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Phenetic study of the Microsorum punctatum complex (Polypodiaceae)

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Microsorum punctatum (L.) Copel. is a widespread species of ferns. It is distributed in the tropics and subtropics of Asia and Africa from sea level to about 2800 m elevation. At present, the taxonomic status and boundary of this species is still uncertain due to the great variation in frond form, size, and venation patterns. These variations depart from previous systematic treatments, so this species group was proposed as a species complex worth investigating. Cluster analysis and canonical discriminant analysis were performed on 694 herbarium specimens deposited at BCU, BKF, BM, K, B, L, and P and consisting of 20 taxa of Microsorum punctatum complex. Twenty-five quantitative and 31 qualitative characters were employed. Cluster analysis separated these 20 taxa into seven groups, viz. M. thailandicum, M. membranaceum, M. musifolium, M. glossophyllum, M. siamense, M. steerei, and M. punctatum. The seven cluster groups are discussed. From a canonical discriminant analysis using the 20 taxa as priori groups, it can be concluded that M. siamense, M. thailandicum, M. membranaceum, and M. musifolium are obviously distinct taxa, while characteristics of M. glossophyllum and Pleopeltis megalosoides are mixed and should be proposed as the same species. Likewise, specimens of M. steerei, P. tonkinense, and P. playfairii should be recognized as M. steerei. Moreover, the specimens of the species previously recognized as the other 10 synonyms of M. punctatum are not distinct from M. punctatum and are treated here as members of M. punctatum. The four most important characters that have separated the seven species are stipe length, density of sori, number of sori rows between adjacent secondary veins, and diameter of sori. These quantitative characters, together with some qualitative characters were coded into a database using Descriptive Language for Taxonomic Analysis (DELTA) software. An identification key to species of the Microsorum punctatum complex was then created using the associated program INTKEY. The differentiation between the taxa studied is discussed.
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R
ESEA RCH ARTI CLE
doi: 10.2306/scienceasia1513-1874.2012.38.001
ScienceAsia 38 (2012): 112
Phenetic study of the Microsorum punctatum complex
(Polypodiaceae)
S. Petchsria,b, T. Boonkerda,b,, B.R. Baumc
aBiological Sciences Program, Faculty of Science, Chulalongkorn University, Bangkok 10330 Thailand
bDepartment of Botany, Faculty of Science, Chulalongkorn University, Bangkok 10330 Thailand
cAgriculture & Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Ottawa, Ontario, K1A 0C6,
Canada
Corresponding author, e-mail: Thaweesakdi.B@chula.ac.th
Received 4 Jul 2011
Accepted 15 Feb 2012
ABSTRACT:Microsorum punctatum (L.) Copel. is a widespread species of ferns. It is distributed in the tropics and
subtropics of Asia and Africa from sea level to about 2800 m elevation. At present, the taxonomic status and boundary
of this species is still uncertain due to the great variation in frond form, size, and venation patterns. These variations
depart from previous systematic treatments, so this species group was proposed as a species complex worth investigating.
Cluster analysis and canonical discriminant analysis were performed on 694 herbarium specimens deposited at BCU,
BKF, BM, K, B, L, and P and consisting of 20 taxa of Microsorum punctatum complex. Twenty-five quantitative and
31 qualitative characters were employed. Cluster analysis separated these 20 taxa into seven groups, viz. M. thailandicum,
M. membranaceum,M. musifolium,M. glossophyllum,M. siamense,M. steerei, and M. punctatum. The seven cluster
groups are discussed. From a canonical discriminant analysis using the 20 taxa as priori groups, it can be concluded that
M. siamense,M. thailandicum,M. membranaceum, and M. musifolium are obviously distinct taxa, while characteristics of
M. glossophyllum and Pleopeltis megalosoides are mixed and should be proposed as the same species. Likewise, specimens
of M. steerei,P. tonkinense, and P. playfairii should be recognized as M. steerei. Moreover, the specimens of the species
previously recognized as the other 10 synonyms of M. punctatum are not distinct from M. punctatum and are treated here as
members of M. punctatum. The four most important characters that have separated the seven species are stipe length, density
of sori, number of sori rows between adjacent secondary veins, and diameter of sori. These quantitative characters, together
with some qualitative characters were coded into a database using Descriptive Language for Taxonomic Analysis (DELTA)
software. An identification key to species of the Microsorum punctatum complex was then created using the associated
program INTKEY. The differentiation between the taxa studied is discussed.
KEYWORDS: systematics, fern, numerical taxonomy, morphometric analysis
INTRODUCTION
Microsorum punctatum (L.) Copel. (Polypodiaceae)
is a common species of fern in Africa and Asia and
occurs naturally in various forest types of tropics and
subtropics from sea level up to 2800 m elevation 1,2.
The most recent treatment of Microsorum treated
several species as synonyms of this species3. For
example, M. musifolium Copel. and M. glossophyl-
lum Copel. were previously recognized as distinct
species2. Holttum4pointed out that dried herbarium
specimens of M. punctatum are very similar to the
narrow-frond specimens of M. musifolium and it is
not easy to distinguish them. Moreover, Nooteboom1
noted that M. musifolium and M. glossophyllum are
connected with M. punctatum by many intermediates.
He also pointed out that M. membranaceum and
M. steerei are very close to M. punctatum and that they
might be conspecific or in part varieties.
Another related species, M. polycarpon (Cav.)
Tardieu has different morphological characters of
their fronds from M. punctatum; but currently was
considered as a synonym of M. punctatum by some
pteridologists1,4. Likes Polypodium punctatum L.
subsp. subirideum Christ and P. punctatum subsp.
subdrynariaceum Christ were firstly proposed in 1906
and were finally treated as synonyms of M. punctatum
in 19982,3.
Recently, M. thailandicum Boonkerd & Noot.,
and M. siamense Boonkerd were discovered from
limestone mountains in southern Thailand5,6. These
two species were noted that they resemble to
the widespread polymorphic M. punctatum and
M. steerei.
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2ScienceAsia 38 (2012)
Moreover, these taxa, especially cultivated plants,
exhibit variations in frond forms (e.g., irregularly
lobed). Some of these forms have been described as
cultivars, i.e., M. punctatum (L.) Copel. ‘Serratum’.
These variants are not included in the previous recog-
nized systematic treatments.
These examples show that M. punctatum and
related species are not clearly circumscribed or delim-
ited. It seems that the taxonomic status of M. punc-
tatum and its related taxa is still unclear. Nooteboom
placed these collective taxa, thus as a species complex
and suggested that they are worth investigating (per-
sonal communication, 9 September 2005).
It can be seen that members of this species com-
plex have a history of circumscriptional uncertainty,
suggesting the need for further taxonomic evaluation.
The objectives of this work were as follows: (1) to
investigate the morphometric relationship and deter-
mine the suitability of the species circumscriptions
of M. punctatum and of other related taxa, and (2)
to determine the importance of some macro- and
microscopic characters that can be used to distinguish
these taxa. With these objectives in mind, both cluster
analysis and discriminant analysis were performed
based on both qualitative and quantitative characters
(i.e., 56 characters) examined from 694 herbarium
specimens.
MATERIALS AND METHODS
Plant materials
In this study, we examined about 1500 herbarium
specimens kept in B, BCU, BKF, BM, K, L, and P
(herbarium abbreviations according to Ref. 7). Six
hundred and ninety four complete specimens of 20
taxa were used (Table 1), considering each specimen
as an operational taxonomic unit (OTU).
Data analysis
Twenty-five quantitative characters (Fig. 1 and
Table 2) and thirty-one qualitative characters
(Table 3) were subjected to cluster and canonical
discriminant analysis (data matrices are available
from the corresponding author). Cluster analysis was
carried out based on both quantitative and qualitative
characters using the Gower similarity coefficient8and
UPGMA clustering in the MVSP program (Kovach
Computering Services, MVSP Plus, version 3.1). The
characters used in the analysis were assumed to be of
equal importance and were not weighed.
A subset of characters that maximized differences
among the groups determined by cluster analysis was
selected by stepwise discriminant analysis9. To char-
Fig. 1. Measurment of rhizome, scale and frond part. A plant, B. venation, C. sori distribution, D.
rhizome, E. scale, F. sporangium and G. spore. Noted: The alphabet AC = Costal areole.
LMW
B.
Type 1.
Type 2.
NSR
DSO
PAW
SCW
DBSV
ALB
F.
DSR
NAC
APS
ALA
RHD
PDL
STD
RDL
PHS
PHS
SPW
G.
SPL
A.
STL
LML
E.
SCL
AC
AC
D.
C.
Fig. 1 Measurment of rhizome, scale and frond part. A.
plant, B. venation, C. sori distribution, D. rhizome, E. scale,
F. sporangium, and G. spore. Noted: The alphabet AC =
Costal areole.
acterize mean differences among the taxa, canonical
discriminant analysis was used to acquire insight into
group differences and to estimate character weights
from correlations between canonical variables and
original variables, using the procedure CLASSIFY
in SPSSPC -FW, release 10.0 10 . In comparison, the
groupings consisting of the 20 taxa of M. punctatum
complex were used as a priori group for a series of
discriminant analysis. To summarize the range of
variation between and within the segregated groups
on each character, univariate analysis was performed.
Boxplots of the most important characters drawn from
the magnitude of their value in F test were carried out
using SPSSPC -FW10.
For the construction of a key to species, qualita-
tive characters of the segregated taxa were tabulated
and compared with the results from morphometric
analysis for their importance in discriminating the 20
taxa. Subsequently, the best characters for separating
the segregate genera as suggested by DELTA11 were
used.
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ScienceAsia 38 (2012) 3
Table 1 List of the 20 taxa of the Microsorum punctatum (L.) Copel. complex and their present status.
No. Taxon Taxon according to Nooteboom (1997), Taxon according to the present study
Boonkerd and Nooteboom (2001),
and Boonkerd (2006)
1. M. siamense Boonkerd M. siamense Boonkerd M. siamense Boonkerd
2. M. thailandicum Boonkerd & Noot. M. thailandicum Boonkerd & Noot. M. thailandicum Boonkerd & Noot.
3. M. membranaceum (D. Don) Ching M. membranaceum (D. Don) Ching M. membranaceum (D. Don) Ching
4. M. glossophyllum (Copel.) Copel. M. punctatum (L.) Copel. M. glossophyllum (Copel.) Copel.
5. Pleopeltis megalosoides Alderw. M. punctatum (L.) Copel. M. glossophyllum (Copel.) Copel.
6. M. musifolium (Blume) Copel. M. punctatum (L.) Copel. M. musifolium (Blume) Copel.
7. M. steerei (Harr.) Ching M. steerei (Harr.) Ching M. steerei (Harr.) Ching
8. P. tonkinense Baker M. steerei (Harr.) Ching M. steerei (Harr.) Ching
9. P. playfairii Baker M. steerei (Harr.) Ching M. steerei (Harr.) Ching
10. M. punctatum (L.) Copel. M. punctatum (L.) Copel. M. punctatum (L.) Copel.
11. M. punctatum ssp. subirideum Christ M. punctatum (L.) Copel. M. punctatum (L.) Copel.
12. M. punctatum ssp. subdrynariaceum Christ M. punctatum (L.) Copel. M. punctatum (L.) Copel.
13. Polypodium irioides Poir. M. punctatum (L.) Copel. M. punctatum (L.) Copel.
14. M. validum (Copel.) Ching M. punctatum (L.) Copel. M. punctatum (L.) Copel.
15. P. glabrum Wall. M. punctatum (L.) Copel. M. punctatum (L.) Copel.
16. P. millisorum Baker M. punctatum (L.) Copel. M. punctatum (L.) Copel.
17. M. sessile F´
ee M. punctatum (L.) Copel. M. punctatum (L.) Copel.
18. P. polycarpon Cav. M. punctatum (L.) Copel. M. punctatum (L.) Copel.
19. M. punctatum (L.) Copel. ‘Serratum’ M. punctatum (L.) Copel. M. punctatum (L.) Copel.
20. M. neoquineense Copel. M. punctatum (L.) Copel. M. punctatum (L.) Copel.
Table 2 Twenty five quantitative characters, with their methods of scoring used in this study of the M. punctatum (L.)
Copel. complex. Univariate F values of the different characters used in the canonical discriminant analysis and pooled
within canonical structure using (I) 7 clustering groupings and (II) 20 taxa groupings.
No. Abbrev. Characters Discriminant Function
I II
F-value Sign. Axis 1 Axis 2 F-value Sign. Axis 1 Axis 2
1. LMWaLamina width (mm) - - 0.01 0.09 250.10 0.00 0.06 0.18
2. LMLa,b Lamina length (mm) - - 0.03 0.05 - - 0.03 0.04
3. STL Stipe length (mm) 4682.59 0.00 0.87c0.00 1491.98 0.00 0.86 0.02
4. STD Stipe diameter (mm) 249.03 0.00 0.02 0.05 63.67 0.00 0.03 0.06
5. DMP Diameter of phyllopodia (mm) 237.82 0.00 0.07 0.03 57.78 0.00 0.08 0.02
6. PDL Phyllopodia length (mm) 161.23 0.00 0.08 0.09 32.27 0.00 0.09 0.09
7. RHDa,b Rhizome diameter (mm) - - 0.04 0.04 - - 0.05 0.04
8. RDL Distance between closest phyllopodia (mm) 359.81 0.00 0.13 0.20 101.46 0.00 0.13 0.20
9. SCL Scale length (mm) 140.01 0.00 0.00 0.19 29.23 0.00 0.00 0.18
10. SCWbScale width (mm) 133.98 0.00 0.00 0.12 - - 0.03 0.08
11. ALA Angle of frond apex 169.30 0.00 0.10 0.13 34.05 0.00 0.10 0.14
12. ALB Angle of frond base 731.54 0.00 0.15 0.03 165.90 0.00 0.15 0.02
13. DSO Diameter of sori (mm) 278.22 0.00 0.01 0.37 52.30 0.00 0.01 0.36
14. DSRa,b Diameter of sporangium (mm) - - 0.00 0.05 - - 0.01 0.05
15. NAC Number of annulus cell 423.82 0.00 0.03 0.02 126.09 0.00 0.03 0.01
16. SPL Spore length (mm) 219.42 0.00 0.02 0.19 44.13 0.00 0.02 0.19
17. SPWbSpore width (mm) 146.42 0.00 0.01 0.16 - - 0.00 0.07
18. DSL Density of sori per cm2203.27 0.00 0.00 0.43 41.00 0.00 0.00 0.42
19. PLS Distance between lowest sori and frond base (mm) 178.92 0.00 0.05 0.12 47.79 0.00 0.06 0.12
20. PHS Distance between highest sori and frond apex (mm) 128.51 0.00 0.02 0.06 30.72 0.00 0.02 0.07
21. NSR Number of sori rows between closest secondary vein 1325.89 0.00 0.02 0.57 439.81 0.00 0.02 0.58
22. DBSV Distance between closest secondary vein (mm) 190.00 0.00 0.02 0.22 38.34 0.00 0.02 0.23
23. APS Angle between primary and secondary vein 529.00 0.00 0.02 0.29 85.67 0.00 0.02 0.32
24. SAW Secondary areole width in mm 153.54 0.00 0.01 0.01 36.12 0.00 0.01 0.03
25. PAW Primary areole width (mm) 314.46 0.00 0.05 0.38 72.90 0.00 0.05 0.37
a,b A character followed by a superscript letter indicates a character not selected by stepwise discriminant analysis to be
used in further canonical discriminant analysis: a=I, b=II.
cNumbers in bold are the important variables associated with each axis.
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4ScienceAsia 38 (2012)
Table 3 Comparison of 31 qualitative characters of the seven clustering groupings.
No. Character Taxa
I II III IV V VI VII
1. Shape of rhizome in transverse section: (1) approximately cylindrical, 1 1,2 1,2 1,2 1 1,2 1,2
(2) dorso-ventrally slightly flattened or flattened
2. Rhizome surface: (1) not waxy, (2) at least sometimes waxy under the 1 1 1 2 1 2 2
scales or often waxy
3. Differentiation of vascular bundle sheaths: (1) vascular bundle sheaths 1 1 2 2 1 1 1
be parenchymatous, (2) vascular bundle sheaths be collenchymatous
4. Attachment of scales: (1) pseudopeltate, (2) peltate 1 1 2 1 2 1 1
5. Density of scales: (1) densely set, (2) apically densely set or 1 2 2 1 1 2 2
moderately densely set
6. Spreading of scales: (1) distinctly or slightly spreading, (2) appressed 1 1 2 1 1 1 1
7. Scales shape: (1) narrowly ovate to ovate, (2) triangular 1,2 1,2 1 1 1,2 1,2 1,2
8. Scales margin: (1) entire, (2) dentate to denticulate 2 1 1 2 2 2 2
9. Scales apex: (1) acute (2) acuminate to slightly caudate 2 1,2 2 2 2 1,2 2
10. Presence of hyaline margin on rhizome scales: (1) absent, (2) present 1 2 2 1 1 1 1
11. Indumenta type of central region of scales: (1) dark black on central 1 2 1 1 2 1 1
region or glabrous, (2) bearing multiseptate hairs at least when young
12. Scales translucence: (1) opaque and blackish, (2) translucent and 2 2 2 1 2 2 2
brownish
13. Phyllopodia distinctness: (1) distinct, (2) obscure 1 1 2 1 1 1 1
14. Lamina texture: (1) membranaceous, (2) firm herbaceous, 3 1 2,3 2 3 3 2,3
(3) subcoriaceous-coriaceous
15. Frond colour when living: (1) light to dark green, (2) iridescent 2 1 1 1 2 1 1
blue-green
16. Lamina shape: (1) linear, (2) (narrowly) elliptic, (3) (narrowly) ovate, 1 1–3 4 4 2 4 1–4
(4) narrowly obovate to oblanceolate
17. Lamina base: (1) cuneate, (2) angustate, (3) truncate to obtuse, 5 2 3 1 4 2 2
(4) attenuate, (5) abruptly narrowed
18. Margins of lamina: (1) margin entire, (2) margin entire, undulate 1 1 1 1 2 1 1,2
19. Presence of indumentum on fronds: (1) with only scales, (2) with a few 3 3 2 1 3 3 2
scales and short glandular hairs, (3) with only short glandular hairs
20. Presence of stipe: (1) present, (2) absent or obscure 1 1 2 1 1 2 1
21. Midrib character: (1) slightly raised or raised on both surface, 2 1 1 1 1 1 1
(2) raised on lower surface, grooved on upper surface
22. Venation general pattern: (1) type 1: connecting veins forming a row of 2 1 1 1 1 1 1
about equally sized areoles between two adjacent vein and no prominent
veinlet situated parallel to the veins, (2) type 2: the first
connecting vein forming one row of small primary costal areoles
parallel to the costa, other larger, areoles in a row between two veins
23. Visibility of veins: (1) all veins distinct, (2) all veins indistinct 2 1 1 1 2 2 2
or secondary and smaller veins more or less immersed and vague
(at least in living specimen)
24. Branching of included free veinlet venation: (1) free veinlet 2 2 1 1 1 2 1
simple and once-forked, (2) free veinlet simple, once or twice forked,
25. Sori distribution pattern: (1) mostly irregularly scattered on simple 1 2 3 3 1 3 3
free or on 2 or 3 connecting veins, (2) forming into 2–4 irregular rows
parallel to each pair of secondary veins, (3) forming more than (2-)
3–10 (-15) irregular rows parallel to each pair of secondary veins
26. Visibility of hydathodes: (1) distinct, (2) Indistinct 2 2 2 2 1 2 2
27. Sori position on lamina surface: (1) superficial, (2) slightly immersed 2 1 1 1 1 1,2 1
28. Sori distribution: (1) on the whole surface of the lamina or restricted 2 1,2 1,2 2 2 2 1
up to the distal 1/2, (2) absent from the basal parts for 1/5–4/5 of
total length of lamina
29. Spores shape: (1) plano-convex, (2) concavo-convex 2 2 2 1,2 2 2 1
30. Spores colour: (1) hyaline, (2) yellowish hyaline, (3) yellow 2 3 1 1 3 1 1,2
31. Spore surface: (1) plain to slightly verrucate, (2) irregularly rugate 1 2 1 1 1 1 1
I=Microsorum thailandicum, II =M. membranaceum, III =M. musifolium, IV =M. glossophyllum, V =
M. siamense, VI =M. steerei, and VII =M. punctatum.
QUANTITATIVE CHARACTERS
Cluster analysis
Firstly, 31 qualitative and 25 quantitative characters
were used in this analysis and as a result the den-
drogram showed the segregation of 694 specimens
into seven groups at the 0.90 phenon level of Gower
similarity coefficient (Fig. 2). Specimens classified as
group I to group III consisted of M. thailandicum (1),
M. membranaceum (2), and M. musifolium (3), respec-
tively. All specimens of M. glossophyllum (4) and
Pleopeltis megalosoroides (5) were placed in group
IV. Group V included all specimens of M. siamense
(6), while those of M. steerei (7) and its synonyms,
i.e., Polypodium tonkinense (8), and P. playfairii (9)
were placed in group VI. The last group, group VII,
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ScienceAsia 38 (2012) 5
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Gower Similarity Coefficient
I
II
III
IV
VII
VI
V
1
a
2
3
6
4 - 5
7 - 9
10 - 20
Fig. 2 ualitative and 25 quantitative characters of
Fig. 2 UPGMA clustering of 694 OTUs based on Gower Similarity Coefficient calculated between means of 31 qualitative
and 25 quantitative characters of the M. punctatum (L.) Copel. complex (a =numbers correspond to those in Table 1,
I=Microsorum thailandicum, II =M. membranaceum, III =M. musifolium, IV =M. glossophyllum, V =M. siamense,
VI =M. steerei, and VII =M. punctatum).
consisted of specimens of M. punctatum (10) and its
eleven synonyms (11–20).
Likewise, the results obtained from only 25 quan-
titative characters showed segregated groups of 694
specimens at a high phenon level of Gower simi-
larity coefficient (Fig. 3). Specimens of M. mem-
branaceum (Group II in both of 31 qualitative and
25 quantitative characters analysis), M. musifolium
(III), M. glossophyllum and Pleopeltis megalosoides
(IV), M. thailandicum (I), and M. punctatum and its
eleven synonyms (VII) were classified as group A,
B, C, D, and F, respectively. Group E, on the other
hand, was consisted of 2 subgroups: M. siamense
(V) and M. steerei (VI). First subgroup is solely
M. siamense (V). The second subgroup is composed
of all specimens of M. steerei and its synonyms (VI).
Specimens of both subgroups, however, are separated
from each other.
From both results, it can be seen that the spec-
imens of M. thailandicum,M. membranaceum, and
M. musifolium are distinct species. The speci-
mens of Pleopeltis megalosoides, which was previ-
ously treated as a synonym of M. glossophyllum by
Bosman3, and the specimens of M. glossophyllum
were placed together in both analyses (Fig. 2 and
Fig. 3). Then, the group of these specimens should be
considered a distinct species, i.e., M. glossophyllum.
Likewise, M. steerei and its two synonyms, Poly-
podium tonkinense and P. playfairii1,3, were placed
in the same group in both results and should also con-
stitute distinct species. The largest group, M. punc-
tatum and its previously treated synonyms, is also
recognized here as a distinct species from the result of
both cluster analysis. Theses cluster groupings were
supported by previous taxonomic work3,5,6,12 .
The results from the second cluster analysis in-
dicated that M. siamense and M. steerei are closely
related species (Fig. 3) since some quantitative char-
acters of these two species overlapped. However, they
do separate when both qualitative and quantitative
characters were employed (Fig. 2) or only qualitative
characters, such as rhizome surface (2), attachment of
scales (4), frond colour when living (15), lamina base
(17), margins of lamina (18), the presence of distinct
hydathodes (26), spores colour (30) were in consid-
eration (Table 3). Our findings clearly emphasize
www.scienceasia.org
6ScienceAsia 38 (2012)
Gower Similarity Coefficient
13
15
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0.76 0.8 0.84 0.88 0.92 0.96 1
Gower Similarity Coefficient
2
a
3
4 - 5
7 - 9
10 - 20
1
6
A
B
C
F
E
D
Fig. 3 uantitative characters of the M. punctatum (L.)
Fig. 3 UPGMA clustering of 694 OTUs based on Gower Similarity Coefficient calculated between means of 25 quantitative
characters of the M. punctatum (L.) Copel. complex (a =numbers correspond to those in Table 1, A =M. membranaceum,
B=M. musifolium, C =M. glossophyllum, D =Microsorum thailandicum, E =M. siamense,M. steerei, and F =
M. punctatum).
the importance of both qualitative and quantitative
characters in plant classification and identification.
Canonical discriminant analysis
The seven-cluster groupings: When using the re-
sults of the groupings from cluster analysis as a priori
groups, specimens of the 20 taxa can be separated
into 7 groups (Fig. 4a) on canonical axis 1. They,
however, were not clearly distinct on canonical axis
2. The nature of the group differences is demonstrated
by the pooled within a canonical structure (Table 2).
Canonical variable 1 is 99.1% correlated with 21
characters and the variance explained is 79.0% (data
matrix available on request from the corresponding
author). It is most highly associated with stipe length
(3). Canonical variable 2 explains 11.6% of the total
variance. This axis is most highly associated with
DSO (13), SPW (17), DSL (18), as well as NSR (21).
The four variables LMW (1), LML (2), RHD (7), and
DSR (14) were not selected by stepwise discriminant
analysis to be used in further canonical discriminant
analysis (Table 2).
The canonical plot on two canonical axes (Fig. 4a)
shows the division of specimens into 7 groups on
canonical axis 1, but was not clearly distinct on
canonical axis 2. It is evident from both cluster
analysis (Fig. 2) and canonical discriminant analysis
(Fig. 4a) that the 694 specimens collectively grouped
in to 7 distinct species, namely M. thailandicum (I),
M. membranaceum (II), M. musifolium (III), M. glos-
sophyllum (IV), M. siamense (V), M. steerei (VI), and
M. punctatum (VII).
Regarding M. membranaceum, this species
clearly separated from M. punctatum in both cluster
analysis (Fig. 2) and canonical discriminant analy-
sis (Fig. 4a). Although Nooteboom 1suggested that
M. membranaceum might be a variety of M. puncta-
tum. They also clearly differ in at least 8 qualitative
characters, namely rhizome surface (2), scale margin
(8), presence of hyaline margin on rhizome scales
(10), lamina texture (14), visibility of veins (23), spore
shape (29), spore colour (30) and spore surface (31).
As regard to their habitats M. membranaceum usually
occurs above 600 m altitude whereas M. punctatum
has a much wider vertical distribution from sea level
to 2800 m altitude1. These two species also differ in
life-span of fronds. M. membranaceum usually sheds
its fronds annually while M. punctatum has perennial
fronds1.
In previous treatment of the microsoroid ferns,
Nooteboom1recognized M. musifolium as a synonym
of M. punctatum and noted that this species is a
www.scienceasia.org
ScienceAsia 38 (2012) 7
20100-10-20
6
4
2
0
-2
-4
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(a)
Canonical Axis 1
Canonical Axis 2
VI
V
IV
III
VII
I
II
20100-10-20
6
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0
-2
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(b)
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Canonical Axis 2
5
1
2
3
7
6
4
Fig. 4 The ordination of 694 herbarium specimens of the
M. punctatum (L.) Copel. complex A7 clustering groupings,
B20 taxa groupings :- 1 =Microsorum thailandicum ();
2=M. membranaceum (); 3 =M. musifolium (J); 4 =
M. glossophyllum (); 5 =M. siamense (); 6 =M. steerei
(); and 7 =M. punctatum ().
form with broader leaves and more connecting veins,
connected with M. punctatum with many interme-
diates. The results of cluster analysis (Fig. 2 and
Fig. 3) and canonical discriminant analysis (Fig. 4)
clearly indicated the distinction between these two
taxa. This finding also corresponds with the differ-
ence in qualitative characters of at least 8 charac-
ters, namely, rhizome surface (2), differentiation of
vascular bundle sheaths (3), attachment of scales (4),
spreading of scales (6), scale margin (8), lamina base
(17), visibility of veins (23), and spore shape (29).
Hence M. musifolium should be recognized as its own
species.
Microsorum glossophyllum is another species that
was previously treated as a synonym of M. puncta-
tum1,3. In contrast, our results from cluster analysis
(Fig. 2 and Fig. 3) and canonical discriminant analysis
(Fig. 4) strongly supported the segregation of this
species from M. punctatum. The two species also
differ in at least 7 qualitative characters (Table 3),
namely differentiation of vascular bundle sheaths (3),
density of scales (5), scale translucence (12), lamina
base (17), presence of indumentum (19), visibility
of veins (23), and sori distribution (28). Thus it is
thus clear that M. glossophyllum constitute a distinct
species, as Bosman has suggested3.
Likewise, the dendrogram of cluster analysis us-
ing only 25 quantitative characters (Fig. 3) showed
the separation of M. siamense and M. steerei into
two subgroups. The result of canonical discriminant
analysis, also, using the same set of data (Fig. 4a)
showed a distinction between M. siamense (V) and
M. steerei (VI). It can be seen from Fig. 4a that
the phenetic gaps between groups are larger than
those within groups. This finding agrees with the
results of morphological studies based on qualitative
characters (Table 3) that the two taxa differ in at least
7 characters, i.e., rhizome surface (2), attachment of
scales (4), frond colour when living (15), lamina base
(17), margins of lamina (18), the presence of distinct
hydathodes (26), and spores colour (30).
It was earlier suggested that M. steerei might be
a reduced form or a variety of M. punctatum1. Our
results based solely on qualitative characters (Table 3)
and cluster analysis using both qualitative and quanti-
tative characters (Fig. 2) agreed well with this sugges-
tion. However, cluster analysis using only quantitative
characters (Fig. 3) and canonical discriminant analysis
using the same data set (Fig. 4a) showed distinction
between these two species. This discrepancy needs
further investigation, especially on molecular data,
which is beyond the scope of this study.
The 20 taxa groupings: Twenty taxa (Table 1) were
used as a priori group. Twenty-five quantitative
characters were used in this analysis (Table 2). The
canonical variable 1 is 99.1% correlated with 20
characters and the variance explained by it is 76.6%.
It is most highly associated with stipe length (3).
Canonical variable 2 accounted for 11.3% of the total
variance which is the axis most highly associated with
DSO (13), DSL (18) and NSR (21). According to
the stepwise discriminant analysis five variables, viz.
LML (2), RHD (7), SCW (10), DSR (14), and SPW
(17) were not selected for further use in canonical
discriminant analysis (Table 2).
The canonical plot on two canonical axes
(Fig. 4b) also shows the separation of 694 herbarium
specimens of the 20 taxa (Table 1) into 7 groups
on canonical axis 1, but is not clearly distinct on
canonical axis 2. Specimens of M. thailandicum,
www.scienceasia.org
8ScienceAsia 38 (2012)
M. membranaceum,M. musifolium and M. siamense
were included to group 1(), 2(), 3(J) and 5(),
respectively, whereas group 4() is comprised of
all specimens of M. glossophyllum and its previous
treated synonyms3. All specimens of M. steerei,
Polypodium tonkinense and P. playfairii were mixed in
group 6(o). Likewise, members of the 11 previously
treated synonyms of M. punctatum13were pooled
into the largest group, i.e., group 7().
Based on the results of cluster analysis together
with canonical discriminant analysis, it is reasonable
to divide the 20 taxa (Table 1) of the M. punctatum
complex into seven distinct taxa, viz. M. thailandicum,
M. membranaceum,M. musifolium,M. siamense,
M. glossophyllum,M. steerei, and M. punctatum,
respectively.
Boxplots of the four most important characters,
namely stipe length (STL), density of sori/cm2(DSL),
number of sori rows between closest secondary vein
(NSR), and diameter of sori (DSO) that collectively
segregated these seven taxa are presented in Fig. 5.
Classificatory discriminant analysis
The stepwise discriminant analysis suggested that
twenty-one of the twenty-five quantitative characters
were essential for the division of the seven taxa
(Table 2).
The linear discriminant function classification re-
sult appears to be 100.0% correct. For this reason,
the linear discriminant function (Table 4) can be used
to identify unnamed specimens in the M. punctatum
complex. To employ the discriminant function in
Table 4 for identification, it is necessary to multiply
each character score by its coefficient in each column.
Then the total in each column is calculated, the
column with the highest total is the taxon to which
the specimen belongs. This method of identification
is different from traditional keys; however, it can be
applied in a complementary manner.
QUALITATIVE CHARACTERS
Table 3 shows the summary of thirty-one qualitative
characters of the seven recognized taxa. Rhizome
scales character (the presence of hyaline marginal
region) is useful in separating the Microsorum punc-
tatum complex into 2 groups. Group I is composed
of 2 species, namely M. membranaceum and M. musi-
folium. Scale attachment, shape and texture of lamina,
and spore surface are valuable characters to segregate
each species. Group II included all specimens of
M. punctatum,M. glossophyllum,M. steerei,M. thai-
landicum and M. siamense, the last two species can
be separated from the other species by the presence
Stipe length (mm)
200
150
100
50
0
595
167
523
542
552
514
VIIVIVIVIIIIII
Page 1
VIIVIVIVIIIIII
Number of sori rows
15
12
10
8
5
2
630
673
675
677
679
568 570
583
569
579 584
587
574
590
385
356
306
345
381
397
4154
29
58
504 523
526
Page 1
Density of sori / sq. cm
80
60
40
20
0
676 625
657
619
652
314
569
570
477
VIIVIVIVIIIIII
Page 1
VIIVIVIVIIIIII
Diameter of sori (mm)
4
3
2
2
2
1
0
64
615
619
304
147 2
306
317
324
470
Page 1
Fig. 5 Boxplots of the four most important characters of the
M. punctatum (L.) Copel. (I =Microsorum thailandicum,
II =M. membranaceum, III =M. musifolium, IV =
M. glossophyllum, V =M. siamense, VI =M. steerei, and
VII =M. punctatum).
of iridescent blue-green fronds. M. glossophyllum can
also be separated from M. punctatum by using charac-
ter of lamina indumenta and sori distribution pattern.
It seems likely that only quantitative characters are
not clearly placed the seven taxa into distinct taxa.
www.scienceasia.org
ScienceAsia 38 (2012) 9
Table 4 Classification Function Coefficients of seven clustering groups obtained from cluster analysis based on 21
quantitative characters.
Character Categories
I II III IV V VI VII
STL 0.39 3.90 0.53 0.01 0.85 1.43 2.70
STD 1.21 7.83 2.91 0.42 2.57 4.57 3.04
DMP 1.51 3.62 2.21 1.90 2.48 4.36 0.91
PDL 0.31 5.84 2.85 1.55 0.87 0.75 2.91
RDL 0.40 0.76 2.00 2.39 1.06 0.94 1.75
SCL 2.54 2.01 3.61 4.33 0.64 4.40 1.27
SCW 3.51 0.31 7.12 0.53 10.06 1.81 0.89
ALA 0.11 0.34 1.01 0.79 1.08 0.81 0.28
ALB 0.29 1.22 2.55 0.61 0.29 0.37 1.14
DSO 34.49 46.71 37.64 47.31 35.85 36.55 35.08
NAC 22.50 13.95 15.33 15.60 15.91 16.56 14.76
SPL 1.21 1.70 1.49 2.17 1.56 1.47 1.63
SPW 1.70 1.57 1.46 1.04 1.52 1.51 1.21
DSL 0.60 0.71 1.05 0.77 0.58 0.89 0.98
PLS 0.01 0.02 0.01 0.01 0.02 0.01 0.00
PHS 0.38 0.61 0.90 0.71 0.57 0.72 0.61
NSR 1.94 0.31 0.66 0.14 0.57 0.44 1.69
DBSV 2.87 0.54 1.51 0.52 0.45 0.69 0.33
APS 1.95 1.13 1.23 1.32 1.23 1.91 1.68
SAW 1.06 3.16 0.11 0.90 1.11 1.66 2.84
PAW 2.98 1.54 0.88 2.40 2.76 2.38 4.26
(Constant) 497.09 569.17 363.41 359.65 325.02 394.29 445.24
I=Microsorum thailandicum, II =M. membranaceum, III =M. musifolium, IV =M. glossophyllum, V =
M. siamense, VI =M. steerei, and VII =M. punctatum.
Table 5 Means and standard deviation of 25 quantitative characters of the seven clustering groupings of the M. punctatum
(L.) Copel. complex.
Character I II III IV V VI VII
mean ±SE mean ±SE mean ±SE mean ±SE mean ±SE mean ±SE mean ±SE
LMW 16.97 1.48 72.64 26.81 96.53 16.45 77.66 10.77 37.63 3.33 39.33 10.58 57.84 18.38
LML 304.54 52.37 536.25 184.31 831.55 156.73 958.36 134.75 230.62 3.11 317.22 66.39 735.91 437.00
STL 34.19 1.05 138.77 6.47 0.48 0.11 19.28 1.65 42.01 1.27 62.34 4.61 101.38 6.96
STD 2.92 0.18 4.75 0.97 5.75 1.04 5.37 1.21 2.49 0.11 1.54 0.36 4.53 0.92
DMP 2.84 0.29 3.46 1.05 5.56 1.33 6.19 1.43 1.79 0.11 3.01 0.55 4.39 1.06
PDL 3.41 0.32 2.67 0.88 6.13 1.25 5.87 1.45 3.19 0.39 3.28 1.15 4.40 1.27
RHD 4.93 0.25 5.56 1.47 6.98 1.40 7.51 1.51 3.63 0.32 4.99 0.88 5.74 1.25
RDL 1.63 0.19 2.83 1.32 12.45 2.99 11.60 2.70 4.00 0.20 3.48 1.64 8.20 1.88
SCL 3.02 0.20 4.42 1.34 3.64 0.40 4.01 0.64 2.60 0.27 4.03 1.03 3.30 0.62
SCW 1.15 0.08 1.46 0.36 1.42 0.21 1.25 0.18 1.24 0.06 1.04 0.19 1.18 0.24
ALA 7.44 1.24 14.25 4.36 23.16 5.17 19.32 3.56 19.33 0.82 16.71 5.53 12.96 3.03
ALB 1.78 0.83 2.02 1.09 18.93 5.98 6.04 1.52 2.50 0.55 1.50 0.66 4.20 2.52
DSO 1.31 0.06 1.93 0.39 1.39 0.15 1.85 0.41 1.37 0.04 1.33 0.13 1.28 0.17
DSR 0.18 0.01 0.19 0.02 0.16 0.01 0.19 0.02 0.18 0.01 0.16 0.03 0.17 0.15
NAC 28.33 1.80 19.41 1.76 18.90 0.75 19.78 0.97 20.83 0.75 21.56 1.37 19.26 1.00
SPL 45.22 1.89 58.75 8.14 47.71 4.89 57.97 4.68 51.25 1.37 50.64 4.16 51.70 4.11
SPW 31.94 3.70 36.60 5.57 33.33 3.62 32.53 3.94 33.75 1.37 33.27 3.98 31.39 3.11
DSL 19.11 2.37 9.87 3.78 39.50 6.59 17.34 4.63 12.00 1.26 25.97 7.92 38.42 9.97
PLS 138.91 17.19 196.78 113.64 421.81 112.53 517.69 144.79 59.44 2.81 156.98 43.55 381.50 120.39
PHS 5.01 1.75 16.79 6.07 20.05 4.38 17.66 7.57 11.22 1.15 16.90 7.13 14.38 6.62
NSR 7.22 0.67 2.24 0.46 8.08 1.47 4.01 0.87 3.33 0.52 4.10 0.88 9.35 1.96
DBSV 16.22 0.63 10.03 2.36 13.27 1.95 8.66 1.68 7.10 0.64 9.05 1.87 13.27 2.63
APS 49.78 1.72 21.51 3.57 27.74 3.60 29.10 4.84 29.50 1.52 44.73 6.04 34.02 5.43
SAW 1.80 0.13 6.80 1.58 7.28 1.40 6.94 1.43 4.29 0.27 3.96 0.78 6.64 1.95
PAW 1.09 0.11 4.14 1.97 4.24 1.33 3.11 1.73 1.46 0.13 2.16 0.59 1.10 0.56
I=Microsorum thailandicum, II =M. membranaceum, III =M. musifolium, IV =M. glossophyllum, V =
M. siamense, VI =M. steerei, and VII =M. punctatum.
www.scienceasia.org
10 ScienceAsia 38 (2012)
Qualitative and quantitative characters were therefore,
collectively used to construct a key to these seven
segregated taxa.
CONCLUSIONS
The results from both cluster and discriminant anal-
yses revealed that M. membranaceum,M. siamense,
and M. thailandicum are distinct taxa. Likewise,
M. musifolium and M. glossophyllum, which were
previously included as synonyms of M. punctatum
(L.) Copel. by Nooteboom4, should be treated as
two distinct species as suggested by the results from
our study. This study supports the recognition of
M. steerei and its previous treated synonyms, i.e.,
Polypodium tonkinense, and P. playfairii. Finally, we
also found that the other synonymous species and
infraspecific taxa related to M. punctatum (Table 1)
should be placed in M. punctatum.
To sum up, the results from our study provide jus-
tification for the recognition of seven species, namely
M. thailandicum,M. membranaceum,M. musifolium,
M. siamense,M. glossophyllum,M. steerei, and
M. punctatum within the M. punctatum complex.
IDENTIFICATION LIST
Only specimens with an identifiable collector and
collection number are mentioned. Numbers after the
colon refer to the species numbers as given in the
following list. (T) denotes a type specimen.
1. Microsorum steerei
2. M. siamense
3. M. thailandicum
4. M. membranaceum
5. M. glossophyllum
6. M. musifolium
7. M. punctatum
Abeysiri 55: 4 — Abraham 666: 7 — d’Alleizette
herb 460 T: 1 — Allen 7, 173: 7 — Andrews 108: 7 —
Antun-Cupffert 337: 7 — Ashton 19 060: 7 — Avon
370: 7.
B200091734: 1 B 200 099 607, 200 099 652,
200 099749: 7 — J.M. Baker 84: 7 — Bakhuizen van
den Brink 5739: 7 — Bakshi 207: 7 — Balansa 45,
70, 107 (T), 148 (T), 198: 1; 1990: 4 — van Balgooy
2329, 4628: 7 — Balker 339: 4 — Balslev 342: 7 —
Banerji et al 1313, 2604, 26 957: 4 — Barrett (&Dorr)
201: 7 — Beckett 648: 7 — Beddome 48/341: 7 (T);
67, 101, 159, 177, 339: 4; 1911: 6; 1991: 7 — Beer
7768: 7 — Ben 438: 7 — Benson 106, 1293: 7 —
Bernhardi 234: 6 — van Beusekom et al 258, 683:
7 — Bhargava et al 2347, 2836, 6356: 7— Bidgood
4775: 7 — Blackwood 188: 5 — Bliss 41, 51, 189:
4 — Bloembergen 18, 3424: 7 — Blume (n.v.): 6
(T)— Bon 200, 1274: 1; 2119: 7 — Boonkerd 1442:
3 (T) — van Borssum Waalkes 3053, 3228: 7 — Bos
4106, 4218: 7 — Botavae 74: 7 — Bourdy 306: 7 (T)
— Braker 4136: 7 — Braithwaite 2306, 2570, 4136,
4370: 7; 4721, 4866: 5 — Brass 535, 673, 2756:
7; 11 319, 23 055: 5; 24 220: 7; 24 483: 5; 25 458,
29 373: 7; 27 212, 29 549, 29 786, 30 498, 31 569,
32 403: 5 — Brooks 357: 6; 416: 7 — Brownlie 1304,
1384, 8454, 16 074: 7 — Bryan 1114, 1167: 7 —
Bulmer 103 837: 5 — Bunk 384: 7 — B¨
unnemeijer
1848, 3519, 4315, 12 427: 7 — Buwalda 4159, 6052,
6336, 6978: 7.
Cadet 3824: 7 — Cadi`
ere 30: 1; 98: 7 — Carr
7660: 5; 12 148: 7; 13 015, 13 340: 5 Caruallo
2279, 4235: 7 — Castro 5910: 7 — Cavalerie (&
Fortunat) 2635:7; 3390: 4; 4012: 7 — Chapman 581,
3132: 7 — Chase 5220: 7 — Chevalier 21 088: 7 —
Christ 1940: 1 — Christensen 529: 7; 1339, 3418: 1
— Chusan 1847: 7 — Clarke 8739, 21 388, 27 186,
33 720: 4; 40 735, 43 399: 7 Clemens & Clemens
7133: 5 — Combes 4097: 7 — Conn 152: 5; (&
Kairo) 157: 5 — Coode 6251: 7 — Cooks 1909: 7
— Copeland 275: 7; 388: 5 (T); 1535: 7; 1537: 6;
1776, 15 356: 7 — Corner 30 247: 6 — Coveny 6900:
7 — Croft 151:5; 199: 4; 203, 451, 533, 568:5; 1129:
7; 1728:5; 61 266, 61 160, 61 578, 65 453: 7; 65 719:
5 — Cusclah 17 347: 7.
Darbyshire 624: 7 — Darnaedi 71, 2107: 7 —
Dawkins 389: 7 — Decary 17 754: 7 — Deighton
6056: 7 — Deplanche 23, 198: 7 — Dickason 7637,
7986: 7; 8118, 9029: 4 — D¨
ummer 472: 7.
EBL 1537: 6 — Edelfelt 220: 5 — Edward 38,
2411: 1 — Elbert 913: 7 — Elmer 5873: 4; 5884 (T),
7854, 7991, 8263: 7 ; 8367: 4; 9946: 7; 10 500: 6;
10 920, 13 813, 13 598, 16 863: 7; 20 871: 6; 22 330:
7 — Endert 1889, 2358: 7; 4022: 6; 15 062: 7 (T)—
Ernst 11 045: 7 — Esquirol 2245: 4; 3601: 7 — Eyma
3254: 7.
Faden 69–946: 7— Flenley 2084: 5 — Floto
7237: 7 — Floyd et al 5682: 7; 5974, 6820: 5 —
Forster 10 852: 7 — Fosberg 14 149: 7 — Foxworthy
42 135: 7 — Franck 11 448: 7 — Fris et al 7120: 7 —
Fung 20 053: 7.
Gamble 884, 1925, 4000, 4847, 6366, 6367,
9699, 14 409, 14 870: 4; 16 350: 7 Gardner 1145,
1298: 4 H.B.G. Garrett 59, 59b: 4; 288: 7; 391: 4
— Gay 132: 6; 405, 1031: 7; 1806: 5 — Germain 40:
7 — Giesenhagen 1910: 1 — Glover 263: 7 — Gough
3243, 6055, 8289, 16 350: 7 — Gutierrez 117367: 7
— Gutzwiller 1305: 7.
Hafashimane 26, 357: 7 — Haines 5379: 4 —
Hancook 342, 357, 1892: 6 — Hartley 11 536: 7 —
Haniff 21 028: 6 — Henderson 19 704, 19 708: 6 —
www.scienceasia.org
ScienceAsia 38 (2012) 11
Table 6 Key to species: The following is a simplified key to identify taxa in the M. punctatum (L.) Copel. complex based
on the output obtained from DELTA12 .
1a. Fronds linear to narrowly elliptic, lamina base abruptly narrowed, living fronds iridescent
blue-green; sori mostly irregularly scattered on simple free or on 2 or 3 connecting veins . . . . 2
1b. Fronds linear to ovate or narrowly obovate to narrowly oblanceolate, living fronds light to
dark green; sori forming irregular rows parallel to each pair of secondary veins . . . . . . . . . . . . . 3
2a. Lamina up to 4 cm width; scale peltate; hydathodes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. siamense
2b. Lamina less than 2 cm width; scale pseudopeltate; hydathodes absent . . . . . . . . . . . . . . . . . . . . . . M. thailandicum
3a. Rhizomesurfacenotwaxy............................................................ 4
3b. Rhizome surface at least sometimes waxy under the scales or often waxy . . . . . . . . . . . . . . . . . . 5
4a. Scales pseudopeltate;stipepresent..................................................... M. membranaceum
4b. Scalespeltate;stipeabsentorobscure.................................................. M. musifolium
5a. Frond up to 100 cm long; stipe present, more than 2.0 mm diam . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5b. Frond less than 40 cm long; stipe absent or obscure, less than 1.5 mm diam . . . . . . . . . . . . . . . . M. steerei
6a. Distance between lowest sori and frond base up to 50 cm; all veins distinct; scales opaque
andblackish......................................................................... M. glossophyllum
6b. Distance between lowest sori and frond base less than 35 cm; all veins or secondary and
smaller veins more or less sunken and obscure (at least in living specimen); scales translucent
andbrownish........................................................................ M. punctatum
Hennipman 3065, 5112, 5462, 5981: 7 — Henry 339:
4; 1895: 1, 1951: 1; 10 899: 7 — Hepper 8682: 7 —
Heward 183: 7 — den Hoed 909: 7 — Holstvoogd
772: 7 — Holttum 15 702: 7 — Hoogland et al 4497,
6877: 5; 10 588: 7 — Hooker 526: 7; 1145, 1298 : 4;
1803, 1820, 1867, 3799: 7 — E.G. Hose & G. Hose
1822, 1827, 1894: 6; 4823: 7 — Humblot 666: 7 —
J. Hutchison 1139: 7.
Inaeteay 173: 7 — Inder 4022: 6 — Ismail 2744:
7 — Iwatsuki et al 3252: 6; 9600: 4; 10 900: 7;
15 642: 4.
Jackson U123: 7; 51: 5 — Jacobson 10: 7
— Jacquemont 600: 4 — Jacques-Fe’lix 864: 7 —
Jalconer 68: 4 — Jaman 4036: 6 — Jarrett 68, 766,
784: 7 — Jati 10: 4; 875: 7 — Jeffrey 208: 7 — Jermy
(& Rankin) 3573, 3628, 8220: 7 — J.D.H. 750, 2223:
7 — J.J. 6061: 4 — Johns 7995: 5 — de Joncheere
1113, 1325: 7 — Jones & Onochie FHI 16952: 7.
Kampu 1, 2, 3, 4, 5: 1 — Kandau 62 458: 7 —
Katendo 1187: 7 — Kato et al B3252: 6 — Keke
902: 4 — Khasya 1867: 4 — Khwaunju 1259: 4 —
G. King’s collector (=Kunstler) 192: 6; 5069: 7 —
Køie & Sandermann Olsen 1149: 7 — Konar 56: 4 —
Korthals 33, 113, 148, 165, 185, 196, 273, 527, 684,
973: 7 — Kostermans 59: 7 — Kulong 11 582: 7.
Lace 4894: 4 — Lam 1108: 7 (T); 1365: 5; 3717:
7 — Larsen et al 2314: 4; 2597, 3096, 3374, 5078: 7
— Laumonier 876: 4; 1796: 6; 6150, 9085, 10 720,
26 980: 7 — Lauterbach 567: 6 — Lecerber 2042,
2617: 7 — Ledermann 7695, 8549, 8743: 6 — van
Leeuwenberg 1785, 2542, 5032, 6651: 7; 9853: 5;
10 647, 13 492: 7 — Leland et al 65 641: 5 — Leonard
1618: 7 — Linder 759: 7 — Littke 394: 7; 469: 6 —
Loher 867: 1 — Louis et. al. 950, 1417, 1932: 7 —
Lucas et al 230: 7 — Lungchow 83: 7 — L ¨
utjeharms
4750, 4990, 4999, 5019, 5151, 5251, 5293: 7.
Maddine 1867: 4 — Madhusoodanan CU 29683:
4 — Maitban 281: 7 — Manickam 31 220: 7; 31 442:
4 — Marie 10 775: 7 — Matthew 2, 4, 1907, 1928: 7;
1967: 4 — McDonald 3829: 5 — Melsetter 46 915:
7 — Melville et al 2023, 3669: 7 — Mense 343: 4
—Merrill 7331: 7; 11 691: 4 — van Mettenius 276:
5 — Meyer 7997: 7 — Mitchell 378: 7 — Mildbraed
198, 4426, 6524: 7 — Miller 1364: 7 — Mitchell 132,
154: 7 — Mooney 128: 4; 2383: 7 — Mooze 30: 7
— Moseley 1874, 3412 (T), 6447: 7 — Mousset 20,
7166: 7 — Murata et al 16 387, 17 674: 7.
Nakaike 408: 5 — N.C. Nair (BSI-series) 51 452:
4 — Narasimtan 165 111: 4 — de N´
er´
e 332, 1412:
7 — Nooteboom 5915: 7 — Noumea 29, 199: 7 —
Nuttall 1867: 7.
Pannell 11 385: 7 — Pancher 186, 506: 7 —
Parris 6900: 7; 7751, 9251, 9479: 5; 11 051, 11 720:
7 — Pascal 923: 7 — Paul 64 665: 7 — Paush 1931:
7 — Perrier 1747, 6149: 7 — P´
etelot (Colani) 1339,
1789: 1; 2898, 4101: 7; 4871, 4911: 1 — Phengkhlai
683: 7 — Piggott 2103, 2973: 7 — Phustouve 34: 7
— Playfair 383: 1 (T) — Pleyte 57, 265: 7 — Preuss
2: 7 — du Puy 7: 7 — P.V. fern 1: 2 (T).
Ramamoorthy 256: 4 — Ramos 973, 14 779,
14 862, 31 419: 7 — Rehmann 8674: 7 — Ridsdale
5567: 7 — Ridley 1917, 6554, 8935: 7 (T)— Robin-
son 1954: 7 — Rodin 177, 245, 569: 7 — Rochers
2634: 7 — Rosenstock 20, 99 678: 7 — van Royen
3474, 4617: 7 — Rudatis 1369: 7.
Saldanha 421, 641, 717, 820, 14457, 14 800,
www.scienceasia.org
12 ScienceAsia 38 (2012)
15 068, 17 959: 4; 12 517, 16 392: 7 Sands 1780:
5; 2118, 2695, 6730: 7 — Sangster 630: 7 — Savatier
987: 7 — Savi 340: 7 — Schelpe 5032, 5225: 7 —
Schlechter 2764: 6; 16 304: 7 — Schmitz 5169A,
F7: 7 — Schodde 3026: 7 — Schultze 104: 7 — M.
Semesle 580: 7 — Seemann 728: 7 — Sermolli 5232,
7219, 7244: 7 — Shimizu 10 102: 4 — Sinchal 339:
4 — Skinn 283: 7 — Skinner 4828: 4 — Sledge 543,
832: 4 — Sloover P195: 7 — Smith, J. s.n.: 7 ; 1187:
4; 1859: 6 — S.P. 2, 6: 3; 20, 26, 39, 60, 62: 2; 49, 64,
71, 88, 120, 121: 3 — Stachey & Winterbottom 1: 4
— Steere s.n.: 1 (T) — Stevens 58 710: 7 — Steward
et al 21 169: 4 — Sunanda 9557: 7 — Surbeck 1082:
7 — Suruhoe 82: 1.
Tagawa 1853: 7 — Tchinaye 89: 7 — Tem
11 209: 7 — Tessier 19067: 7 — Thollon 1304: 7
— Thomas 145, 1369, 11 536: 7 (T) — Thorold TN6,
28, 87: 7 — Topping 4200: 4 — Treutler 246, 661: 4
— Tsiang 29 192, 36 090: 7 — Turneau 836: 6, 904:
7; 905: 6 — Tutcher 10 771: 7 — Tweedie 2432: 7.
Unesco Limestone Exp. 635: 6.
Vanoverberg 3678: 6 — Veldkamp & Stevens
5911, 6793: 5 Verdcourt 147, 3919: 7; 5113: 5
— Vesco 1847: 7 — Viane 16, 828: 7 — Vidal 4041:
7 — Vieillard 459, 10 775: 7 — Vinas 5974, 60 249:
5 — Vink 16534, 17 568: 5 — de Vriese 26, 32:7; 60:
6; 325: 7.
Wace 4, 42: 7 — Walker 25: 4; 132: 7 —T.G.
Walker T7884, T12316: 7 — Wallace 191: 7 —
Wallich 273, 281: 7 (T); 282: 4; 1837: 7 — Wakefield
1435: 7 — Watt 101 087: 4 — Wenzel 1216, 2611:
7 — de Wilde et al 3734, 3876: 7; 12 385: 6 —
Whitmore 432: 7; 1045: 5 — Widjaja 4293: 5 —
Womersley, J.S.6820, 11 092: 5.
Yapp 296: 7; 575: 6 — Ying 1657: 7.
Zollinger 935, 1028, 1028: 7; 3005: 6.
Acknowledgements:We are indebted to the directors
and curators of B, BCU, BKF, BM, K, L, and P for
making their collections available for this study. This
work is partially supported by a grant from Chulalongkorn
University, Bangkok: the Centre of Excellence in Biodi-
versity, Chulalongkorn University graduate scholarship to
commemorate the 72nd anniversary of His Majesty King
Bhumibol Adulyadej, the National Research University
Project of CHE and the Ratchadaphiseksomphot Endow-
ment Fund (CC270A) and the TRF/BIOTEC Special Pro-
gram for Biodiversity Research and Training program (grant
No. T 145007).
REFERENCES
1. Nooteboom HP (1997) The microsoroid ferns (Polypo-
diaceae). Blumea 42, 261–395.
2. Bosman MTM (1991) A Monograph of the Fern Genus
Microsorum (Polypodiaceae). Rijksherbarium/Hortus
Botanicus, Leiden, the Netherlands.
3. Bosman MTM, Hovenkamp PH, Nooteboom HP
(1998) Microsorum Link. In: Kalkman C, Noote-
boom HP (eds) Flora Malesiana. Rijksherbarium/Hor-
tus Botanicus, Leiden, the Netherlands, pp 90–131.
4. Holttum RE (1965) Ferns of Malaya. A Revised Flora
of Malaya. Vol. 2. Govt. Printer, Singapore, pp 1–643.
5. Boonkerd T (2006) A new species of Microsorum
(Polypodiaceae) from Thailand. Blumea 51, 143–5.
6. Boonkerd T, Nooteboom HP (2001) A new species of
Microsorum (Polypodiaceae) from Thailand. Blumea
46, 581–3.
7. Holmgren PK, Holmgren NH (1998) Index Herbario-
rum: A global directory of public herbaria and associ-
ated staff. http://sweetgum.nybg.org/ih/.
8. Gower JC (1971) A general coeflicient of similarity and
some of its properties. Biometrics 27, 857–71.
9. Sneath PHA, Sokal RR (1973) Numerical Taxonomy:
The Principles and Practice of Numerical Classifica-
tion. W. H. Freeman and Company, San Francisco,
USA.
10. Anonymous (1999) SPSS for Windows Release 10.0,
Standard version [Computer software], SPSS Inc,
Chicago.
11. Dallwitz MJ, Paine TA, Zurcher EJ (1993) User’s
Guide to the DELTA System: a General System for Pro-
cessing Taxonomic Descriptions. http://biodiversity.
uno.edu/delta/.
12. Tagawa M, Iwatsuki K (1989) Polypodiaceae. In:
Smitinand T, Larsen K (eds) Flora of Thailand.
Chutima Press, Bangkok, pp 486–580.
www.scienceasia.org
... Microsorum punctatum (L.) Copel (Polypodiaceae) is a common fern species found in Africa and Asia, in various forest types of tropics and subtropics from sea level up to 2800 m elevation [8,9]. The plant is a common medium sized epiphytic herb occurring naturally in semi exposed as well as shaded sites. ...
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A new species, Microsorum siamensis, is described and illustrated.
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The tremendous advances in electronic data processing are likely to result in revolutionary changes in the theories and practices of taxonomy. The process of classification is being removed from speculations regarding the origin of the taxa being classified. A natural classification is one whose taxa share the largest number of properties and which is most useful for a wide range of purposes. The principles of numerical taxonomy are stated briefly and illustrated by means of diagrammatic examples. The relative roles of the taxonomist and computer are discussed and estimates given of computer time and costs involved in numerical taxonomic work. The numerical taxonomic work done in botany so far is discussed and the paper concludes with a brief mention of several problems of numerical taxonomy with regard to botanical work. These are: scarcity of characters, correlations between cytogenetic work and phenetic similarities, and problems raised by hybridization.
Book
The DELTA system is a flexible data-coding format for taxonomic descriptions, and an associated set of programs for producing and typesetting natural-language descriptions and keys, for interactive identification and information retrieval, and for conversion of the data to formats required for phylogenetic and phenetic analysis. This manual is a comprehensive guide to the data format and the program directives.
Article
A general coefficient measuring the similarity between two sampling units is defined. The matrix of similarities between all pairs of sample units is shown to be positive semidefinite (except possibly when there are missing values). This is important for the multidimensional Euclidean representation of the sample and also establishes some inequalities amongst the similarities relating three individuals. The definition is extended to cope with a hierarchy of characters.
1274: 1; 2119: 7 — Boonkerd 1442: 3 (T) — van Borssum Waalkes 3053
  • Bhargava
Bhargava et al 2347, 2836, 6356: 7— Bidgood 4775: 7 — Blackwood 188: 5 — Bliss 41, 51, 189: 4 — Bloembergen 18, 3424: 7 — Blume (n.v.): 6 (T)— Bon 200, 1274: 1; 2119: 7 — Boonkerd 1442: 3 (T) — van Borssum Waalkes 3053, 3228: 7 — Bos 4106, 4218: 7 — Botavae 74: 7 — Bourdy 306: 7 (T) — Braker 4136: 7 — Braithwaite 2306, 2570, 4136, 4370: 7; 4721, 4866: 5 — Brass 535, 673, 2756: 7; 11 319, 23 055: 5; 24 220: 7; 24 483: 5; 25 458, 29 373: 7; 27 212, 29 549, 29 786, 30 498, 31 569, 32 403: 5 — Brooks 357: 6; 416: 7 — Brownlie 1304, 1384, 8454, 16 074: 7 — Bryan 1114, 1167: 7 — Bulmer 103 837: 5 — Bunk 384: 7 — Bünnemeijer 1848, 3519, 4315, 12 427: 7 — Buwalda 4159, 6052, 6336, 6978: 7. Cadet 3824: 7 —Cadì ere 30: 1; 98: 7 — Carr 7660: 5; 12 148: 7; 13 015, 13 340: 5 — Caruallo 2279, 4235: 7 — Castro 5910: 7 — Cavalerie (& Fortunat) 2635:7; 3390: 4; 4012: 7 — Chapman 581, 3132: 7 — Chase 5220: 7 — Chevalier 21 088: 7 — Christ 1940: 1 — Christensen 529: 7; 1339, 3418: 1