136
Chapter 5
Phenetic and cladistic analyses of Australian
Bulbostylis Kuuth
Introduction
This chapter focuses on the limits of Australian taxa within the genus Bulbostylis
Kunth nom. cons.
Bulbostylis is a large and mostly pantropical genus comprising around c. 200
species, with the centre of diversity in the African continent (Haines and Lye 1983;
Goetghebeur and Coudijzer 1985); the generic rank is now widely accepted (see
Chapter 1 for historical perspective). Currently, there are six species of Bulbostylis
recognised that occur in Australia; B. burbidgeae K.L. Wilson, B. turbinata
S.T.Blake, and B. pyriformis S.T.Blake are endemic, while B. barbata (Rottb.)
C.B.Clarke and B. densa (Wall.) Hand.-Mazz. have a widespread distribution.
Bulbostylis humilis (Kunth) C.B.Clarke (sYn B. striatella C.B.Clarke; see World
Checklist of Mononcotyledons) is a relatively recent introduction to Australia, and
known in the Arrnidale - Glenn Innes area of New South Wales.
Intermediate morphology due to clinal variation or hybridisation is COlnn10n for
some species within the genus e.g. B. schoeinoides complex (Gordon-Gray 1988),
B. hispidula complex (Haines and Lye 1983; Lye 1995), B. densa complex (Haines
137
and Lye 1983; Gordon-Gray 1995; Lye 1996), creating uncertainties in species lin1its
and confusion for the identification of specimens.
Bulbostylis densa (as Bulbostylis capillaris var. trifida (Kunth) C.B.Clarke) was
separated from B. capillaris (L.) Kunth ex C.B.Clarke due to morphological
differences between the American and Asian/Pacific specimens. Blake (1941),
however, commented that the Australian material that Clarke assigned with the
Indian B. capillaris var. trifida, was indistinguishable from the American
B. capillaris var. capillaris. Since then, other species and subspecies have been
separated from and within B. densa (Haines and Lye 1983), e.g. B. pusilla (Hochst.
ex A.Rich) C.B.Clarke and B. densa subsp. afromontana (Lye) R.W.Haines were
split to delimit the African variation. Collections from Kwazulu-Natal, South Africa,
which have nuts at the plant base that are distinctly larger than the aerial counterparts
(i.e. the plants are amphicarpic), do not fit the current description of B. densa or
similar species (Haines and Lye 1983; Gordon-Gray 1995). In addition, two
collections, one from China and the other from Queensland, Australia (E. sp. afC
densa 1), have morphology similar to B. densa s.s., but with different nut
characteristics; the Queensland collection does not fit descriptions of any accepted
Australian species. It is necessary to compare Australian (including Paci fic), African,
and Asian material of B. densa, \vith the American material of B. capillaris, to assess
species limits of B. densa in Australia.
Bulbostylis barbata is another cosmopolitan species where the morphological
variation needs to be compared globally. Bulbostylis barbata subsp. pulchella
(Thwaites) T.Koyama was separated to demarcate the variation in plants from
southern India and Indo-China generally.
A putative new species of Bulbostylis (B. sp. aff. barbata), with distinctly
piliferous glume margins, and hairy sheaths and leaf blades, was collected fronl the
Kakadu National Park in the Northern Territory. These plants grow interspersed with
plants of B. barbata and have a similar growth habit and inflorescence-synflorescence structure. Although the nuts of the putative new species are similar to
those seen in B. barbata there are apparent differences between the collections of
both taxa. The species limits for B. sp. aff. barbata required testing before a new
species can be defined.
Blake (1941) described B. pyriformis S.T.Blake and commented on the style base
that mayor may not persist on the nut. This feature of the style base has been a cause
of great confusion in the placement of B. hispidula (Vahl) R.W.Haines, as is evident
fron1 the many nomenclatural synonyms (World Checklist of Monocotyledons 2004);
the persistence of the style base on the nut was a key character in assigning taxa to
Bulbostylis, with non-persistence characteristic for Fimbristylis. Embryo morphology
provided the evidence that united specimens of B. hispidula with Bulbostylis through
sharing the Bulbostylis-type embryo (Van der Veken 1965). Some specimens of
Bulbostylis turbinata also show variability in the persistence of the style base on the
nut. The Australian B. pyriformis shares similar characteristics in general plant
morphology and nut micromorphology with taxa of the B. hispidula complex. It was
necessary to compare the Australian material with SOlue of the African B. hispidula
subspecies to test the species limits.
\Nilson (1980) described Bulbostylis burbidgeae as a new species endeluic to
Australia. Two separate collections, P.K. Latz 11364 (NSW 452329) (B. sp. aff.
puberula) and C.R. Dunlop (DNA 14302, NSW) (B. sp. aff. burbidgeae)
superficially resemble B. burbidgeae, but have nuts that are quite distinct from each
139
other and from those typical of B. burbidgeae. Both collections are fron1 the
Northern Territory and therefore outside the known Western Australian distribution
range for B. burbidgeae. The collections of B. sp. aff. puberula and B. sp. aff.
burbidgeae need to be compared to the other Australian species and to B. puberula
(Poir.) C.B.Clarke, to assess the species boundaries.
As a recent introduction to Australia (Wilson 1993), the African species
Bulbostylis humilis, needs to be included to assess the extended range of distribution.
The limits of all species and putative species of Australian Bulbostylis were tested
using phenetic analysis. The relationships of those species and of the genus as
sarrlpled here were then assessed for monophyly in the cladistic analysis.
Materials and methods
Taxa
All Australian taxa currently recognised as Bulbostylis, i.e. B. barbata, B.densa,
B.turbinata, B. pyriformis and B. burbidgeae (Wilson 1980, 1993), formed the basis
of the phenetic study. Putative new species, i.e. B. sp. aff. barbata,
B. sp. aff. burbidgeae, B. sp. aff. puberula and B. sp. aff. densa l, were included for
species level assessment (Table 5.1). Overseas specimens for the widespread
Bulbostylis barbata and B. densa were included with Australian material in the
phenetic analyses to define the species on a global level. Representative specilnens of
B. capillaris (TYPE species for the genus) B. humilis, B. puberula, and samples from
the B. hispidula complex (B. hispidula (Yahl) R.W.Haines subsp. pyr{formis
Table 5.1 Specimens sampled as the focus group in the phenetic assessment of Australian
Bulbostylis. The 'OTU' corresponds to the label used in phenetic analyses. States are given for
Australian collections and the Country of origin for all other samples collected overseas. N.T. =
Northern Territory, W.A. = Western Australia, S.A. = South Australia, Qld = Queensland, N.S.W. =
New South Wales, P.N.G = Papua New Guinea. See Appendix 1 for specimen details.
Species
Bulbostylis
sp. aff.
barbata
Bulbostylis
barbata
Bulbostylis
turbinata
Bu/bostylis
densa
OTU
State or
Country
baffba1
N.T.
baffba2
N.T.
baffba3
N.T.
baffba4
N.T.
baffba5
N.T.
baffba6
N.T.
baffba7
N.T.
bba1
Qld
bba2
N.T.
bba3
W.A.
bba4
N.S.W.
bba5
N.T.
bba6
N.S.W.
bba7
N.T.
bba8
W.A.
bba9
N.T.
bba10
N.T.
bba11
W.A.
bba12
Qld
bba13
Singapore
bba14
USA
Kenya
bba15
India
bba16
America
bba17
India
bba18
bba19 South Africa
Thailand
bba20
btl
S.A.
bt2
W.A.
bt3
N.T.
bt4
N.T.
bt5
W.A.
bt6
N.T.
bt7
W.A.
bt8
W.A.
bt9
Qld
bt10
N.T.
bde1
N.S.W.
bde2
N.S.W.
bde3
Qld
bde4
Qld
bde5
Qld
Collector
Clarke K.L 184, Bruhl J.1., Wilson K.L., Cowie J.D.
Rice B.L.
Clarke K.L 245, Bruhl J.J., Wilson K.L., Cowie J.D.
Clarke K.L 241, Bruhl J.J., Wilson K.L., Cowie J.D.
Bruhl J.J. 369A
Clarke K.L 251, Bruhl J.1, Wilson K.L., Cowie J.D.
Clarke K.L 239, Bruhl J.J, Wilson K.L., Cowie J.D.
Wilson K.L. 5442
Beauglehole A.C. 26084
Mitchell A.S. 1150
Tindale M.D. 2058
Bruhl J.1., Hunter J.T., Egan J. 1269B
Bell D.B.
Latz P.K. 8263
Clarke K.L 160, Bruhl J.J, Wilson K.L.
Knight F. 14185
Clarke K.L 221, Bruhl J.J, Wilson K.L., Cowie J.D.
Clarke K.L 113, Bruhl J.J, Wilson K.L.
Clarke K.L 100, Bruhl J.J.
Burkill H.M., Shah M. HMB235
Hill S.R. 24361
Napper D.M., Kanuri 2079
Raizada M.B.
Correll D.S. 52337
Rajn R.R.V.
Polhill R. 847, Paulo S.
Larsen K. 1299, Smitinand T., Warncke E.
Cleland J.B.
Mitchell A.A. 479
Latz P.K.7126
Latz P.K. 6339
Royce R.D. 1491
Latz P.K. 7087
George A.S. 820
Payne A.L. PRP 1854
Harris P.L. 342
Beaug1ehole A.C. 26568
Bruhl J.J., Quinn F.C. 1197
Hunter J.T., Hunter V. 2737
Forster P.I. PIF8482
Blake S.T. 21453
Bean A.R. 1570
Table 5.1 cont'd
bde7
Qld
Hubbard C.E. 3128
McKee H.S. 9317
bde8
Qld
bde9
N.S.W.
Williams J.B.
Gray M. 3255
bdel0
N.S.W.
bdell
P.N.G.
Croft 34706, Lelean
bde12
Robbins R.G. 2660
P.N.G.
bde13 Philippines Ramos M., Edaro G.
bde15
Sri Lanka Davidse G. 7614
Eulbostylis sp. aff. baffdl
Bean A.R. 3236
Qld
baffd2
Field survey team 820
densa 1
China
Eulbostylis sp. aff. bde16 South Africa Meeuse A.D.J. 10158
densa 2
bde17 South Africa Scheepers J.C. 1141
bde18
Swaziland Haines R.W. 7048
Eulbostylis
bbul
W.A.
Hart R.P. 2092
burbidgeae
bbu2
W.A.
Carolin R.7640
bbu3
W.A.
Mitchell A.A. 1929
bbu4
Burbidge N. 1102
W.A.
bbu5
N.S.W.
Payne A.L. PRP976
Eulbostylis sp. aff. baffbu
N.T.
Dunlop C.R. 4725
burbidgeae
Latz P.K. 11364
Eulbostylis sp. aff. baffpu
N.T.
puberula
Eulbostylis
bpyl
pyriformis
N.S.W.
Johnson L.A.S.
bpy2
N.T.
Latz P.K. 10622
bpy3
N.S.W.
Hunter J.T., Bell D.B.
bpy4
N.T.
Latz P.K. 488?
bpy5
N.S.W.
Wilson K.L. 1479A
bpy6
N.T.
Latz P.K. 9852
bpy7
Sharpe P.R. 232
Qld
bpy8
Qld
Bean A.R. 4227
142
(Lye) R.W.Haines and B. hispidula subsp. senegalensis (Chenn.) Vanden Berghen)
were restricted to use in cladistic analysis.
Phenetic study
As Bulbostylis was recovered as a distinct group in the main phenetic analysis
perfonned in Chapter 3, only taxa from Bulbostylis were included in the phenetic
analyses for this chapter.
Pattern analyses
Additional OTUs were added to the Bulbostylis OTUs in the main data set (in
Chapter 3). A total of 70 specimens (OTUs) of Bulbostylis fonned the basis for the
phenetic study (Table 5.1 see also Appendix 1 for full species list), where 20
quantitative and 89 qualitative morphological characters (Table 5.2) were analysed in
PATN (Belbin 1993).
Data were subjected to ordination, cluster and network analyses as detailed in
Chapter 2, and the combined data set, analysed using the Gower Metric sin1ilaritycoefficient, is presented here.
Groups that were clear-cut in the first analysis for the genus were relnoved and the
data re-analysed as subsets to assess the remaining taxa. Two-dimensional scatter
plots were used to present the ordination results. Boundaries of the 3-dimensional
ordinations were outlined in the corresponding 2-dimensional scatter if the 2dinlensional groupings were indistinct.
Table 5.2 Attribute codes and definitions used in the main phenetic analyses for
the Australian Bulbostylis, including corresponding initial weight values. Weight
values changed in subset analyses.
Attribute
Description
charI
Mean aerial spikelet width in mm (spikelets with mature fruit) at the
widest point
Mean aerial nut length in mm from base of stipe to nut apex (excluding
persistent style base)
Mean aerial nut width in mm at widest point
Aerial nut length:width (ratio 1:W/L(x) (to decimall/x), ratio
coefficient
Mean aerial nut 'stipe' length in mm
Stipe length/nut length (proportion)
Mean aerial anther length in mm (including appendages)
Mean aerial style length in mm (including style base to base of style
arm junction)
Mean aerial style width in mm (at mid third)
Style length:width (1 :W/L(x) to decimal l/x), ratio coefficient
Mean aerial stylebase length in mm (from base to constriction at style
junction)
Mean aerial stylebase width in mm (at widest point)
Style base length:width (1 :W/L(x) to decimal I/x); ratio coefficient
Mean aerial glume length in mm (from base of nerve to apical point)
Mean aerial glume width in mm (at widest point)
Glume length:width (1:W/L(x) to decimal l/x); ratio coefficient
Mean leaf width in mm (at mid third)
Mean culm width in mm (at mid third)
Mean root width in mm (one em below plant base)
Mean inflorescence-synflorescence length in mm (from base of main
bract to furthermost point of spikelets)
Basal spikelets O-absent: always only aerial; I-present: basal spikelets
(morphologically distinct) as well as aerial spikelets
Sub-radical spikelets (Wilson 1980), spikelets that are aggregated near
the plant base that are morphologically similar to the aerial spikelets:
the nuts are indistinct from aerial nuts
Nut shape in transverse section is plano-convex; dorsal/ventral sides of
a 3-angled fruit with the adaxial face distinct from the rest, being
broader than the abaxial faces, often +/- rounded
Nut shape in transverse section is strongly triqetrous with deeply
concave faces
Nut shape in transverse section is triquetrous, having 3-angles, with
faces being concave
Nut shape in transverse section is trigonous, 3-angles with faces
somewhat flattened
Nut shape in transverse section is rounded trigonously, with 3 equal
sides but well rounded edges and faces (convex)
Nut outline obovate (2:1 or 3:2)
Nut outline widely obovate (6:5)
Nut outline very widely obovate (1: 1)
Nut outline pyriform (pear-shaped)
Nut outline obtrullate
Nut outline widely obtrullate (6:5)
Nut outline very widely obtrullateI: 1)
char2
char3
char4
char5
char6
char7
char8
char9
charlO
charI 1
charI 2
char13
charI 4
charI 5
charI 6
charI 7
charI8
charI 9
char20
char21
char22
char23
char24
char25
char26
char27
char28
char29
char30
char31
char32
char33
char34
Weight
1
1
1
1
1
1
1
1
1
0.5
0.5
0.2
0.2
0.2
0.2
0.2
0.125
0.125
0.125
0.125
0.125
0.125
0.125
Table 5.2
(cont'd)
char35
char36
Nut outline napiform
Nut epidermis without protuberances (apparent at 50x magnification
under a dissecting microscope
Nut epidermis rugulose (minutely rugose)
Nut epidermis rugose with rounded waves
Nut epidermis rugose with acute waves (apex acute from a central
raised silica body
Nut epidermis sub-puncticulate, from single raised cells that are not
prominent and are scattered over the surface
Nut epidermis puncticulate, from prominent single cells raised evenly
over surface
Nut epidermis with rows of warts on face, usually 2 vertical rows on
each face
Nut epidermis reticulate, from distinct and raised cell walls
Nut epidermis finely hexagonal, giving a honeycomb appearance,
obvious at lOx magnification under a dissecting microscope
Nut epidermal cells isodiametric; almost square to just rectangular
Nut epidermal cells oblong longitudinally (2: 1)
Nut epidermal cells narrowly oblong longitudinally (6; 1;3; 1)
Stamen number: 1
Stamen number: 2
Stamen number: 3
Sheath glabrous (hairs absent)
Sheath with short to medium hairs (60-100 /lm)
Sheath with bristly hairs almost erect from surface (> 100-1000 /lm)
Glume margins entire
Glume margins ciliolate; small fine hair-like projections from the
margins, sometimes only distally (5 div @ 50x - sometimes only
distally)
Glume margins fimbriolate; small, flattened projections from margins,
sometimes only distally (100 /lm)
Glume margins piliferous; fine, long, loose hairs arising from the
margins (1 000 /lm)
Glume apex rounded
Glume apex acute (muticous)
Glume apex sub-mucronulate
Glume apex mucronulate
Glume apex mucronate
Glume apex acuminate
Glume apex aristate
Glume outline ovate
Glume outline trullate (kite-shaped)
Glume outline narrowly triangular
Glume outline linearly triangular
Glume apex reflexed at maturity
Glume apex not reflexed at maturity
Glume nerve muticous
Glume nerve to a mucro point
Glume nerve excurrent (greater than 0.5 mm)
Glume abaxial surface glabrous
char37
char38
char39
char40
char41
char42
char43
char44
char46
char47
char48
char49
char50
char51
char52
char53
char54
char55
char56
char57
char58
char59
char60
char61
char62
char63
char64
char65
char66
char67
char68
char69
char70
char71
char72
char73
char74
char75
0.125
0.125
0.125
0.125
0.125
0.125
0.125
0.125
0.125
0.2
0.2
0.2
0.2
0.33
0.33
0.33
0.33
0.33
0.33
0.25
0.25
0.25
0.25
0.143
0.143
0.143
0.143
0.143
0.143
0.143
0.25
0.25
0.25
0.25
0.5
0.5
0.33
0.33
0.33
0.167
Table 5.2
(cont'd)
char76
char77
Glume abaxial surface with nerve only scabrid
Glume abaxial surface scabrid over lower half of glume (even isolated, sparse, dense toothed hairs)
Glume abaxial surface scabrid over most of the glume back (20-40 fim)
Glume abaxial surface with short hairs (100 fim)
Glume abaxial surface bristly, with erect hairs (> 1000 fim)
Glume arrangement on the rachilla distichously spiral (glumes opposite
each other and glume pairs ascending arranged spirally)
Glume arrangement on the rachilla tristichous
Non fertile glume number at the base of each spikelet: 0
Non fertile glume number at the base of each spikelet: 1
Leaf to culm ratio: 1to 1
Leaf to culm ratio: 2t03
Leaf to culm ratio: 1t02
Leaf to culm ratio: lt03
Leaf to culm ratio: 1 to 4
Culm surface glabrous
Culm surface scabrid (includes distally)
Culm surface with short hairs that are almost erect (c. 100 Jlm)
Culm surface bristly with stiff erect hairs (includes distally)
Leaf abaxial surface glabrous
Leaf abaxial surface with scabrid margins
Leaf abaxial surface scabrid over the abaxial surface
Leaf abaxial surface with erect to ascending hairs (c. 100 fim)
Leaf abaxial surface bristly/hispid (1000 Jlm - erect to outwardly
ascending)
Inflorescence-synflorescence mostly solitary (HF 1), or 1-2
coflorescences (Cofl)
Inflorescence-synflorescence as anthelodia, main florescence (HF 1)
plus multiple primary coflorescences (Cofl) on lengthened epipodia
(rays); some coflorescences may be sessile, but not all
Inflorescence-synflorescence as ramified (compound) anthelodia
Inflorescence-synflorescence as 'heads' of 3-7 sessile spikelets
Inflorescence-synflorescence hemispherical 'head' of> 7 sessile
spikelets
Inflorescence-synflorescence bracts present and glume-like
Inflorescence-synflorescence bracts present and leaf-like
Inflorescence-synflorescence bracts shorter than inflorescencesynflorescence length
Inflorescence-synflorescence bracts equals the inflorescencesynflorescence length
Inflorescence-synflorescence bracts longer than the inflorescencesynflorescence length
Inflorescence-synflorescence prophyllar buds or spikelets present
Inflorescence-synflorescence prophyllar buds or spikelets absent
char78
char79
char80
char8l
char82
char83
char84
char85
char86
char87
char88
char89
char90
char91
char92
char93
char94
char95
char96
char97
char98
char99
chariOO
charlO 1
charl02
charl03
charI04
charl05
charI06
charl07
charl08
charl09
charI 10
0.167
0.167
0.167
0.l67
0.167
0.5
0.5
0.5
0.5
0.2
0.2
0.2
0.2
0.2
0.25
0.25
0.25
0.25
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.5
0.5
0.33
0.33
0.33
0.5
0.5
Cladistic analysis
Ingroup
Species of Crosslandia and Abildgaardia that were defined in Chapters 3 and 4
were combined with the terminal taxa of Bulbostylis as determined in the phenetic
analyses of this chapter. Samples of Bulbostylis humilis, B. puberula, B. hispidula
subsp. pyriformis and B. hispidula subsp. senegalensis were added to the ingroup
data. Terminal taxa of Fimbristylis used in previous analyses were maintained in this
cladistic analysis (Table 5.3).
Outgroup
The outgroup in the cladistic analysis for this chapter comprised Arthrostylis
aphylla, provisional Actinoschoenus compositus, Trachystylis stradbrokensis
(Donlin.) Klik., Schoenoplectus tabernaemontani (C.C.Gme!.) Palla (= S. validus
Yahl), Schoenoplectiella lateriflora (J .F.Gmel.) Lye (= Schoenoplectus lateriflorus),
and Schoenoplectiella laevis (S.T.Blake) Lye (= Schoenoplectus laevis) (Appendix
1).
Characters and homology
Guaglianone (1970) observed intraprophyllar buds within the inflorescence
prophylls of species in Bulbostylis, and proposed the presence of intraprophyllar
buds as a generic separator between Bulbostylis and Fimbristylis; intraprophyllar
buds are absent in Fimbristylis. All specimens of Bulbostylis used in this study were
examined for the presence of intraprophyllar buds or spikelets.
Tablle 5.3 Taxa included in the cladistic analyses to assess the relationships of
Australian species of Bulbostylis. Species fron1 Crosslandia and Abildgaardia included
here were defined in Chapter 3 and 4 respectively. See Table 5.1 for Bulbostylis specimen
list and Appendix 1 for specimen details.
Taxa
No. specimens
sampled
Ingroup
Abildgaardia macrantha (provisional)
Abildgaardia mexicana
Abildgaardia odontocarpa (provisional)
Abildgaardia ovata
Abildgaardia oxystachya (provisional)
Abildgaardia pachyptera (provisional)
Abildgaardia schoenoides
Abildgaardia triflora
Bulbostylis barbata
Bulbostylis burbidgeae
Blllbostylis capillaris
Bulbostylis densa
Bulbostylis hispidula subsp. pyriformis
Bulbostylis hispidula subsp. senegalensis
Blllbostylis puberula
Bulbostylis pyriformis
Bulbostylis humilis
Bulbostylis turbinata
Bulbostylis sp. aff. barbata
Bulbostylis sp. aff. burbidgeae
Bulbostylis sp. aff. densa 1
Bulbostylis sp. aff. densa 2
Bulbostylis sp. aff. turbinata 1
Bulbostylis sp. aff. turbinata 2
Bulbostylis sp. aff. puberula
Crosslandia anthelata (provisional)
Crosslandia setifolia
Crosslandia spiralis (provisional)
Crosslandia vaginata (provisional)
Fimbristylis bahiensis
Fimbristylis blakei
Fimbristylis cinnamometorum
Fimbristylis depauperata
Fimbristylis fimbristyloides
Fimbristylis furva
Fimbristylis hygrophila
Fimbristylis microcarya
Fimbristylis schultzii
Fimbristylis sp L.
Fimbristylis variegata
10
5
2
13
13
11
12
4
20
5
4
15
3
2
7
8
3
8
7
1
2
3
1
1
1
5
18
3
14
4
2
5
2
4
2
2
2
2
2
1
oオエセ イッ ー
Actinoschoenus compositus (provisional)
Arthrostylis aphylla
Schoenoplectiella laevis
Schoenoplectiella lateriflora
Schoenoplectus tabernaemontani
Trachystylis stradbrokensis
4
4
5
5
3
7
14S
Embryo morphology
Van der Veken (1965) and Goetghebeur (1986) sampled embryos of some species
of Bulbostylis, reporting variation in general embryo shape and size, and primordial
leaf development. Bulbostylis barbata was the only Australian species sampled in
both studies. In this study, embryos from representatives of each of the species (and
subspecies) were sampled and compared (Appendix 1). Second and third (ifpresent)
prinliordial leaves were not scored due to difficulties in observing these structures in
many of the embryos. Tissues in small embryos were much denser than in the larger
embryos sampled, and I was unable to clear some small embryos sufficiently to
define the inner layers (e.g. B. sp. aff. barbata, and B. sp. aff. densa l). Alternative
methods in pre-treating the embryo before clearing, or using a different clearing
medium, may be necessary in future work.
Anatomy
Leafblade and culm anatomy were sampled across the species of Bulhostylis
studied to compare general shape and tissue arrangement. General anatomy
(Metcalfe 1969, 1971) and photosynthetic pathway in Bulbostylis (Goetghebeur
1986; Bruhl 1995); has been reported to be the same as commonly found in
Fimbristylis; sampling tested the uniformity in this study.
PA up* analyses
Parsimony analysis was performed on 47 terminal taxa and 155 characters using
heuristic searches (hsearch swap=TBR addseq=rand nreps=l 000 hold=5
multrees=yes). Branch support was assessed using Bootstrap analysis (1000
bootstrap replications) because the computational time required to calculate the
Bren1er support indices past 3 extra tree length steps was too protracted, even when
limiting the addition-sequence replications to 10. Characters were traced in
MacClade and the most relevant characters are presented in the cladogram.
Results
Phenetic study
Representative OTUs for the genus Bulbostylis formed a distinct group in the
initial main analysis (see Chapter 3), with some species groups of OTUs (B. sp. aff.
barbata, B. barbata, B. burbidgeae and B. sp. aff. puberula) apparent in the
Bulbostylis cluster at the broad level in 2-dimensional analysis (Figure 5.1).
"Then additional samples of Bulbostylis were added to the first main analysis and
re-analysed (see Materials and methods, this chapter), distinct species groups were
fornled by the OTUs in the 2-dimensional ordination (stress = 0.18; Figure 5.2).
Characters that were most strongly correlated with the groups formed within the
ordination were consistent with the sYnf10rescence type, nut epidermal patterning,
number of stamens, anther length, nut stipe length to nut length ratio, hairiness (or
absence of) of culms and glumes, and shape of glumes (Figure 5.3). The
hemispherical heads of sessile spikelets associated with B. barbata and B. sp. aff.
barbata and the mostly primary rayed anthelodium (i.e. spikelets on lengthened
epipodia) of B. pyriformis and B. turbinata were correlated with the separation of the
groups.
• Crosslandia
III Abildgaardia
l
.. B. sp. aff. barbata
X B. barbata
:( B. turbinata
• B. densa
+ B. burbidgeae
<> B. sp. aft. puberula
'
I'
J
I
B. sp. aff. burbidgeae
<> B. pyriformis
o Fimbristylis
•
•
•.#••
• ..,_
:;• •• ••
•
.
..... •
# ••
••
D
D
D
D
Den
..
%{;} D
0
Figure 5.1. MDS ordination in 2-dimensions (stress = 0.17) from primary phenetic analysis
(see Chapter 3) highlighting Bulbostylis. Species groups for B. sp. aff. barbata, B. barbata
and B. burbidgeae are distinct in this broad level ordination. See Table 5.1 and Appendix 1
for OTU and specimen details.
.
n
.. ·r· ·
•
II , :
•
•
Figure 5.2. MDS ordination for OTUs of Bulbostylis (stress = 0.18). Lines separating OTUs in B.
densa (amphicarpic specimens) and B. sp. aff. densa indicate the clear-cut groups observed in 3dimensions (stress = 0.12). See Table 5.1 and Appendix 1 for OTU and specimens details.
'.66.6
46,4307;:(3.100 +67 -gO -51 <>75 D1?j
Stipe length/nut length
Inflorescence: primary 'anthela'
Glunne shape: ovate
Nut width
•
•
Culm glabrous
Glume back glabrous
Stamens: 3
o
Anther
length
IT
Inflorescence: hemispherical
Glume shape: trullate
Nut epidermis: reticulate
Nut cell outline: isodiametric
Figure 5.3 Characters that correlate (>80%) to group formation in the ordination shown in Figure
5.1. Inflorescence-synflorescence of many sessile spikelets forming a hemispherical 'head', glume
shape: trullate, nut epidermis being reticulate with isodiametric cells, plus anther length separated
Bulbostylis barbata and B. sp aff. barbata from the other OTU groups. Culm and glume backs
glabrous contributed to separating the B. densa group and stamens numbering 3 correlated
strongest with the B. burbidgeae OTUs. The group containing OTUs for B. turbinata and
B. pyriformis were consistent with the correlated characters of inflorescence-synflorescence:
primary 'anthela', highest stipe length-nut length ratio; B. pyriformis has the greatest nut widths.
See Table 5.2 for attribute definitions.
QエセY b
Bt2
Bt
"'?';$l_ _
M M Mセ
Bt7
Bt8
Bt1
I.
I
•
B. aff. barbata
B. barbata
B. turbinata
B. densa
1)(
B. aft. densa
• B. burbidgeae
;+ B. aff. puberula
• B. aff. burbidgeae
!
I
B. pyriformis
Figure 5.4. Minimum spanning tree (MST) for OTUs of Bulbostylis corresponding to ordination in
Figure 5.2. Borders indicate greater separation seen in 3-dimensions (stress = 0.12). See Appendix
1 for specilnen details.
0.0359
baffba1
baffba4
baffba6
baffba7
baffba2
baffba5
baffba3
btl
bt4
bt7
bt5
bt8
bt6
bt2
bt3
bt9
bt10
baffbu
bbu1
bbu2
bbu3
bbu4
bbu5
baffpu
bpy1
bpy2
bpy3
bpy7
bpy8
bpy4
bpy5
bpy6
bba1
bba2
bba3
bba8
bba4
bba6
bba5
bba10
bba9
bba11
bba20
bba12
bba7
bba13
bba14
bba15
bba16
bba17
bba18
bba19
bde1
bde2
bde8
bde10
bde9
bde13
bde15
bde12
bde3
bde4
bde7
bde6
bde5
baffd1
baffd2
bde16
bde17
bde18
0.1381
I
0.2403
1
1}_
0.3426
1
1
0.4448
0.5470
I
1
4}_1_
6}_
I
7}/_1_
2} _ _ I
5}_1_1
3} _ _ 1
_
28}
_
31} _ _
34}_1_
I
I
32} _ _ 1 _ _
1_ I
35}
33}
1_1_
29}
1
30)
1
36}
37}
62}
56}_
57)1_
58}_1
59}_11_
60) _ _ 1
61 )
63)
_
64}_
65}1_
69)_
I
_
1
1_1 _ _
1
_
_
I
7o}_I_I_I_
1
66}
67)_
II
68}_1 _ _ 11
-
_
8} _ _
9}_1_
10} _ _ I_ _
15}
111) _ _
13}_I
_
12}_
17}_I _ _
16} _ _ 1 _ _
18}__
I
27} _ _ 1_ 1
19} _ _ 1_1
14}
20}
21}__
I
_
1_
1
I
22}_1_1
1 __
23}
_
1_
24}
25)
1
1_1
26}
38)___
39} _ _ 1____
45}__
47} _ _ 1_
46) _ _ 1__
49)
I
50)
1_1_
48}
1__
40}______
41}_
44}_1__
1_
43}
1_____
42}
51)
I
152}
1
53}__
1
54}_ I
1
55}_1_1
_
I
1
I
I
1
1
1
I
I
1
I
1
1
I
1
I
1
1
1
1
I
1
I
I
0.0359
0.1381
0.2403
0.3426
0.4448
_
0.5470
Figure 5.5. WPGMA Hセ = -0.1) phenogram that corresponds with the ordination in Figure 5.2, for
all OTUs of Bulbostylis. OTUs form six groups, the putative species B. sp. aff. barbata is clearly
separated from B. barbata. See Table 5.1 and Appendix 1 for OTU and specimen details.
155
Species groups in the ordination were generally supported by network (Figure 5.4)
and cluster analyses (Figure 5.5). Stronger grouping was obtained in the ordination
for 3-dimensions (stress = 0.12) as indicated in the 2-dimensional scatter plot (Figure
5.2).
Individual OTUs, baffbu (B. sp. aff burbidgeae), baffpu (B. sp. aff. puberula), bt9
and btl 0 (B. turbinata), were separated from other groups, and from each other, in
the 3-dimensional ordination. Although the separation is evident in 2-dimensions for
bafjbu and baffpu, this is not the case for OTUs bt9 and btl 0, which appear as if
distinctly clustered with the other B. turbinata OTUs. Within the phenogranl (Figure
5.5), baffbu is clustered with OTUs of B. turbinata and baffpu is broadly included
with the B. burbidgeae OTU group. However, the dissimilarity ofbaffpu to OTUs of
B. burbidgeae is present in the phenogram.
Discrete groups of OTUs were retrieved as the species groups B. sp. aff. barbara,
B. barbata, B. pyriformis, B. burbidgeae and B. turbinata (excluding bt9, btl 0)
within 3- and 2-dimensional ordinations (Figure 5.2), and generally in the phenogram
(Figure 5.5). The remaining OTUs formed the B. densa group that included baffd 1,2
(B. sp. aff. densa) and bde16, 17, 18 (African amphicarpic samples of B. densa).
Bulbostylis densa group
In the subset analyses of the B. densa group, OTUs for the amphicarpic specimens
colIected from Africa formed a group separate to the main B. densa OTUs, and to the
OTUs of B. sp. aff. densa. The three groups are more robust in 3-dimensions (stress
= 0.12), indicated by the drawn boundaries around the specific groups in the 2dinlensional scatter plot (Figure 5.6). There were 16 characters with greater than
•
•
•
•
•
• • ••
•
•
•
•
•
• B. densa
• B. aft. densa
B. densa amphicarpic
-
-
-
-
Figure 5.6 MDS ordination in 2 dimensions (stress = 0.18) for the Bulbostylis densa group from
the prirrlary Bulbostylis analysis (see Figure 5.2). OTUs for the amphicarpic material from
Africa form a group separate to the main B. densa group and the separate OTUs of B. sp. aff.
densa. The boundaries shown indicate the distinct OTU groups in the 3-dimensional ordination
(stress =: 0.12). See Table 5.1 and Appendix 1 for OTU and specimen details.
.7 .60 84 010
Anther L
Basal spikelets: present
Synflorescence: solitary or few
rayed
Glume apex: mucronate
Glume margins: fimbriolate
::4(
56 .30
+ 63
-21 セ ... 99 <> 57
44 659,46,36,43
28
Empty glumes: 1
Glume apex: rounded
Nut epidermal sculpturing: reticulate
Nut epidermal cells: isodiametric
•
o
Nut outline: obovate
Glume apex: acute
•
Nut outline: very
widely obovate
Glume apex: rounded
Style L/W
Nut epidermal cells: finely hexagonal
Figure 5.7 Characters correlated (> 70 0/0) with the ordination in Figure 5.6 for OTUs of the
ウゥャケセ ッ「 オb
densa group. Characters with >80 % correlation to the ordination were anther
length, glume apex: acute, empty glume: 1, style length to width ratio, glume margins: ciliolate,
and nut shape: obovate. L=length. See Table 5.2 for attribute definitions.
Bd18
WQ、bセ
Bd1
•
Bd16
\Bd2
Bd15":7
•
Bd10
Bd4
Bd13
セR、ヲ 。b
.B. densa
• B. aff. densa
Baffd1
B. densa amphicarpic
Figure 5.8 Minimum spanning tree (MST) with linkages for the Bulbostylis densa group plotted
onto the 2-dimensional ordination in Figure 5.6. See Appendix 1 for specimen details.
0.1600
bde1
bde2
bde3
bde4
bde7
bde5
bde8
bde9
bde10
bde13
bde15
bde12
bde16
bde17
bde18
baffdl
baffd:2
1
1)_
2) 1
3)
4)_
6)_1
5)
7)_
8)_1_
9)_1
11)
12)
10)
15)_
16}_ I
17} I_I
13}
14}
1
0.1600
0.2324
0.3048
I
I
0.3772
1
0.4496
0.5220
I
I
1
1
1
0.2324
I
0.3048
I
0.3772
1---I
I
1
1
0.4496
0.5220
Figure 5.9 WPGMA (P = -0.1) phenogram for the Bulbostylis densa subset (see Figure 5.2 for
all species of Bulbostylis) that best correlates with the ordination (Figure 5.6) and Minimum
Spanning Tree (MST) (Figure 5.8). Operative Taxonomic Units (OTUs) for B. densa with
amphicarpic nuts (bde16-18) are grouped together separate to the remaining OTUs for B. densa.
Similarly, OTUs for B. sp. aff. densa (baffdl, baffd2) group separately. See Table 5.1 and
Appendix 1 for OTU and specimen details.
160
セGj\PW
correlation to the three groups seen in the ordination scatter plot (Figure 5.7).
Characters with >800/0 correlation to the ordination were anther length, glume apex:
acute, empty glume number: 1, style length to width ratio, glume margins: ciliolate,
and nut shape: obovate. Linkages between OTUs in the minimum spanning tree
(Figure 5.8) support the major groups (B. densa S.s., B. densa 'amphicarpic', B. sp.
aff. densa) and the minor groups (within B. densa s.s.) observed in the phenogram
(Figure 5.9).
Bulbostylis turbinata-B. sp. aff. burbidgeae group
Subset analysis of the B. turbinata-B. sp. aff. burbidgeae group of OTUs indicates
that B. sp. aff. burbidgeae bt9 and btl a are discrete units from the main OTU group
of B. turbinata in the 3-dimensional ordination (stress
=
0.1), and to a lesser extent in
2-dimensions (Figure 5.10). There were 24 characters with >70% correlation to the
ordination; characters with >80% correlation were inflorescence prophyllar
branching: present, inflorescence prophyllar branching: absent, culm width, style
length, glume width and style base width (Figure 5.11). The minimum spanning tree
OTU linkages (Figure 5.12) correspond to the groups from the ordination and cluster
analysis (Figure 5.13).
Terminal taxa as recognised in the phenetic analyses, i.e. Bulbostylis sp. aff.
barbata, B. barbata, B. turbinata, B. sp. aff. turbinata 1, (bt9), B. sp. aff. turbinata 2
(btl 0), B. pyriformis, B. burbidgeae, B. densa, B. sp. aff. densa 1, B. sp. aff. dellsa 2
(African amphicarpic), B. sp. aff. burbidgeae, and B. sp. aff. puberllla, were
cornbined with samples from B. humilis, B. capillaris, B. hispidula subsp.
senegalensis, and B. hispidula subsp. pyriformis for use in cladistic analysis.
' . 8 . turbinata
.819
148t10
X 8. sp. aff. burbidgeae
•
•
•
•
i
•
• •
•
Figure 5.10 MDS ordination in 2-dimensions (stress= 0.18) for OTUs of the B. turbinata group
from Figure 5.2 The OTUs bt9, bt10 and B. sp. aff burbidgeae (baffbu) are separated from the
main group of B. turbinata OTUs. The boundaries indicate the stronger group resolution seen in
the 3-dimensional ordination (stress = 0.11). See Table 5.1 and Appendix 1 for OTU and
specimen details.
,• 109 • 110
18 0 12 ;:( 8 • 15
Aerial style L
Syn tlorescence
prophyllar buds: present
•
•
Syntlorescence
prophyl1ar buds: absent
CulmW
o
Aerial style base W
•
Aerial glume W
Figure 5.11 Attributes correlated (>80%) with the ordination in Figure 5.10 for OTUs of the
Bulbostylis turbinata group. Key attributes were style length, style base width, aerial glume
width, culm width, synflorescences prophyllar bud, or growth, absent/present (polymorphic).
L=length, W=width. See Table 5.2 for attribute definitions.
I
• B. turbinata
I.Bt9
!.Bt10
I
X B. sp. aft burbidgeae
I
Bt2
Bt1
Figure 5.12 Minimum spanning tree (MST) with linkages for the Bulbostylis turbinata group
plotted onto the 2-dimensional ordination of Figure 5.10. See Appendix 1 for specimen details.
0.1480
btl
bt2
bt3
bt6
bt4
bt7
bt5
bt8
bt10
bt9
baffbu
1)
1
0.2322
1
7)
0.4006
I
0.4848
1
--:-
1
11 _ _
I
I
T
5)
1
8)
1 _ _- . , . -
10)
1 __
9)
11)
0.5690
I
I
--:-1
2)
3)
6)
4)
0.3164
1
_
""--1----.,.-1-----:--1------,1,------------:1-
I
1
0.1480
0.2322
0.3164
0.4006
0.4848
0.5690
Figure 5.13 WPGMA Hセ]
-0.1) phenogram for the Bulbostylis turbinata subset (see Figure 5.2
for all OTUs of Bulbostylis) that fits the ordination (Figure 5.10) and MST (Figure 5.12).
Bulbostylis OTUs group together and the OTUs bt9, btl0, and B. sp. aff. burbidgeae (baffbu)
remain separate. See Table 5.1 and Appendix 1 for OTU and specimen details.
165
Cladistic analysis
One hundred and twelve most parsimonious trees were retrieved (Tree
length=1210, CI=0.4455, HI=O.5545, RI=O.5984, RC=O.2666) from a heuristic
search. Tree 1 had similar topology to the tree obtained from strict consensus, and
was selected to show branch support and character/branch associations (Figure 5.14).
Two broad sister clades, the Bulbostylis-Fimbristylis clade (A) and the
Abildgaardia-Crosslandia clade (B), form an internal clade sister to Fimbristylis
bahiensis, which is, in tum, sister to Fimbristylis variegata (Figure 5.14).
Fimbristylis continues to be retrieved as a non-monophyletic group (see also
Chapters 3 and 4), seen by F. sp. Land F. blakei being placed sister to the
Crosslandia-Abildgaardia clade. Crosslandia is not a monophyletic group in this
analysis, as the provisional C. vaginata is placed sister to species of Abildgaardia.
Within clade A, all taxa sampled from Bulbostylis formed a clade
(Bootstrap=83%) sister to Fimbristylis fimbristyloides, and combined on branch C as
a group sister to the clade Fimbristylis 1 (internal branch D, contains F. depauperata
from the TYPE section of Fimbristylis section Fimbristylis). Both clades from C and
D are nested within species assigned to Fimbristylis.
The Bulbostylis clade (Bootstrap=83%) was formed by two main clades where
B. humilis (B4) is sister to the remainder of the species (B3, B2 and B 1); the latter
with moderate branch support (Bootstrap=74%). B. barbata (B3) was sister to the
terminal groups B2 (B. pyriformis-B. turbinata-B. burbidgeae-B. hispidula clade)
and B 1 (B. capillaris-B. densa-B. puberula clade). The strict consensus shows
branch collapse for most of the terminal taxa within the Bulbostylis clades Bland
B2; only the terminal B. puberula and B. capillaris (with weak branch support), and
B. sp. aff. barbata-B. burbidgeae-B. sp. aff. burbidgeae branches persist. Moderate
Figure 5.14 Cladogram for tree 1 of 112 shortest trees (tree length = 1490) in the assessment
ofmonophyly for Australian species of Bulbostylis. Bulbostylis forms a monophyletic gTOUp
sister to Fimbristylis fim bristyloides, fonning clade C, which is sister to Fimbristylis 1 in
clade B. Fimbristylis depauperata that is from the TYPE section of the genus, is placed in
clade B. Abildgaardia and Crosslandia form a broad group that is sister to Fimbristylis sp. L
and F. blakei; all grouped in clade D. Crosslandia is not monophyletic in this analysis.
Within the Bulbostylis clade four main groups frequently occur (B 1, B2, B3, B4), however,
only 3 groups were retrieved from strict consensus of the 112 most parsimonious trees: B4,
B3 and B 1-2. Bootstrap support values are given below the branches. A, B, C, and D
indicate the intenlal branch for the main clades. Dashed lines indicate collapsed
branches in the tree from strict consensus. See Appendix 1 for specimen details and
Appendix 2 for characters.
A
イMGhヲi!Kfゥュ「ウエケャ
variegata
153-3; 1-2; 14-2; 15-2; 28-2; 35-1
r--tlt-I- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 153-3
[セZ
セ QLR[ UM
Qセ [ TMSL [
24-2
Abildgaardia macrantha
35-1; 41-2
-
-
Abildgaardia pachyptera
15-1,2; 41-1
GセARLS[
セQ
28-1
Crosslandia vaginata
.Ill!
14-1,3; 1A-IP; 35-4; 73-1,3
1t;11-1
Fimbristylis hygrophila
28-2
35-1
Abildgaardia schoenoides
---l.I}I.
f741l14-1,3,4;
15-1,2
Abildgaardia odontocarpa
_ 41-2; Qセ[S
Yセ
35-3
Abildgaardia oxystachya
1111111-1,1'+-'j,4;
••
150-2
86-4
115-1,2; 35-1; 41-1
[SMセQ
Abildgaardia mexicana
681'"+++
I
RセQM
15-3
Abildgaardia ovata
35-3; 41-1
"+-_ __.0011 135 -3
B
_
Crosslandia spiralis
150-1
25-2
1セQ
Crosslandia anthelata
66
14-3
15-1
73-3
I
Crosslandia setifolia
115- 1,2
III
""+
14-34· 34-4
ᄋQM iQ セ
'
511
I
"
Fimbristylis furva
1111
14-3; 15-3; 24-1; 25-2
[RMQャセ
C
I
Fimbristylis fimbristyloides
153-2,3; 15-3; 24-1; 25-3
II
r++IIII-·15--2-,-3;-7-3--3--------------
---
35-1
F. cinnamometorum
15-3,4; 24-2; 25-2; 41-4; 57-5; 72-2
IIIII
A
Fimbristylis Sp. L.
Fimbristylis blakei
II
1-2; 14-1
15-2
r+-'------------I
153-2,3
83"4
セZ
.
Bulbostylis barbata
Bulbostylis sp. aft. turbinata 2
-.It----------
.
•
74
Bulbostylis pyriformis
.14-3
.!
:,.! - + 1 - - - - : : 14-3,4
.セUSMᄋA .
••
'.
Bulbostylis turbinata
セZ
Bulbostylis sp. aft. barbata
.
8_3_-1_2
_
Bulbostylis sp. aft. burbidgeae
•
73·;..
_
_
セZゥUᄋ
Bulbostylis hispidula subsp.
senegalensis
B. hispidula subsp. pyriformis
Bulbostylis sp. aft. densa 2
.................. ...
•• •• II
.
: . 153-23·15-3
'
Bulbostylis densa
Bulbostylis sp. aft. densa 1
rtt-II---
:
セQTMRL [
OJ
N
Bulbostylis burbidgeae
25-1
:
Bulbostylis sp. aft. puberula
--------
86-1
+0
Bulbostylis puberula
15-1
Bulbostylis capillaris
I
Fimbristylis microcarya
D
Fimbristylis disticha
111111
II
15-4
24-1
25-3
41-2
57-5
Bulbostylis humilis
Bulbostylis sp. aft. turbinata 1
150-3
11-3
24-2
25-1
-
Fimbristylis schultzii
+I
153-3"++111-- ---.1-5--2--11 2
I
Fimbristylis depauperata
'
rrtt:nfl'lll:1-'4-1--2- - - - - Actinoschoenus compositus
153-3 72 .,11------ Arthrostylis aphylla
15-2
1-2
11-1
28-2
Abildgaardia triflora
25-3,35-5,41-4
II
QMRセM M M M M M M M ᄋM Mi K QLN
Fimbristylis bahiensis
iセ
153-1
141-1
III
15-4; 24-3; 35-1
Itl1-1; 72-1
1114-1; 41-4
+-------
11
15-397 1'1
50-291t11111-2; 41-1; 73-3
1-1
14-3
o
Trachystylis stradbrokensis
S-
Schoenoplectus tabemaemontani
oc
Schoenoplectiella lateriflora
Schoenoplectiella laevis
167
Bootstrap support (73%) for the terminal branch B. aff. burbidgeae and
B. burbidgeae contradicts the branch collapse obtained from strict consensus (Figure
5.14).
Observations
Injlorescence-synjlorescence structure
lV[ost species of the Bulbostylis included in the study possess prin1ary-rayed
spikelets (coflorescences). Solitary spikelets rarely occur, and if spikelets are solitary
then there are at least some rayed spikelets present within the plant (Figure 5.15 A).
Secondary orders within the synflorescence were frequent in B. densa, and B. sp. aff.
den.fJa 1, B. sp. aff. densa 2, B. puberula, B. pyriformis and B. hispidula subsp.
senegalensis. Although intraprophyllar buds were often observed in the open rayed
synflorescence, it was rare for the buds to mature and develop into spikelets.
The 'head' of sessile spikelets in specimens of B. barbata and B. sp. aff. barbata
(Figure 5.15 B, C) is formed from multiple 'branched' sessile primary
coflorescences, plus sessile spikelets that develop from within the inflorescence
prophylls of the coflorescences. Spikelets arising from the intraprophyllar growth are
distinct and contribute to the density of the 'head' of spikelets that occur in the two
species. This synflorescence type of terminal capitulum (56-10) was not a
synapomorphy; B. sp. aff. barbata was grouped within the 'rayed' synflorescence
types of reduced anthelodium and reduced ramified anthelodium seen in the
B. pyriformis-B. turbinata-B. hispidula clade (Figure 5.14).
Style bases in the Australian Bulbostylis pyriformis, and occasionally in
B. turbinata, may persist on, or fall from the nut; the style always detaches from the
Figure 5.15 Variation ofsynflorescence structure for some species of Bulbostylis.
A. Most Bulbostylis in the study possess 2-4 primary rayed spikelets (primary
coflorescences on lengthened epipodia) as seen in B. sp. aff densa 1 (baffd2
pictured); 1-2 secondary coflorescences on rays (lengthened epipodia) may be
present in the B. densa group, including B. puberula. Scale=5 mm. B. Sessile
spikelets plus spike1ets from prophyllar buds form a head in B. barbata and B. sp.
aff. barbata (baffba1 pictured). Sca1e=5 lum. C. Representation of the prophyllar
synflorescence structure seen in B. Sessile spike1ets arise from the axils within the
prophyll and contribute to the ramification of the synflorescence 'head' as indicated
by 2° and 3° orders. D. 'Subradical' spikelets (arrow) may be present in B. barbata
(bba8 pictured), B. turbinata, and B. burbidgeae; these spike1ets have similar
morphology to the aerial spikelets. Scale=10 mm. See Table 5.1 and Appendix 1 for
OTU and specimen details.
A
169
style base leaving the style base on the nut, even if only for a short time. This is in
contrast to style bases in the B. Izispidula group, where the style may persist on the
nut, or fall from the nut intact with the style. In some specimens of B. hispidula
subsp. senegalensis, all the fallen styles observed had the style base intact. The large
bulbous style base present in B. pyriformis and B. turbinata usually protrudes from
the umbonate nut apex and tends to be easily removed. All other species scored in
this study have smaller style bases that sit firmly at the apex of the nut.
Amplzicarpy
The spikelets observed at the base of the plant in B. Izumi/is show different
morphology (73-3) to the basal spikelets of the African B. densa (B. sp. aff. densa 2).
In B. Izumilis the basal spikelets are attenuate and florets may be bisexual. In the
African B. densa (B. sp. aff. densa 2), the spikelets at the base of the plant are
clustered in groups of 2 or 3 at the soil level (73-4) and have glumes that are much
smaller than those in the aerial spikelets or in B. Izumi/is. The basal glumes in
B. densa may fall early and leave the nut exposed. Both types of basal spikelets are
amphicarpic, with nuts in the basal spikelets being larger than in the aerial spikelets;
the glumes also differ. Subradical spikelets (73-2) in some B. barbata, B. turbinata,
and B. burbidgeae differ from classic amphicarpic plants, as the nuts and glumes
reselnble their aerial counterparts in size and shape; the spikelets occur on shortened
culn1s (Figure 5.15 D).
Nut sculpturing
Nut epidermal shape and protuberances (or lack of) can be useful in identifying
species (Figures 5.16-20), but were only broadly associated with the internal
170
Bulbostylis clades (Figure 5.14). Species with nuts that have papillate or granulate
sculpturing (Figures 5.16-18) were split between the Bland B2 clades, so that nuts
with vertically elongated epidennal cells (Figure 5.19) were interspersed. Bulbostylis
burbidgeae nut epidennal cell walls are barely to mildly sinuose, as are the
B. hispidula and B. pyriformis saJnples (Figure 5.19). In contrast, Bulbostylis barbata
and B. sp. aff. barbata have epidennal cell walls that are very strongly sinuose
(Figure 5.20). Samples from the Bulbostylis turbinata group fall between the two
extremes (Figure 5.18).
EmblYo morphology
All species within this study, that fonned the clade Bulbostylis share the general
Bulbostylis-type embryo (synapomorphy 150-3; Figure 5.14). Variation in the size
and general shape of embryos was observed (Figures 5.21-22). The embryos from
B. sp. aff. densa 1 and B. sp. aff. barbata were the smallest sampled, with the very
dense cellular contents obscuring visibility of the primordial leaf or leaves (Figure
5.21 B, D). Embryos from B. hispidula subsp. pyriformis (Figure 5.22 A, B) and B.
hispidula subsp. senegalensis (Figure 5.22 C, D) are conspicuously larger than
embryos of the other species; the basal orientated shoot and root are prominent, and
the second primordial leaf is well-developed and almost the same size as the first
leaf. The embryo of Bulbostylis pyriformis is slightly smaller than that in B. hispidula
and the second primordial leaf was visible, although not well-developed (Figure 5.22
E, F)" Bulbostylis humilis (Figure 5.22 G, H) has an embryo size and internal
structure similar to that seen in the B. hispidula specimens.
Figure 5.16 Scanning electron micrographs (SEM) showing the variation of nuts for some
samples of the Bulbostylis densa group. A. B. densa (bde2) with B. epidermis at higher
magnification; C. B. densa (bde3); and D. B. sp. aff. densa 1 (baffd1) with E. epidermis at
higher magnification, showing epidermal cells with minute central silica body. Epidermal
cell walls are sinuose. Scale bar for A, D=100 J.lm; B, E=50 J.lm; C=200 J.lm. See Table 5.1
and Appendix 1 for OTU and specimen details.
Figure 5.17 Scanning electron micrographs (SEM) showing the variation of nuts for samples
of the Bulbostylis burbidgeae group. A. B. burbidgeae (bbu5) with B. epidermis at higher
magnification; C. B. sp. aff. burbidgeae (baffbu); with D. epidermis at higher magnification;
E. B. sp. aff. puberula 1 (baffpu) with F. epidermis at higher magnification, showing
epidermal cells forming angular ridges. Scale bars A, D=100 /-!m; B, E=50 /-!m; C=200 /-!m.
See Table 5.1 and Appendix 1 for OTU and specimen details.
Figure 5.18 Scanning electron micrographs (SEM) showing the variation of nuts for samples
of the Bulbostylis turbinata group. A. B. turbinata (bt3) with B. epidermis at higher
magnification; C. B. turbinata (bt?) with D. epidermis at higher magnification;
E. B. turbinata (bt9) with F. epidermis at higher magnification; G. B. turbinata (btlO) with
H. epidermis at higher magnification. OTUs bt9 and btlO formed a group separate to other
B. turbinata samples (see Figure 5.2), consistent with the nut differences pictured. Scale bars
for A, C, E, G=IOO Jim; B, D, F, G=50 Jim. See Table 5.1 and Appendix 1 for OTU and
specimen details.
Figure 5.19 Scanning electron micrographs (SEM) showing the variation of nuts for samples
from Bulbostylis pyriformis and the B. hispidula complex. A. B. pyriformis (bpy5) with
B. epidermis at higher magnification; C. B. hispidula subsp. pyriformis (M. Richards
23175B); with D. epidermis at higher magnification; E. B. hispidula subsp. senegalensis
(J T Davey 10) with F. epidermis at higher magnification. All samples have vertically
elongated epidermal cells that may be raised so that the nut sculpturing is rugose. Scale bars
A=200 11m; C, E=500 11m; B, D, F=50 11m. See Table 5.1 and Appendix 1 for OTU and
specimen details.
Figure 5.20 Scanning electron micrographs (SEM) showing the differences between nuts of
Bulbostylis barbata and B. sp. aff. barbata. A. B. sp. aff. barbata (baffba5) with B. epidermis
at higher magnification; C. B. barbata (bba5); and D. epidermis at higher magnification,
showing wax plates on the surface that break away to reveal strongly sinuose cell walls
around the isodiametric cells, creating a reticulate pattern over the surface of the nut. Scale
B, D=50 セュN
See Table 5.1 and Appendix 1 for OTU and specimen
bar A, C=200 セュ[
details.
Figure 5.21 A. Embryo morphology in Bulbostylis. B. capillaris (G. Davidse) B. B. sp. aff.
densa 1 (baffd1) C. B. sp. aff. barbata (baffba4) D. B. barbata (bba18) E. B. sp. aff.
burbidgeae (baffbu) F. B. puberula (G. Davidse 9037). Collector and collection number, or
specimen OTU label are given in brackets. Open arrow=shoot, closed arrow=root. Scale
bar=100 J.!m (for all images). See Table 5.1 and Appendix 1 for specimen details.
Figure 5.22 Variation in Bulbostylis embryo size, shape, and development of second
primordial leaf. A. B. hispidula subsp.pyriformis (M Richards 23175B) and B. the welldeveloped second primordial leaf (long arrow). C. B. hispidula subsp. senegalensis
(J T Davey 10) and D. the second primordial leaf is well-developed. E. B. pyriformis (bpy2)
with a slightly smaller sized embryo and F. second primordial leaf visible but not well
developed. G. B. humilis (C.P. Strong et al.) embryo with H. second primordial leaf welldeveloped. Open arrow=shoot, closed arrow=root. Collector and collector number, or OTU
label are given in brackets. Scale bar=100 !-lm. See Table 5.1 and Appendix 1 for specimen
details.
A
B
c
D
E
F
H
Figure 5.23 Culm and leaf blade transverse sections in Bulbostylis. A. Culm and B. leaf
blade sections of B. capillaris (G. Davidse); C. culm and D. leaf blade sections of B. sp. aff.
densa 1 (A.R. Bean 3236); E. culm and F. leaf blade B. sp. aff. densa 1 (Field survey team
820); G. culm and H. leaf blade sections of B. densa (bde2). Collector and collection
number, or OTU label are given in brackets. Scale bar=100 !lm. See Table 5.1 and Appendix
1 for specimen details.
A
B
c
E
F
G
H
Figure 5.24 Culm and leaf blade transverse sections for Bulbostylis. A. Culm and B. leaf
blade sections of B. burbidgeae (bbu5). Variation observed in C. culm and D. leafblade
sections of B. barbata (K.L. Clarke 187 et al.), and E. culm and F. leaf blade for OTU bbal.
G. Culm and H. leaf sections for B. sp. aff. barbata (baffba1). Collector and collection
number, or OTU label are given in brackets. Scale bar=100 セュN
See Table 5.1 and
Appendix 1 for specimen details.
A
B
c
D
Figure 5.25 Culm and leaf blades transverse sections for Bulbostylis.
A. Culm and B. leaf blade sections of B. puberula (G. Davidse 9037); C. culm and D. leaf
blade sections of B. humilis (C.P. Strong et al.) Collector and collection number are given in
brackets. Scale bar=100 J.lm. See Table 5.1 and Appendix 1 for full specimen details.
181
Vegetative anatomy
The leaf blade anatomy of all species of Bulbostylis in this study conforms to the
C4 fimbristyloid photosYnthetic pathway (Figures 5.23-25). Leaf blades in transverse
section are either sub-triangular in outline, or channelled
H」。ョ ャゥ」オ 。エ・Iセ
usually with
two shallow or acute ribs. Most species sampled have three vascular bundles within
the leaf, occasionally five in B. barbata (Figure 5.24). A hypodermis is absent from
the leaves of all sampled species and the adaxial epidermal cells are inflated roughly
four times that of the abaxial epidermis. Culms are mostly regularly grooved and
almost circular in outline, to irregularly circular and barely wavy (Figures 5.23--25).
All species sampled have numbers of vascular bundles equal to the nun1ber of
sclerenchyma strands. The vascular bundles form one concentric ring below the
epidermal layer. Sclerenchyma strands are often bulbous, forming the ridges of the
channels, but may be square to just rectangular (Figures 5.23-25).
Discussion
The group recovered in phenetic analyses as Bulbostylis sp. aff. barbata is a
distinct new species, as indicated by its placement in the combined minor clades B 1
and B2, sister to B. barbata (see Figure 5.14). The variation observed in B. barbata
is consistent across the global geographic range. I have not seen the TYPE specimen
for B. barbata subsp. pulchella, but the representative samples from India included in
this study (Appendix 1) do not differ greatly from the other sampled specimens (as
defined in phenetic analyses). To assess the limits of the species and subspecies, it is
recommended that B. barbata subsp. pulchella be compared with a wider sample.
182
High levels of homoplasy for many of the Bulbostylis sampled could explain the
lack of overall terminal branch support within the cladistic analyses of this chapter.
In general, species of Bulbostylis form a monophyletic group sister to the clade of
main Fimbristylis species (Fl) and was in direct contrast to placement of Bulbostylis
in Chapters 3 and 4, where the two species of Bulbostylis (B. barbata and B. dellsa)
were nested within the same Fimbristylis species that form the Fl clade of this
analysis. The consistency of the embryo type and vegetative anatomy appears to have
been important in stabilising the results, despite extensive homoplasy across many of
the morphological characters.
The presence and development of the second and third primordial leaves in the
embryo could provide a strong character for grouping species into sections if these
structures could be viewed in all the sampled embryos. The sampled species of
Bulbostylis with larger nuts (B. hispidula, B. humi/is) had larger embryos and a
prominent second primordial leaf. Bulbostylis pi/osa falls into this category, having
the largest embryo sampled in Van der Veken's (1965) study of BulbosZvlis, and was
shown to have the second primordial leaf well-developed and almost as large as the
first leaf. Bulbostylis breviculmis (a sYnonYm of B. striatella) was shown to have a
poorly developed second leaf (Van der Veken 1965), differing from the embryo
observed in B. humilis within this study. These differences question the SYnonon1Y of
B. breviculmis, or the consistency, and therefore usefulness as a character, of the
development of the second primordial leaf.
The smaller embryo (although larger than the other Australian species), and nut
size in general, plus the less developed second leaf in the Australian B. pyriformis
contribute to maintaining species level status separate from B. hispidula. Clearly
there are general similarities in nut shape, epidermal patterning (see Figure 5.19), and
183
synflorescence morphology between specimens of B. pyriformis and the B. hispidllla;
however, a more comprehensive study is needed to assess the broad similarities
between all entities of the B. hispidula complex and B. pyriformis.
Those species with the smaller embryos, where the primordial shoot and root is
less prominent, are mostly grouped in the B 1 clade (i.e. B. densa group, B. puberula,
and B. capillaris). If the inner organs could be scored, however, the uncertainty with
the placement of taxa that currently fall into the B2 clade may be resolved. These
taxa, B. sp. aff. barbata, B. sp. aff. burbidgeae, B. sp. aff. puberula and
B. burbidgeae, seem 'misplaced' due to general embryo morphology and nut
characters. In samples with an abundance of fruits, sectioning embryos elnbedded in
paraffin \vax, would allow the scoring of the internal organs to assess specific
groups.
Prophyllate spikelets, seen within the synflorescence for B. barbata and B. sp. aff.
barbata, or prophyllate buds that were present in most of the Bulbostylis samples in
the phenetic study, and as reported by Guaglianone (1970), have also been described
for some species of Schoenoplectus (e.g. S. calfornicus) and Rhynchospora (e.g.
R. corymbosa, R. brownii) (Kukkonen 1986; Vegetti 2003). Extending the sample
across species in Bulbostylis is recommended to investigate the usefulness of the
character more thoroughly.
Spikelets found at the base of the plant have been recorded previously for
B. humilis (syn. B. striatella), B. heterostachya Cherm. (Chermezon 1929),
B. glaberrima Kilk. (Haines 1971), B. basilis Fosberg, B. schaJJneri (Boeck.)
C.B.Clarke, B. sphaerocarpa (Boeck.) C.B.Clarke (Fosberg 1977), and B. funckii
(Steud.) C.B.Clarke (Goetghebeur and Grager 1993). Amphicarpy occurs across a
184
number of genera and has been reported in Trianoptiles i.e. T capensis (Steud.)
Harv., T solitaria (C.B.Clarke) Levyns (Levyns 1943; Haines and Lye 1977),
Schoenoplectus i.e. S. erectus (Pair.) Palla ex J.Raynal subsp. raynalii (Schuyl,er)
Lye, S. lateriflorus (J.F.Gmel.) Lye subsp. lateriflorus, S. microglumis Lye,
S. articulatus (L.) Palla, S. senegalensis (Hochst. ex A.Rich.) Palla, S. leucanthus
(Boeck.) J.Raynal, S. proximus (Steud.) J.Rayna1, Eleocharis i.e. E. minima Kunth
(Browning 1992), E. caespitosissima Baker (Bruhl 1994), and Crosslandia (see
Chapter 3). The African material of B. densa (baffd2: bde 16, 17, 18) has classic
amphicarpic nuts, and when combined with the generally larger anther length,
fimbriolate glume margins of the mostly mucronate glumes, and the inllorescence of
mainly solitary or reduced anthelodia (anthela of 2-3 rays), forms a group separate to
the other B. densa OTUs. It is necessary to compare the African amphicarpic
material more broadly with other species of the B. densa complex as per Haines and
(1983), and Gordon-Gray (1995) to fully assess the species boundaries.
The variation in floret sex of the radical spikelets seen in B. humiliswas reported
as common among species of Bulhastylis by Chermezon (1929). Bisexual or female
florets nlay occur in the radical spikelets, and the stamen number may be variable
compared to the more consistent numbers in aerial spikelets.
Spikelets that may be present near the base of the plant as well as the aerial
spikelets in B. barbata and B. turbinata do not exhibit amphicarpic features, and are
tenned subradical by Wilson (1980). None of the examined overseas specimens of
B. barbata have subradical spike1ets, the presence of which is a likely result of the
extreme environmental conditions in which they grow. Most of the specimens that
exhibit greatly reduced culms have been collected from the dry desert regions of
central and north-western Australia (see Appendix 1). Subradical spikelets usually
185
produce fruit earlier than aerial spikelets, but this could be due to the later
developlnent of aerial spikelets once conditions are more favourable. Experiments
would be needed to test the effects of harsh environments on the production of
subradical spikelets in Bulbostylis barbata. Further study could concentrate more
specifically on the variation within B. barbata, by expanding the sample size and
including molecular data within the study to fully explore the presence of subradical
spikelets.
RaYnal (1976) used the potential to develop amphicarpy in species of
Schoenoplectus to redefine Schoenoplectus section Supini (Cherm.) J.RaYnal.
Assessing all the species currently assigned to Bulbostylis could provide similar
results, when used in conjunction with data from embryo morphology and anatomy,
and general vegetative morphology. Using Clarke's (1908) classification of
Bulbostylis, all the species known then to be amphicarpic, are found in sections I and
II.
The putative new species Bulbostylis sp. aff. barbata can now be named using the
results of the phenetic and cladistic data as support. A distinctly smaller nut, di fferent
from all other Australian species, and the hairy glume margins of the almost hyaline
glumes, supports the recognition of this new species, known only from K.akadu
National Park, Northern Territory.
Given the limited sample size of the unknown identities, B. sp. aff. densa 1, bt9,
btl 0, B. sp. aff. puberula and B. sp. aff. burbidgeae, and the broad variation within
the Bulbostylis densa and B. hispidula groups, it seems necessary to explore the
limits of taxa in these groups more broadly by increasing the sample size and the
nunlber of species in subsequent analyses. Comparing the unknowns to other
overseas species is also necessary to exclude the possibility of extended ranges,
possibly introduced into Australia via human movement.
A new combination is now provisionally put forward prior to valid publication.
Nomenclature of Bulbostylis in Australia
Genus: Bulbostylis Kunth (nom. cons.) Enumeratio Plantamm 2: 205 (1837)
Bulbostylis capillaris (L.) Kunth ex C.B.Clarke in 1.D. Hooker, Fl. Brit. India
6:652 (1885)
BasionYll1: Scirpus capillaris L.
TYPE:
1. Bulbostylis barbata (RoUb.) C.B.Clarke
BasionYll1: Scirpus barbatus Rottb.
B. eustachyii Eardley
2. Blilbostylis bllrbidgeae K.L. Wilson
3. Bulbostylis dellsa (Wall.) Hand.-Mazz.
BasionYll1: Scirpus densus Wall.
Bulbostylis capillaris var. trifida (Nees) C.B.Clarke
4. Blilbostylis humilis (Kunth) C.B.Clarke
BasionYll1: Isolepis humilis Kunth
Fimbristylis arenaria Nees
Isolepis breviculmis Kunth
Scirpus arenarius (Nees) Boeck.
Bulbostylis breviculmis (Kunth) C.B.Clarke
Isolepis humillima Hochst. ex C.B.Clarke
Bulbostylis striatella C.B.Clarke
Abildgaardia humilis (Kunth) Lye
Abildgaardia striatella (C.B.Clarke) Lye
5. Bulbostylis kakadu K.L.Clarke & 1.1.Bmhl sp. nov. ined.
6. Blilbostylis pyriforl11is S.T.Blake
7. Bulbostylis tllrbillata S.T.Blake
Specimens with uncertain species limits, B. sp. aff. densa 1, B. sp. aff. densa 2
(African amphicarpic), B. sp. aff. turbinata 1, B.sp. aff. turbinata 2, B. sp. aff.
burbidgeae, and B. sp. aff. puberula, need to be assessed against a broader sample of
species from Bulbostylis prior to publication.
187
Species ofuncertain standing
Bulbostylis pi/osa (Steud.) Beetle nom. illeg. Leaflets of Westem Botany 4: 45
(1944)
Basionym: Isolepis pi/osa Steud. Type: Hrbr. Drummond IV nr. 360 (nisi schedula
commutata) N. Holi.
The TYPE specimen is not located in Australia, and until the TYPE sheet can be
examined, placement of this taxon within Australian species cannot be determined.
188
Chapter 6
Testing monophyly of the tribe Abildgaardieae I.Jye
Introduction
The aim of this chapter is to test monophyly for the tribe Abildgaardieae Lye by
subjecting representatives of all genera assigned to the tribe by Goetghebeur (1986,
1998) and Bruhl (1995) (Table 6.1) to cladistic analysis. The genera, Abildgaardia
Vahl (the
TYPE
genus), Fimbristylis Vahl (including Tylocarya Nelmes (Kern 1974;
Simpson 1993; Goetghebeur 1998), or Tylocarya treated as a distinct genus
(Goetghebeur 1986; Bruhl 1995), Bulbostylis Kunth, Crosslandia W.Fitzg., Nemum
Desv. ex Ham., and Nelmesia Van der Veken, are the focus of the study.
The general history of the tribe Abildgaardieae was outlined in Chapter 1, as were
problenl areas where disagreement on generic boundaries persists. The position of
Abildgaardia, assigned as a section (or series) of Fimbristylis (Koyama 1961; Kern
1974; Simpson 1993): as Monostachyae Ohwi, or as a genus (Vahl1805; Kral 1971;
Haines and Lye 1983; Goetghebeur 1986; Bruhl 1995; Gordon-Gray 1995;
Goetghebeur 1998), is specifically relevant in this study because of the large number
of species that occur in, and are endemic to, Australia (see Chapter 4). The status of
Tylocarya is either as a monotypic genus (Bruhl 1995), or more generally accepted,
as a species of Fimbristylis, F. nelmesii lKern (Kenl 1974; Simpson 1993;
Goetghebeur 1998).
189
Although Bulbostylis currently has wide acceptance as a genus, separate fronl
Fimbristylis, the distinction is tenuous. I(oyama (1961) classified Bulbostylis as a
subgenus of Fimbristylis, and Lye (in Haines and Lye 1983) placed Bulbostylis as a
subgenus within Abildgaardia, although both authors have since reverted to using
Bulhostylis at the generic rank (Lye 1995; Simpson and Koyama 1998).
The genus Nemum is thought to be close to Bulbostylis due to a similar embryo
type and the presence of long coarse hairs that may be present at the mouth of the
sheath-leafjunction (Raynal 1973); these two genera are otherwise quite different.
Arthrostylis R.Br. and Actinoschoenus Benth. have been variously cOlnbined, as
Arthrostylis (Kunth 1837; Bentham 1861; Thwaites 1864; Kiikenthal 1(44) within
Fimbristylis (Boeckeler 1874; von Mueller 1875; Clarke 1893; Fitzgerald 1918; Ken1
1955; 1974; Latz 1990: recommending placement into Actinoschoenus). Rye (1992),
in her treatment of the Kimberley Flora (Western Australia), placed the unnamed
species into the reinstated genus Actinoschoenus. Trachystylis S.T.Blake has also
been referred to Fimbristylis as F. stradbrokensis (Domin) J.Kern (Ken1 1959).
Species and generic limits for Crosslandia (Chapter 3) and Abildgaardia (Chapter
4), and species limits for Bulbostylis in Australia (Chapter 5) were defined in the
preceding chapters. A comprehensive assessment of species and generic limits for
Bulhostylis and Fimbristylis was not possible because of the large number of species
assigned to each genus, c. 200 and c. 300, respectively (World Checklist of
Monocotyledons 2004). Representative samples from Bulbostylis and Fimbristylis
were included in the cladistic study. To complete the cladistic data set for the tribe
Abildgaardieae, data were collected from species of Nemum and from the monotypic
genera Nelmesia and Tylocarya.
190
Materials and methods
Ingroup
To assess monophyly for the tribe, and therefore the relationships of genera within
the tribe, all terminal taxa previously defined in the Crosslandia, Abildgaardia and
Bulbostylis chapters (3, 4, and 5) were included in the tribal assessment. When
combined with samples from Nemum spadiceum (Lam.) Desv.ex Ham.,
N. megastachyum (Cherm.) J. RaYn., N. equitans (Kuk.) J. RaYn., Nelmesia
melanostachya Van der Veken, Tylocarya cylindrostachya Nelmes (= F. nelmesii),
and selective species of Fimbristylis, the ingroup represented all genera currently
accepted into the tribe Abildgaardieae.
Previous analyses revealed that Fimbristylis is not a monophyletic group (see
Chapters 3, 4, and 5), however, it was not possible to extend the sample species for
this analysis due to time constraints. Representative taxa from Fimbristylis used in
earlier work within this thesis were maintained for cladistic analysis in this chapter.
A total of 52 species across 8 genera (or 7 if Tylocarya is excluded as a separate
genus) fOlmed the basis for the final cladistic study (Table 6.1, see also Appendix I
for specimen details).
Outgroup
Outgroup taxa used to polarise data were unchanged (Table 6.1; see also Appendix
1). The taxa comprised Arthrostylis aphylla, provisional Actinoschoenus compositus,
Trachystylis stradbrokensis from the provisional Arthrostylideae (Goetghebeur 1986;
Bruhl 1995) or Schoeneae (Goetghebeur 1998), plus Schoenoplectus
tabernaemontani (C.C.Gme!.) Palla (= S. validus), Schoenoplectiella lateriflora
Table 6.1 Taxa included in cladistic analysis to assess monophyly of the tribe Abildgaardieae.
Species from Crosslandia, Abildgaardia, and Australian species of Bulbostylis included here were
defined in Chapter 3, 4, and 5 respectively. See Appendix 1 for specimen details.
Taxa
Ingroup
Abildgaardia macrantha (provisional)
Abildgaardia mexicana
Abildgaardia odontocarpa (provisional)
Abildgaardia ovata
Abildgaardia oxystachya (provisional)
Abildgaardia pachyptera (provisional)
Abildgaardia schoenoides
Abildgaardia triflora
Bulbostylis barbata
Bulbostylis sp. aff. barbata
Bulbostylis burbidgeae
Bulbostylis sp. aff burbidgeae
Bulbostylis capillaris
Bulbostylis densa
Bulbostylis sp. aff. densa 1
Bulbostylis sp. aff. densa 2
Bulbostylis hispidula subsp. pyriformis
Bulbostylis hispidula subsp. senegalensis
Bulbostylis humilis
Bulbostylis puberula
Bulbostylis sp. aff. puberula
Bulbostylis pyriformis
Bulbostylis turbinata
Bulbostylis sp. aff. turbinata 1
Bulbostylis sp. aff. turbinata 2
Crosslandia anthelata (provisional)
Crosslandia setifolia
Crosslandia spiralis (provisional)
Crosslandia vaginata (provisional)
Fimbristylis bahiensis
Fimbristylis blakei
Fimbristylis cinnamometorum
Fimbrisrylis depauperata
Fimbristylisfimbristyloides
Fimbristylis furva
Fimbristylis hygrophila
Fimbristylis microcmya
Fimbristylis schultzii
Fimbristylis sp L.
Fimbristylis variegata
Nelmesia melanostachya
Nemum equitans
Nemum megastachyum
Nemum spadiceum
Tylocarya cylindrostachya
Outgroup
Actinoschoenus compositus (provisional)
aイエィ ッウセケャゥ
aphylla
Schoenoplectiella laevis
Schoenoplectiella lateriflora
Schoenoplectus tabernaemontani
Trachystylis stradbrokensis
No. specimens
sampled
10
5
2
13
13
11
12
4
20
7
5
1
4
15
2
3
3
2
3
7
1
8
8
1
1
5
18
3
14
4
2
5
2
4
2
2
2
2
2
1
1
2
2
4
1
4
4
5
5
3
7
192
(J.F.Gmel.) Lye (= Schoenoplectus lateriflorus) and Schoenoplectiella laevis
(S.T.Blake) Lye (= Schoenoplectus laevis) in the tribe Scirpeae (Bruhl 1995) or
Fuireneae, (Goetghebeur 1986, 1998) depending on the system of classification
accepted.
Characters and homology
Additional characters were added to the cladistic data set when species from
Nemum, Nelmesia and Tylocarya were sampled. These new characters were mainly
associated with the spikelets, i.e. persistence of glumes to the rachilla in species of
Nemum, presence of an intraspicular prophyll in Nelmesia, and variation in
anatomical attributes (Appendix 2).
Embryo morphology and anatomy
Most specimens on loan for Nemum and Nelmesia (ISOTYPE!) could not be
sanlpled for characters from leafblade and culm anatomy, or embryo morphology
due to the limited amount of material available; missing data (mainly
embryographical and anatomical data) were obtained from the literature (Van der
Veken 1965; Metcalfe 1971; Rayna11973; Goetghebeur 1986).
Prepared slides for leaf blade and culm anatomy were available for Tylocarya frOlTI
a previous study by Bruhl (1990), however, embryo morphology for Tylocarya was
obtained from Van der Veken (1965) and Goetghebeur (1986). Only material that
was already loose on the sheets was used to obtain floret measurements, and for
embryo morphology and SEM treatment (not TYPE sheets), ifneeded. Sampling
across the taxa allowed leafblade and culm anatomy to be compared where possible.
Scanning electron microscopy of the nut epidermis enabled the comparison of nut
193
characters, especially the micromorphological attributes e.g. epidermal cell shape
and type 0 f protuberance, between taxa.
Analyses
Data for 55 terminal taxa and 152 characters were subjected to parsimony
analsysis within PAUP* using heuristic techniques (hsearch swap=TBR addseq=rand
nreps= 1000 hold=5 multrees=yes). Branch support was assessed using Bootstrap
analysis (1000 bootstrap replications), as the computational time required to calculate
the Bremer support indices past 3 extra tree length steps was protracted, even when
limiting the addition-sequence replications to 10. Characters were traced in
MacClade and the most relevant characters presented in the cladogram.
Results
Cladistic analysis
A. heuristic search produced 91 most parsimonious trees (Tree length=1482,
CI==0.3947, HI=0.6053, RI=0.5814, RC=0.2295). Tree 1 was one of the retrieved
trees with similar topology to the strict consensus, and was selected to show branch
support and character/branch associations.
Taxa from the' Arthrostylideae', used with Schoenoplectus (including
Schoenoplectiella) in the outgroup for Chapters 3, 4, and 5, violated the assunlption
for monophyly of the ingroup in this analysis. All taxa from the ingroup, plus
members of the provisional Arthrostylideae, formed a broad clade sister to the
outgroup species of Schoenoplectus, with strong branch support indicated
(Bootstrap=87%) (Figure 6.1). Arthrostylis aphylla and Actinoschoenus compositlls
.
0
Schoenoplectu セ[
tabernaemontani
1' 109-2
III'
13-3; 28-2
44-1; 73-3
151-5; 152-12
11 11 -2; 74-3; 123-1,2
90
II
1 11 25-1; 73-4
II
Wセ R
II
F.mbristylis depauperata
1 11 11-2; 13-2
II
;j
0
...
'-
C)
;j
0
Nemum megastachyum
88 •
151-4
C
Q.
SchoenoplectiE lIa
lateriflora
Schoenoplectiella
laevis
Nemum spadiceum
"'11
111111
11I11
Tylocarya cylindrostachya
111122-1,2; 152-1
I
F1
III
Bulbostylis hurnilis
11119-1,2; 74-3; 123-1
II
Bulbostylis barbata
セ
"19-1,2
74-1,2
Bulbostylis sp. aft. turbinata 2
11111111
'19-2
セB iQセ
81
.......· .
: 1
··.
·"0.. ....
:
Bulbostylis pyr,formis
19-2
•
0
•
0
Bulbostylis sp. aft. puberula
·.0
.. "r
..
'
セV[YQ I:
.!
Bulbostylis turlJinata
174 -12
o
:
....
Bulbostylis sp. aft. turbinata 1
119-2
25-1
86-2
100-2,5
l117-5
8
122-1
98-2
151-3
152-4
28-1
Bulbostylis kakadu
Bulbostylis burbidgeae
74-12
A
I/)
0
.0
:i
B. hispidula sutsp. senegalensis
122-2
.1 74 -4
(Xl
Bulbostylis sp. <1ft. densa 2
···1
............i :.1
19-1
Z
B. hispidula subsp. pyriformis
: セiᄋ 19-2
.... . ·1
Nセ
.!!!
Bulbostylis sp. aft. burbidgeae
19-2
0
Bulbostylis densa
Bulbostylis sp.
0
: 641
3ft.
densa 1
Bulbostylis capillaris
B. puberula
Fimbristylis fimlJristyloides
II
11
Fimbristylis sctlultzii
151-1
152-2
III
セ
12lt
SMRUQ セ
Fimbristylis furva
Fimbristylis dis ticha
79 111
QiRLセM[
ュゥLセイッ」。 ケ
Fimbristylis. bahiensis
152-1
セGQSM [
151-7
Firnbristylis
"11-2; 13-2
II
I
F2
Flmbristylis cinnamometorum
13-2; 73-2; 152-2,6
5-L
Fimbristylis variegata
152-6
M
6€
Actinoschoenus compositus
X
Arthrostylis aphylla
X
28-2
73-1; 152-9
III
1 '11 19-2; 123-2; 152-1,7
I
B
53 II
'151-1
Fimbristylis blakei
1123-1
I
III
'25-2
Crosslandia vaginata
11174-1,3; 151-1; 152-6,7,8
II
Abildgaardia macrantha
Abildgaardia ovata
p0.-
I
..
riJ"
Abildgaardia oxystachya
T152 -8
111111
..
100
C2
Fimbristylis hygrophila
28-2
151-1
-
--.
Fimbristylis sp, L
Nelmesia melanostachya
86-6
100-5; 151-3;
152-4
111 109-1' 130-5
61
_
[RMVBUセ
*
Abildgaardia schoenoides
'
Abildgaardia sp. aft. schoenoides
セ
....
86187130 セ
151 b
.. 152 17
Ifo
64-1
Abildgaardia odontocarpa
L.-4
1152-11
I
rT137-2
55 •.
..
'137-3
-'152-11
Abildgaardia mexicana
QLセANiZャMエゥェ
151-1
1526
14
I
Nemum equitans
52-4
セij Av [Q ォゥZ
'"
Abildgaardia sp. aft. pachyptera
III 25-3; 26-2
........
"'74 3
Abildgaardia pachyptera
Abildgaardia triflora
58 .,
1
1152-5
Abildgaardia sp. aft. odontocarpa
Trachystylis stradbrokensis
2-1
Crosslandia s:Jiralis
Crosslandia setifolia
75
Crosslandia anthelata
X
195
were nested with the species F. variegata and the C 4 Fimbristylis bahiensis, and were
sister to the F3-Abildgaardia-Crosslandia clade. Within this clade Trachystylts
(also assigned to the outgroup) was paired with Nemum equitans as a group sister to
the Abildgaardia clade.
Two main clades, A and B, were sister to the clade Fimbristylis depauperata-
Tylocarya cylindrostachya (clade C), and all were sister to the Nemum .spadiceumN. megastachyum clade that showed strong branch support (Bootstrap=87%). The
species of Nemum sampled, therefore, did not form a monophyletic group.
In clade A, all the species of Bulbostylis sampled formed a monophyletic group,
with moderate branch support (Bootstrap=77%), sister to the clade Fimbristylis 2
(F. jimbristyloides, F. schultzii, F. cinnamometorum, F. furva, F. disticJza,
F. microcarya). Clade B, however, contained the smaller F3 clade (F. s1'. L. and F.
blakei - with weak support), which was sister to the Abildgaardia-Crosslandia
clade, which included Fimbristylis hygrophila (= Abildgaardia hygrophila),
Nelmesia, Trachystylis and Nemum equitans.
The previously monophyletic Abildgaardia (Chapter 4) was rendered nonmonophyletic by the placement of Nelmesia melanostachya, which was nested within
the Australian endemics. The Abildgaardia-Nelmesia branch showed weak
Bootstrap support (67%) and the terminal arrangement of taxa did not collapse in the
consensus tree, as indicated by solid lines in Figure 6.1. There are no obvious
sYnapopmorphies that unite Nelmesia and Abildgaardia other than the large stipitate
nut.
1%
Species assigned to Crosslandia in Chapter 3 formed a monophyletic group that
included C. setifolia, and the provisional C. anthelata and C. spiralis, and received
moderate branch support (Bootstrap=73%). Crosslandia vaginata (= Ahildgaardia
vaginata) was basal in the sister group to Crosslandia in this tribal analysis (Figure
6.1) and the Bu/bosty/is treatment (Chapter 5), although there was no internal support
for the placement here.
Fimbristylis hygrophila (= Abildgaardia hygrophila) persists as a sister to the
species of Abildgaardia and Nelmesia melanostachya in the tribal analysis, however,
internal branch support was absent for the placement, as was support for the
placement of Nemum equitans and Trachystylis.
Characters
There are no unambiguous synapomorphies that clearly delimit the internal clades
due to the poor tree resolution caused by the high level of homoplasy within the data
set. Characters from embryo morphology, such as the Schoenoplectus-type embryo
(151-5), the germination pore parallel to the first primordial leaf (154-1), plus
anatomy of the culm (44-1, 51-1), separate the outgroup species of Schoenoplectus
and Schoenoplectiella from the remaining taxa.
The only strongly robust group, other than the outgroup (i.e. Schoenoplectus), is
the clade of Bulbostylis. The synapomorphies for the Bulbostylis clade are: pilose
hairs at the sheath-leafjunction (11-3); leaf vascular bundle number 5 or less (25-1);
a minutely triangular style (117-5); style base persistent on the nut (always separates
from the style) (122-1), although the specimens of B. hispidula may have deciduous
style bases (it falls in tact with the style) (122-2); and Bulbostylis-type embryo (1514).
197
Observations
Injlorescence-synjlorescence
The inflorescence-synflorescence structure is very homoplastic, even when the
structure of the head of sessile spikelets is broken down into different structural
types. The simplest 'head' of3 sessile spikelets (57-4) occurs in Abildgaardia
(A. mexicana), Fimbristylis bahiensis and Bulbostylis (B. humilis) (see Figure 4.12).
A synflorescence head fonned as a compressed spike (multiple primary sessile
coflorescences, of one spikelet per coflorescence, where the tenninal spikelet sits
above the sessile coflorescences (57-5), was observed in Fimbristylis (F. schultzii)
and Actinoschoenus. The multiple branched reduced anthelodia, where all spikelets
are sessile (branching as rays) is highly reduced but discemable under the dissecting
microscope (57-8), is seen in Crosslandia setifolia and Arthrostylis aphylla (see
Figure 3.16). Crosslandia setifolia was the only taxa sampled that produced lateral
heads (see Figure 3.17), where a primary coflorescence has developed into a
secondary main florescence of a head of sessile spikelets. The 'prophyllar' head is a
combination of the multiple branched reduced anthelodium and prophyllate
branching from primary and sometimes secondary inflorescence branches (57-10),
(see Figure 5.15 B, C); this type is restricted to Bulbostylis in this study (B. barbata
and B. kakadu). The only other genus in the study with intraprophyllar growth within
the inflorescence is Schoenoplectus-Schoenoplectiella, which differs in structure
through the paniculodium base plan. In Schoenoplectiella laevis, prophylls were not
restricted to the production of spikelets, as some prophylls were fertile, possessing a
solitary nut in the axil without any other bract visible; unique for taxa within the
study. The most common synflorescence type was the reduced anthelodium with a
198
sessile main primary florescence and primary coflorescences supported on rays
(lengthened epipodia) (see Figures 3.14, 4.12, 4.15 A), which was found across most
genera of the ingroup. Florescence ramification (57-7) within the synflorescence was
common in members of Fimbristylis (F. blakei, F. sp. L., F. cinnamometorum; welldeveloped in F. microcarya, F. depauperata, F. furva and sometimes
F. cinnamometorum; and in some species of Bulbostylis (e.g. B. densa, B. pyriformis,
and may be well-developed in B. puberula and B. hispidula subsp. senegalensis).
The solitary spikelet is the simplest of all the inflorescence-synflorescence
structural types, and, within the study, is most common in Abildgaardia (see Figure
4.12 A). The spike in Nelmesia melanostachya superficially resembles a solitary
spikelet, but the intraspicular prophyll within the solitary spike is exceptional for the
tribe (Figure 6.2). The prohpyll is in place where a lateral branch arises as a solitary,
sessile floret (i.e. single floret spikelet); this inflorescence type is not homologous
with the solitary spikelet in Abildgaardia. Despite the difference in the structure of
the inflorescence, Nelmesia was placed with species of Abildgaardia in many of the
trees retrieved, although there was no support for the placement (Figure 6.1).
Florets within all the studied taxa comply with the basic scirpoid floral
arrangement: being tetracyclic, 2-3 carpels, 1-3 stamens, 0-6 perianth bristles
enclosed by a glume (floral bract) (Vrijdaghs et a1. 2005). Perianth was absent frOlTI
all the sampled ingroup taxa, with the exception of one collection of Abildgaardia
schoenoides (see Chapter 4). Perianth may be present in SchoenoplectusSchoenoplectiella as bristles with retrorse barbs; Schoenoplectus tabernaemontani (=
S. validus) has three perianth bristles present (see Figure 6.8 A). The unique perianth
in the Abildgaardia schoenoides collection differed in having 2 perianth bristles with
antrorse barbs (see Figure 4.13).
Figure 6.2 Nelmesia melanostachya ISOTYPE showing general habit, including solitary spikes
where the lateral spikelet is reduced to a single floret. The insert shows a single floreted spikelet
bearing a mature nut; the large prophyll is obvious and sits between the nut and the rachis. Scale
bar=2 mm. See Appendix 1 for specimen details.
200
Nut epidermal pattern
The nut epidennis varies in the shape and orientation of epidennal cells, or
sculpturing from single silica bodies (puncticulate to granulate), groups of multiple
raised cells fonning various shapes (turbercules), cells raised in ridges that may be
broken or continuous (rugose), raised cell walls (reticulate), or cells with a sunken
lun1en (pitted), and is distinct at the species level. There is some consistency in the
epidennal cell size and shape as seen in the group Abildgaardia - excluding
Nelmesia, (see Chapter 4), where epidennal cells are distinctly rounded (A. ovata,
A. oxystachya, A. schoenoides, A. sp. aff. schoenoides, A. odontocarpa, A. sp. aff.
odontocarpa) or barely hexagonal in shape (A. pachyptera, A. macrantha, A. triflora
and A. mexicana). Large tubercules are common across the species and vary in size
and shape, and occasionally may be few or absent in A. pachyptera, A. macrantha
and A. triflora. Cell walls are not sinuose.
The Bulbostylis species sampled may have nut epidennal cells that are
isodiametric or longitudinally rectangular, and barely rectangular to linearly
rectangular in shape (see Chapter 5). Cellular protuberances may be absent (e.g.
B. barbata, B. kakadu) or individual cells may have a central raised silica body
producing a puncticulate or granulate surface, depending on the size of the silica
body (e.g. B. densa, B. burbidgeae, B. turbinata). Alternatively, the longitudinal
rectangular cells are raised to some extent, giving degrees of rugose patterning as
transverse wrinkles that may be continuous or broken (e.g. B. puberula,
B. pyriformis, B. hispidula). The cell walls are sinuose to some extent (finely-distinctly) in all but B. humilis.
201
Even in the limited Fimbristylis sample the variation in the nut epidetmis is
evident. Epidermal cells vary from isodiametric hexagonal cells to barely circular,
longitudinally rectangular, or transversely rectangular; cell walls may be straight,
barely sinuous to sinuous. Protuberances, as single raised cells to multiple raised
cells with various distribution patterns over the surface, occur frequently across the
species sampled. Nut epidermal features do not seem to influence the group
arrangement for the species of Fimbristylis F2 (Figures 6.3-4), although the poor
resolution in the tree topology could mask the usefulness of the character at the
sectional level. There are no similarities in the nut epidermal sculpturing for the
paired Tylocarya and Fimbristylis depauperata (Figure 6.5). The nut in Tylocarya is
smooth, with hexagonal shaped epidermal cells that have strongly sinuose walls, and
contrasts with the striated epidermal cells in F. depauperata.
The nut epidermis in Nemum spadiceum, N. megastachyum and N. equitans is
distinct in the completely smooth nut surface (Figure 6.6) that is lustrous and
coloured black, dark brown or grey-brown. Taxa from the Arthrostylideae
(Arthrostylis, Actinoschoenus and Trachystylis) that fall with the ingroup in this
analysis have variable nut characters (Figure 6.7). The most striking characters are
the bulbous base of the nuts in the provisional Actinoschoenus compositus (=
Fimbris(ylis composita) (Figure 6.7 C), and the minutely papillose epidermis (Figure
6.7 D). Some species of Fimbristylis have an external gynophore that is brown and
attached at the base of the nut (e.g. F. depauperata, F. fimbristyloides, F. schultzii
and F. bahiensis), however, none is as large as the brown spongy structure seen in
Actinoschoenus compositus. The papillose nut epidermis also in A. compositus has
not been seen among any of the other taxa studied.
Figure 6.3 Scanning electron micrographs (SEM) showing the variation of nut outline and
epidermal sculpturing in some species of Fimbristylis. A. Nut for F. furva (ff2) and B. epidermal
sculpturing at higher magnification, with cells irregularly longitudinal and cell walls that are
distinctly sinuose. C. Nut for F. microcarya (K.L. Clarke 319, L. Little) and D. at higher
magnification, showing epidermal cells that are horizontally elongated and cell walls that are
very fmely sinuose. The waxy covering is not plate-like but continuous over the surface. E. Nut
for F. schultzii (K.L. Clarke 153 et al.) and F. at higher magnification, showing epidermal
sculpturing and hexagonal cell shape. Scale bars A=100 Jlm; C, E=200 Jlm; B, D, F=50 Jlm. See
Appendix 1 for OTU and specimen details.
Figure 6.4 Scanning electron micrographs (SEM) showing the variation of nut outline and
epidermal sculpturing in some species of Fimbristylis. A. Nut for F. disticha (fd2) and
B. epidermal sculpturing at higher magnification, with cells mostly circular and straight cell
walls. C. Nut for F. cinnamometorum (fc5) and D. at higher magnification, showing epidermal
cells that are horizontally elongated and protuberances also elongated horizontally. The waxy
covering is not plate-like but continuous over the surface. E. Nut for F. fimbristyloides (ffi3.)
showing the truncate base and F. at higher magnification, epidermal sculpturing and hexagonal
to circular cell outline. Scale bars A, C, E=100 11m; B, D, F=50 11m. See Appendix 1 for OTU
and specimen details.
Figure 6.5 Scanning electron micrographs (SEM) and light micrograph (LM) showing the
variation of nut outline and epidermal sculpturing in species of Fimbristylis (including
Tylocarya). A. SEM of nut for F. blakei (fbI) and B. epidermal sculpturing at higher
magnification, with cells mostly hexagonal to circular in outline. C. LM of nut for
F. depauperata (K.L. Clarke 305, L. Little) showing the square epidermal cells arranged in
rows. D. SEM of nut for Tylocarya cylindrostachya (A.F.G. Kerr 21294) (= F. nelmesii)
showing the nut outline and E. at higher magnification, the epidermal surface that is smooth
with hexagonal cells with sinuose walls. OTU or collector and collection number are given
in brackets. Scale bars A, D=200 /lm; C=20 /lm; B, E=50 /lm. See Appendix I for OTU and
specimen details.
Figure 6.6 Scanning electron micrographs (SEM) showing the nut outline and epidermal
surface in two species of Nemum. A. Nut for N. spadiceum (E.A. Robinson 4677) and
B. epidermal aurface at higher magnification. C. Nut for N megastachyum (Germain 4420)
and D. at higher magnification. The nuts of both species are similar in outline and the
epidermal surface is lineolate (marked with fme lines), as the cells are barely discemable at
higher magnification. Collector and collection number are given in brackets. Scale bars
A=lOO Jim; C=200Jim; B, D=50 Jim. See Appendix 1 for OTU and specimen details.
Figure 6.7 Scanning electron micrographs (SEM) showing the variation of nut outline and
epidermal sculpturing in species from the provisional tribe Arthrostylideae. A. Nut for
Arthrostylis aphylla (G.N Batianoff10089) and B. epidermal sculpturing at higher
magnification, with cells circular in outline. C. Nut for provisional Actinoschoenus composita
(K.L. Clarke 178 et al.) and D. at higher magnification, showing the small papillae that cover
the nut surface, which are unique among taxa within the study. E. Nut for Trachystylis
stradbrokensis (E.J Thompson 78) showing the nut outline and E. at higher magnification, the
epidermal cell pattern is reticulate-foveate. Collector and collection number are given in
B, D, F=50 Jlm. See Appendix 1 for specimen details.
brackets. Scale bars A, C, E=500 セュ[
Figure 6.8 Scanning electron micrographs (SEM) showing the variation in nut outline and
epidermal sculpturing for outgroup species of Schoenoplectus and Schoenoplectiella. A. Nut
for Schoenoplectus tabernaemontani. (K.L. Wilson 4278) and B. epidermal surface at higher
magnification, with cells narrowly elongated in outline. Perianth with retrorse barbs occur in
this species. C. Schoenoplectiella lateriflora (P.K. Latz 3761) showing the rugose nut surface
and D. the vertically linear epidermal cells at higher magnification. E. Nut for S. laevis (P.M
Milthorpe 1777A, G. M Cunningham) showing the smooth epidermis and F. at higher
magnification. Collector and collection number are given in brackets. Scale bars A=500 セ[
C, E=200 Jim; B, D, F=50 Jim. See Appendix 1 for OTU and specimen details.
208
In the outgroup species sampled for Schoenoplectus and Schoenoplectiella, the nut
shows variable nut sculpturing (Figure 6.8). Schoenoplectus tabernaemontani and
Schoenoplectiella laevis have smooth nuts, while the nuts in S. lateriflora are tightly
rugose. All nuts for the three species have vertically linear cells, which are raised in
S. lateriflora.
Embryo
In species of Fimbristylis, the Fimbristylis-type embryo, although consistent in the
orientation of the primordial shoot and root, is variable in size and shape across the
species sampled (Figure 6.9). Tylocarya cylindrostachya (or fゥュ「イ ウセカャゥ
nelmesia)
was shown to have a variant of the Fimbristylis-type embryo (Goetghebeur 1986)
that is a sY11apomorphy for the Tylocarya-F. depauperata clade (Figure 6.1). Species
of Crosslandia (including C. vaginata) also share the Fimbristylis-type embryo and
are placed separate to species currently assigned to Fimbristylis (see also Chapter 3).
The embryo type in species of Nemum is not typical and varies between the
Abildgaardia-type and Bulbostylis-type (Figure 6.10) (see also Chapters 4 and 5).
The primordial shoot and root are of roughly equal size, or the shoot may be slightly
larger (as in the Abildgaardia-type); the embryo size itself is closer to the
Bulbostylis-type. In Nemum equitans the embryo is similar in outline to the
Bulbostylis-type and is trigonous from the top view of the embryo (Figure 6.10),
although the root is not prominent and the second leaf not detectable, possibly
obscured by the cellular contents surrounding the organs. The elliptic rather than
rounded or trigonous outline in the top view of the embryo in sampled Ii. spadiceum
and N. megastachyum specimens coincides with the di-stigmatic style of both
species; Nemum equitans has tri-stigmatic styles. The variation of the eJnbryo
209
features seen in Nemum was reflected in the analysis, as Nemum equitans and
Trachystylis stradbrokensis fonned a minor clade, well removed from the other
species of Nemum, despite having different embryo types - the Nemum-type and
Carex-type (Figure 6.10) respectively.
Nelmesia melanostachya is shown to have a variation of the Abildgaardia-type
embryo (Van der Veken 1965; Goetghebeur 1986) (Figure 6.10, see also Chapter 4),
where the primordial shoot is basal and larger than the parallel root. The Inain
difference between the Nelmesia-type and Abildgaardia-type embryos is the size,
although the embryo of Nelmesia in this study has been extrapolated from Van der
Veken (1965) and Goetghebeur (1986) and, therefore, may not be a true
representation of the embryo size. The Nelmesia-type embryo is an autapomorphy
for Nelmesia and disrupts the Abildgaardia-type embryo sYnapomorphy for the
species otherwise grouped as Abildgaardia (see also Chapter 4).
The Bulbostylis-type embryo was one of the sYnapomorphies for the Bulbostylis
clade (also see Chapter 5), in contrast to the Fimbristylis-type embryo, which was
homplastic across clades C, A, and B (Figure 6.1) and variable across the sampled
taxa.
Taxa in the Arthrostylis-Actinoschoenus s.l. clade share the Schoenus-type
embryo (Figure 6.10) as a sYnapomorphy on the internal branch that unites the four
taxa (Figure 6.1).
A
F
G
H
Figure 6.9 Light micrographs of whole cleared embryos showing the variation in shape and size for
some species assigned to Fimbristylis, plus schematic embryos for Tylocarya and Nelmesia.
Fimbristylis depauperata (KI. Clarke 305, I. Little) A. side view, B. frontal view, and C. top view of
shoot with the second primordial leaf in view (thin arrow) directly behind the first leaf. D. F shultzii
(KI. Clarke 108 et al.), E. Ffurva (KI. Clarke 210 et al.), F. F disticha (fdl) and
G. F cinnamometorum (fc2) share the Fimbristylis-type embryo. H. The embryo for Tyolocarya is a
variant of the Fimbristylis-type. I. In Nelmesia the embryo is a variation of the Abildgaardia- and
Bulbostylis-types. Scale bars=lOO Nュセ
Solid arrow=root, open arrow=shoot, thin arrow=second
primordial leaf. OTU label or collector and collection number are given in brackets. See Appendix 1
for specimen details. Embryo schematics H and I are adapted from Vander Veken (1965) and
Goetghebeur (1986).
Figure 6.10 Light micrographs of whole cleared embryos for some species from the outgroup
used in cladistic analyses: Actinschoenus, Trachystylis and Schoenoplectiella.
A. Actinoschoenus composita (K.L. Clarke 178 et al.) has the Schoenus-type embryo
(indicated by the arrows) with the embryo outline wide and saucer shaped in side view,
B. Trachystylis stradbrokensis (S T Blake 13201) has a Carex-type embryo with a widened
cotyledon. C. The distinctive Schoenoplectus-type embryo is shown for Schoenoplectiella
laevis (K.L. Wilson 8041 et al.), where the cotyledon extends past the primordial shoot, and
D. at higher magnification showing the genn pore parallel to the first primordial leaf. Scale
Solid arrow=root, open arrow=shoot. Collector and collection number are
bars=100 セュN
given in brackets. See Appendix 1 for specimen details.
212
Anatomy
Leafblade and culm anatomy show 47 of the 55 taxa sampled, all from the
ingroup, share the C 4 fimbristyloid photosynthetic pathway. The C 3 photosynthetic
pathway arises several times amid the ingroup in the tribal analysis, and taxa with C 3
anatomy (Arthrostylis aphylla, Actinoschoenus compositus, Fimbristylis variegata
and Trachystylis stradbrokensis (Figure 6.11) are placed with C 4 species
(Fimbristylis bahiensis and Nemum equitans, respectively).
Despite variation in the general shape of transverse sections of leafblade and
culrn, in the number of vascular bundles, the shape and number of sclerenchyma, and
the shape and arrangement of parenchyma among the C 4 species, there was a general
consistency with all having only sclerenchyma strands in leaf blades and culms. The
exception is seen in the sections of Tylocarya (Figure 6.12). Leafblade anatomy in
Tylocarya shows similar structure to some of those seen in Fimbristylis and
Abildgaardia. In addition to the usual abaxial row of sclerenchyma strands that occur
below the epidermis, Tylocarya varies in that adaxial strands of sclerenchyma are
present and associated with the largest vascular bundles (four in this sample,
excluding the usual comer support) (Figure 6.12). Adaxial strands were observed in
only one other species in this study, and that was F. fimbristyloides (Figure 6.12).
Culm anatomy in Tylocarya, however, is distinct from any of the other specimens
sampled; there are many layers of vascular bundles arranged in rough concentric
rings (3 developed and the 4th newly formed), decreasing in size as the newer
bundles develop below the outermost tissue layers. The large dome-shaped bundles
of support sclerenchyma are in direct contact with many of the newest vascular
bundles in the outermost ring, and are clearly girders and not strands. The vascular
213
bundles have extra sclerenchyma support, with an inner cap of approximately six
strands, with son1e extra rows of sc1erenchyma supporting the largest bundles that
have been pushed inwards (Figure 6.12 B).
Nemum spadiceum was the only species of Nemum sampled for anatomy. The
culIn is highly sc1erified, showing an almost continuous undulating band of fibres.
The central pith is absent and prominent air spaces occur between most of the
vascular bundles. The number of vascular bundles arranged in the single ring
corresponds to the crescentiform section of the undulations, with stomata protruding
through and above the sclerenchyma fibres in the narrow fibre regions (Figure 6.12
E). The leaf blade outline in transverse section is almost elliptic, possessing only four
vascular bundles (not presented). The hypodermis is restricted to a couple of cell
layers and is three to four cells wide in the adaxial central region. The leaf margins in
both Nemum and Nelmesia are folded in and joined at the leaf sheath junction (see
Metcalfe 1971).
The transverse culm outline in Arthrostylis aphylla is four-sided. Tannins are
present within the epidermal cells, forming a broken line between the strands of
sclerenchyma. The vascular bundles are found around the perimeter of the culm, and
do not correspond in number to the many small mounds of sclerenchyma strands
(Figure 6.11 A, B).
Actinoschoenus compositus is the only sampled species with sclerenchyma girders
and strands present within the culms. The prominent ribs have equally prominent
thickly V-shaped or crescentiform strands adjacent to the smaller vascular bundles
(Figure 6.11 C, D). The deep channels have twin stomata at the sides near the base of
the channel, and hairs are prominent near the outer margin. The epidermal layer is
A
c
Figure 6.11 Culm and leaf blade transverse sections for some species from the provisional tribe
'Arthrostylideae' selected as outgroup taxa for use in cladistic analyis, showing the typical
outlines, arrangement of sclerenchyma strands per vascular bundle, and C3 anatomy.
A. Arthrostylis aphylla (A. Gunness AGL1965) 4-sided culm and B. at higher magnification,
showing the greater number of small mounded sclerenchyma strands than vascular bundles.
C. Actinoschoenus composita (K.L. Clarke 178 et at.) culm that is regularly, deeply ribbed, with
vascular bundles alternately large and small; at higher magnification D. thin rectangular girders
are associated with the larger bundles and large crescentiform sclerenchyma strands are opposite
the smaller vascular bundles. Twin stomata oppose each other at the base of each channel
formed by the rib (indicated by the arrow heads). E. In Trachystylis stradbokensis (s. T
Blake1320l) the culm is distinctly triangular in outline and F. the leaf blade is subtriangular,
with vascular bundles completely immersed within the chlorenchyma. Scale bars A, C=500 Jim;
B, E, G=20 Jim; D, F=50 Jim. Collector and collection number are given in brackets. See
Appendix 1 for specimen details.
B
G
Figure 6.12 Culm and leaf blade transverse sections for Fimbristylis, Tylocarya, and Nemum.
A. Culm and B. leaf blade sections of Fimbristylis depauperata (K.L. Clarke 305, L. Little).
Tylocarya cylindrostachya (A.G.F. Kerr 21294) (= F. nelmesii) culm sections at C. low
magnification and D. at high magnification, plus leaf blade sections at E. low magnification
and F. at higher magnification, showing the variation and detail of vascularisation. G. Culm
section for Nemum spadiceum (E.A. Robinson 4676) showing the continuous ring of
undulating sclerenchyma. Collector and collection number are given in brackets. See
Appendix 1 for specimen details. Scale bars A, D=20 Jlm; B, E= 500 Jlm; C, F, G=50 Jlm.
216
densely stained with tannins, seen as a continuous line, and has a thick cuticle layer
(Figure 6.11 E).
In Trachystylis, both the leaf and culm are triangular in outline, and both are dense
with tannin-filled cells. Some tannin is deposited in the epidennal tissues in both
culms and leaves, although not so densely. The three vascular bundles of the leaf
blade are completely surrounded by the chlorenchYma (Figure 6.11).
Tannin deposits in the epidennallayer do not occur in any of the other taxa
sampled, including the outgroup, and were only found within the members of the
provisional Arthrostylideae. In all of the other taxa studied, the tannin deposits were
mostly observed within the chlorenchYma tissue.
Discussion
It is not possible to draw substantial conclusions from the main cladistic analysis,
other than that the tribe Abildgaardieae, as it is currently accepted by Bruhl (1995) or
Goetghebeur (1998), does not fonn a monophyletic group. The weak branch support
in earlier analyses (see Chapters 3, 4, 5) for Arthrostylis aphylla, ActinoschoeJlllS
compositus and Trachystylis stradbrokensis with the outgroup clade gave an
indication of the instability of the outgroup. Including Eleocharis within the current
outgroup could resolve this instability, however, the lack of leafblade characters and
the solitary spikelets limit the usefulness of the genus as an outgroup here. Ghamkar
et a1. (2006, in press), in a molecular study of the tribe, found that samples from
Actinoschoenus (Fimbristylis composita Latz), K.L. Clarke et al. 214, K.L. Clarke et
al. 213 (NE and NSW) and Arthrostylis, K.L. Clarke et al. 212, K.L. Clarke et al. 183
(NE and NSW), all collected from Northern Territory, were nested within
Fimbristylis; surprising considering the major ditTerences seen in this study for
217
vegetative anatomy and embryo morphology bet\veen the sampled Actinoschoenus
and Arthrostylis, and the species of Fimbristylis (see Figures 6.9-10 and 6.11-] 2).
Clearly, Arthrostylis, Trachystylis, plus overseas and Australian Actinoschoenlls need
to be assessed more thoroughly to fully determine their position, especially
considering the taxonomic history of these taxa as species of Fimbristylis.
Species of Bulbostylis and Fimbristylis in clade A of this study are all C 4 taxa,
however, any conclusions drawn regarding uniformity of the photosynthetic pathway
across these genera are limited by the small samples sizes used in this study. The C 3
taxa that occur within the predominantly C 4 clade 13 of the Abildgaardia-Crosslandia
group do not show any relative grouping patterns to explain their positioning within
the cladogram. Stock et al. (2004) demonstrated a connection between phylogeny
and geographical distribution in the tribes Cypereae, Scirpeae and Schoeneae; the
sample size for the Abildgaardiaeae was too small to be informative. Schoenoplectus
has both C 3 and C 4 species in the genus (Stock et al. 2004), however, Bruhl and
Wilson (2005, in press) suggest that the C4 S. pulchella sampled by Stock et al. may
be rnisidentified, as all other species of Schoenoplectus are reportedly C 3 . The
species of Shoenoplectus and Schoenoplectiella sampled in this study i.e.
Schoenoplectus tabernaemontani (= S. validus), Schoenoplectiella lateriflora (=
Schoenplectus lateriflorus) and Schoneoplectiella laevis (= Schoenoplectus laevis)
are all C 3 taxa. Alternatively, Schoenoplectus pulchella may be misplaced. The study
by Bruhl and Wilson (2005, in press) reports the presence of C 3 and C 4 species in
Abildgaardia. However, A. hygropyhila is the only C 3 species currently accepted in
the genus, and is misplaced in Abildgaardia (see Chapter 4). Future studies in
Fimbristylis could determine if C 3 species such as Fimbristylis variegata (with a
Schoenus-type embryo and a prior history with Abildgaardia) should be removed
218
frOlTI Fimbristylis that is currently accepted as containing both C 3 and C 4 species
(Bruhl and Wilson 2005, in press).
The placement of Tylocarya with the representative species for the fゥュ「イウセケャゥ
TYPE section, Fimbristylis section Fimbristylis (F. depauperata) supports the current
acceptance of Tylocarya as a species of Fimbristylis (F. nelmesii) (Kern 1974;
Sinlpson 1993; Goetghebeur 1998). The differences in the culm anatomy and the
variation in the Fimbristylis-type embryo need to be resolved against a broader
Fimbristylis sample, especially when these two species have separated from the other
species of Fimbristylis to be sister to most of the ingroup taxa.
The remaining species of Fimbristylis did not conform to Kern's (1974) sections.
The only terminal branch in Fimbristylis with moderate support (Bootstrap=70%)
was the paired F. disticha (section Fuscae) and F. microcarya (section
Trichelostylus), which are clearly classified in separate sections. A similar result was
obtained in the combined trnL-F and ITS regions data sets of Ghamkhar et a1. (2005,
in press), where a larger sample of species of Fimbristylis were included across the
analysis, but sectional groups for the genus were not retrieved. A broader sampling
of the embryos across the species assigned to Fimbristylis could provide more
natural sectional groups, as there were distinct differences between the Fimbristylis
embryos observed in this study and those by Van der Veken (1965) and Goetghebeur
(1986).
Although intraprophyllar buds were observed in the Bulbostylis species studied
(excluding B. striatella), their presence was not captured in the overall
inflorescence-synflorescence structure; the buds can remain dormant and appear to
be absent in some specimens. The difficulty also lies in the amount of material
2\9
available for examination, as some specimens with only buds may be damaged or
destroyed during examination. Nevertheless, the presence or absence 0 f
intraprophyllar buds, or growth, in species assigned to Bulbostylis could be a useful
distinguishing character at the sectional level within Bulbostylis, if not at the higher
rank of genus. There is potential for future work on this.
It is not surprising that Lye (in Haines and Lye 1983) used the appearance of the
embryo to place Bulbostylis as a subgenus of Abildgaardia. The arrangement of the
shoot and root primordia positioned basally, and the well-developed second
primordial leaf are strikingly similar in all species of Abildgaardia and some species
of Bulbostylis (B. hispidula, B. striatella and B. pilosa). However, the overall size of
the embryo in Abildgaardia is consistently larger and the shoot is always more
prominent than the root, while the reverse is true in all of the Bulbostylis species
sampled in this study and those studied by Van der Veken (1965) and Goetghebeur
(1986). The species of Bulbostylis examined by Van der Veken and Goetghebeur
were: B. caespitosa Peter (= B. oritrephes (Rid!.) C.B.Clarke), Fimbristylis cioniana
Savi (= B. cioniana (Savi) Lye, B. coleotricha (Hochst. ex A.Rich) C.B.Clarke, B.
conifera (Nees) Kunth, B. fendleri C.B.Clarke, B. lanifera (Boeck.) Kilk., B. pringlei
(Britt.) Beetle (= B. schaffneri (Boeck.) C.B.Clarke), B. vanderystii Cherm.,
B. melanocephala (Rid!.) C.B.Clarke, and B. oligostachya (Hochst. ex A.Rich.)
C.B.Clarke. The Bulbostylis-type embryo united the species of Bulbostylis, however,
there are many species placed in this genus that require sampling; the embryo size,
and the number and development of primordial leaves have potential for assessing
the sectional limits of the genus.
The perianth bristles observed in Abildgaardia schoenoides (see Chapter 4 and
Figure 6.3) are similar to those in Eleocharis, as both may have antrorse barbs
220
(Dahlgren et al. 1985; Wilson 1993), in contrast to the retrorse barbs in the species of
Schoenoplectus with a perianth (see Figure 6.3). Many genera are composed of
species with or without perianth (e.g. Schoenoplectus, Schoenus and Rhynchospora)
and considering the rarity of the perianth within the tribe, the novel observation
seems merely to be a remnant feature. The fact that the bristles were not welldeveloped in every floret (although many florets had aborted) and that the specin1en
is distinctly grouped with the other samples for A. schoenoides, adds support to the
remnant hypothesis; it is less likely to me that the presence of perianth in this
material is a reversal.
The surprise placement of Nelmesia within the Abildgaardia clade is not such a
surprise when these results are compared to the systematic study of Goetghebeur
(1986), where Nelmesia was placed in the same clade as Abildgaardia. The sample
size of one collection, missing data, and the many autapomorphies associated with
Nelsmesia, could have contributed to the placement. Nelmesia is known only from
the Belgian Congo and placement within the Australian species of Abildgaardia is
not a likely scenario. The intraspicular prophyll (cf. Haines 1967) present in Carex
L., Kobresia Willd., Schoenoxiphium Nees (Snell 1936; Kern 1958; Tilllonen 1998;
Starr et al. 2004), and Lipocarpha R.Br. (Goetghebeur 1998), although, the prophyll
is modified into a utricle in genera of Carieaceae and the mixed floret sex, differs
from the bisexual florets of Nelmesia with the non-modified prophyll. Lateral
branches are consistently one-flowered spikelets in Nelmesia while the taxa shown in
Kern (1958), Timonen (1998) and Starr et al. (2004) may have lateral branches with
varying numbers of florets within the spikelet. Expanding the sample size to include
taxa with features similar to Nelmesia and more collections of the species, could
resolve some of the problems in assessing monophyly.
221
\Vith Nelmesia excluded from the analysis, as occurred in Chapter 4, the species
that form Abildgaardia are monophyletic and in a clade separate to most species of
Fimbristylis, including F. depauperata, the representative for the
TYPE
section,
Fimbristylis section Fimbristylis. Despite the problems within Fimbris(ylis, in the
full analysis, Abildgaardia does not form a clade with the species of Fimbristylis
(excluding F. blakei and F. sp. L (Kimberley Flora). If Abildgaardia is considered as
a section in Fimbristylis, then my study shows that Bulbostylis and Crosslandia
would need to be demoted from the generic rank to the rank of section as well.
Actinoschoenus and Trachystylis would also need to be reassigned as sections of
Fimbristylis, if the results from my analysis are interpreted as sections.
The Carex-type embryo is considered to be closest to the ancestral form
(Goetghebeur 1998), and is unique to Trachystylis in this study. Actinoschoenlls
thouarsii Benth., A. filiformis and A. repens were shown to have the Carex-type
embryo (Van der Veken 1965; Goetghebeur 1986), however Actinoschoenus
compositus and Arthrostylis aphylla (see also Goetghebeur 1986) share the
Schoenus-type embryo. It is worth noting that Trachystylis with the Carex-type
embryo and C 3 anatomy, both considered as ancestral features, was not placed in any
of the basal positions within the clades, but was among taxa with the derived embryo
types and anatomy. I cannot see why Trachystylis stradbrokensis and Nemum
equitans were placed together within the Abildgaardia-Crosslandia clade, other than
the lack of informative characters from the many autapomorphies present in both
taxa, inhibiting assessment of the relationships. Expanding the sample to include
more species of Nemum and Actinoschoenus, plus other taxa from the tribe
Schoeneae Dumort., the alternative valid tribe in which Trachystylis, Arthrostylis and
AClinoschoenus are placed (Goetghebeur 1998), may assist in resolving the lack of
222
monophyly achieved in this analysis. In addition, selecting a broader outgroup
sample to include other closely related genera and so aid optimisation 0 f the tree
(Grandcolas et al. 2004), starting with Eleocharis R.Br., is recommended.
There is still much work to be done to assess the limits of the genera with the
largest number of species, i.e. Fimbristylis and Bulbostylis, and the tTIonotypic genera
that have only minimal collections, i.e. Nelmesia and Tylocarya.
The lack of resolution for relationships in the current study is due in part to
problems at both ends of the sampling spectrum, confounded by high levels of
hOlTIoplasy and the difficulty in defining adequately characters for the cladistic
analyses. Problematic data is not new, and many articles have been written about the
use of characters in cladistic studies in an attempt to work through some of the lack
of cladistic or phylogenetic resolve in analysis (Rieger 1979; Scotland and Williams
1993; Thiele 1993; Donoghue and Ackerly 1997; Poe and Wiens 2000; Wiens 2000;
Desutter-Grandcolas et al. 2005). The next step that could be taken is the merging of
the plant morphology, embryo morphology and anatomical data from this study with
the nlolecular study for the tribe, and then seeing if the combined data set is more
stable.
223
General conclusions
The tribe Abildgaardieae, as currently delimited, does not form a monophyletic
group. The data analysed were obtained from morphology, vegetative anatomy and
en1bryo morphology. Monophyletic groups were retrieved for some genera in
Chapters 3, 4, and 5, where Crosslandia, Abildgaardia, Fimbristylis, and Bulbostylis
fanned the 'ingroup'.
Species of Crosslandia formed a monophyletic group: C. setifolia, C. anthelata
ined., C. spiralis ined. (Fimbristylis spiralis) and C. vaginata ined. (Abildgaardia
vaginata). Crosslandia vaginata, although consistently retrieved, had little support
and in the Bulbostylis analysis (Chapter 5) and the 'whole tribe' analysis (Chapter 6)
was placed as sister to the Crosslandia clade; it is important to note that Crosslandia
vaginata did not fall within the Fimbristylis or Abildgaardia s.s. clades (although it
was placed in the broad Abildgaardia-Crosslandia clade). The variation in embryo
morphology (Fimbristylis-and Schoenus-types) and inflorescence-synflorescence
structure indicates that the sample size needs to be increased to fully define the limits
before validly publishing new combinations. Extending the molecular sample is also
recommended to explore the genetic variability across the geographical range. The
remaining three species of Crosslandia have support for their placement as separate
species within Crosslandia.
There is no evidence to support maintaining Abildgaardia as a section of
Fimbristylis, as the species of Fimbristylis did not form a monophyletic group in any
of the analyses, and Bulbostylis was placed more closely to Fimbristylis than were
the species of Abildgaardia. A well-supported, monophyletic group was formed by
224
species of Abildgaardia in the analyses in Chapters 3, 4, and 5, but not in the final
analysis in Chapter 6, where Nelmesia melanostachya rendered the group nonmonophyletic. Nevertheless, the Abildgaardia clade did not fall within the species of
Fimbristylis, although two species of Fimbristylis (F. blakei and F. sp. L) were
grouped in the same broad clade as Abildgaardia and Crosslandia. Abildgaardia
hygrophila was not supported as a species of Abildgaardia, even though it is placed
near the provisional Crosslandia vaginata and the other species of Abildgaardia.
Similarities between A. hygrophila (= Fimbristylis hygrophila) and species of
Abildgaardia, as defined here, are purely superficial. The fact that this species was
not grouped with any of the species of Fimbristylis begs for further investigation into
the correct placement of this C 3 species bearing a Fimbristylis-type embryo. There
was no support for the inclusion of Abildgaardia baeothryon within Abildgaardia. A
study of the embryo and anatomy of Abildgaardia papillosa is recommended
because of the findings for Fimbristylis bahiensis (= A. baeothryon) in this study and
the affinity between the two species. Broader sampling to capture the variation
between Abildgaardia oxystachya and A. pachyptera to define their limits is needed
prior to publishing. In contrast, species status for the samples A. sp. aff. odontocarpa
and A. sp. aff. pachyptera is merited and descriptions are being prepared. Meanwhile,
detelmining where the name A. schoenoides R.Br. should be applied and if
Fimbristylis squarrulosa (TYPE) is a synonym of A. schoenoides must be sought by
exanlining the A. schoenoides TYPE specimen held at BM. Comparing the TYPES
assigned to both names with the groups retrieved from phenetic analyses is necessary
to determine the correct application of the names.
Species of Bulbostylis formed a monophyletic group that was well supported in
Chapters 5 and 6. Bulbostylis kakadu ined. is a distinct species separate to B. barbata
225
and a description can now be prepared for valid publication.. Any relationship
between the Australian B. pyriformis and the B. hispidula group was inconclus lve. A
more thorough study is recommended to explore fully the relationship between all
the entities of the B. hispidula complex and the Australian B. pyriformis.
No sections from Kern's (1974) classification of Fimbristylis were retrieved in this
study, however, the placement of Tylocarya with Fimbristylis depauperata (from the
TYPE
section, Fimbristylis section Fimbristylis) supports Kern's (1958) decision to
place Tylocarya in Fimbristylis as F. nelmesii.
セャ・ュ「 イウ
of the provisional Arthrostylideae loosely formed the outgroup beside
species of Schoenoplectus and Schoenoplectiella. However, the shift of
Actinoschoenus, Arthrostylis and Trachystylis into the ingroup when Nemum,
Nelmesia and Tylocarya were added, reflected the tenuous support for the outgroup
placement in previous chapters (Chapters 3, 4, and 5). Actinoschoenus and
Arthrostylis require further investigation to explore the species and generic limlts - a
study is currently underway. Expanding the study to include other melllbers of the
Schoeneae, considered close to Actinoschoenus and Arthrostylis, is necessary. Better
tree topology may result by adding these taxa and a broadened sample of species
fronl Fimbristylis and Bulbostylis, where work is also required to assess species and
generic limits.
The tribe Abildgaardieae may need to be reclassified in the near future. Expanding
Abildgaardieae to include Actinoschoenus, Arthrostylis and Trachystylis is one
option, or defining smaller tribal groups where the name Abildgaardieae is applied to
taxa in the broad Abildgaardia-Crosslandia clade, and Fimbristylideae Cherm ex
Raynal reinstated to accommodate taxa in the Fimbristylis-Bulbostylis clade is a
226
second option. However, further cladistic studies to resolve monophyletic groups are
needed.
Combining the morphological, anatomical, and embryographic data from this
study with the molecular data from a sister study by Ghamkar (2004), to assess
congruence and monophyly of the combined data, may move towards resolving
monophyletic groups.
In the tribe Abildgaardieae and the Cyperaceae in general, the search for
monophyletic groups to develop natural systems of classification may well rely on
more collaborative projects, especially where genera with large numbers of species
cover vast areas globally - and all of us with limited resources.
227
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Appendix 1. List of all specimens used in phenetic and/or cladistic analyses. The OTU label
corresponds to the code for each specimen used in phenetic analyses. Herbarium codes (Herb.
Code.) are provided for all sampled specimens followed by the sheet number where available.
Specimens collected in Australia show the state in which they were collected, if collected
overseas then the country or continent is given. Specimens used for Scanning electron
microscopy (SEM=*), embryo morphology (embryo=#) or leaf blade and/or culm anatomy
(anatorny=+) are indicated in the phenetic code column next to the OTU label, if given.
N.T.=Northem Territory, W.A.=Westem Australia, Qld=Queensland, Vic=Victoria.
(JJB)=prepared sections provided by J.J. Bruhl. Names given here are prior to analyses and are
based on Abildgaardia and Tylocarya as genera.
Generic groups studied
and species names
OTU
label
Herb. code State or Collector
Country
Ingroup
Cross/audia W.Fitzg.
Crosslandia anthelata nom. C18
provo Goetgh. ined.
C19#
C20#
C21
C22
C23
Crosslandia setifolia
W.Fitzg.
C1
C2
C3
C4
C5
C6
C7
C8
C9
C10
C11
C12
C13
C14
MEL
1590282
BRI 313723
NE 61439
NSW
452330
NSW
452331
CANE as
CBG
8309188
BRI 313715
MEL
2048535
NSW
452305
MEL
2048435
BRI 313716
NSW
460200
CANB
264103
NE(exNSW
416310)
NSW
303698
NSW
460198
MEL
2048448
BRI 313711
MEL
2048434
NE
N.T.
Dunlop C.R. 6854, Wightman G.
N.T.
N.T.
N.T.
Dunlop C.R. 3446
Bruhl J.J., Hunter Y., Egan J. 1268
Dunlop C.R. 3408
N.T.
Wilson K.L. 5150, Dunlop C.R.
N.T.
Thompson H.S. 403
N.T.
Chippendale G. 1268
W.A.
Poulton G. 5
W.A.
Wilson K.L. 4885
N.T.
Craven L.A. 7928, Whitbread G.
N.T.
N.T.
Blake S.T. 17420
Cowie J.D. ill 4639
W.A.
Pullen R.
N.T.
Wilson K.L. 5260
W.A.
Wilson K.L. 4859
W.A.
Wilson K.L. 4803
N.T.
Lazarides M. 8, Adams L.G.
W.A.
N.T.
Burbidge N. 5703
Dunlop C.R. 6789
W.A.
Clarke K.L. 166, Bruhl J.J., Wilson
K.L.
237
C15
C16
C17
Fimbristylis spiralis R.Br.
Fl#
F2
F3
Abildgaardia vaginata R.Br. Avl
Av2#
Av3#
Av4
Av5
Av6
Av7+
Av8
Av9#
Avl0
Avl1
Av12
Av13
Av14
N.T.
Blake S.T. 16585
W.A.
Van Rijn P.l. 19
N.T.
N.T.
Clarke K.L. 155, Bruhl 1.1., Wilson
K.L., Cowie I.D.
Specht R.L. 235
N.T.
N.T.
Dunlop C.R. 2957
Leach G. 3601, Cowie I.D.
Qld
N.S.W.
Blake S.T. 15540, Webb LJ.
Floyd A.G.F. AGF2205
Qld
N.S.W.
Qld
Brass L.l. 18362
O'Hara 1. 3472 and Coveny R.
Blake S.T. 8598
N.T.
Cowie I.D. 6801
N.T.
N.S.W.
Qld
Brennan K. 2588
Bell D.M.
Forster P.I. PIF9732, Machim P.
Qld
Qld
Forster P.I. PIF16257
Blake S.T. 8222
Qld
Blake S.T. 22499
Qld
Qld
Brass L.l. 1924
Sharpe P.R. 5299 and Bird L.
K 2525
Brazil
Mattes Silva L.A. 394, Ribeiro AJ.,
da S. Brito H.
K2528
Brazil
Almeida de 1esus 1. 1466
CANB
159880
CANB
78686
NE
MEL
2048472
DNA 36442
NSW
422184
BRI300861
NSW
285362
BRI300844
BRI 300865
MEL
2048450
DNA
128735
DNA 72733
NE 54172A
MEL
716909
BRI 601359
MEL
2048451;
BRI300850
CBG
8309188
CANB
506846;
BRI300856
BRI 300848
BRI
549964;
CANB
505073
Abildgaardia Vahl
Abildgaardia baeothryon
A.St.-Hil.
*#+
(all as Fimbristylis bahiensis #+
Steud.)
KEW 2529 Brazil
Brazil
K 2527
Abildgaardia hygrophila
(Gordon-Gray) Lye
Abildgaardia macrantha
(Boeck.) comb. provo
Goetgh.
*
NE 71642
#+
NU
AmI
CANB
280535
South
Africa
South
Africa
W.A.
Bento Pickel D. 3140
Mori S. 10389, dos Santos S., White
I.
Ward C.l. 2794
Tinley K.L. 307
Hartley T.G. 14405
238
NSW
454230
Am3* NSW
452333
BRI301881
Am4
MBA
Am5
NSW
Am6
338994
Am7*# BRI437357
DNA
Am8
122604
CANB
Am9
242626
Aml0 NE
N.T.
Cowie J.D. 6202 & Booth R.
N.T.
Wilson K.L. 4971
N.T.
Qld
Qld
Qld
N.T.
Dunlop C.R. 4102
Clarkson J. 8324
Wilson K.L. 8073, Clarkson J., Jacobs
S.W.L.
Clarkson J. 6624
Cowie J.D. 5260, Taylor S.
N.T.
Dunlop C.R. 3453
N.T.
*#+
Mexico
Clarke K.L. 249, Bruhl J.1., Wilson
K.L., Cowie J.D.
Pringle C.G. 3127
Mexico
Breedlove D.E. 54895, Davidse G.
Mexico
Kral R. 25115
Mexico
Pringle C.G. 9294
Mexico
GomilezS.1116
Mexico
Arsene G.
Qld
Blake S.T. 13582 (HOLOTYPE)
Am2
Abildgaardia mexicana
(Palla) Kral
Fimbristylis odontocarpa
S.T.Blake
F. sp. aff. odontocarpa
Abildgaardia ovata
(Burm.f.)
Kral
MEL
268631
MO
3524080
MO
1917406
MO
3058759
MO
3632024
MEL
2050681
Aodl BRI341121
MEL
2048459
MEL
2048468
Aod2* BRI574804
#+
Aaffod BRI 302127
*#+
Aovl BRI 476426
(ISOTYPE)
(ISOTYPE)
Qld
Turpin G.P., Thompson EJ.
W.A.
Carey J.
Qld
Specht R.L. 408, Reeves R.D.
Aov2* BRI457532 Qld
Aov3 MEL
Qld
2048512
N.S.W.
Aov4 NSW
468324
Aov5 NSW 87198 N.S.W.
Aov6 NSW
N.S.W.
468325
N.S.W.
Aov7 MEL
2048525
Aov8* NSW
N.S.W.
468326
Aov9 BRI384713 Qld
Aov10 BRI 591266 Qld
Aovll NE
Qld
*
Batianoff G.N. 11056
O'Shanessy P.A. 1656
Wilson K.L. 5818
Johnson L.A.S.
Rodd A.N. 2277
Mueller F.
Rodd A.N. 2434
StanleyT.8019
Neldner V.J. 3905
Clarke K.L. 99, Bruhl J.J., Wilson
K.L.
239
A. ovata (cont'd)
Abildgaardia oxystachya
(F.Mue:ll.) comb. provo
Goetgh.
CANB
MEL
2050881
MEL
2050886
MEL
2050863
MEL
2050864
Aox1* NSW
452336
Aox2
Aox3*
#
Aox4+
Aox5*
+
Aox6+
Aox7
Aox8
Aox9
Aox10
Aox11
Aox12
+
Abildgaardia pachyptera
(S.T.Blake) comb. provo
Goetgh.
Aox13
ApI
Ap2
Ap3
Ap4
Ap5
Ap6
Ap7
Ap9+
A. sp. aff. pachyptera
Abildgaardia schoenoides
R.Br.
Timor
India
Wiriadinata H. 449
Mueller F.
U.S.A.
Medley Wood J. 4918
India
Thomson G.
India
Unknown
W.A.
Latz P.K. 4038
BRI302157 Qld
NE 61223 N.T.
Blake S.T. 15725, Webb L.J.
Bruhl J.J. 1252
NE
NSW
452337
NE
N.T.
N.T.
Latz P.K. 8667
Wilson K.L. 5369
W.A.
BRI336290
BRI 386514
CANB
280532
CANB
353598
DNA
123491
CANB
505055;
BRI203628
DNA 21755
NE 61427A
MEL
1615159
MEL
252996
NSW
452342
NSW
452335
NSW
452334
MEL
2048462
NE
Qld
W.A.
W.A.
Clarke K.L. 124, Bruhl LT., Wilson
K.L.
Blake S.T. 19620
Cane S. 53
Hartley T.G. 14357
W.A.
Carr G.W. 4377, Beauglehole A.C.
N.T.
Booth R. 618K.L.
Qld
Blake S.T. 13611
N.T.
N.T.
N.T.
Wightman G. 424 and Dunlop C.R.
Hunter J.T. 1547, Bruhl J.1.
Dunlop C.R. 9041
N.T.
Jones M., Booth R. 24
W.A.
Dunlop C.R. 5339
N.T.
Wilson K.L. 5109, Taylor S.
N.T.
Wilson K.L. 5207
N.T.
Chippendale G
N.T.
Clarke K.L. 253, Bruhl LT., Wilson
K.L., Cowie I.D.
Clarke K.L. 181, Bruhl LT., Wilson
K.L., Cowie I.D.
Adams L.G. 1715
Ap10+ NE
N.T.
Ap11
N.T.
CANB
166722
Aaffpa NE
ch
As1
NSW
468344
N.T.
W.A.
Clarke K.L. 201, Bruhl J.1., Wilson
K.L., Cowie I.D.
Wilson K.L. 4888
240
A. schoenoides (cant' d)
Abildgaardia sp. aff.
schoenoides
Abildgaardia triflora (L.)
Abeywickr.
As2
BRI
480019;
NSW
468346
As3
NSW
468347
As4
NSW
468343;
MEL
1601269
As7+ BRI 533192
As8
NSW
229593
As9*+ NE
W.A.
Dunlop C.R. 7838
W.A.
Mitchell A.A. 2129
N.T.
Dunlop C.R. 865], White N.G.
Qld
Qld
Bruhl 1.1. 487
lacobs S.W.L. 5903
Qld
N.T.
Clarke K.L. 70, Bruhl 1.1., Wilson
K.L.
Clarke K.L. 157, Bruhl 1.1., Wilson
K.L.
Clarke K.L. 216, Bruhl 1.1., Wilson
K.L., Cowie J.D.
Clarke K.L. 120,1 Bruhl 1.1., Wilson
K.L.
Clarke K.L. 230, Bruhl 1.1., Wilson
K.L., Cowie J.D.
Perry R. 222.
N.T.
Bruhl 1.1. 1261, Hunter 1.1'., Egan 1.
N.T.
Zaire
Dunlop C.R. 5863, Craven L.A.
Malaisse F. 400, Goetghebeur P.
AsI0* NE
W.A.
Asll+ NE
N.T.
As12+ NE
W.A.
As13* NE
+
As14 CANB
16035
As5*# NE 61432
+
As6*# BRI329001
*
MO
4579353
#+
N.T.
Tanzania Greenway P.l. 1859
South
Ward C.l. 1708
Africa
Sri Lanka Koyama T. 13910, Koyama M.
Fimbristylis VahI
Fimbrislylis blakei Latz
fbI *#+ DNA
N.T.
D0120436
fb2
BRI435191 N.T.
(as Fimbristylis sp.)
#+
Fimbrislylis
cinnamometorum
(Yahl) Kunth
fcl+
DNA
A0086876
DNA
134230
NE
fc2#+
NE
N.T.
fc3#
fc4
NE
NE
Qld
N.T.
fc5*#+ NE
Dunlop C.R. 10015, Latz P.K.
Latz P.K. 10375
N.T.
Latz P.K. 11214
N.T.
OrrT.M.442
Qld
Clarke K.L. 61, Bruhl 1.1., Wilson K.L
Clarke K.L. 132, Bruhl 1.1., Wilson
K.L., Cowie J.D.
Clarke K.L. 276
Clarke K.L. 228, Bruhl 1.1., Wilson
K.L., Cowie I.D.
Clarke K.L. 139, Bruhl 1.1., Wilson
K.L.
W.A.
241
Fimbristylis depauperata
#+
NE
Qld
Clarke K.L. 305, Little L.
Qld
Clarke K.L. 263, Bruhl J.1., Wilson
K.L.
Dallachy
R.Br.
NE
Fimbristylis fimbristyloides
ffil
(F.Muell.) Druce
Fimbristylis furva R.Br.
ffi2+
ffi3#
ffl
MEL
269463
BRI336296
QRS 43342
NE
f£1*#+ NE
#+
NE
Fimbristylis fusca (Nees)
Qld
Qld
N.T.
Qld
N.T.
Blake S.T.18678
Flecker H.
Clarke K.L. 267, Bruhl J.1., Wilson
K.L., Cowie J.D.
Clarke K.L. 52, Bruhl jNQ セ
Wilson K.L
Clarke K.L. 210, Bruhl J.1., Wilson
K.L., Cowie J.D.
Dunlop C.R. 5922 Craven L.A.
ffu1
BRI329000 N.T.
ffu2
fsL1
BRI 301745
PERTH
2272911
PERTH
2272962
PERTH
2272911
N.T.
W.A.
Blake S.T. 16571
Kenneally K.F. 11171
W.A.
Kenneally K.F. 11168
W.A.
Kenneally K.F. 11167
NE
W.A.
NE
N.T.
NE
NE
Qld
W.A.
NE
W.A.
NE 65205
South
Africa
Clarke K.L. 131,
K.L.
Clarke K.L. 268,
K.L.
Clarke K.L. 319,
Clarke K.L. 108,
K.L.
Clarke K.L. 153,
K.L.
Browning J. 834
Benth.
Fimbristylis sp. L
(Kimberley flora)
fsL2+
#
Fimbristylis microcarya
#+
F.Muel1.
Fimbristylis schultzii Boeck. *
Fimbristylis variegata
*#
Gordon-Gray
Bruhl J.J., Wilson
Bruhl J.J., Wilson
L. Little
Bruhl J.J, Wilson
Bruhl J.J., Wilson
Bulbostylis Kunth
Bulbostylis barbata (Rottb.) bba1+ DNA 24124 Qld
Wilson K.L. 5442
C.B.Clarke
bba2
bba3
bba4
bba5
bba6#
DNA 49868
DNA 62627
CANB
410874
NE 61441
NE 60472
N.T.
W.A.
N.S.W.
Beauglehole A.C. 2608
Mitchell A.S. 1150
Tindale M.D. 2058
N.T.
N.S.W.
Bruhl J.J., Hunter J.T, Egan J. 1269B
Bell D.B.
bba7
bba8
BRI 316049 N.T.
W.A.
NE
bba9
N.T.
bba10
CANB
325418
NE
bba11
NE
W.A.
N.T.
Latz P .K. 8263
Clarke K.L. 160, Bruhl J.J., Wilson
K.L.
Knight F. 14185
Clarke K.L. 221, Bruhl
K.L., Cowie J.D.
Clarke K.L. 113, Bruhl
K.L.
jNQ セ
Wilson
jN セ
Wilson
242
B. barbata (cont'd)
bba12
NE
bba13
bba14
K
MO
4280627
bba15 EA
bba 16 NSW
468581
bba17 NSW
bba18# BRI 512812
Qld
Clarke K.L. 100, Bruhl J.J., Wilson
K.L.
Singapore Burkhill H.M., Shah M. HMB235
U.S.A.
Hill S.R. 24361
Kenya
India
Napper D.M., Kanuri 2079
Raizada M.B
U.S.A.
India
Correll D.S. 52337
Rajn R.R.V
Po1hill R. 847, Paulo S.
+
bba19
EA
bba20
L 2623
South
Africa
Thailand
NE
N.T.
+
Bulbos(ylis sp. aff. barbata
Bulbostylis burbidgeae
K.L.Wilson
Bulbostylis sp. aff.
burbidgeae
Bulbostylis capillaris (L.)
Kunth ex C.B.Clarke
baffba1 NE
N.T.
+
baffba2 DNA 22621 N.T.
baffba3 NE
N.T.
baffba4 NE
#
baffba5 CANB
421033
baffba6 NE
N.T.
baffba7 NE
N.T.
bbu1
W.A.
Clarke K.L. 251, Bruhl J.1., Wilson
K.L., Cowie I.D.
Clarke K.L. 239, Bruhl J.1., Wilson
K.L., Cowie I.D.
Hart R.P. 2092
W.A.
Carolin R. 7640
W.A.
Mitchell A.A. 1929
W.A.
W.A.
Burbidge N. 1102
Payne A.L. PRP976
N.T.
Dunlop 4725
U.S.A.
Davidse G.
U.S.A.
Curtiss A.H.
PERTH
4275098
bbu2
PERTH
1083007
bbu3
PERTH
5223741
bbu4
BRI 311667
bbu5#+ NSW
452309
baffbu DNA 14302
NSW
452328
MO
+
N.T.
U.S.A.
Styains A.H.
#
bde1
MEL
268550
MEL
268552
L 65104
NE 60798
U.S.A.
N.S.W.
Horr W.H. E170
Bruhl J.J. 1197, Quinn F.e.
bde2
NE 63629
N.S.W.
Hunter J.T., Hunter V. 2737
+
Bulbostylis densa (Wall.)
Hand.-l'vlazz.
N.T.
Larsen K. 1299, Smitinand T., Wamc1
E.
Clarke K.L. 187, Bruhl J.1., Wilson
K.L., Cowie I.D.
Clarke K.L. 184, Bruhl J.1., Wilson
K.L., Cowie I.D.
Rice B.L.
Clarke K.L. 245, Bruhl J.1., Wilson
K.L., Cowie I.D.
Clarke K.L. 241, Bruhl J.1., Wilson
K.L., Cowie I.D.
Bruhl J.J. 369A
243
B. densa (cont'd)
Bulbostylis sp. aff. densa 1
BulbostyUs sp. aff. densa 2
bde3
NSW
468227
bde4
BRI 304640
bde5
BRI 472840
CANB
bde7
50320
BRI311651
bde8
CANE
117835
BRI407433
bde9
bdel0 CANB
282965
bdell CANB
410867
bde12 CANB
87831
bde13 MEL
268539
bde15 BRI 342062
baffd 1* BRI 506795
#+
baffd2 MO
+
4501576
bde16 PRE
bde17
PRE
bde18
Bulbostylis hispidula subsp. *#+
pyriformis (Lye)
R.W.Haines
*#+
PRE
K
Bulbostylis hispidula subsp. *#+
senegalensis (Chenn.) Van
den Berghen
Bulbostylis puberula
C.B.Clarke
#+
Qld
Forster P.I. PIF8482
Qld
Qld
Qld
Blake S.T. 21453
Bean A.R. 1570
Hubbard C.E. 3128
Qld
McKee H.S. 9317
N.S.W.
N.S.W.
Williams J.B.
Gray M. 3255
P.N.G.
Croft, Lelean 34706
P.N.G.
Robbins R.G. 2660
Philippine Ramos M., Edaro G.
s
Sri Lanka Davidse G. 7614
Bean A.R. 3236
Qld
China
Field survey team 820
South
Africa
South
Africa
Swaziland
Kenya
Meeuse A.D.l. 10158
Scheepers J.C. 1141
Haines R.W. 7048
McCallum-Webster
K
NSW
452317
K
Tanzania Richards M. 23175B
Eritrea
Pappi A.
Sudan
Davey J.T. 10
MO
5018124
K
Senegal
Vanden Berghen C. 7484
K
CANB
67185
L 65128
CANB
216795
K
K
DNA 70798
Bulbostylis sp. aff. puberula baffpu NSW
452329
Bulbostylis pyriformis
bpy1
NSW
400826
S.T.Blake
Sri Lanka Davidse G. 9037, Sumithraarachchi
D.B.
Jacobs M. 5691
Borneo
Singapore Sinclair J.
Sri Lanka Clayton D. 5112
Thailand
Brunei
Indonesia
N.T.
van Beusekom C.F. 2247, Smitinand 1
Coode M.J.E. 774, Kirkup D.W.
Elsol J.A.
Latz P.K. 11364
N.S.W.
Johnson L.A.S.
244
bpy2#+ NSW
452308
NE 60921A
bpy3
BRr 311664
bpy4
NSW
bpy5
452307
NSW
bpy6
400827
BRr 9336
bpy7
BRr 542784
bpy8
PRE
Bulbostylis Izumi/is (Kunth)
C.B.Clarke
2827DD
PRE
2927BC
NE 58064
#+
bt2
DNA 62665
Blllbostylis turbinata
S.T.Blake
DNA 51665
bt3
DNA 72393
bt4
PERTH
bt5
2091526
bt7
PERTH
2073552
PERTH
bt8
5221005
BRr 476931
bt9
btl0
MEL
1620712
Ne/mesia Van der Veken
Nelmesia melanostachya
NY
Van der Veken
Nemum Desv. ex Hamilt.
Nemum equitans (Kilk.)
#
NU
J.Raynal
EA
EA
NemUln spadiceum (Lam.)
Desv.
#+
NU
*
NU
*
K
Nemum megastachyum
(Cheml.) J.Raynal
EA
Ty/ocarya Nelmes
Tylocarya cylindrostachya *+(JJB L 65198
Nelmes
)
Outgroup
Actinoschoenus compositus *#+
NE
(Latz) nom. provo ined.
NE
B. pyriformis (cont' d)
N.T.
Latz P.K. 10622
N.S.W.
N.T.
N.S.W.
Hunter J.T., Bell D.B.
Latz P.K. 488?
Wilson K.L. 1479A
N.T.
Latz P.K. 9852
Qld
Qld
South
Africa
South
Africa
N.S.W.
W.A.
Sharpe P.R. 232
Bean A.R. 4227
Jarman N. 134
N.T.
N.T.
W.A.
Latz P.K. 7126
Latz P.K. 6339
Royce R.D. 1491
W.A.
George A. S. 820
W.A.
Payne A.L. PRP 1854
Qld
N.T.
Harris P.L. 342
Beauglehole A.C. 26568
Congo
Gerard 57 (ISOTYPE)
Zambia
Robinson E.A. 2681
Zambia
Zambia
Zambia
Robinson E.A. 3912
Greenway P.J.
Robinson E.A. 5165
Zambia
Zambia
Congo
Robinson E.A. 4676
Robinson E.A. 4677
Germain 4420
Thailand
Kerr A.F.G. 21294
N.T.
Clarke K.L. 178 Bruhl J.J., Wilson
K.L., Cowie I.D.
Clarke K.L. 231 Bruhl J.1., Wilson
K.L., Cowie I.D.
Clarke K.L. 235 Bruhl J.1., Wilson
K.L., Cowie I.D.
Clarke K.L. 211 Bruhl J.1., Wilson
K.L., Cowie I.D.
Cowie I.D. 5643, Brennan K.
N.T.
NE
N.T.
*#+
NE
N.T.
*
NE
N.T.
Ruch M. 9
Strong C.P., Fletcher J.R., Sharp G.C.
Mitchell A.A. 479
245
A. compositus (cont'd)
NE
Arthrostylis aphylla R.Br.
NSW
N.T.
196791
MEL
N.T.
1619328
BRI 311099 Q1d
Clarke K.L. 227 Bruhl J.1., Wilson
K.L., Cowie J.D.
Wilson K.L. 7520
Cowie J.D. 4643
Brass L.J. 18669
BRI429959
BRI 399429
NSWex
Herbario
Kewensis
24110
NSW
259584
NSWex
AD 75517
NSW
NE 30785
NE 58440A
NSW
BatianoffG.N.10089
Qld
Gunness A. AG 1937
Q1d
McKee H.S. RSNH 24110
New
Hebrides
*
NSW
NSW
N.T.
Qld
*
NSW
*
*+
Schoenoplectus
tabernaell10ntani
(C.C.Gmel.) Palla
(=S. validus Vahl)
Schoenoplectiella lateriflora
(J.F.Gmel.) Lye
(=s. lateriflorus G.F.Gme1)
NSW
NSW
Schoenoplectiella laevis
(S.T.Blake) Lye
(=s. laevis S.T.B1ake)
*
*#+
Trachystylis stradbrokensis
(Domin.) Kuk.
N.T.
*
#+
*
*
NSW
NSW
251326
NSW
NSW
338962
BRI 156925
BRI 156923
BRI 399437
BRI 10875
MEL
716466
MEL
2048446
BRI227559
BRI541420
BRI 156931
CANB
193493
Vic
Wilson K.L. 6804
S.A.
Hunt D. 1788
Q1d
N.S.W.
N.S.W.
W.A.
Wilson K.L. 4278
Wallace B.J.
Bruhl J.J. 150
Jacobs S.W.L. 4226
Latz P.K. 3761
Wilson K.L. 3389, Sharpe P.R.,
Johnson L.A.S., B1axell D.
Philippine Ramos M.
s
Unknown
India
Dunlop C.R. 5405
W.A.
W.A.
N.S.W.
N.T.
Qld
Q1d
Qld
Q1d
Q1d
Q1d
Brennan K.
Milthorpe P.L. 1777A, Cunningham
G.M.
Wilson K.L .5399 and Scarlet C.
Wilson K.L. 8041, Clarkson J. and
Jacobs S.W.L.
White C.J.
Q1d
Blake S.T. 13201
Gunness A. Ag 1965
Durrington L 3 12
Sharpe P.R. 5199, Wilson K.L.,
S.W.L.
Blake S.T. 23328
Q1d
Q1d
Q1d
Qld
Sharpe P.R. 2280, Dowling R.
Thompson E.1. 78
Blake S.T. 15945
Brass L.J. 18660
j。」ッ「セ
246
Appendix 2 Full character list used in the assessment of monophyly for the tribe
Abildg,aardieae. All characters used in cladistic analyses in Chapters 3, 4, and 5 are subsets
from this list. Characters presented in the cladograms from subset analyses are given in square
brackets after the tribal character name, with the related chapter indicated as: Chapter 3==!,
Chapter 4=/\ and Chapter U]セ
following the subset character number.
1. Longevity whether
1. annual
2. perennial (with remains of old sheaths and or culms)
Some young perennial plants may appear to be annual therefore care must be taken when
scoring this character - if possible check plants in the area when collecting and note
2. Perennial rhizome whether
1. caespitose - indistinct due to compaction giving clumped base, detectable by persistent sheaths
from last years growth
2. base clumped but rhizome visible sometimes growing vertically
3. base not distinctly clumped but has obvious 'running rhizome' giving smaller clumps spread
4. distinct thick horizontal rhizome with many scales, not spreading widely
5. very distinct and elongated horizontally - base widely spreading
3. Sheath surface cover
1. glabrous
2. scabrid backs
3. scabrid margins
4. medium to dense cover of short to medium hairs
5. mixture of short and long hairs
6. short plus or minus horizontal hairs 60-100 ,. lIn
7. sparsely bristly (hispid - hairs horizontal or nearly so)
8. densely bristly
9. sparse to medium distribution oflong flexuose hairs (c. 300 Ilm, as in Arthrostylis aplzyl/a)
10. dense matt oflong hairs
4. Sheath fitting whether at maturity
1.. tight fitting around culm, especially evident at sheath apex - sheath closed (as in Sclzoenoplectus
mucronatus)
2. fits against culm but not really tight and not loose
3. sheath reduced to near base of plant and is open - recognised from sheath margins (as in
F. fimbristyloides)
4. seems absent and reduced to extreme culm base (leaf blade seems to go all the way to the base of
the plant)
5. open at apex but not loose (as in Abildgaardia oxystachya)
6. sheath loose evenly around the culm and the sheath length, not more so around the apex - sheath
closed (as in Schoenoplectus tabernaemontani)
'7. loose and open around culm, especially at sheath apex (as in Crosslandia setifolia and
Abildgaardia vaginata)
8. sheath apparently open along it's length, at least at maturity - if fused then only in young culms
5. Sheath margins texture
1. sheath margins barely discemable
2. hyaline (thin and translucent (transmits light - very easily damaged))
3. membranous (thin and semi-translucent (like frosted glass), membrane-like)
4. thinly chartaceous (thinner than chartaceous)
S. chartaceous (papery, opaque (light not transmitted) and thin)
6. subcoriaceous (thickish and strong)
7. coriaceous (thick and leathery but flexible)
6. Sheath bases whether
1. sheath base intact as interveinal tissue persists
2. breaking down to fibres from remaining nerves - distinct
7. Sheath backs texture
1. hyaline (thin and translucent)
2. membranous (thin and semi-translucent - membrane-like)
3. thinly chartaceous (papery, opaque (not transparent, dull not shining) and thin) damaged with
forceps not flexible
4. distinctly chartaceous (colour and feel of thin papyrus)
247
5. scarious (thin and dry appearing shrivelled)
6. subcoriaceous (thick and leathery) not easily damaged
7. cartilaginous (hard and tough but flexible)
8. fibrous (having loose woody fibres)
9. pannose (with a felty texture)
8. Sheath apex shape excluding hair extensions
1. pointed (apex longest on margin receding back to culm giving pointed triangular affect)
2. truncated (margins abrubtly end and are the same width the length of the sheath)
3. rounded (margins rounded and are or at least close to the full width of margins along rest of
sheath)
4. tapered (margins narrowed at leaf junction gradually widening to full margin width of sheath
along rest of length)
5. auriculate (sheath tapering then with rounded ends forming auricle)
6. extended (extends beyond sheath-blade junction abaxially)
9. Sheath orifice adaxial
1. open when mature
2. deep V
3. fused at apex
10. Sheath colour in dried material
1. cream
2. straw coloured (golden)
3. yellow-brown
4. light brown
5. dark golden brown
6. pale orange brown
7. orange brown
8. pink-brown
9. red brown
J[ O. mid brown
1l1. dark red brown
12. dark brown (at least nerves are very dark)
11. Sheath leaf junction whether [121\]
1. glabrous
2. short hairs unrelated to ligule (hairs restricted to sheath apex margin and not across the width of
the blade junction)
3. pilose (at sheath apex but not continuing across the full width of blade junction)
4. long coarse hairs
12. Leaf to culm ratio (mature culms and leaves, with or without leaf blade)
1.1:4
2. 1:3
3. 1:2
4.2:3
:5. 1: 1
13. Ligule (whether present)
1. absent
2. as a fringe of stout hairs (near the sheath apex adaxial across the blade or subulate point as in
Fimbristylis depauperata)
3. as a membranous flap (formed as a continuation of the sheath margins adaxial across the blade or
subulate point as in Schoenoplectus)
14. Leafblade whether present or absent
1. absent (binseniform)
2. reduced to subulate points only
3. mixture (of absent or reduced and well developed blades within an individual)
4. present (always)
15. Leaf number per culm (includes bladeless sheaths associated with an individual culm)
1. one per culm
2. two to three per culm
3. greater than three and up to five per culm
4. greater than 5 less than 10
5. greater than 15
248
doesn't include open sheaths that are restricted to base of 'groups' of culms
16. Leafblades shape (see Radford 1974 p. 129)
1, narrowly elliptic (with widest axis at midpoint of structure and with margins synunetrically
curved; more then L W 6:1-3:1)
2. linear (with widest axis at midpoint of structure and with margins essentially parallel; more than
L W 12:1)
3. linear-Ianceolate
4. ovate (with widest axis below middle and with margins symmetrically curved; L W 2: 1-3:2)
5, ovate-Ianceolate
6, lanceolate (with widest axis below middle and with margins symmetrically curved; more than L
W 6:1-3:1)
7, obovate (inversely ovate)
8. falcate (broad blade that is sickle-shape, arcing back from centre of plant)
17. Leafblades habit whether
1. erect (follows line of and is usually intermingled with culms)
2. erect then recurved near apex
3. curly ascending (slightly horizontal then upright)
4. ascending (slightly horizontal and then upright from mid ofleafusually at side of culms)
5. ascending spreading
6. loosely ascending as leaves all 'mishappened'
7. strongly falcate (leaves bent backwards from near base of plant)
18. Leafblade shape in transverse section (along mid-third)
1. sub-triangular (adaxially concave and abaxial midrib distinct as a point with convex sides)
2. concave triangular (abaxial sloping concave faces)
3. shallow channel (crown-like abaxial central rib as point and concave sides giving shallow
channelled appearance from the tlrree points adaxial side usually concave as in B. sp aff.
barbata.)
4. strongly channelled (with deep channels as in B. turbinata)
5. thickly crescentiforrn
6. v-shaped
7. thickly v-shaped
8. U-shaped
9. crescentiform
10. half-circular or obliquely so
11. depressed elliptic
12. thinly crescentiforrn (as in F. schultzii)
13. broadly linear (almost flat - horizontal and thin
14. fused at the margins and is almost subcylindrical
15. fused at the margins and is almost triangular
19. Leafblade width (at midpoint of blade)
1. to 0.3 mm
2. greater than 0.3 to 0.5 mm
3. greater than 0.5 to 0.73 mm
4.0.75 to 0.8 mm
5.0.85 to 1.4 mm
6.1.5 to 2.0 mm
7.2.1 to 3.0 mm
8. greater than 3.0 to 4.8 mm
20. Leafblade vestiture
1. glabrous
2. minutely scabrid margins
3. scabrid margins
4. scabrid over abaxial surface
5. hairy not scabrid (hairs almost horizontal to slightly antorse)
6. densely hirsute backs
7. sparsely bristly
8. bristly hispid
21. Leaf anatomy sclerenchyma presence
1. apparently absent
2. in abaxial area only
249
3, in abaxial and to a lesser extent adaxially (not including sclerenchyma at leaf margins)
22. Leaf anatomy sclerenchyma
1. present as strands (not in contact with vascular bundle but adjacent to)
2. present as girders (in contact with vascular bundle)
3. present as cap above the phloem
4. present as a cap on the inner side of the vascular bundle
23. Leafblade anatomy sclerenchyma shape for abaxial surface
1. square to square with concave sides
2. rhombic (with upper epidermal edge shorter than inner one)
3. reverse rhombic (with upper epidermal edge longer than the inner edge)
4. low mound
5. dome (wide base towards VB)
6. high dome
7. oval-elliptic
8. circular-rounded
9. crescentiform
10. pulviniform (rounded rectangular)
11. rectangular
12. triangular (point towards VB)
24. Number of sclerenchyma strands or girders (compared to vascular bundle number)
1. less than the number of vascular bundles
2. equals the number of vascular bundles
3. greater than number of vascular bundles
25. Leaf anatomy vascular bundle number
1. less than five
2. five to seventeen
3. greater than twenty
26. Leaf anatomy vascular bundles, whether
1. vascular bundles form one layer below the abaxial epidermis
2. vascular bundles form a partial or complete second row as new VB's form near the abaxial
epidermal region
27. Leaf anatomy size of vascular bundles., whether
1. same size
2. 2 sizes with midrib VB being the larger
3. varying sizes with largest bundle at midib
28. Leaf and culm anatomy type of vascularisation [27'''; 26!]
1. C4 fimbristyloid (primary and secondary bundles have PCR tissue interrupted laterally by the
metaxylem vessel elements, the mestome sheath complete and surrounded by PBS)
2. C3 type (having two sheath layers, formed by the mestome sheath which is surrounded by large
achlorenchymatous parenchyma sheath)
The inner border parenchyma cells are large and chlorenchymatous, constituting the PCR
tissue, and interrupted laterally by the metaxylem vessel elements; the mestome sheath of small,
achlorenchymatous, thick-walled cells; and a complete (unless interrupted by sclerenchyma) PBS,
which is usually smaller and less chloroplast laden than the surrounding PCA tissue (a PBS also
surrounds the secondary bundles). Definitions from Bruhl (1990).
The C4 anatomical types are described in terms of primary vascular bundles, the latter being
recognised by the possession of meta- and proto-xylem, often associated with a protoxylem lacuna.
C4 has either fimbristyloid, chlorocyperoid, eleocharoid, or rhynchosporoid type
Fimbristyloid C 4 comprises three bundle sheaths in primary and secondary bundles: the inner
border parenchyma cells are large and chlorenchymatous (=PCR tissue) and is interrupted laterally
by the metaxylem vessel elements; the mestome sheath of thick-walled, achlorenchymatous cells;
and a complete PBS of smaller cells that are less chloroplast laden than the surrounding PCA
tissue.
Chlorocyperoid C4 anatomy is essentially similar, but here the PBS is restricted to one or a
few cells lateral to the metaxylem vessel elements, or sometimes completely absent (being always
absent from secondary bundles).
The border parenchyma cells also constitute the PCR tissue in eleocharoid C4 anatomy, but are
usually not interrupted by the metaxylem vessel elements, and the PBS is absent.
The mestome sheath constitutes the PCR site in rhynchosporoid C4 species, and the PBS is
present but irregularly incomplete.
250
29. Leaf anatomy parenchymatous bundle sheath (peA) [27!]
1. colourless (no chlorophyll present) as seen in Schoenoplectlls and Arthrostylis
2. contains chlorophyll
30. Leaf vascular bundles whether
1. embedded within chlorenchymatous tissue
2. not embedded within chlorenchymatous tissue but intrudes into the clear parenchyma tissue
3. chlorenchymatous tissue apparently absent, or at least highly reduced and undiscemable (tannin
cells prevalent)
31. Leaf chlorenchyma shape
1. globular parenchyma surrounding VB
2. elongated parenchyma near epidermal area above VB and globular around lower part of VB
3. elongated cells surround VB
4. obvious rectangular palisade surrounding the VB
32. Leaf anatomy mesophyll between vascular bundles, whether
1. completely radiate (chlorenchyma arranged in a distinct ring around all major and minor vascular
bundles)
2. incompletely radiate (individual vb's partially encircled by chlorenchymatous cells that are
radiately arranged - no radial vb's occur at the xylem end of the vb, radiate around the phloem
end)
33. Leaf anatomy bulliform cells, whether
1. absent
2. present
34. Leaf anatomy epidermal cells, whether
1. epidermal cells the same size or only slightly larger than abaxial counterparts
2. adaxial epidermal cells about twice as large as abaxial epidermis
3. adaxial epidermal cells mostly three to four times as large as abaxial
4. adaxial epidermal cells about 6 times larger than abaxial cells
5. central adaxial epidermal cells three to four times the size of abaxial epidermis
6. central adaxial epidermis cells about six times the size of abaxial epidermis
1'. central adaxial epidermal cells about 10 times larger than abaxial epidermal cells
Adaxial epidermal cells situated at the midrib that are much larger than the neighbouring
epidermal cells - scored as present when obviously enlarged compared to abaxial epidermal cells
35. Leaf anatomy adaxial hypodermis, whether
1. absent, as epidermal cells only present
2. present, as 1 row often incomplete
3. present, as 2 rows (sometimes 2nd row incomplete)
4. present, as 3 rows (sometimes incomplete)
:;. many rows present ie greater than four rows
36. Leaf anatomy air cavities, whether
]. absent
2. present
37. Leaf anatomy stomata
1. raised (protrude above cuticle layer - above leaf surface)
2. flush (with leaf surface - discrete between epidermal cells)
3. sunken (into leaf surface at bottom of dissecting ribs or pits etc)
38. Culm whether noded vertically
1. not stalked (no obvious ascending nodes present)
2. stalked (node is obvious and culms appear stalked)
39. Culm outline in transvers section, whether
1. narrowly elliptic
2. elliptic no dissections or wavy margins
3. elliptical smooth dissected (deep dissections around the girth of the culm)
4. elliptic wavy no dissections
5. elliptical wavy or undulating with dissections evenly around the margin
6. elliptical deeply undulating
7. elliptic with distinct ribs (grooved)
8. irregularly elliptic with wavy margins
9. irregularly fusiform shaped (waves caused by protruding sclerenchyma bundles as in
Fimbristylis microcarya)
251
40.
41.
42.
43.
44.
10. irregular quadrangle 4 sides, with the parallel sides having one shorter than the other (see
Arthrosytlis aphylla)
11. transversly oblong
12. irregular no definite shape
13. irregularly 7-ribbed
14. irregularly 9-ribbed
15. sub-symmetrical 6-ribbed
16. sub-symmetrically 7 ribbed, distinct
17. sub symmetrically 8-ribbed
18. circular
19. circular undulating
20. circular with distinct ribs (see Actinoschoenus composita)
21. pentagonal (having 5 sides)
22. acutely hexagonal
2:3. irregular hexagon with ribs (6 main points with 6 subpoints forming ribs)
24. depressed triangular with convoluted margins
2: 5. triangular
26. triquetrous (3-angled with concave sides (acutely triangular Metcalfe 1971)
Culm width (at mid third section)
1. to 0.3 mm
2. 0.31 to 0.8 mm
3.0.85 to 1.1 mm
4.1.15 to 1.3 mm
5. 1.5 to 1.6 mm
6. 1.7 to 2.4 mm
7.2.5 to 3.0 mm
8.3.2 to 4 mm
9. greater than 4 mm
Culm cover
t. glabrous (often scabrid at inflorescence junction)
2. scabrid (minute prickle hairs isolated to sparse cover)
3. scabrid (minute prickle hairs with dense cover)
4. hairy not scabrid
5. toothed ascending hairs (40-60 flm)
6. finely pubescent (short interlocking hairs)
'7. hairs mixed (short and long ?toothed)
8. bristly (distally)
9. bristly along the culm length
10. long hairs (c. 600 flm)
dense - A<B (when A is distance between trichomes and B is trichome height);
sparse - A=B to 5xB
isolated - A>5xB
Culm anatomy, total number of vascular bundles is [41-]
1. less than fifteen
2. greater than fifteen less than thirty
3. greater than thirty less than thirty-five
4. greater than forty
Culm anatomy, size of vascular bundles
1. all same size
2.2 sizes alternating
3. 2 sizes not alternating
4. varying sizes alternating evenly
5. varying sizes not alternating evenly
Culm anatomy, number of rings of vascular bundles (VB's)
1. not arranged in rings apparently 'unorganised' within the culm
2. ring of vascular bundles around the outer culm edge and dispersed throughout the culm
3. single ring of bundles all of similar size or alternating size
4. one complete ring and a second semi ring (with the inner VB's being primary and pushed inward
when secondary VB's form in the outer region)
252
45.
46.
47.
48.
49.
5. two complete rings of vascular bundles (the inner ring formed from a lesser number of primary
VB's when secondary VB's develop in the outer ring)
6. multiple rings (older VBs pushed inward while newest VBs are small and numerous near the
culm margin)
Culm anatomy sc1erenchyma [421\]
1. present as strands (not in direct contact with vascular bundle)
2. present as girders (in contact with vascular bundle)
3. present as a cap to four cells thick above the phloem
4. present as a cap to 9 cells thick at the base of the vascular bundle
5. present as multiple layers around the vascular bundle
See Metcalfe 1971
Culm anatomy sc1erenchyma strand number per VB
I . less than number of VB in first ring
2. equals number of vascular bundles in first ring (or equals the number in both rings)
3. greater than the number of vascular bundles in first ring
Culm anatomy sc1erenchyma shape
1. square strand
2. rhombic (bottom wider than epidermal section)
3. reverse rhombic (upper edge wider than inner one)
4. low mound to dome shaped strand (wide base towards VB)
5. high dome strand
6. reverse high dome strand (widest part on epidermis)
7. pulviniform strand (rounded rectangluar)
8. elliptical strand
9. circular strand
la. bulbiform strand
11. crescentiform strand
12. rectangular strand
13. stilted rectangular strand (edges with legs protruding into the parenchyma as in Arthrostylis
planiculmis)
14. thickly v-shaped strand (see Actinoschoenus composita)
15. triangular strand (point towards VB)
16. continuous around the culm
17. reverse high dome girder (widest at the VB)
18. roughly circular girder
19. pulviniform girder
20. rectangular girder (see Actinoschoenus composita)
21. triangular girder (point to VB apex)
22. crescentiform girder on inside of VB (as seen in Schoenoplectus tabernaemolltani and
Eleocharis)
23. cap above phloem to four cells thick
24. cap at base of vascular bundle to 9 cells thick
follows Metcalf 1971
Culm anatomy sc1erenchyma cap on vascular bundle
1. absent
2. present up to 4 cells
Culm anatomy photosynthetic parenchyma shape
1. elongate rounded rectangular cells usually arranged roughly in two to four rows beneath the
epidermis
2. roughly three to four rows of rounded cells sometimes stretched and packed tightly
3. irregularly shaped shorter parenchyma stacked like brickwork in alternating rows
4. shorter rounded irregularly rectangular single upper row and rounded cells beside and below
5. single row of distinct palisade upper and rounded cells beside and below
6. rounded cells packed tightly in a single row above vascular bundle, but more may be present
between the bundles (sometimes slightly stretched but not palisade)
50. Culm vascular bundles
1. not fully immersed within the chlorenchyma tissue as all protrude into the pith (especially
evident in C 3 culms)
253
51.
52.
53.
54.
55.
56.
57.
2. younger vascular bundles immersed within the chlorenchYma tissue while mature bundles
protmde into the pith (especially evident in C 4 culms)
3. younger vascular bundles partly immersed while mature bundles fully immersed (see Metcalfe p:
394)
4. chlorenchYma apparently absent or highly reduced
Culm anatomy central clear parenchYma present as medulla {UPセ}
1. absent (no pith cells visible)
2. absent, breaks down to strands between vascular bundles only
3. present and is distinct
4. present but breaking down in the centre
Culm anatomy stomata when sunken
1. single stomata at bottom of groove or dissection
2. twin stomata near on side walls of groove or dissection
Culm anatomy stomata
1. none apparent or at least very few
2. raised
3. flush (located between sclerenchYma)
4. sunken (at base of dissection)
Root width {USセ}
1. to 0.13 mm
2.0.15 to 0.45 mm
3. 0.46 to 0.55 mm
4.0.56 to 0.8 mm
5. 0.81 to 0.85 mm
6. 0.89 to 1.0 mm
7. 1. 1 to 2.0 mm
8.2.1 to 3.0 mm
Root colour
I. pale cream yellow brown
2. distinctly yellow
3. light brown (straw coloured)
4. orange brown
.5. mid brown
6. red brown
7. grey brown
8. dark brown to black
Root cover
1. glabrous
2. few hairs
3. villous (many long hairs not matted)
4. tomentose (thickly matted)
Inflorescence-synflorescence stmcture
1. spike (intraspicular prophylls indicate lateral branches with a solitary floret, as in Nelmesia)
2. solitary (main florescence only)
3. highly reduced anthelodium (main florescence plus one coflorescence either rayed or sessile)
4. reduced anthelodium (main florescence plus multiple 'rayed' coflorescences - usually 2 to 3)
5. sessile reduced anthelodium (main florescence plus sessile coflorescences - usually 2 giving 3
spikelets)
6. compressed reduced paniculodium forming a 'head' (compressed spike ie main florescence plus
multiple lateral primary coflorescences ie reduced paniculodium as in Fimbristylis schultzii)
7. highly reduced secondary anthelodium (with one second order main florescence (HF2), either
sessile or on lengthened epipodia (ray)
8. ramified reduced anthelodium (spikelets of second order or higher branching that are mostly
'rayed' or sometimes sessile)
9. many spikeleted 'head' of sessile spikelets formed by primary secondary growth and their
secondary lateral growth i.e. HF 1, Cofl + Cof2 (anthela type florescences with reduced epipodia
as seen in Crosslandia WA)
10. lateral hemispherical 'head' on long mesopodia (one or two primary 'rays' supporting lateral
'heads' formed from secondary main florescence (HF2) plus sessile secondary coflorescences
(Cofl2) and their sessile lateral branches (Cofl3)
254
11. terminal 'head' from many sessile spikelets formed from lateral rays (Cofl) and ramification
from intraprophyllar growth (as in Bulbostylis barbata)
12. pseudolateral digitate sessile or on lengthened epipodia (reduced paniculodium)
13. second order pseudolateral digitate lateral and intraprophyllar growth (paniculodium)
Synflorescence structures in this study are based on the premise of consistent reduction ie
paracladia are absent from the main florescence therefore are also absent from the coflorescences;
branches arising are considered as secondary lateral branches (ie ramification)
58. Inflorescence-synflorescence number of usual primary coflorescence branches 'rayed' or sessile {UWセ[
54"']
1. absent
2. one
3. two to four
4. five to eight
5. eight to fourteen
6. greater than fifteen
59. Inflorescence (maximum number of orders consistently present)
1. primary (HF 1 and Cofll includes solitary and simple anthelas)
2. secondary (HF2 lateral florescence growth arising from primary florescence parts with its own
lateral growth)
3. secondary intraprophyllar (growth from within primary prophylls)
4. tertiary (HF3 lateral florescence growth arising from secondary florescence parts with its own
lateral growth)
5. tertiary intraprophyllar (growth arising from with secondary prophylls)
6. fourth (HF4 lateral florescence growth arising from tertiary florescence parts)
Orders were determined using compound rayed specimens and then extrapolated to taxa with
congested 'heads'. Orders were determined using the terminal spikelets on each culm or ray. For
example if a rayed spikelet (i.e. Cofl) developed lateral spikelets (either sessile or on rays) then
that spikelet became the terminal spikelet (HF2) for that ray and the lateral spikelets are CoO. Care
is required when assessing congested 'heads' of spikelets to ensure that congestion is from the same
florescence order. See Crosslandia Figures 3.12-3.17 and Bulbostylis Figure 5.15 for
synflorescence type detail.
60. Inflorescence-synflorescence position whether [56"']
1. terminal (and ascending on the culm)
2. pseudolateral (pushed laterally by the main bracts that usually continue to ascend in line with the
culm, although when large spikelet numbers present (as in Schoenoplectus) the main bract is
reflexed. In Trachystylis the coflorescence appears in the terminal growing position and the main
florescence is pushed sideways)
61. Inflorescence synflorescence, whether open or contracted due to ray length
1. consistently open (solitary spikelet or spikelet 'rays' long and spreading - looks gangly)
2. open (rays lengthened, but not extremely long - as in Bulbostylis turbinata)
3. consistently contracted (spikelets on short rays that can be easily seen giving shortened
appearance but not capitate; as in Fimbristylis complanata)
4. mixed (some single spikelets on 'rays' while others may be sessile or on shortened rays)
5. congested (multiple sessile spikelets on severly restricted epipodia, forming heads or finger
clusters; see inflorescence-synflorescence structure for different types)
62. Inflorescence-synflorescence length
1. to 35 mm
2.38 to 60 mm
:.. greater 60 to 75 mm
4.80 to 100 mm
5. 105 to 120 mm
6. 125 to 170 mm
64. Inflorescence-synflorescence bracts whether [60"']
]. absent
2. present
65. Inflorescence-synflorescence bracts (when present)
1. g1ume-like and clearly associated with inflorescence when not solitary (may have an apiculate
extension as seen in some lower glumes on spikelets)
2. leaf-like and ascending (growing upright at roughly 45 degrees or sometimes loosely erect)
3. leaf-like and spreading (between 45 and 90 degrees separation from culm)
255
66.
67.
68.
69.
70.
71.
72.
4. leaf-like and distinctly erect
5. leaf-like and reflexed downwards due to many aggregated spikelets (as in fゥュ「イウセカャ
scllllltzii)
6. culm-like and continuing in line with the culm (or sometimes obviously bent backwards due to
many aggregate spikelets)
Inflorescence-synflorescence bracts length
1. shorter than inflorescence
2, equals inflorescence
3. longer than inflorescence
4. much longer than inflorescence
Inflorescence-synflorescence primary bract number [63"']
1. one
2. two (closely alternately opposite)
3. three (ascending alternately of roughly equal length)
4. many (of roughly equal length)
Spikelet axis whether
1. monopodial (rachilla of spikelet has one axis ie growth continuous from one growing point)
2. sYmpodial (rachilla formed from multiple reduced axes)
Crosslandia with sYmpodial growth has distichous glumes which when glumc pulled away
breaks the section of rachilla away as well. The nut is partly surrounded by the glume margins of
the opposite (lower) glume. The nutlet is buried deep within the rachilla section. Basal and aerial
spikelets show the same pattern even though the spikelets are morphologically different.
Aerial spikelet prophylls whether
1. present (distinct bract-like extension between nut and rachilla)
2. absent
Aerial spikelet rachilla shape of wings [641\]
1. wingless or minute
2. reduced and rounded (due to compaction of the rachilla)
3. reduced and truncate (due to compaction of the rachilla)
4. reduced and tapered to a point
5. distinct (elongated) rounded - broad with rounded apex (rachilla expanded where fertile)
6. distinct oblong - narrow with curved apex (rachilla expanded where fertile)
7. distinct truncate (apex terminates abrubtly)
8. distinct narrowly triangular - narrow with pointed apex
Spikelet sex in aerial spikelets (excluding lowest empty glumes)
1. hermaphrodagamous (bisexual florets only spikelet)
2. hermaphrodandrous (male florets proximal, bisexual florets distal in spikelet)
3. gynehermaphroditic (bisexual florets proximal, female florets distal in the spikelet)
4. gynagamous (female florets only in spikelet)
5. gynecandrous (male florets proximal, female florets distal in spikelet)
6. androgynous (male florets only in spikelet)
7. gynehermaphrodandrous (male, bisexual mid, female distal)
Aerial spikelet outline
1. glumes angular - widely dome shaped (truncated proximally and broadly rounded at the apex)
2. elliptic (2: 1 to 3:2 widest point at centre)
3. loosely narrowly elliptic (6: 1 to 3: 1 narrow at base and apex widest point at centre - glumes not
tightly imbricate or angular)
4. angularly narrrowly elliptic (6: 1 - glumes somewhat reflexed)
5. obliquely ovate (curving on one side of spikelet due to rachilla twisting)
6. smoothly ovate (2: 1 to 3:2 wider at the base narrowing towards apex)
7. loosely ovate
8. angularly ovate (glumes apex reflexed)
9. cylindrical (as in Schoenoplectiella laevis)
10. smoothly narrowly cylindrical (glumes tightly imbricate - 6: 1 as in Nelmesia melanostachya)
11. loosely narrowly cylindrical
12. loosely lanceolate (glumes not tightly imbricate 3: 1 to 6: 1 wider at the base narrowing towards
the apex)
13. loosely obliquely lanceolate (curving on one side due to twisting rachilla)
14. angularly lanceolate
15. narrowly oblong (3: 1 to 6: 1 base and apex same width)
16. triangular (2: 1 to 3:2 wide at base to pointed apex)
256
17. narrowly triangular (6: 1 to 3: 1 wide at base to pointed apex)
18. smoothly linear (glumes tightly imbricate - 12: 1 narrower than narrowly oblong sides even)
19. angularly linear (glume nerves long and recurved)
73. Number of fertile florets (male or female) per aerial spikelet {WRセ}
1. 1 to 2 (as in Actinoschoenus, Arthrostylis, Trachystylis)
2. greater than 2 to 4
3. many (greater than 4 up to 25 sometimes many glumes but few nuts as in Abildgaardia)
4. numerous greater than 25 (as in Nemum)
74. Spikelets whether morphologically different to aerial spikelets {WSセ[
66!]
1. aerial only (always on lengthened culms)
2. aerial plus subradical (subradical culms distinctly shortened and spikelets near base, but spikelets
otherwise identical to aerial counterparts)
3. Aerial plus basal (basal spikelets absent or very highly reduced and with different morphology
and floret sex to aerial counterparts)
4. aerial plus subterranean (spikelets reduced to one or two nuts close to or below ground level amphicarpy)
Basal spikelets are those that differ in morphology and frequently sexuality, from aerial
spikelets, usually maturing before their aerial counterparts i.e. amphicarpic. As seen in
Crosslandia, Fimbristylis spiralis and occasionally in Abildgaardia vaginata, plus some species of
Bulbostylis and Schoenoplectiella.
75. Basal spikelet sex (excluding lowest empty glumes whether)
1. hermaphrodagamous (bisexual [perfect] florets only in spikelet)
2. gynehermaphroditic (bisexual florets [perfect] proximal, female florets distal in the spikelet)
3. gynagamous (female florets only in spikelet)
76. Basal spikelet (shape)
1. irregularly widely ovate (due to extreme reduction of number of florets - usually one to two)
2. narrowly elliptic (6: 1 to 3: 1 narrow at base and apex widest point at centre)
3. lanceolate (3: 1 to 6: 1 wider at the base narrowing towards the apex)
4. narrowly oblong (3: 1 to 6: 1 base and apex same width)
5. oblanceolate (6: 1 to 3: 1 narrow at base and wider at apex due to spreading glumes)
6. narrowly triangular (6: 1 to 3: 1 wide at base to pointed apex)
7. linear triangular (12: 1 wider at base than the tapered apex)
8. linear (12: 1 narrower than narrowly oblong with sides even)
77. Basal spikelet (floret numbers)
1. one to two
2. greater than two but less than four
3. greater than four to many
78. Basal spikelets whether
1. Basal spikelets always sessile (no culm present)
2. Basal spikelets mostly sessile although some with very short culms present (less than 3 mmlong)
3. Basal spikelets mostly on reduced culms (at first appearing sessile and clumped at plant base
although rarely some longer culms with gynagamous spikelets present)
4. Basal spikelets mostly on shortened culms greater than 5mm long (often c. 15 mm) and restricted
to plant base
79. Spikelet glumes whether
1. always falls with mature nuts
2. persists on the spikelet after nuts mature
80. Aerial glume length
1. to 2.12 mm
2. 2.2mm to 3.5 mm
3.3.6 to 3.8 mm
4.3.9 to 4.1 mm
5.4.2 to 4.55 mm
6.4.6 to 4.85 mm
7. 4.9 to 5. 1 mm
8.5.2 to 5.75 mm
9.5.8 to 5.95 mm
10. 6.0 to 6.25 mm
11..6.3 to 6.55 mm
12. 6.6 to 6.8 mm
257
13.6.9 to 7.1 mm
14.7.2 to 8.4 mm
15.8.5 to 10.1 nun
16. 10.2 to 11.6 mm
17. 12 to 13.7 mm
18. 14 to 15 mm
81. Aerial glume width [74/\]
1. to 0.7 mm
2.0.75 to 0.9 mm
3. greater than 0.9 to 1.46 mm
4. 1.5 mm to 1.7 mm
5. greater than 1.7 mm to 1.85 mm
6. greater than 1.85 mm to 2.05 mm
7.2.05 mm to 2.4 mm
8. 2.45 mm to 2.55 mm
9. 2.6 mm to 2.9 mm
10. greater than 2.9 mm
82. Aerial glume back colour (excluding tannins)
1. no real colour as is translucent (some tannin may be present mainly near the glume base)
2. cream
3. straw
4. yellow brown
5. pale orange brown
6. orange brown
7. light brown
8. pink-brown
9. mid brown
10. red brown
11. deep burgundy (as in Nemum)
12. burgundy black (very dark burgundy appearing almost black)
83. Aerial glume margin colour whether
1. darker than gume back
2. same colour as glume back and sides
3. colour lighter than glume back
4. consistantly colourless and distinct
84. Aerial glume margins {XSセ}
1. entire (without indentations, incisions, or trichomes along margins)
2. lacerate (margins irregularly cut, appearing torn)
3. with antrorse prickle hairs pointing towards apex
4. minutely ciliolate «20 11m)
5. ciliolate (with tiny or small trichomes protruding from margins c. 20 11m)
6. short hairs
7. hispid or almost so (as a continuation of glume back indumentum)
8. fimbriolate (minutely fimbriate flattened projections)
9. fimbriate (fringed margins with flattened processes)
10. ciliate at apex only
11. loosely ciliate (long lax hairs that look mishappen)
12. piliferous (with long conspicuous trichomes c. 200 11m, that are lax or flexuose, and protruding
from margins)
85. Aerial glume whether margin in transverse section
1. is continuing in line with the glume sides, not inrolled or splayed
2. is inrolled between 1/2 and upper 1/3 of glume
3. margins only slightly curved backwards or flattened with glume still generally boat shaped
4. is splayed so that the margin is between 45 and 90 degrees - flattening out giving a narrow keel
with splayed sides
5. is almost flattened (only the nerve is raised; the glume sides are flattened against the glume
below, although the basal area around the nutlet may be boat shaped)
6. is revolute (margin strongly recurved and bending backwards)
86. Glume texture whether [77!]
1. hyaline (thinner than membranous and very delicate, usually colour is absent)
258
87.
88.
89.
90.
2. membranous (almost transparent and usually colourless nut can be seen through the glume as in
Bulbostylis sp. aff barbata)
3. finely chartaceous (thicker than membranous light may be seen but is dulled and papery, has
some flexibility but easily damaged with forceps, as in Fimbristylis blakei)
4. tougher than chartaceaous as is more flexible but damaged with forceps (as in Actinoschoen/ls)
5. chartaceous (thicker than finely chartaceous as light does not pass through but may still be
damaged, as in Crosslandia anthelata)
6. fine leathery (not overly thickened but is quite tough and flexible not easily damaged unless
pulled, as in some Fimbristylis)
7. subcoriaceous (quite tough and hard to bend, thick and not at all transparent, as in species of
Abildgaardia )
Aerial glume margins {XVセ[
80"']]
1. margins hyaline (light passes directly through)
2. membranous (light is opaque)
3. finely chartaceous and indistinct, or almost so, from rest of glume
4. indistinct from rest of glume (with same texture as glume backs which are subcoriaceous as in
Abildgaardia)
Aerial glume apex outline
1. rounded (margins and apex forming a smooth arc)
2. retuse (lobe rounded; sinus depth to 1/16 distance to midpoint of blade; margins convex)
3. emarginate (lobe rounded; sinus depth 1/16 to 1/8 distance to midpoint ofblade;margins straight
or convex)
4. obtuse (margins straight to convex, forming a terminal angle more than 90 degrees)
5. acute (base cuneate - margins straight to convex forming a terminal angle 45-90 degrees;
muticous)
6. acuminate (base narrowly cuneate - margins straight to convex forming a terminal angle of less
than 45 degrees; muticous)
7. sub-mucronate (nerve less than 0.1 mm, but not muticous)
8. mucronulate (1: 1 I w nerve 0.1 mm)
9. mucronate (with nerve less than 3: 1 length/width, straight and stiff between 0.1 and 0.3 mm)
10. apiculate (more than 3: 1 length/width, usually slightly curled and flexuous; used for nerve
extension greater than 0.3mm but equals or less than 0.7 mm)
11. aristate (more than 3: 1 length/width, usually prolonged, straight and stiff; used here for
excurrent nerve forming awn-like projection greater than 0.7 mm long)
Aerial glume general shape [80!]
1. oblong (2: 1 to 3:2 with widest axis at midpoint of structure and with margins essentially parallel)
2. narrowly oblong (6:1 to 3:1 with widest axis at midpoint of structure and with margins
essentially parallel)
3. spathulate
4. linear (more than 12: 1 with widest axis at midpoint of structure and with margins essentially
parallel)
5. ovate
6. widely ovate
7. very widely obovate (1: 1 with apex curving in a wide arc)
8. narrowly emarginate glume apex narrowing but often rounded at end with nerve extending and is
usually reflexed (see Fimbristylis schultzii)
9. emarginate (glume apex almost as wide as base and rounded, dipping into but not attached to
nerve)
10. obtuse (nerve terminates below glume apex and apex is broad and round)
1 Jl. trullate (2: 1 to 3:2 kite shaped with widest point near base)
12. lanceolate (more than 6: 1 to 3: 1 with widest axis below middle and with margins symmetrically
curved)
13. oblanceolate (more than 6:1 to 3:1 reverse of laneeolate)
14. narrowly trullate (6:1 to 3:1 with widest axis below middle and with straight margins, trowel
shaped)
15. triangular (2:1 to 3:2 with 3 sides and 3 angles)
16. narrowly triangular (6: 1 to 3: 1 with 3 sides and 3 angles)
17. linear triangular (more than 12: 1 with 3 sides and 3 angles)
Aerial glume apex whether
1. not recurved at maturity (usually looks quite sleek and neat)
2St)
2. distinctly straight - not bent backwards or curving forwards
3. slightly reflexed backwards, but not strongly recurved
4. strongly and consistently recurved at maturity (as in Abildgaardia schoenoides)
91. Aerial glume nerve whether
1. muticous (nerve does not extend pass glume apex, frequently finishes abrubtly beneath glume
apex)
2. submruco point (c.0.05 to 0.1 mm)
3. nerve to mucro point (to 0.1 to 0.5 mm)
4. nerve excurrent (greater than 0.5 mm)
92. Aerial glume abaxial surface
l. glabrous
2. nerve only scabrid
3. sparsely scabrid
4. scabrid over most of the surface
:5. short antrorse appressed hairs
6. antrorse hairs (c. 100um long at 45 degrees to glume back)
7. bristly
8. tomentose
93. Glume epidermal cells shape in glume sides (at 50 x magnification)
1. indistinct
2. rectangular 1:2 to 1:3 longitudinally lengthwise with strongly sinuose walls
3. rectangular 1:2 to 1:3 longitudinally lengthwise with straight walls
4. linearly rectangular 1:6 longitudinally
5. linear longitudinally
6. irregularly elongate
94. Glume epidermal cells whether
1. tanin idioblasts absent
2. tanin idioblasts present (cells filled with tanins but not raised)
95. Glume epidermal tanins shape
1. present as cellular 'dots'
2. present as squares
3. present as rectangular c. 3: 1
4. minute striations which are very faint
5. present as long striations (vertical lines along length of glume)
6. present as joined striations forming almost continuous lines
7. forming continuous 'colour'
96. Aerial glumes tanin cover
1. restricted mainly to apex
2. sparse and restricted mainly to glume base
3. sparse and restricted to glume backs and sides, not on margins
4. sparse mostly occurring near outer margins only
5. sparse mostly occurring on glume sides and margins (rarely in nerve area)
6. evenly over glume but not all cells gives speckled appearance
7. dense over glume sides and margins
8. dense and continuous over glume back and thinning on sides
97. Glume epidermal cells whether
1. raised' gland' cells (as seen in Fi11lbristylis cinna11lo11letoru11l)
2. devoid of any obvious raised gland-like cells
98. Aerial glume shape in cross-section [91/\]
1. single highly thickened nerve giving distinct keel (see Fi11lbristylis schultzit')
2. narrowly keeled (as in Bulbostylis barbata, narrow V of 3-nerves forming a keel)
3. narrow U of 3 to 5 nerves
4. broad V (from multiple nerves but forming a distinct V as in Abildgaardia vaginata)
5. nerve area broad 5 to 7 nerves forming U but margins continuous texture with sides (sometimes
glume flattened but nerves distinct)
6. nerve area broad (forming U with distinct margins from glume sides, usually consists of 5 to 7
nerves forming rounded V-bottom)
7. nerves indistinct (glume forms wide, shallow V with margins not flattened as in 2; nerves usually
indistinct or only 1 fine nerve visible)
8. nerve distinct and broad with glume broad and shallow curved or almost flat
260
9. nerve area often indistinct or as apparent single slim nerve glume relatively flat at least above
99. Aerial glumes glandular cover
1. mostly restricted to apex
2. mostly restricted to margins
3. over most of surface
100. Aerial glume arrangement
1. distichous (attached directly oppositely ascending, sometimes rachilla twists to give spiraldistichous impression but glumes in definite rows)
2. sub-distichous (glumes at least distichous at first then twisting, but not fully spiral spirodistichous) glumes appear spirally arranged and not in rows but spikelet is slightly
depressed in cross section
3. almost alternately opposite (decussuate but ascending minutely acropetally as seen in Bulhostylis
barbata)
4. opposite decussate (as in Trachystylis where empty glumes are paired and 90 degrees from distal
fertile pair)
5. tristichous where glumes attached ascending in a tristichous spiral
6. spiral (glumes attached in a close ascending spiral acropetally)
101. Aerial glumes (number neutral (empty) per spikelet)
1. none
2. one
3. two
4. three
5. four
6. five
7. six or seven
102. Basal glume (length)
1. less than 2 mm
2. greater than 4 mm
103. Basal glumes (margins)
1. entire
2. ciliate
3. ciliolate
4. fimbriate
5. fimbriolate
6. involute
7. lacerate
104. Basal glumes (general outline)
1. linear (12: 1 - widest axis at midpint of structure and margins essentially parallel)
2. narrowly oblong (6:1 to 3:1)
3. lanceolate (more than 6: 1 to 3: 1 - widest axis below middle and margins symmetrically curved)
4. ovate (2:1 to 3:2 - see lanceolate)
5. widely obliquely ovate
6. narrowly trullate (more than 6: 1 to 3: 1 - with widest axis below middle and with straight
margins)
7. triangular (2:1 to 3:2 - with 3 angles and 3 sides)
8. narrowly triangular (6: 1 to 3: 1 - with 3 sides and 3 angles)
9. linear-triangular (more than 12:1 - with 3 sides and 3 angles)
105. Basal glumes abaxial surface
1. glabrous
2. nerve scabrid
3. back scabrid
106. Basal glume epidermal cells whether
1. tanin idioblasts absent
2. tanin idioblasts present
107. Basal glume epidermal cells tanin cover
1. restricted to glume base
2. sides of glumes
3. over entire glume (giving striated appearance)
108. Perianth whether present
1. absent
261
2. present as bristles
Hypogynous bristles or scales are absent in members of the Abildgaardieae, however, there is
one specimen of Abildgaardia schoenoides collected from Kakadu NP that has perianth present.
109. Bristle hairs, whether
1. antrorse
2. retrorse
110. Perianth number when present
111. Stamen number in aerial male or bisexual florets
1. one
2. two
3. three
4. four
5. five
6. six
112. Anther length, including apiculum
1. to 0.15 mm
2.0.2 to 0.75 mm
3.0.8 to 0.9
4. 0.95 to 1.50 mm
5. 1.55 mm to 1.65
6. 1.70 to 1.85
7. 1.9 to 3.0
8. 3.0 to 3.50
9. 3.60 to 4.2
10.4.3 to 6.7 mm
113. Anther apiculum, whether connective tissue extends past the antheridium
1. indistinct (either absent or to 0.05 mm in length)
2. distinct (greater than 0.05 mm to 0.2mm)
3. prominent (greater than 0.2 mm)
114. Aerial style length (measured from base of stylebase to base of stigmas)
1. to 1.25 mm
2. 1.30 to 2.75 mm
3. 2.80 to 3.35 mm
4. 3.4 to 3.8 mm
5. 3.9 to 4.30
6.4.40 to 4.9
7. 5.0 to 5.50 mm
8. 5.6 to 6.2 mm
9.6.5 to 7.9 mm
10.8.0 to 12.60 mm
115. Aerial style width [106!]
I. to 0.1 mm maximum measurement
2. greater than 0.1 to 0.15 mm (0.1 is minimum measurement)
3. greater than 0.15 mm to 0.2 mm
4. greater than 0.2 mm to 0.25 mm
5. greater than 0.25 mm to 0.30 mm
6. greater than 0.30 mm to 0.34 mm
7.0.35 mm to 0.45 mm
116. Aerial style surface cover, (excludes style-base)
1. glabrous
2. isolated fimbriola 40-80um
3. sparse fimbriola 40-80um
4. dense fimbriola 40-80um
5. isolated fimbria 100-140um, scattered along style often nearer the base and missing from apex
6. sparse fimbria 100-140um, with the distance between each process greater than the length of
each process
7. dense fimbria 100-140um, with the distance between each process less than the length of the
process
8. coarsly fimbriate 220 セ ュ
long and 100 セュ
apart dense becoming very dense towards base
9. ciliate
262
10. densely matted with long hairs excluding style-base
117. Aerial style outline in transverse section
1. flattened and broad
2. distinctly flattened (strap-like, as seen in many Fimbristylis species)
3. terete (apparantly, as seen in Trachystylis)
4. finely triangular (as seen in Crosslandia)
5. minutely triangular (as seen in Bulbostylis barbata)
6. distinctly triangular (sometimes slightly flattened but with 3rd angle distinct)
7. broadly triangular with distinct flattened face giving flat appearance
118. Aerial style base length (in millimetres)
1. from 0.05 to 0.2 mm
2. to 0.5 mm
3. greater than 0.5 and less than 1.5 mm
4. greater than 1.5 to 3.5 mm
119. Aerial style base width (in millimetres)
1. to 0.15 mm
2. 0.2 to 0.5 mm
3. greater than 0.5 to 1.0 mm
4. greater than 1.0 mm
120. Style-base shape
1. not widened or distinct
2. globular (as seen in Bulbostylis barbata)
3. bulbous
4. narrowly triangular (as in Crosslandia and Abildgaardia vaginata)
5. regularly triangular (no face distinctly concave or convex)
6. triangular (with adaxial faces concave and abaxial face flat as in Abildgaardia macrantha)
7. conical
8. broadly triangular (somewhat rounded not with distinct triangular edges)
9. broad and convex (on abaxial side away from rachilla giving a depressed appearance)
10. squarish to barely triangular and depressed (as seen in Fimbristylis depauperata continues in
line with style but is distinct)
121. Aerial style-base surface cover [110!]
1. glabrous
2. sparse rounded fimbriola (20 11m in length)
3. isolated to sparse fimbriola (40-60 11m)
4. dense fimbriola (40-60 11m)
5. fimbriate (100-140 11m)
6. coarsely fimbriate (c. 220 11m and 100 11m apart thick coarse looking very dense on style base)
122. Aerial or basal style-base, whether persistent or deciduous [116/\]
1. persistant (style-base always separates from style and often persists on nut apex, but not always)
2. deciduous (style-base remains connected to style when abscissed from nut apex)
123. Aerial style stigma number
1. two
2. three
124. Aerial style stigmas relative length, when compared with style length
I. stigmas (less than length of style)
2. approximately equals (stigmas=style L)
3. stigmas greater than style length
125. Aerial style stigmatic processes observed at lOx magnification
1. minute papillae
2. small fine papillae
3. fimbriolate
4. fimbriate
5. woolly (distinctly)
126. Aerial style stigma colour
1. white
2. golden brown
3. red brown
4. deep red-brown
127. style (surface cover, excludes style-base)
263
1. glabrous
2. occasional fimbriola
3. sparse
4. hairs denser at base and sparse or absent towards style apex
128. Basal style (shape in transverse section)
1. ligulate
2. terete
3. narrowly triangular
4. minutely triangular
129. Basal stylebase (indumentum cover) whether
1. absent as stylebase is glabrous (at least appearing so)
2. occasional fimbriola
3. sparse fimbriola occuring near style base to nut abscission zone
4. dense fimbriola 40-120 セ ュ
130. Aerial nutlet length [1241\]
1. less than 0.65mm
2. 0.65 to 1.3mm
3. 1.35 to 2.0mm
4. greater than 2 and less than 3mm
5. greater than 3 less than 4.5mm
131. Aerial nutlet width [1251\]
1. to 0.60 mm
2. 0.61 to 1.15 mm
3. greater than 1.15 mm to 1.34 mm
4. 1.35 mm to 1.50 mm
5. greater than 1.50 mm to 1.75 mm
6. greater than 1.75 mm less than 1.90 mm
7. 1.90 mm to 2.50 mm
132. Aerial nutlet stipe length
1. zero as no constriction observed
2. to 0.2 mm
3. greater than 0.2 to 0.28
4. 0.3 to 0.4
5. greater than 0.4 to 0.8
6. greater 0.8 to 1.0 mm
7. greater than 1.0 to 1.10
133. Aerial nut hypogynophore [127 /\]
1. apparently absent or highly reduced enclosed within the fruit wall (may appear as a 'button')
2. distinct as a stalk not enclosed by the fruit wall of the nut (usually brownish) with filaments
attached at the base
3. completely enclosed within the 'stipe' of the nut fruit wall and not apparent from the outside
(with filaments attached at the base of the stipe)
Hypogynophore is a stalk directly below the ovary and may be apparent as separate from the
nut proper or enclosed within the stipe of the nut. On some nuts the hypogynophore may be highly
reduced or appear absent. Stamens and perianth parts are attached at the base of the
hypogynophore.
In Abildgaardia odontocarpa the hypogynophore is present in the stipe of the nut and not
evident from the outside - separating from the seed proper when the wet nut is opened. The
hypogynophore adheres to the inner area of the nut that encased it.
134. Aerial nutlet shape in transverse section (at mid-third of organ) [123!]
1. biconvex (lenticular depressed due to two faces)
2. cylindrical (without distinct sides appearing rounded - ribs not evident)
3. rounded trigonous
4. distinctly plano-convex (distinct dorsal ventral sides in 3-sided fruit ie has definite face where
fruit sits against the rachilla evenly-flatly - usually slightly larger than the other 2 faces that form
an almost single convex face eg A macrantha)
5. sub-trigonous
6. trigonous (3-sided, of roughly equal size, faces not concave)
7. triquetrous
8. strongly triquetrous
264
135. Aerial nutlet nut apex (excluding any persistent stylebase)
1. nut apex with a distinct extension or point formed from style base (as in Schoenoplectus)
2. nut apex umbonata (sometimes nut apex may have a small point but it is not formed from the
stylebase and is usually no more than 0.1 nun in length)
3. nut apex generally rounded
136. Aerial nutlet outline excluding external gynophore
1. ovate
2. elliptic (2: 1 to 3:2 as in Crosslandia)
3. widely elliptic (6:5)
4. obovate (2: 1 to 3:2)
5. widely obovate (6:5 as in Bulbostylis barbata)
6. very widely obovate (1: 1 as seen in Schoenoplectus)
7. pyriformis (pear shape - nut with truncate base with style-base protruding, apex umbonate)
8. obampulliformis (compressed in 3-dimensions - apex somewhat elliptic with slight contriction
near base but not like stipe in Abildgaardia)
9. obcordate (as in Abildgaardia pachyptera nut wings give the distinct shape with wings
pronounced or not, sometimes entire or with notches)
10. obtrullate (3:2)
11. widely obtrullate (6:5)
12. very widely obtrullate (1: 1)
13. napiform (more tapered than obtrullate with smoother lines)
14. widely napiform (starting as widley obovate but with tapered base)
] 5. clavate (club shaped as in A macrantha as stipe forms club handle)
16. capitate (head like as in A ovata as stipe forms neck)
17. strongly capitate (where nut severly restricted half way forming tight head at apex on a long
narrow stalk or stipe)
1. 8. mace shaped (from pronounced horns from deep notches on strongly constricted head with
extended stipe)
137. Aerial nutlet wings, whether present or absent on the nut (see Lye 2000 p:625)
1. absent as there are no pronounced projections from face edges
2. present (in the horizontal plane ie not on the convex face sometimes greatly reduced and
notched)
3. as three distinctly broad notched protusions from the face edges and top (looks mace·-like)
138. Aerial nutlet colour
1. white
2. pale pink, with white 'bloom' on outer surface
3. cream
4. straw
:5. yellow
6. dark golden brown
7. port wine (may have cream-pink face and port wine ribs as seen in Bulbostylis barbata)
g. light grey
9. dark grey
10. light brown
11. grey brown
12. dark brown
13. black
139. Aerial nutlet surface whether (follows Radford et al. 1974)
1. glaucous (covered with a bloom or smooth waxy coating)
2. glaucescent (sparingly or slightly glaucous; does not include waxy coating removed or damaged)
3. dull (light not reflected back; surface not coating with wax but not highly lustrous)
4. lacquered (nut appearing as iflacquered; some light reflected but not lustrous)
5. glistening (especially evident in white nuts)
6. shining (nitid or laevigate; appears lustrous or polished)
140. Aerial nutlet epidermal cell outline (at 50x magnification)
1. indistinct
2. minutely ovate (need measurements)
3. appearing circular to hexagonal (c. 20 )lm in size and at 500x are actually hexagonal)
4. widely elliptic to roughly circular (40-80 )lm sometimes squarish, at 500x are actually
hexagonal)
265
5.
6.
7.
8.
9.
distinctly hexagonal (giving nut surface a honecombe appearance)
cells almost square to just rectangular transverely 1 by 1(or 1 by 2)
transversly narrowly oblong (20 x 60 /lm to 20 x 120 /lm)
transversely rectangular 40 x 60 /lm (in distinct longitudinal rows)
transversly rod-shaped having tapered ends (walls straight)
10. longitudinally oblong (40 x 20/lm)
II. longitudinally narrowly oblong 6: 1 (120 x 20 /lm), 3: 1 (60 x 20 /lm)
12. aciculated (marked with very fine longitudinal irregular streaks, as if produced by the point of a
needle)
141. Aerial nutlet surface patterning (including protuberances follows Lye 2000)
1. smooth (no apparent pattern on nutlet surface, epidermal cells indistinct at lOx magnification)
2. subpuncticulate (some single papillae raised but not over all of surface or not prominent)
3. puncticulate (single papilla 5-15 urn diameter raised to form minute bumps prominent in cells
over the surface of the nutlet)
4. subpusticulate (1-3 cells raised in groups forming low mounds and are not prominent)
5. small tubercules (from 1-3 cells prominent)
6. pusticulate (large rounded tubercules)
7. tuberculate (from multiple cells - prominent greater than 4, protruding conical outgrowths or
papillae formed over more than one epidermal cell, usually 20-100 urn diameter)
8. verrucate (usually flat topped and very distinct and upright)
9. continuously rugulose
10. discontinuously mildly rugose
11. continuously mildly rugose over nut excluding stipe
12. rugose (longitudinally elongate epidermal cells that are raised to form prominent undulating
transverse wrinkles)
13. continuously acutely rugose
14. longitudinally grooved striated (grooves in prominent longitudinal rows)
15. transversly oblong cells in indistinct longitudinal rows
16. transversly interlocking rod-shaped in roughly longitudinal rows
17. reticulate ('netted' epidermal cells with defined walls but not in distinct rows)
18. scalariform (ladderlike and almost in rows)
19 mostly individual cells sunken
20, individual epidermal cells raised (not forming tubercules and not puncticulate)
21. reticulate-foveate (cell walls raised and thickened)
22. alveolate (cell depression but cell walls not raised and distinct)
142. Aerial nutlet protuberance or pattern distribution
1. absent
2. occasional
3. uneven distribution (sparse)
4. restricted to vertical rows (usually two) down the fruit wall face and nut trigonous ribs
5. bordering margins of nutlet (i.e. along face ribs)
6. sparse distribution even over upper three quarters of area of nutlet (no constriction)
7. sparse distribution spread evenly over all of nutlet (no constriction)
8. sparse distribution over nutlet surface excluding stipe (constriction)
9. dense distribution over upper three quarters of nutlet (not constricted towards base)
10. dense distribution spread evenly over nutlet surface excluding stipe (constriction)
11. dense distribution evenly of nutlet surface (constriction not present or at least minimal)
143. Basal nut length
1. to 1.2 mm
2. from 1.3 to 2.1 mm
144. Basal nut width
1. from 0.75 to 1.25 mm
145. Basal nutlet, whether distinctly beaked at nut apex
1. nut with a distinct beak formed from style base (as in Schoenoplectus)
2. distinct beak absent (sometimes nut apex may have a small point but it is not formed from the
stylebase and is usually no more than 0.1 mm in length)
146. Basal nutlet shape in transverse section
1. cylindrical (dorsal ventral sides not easily determined)
2. plano convex (dorsal ventral sides obvious in fruit)
3. sub trigonous to trigonous
26()
147. Basal nutlet outline shape
1. elliptic (2:1 to 3:2)
2. narrowly elliptic (3: 1)
3. widely elliptic (6:5)
4. obovate (2:1 to 3:1)
5. widely obovate
148. Basal nutlet colour
1. cream
2. golden
3. dark golden brown
4. dark grey
5. dark brown
6. black
149. Basal nutlet surface protuberances (follows Lye 2000) [138!]
1. absent
2. rugose
3. puncticulate (each cell with a raised silica body)
4. subpusticulate (individual epidermal cells raised -not forming tubercules and not puncticulate as seen in Crosslandia nutlets)
5. groups of 1-3 cells raised but not prominent (giving 'chequered appearance of raised areas over
entire nut surface)
6. cells walls raised and thickened over nut surface
7. small tubercules (from 1-3 cells) raised evenly over nutlet surface
150. Basal nutlet surface, whether [139!]
1. glaucous (covered with a bloom or smooth waxy coating)
2. glaucescent (sparingly or slightly glaucous)
3. dull
4. lacquered
5. shining (nitid, laevigate, lustrous or polished)
151. Embryo, general type {QUPセ[
145"\]
1. Fimbristylis-type (root orientation lateral and smaller than basal orientated shoot)
2. Abildgaardia-type (root orientation basal and smaller than basal orientated shoot, but larger than
Fimbristylis-type)
3. Bulbostylis-type (root orientation basal and the same size or larger than the basal orientated
shoot)
4. Nemum-type (root orientation basal and smaller in size to the basal orientated shoot)
5. Schoenoplectus-type (root orientation lateral distal (apical) beneath mushroom shaped cotyledon
and and smaller than basally pointing shoot but midway along cotyledon apical extension)
6. Carex-type (root terminal and larger than the inconspicuous lateral shoot)
7. Schoenus-type (shoot and root distinctly sub basal)
152. Embryo cotyledon outline
1. narrowly top-shaped (wide at the apex and gradually narrowing to a 'point' as in Fimbristylis Atype embryo)
2. reverse dome shaped (with base rounded and almost parallel sides as in Fi1llbristylis
cinna1llo1lletorum Fimb B type)
3. roughly reverse dome shaped with cotyledon having a saddle (as in Fimbristylis disticha Fimb C
type)
4. broadly top-shaped with base widely rounded due to shoot and root size (as in Bulbostylis-type
embryo)
5. very widely top shaped with base wide due to shoot root (as in Ne1llum megastaclzYllln)
6. inversely bell shaped (as in Crosslandia type embryo Fimb D type embryo)
7. broadly inverse bell-shaped (as in Abildgaardia-type embryo)
8. very broadly inversely bell shaped (cotyledon very wide brimmed and not deep almost hat-like
as in Abildgaardia oxystachya base rounded)
9. saucer shaped (as in Actinoschoenus broad and compressed at poles)
10. ellipsoid (as in Carex-type embryo - seen in Actinoschoenus, see Kern 1974)
11. subpyramidal (distinctly 3-sided not rounded and sharply pointed at the base as in Abildgaardia
mexicana)
12. mushroom shaped with cotyledon distal extension (as in Schoenoplectus-type embryo)
267
The outline of the embryo can depend on the view i.e. from the side the embryo may look inversely
bell shaped but from the front position may look ellipsoid.
153. Cotyledon shape from distal or proximal view
1. narrowly elliptic (cotyledon appearing somewhat flattened on the sides)
2. elliptic
3. circular
4. cotyledon almost triangular-trigonous
5. triangular
154. Embryo morphology, orientation of the germ pore compared to the first leaf primordia {QUSセ}
1. parallel with the first leaf primordia
2. perpendicular
3. distinctly open and circular in the centre (as seen in Fimbristylis schllltzii)
268
Appendix 3 Glycerin jelly for semi-permanent slides (Kearns and Inouye 1993).
Ingredients
Quantities
Distilled water
35 mL
Glycerin
30mL
Gelatin
10 g
Phenol -- crystalline (preservative)
1g
N.B. phenol is a known carcinogen.
Method: In a beaker, dissolve the gelatin in distilled water by heating gently. Add the glycerin
and phenol, stirring while on low heat. Avoid creating bubbles in the mixture. When dissolved
pour slowly into two new Petri dishes and allow to cool. If preferred the phenol can be omitted,
but the jelly will need to be kept refrigerated to prevent mould growing.
Preparing semi-permanent slides: Take a cleaned microscope slide and place onto it a small
cube of glycerin jelly c. 5 x 5 mm. Place the prepared sections onto the top of the jelly in a small
amount of distilled water. Gently place a cover slip so that it balances on top of the jelly. Heat
the slide simply by placing onto a dissecting microscope slide that has under lighting and heat
until melted and the coverslip is sitting flat on the slide. Allow to cool before storing in slide
boxes.