Table Of ContentA new fern species for Queensland: Diplazium
squamuligerum (Rosenst.) Parris (Woodsiaceae)
Daniel J. Ohlsen1 & Ashley R. Field2
Summary
D.J.Ohlsen & A.R.Field (2013). A new fern species for Queensland: Diplazium squamuligerum
(Rosenst.) Parris (Woodsiaceae). Austrobaileya 9(1): 114-125. The taxonomic status of an unknown
fern species from the Atherton Tableland, north-east Queensland, hitherto attributed to Asplenium
L., was investigated. Phylogenetic analysis of trnL-F and rbcL chloroplast DNA sequences supported
classification in Diplazium Sw., a finding supported by closer examination of scale features.
Inspection of Diplazium type material determined that the Australian material belongs to Diplazium
squamuligerum (Rosenst.) Parris, a species previously described from Papua New Guinea. A thorough
description of this species and an amended key to the Diplazium species of Australia are provided.
This study highlights the value of molecular study and close inspection of scale features for fern
identification. Taxonomic revision in Diplazium is also discussed in light of the findings presented.
Key Words: fern, Athyriaceae, Woodsiaceae, Diplazium, Diplazium squamuligerum, Australia flora,
Queensland flora. Wet Tropics bioregion, new species record, taxonomy
'D.J.Ohlsen, School of Botany, The University of Melbourne, Parkville, Victoria 3010, Australia.
Email: [email protected]
2A.R.Field, Queensland Herbarium, Department of Science, Information Technology, Innovation and
the Arts, Brisbane Botanic Gardens, Toowong, Queensland 4066, Australia &Australian Tropical
Herbarium, James Cook University, Cairns Campus, Smithfield, Queensland 4878, Australia. Email:
A shley. F ield@science. dsitia. qld. gov. au
Introduction
The Wet Tropics of north-east Queensland, known from only three sites in Australia, was
from Ingham north to Cooktown, contains described (as Polypodium sinuosa Wall, ex
the highest fern species diversity in Australia. Hook.) from Indonesia in 1863, but was not
257 leptosporangiate fern species occur in collected in Australia until 1985 when it was
the region, which is 66% of the total for found on Moa Island, Torres Strait. It was
Australia (McCarthy 1998). While 37 of these later collected on Cape York in 1987, and was
are considered to be endemic to this region, formally recognised as an Australian species
the vast majority are widespread being (Andrews 1990). Entirely new taxa, endemic
shared with neighbouring tropical islands, to north-east Queensland have also gone
particularly New Guinea which shares 56% unnoticed until relatively recently. These
of tropical Australian leptosporangiate fern include Antrophyum jagoanum D.L.Jones
species (McCarthy 1998). Several such species & Bostock, Diplazium bostockii D.L.Jones,
are extremely rare in Australia, known only and Lastreopsis windsorense D.L. Jones
from one or a few small populations. Notable (McCarthy 1998). However, the most recent
examples arq AspleniumpeUucidiim Lam. and new fern species confirmed for Australia is
Hymenasplenium unilaterale (Lam.) Hayata, Oleandra musifolium Blume. In contrast to
both restricted to a single known population the previous cases, populations of this species
in Australia (Brownsey 1998). have been known in Australia for over 60
years, but under the misapplied name of O.
The rarity of some taxa has delayed their
neriiformis Cav. (Hovenkamp & Ho 2012).
discovery until relatively recently in Australia.
Lecanopteris sinuosa (Wall, ex Hook.) Copel., Four fern specimens collected from two
areas on the Atherton tableland, in north-east
Queensland, hitherto identified as Asplenium
Accepted for publication 29 July 2013 sp. indet. (A. sp. ‘RE.I. Road’ in BRI)
Ohlsen & Field, Diplazium squamuligerum 115
(Aspleniaceae) on herbarium specimens, also features such as differences in the arrangement
appear to represent a new taxon for Australia of vascular bundles in the stipe, and by the
(Fig. 1). This taxon is clearly morphologically possession of non-clathrate scales (Kato &
distinct from the 30 Australian Asplenium Kramer 1990), which are often only obscurely
species treated by Brownsey (1998). It was differentiated from the clathrate scales
first collected in 1983: once from the North observed in Aspleniaceae. Two Woodsiaceae
Johnstone Logging Area (Lockyer s.n.), which genera occur in north-east Queensland:
is thought to be from a population on the west Deparia Hook. & Grev. and Diplazium Sw.
side of Topaz National Park (N.P.) (B.Gray sensu lato (including Callipteris Bory).
pers. comm.), and twice from Maalan, 20
This study sequenced two chloroplast
km further south. The fourth and most recent
regions, rbcL and trnL-L, of a fresh collection
collection, in 2005 (Sankowsky & Sankowsky
of Asplenium sp. ‘RE.I. Road’, in order to
2637), was from near the end of P.E.I. Road
determine this taxon’s affinity amongst the
in Topaz N.P, within one km of the suspected
ferns, and to resolve its taxonomic status.
location of the Lockyer collection.
Materials and methods
Determining the taxonomic status of
taxa in Aspleniaceae is often fraught with Collections of Asplenium and Diplazium
difficulty. Aspleniaceae is the largest fern in Australian herbaria, and selected type
family with over 700 species occurring material from extra-Australian herbaria were
worldwide (Kramer & Viane 1990; Smith examined. One specimen of Asplenium sp.
et al. 2006). This makes verification of ‘P.E.I. Road’ was collected (iOhlsen 461, BRI,
identity or novelty challenging for unresolved MELU) in the vicinity of the most recent P.E.I.
Australian collections; the putative taxon Road collection {Sankowsky & Sankowsky
must be compared with a large number of 2637, BRI).
type specimens from overseas. In addition to
DNA isolation, amplification and
the large species diversity in Aspleniaceae,
sequencing.
hybridisation and polyploidy are frequently
DNA was extracted from 20 mg of silica gel-
encountered (see Lovis 1977). Hybrids and
dried leaf tissue. Leaf tissue was ground using
polyploids are often morphologically distinct
a mortar and pestle with the aid of acid washed
from parental lineages (Kramer & Viane
grinding sand (Ajax Finechem, Australia).
1990), obscuring their origins and giving the
DNA was isolated from ground samples
impression of completely new lineages.
using a DNeasy Plant Mini Kit (QIAGEN,
While identification within Aspleniaceae Germany), following the manufacturer’s
is often difficult, members of the family instructions. DNA was eluted in 100 pL of the
are generally easy to distinguish from supplied elution buffer.
other families. Those in Aspleniaceae are
The chloroplast DNA markers rbcL and
distinguished by linear sori, clathrate scales
trnL-L were sequenced. These regions were
on the stipes, and stipes which have two
chosen because 1) both are routinely used
C-shaped vascular bundles at the base which
in fern systematics, enabling comparison
unite apically up the stipe to form a ‘butterfly-
of sequences obtained from this species to
like’ shape (Kato & Kramer 1990). Species
those of many other species, and 2) effective
of Aspleniaceae are morphologically most
phylogenetic placement (at least to genus
similar to some members of the Woodsiaceae
level) can be achieved with both regions.
sensu Smith et al. (2006) (=Athyriaceae
rbcL is a gene enabling comparison across
sensu McCarthy (1998)), which also possess
multiple plant lineages, and the trnL-L region
elongate indusiate sori. The majority of
contains an intron and intergenic spacer,
these Woodsiaceae species are most easily
which have faster mutation rates, enabling
distinguished from Aspleniaceae by being
differentiation between closely related species
large terrestrial plants; however, they are
and populations.
consistently distinguished by more subtle
116 Austrobaileya 9(1): 114-125(2013)
Fig. 1. Diplazium squamuligerum (syn. Asplenium sp. RE.I. Road) from Topaz National Park, Atherton Tableland,
Queensland (Ohlsen 461 et al. [BRI, MELU]). (a) a mature frond with elongate sori, (b) a fully developed scale from
the base of the stipe, (c) D. squamuligerum in habitat (two plants are shown: top left, and bottom beside the creek
pool), (d) detail of the rachis and stipe wing, and the protuberances, some with scales developing upon them.
Chloroplast DNA markers were TTA CTA GCT TCA CG-3’) (Schuettpelz &
amplified by Polymerase Chain Reaction Pryer 2007). The trnL intron, trnL 3’-exon
(PCR), performed on a MyCycler thermal and trnL-F intergenic spacer were amplified
cycler (Bio-Rad, USA). Reaction mixtures using the primers F (5’-ATT TGA ACT GGT
comprised 5 pL of 5x MyTaq Reaction Buffer GAC ACG AG-3’) (Taberlet et al. 1991) and
containing 5mM of each dNTP and 15 mM Fernl (5’-GGC AGC CCC CAR ATT CAG
MgCl2 (Bioline, Australia), 50 pg BSA GGR AAC C-3’) (Trewick et al. 2002). PCR
(Thermo Fisher Scientific, Australia), 0.125 thermocycling conditions involved an initial
pL (0.625 Units) MyTaq DNA Polymerase denaturation step of 95°C for 1 minute,
(Bioline, Australia), 10 pmol of each primer, followed by 33 cycles of 95°C for 1 minute,
2.0 pL of extracted DNA, and distilled water 55°C for 1 minute, and 65°C for 4 minutes,
added to make a total volume of 25 pL. No and a final extension time of 65°C for 5
amplification occurred without the addition minutes. DNA concentrations were quantified
of BSA. The rbcL gene was amplified using by electrophoresis against Hyperladder I
the primers ESRBCL1F (5’-ATG TCA CCA and EasyLadder I (Bioline, Australia) and
CAA ACG GAG ACT AAA GC-3’) and PCR products were purified using illustra
ESRBCL1361R (5’-TCA GGA CTC CAC ExoSTAR 1-step enzymatic purification (GE
Ohlsen & Field, Diplazium squamuligerum 117
Healthcare Life Sciences, UK). Purified PCR Asplenium marinum L. were also included as
products were then sent to the Australian outgroups.
Genome Research Facility (AGRF),
Species identification.
Melbourne Branch, where sequencing
Online images of Diplazium types (B,
reactions, and capillary separation, using the
MICH) were examined, to determine whether
96-capillary analyser AB 3730x/ sequencing
this taxon could be assigned to an existing
platform, were performed.
Diplazium species, or if it was an undescribed
Sequence editing, alignment, and analysis. species.
Sequences were edited using Sequencher
Common Abbreviations used in Specimen
v. 3.0 (Gene Codes Corporation, Ann
Citations
Arbor, MI, USA) and the NCBI nucleotide-
nucleotide BLAST (blastn) tool (Altschul LA (Logging Area); NP (National Park).
et al. 1997) was then used to evaluate the
Results
similarity of both rbcL and trnL-L sequences
to existing sequences in GenBank. After the Searches using the BLAST tool of both
search using the BLAST tool was performed, trnL-L and rbcL sequences, determined that
available rbcL and trnL-L sequences of sequences from the P.E.I. Road specimen had
Diplazium species from GenBank were the greatest similarity to species of Diplazium
gathered (Appendix 1) and aligned manually in the family Woodsiaceae. 37 Diplazium
in Se-Al Sequence Alignment Editor v. 2.0all rbcL sequences were used to produce an
(Rambaut 2002). Outgroup sequences were alignment 1188 base pairs long, which
also gathered from GenBank (Appendix 1). contained 110 parsimonious characters.
Two most parsimonious trees were obtained
Parsimony analyses were run in PAUP*
(length=398 steps, CI= 0.626) one of which is
v 4.0(310 (Swofford 2000). Gap characters
shown (Fig. 2a). This had a topology identical
in the alignment were treated as a fifth
to the strict consensus tree, except for the
character state. For multiple base indels,
placement of D. doderleinii (Luerss.) Mak.,
characters were excluded from analyses so
which in the consensus tree is placed in a
that indels were represented only by a single
polytomy with the clade of D. amamianum
gap character, when variability did not occur
Tagawa + D. hachijoense Nakai and the clade
within indels. Question marks, the character
of D. taiwanense Tagawa + D. virescens
recognised in PAUP for missing data, were
Kunze. 10 Diplazium trnL-L sequences were
used to fill gaps, where needed, when indels
used to produce an alignment of 753 included
fell across otherwise variable regions of the
base pairs, which contained 139 parsimonious
alignment. A heuristic tree search was used,
characters. A single most parsimonious tree
with delayed character-state optimization
was obtained (length=302 steps, 0=0.831)
(DELTRAN) and starting trees obtained
(Fig. 2b). Maximum parsimony analyses of
by a closest addition sequence followed by
rbcL (Fig. 2a) and trnL-L (Fig. 2b) shows that
tree bisection-reconnection (TBR) branch
the P.E.I. Road taxon is nested well within
swapping. Bootstrap support for tree nodes
Diplazium sensu lato. Of all Diplazium
was determined for each analysis, with 100
species currently sequenced for rbcL, it is
replicates. Separate parsimony analyses were
most closely related to D. proliferum (Lam.)
performed for each chloroplast DNA region.
Thouars. (=Callipteris prolifera (Lam.) Bory
Athyrium felix-femina (L.) Roth was chosen as
sensu Jones [1998]) with high bootstrap
an outgroup, based on the sister relationship
support (98%). It differs from D. proliferum
of Athyrium to Diplazium in past molecular
by 12 base pairs in rbcL.
phylogenies (Sano et al. 2000; Wang et al.
2003; Schuettpelz & Pryer 2007) and for Inspection of Diplazium type material
the rbcL analysis, in which it is possible revealed that the P.E.I. Road taxon matches
to align distantly related taxa, the more Asplenium varians var. squamuligerum
distantly related Blechnum occidental L. and Rosenst., from New Guinea, now treated as
118 Austrobaileya 9(1): 114—125(2013)
6 I Diplazium amamianum
99 ' Diplazium hachijoense
1 2
— Diplazium doederleinii
Diplazium taiwanense
- Diplazium virescens var. conterminum
- Diplazium virescens
2
|— Diplazium crassiusculum
Diplazium donianum var. aphanoneuron
99 1
^ Diplazium lobatum
1
Diplazium dilatatum
a
87 Diplazium hayatamae
Diplazium deciduum
1
Diplazium fauriei
U- Diplazium mettenianum
91
Diplazium griffithii
— Diplazium pullingeri
L-=— Diplazium kawakamii
8
- Diplazium centripetale
—— Diplazium incomptum
4
76 -9 | -D12ip lazium longicarpum
97 1---Diplazium subserratum
93 Diplazium proliferum
Diplazium squamuligerum
98
p
5 Diplazium chinense
L_6
87 Diplazium subtripinnatum
4 8 (cid:9632) Diplazium esculentum
10
— Diplazium nipponicum
(cid:9632) Diplazium sibiricum var. glabrum
Diplazium squamigerum
73 9 Diplazium bombanasae
— Diplazium cristatum
Diplazium lonchophyllum
95
15 - Diplazium plantaginifolium
13 | Diplazium okudairae
' Diplazium wichurae
93 —— Diplazium pinfaense
18 - Athyrium felix-femina
70
43
-Blechnum occidentale
-Asplenium marinum
(a)
Fig.2a. Parsimony analyses of rbcL showing the phylogenetic position of Australian Diplazium squamuligerum.
Branch lengths are given above braches and bootstrap support values greater than 70% are given below branches.
Ohlsen & Field, Diplazium squamuligerum 119
12
(cid:9632) Diplazium pinfaense
24
100 r^- Diplazium wichurae
11
100
L^- Diplazium heterocarpum
Diplazium hainanense
7
99
Diplazium hachijoense
7
89
16 Diplazium esculentum
15 Diplazium nipponicum
15.
90 17 Diplazium mettenianum
46
Diplazium squamuligerum
59
Diplazium ovatum
Athyrium filix-femina
(b)
Fig.2b. Parsimony analyses of trnL-F. Branch lengths are given above braches and bootstrap support values greater
than 70% are given below branches.
120 Austrobaileya 9(1): 114—125(2013)
Diplazium squamuligerum (Rosenst.) Parris. and protuberances; protuberances eventually
The original diagnosis treated it as a variety bearing light brown scales to 1 mm long,
of the unrelated African Asplenium varians with darker brown veins and bifid marginal
Wall, ex Hook. & Grev. (Rosenstock 1913). teeth. Lamina bipinnate-tripinnatifid, 3-10
The diagnosis is very brief, outlining only cm long, 1.5-4 cm wide, dark green above,
some immediately obvious features which paler below, occasionally proliferous from the
distinguish D. squamuligerum from A. rachis. Primary pinnae 5-25 mm long, 2-10
varians. Many of the morphological features mm wide, most with an acroscopic secondary
that are quite distinctive are not mentioned. pinna to 5 mm long and an elliptic apical
The following description describes segment 5-10 mm long; margins serrate;
Diplazium squamuligerum in greater veins free. Sori to 4 mm, elongate along most
detail, including several distinctive features veins; indusium entire, narrow, light brown
overlooked by the earlier diagnosis. (Fig. la, b, d)
Taxonomy Specimens examined: Australia: Queensland. Cook
district: North Johnstone LA, July 1983, Lockyer s.n.
Diplazium squamuligerum (Rosenst.) (CANB); P.E.I. Road, Topaz NP, July 2005, Sankowsky
Parris, Kew Bull. 41 (1): 69 (1986); Asplenium & Sankowsky 2637 (BRI); North Johnstone LA, Jan
2013, Ohlsen 461 etal. (BRI, MELU); Portion 545 Parish
varians var. squamuligerum Rosenst., Repert.
of Dirran, Malian Road, July 1983, Gray 3133 (CNS).
Spec. Nov. Regni Veg. 12: 528 (1913); A.
squamuligerum (Rosenst.) Hieron, Hedwigia Distribution and habitat: In Australia
61:5 (1919).Type: PapuaNew Guinea: Morobe Diplazium squamuligerum is known only
Province: Sattelberg Hinterland, 1400-1500 from two areas on the Atherton Tableland,
m [I.C.]Keysser 228, April 1913 (holo: S, S-P- north-east Queensland; however, it also
7625, online image!; iso: UC 378428, online occurs in Papua New Guinea. It grows in
image!; MICH 1190092, online image!). mixed mesophyll rainforest, on metamorphic
rocks or between tree roots in volcanic soil,
Rhizome erect, to 1.5 cm tall, scaly; scales to
in steep embankments of small, slow-flowing
1 mm long, dark brown with toothed margins.
creeks (Fig. lc).
Fronds arcuate, 5-15 cm long, 1.5-4 cm wide.
Stipe 0.5-5 cm long, dark green, enveloped by Notes: Wings of the axes often become
green tissue which elongates to produce wings obscure when dried, but are obvious in fresh
material.
Key to Australian species of Diplazium (naturalised taxa indicated *)
1 Veins anastomosing.2
.
1 Veins free.5
2 Lamina simple.D. cordifolium
2. Lamina pinnate to tripinnate.3
3 Lamina pinnate, rachis proliferous.D. proliferum (syn. Callipteris prolifera)
3. Lamina bipinnate or tripinnate, rachis not proliferous.4
4 Secondary pinnae 1-1.5 cm wide, margins incised more than half-way to
the pinnule midvein.D. dietrichianum
.
4 Secondary pinnae 1.5-2.5 cm wide, very shallowly lobed.*D. esculentum
5 Lamina pinnate.6
5. Lamina bipinnate or more divided.7
6 Lamina apex a single pinna similar to lateral pinnae.D. pallidum
6. Lamina apex formed by reduction of lateral pinnae.D. dameriae
Ohlsen & Field, Diplazium squamuligerum 121
7 Fertile laminae < 250 mm long, stipe and rachises winged, wing extending
to form scale bearing protuberances, pinnae with a basal acroscopic
secondary pinna or lobe.D. squamuligerum
7. Fertile Laminae > 250 mm, stipe, rachis and pinnae not as above.8
8 Lamina bipinnate.9
8. Lamina tripinnate or more divided.10
9 Basal lobes of secondary pinnae longest, abaxial surface of pinnae
glabrous.D. dietrichianum
9. Basal lobes of secondary pinnae similar to or smaller than the rest, abaxial
surface bearing red glandular hairs.D. dilatatum
10 Lamina pale green, pinnules less than 5 mm long and 3 mm wide.D. assimile
10. Lamina dark green, pinnules greater than 10 mm long and 4 mm wide.11
11 Apex of pinnules acute to caudate.D. bostockii
11. Apex of pinnules obtuse.12
12 Pinnules of fertile lamina dissected < one third of the distance to the
midvein, pinnules decurrent on basiscopic margin.D. australe
12. Pinnules of fertile lamina dissected > half-way to the midvein, pinnules
not decurrent.D. queenslandicum
Discussion Prior to this study, 12 species of Diplazium
were known in Australia (one species, D.
The identification of the PE.I. Road taxon as
esculentum is naturalised). All of these
a species of Diplazium by chloroplast DNA
Diplazium species are large terrestrials, with
sequence is also supported by morphology.
fronds over 50 cm long. The small size of D.
Populations of D. squamuligerum in Australia
squamuligerum is therefore unique amongst
have elongate sori and scales that are borne
Australian Diplazium, and unusual amongst
on small protuberances: both features typical
all Diplazium (Kato & Kramer 1990). Its
of Diplazium (Jones 1998). The scales are
position in the chloroplast phylogeny, as sister
characterised by bifid teeth on a darkened
to D. proliferum is supported by its possession
margin and conform to the ‘Callipteris type’
of the ‘Callipteris type’ scales, that are also
(Fig. lb). This scale type is known only from
possessed by D. proliferum (Jones 1998).
the segregate genus Callipteris and a few
Both D. squamuligerum and D. proliferum
other Diplazium species (Pachebo & Moran
are also proliferous from the rachis, albeit
1999; Pachebo & Moran 2003).
rarely so in D. squamuligerum. Otherwise
These specimens were repeatedly these species are quite dissimilar in general
misidentified as a species of Asplenium, appearance. D. proliferum is a very large fern
rather than being correctly identified as a with pinnate fronds to over 2 metres in length
Diplazium. Hence this study is another case whereas the fronds of D. squamuligerum are
which highlights the value of DNA sequences at least 2-pinnate and less than 15 cm long.
for phylogenetic placement and identification D. squamuligerum also has free venation and
when morphology may be misleading winged axes (Fig. Id); the latter feature is not
(Gastony et al. 2001). It also begs caution in shared with any other Australian Diplazium
identifying species by overall morphology species.
without close inspection of more subtle but
The current phylogeny of Diplazium is
taxonomically informative features such as
limited in the number of species sampled.
scale characters.
Chloroplast DNA sequences of only 38
species are publicly available, all of which
122 Austrobaileya 9(1): 114—125(2013)
are incorporated in the phylogenies presented adjacent to one of approximately ten locations
here. This is a low proportion of the 400 species of Dryopteris wattsii M.McKeown, Sundue &
of Diplazium in total (Kato & Kramer 1990). Barrington. In addition to these exceptionally
So it is highly likely that another unsampled rare species, other uncommon fern species
species of Diplazium is more closely related such as Dicksonia herbertii W.Hill, Oleandra
to D. squamuligerum than D. proliferum, musifolia, and Pteridoblechnum acuminatum
especially given the large number of base (C.T.White & Goy) Hennipman and P.
pair differences between the two species. neglectum (F.M.Bailey) Hennipman also occur
Several Diplazium species, which occur in at or near the D. squamuligerum population
New Guinea, or possess the ‘Callipteris type’ localities. Likewise, the Maalan population is
scales, are yet to be sequenced, and may be within 7 km of one of only three populations
more closely related. of Asplenium normale D.Don in Australia.
The occurrences of multiple, very localised
Diplazium has received little attention
fern species near sites of D. squamuligerum
from molecular study, as is demonstrated by
highlights the high conservation priority that
the low number of sequences available on
should be given to these areas, especially
GenBank. Further molecular work, involving
considering the high diversity of seed plant
a more comprehensive sampling of species,
species also present. It also suggests that a
is required in Diplazium not only to gain
number of equally localised species may have
a better understanding of the systematics
been lost due to vegetation clearing around
and biogeography of this genus, but also to
these populations.
aid taxonomic revision, particularly at the
generic level. It has been suggested, based on Diplazium squamuligerum is currently
a recent molecular study using a rather small one of the most threatened fern species in
number of species (Wang et al. 2003), that Australia. While it is reserved in Topaz NP,
several genera which had been segregated all populations of this species are small in
from Diplazium be placed back into that area and size and are particularly vulnerable
genus. This included Callipteris, a genus still to stochastic events. Periods of particularly
accepted in Australia (Jones 1998). However, high rainfall, with increased runoff due to
revision of Diplazium cannot be completed surrounding cleared land, can swell the small
until more species sampling is undertaken. creeks along which it occurs. Consequently
The close relationship of D. squamuligerum plants face the danger of being uprooted from
to D. proliferum suggests that the ‘Callipteris the creek embankments. This species would
type’ scale may define a monophyletic group. also be extremely susceptible to desiccation.
It has been suggested that this feature may It occurs in very high rainfall areas, and along
be used to recircumscribe Callipteris, to small slow flowing creeks, that provide the
include the few Diplazium species, such as constant soil moisture and high humidity that
D. squamuligerum, that have this scale type, is probably required by the plant. Extended
and also have free venation (Pachebo & dry periods may reduce such small creeks and
Moran 2003). However, the monophyly of limit water access to plants. Such stochastic
all species possessing this scale type needs weather events are expected to become more
to be demonstrated before any revision is extreme in the future with ongoing climate
undertaken (Pachebo & Moran 2003). change (Abbs et al. 2006; Mpelasoka et al.
2008; Walsh et al. 2004).
Identification of this species as D.
squamuligerum provides an additional The one collection from Papua New
example of a fern species that is localised Guinea that exists in Australian herbaria
in a few sites on the Atherton Tableland. (Brass 25731 [CANB]) was collected from
Populations at Topaz NP are within close Mt Pabinama, Normanby Island, which is
proximity to other localised fern species. They 550 km south-east from the type locality,
are within 3 km of the only known locality of and suggests that D. squamuligerum is
Hymenasplenium unilateral in Australia, and more widespread in Papua New Guinea.
Ohlsen & Field, Diplazium squamuligerum 123
A conservation status of Vulnerable is Gastony, G.J. & Johnson, W.P (2001). Phylogenetic
recommended for D. squamuligerum placements of Loxoscaphe thecifera
(Aspleniaceae) and Actiniopteris radiata
in accordance with the distribution and
(Pteridaceae) based on analysis of rbcL
abundance characteristics of other taxa listed
nucleotide sequences. American Fern Journal
as Vulnerable under the Nature Conservation 91: 197-213.
Act 1992 (Queensland), the Environment
Hovenkamp, P.H. & Ho, B-C. (2012). A revision of the
Protection and Biodiversity Conservation Act
fern genus Oleandra (Oleandraceae) in Asia.
1999 (Australian Commonwealth) and also PhytoKeys 11: 1-37.
in line with the criteria of the International
Jones, D.L. (1998). Athyriaceae. In P.M. McCarthy
Union for Conservation of Nature (IUCN). A (ed.). Flora of Australia Ferns, Gymnosperms
conservation advice report is currently being and Allied Groups 48: 418-429. ABRS/CSIRO:
prepared for D. squamuligerum in Australia. Melbourne.
Kato, M. & Kramer, K.U. (1990). Subfamily
Acknowledgements
Athyrioideae. In K.U. Kramer & PS. Green
We would like to express our gratitude (eds.). The families and genera of vascular
plants Pteridophytes and gymnosperms 1: 130-
to Bruce Gray, who assisted the recent
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collection of Diplazium squamuligerum. All
sequencing was performed in the Cookson Kramer, K.U. & Viane, R. (1990). Aspleniaceae. In
K.U. Kramer & PS. Green (eds.). The families
Laboratory, School of Botany, The University
and genera of vascular plants Pteridophytes
of Melbourne, and was funded by a BushBlitz and gymnosperms 1: 52-56. Springer-Verlag:
Research Grant. Much appreciated advice in Berlin.
sequencing D. squamuligerum was provided
Lovis, J.D. (1977). Evolutionary patterns and processes
by Lara Shepherd and Leon Perrie. We are also in ferns. Advances in Botanical Research 4:
thankful to Leon Perrie, Nada Sankowsky, 223-415.
Darren Crayn, and to the anonymous
McCarthy, PM. (1998). Flora of Australia Ferns,
reviewers for their helpful comments on the Gymnosperms and Allied Groups 48. ABRS/
manuscript. CSIRO: Melbourne.
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