar to the illustration of the oospore wall of the type specimen of N. lhotzkyi (Nordstedt 1889). Figs 47, 48. Oospore of a specimen of p637, Nitella heterophylla
A.Braun, from Condinar Pool, Pilbara region, Western Australia (M. Lyons and S. Lyons 3000, 5.ix.2003, PERTH). Fig. 47. Whole oospore (325 mm  256 mm),
showing fibrous oospore wall and very large flanges on the 6 or 7 striae. Scale bar = 100 mm. Fig. 48. Detail of oospore wall, showing felted, fibrous construction
of both fossa and flange. Scale bar = 20 mm. Figs 49, 50. Oospore of a specimen of p464, Nitella leptostachys A.Braun, from a Creek near Mount Barker, Western
Australia (M.T. Casanova, 2.x.2002, MEL). Fig. 49. Whole oospore (261 mm  233 mm), showing reticulate ornamentation and flanges on the 8 or 9 striae. Scale
bar = 100 mm. Fig. 50. Detail of flanges, 20–25 mm high, showing smooth surface with grains, rather than the ridges of the reticulum. Scale bar = 20 mm.
210 Australian Systematic Botany M. T. Casanova

51 52

53 54

55 56

57 58

Figs 51–58. Figs 51, 52. Oospore of a specimen of p821, Nitella sp. 4 aff. leonhardii R.D.Wood, from the Blue Mountains, New South Wales (J.L. Porter 297, 24.
i.2006, MEL). Fig. 51. Whole oospore (280 mm  258 mm), showing small size, reticulate ornamentation and ~6 striae. Scale bar = 100 mm. Fig. 52. Detail of
oospore wall, showing reticulate ornamentation that extends onto the ridges on the striae. Scale bar = 20 mm. Figs 53, 54. Oospore of a specimen of p299, Nitella sp.
5 aff. leonhardii R.D.Wood, from Freshwater Lake, Grampians region, Victoria (M.T. Casanova, 3.i.12, MEL). Fig. 53. Whole oospore (342 mm  303 mm),
showing reticulate ornamentation continuing onto the 8 ridges on the striae. Scale bar = 100 mm. Fig. 54. Detail of oospore wall, showing the deep reticulum with ~5
meshes across the fossa and extension of the ornamentation onto the ridges on the striae. Scale bar = 20 mm. Figs 55, 56. Oospore of a specimen of p901, Nitella
myriotricha A.Braun, from Sunnybank, Queensland (C.T. White 3221, 14.viii.1926, BM). Fig. 55. Whole oospore (226 mm  240 mm), showing reticulate
ornamentation and short flanges on the 7 striae. Scale bar = 100 mm. Fig. 56. Detail of oospore wall, showing reticulate ornamentation with ~4 meshes across the
fossa, and smooth flanges 25–30 mm high. Scale bar = 20 mm. This specimen is similar to the oospore of N. myriotricha collected in Victoria in the vicinity of the
type collection, and to the New South Wales material. Figs 57, 58. Oospore of a specimen of p105, Nitella sp. 6 aff. myriotricha A.Braun, from Yamburram Range,
Northern Territory (N. Walsh 3773, 15.v.1994, MEL). Fig. 57. Whole oospore (400 mm  300 mm), showing finely reticulate oospore wall and 9- or 10-flanged
striae. Scale bar = 100 mm. Fig. 58. Detail of oospore wall, showing shallow reticulate ornamentation, with ~12 meshes across the fossa. Scale bar = 5 mm.
An overview of Nitella in Australia Australian Systematic Botany 211

59 60

61 62

63 64

65 66

Figs 59–66. Figs 59, 60. Oospore of the isotype specimen of p312, Nitella tasmanica Muell. ex A.Braun, from South Esk River, Tasmania (C. Stuart 218, 25.
xii.1848, MEL). Fig. 59. Damaged oospore, showing reticulate ornamentation on the fossa wall and striae of ornamented low ridges. Scale bar = 100 mm. Fig. 60.
Detail of oospore wall, showing reticulate ornamentation of ~4 meshes across the fossa. Scale bar = 10 mm. Figs 61, 62. Oospore of the isotype specimen of p059,
Nitella gelatinifera R.D.Wood, from Canning River, Western Australia (L. Preiss, Nov. 1841, MEL). Fig. 61. Whole oospore (170 mm  170 mm), showing
smoothly reticulate ornamentation continuing up the 6 or 7 ridges on the striae. Scale bar = 50 mm. Fig. 62. Detail of oospore wall, showing the uneven nature of the
reticulum of ~4 meshes across the fossa wall, and the ornamented ridges on the striae. Scale bar = 20 mm. Figs 63, 64. Oospore of a specimen of r057, Nitella
gelatinifera var. microcephala (A.Braun) J.C.van Raam, from the Wimmera, Victoria (P. Eckhart 83, MEL). Fig. 63. Whole oospore (187 mm  137 mm), showing
smoothly reticulate oospore wall and 6 or 7 striae with ornamented ridges. Scale bar = 50 mm. Fig. 64. Detail of oospore wall, showing reticulate surface (~2 meshes
across the fossa) and ridged flanges. Scale bar = 20 mm. Figs 65, 66. Oospore of an isotype specimen of r054, Nitella gelatinifera var. cryptostachya (A.Braun)
J.C.van Raam, from Boggy waters, Mersey River, Tasmania (C. Stuart 559, MEL). Fig. 65. Whole oospore (205 mm  168 mm), showing reticulate oospore
wall and 6 or 7 striae with smooth flanges. Scale bar = 50 mm. Fig. 66. Detail of oospore wall, showing papillate–reticulate surface and smooth flanges.
Scale bar = 20 mm.
212 Australian Systematic Botany M. T. Casanova

67 68

69 70

71 72

73 74
An overview of Nitella in Australia Australian Systematic Botany 213

N. penicillata indicated that it was very different from the rest of
the section, with mucronate end-cells, homeoclemous whorls and
lacking mucus (Braun 1852). Braun and Nordstedt (1882) placed
it alongside N. sonderi and N. robertsonii. Its oospores were
described as being covered with little prickles, or toothed scales
similar to those of N. sonderi (Nordstedt 1889) as depicted by
Leitch et al. (1990: fig. 79). Specimens (including type
specimens) of N. penicillata that have been examined are more
closely aligned with species in section Gioallenia (see above)
than those in section Decandollea.
Nitella subgenus Hyella
The unifying characteristic of species in subgenus Hyella is that
the end-cell of the dactyls is long-cylindrical (allantoid) rather
than conical and acute (Fig. 2g), except where there are more than
three cells (pluricellulate; Figs 2f–i). Most species in this group
have both pluricellulate dactyls with three to five cells, and
allantoid end-cells. However, some species have only allantoid
end-cells, and other species have pluricelluate dactyls with
conical, acute end-cells. Branchlets are usually more than
2  furcate, although some taxa with two, one and zero
furcations are present in the group. Wood (1965) did not
divide Hyella into sections. The N. cristata–hookeri complex
(i.e. section Migularia) was not included in this group by Wood
(1965, 1972), presumably because their end-cells are uniformly
conical and acute; however, it is included here.
Key to sections in subgenus Hyella
1. Terminal branchlet segments 3–7 cells long, the cells not tapering, the end-
cell variable, oospores papillate to reticulate................. section Hyella
Terminal branchlet segments of 2–3–4 cells, the cells confluent and
tapering to an acute end-cell, or the terminal two cells together
forming a mucro, oospores with a crest of united flanges at the
apex ......................................................................... section Migularia
Section Hyella
According to Wood (1972) Australia has three species in section
Hyella. However, there is a huge amount of variation among
specimens and there are likely to be more than 10 easily
distinguished species in the group. Almost all of these have
reticulate oospores (Figs 49–70). There are two accepted
monoecious species, N. leptostachys (p464, Figs 49, 50) and
N. leonhardii (van Raam 1995). There is oospore and vegetative
variation in N. leonhardii, and entities that key out to
N. leonhardii from eastern Australia (p821, p299, Figs 51–54)
and New Zealand need to be compared (Casanova and Karol
2008). Dioecious species include N. myriotricha, N. tasmanica
and N. gelatinifera. The initial gathering of N. myriotricha, with
highly divided, soft, mucus-covered branchlets and fertile parts in
contracted whorls, was in Victoria, and specimens from New
South Wales and Queensland (p901, Figs 55, 56) are similar to the
type. However, additional specimens from Northern Territory
(p105, Figs 57, 58) and Western Australia have very different
oospores and differ in their vegetative features as well.
N. tasmanica (p312, Figs 59, 60) and N. gelatinifera (r059,
Figs 61, 62) were united by Wood (1962) under the name
N. tasmanica; however, van Raam (1995) reinstated
N. gelatinifera as a species, recombining Wood’s varieties
microcephala (r057, Figs 63, 64), cryptostachya (r054,
Figs 65, 66) and afoliata (p946, Figs 67, 68) (of N. tasmanica
subspecies gelatinifera) as varieties of N. gelatinifera.
Examination of specimens from throughout Australia supports
this separation, and suggests that some varieties and forms of
N. gelatinifera should be raised further to species. The status of
N. tasmanica ssp. gelatinifera var. beaugleholii is problematic.
Examination of the type material has not confirmed the presence
of pluricellulate dactyls, although the overall morphology is
similar to specimens of N. gelatinifera var. afoliata. This
difference is compounded by the lack of reticulate
ornamentation (p947, Figs 69, 70). On the basis of these two
criteria, N. tasmanica ssp. gelatinifera var. beaugleholii is
misplaced in subgenus Hyella.
Oospores of all the specimens examined so far in subgenus
Hyella (with the exception of the specimens of N. tasmanica ssp.
gelatinifera var. beaugleholii and some specimens that key out to
N. myriotricha) have reticulate ornamentation on the oospores.
Oospores vary in size from ~180 mm long to over 300 mm long and
the reticulum varies in the number of meshes across the fossa, the
presence or absence of flanges, whether the ornamentation
extends up the flanges or not, and the depth and width of the
reticulum. These features can be hard to distinguish when viewed
under a light microscope, so oospores are not as useful for
discriminating among taxa in section Hyella as in other
sections. However, the range of vegetative variation is huge,
and specimens with zero furcations are not likely to be the same
species as specimens that are regularly 2  furcate, on the basis of

Figs 67–74. Figs 67, 68. Oospore of the isotype of p946, Nitella gelatinifera var. afoliata (R.D.Wood) J.C.van Raam, from Rocky River, Flinders Chase NP,
Kangaroo Island (R.D. Wood 60-12-27-1, 27.xii.1960, AD). Fig. 67. Whole oospore (263 mm  236 mm), showing reticulate ornamentation that continues half
way up the flanges on the 7 or 8 striae. Scale bar = 100 mm. Fig. 68. Detail of oospore wall, showing shallow reticulate ornamentation of ~5 meshes across the fossa.
Scale bar = 10 mm. Figs 69, 70. Oospore of the isotype specimen of p947, Nitella tasmanica subspecies gelatinifera var. beaugleholii R.D.Wood, from a Creek 2
miles SW of Snelling[ton] Beach, Kangaroo Island (R.D. Wood 60-12-29-11, 29.xii.1960, NY, AD). Fig. 69. Whole oospore (256 mm  244 mm), showing
apparently smooth ornamentation and low ridges on the 7 striae. Scale bar = 100 mm. Fig. 70. Detail of oospore wall, showing ornamentation of fine grains ~0.5 mm
in diameter, 8–10 per 10-mm distance, continuing up the ridges on the striae. Scale bar = 10 mm. Figs 71, 72. Oospore of the holotype of r112, Nitella reticulata
M.B.Williams, from a Dune pond, Maroubra Beach, New South Wales (M.B. Williams 341, 20.vii.1955, NSW). Fig. 71. Whole oospore (414 mm  328 mm),
showing finely reticulate oospore wall and 6 or 7 striae with reticulate ridges united in an apical crest. Scale bar = 200 mm. Fig. 72. Detail of oospore wall, showing
reticulate surface (~10 meshes across the fossa) and ornamented flanges. Scale bar = 20 mm. Figs 73, 74. Oospore of the holotype of r113, Nitella microteles
M.B.Williams, from a Creek 3 miles east of Nimmitabel, New South Wales (M.B. Williams 108, 7.ii.1955, NSW). Fig. 73. Whole oospore (427 mm  327 mm),
showing two layers, the outer (possibly consisting of the enveloping cells) roughened and the inner flatly granulate, and 6 or 7 striae united in an apical crest.
Scale bar = 100 mm. Fig. 74. Detail of oospore wall, portion of outer wall layer and inner bacculate layer. Scale bar = 20 mm. This oospore was not aggressively
cleaned because it was the only oospore remaining on the type specimen.
214 Australian Systematic Botany M. T. Casanova

75 76

77 78

79 80

81 82
An overview of Nitella in Australia Australian Systematic Botany 215

the evidence that the number of furcations is consistent for species
under culture (Williams 1959; Casanova et al. 2007).
Section Migularia
Wood (1965) placed section Migularia in subgenus Tieffallenia;
however, the presence of pluricellulate dactyls and evidence from
nucleotide sequences (Karol 2004) put it in subgenus Hyella. The
characteristic of central secondary rays is not constant or unique to
this group. The status of extra-Australian species N. imahorii and
N. struthioptila (included in this section by Wood 1965) is outside
the scope of this overview. Wood (1972) amalgamated Williams’
(1959) species (N. reticulata (r112, Figs 71, 72), N. microteles
(r113, Figs 73, 74), N. diffusa and N. cristata var. ambigua (p266,
Figs 75, 76)) with N. cristata (p815, Figs 77, 78) and doubtfully
included N. hookeri in the Australian flora. He also included
N. robertsonii and N. remota (r032, Figs 79, 80) in N. cristata.
Specimens of N. robertsonii from central Victoria and the
Northern Tablelands of New South Wales have been seen and
it clearly belongs in section Gioallenia. N. remota was not well
described and the type material was lost; however, Wood (1972)
saw additional material and included it as a variety of N. cristata.
N. remota is likely to be a species in its own right. Oospores of
species in section Migularia are invariably flanged, and
frequently ‘cristate’, with a crest formed from the united and
extended flanges at the apex of the oospore. The ornamentation
varies from smooth to pitted, reticulate to sparsely verrucate to
almost vermiferous (Figs 71–80); however, the status of the
varieties and forms of N. cristata recognised by Wood (1972)
needs further investigation.
Nitella hookeri was confirmed for Australia (Wood and Mason
1977) and a recent review of the species in New Zealand,
Kerguelen Island and Australia (Casanova et al. 2007) led to
the two Australian varieties being amalgamated into a single
species, N. arthroglochin (p604, Figs 81, 82). Van Raam (1995)
recorded N. microteles for Tasmania on the basis of specimens
that were morphologically similar to N. tricelluaris from New
Zealand; however, oospores of the specimens fitting that
description (p524, Figs 83, 84) are very different from those of
both N. microteles (Figs 73, 74) and N. tricellularis (Casanova
et al. 2007). Hotchkiss and Imahori (1988a) described N. woodii
(clearly related to N. cristata) and recorded the presence of
specialised vegetative reproductive organs. Such organs are
found on Nitella sp. aff. cristata in New Zealand where they
have been referred to as ‘opaque dactyls’ (Wood and Mason
1977), although the New Zealand taxon differs considerably from
most Australian material (Wood and Mason 1977). The
N. cristata complex in Australia and New Zealand is currently
under review.
Several other specimens exist that cannot be keyed out by
using current or historical treatments, and whose oospores differ

83 84

Figs 83, 84. Oospore of a specimen of p524, Nitella sp. 7 aff. tricellularis Nordstedt, from Lake St Clair, Tasmania
(T. Dugdale and S. Talbot 9a, 25.i.2004, MEL). Fig. 83. Whole oospore (460 mm  410 mm), showing 7 or 8 thick ridges on
the striae and verrucate ornamentation. Scale bar = 100 mm. Fig. 84. Detail of oospore wall, showing low verrucae joined in a
vermiferous or incomplete reticulate pattern, thick ridges on the striae, extending into smooth flanges near the apex or crest of
the oospore. Scale bar = 50 mm.

Figs 75–82. Figs 75, 76. Oospore of a specimen of p266, Nitella cristata var. ambigua A.Braun, from Barleyfields Lagoon near Uralla, New South Wales
(J. Divola, 29.ii.1996, MEL). Fig. 75. Whole oospore (327 mm  300 mm), showing the verrucate oospore wall and weak flanges on the 6 or 7 striae that are united
in an apical crest. Scale bar = 100 mm. Fig. 76. Detail of oospore, showing verrucate ornamentation. Scale bar = 20 mm. Figs 77, 78. Oospore of a specimen of p815,
Nitella cristata A.Braun, from a wetland near Interlaken, Tasmania (M.T. Casanova, 30.xi.2005, MEL). Fig. 77. Whole oospore (460 mm  382 mm including
flanges), showing flanges 50–70 mm long on the striae, crest at the apex of the oospore and relatively smooth fossa wall. Scale bar = 100 mm. Fig. 78. Detail of
oospore wall and flange, showing roughened granular surface. Scale bar = 20 mm. This is similar to the oospore of the type material of N. cristata. Figs 79, 80.
Oospore of a specimen of r032, Nitella remota A.Braun ex A.Braun, and Nordstedt from Bangham Forest Reserve, South Australia (R.J. Bates 26076, x.1991,
AD). Fig. 79. Whole oospore (375 mm  303 mm), showing roughened oospore wall and ~6 striae with huge flange united in an apical crest. Scale bar = 200 mm.
Fig. 80. Detail of oospore wall and flanges, showing roughened wall and unevenly verrucate flanges. Scale bar = 20 mm. Every attempt was made to clean this
oospore; however, because of the fragility of the flanges some debris remains on the fossa wall. Figs 81, 82. Oospore of a specimen of p604, Nitella arthroglochin
(A.Braun) M.T.Casanova, from Corralinga Creek, Fleurieu Peninsula, South Australia (E.N.S. Jackson 75, 23.iii.1959, AD). Fig. 81. Whole oospore
(490 mm  440 mm), showing short thick flanges on the 7 striae and verrucate–granulate fossa wall. Scale bar = 100 mm. Fig. 82. Detail of oospore wall,
showing low relief of the verrucate–granulate ornamentation that continues up the flanges. Scale bar = 20 mm.
216 Australian Systematic Botany
significantly from those illustrated here. At least 15 and possibly
20 new species are likely to exist in Australia.
Discussion
In all, 52 species of Nitella (89 taxa) are discussed or listed here
and oospores are illustrated for 41 of them. Coverage of species
across Australia is relatively patchy, because there are only few
collections from northern and arid Australia. Similarly, only a
proportion (30–50%) of the specimens deposited in herbaria are
sufficiently mature, or of sufficient quality, for thorough
comparison of taxonomically important characteristics.
However, there is evidence that there are more species of
Nitella in Australia than has been recognised in previous
treatments. More collections are needed, even from areas that
are relatively well collected, to determine the degree of vegetative
and oospore variation within each species, as well as to document
species that have not yet been described or discovered. An
examination of the database of specimens indicates that
November to December is a good time to collect fertile
charophytes in Tasmania, October to November in southern
mainland Australia, September to March in New South Wales
and southern Queensland, and the early dry season in Northern
Territory, northern Queensland and northern Western Australia.
The species concept put forward in this treatment is based on
several morphological criteria, all of which have been used by
authors in the past (Braun and Nordstedt 1882; Allen 1898;
Nordstedt 1918; Groves and Allen 1935; Williams 1959; van
Raam 1995; Sakayama et al. 2005). The important characters are
(1) sexual state (monoecy/dioecy), (2) oospore characters such as
ornamentation, expression of flanges and size, (3) the number of
furcations and the ratio of branchlet segment lengths and widths,
(4) end-cell and dactyl characters and (5) distribution, at least for
dioecious species (e.g. species that occur in tropical northern
Australia are unlikely to be the same species as those that occur in
Tasmania, similarly there is not much overlap between eastern
and western Australia). Characters such as the degree of
calcification and the amount of mucus on the fertile whorls
appear to vary in relation to environment. The presence of
calcification and the amount of mucus can be diagnostic;
however, their absence is not a diagnostic character. Not all
characters are useful for all subgenera or sections. In subgenus
Tieffallenia, oospore ornamentation is highly variable, whereas in
subgenus Hyella section Hyella, oospore ornamentation is almost
invariably reticulate. This does not mean that all members of
Hyella are the same species; rather, differences among the species
are expressed in the fine detail of oospore morphology as well as in
vegetative features, i.e. in the degree of furcation, the expression
of contracted fertile parts (heads or spikes) and the number
and shape of dactyl cells. The status of species in section
Decandollea is problematic. So far, all monoecious specimens
examined (i.e. N. hyalina) have fibrous oospores and, according
to Karol (2004), invariant rbcL nucleotide sequences. Despite
that, there is variation in their vegetative morphology, from tiny,
inconspicuous plants with few secondary branchlets to large,
mucus-covered plants with highly furcate branchlets and
abundant secondary branchlets. The occurrence of accessory
branchlets is diagnostic for this section, although the
expression of this feature varies within a plant for N. lhotzkyi
M. T. Casanova
and N. ungula. More collections are required to determine
whether there are natural limits to the variation in N. hyalina,
whether separate species can be distinguished, and the
significance of that variation in relation to environmental
influences.
In the present paper, the morphology of oospores of several
Australian type specimens and species is documented. For some
species, the type material was infertile, or if oospores existed they
had been removed. A rearrangement of some of the species in
sections is proposed, and several species are reinstated. The
illustrations of oospores will provide data for comparison with
the oospores obtained from further collections. Where oospores
of type material could not be obtained, it is possible that mistakes
in determinations have been made. However, this treatment
provides a starting point for confirmation of species-level
diversity in Australian Nitella and a framework for testing
relationships among species.
There is overwhelming evidence that, for the genus Nitella,
oospore characteristics are a good guide to species; however, the
amount of variation within each species is not usually well
documented. Where oospore ornamentation differs
significantly it is easy to recognise different species; however,
in subgenus Hyella at least, oospore variation is small and
vegetative variation is large. Early workers in charophyte
taxonomy had only a few specimens to work with and came
from a place where there were relatively few species of Nitella, so
they underestimated the number of species that existed. Later
workers, assuming they would find one or two new species only,
tended to underestimate the amount of work involved and the
diversity present, and rarely had sufficient time for a thorough
study. Wood’s (1972) treatment was extensive, although not
thorough, and was based on an erroneous species concept
(Proctor 1980; Casanova 2005a). As well, there are still too
few good, fertile specimens in Australian herbaria. The higher
the number of good collections, the better we will be able to
determine species-level differences, and the more reliable and
useful taxonomic treatments will be. Charophytes are of great use,
potentially, for the determination of environmental flows and
water regimes. The group is speciose, each species has a specific
set of measureable environmental requirements, they can be
identified to species from vegetative and oospore features, and
the oospores remain in the soil of wetlands and riparian zones
even when they are dry. We can already predict the water
requirements of a wetland on the basis of the charophytes that
are there (Casanova and Brock 1999; Casanova 2004), and need
only to document the range of species and their ecology, for
charophytes to be more useful to water managers.
Acknowledgements
Staff at the Royal Botanic Gardens Melbourne (National Herbarium of
Victoria) and the State Herbarium of South Australia have facilitated this
study in many ways. This study would have been much more difficult (if not
impossible) without the texts and translations of very early references given to
me by Beth Williams, thank you Beth. Discussions with and data provided by
Dr Ken Karol about his work on nucleotide sequences in Nitella have been
important in the development of this paper. Oospore micrographs were taken
at the Scanning Electron Microscope Unit of the University of Melbourne with
the assistance of Dr Simon Crawford. This work has been funded, in part, by
Australian Biological Resources Study Grants to Dr Gerry Kraft and
An overview of Nitella in Australia
M. T. Casanova in 2004, and to M. T. Casanova in 2007–2009. Drs John
Porter, Margaret Brock and Tony Dugdale, Keely Ough, Hans and Annie
Wapstra have provided beautiful specimens of Nitella that have greatly
enhanced this research.
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Manuscript received 25 August 2008, accepted 12 March 2009