Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 June 2020 | 12(9): 16064–16076
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
doi: https://doi.org/10.11609/jott.5004.12.9.16064-16076
#5004 | Received 12 April 2019 | Final
received 11 May 2020 | Finally accepted 06 June 2020
Resolving taxonomic problems in
the genus Ceropegia L. (Apocynaceae:
Asclepiadoideae) with vegetative micromorphology
Savita Sanjaykumar
Rahangdale 1 & Sanjaykumar
Ramlal Rahangdale 2
1 Department of Botany, B.J.
College, Ale, Pune District, Maharashtra 412411, India.
2 Department of Botany, A.W. College,
Otur, Pune District, Maharashtra 412409, India.
1 gauriyana@yahoo.co.in, 2 rsanjay2@hotmail.com
(corresponding author)
Editor: M.K. Vasudeva
Rao, Shiv Ranjani Housing Society, Pune, India. Date of
publication: 26 June 2020 (online & print)
Citation: Rahangdale, S.S. & S.R. Rahangdale
(2020). Resolving
taxonomic problems in the genus Ceropegia L. (Apocynaceae: Asclepiadoideae)
with vegetative micromorphology. Journal of Threatened Taxa 12(9): 16064–16076. https://doi.org/10.11609/jott.5004.12.9.16064-16076
Copyright: © Rahangdale & Rahangdale 2020. Creative Commons Attribution 4.0 International
License. JoTT
allows unrestricted use, reproduction, and distribution of this article in any
medium by providing adequate credit to the author(s) and the source of
publication.
Funding: Self-funded.
Competing interests: The authors declare no competing interests.
Author details: Savita S. Rahangdale is presently Assistant Professor in Botany with an
experience of 20 years in teaching and research. Her major research areas are
Angiosperm taxonomy and medicinal plants.
Sanjaykumar R. Rahangdale
is Associate Professor in Botany. His research interests are, cytology,
genetics, taxonomy and ecosystem conservation. Both are associated with State
Department of Forests as Honorary Botanists towards diversity and conservation
studies.
Author contribution: SSR has done the micro-morphology work of this study
and presentation of the results. SRR has done the field work, collection and
processing of samples, morphological characterization of the taxa, photography
and manuscript editing. The work is done on a mutual benefit basis.
Acknowledgements: Authors are thankful to the authorities of BSI, Pune
and ARI Pune for herbarium references.
The thanks are due to Dr. Aniket Gade, Department of Biotechnology, SGBAU, Amravati for
cladistics analysis and help. The authorities
of respective institutions of authors are sincerely acknowledged for facilities
& support.
Abstract: The genus Ceropegia
L. of Family Apocynaceae, subfamily Asclepiadoideae comprises 213 accepted taxa distributed in
tropical and sub-tropical regions of Africa, India, Australia and neighbouring regions. The taxa are mainly identified on the
basis of flower morphology. A study was undertaken to reveal micromorphology of
26 taxa distributed in Western Ghats of India to solve the identity problems.
The micromorphology is studied with standard microscopic methods in five
replicates of each character and taxa to reveal the parameters, distribution of
stomata, stomatal index, stomatal density, epidermal cell wall pattern and
stomatal measurements. The data obtained was subjected to ANOVA to find out the
experimental mean, standard deviation and standard error. A consensus phylogeny
tree is constructed using the PAST on the basis of Jaccard similarity
coefficient. Results of study revealed that, micro morphological characters,
viz., type of stomata, number and characters of subsidiaries, anticlinal cell
wall pattern, and stomatal index are very significant in delimitation of
closely allied taxa. Beside ‘paracytic’ stomata, ‘tetracytic’, ‘isotricytic’, &
‘anomocytic’ stomatal types, and amphistomatic
distribution are recorded for first time in Ceropegia.
The taxa are separated easily from each other using vegetative micromorphology
and can be identified even in absence of flowers. An identification key and
phylogenetic tree is derived on the basis of vegetative micro-morphology and
gross morphological characters.
Keywords: Epidermal cell wall pattern,
identification key, phylogeny, stomata, stomatal index, subsidiaries.
Abbreviations: SD—Stomatal density | SI—Stomatal
index | WG—Western Ghats.
INTRODUCTION
The genus Ceropegia
L. of family Apocynaceae, subfamily Asclepiadoideae comprises many narrow endemic and
threatened species distributed only in tropical and sub-tropical regions
especially, Africa, India, Australia, and neighbouring
region. The genus comprises of 213
accepted species, eight unplaced, and 36 un-assessed taxa of species and
infra-specific ranks (Plant list 2019; Plant of the World Online 2020). Most of the taxa are endemic to the regions
or locations from where they are described.
In India, it is represented by about 68 taxa comprising of 61 species,
two subspecies and five varieties with recent descriptions by Diwakar & Singh
(2011), Rahangdale & Rahangdale
(2012), Kamble et al. (2012), Sujanapal
et al. (2013), and Kumar et al. (2018).
Of these taxa, 40 species and six varieties are endemic to India, while
34 species and two varieties are endemic to Western Ghats (Jagtap
& Singh 1999; Singh et al. 2015).
The taxa under this genus are
mainly described on the basis of floral characters, while many vegetative
characteristics are similar in many taxa.
The taxonomists in the world have tried to resolve systematic crises among
these species or taxa below rank of species.
Since last decade many new taxa at specific and infra-specific rank are
described (Singh et al. 2015). Still,
many of the Indian taxa are kept under unresolved categories by international
databases (such as The Plant List (2019) as well as POWO (2020)) due to insufficient data related to
allied species and varieties. In India,
there are two major regions of distribution of this genus, viz., Himalaya and
peninsular India. Most species found in
Western Ghats or peninsular India possess tubers, while those found in the
Himalayan region do not have tubers and are non-succulent. In general, all these taxa are described on
the basis of morphological characters; especially on floral morphology as the
allied taxa do have many similarities in vegetative characters. The vegetative micro-morphology was neglected
in most of the cases, may be because of morpho-similarities among the taxa in
vegetative condition. There is
negligible literature available on microscopic characters of vegetative organs;
and Ceropegia species are generally identified
on the basis of habit and flower characters.
Therefore, it is a difficult task to identify the species without
flowers. So, there is need of micro-morphological
as well as anatomical characters that would be helpful in identification of
taxa at species and infra-specific level in vegetative state. Considering these facts and lacunae in the
literature a study was undertaken to reveal micro-morphological characters of
taxa of this genus and provide identification key on the basis of vegetative
and micro-morphological characters.
MATERIALS AND METHODS
Plant Materials
Materials for the present study
comprise a total 26 taxa of the genus Ceropegia
L. from Western Ghats. Fresh materials
were collected from wild plants as well as plants maintained in the nursery for
study. The locations of taxa included in
the study range from Nashik in Maharashtra to Coimbatore in Tamil Nadu through
Wayanad in Kerala. A list of taxa under
study with their status, distribution, and locations of collection for the
present study is given in the Table 1.
In the present study, C. attenuata Hook. var. attenuata,
C. attenuata Hook. var. mookambikae
Diwakar & Singh, C. mahabalei Hemadri & Ansari var. hemalatae
Rahangdale & Rahangdale,
C. mahabalei Hemadri
& Ansari var. mahabalei, C. maharashtrensis Punekar et al. are
retained because these taxa are merged without any solid reasons and without
comparing the characters said in original protologues of above mentioned taxa.
As the present study is aimed to find out more stable characters for correct
distinction between the taxa, the above mentioned taxa are accepted.
Methods
Vegetative morphology
Total 14 vegetative
morphological characters of each taxon are recorded by observation of living
materials. They are mentioned in the
Table 4 to avoid repetition. Only newly
observed characters are mentioned in detail.
Micro-morphology
For epidermal study, five leaves
per sample were picked at 4–6th nodes from base. Epidermal peelings of suitable size were made
from both upper and lower surfaces of each leaf. Peelings were treated with 5% Sodium
Hypochlorite for clearing. Then
epidermal debris was brushed off under observation using Lawrence and Mayo
stereo-zoom microscope NSZ-606 and the peel was finally mounted in 50% Glycerine.
Thereafter, slides were observed under research microscope (Lawrence
& Mayo Model No. XSZ-N107T) for epidermal details. For this, five leaves were taken and from
each leaf five different fields were observed and data was recorded, making
total observation size of 25 sample areas.
The observed parameters are, type of stomata, distribution of stomata on
leaf surface (hypostomatic/amphistomatic),
stomatal density/mm2, stomatal Length (µ), Width (µ), pore length
(µ), characters and number of subsidiaries and epidermal cell wall characters
as per the techniques described by Dilcher (1974) and
Kotresha & Seetharam
(2000). The classification of stomatal
types given by Prabhakar (2004) was followed to recognize stomatal complex,
with number and characteristics of subsidiaries.
Parameters studied
1. Stomatal index (SI):
Stomatal index = [S ÷ (E+S)] ×
100
where, E = no. of epidermal
cells, S = no. of stomata in an unit (mm2) area taken for
observation (0.022mm2 under 400X magnification).
2. Stomatal Density (SD):
Stomatal Density = No. of
stomata per unit area (1mm2)
For this, number of stomata in a
unit area is recorded for total sample size as that for stomatal index
observations.
3. Stomatal length (μ):
The length of stomata is
measured as the longitudinal end to end distance of guard cells, using ocular
meter scale standardized for each magnification with the stage micrometer.
4. Stomatal width (μ):
This is the maximum length in
the middle transect of stoma with the ocular meter scale.
5. Stomatal pore length (μ):
This is length of stomatal
opening observed with the help of ocular meter scale.
Statistical analysis comprising of
mean, variance, standard deviation, and standard error are applied for each
parameter as per method given by Singh & Chaudhary (1985).
Phylogenetic analysis
Phylogenetic analysis was done
on the basis of 42 vegetative morphological and micro-morphological characters
(Table 4). For phylogenetic analysis
PAST ver.03 was used as per Harper (1999), Hammer et al. (2001), Hammer &
Harper (2006). In this, the characters
were represented in the binary format (Table 5), which was used to generate a
phylogenetic tree by Jaccard similarity coefficient model and data replication
method - bootstrap method at 1,000 replications on the basis of a matrix of
characters.
RESULTS
The observations regarding
stomatal distribution, type, subsidiaries character & number, and the
anticlinal wall characters of epidermal cells are presented in the Table 2 and
Image 1. The results are described under
the subheads as mentioned below.
Distribution and type of stomata
The stomata are distributed only
on lower surface of the leaf (hypostomatic) in all
the taxa studied except in C. bulbosa. In both varieties, C. bulbosa
var. bulbosa and C. bulbosa
var. lushii, the leaves are amphistomatic, i.e., stomata are present on both surfaces
of the leaves. Four types of stomata,
i.e., paracytic, anomocytic,
tetracytic, and isotricytic
are recorded in the taxa under study.
Among these types, the paracytic stomata are
most common and found in 16 taxa, the anomocytic are
less common and recorded in four taxa; while the tetracytic
stomata are observed in only C. elegans. Five taxa have mixed stomata; of which, four
taxa viz., C. anantii, C. attenuata
var. attenuata, C. maharashtrensis,
and C. sahyadrica have isotricytic
and tetracytic intermixed with each other, while in C.
rollae paracytic
stomata were found along with anomocytic stomata.
Characters and number of
subsidiaries
The subsidiaries in studied taxa
are either distinct (different than epidermal cells) or indistinct (same as
epidermal cells). The stomatal complexes
in 12 taxa have distinct subsidiaries while 13 taxa have indistinct ones. Among the 16 taxa having paracytic
stomata, 12 taxa have distinct subsidiaries while remaining four have
indistinct ones. In C. rollae, where stomata are of mixed type, i.e., paracytic and anomocytic, the
subsidiaries were distinct along the paracytic
stomata while with anomocytic stomata there were
indistinct subsidiaries. The number of
subsidiaries is also variable from two to five, but generally showed a fixed
number or a range in all the specimens of same taxon. The number is fixed to two subsidiaries in
nine taxa and four subsidiaries in two taxa.
In other 15 taxa, the number of subsidiaries ranges 2–3 in three taxa,
2–4 in three taxa, 3–4 in seven taxa, and 4–5 subsidiaries in two taxa with anomocytic stomata.
The taxa having mixed stomata with isotricytic
and tetracytic types have indistinct subsidiaries.
Anticlinal wall pattern
The patterns of anticlinal walls
of epidermal cells are observed and recorded in all the taxa under study. There are three patterns, ‘straight’,
‘rounded (curved)’, and ‘undulate (wavy)’.
Out of all 26 taxa, 10 have undulate anticlinal walls, eight taxa have
rounded walls, and remaining eight have straight anticlinal walls. The results showed that, anticlinal wall
pattern is not found to be associated with the other characters studied. This character is independent of stomatal
type, size or number and type of subsidiaries.
But, it is important for the differentiation between different taxa.
Stomatal characteristics
The experimental results about
stomatal measurements are presented in Table 3 and described below.
Stomatal Index (SI)
The results revealed that, mean
value of SI for the taxa under study (experimental mean) is 14.75±0.35. The highest SI is found in C. attenuata var. attenuata
(21.80) followed by C. mahabalei var. hemalatae (20.32), and C. sahyadrica
(19.94); while the lowest value of SI is recorded in C. juncea
(8.18) along with C. panchganiensis
(11.24), C. vincifolia (11.81), and C. hirsuta (11.88).
Stomatal Density (SD)
The mean value for stomatal
density/mm2 of leaf area is 252.31±8.50 with the highest value of
403.64±10.60 in C. maharashtrensis followed by
C. anantii (390.91), and C. jainii (370.91), while the lowest SD 98.18±3.40 in C.
juncea subtended by C. bulbosa
var. lushii (105.45), C. bulbosa var. bulbosa
(127.27), and C. hirsuta (152.73). C. juncea is
a succulent taxon with highly reduced leaves and also have lowest value of SI.
Stomatal Size
Regarding the stomatal
dimensions, the largest stomata are recorded in Ceropegia
juncea, while the smallest stomata in the C. attenuata var. mookambikae. The experimental mean for stomatal length is
29.54±0.73µ, stomatal width 19.68±0.67µ and the pore length 19.68±0.64
microns. The stomatal length is highest
in C. juncea (39.0µ), followed by C.
vincifolia (32.5µ), C. bulbosa
(32.25µ), C. oculata (32.0µ), and C.
candelabrum (31.5µ). The succulent
species C. juncea have largest stomata with
respect to length 39.0±1.5µ, width 26.5±1.7µ, and pore length 27.0±0.97µ; while
those values for C. attenuata var. mookambikae are 23.0±0.5µ, 15.0±0.79µ, and
12.0±0.5µ, respectively having the smallest stomata. As per the ratio of length x width, the
stomatal size is ranging from the smallest one of 345µ2 in C. attenuata var. mookambikae
to 1033.5µ2 in C. juncea has
largest stomata with the experimental mean value of 587.3µ2.
Phylogeny
A consensus phylogenetic tree
based on 42 morphological and micro-morphological characters obtained after
1,000 replications revealed that the dendrogram is divided into three clusters
(Figure 1). Cluster I comprises seven
taxa: C. maccannii, C. panchganiensis,
C. rollae, C. attenuata
var. attenuata, C. sahyadrica, C. maharashtrensis,
and C. anantii. All these taxa are clustered with unit
distance of 0.2–0.8.
Cluster II comprises total 15
taxa with unit distance of 0.2–0.7. All
the taxa of this cluster have paracytic stomata. This cluster further has two
sub-clusters. The sub-cluster IIa comprises eight taxa: C. attenuata
var. mookambikae, C. mahabalei
var. mahabalei, C. media, C. candelabrum,
C. evansii, C. fantastica, C. oculata,
and C. lawii.
Taxa in this sub-cluster have two distinct subsidiaries in all the
taxa. From the dendrogram it is observed
that the taxa, C. evansii, C. fantastica, C. oculata, and C. lawii
are closely related having unit distance of 0.2–0.4. Another sub-cluster IIb comprises seven taxa
having indistinct and variable subsidiaries: C. hirsuta,
C. vincifolia, C. odorata,
C. anjanerica, C. jainii,
C. mahabalei var. hemalatae,
and C. concanensis.
The Cluster III is an externally
linked group having four taxa with unit distance of 0.2–0.8. It comprised two varieties of C. bulbosa, C. elegans, and C.
juncea.
DISCUSSION
As the genus Ceropegia
is considered to be xerophytic because of its escape mechanism against the hot
period by perennial tubers. It should be
preferably hypostomatic and the results are in
corroboration with the concept, but the exceptions are both the varieties of C.
bulbosa as they have amphistomatic
leaves. Both the varieties of this taxon
are usually distributed in dry rain-shadowed area of the Western Ghats, even
though they are amphistomatic; this may be due to
more succulent leaves than other taxa.
Metcalfe & Chalk (1950) reported isobilateral leaves in the species
of Ceropegia and Hoya R.Br. having
fleshy leaves. Therefore, present
results also confirm the amphistomatic nature of some
taxa in Ceropegia which have fleshy
leaves. The presence of stomata only on
lower surface of leaf is an adaptation to reduce the rate of
transpiration. It is interesting to note
that except C. candelabrum, C. elegans, and C.
juncea, all the taxa have tubers and their
vegetative growth is confined to the rainy season only, but still they have hypostomatic leaves.
This is an interesting fact and raises further curiosities towards the
taxa. The taxa C. candelabrum, C. elegans, and C. juncea
generally remain in their active growth during summer season also and therefore
their hypostomatic nature is justified, but for other
taxa which grow in high rainfall area and escape the dry spell, especially
about 8–9 months of year except rainy season and still have the hypostomatic leaves.
This fact is interesting and the hypostomatic
leaves must be a qualitative character governed by genes only; because in the
present study beside C. bulbosa three more
taxa, viz., C. candelabrum, C. elegans and C.
juncea, have succulent leaves but have only hypostomatic nature unlike C. bulbosa.
The type of stomata is paracytic in most of the taxa (16 out of 26) under
study. Rubiaceous
type (paracytic) stomata are reported to be common
feature of many genera of Asclepiadaceae (present Asclepiadoideae of Apocynaceae)
(Metcalfe & Chalk 1950; Paliwal et al.
1980). The results of present study
showed similar observations about the type of stomata in Ceropegia,
but besides paracytic stomata, tetracytic,
anomocytic and mixed stomata of isotricytic
+ tetracytic, & paracytic
+ anomocytic are also found. Thus, the variations of type of stomata occur
in the genus.
The results revealed that, mean
value of stomatal index for the taxa under study, i.e., experimental mean is
14.75±0.35. The highest SI is found in C.
attenuata var. attenuata
while the lowest value for SI is observed in C. juncea
(8.18±0.37). The mean value for
stomatal density/mm2 of leaf area is 252.31±8.50 with the highest
value of 403.64±10.60 in C. maharashtrensis
and the lowest 98.18±3.40 in C. juncea. There is no correlation observed among the
taxa with respect to SI values, but for SD the taxa occurring in the dry
climates and with succulent habits have relatively lower values for the
parameter. The SI remain unchanged even
in different seasons in soybean cultivars (Rahangdale
2003) suggesting true genetic nature of this parameter. The SD is the lowest in C. juncea a succulent taxon with highly reduced leaves and
increasing to some extent in C. bulbosa var. lushii, C. bulbosa var.
bulbosa, and C. hirsuta. Low SD helps these taxa to retain more water
in plant body and survive during dry spell of the year. On the other hand, taxa growing under high
rainfall conditions, viz., C. attenuata, C. anantii, C. jainii, C. media, C. mahabalei, C. anjanerica, and
C. maharashtrensis, have high stomatal density
ranging from 325 to 403 stomata/mm2, thereby adapting to the high
annual rainfall of about 2,000–4,000 mm.
It is interesting to note that, C. concanensis
found in the Konkan region, on lateritic rock plateaux
with very negligible soil have intermediate value of stomatal density. This is in accordance with it’s relatively broader leaves and habitat because the rain
water never remain on the sloppy plateaux. Such plateaux show
relatively dry conditions as compared to the little more soil rich habitats
where C. anantii and C. attenuata
occur.
Present study revealed that on
the basis of micro-morphological characters, the taxa with overlapping
vegetative characters can be easily identified.
The type of stomata, number and nature of subsidiary cells and
anticlinal wall pattern are important characters to differentiate between very
closely allied taxa at species as well as infraspecific ranks. This has been shown in an artificial key as
well as the morphological phylogeny tree derived in the present study. These results are in corroboration of the
previous studies in different angiosperm taxa.
Metcalfe & Chalk (1988) reported that stomatal index is independent
of the changes in epidermal cell size brought by the environmental factors, and
therefore, the SI is generally considered as a taxonomic character along with
the others. Many of the works have
considered SI as a strong taxonomic character to delimit the allied taxa of
family Vitaceae (Patil
& Patil 1983, 1984), in Bauhinia L. (Kotresha & Seetharam 1995),
and Cassia L. species (Kotresha & Seetharam 2000). The
leaf micromorphological characters are very significant to compare the fossil
taxa with the allied species of extant taxa (Dilcher
1974). Doyle & Endress (2000)
observed that, the morphological evidences overcome the weak molecular
evidences while assigning the phylogeny of angiosperms. Kotresha & Seetharam (2000) studied epidermal micromorphology of Cassia
L. species and delimited the species within the genus on the basis of stomatal
index, size, and epidermal cell wall structure (especially the anticlinal wall
pattern) and the type of stomata. Thus,
stomatal characters and micro-morphology can play a very significant role in
resolving the taxonomic ambiguities.
Present study is also showing similar outcomes giving significant
information in delimitation of Ceropegia
species and varieties on the basis of micromorphological characters. For example, two varieties of C. mahabalei, viz., var. mahabalei
and var. hemalatae, are different with respect
to many characters including stem scandant, undulate
leaf margins, 2–4 indistinct subsidiaries and rounded anticlinal walls of
epidermal cells in latter, whereas var. mahabalei
have erect stout stem, straight leaf margins, two distinct subsidiaries, and
undulate epidermal anticlinal walls.
These characters are other than the characters which are provided in
original protologue. Variety hemalatae does not match in any characters with C.
oculata.
Similar is the case of varieties of C. attenuata,
viz., var. attenuata and var. mookambikae.
The later variety has paracytic stomata with
two distinct subsidiaries and SI 15.05, while the variety attenuata
has mixed stomata of isotricytic and tetracytic types with 3–4 indistinct subsidiaries stomatal
index 21.80. As per the key provided in
the present study variety mookambikae is more
allied to C. concanensis than with C. attenuata var. attenuata.
The consensus phylogenetic tree
has three clusters. The type of stomata
is defining character for these three clusters and supported by the number
& nature of subsidiary cells. Cluster
I is having anomocytic and mixed type of stomata with
indistinct variable number of (two/three/four/five) subsidiary cells. All the taxa in this cluster are erect
ones. Cluster II comprises total 15 taxa
having paracytic stomata. This cluster further has two
sub-clusters. Sub-cluster IIa comprises eight taxa having two distinct subsidiaries
in all. From the dendrogram it is
observed that the taxa, C. evansii, C. fantastica,
C. oculata, and C. lawii
are closely related having unit distance of 0.2–0.4. Along these C. candelabrum is
separated from others on the basis of having fascicled roots and undulate
anticlinal epidermal walls. Sub-cluster
IIb comprises seven taxa having indistinct and variable subsidiaries. C. mahabalei
var. hemalatae is separated from others on the
basis of scandant habit, undulate leaf margins and
high SI value. The cluster III is an
externally linked group has four taxa clustered with unit distance of
0.2–0.8. It comprised two varieties of C.
bulbosa, C. elegans,
and C. juncea. This is the group of taxa having succulent
stems and leaves. Among them C. juncea is placed as distinct taxon as it has highly
reduced leaves. Among these C. elegans has tetracytic
stomata while others have paracytic ones and
therefore, it is placed little distantly.
The dendrogram depicts clear differences among related taxa on the basis
of micromorphology, especially the type of stomata, anticlinal wall pattern, SI
and supported by gross morphology. In
the present study, the grouping of C. maccannii,
C. panchganiensis, C. sahyadrica, and C. rollae
in first cluster, as well as C. anjanerica,
C. mahabalei, C. jainii, C. media, and C. odorata in second cluster is in corroboration with
the molecular phylogeny done by Surveswaran et al.
(2009) where these taxa were placed in two separate clusters.
It is necessary to discuss here
the delimitation of some of the taxa in present study. C. mahabalei
var. mahabalei and C. mahabalei var. hemalatae
are distinct in gross morphology of stem, leaves, inflorescence and the corolla
beak, in a similar way they also differ in having indistinct 2–4 subsidiaries,
rounded anticlinal walls and higher SI in the later variety. In the dendrogram they are also placed apart
from each other on the basis of above characters. Similarly, two varieties of C. attenuata, viz., var. attenuata
and var. mookambikae, are also placed
distantly in the dendrogram in two different clusters. This is because of the type of stomata,
number & characters of subsidiaries, size of stomata and SI. The most confusing taxa in terms of
vegetative and floral morphology are C. rollae
and C. lawii, as they are
morphologically very similar to each other.
Results of the present study separated them in the dendrogram as well as
in the identification key on the basis of type of stomata, number and feature
of subsidiaries, anticlinal wall pattern, and size of stomata. Thus, their identity can be confirmed on the
basis of micro-morphology even in vegetative state. Similarly, all the above mentioned taxa are
separated on the basis of micro-morphological characters in the artificial key
also. Thus, micro-morphological
characters when combined with the gross morphological characters give better
opportunity for correct identification of the taxa. The inter-specific variations in the stomatal
characters such as stomatal index and stomatal frequency in Vitis
L., Cissus L., and Leea
D. Royen ex L. were studied by Patil
& Patil (1984) and the taxa within these three
genera were delimited on the basis of the stomatal characters and relationship
among them was also established.
Stomatal size and frequency was used to delimit between 56 species of
genera of family Vitaceae by Patil
& Patil (1983) and opined that, the stomatal data
is supporting other characters to delimit the species and also to establish
phylogenetic relationship to some extent.
The results of present study show a similar trend in the genus Ceropegia.
Therefore, on the basis of
studied micromorphological characters combined with other vegetative characters
26 taxa of Ceropegia at species and
infraspecific level are delimited and an identification key is being provided
for easy identification. These
characters provide a very easy way of identification even in absence of flowers
of these taxa.
CONCLUSION
The micro-morphology of taxa in
the genus Ceropegia L. is revealed and is
determinative for the identification of taxa even in a vegetative state in
combination with the other general morphological characters. The type of stomata, number and nature of
subsidiary cells, anticlinal epidermal wall pattern and stomatal index are key
characters for differentiation of the taxa.
Besides paracytic stomata, anomocytic,
tetracytic and mixed, isotricytic
+ tetracytic and paracytic
+ anomocytic stomata are found in the genus Ceropegia.
These characters are crucial in correct identification of the taxa at
species and infraspecific level.
Occurrence of tetracytic and mixed stomata is
a first report in the genus by this study.
It is also confirmed that, C. mahabalei
var. hemalatae and C. attenuata
var. mookambikae are distinct taxa from their
allied taxa, respectively.
Key
for identification of species on the basis of vegetative and
micro-morphological characters
1.
Plants erect ………………………… ……………………………………………………………………………………………………………………………....…………….
2
1.
Plants twiner ...…………………………………… ……………………………………………………………………………………………………………………………….....
3
2.
Leaves broadly ovate ..………………………………………………… ………………………………………………………………………………………………………
4
2.
Leaves lanceolate/linear ...………………………………………………… …………………………………………………………………………………………………
5
4.
Stomata paracytic or anomocytic
……………………………………………………………………… …………………………………………………………………… 6
4.
Stomata mixed ...……………………………………………………………………………… ……………………………………………………………………………………..
7
6.
Stomata paracytic, subsidiaries 2 distinct, epidermal
cell wall undulate …………………………………………………………………. C. lawii
6.
Stomata anomocytic, subsidiaries 4–5 indistinct,
epidermal cell wall straight / rounded ……………… ………………………………….. 8
7.
Stomata para & anomocytic, subsidiaries 2–4,
epidermal cell wall straight ……………………………… …………………………….... C. rollae
7.
Stomata isotricytic & tetracytic,
subsidiaries 3–4 indistinct ……………………………..……… ……………………………… ………………….… 9
8.
Epidermal cell wall straight, SI 17.36 ………………………………………………….………… … ………………………………………………...
C. maccannii
8.
Epidermal cell wall rounded, SI 11.24 ……………………………………………………… …………………………………………………
C. panchganiensis
9.
Epidermal cell wall rounded, SI 19.94, SD 285.45 …………………………… …………………………………………………………………
C. sahyadrica
9.
Epidermal cell wall straight, SI 15.48, SD 403.64 ……………………………… ……………………………………………………………
C. maharashtrensis
5.
Plants robust up to 1m tall, stomata paracytic
...………………………………………………………… …………………… ……………………………… 10
5.
Plants small / delicate, shorter than 0.5m tall, stomata paracytic
/ other types ..........………………………………........……………… …. 11
10.
Stem stout, leaf margin straight, subsidiary 2, distinct, epidermal cell wall
undulate ..................... C. mahabalei var.
mahabalei
10.
Stem scandant, leaf margin undulate, subsidiary 2–4,
indistinct, epidermal cell wall rounded …… C. mahabalei
var. hemalatae
11.
Stomata paracytic, subsidiaries distinct
.……………………….…………………………… …………………………………………………………… 12
11.
Stomata isotricytic & tetractytic,
subsidiaries 3–4 indistinct ……………………….……… ……………………………………………………… ... 13
12.
Subsidiaries 2 or 4, epidermal cell wall undulate
....……………………………………………………………………………………… ………………. 14
12.
Subsidiaries 2–3, epidermal cell wall rounded / undulate
.……………………………………………………………………………………… ……………… 15
14.
Subsidiaries 2, SI 15.05, SD 329.09 ……………………………………… ……………………………………………… C. attenuata
var. mookambikae
14.
Subsidiaries 4, SI 12.11, SD 258.18 ………………………………………………… …… ………………………………………………………… C. concanensis
15.
Leaves hirsute, epidermal cell wall undulate, SI 14.69, SD 338.18 ………………… … ………………………………………………….
C. anjanerica
15.
Leaves sparsely hairy, epidermal cell wall rounded, SI 12.26, SD 370.91 ……………… ………………………………………………………
C. jainii
13.
Epidermal cell wall undulate, SI 21.80, SD 300.0 ……………………………………… … ……………………………..
C. attenuata var. attenuata
13.
Epidermal cell wall rounded, SI 16.71, SD 390.91 ……………………………………… ……………………………………………………………
C. anantii
3.
Plants succulent, leaves ovate/lanceolate/linear/minute, epidermal cell wall
straight ........……………………………………… ......……… 16
3.
Plants non succulent, leaves ovate/ lanceolate, epidermal cell wall otherwise
………………….........……………… ……………………....... 17
16.
Leaves minute, hypo-stomatic, stomata paracytic, subsidiaries 2–4 distinct ……………… ……………………………………………….
C. juncea
16.
Leaves conspicuous, amphi-stomatic, stomata anomocytic, subsidiaries 3–4 indistinct …....…………………… ....….....................
18
18.
Leaves ovate-lanceolate, SI 15.60, SD 127.27 …………………………………………… …………………………………………
C. bulbosa var. bulbosa
18.
Leaves linear, SI 13.10, SD 105.45 ……………………………………………… …………………………………………………………. C.
bulbosa var. lushii
17.
Leaves lanceolate, stomata paracytic
………………………………………………………...………………………………… …………………………………. 19
17.
Leaves broadly ovate, stomata otherwise ……………………………………………………………………………………………… ………..……………
20
19.
Leaves minutely hairy, veins winged below, epidermal cell wall straight, SI
14.85, SD 314.55 ……………… ………………….. C. media
19.
Leaves glabrous, veins not winged, epidermal cell wall rounded, SI 13.51, SD
250.91 ………… …………………………………… C. odorata
20.
Plants almost glabrous, stomata para or tetracytic
………………………………………………………………………………… ………………………. 21
20.
Plants hairy, stomata paracytic
…………………………………………………………………………………………………………………………… …………….. 22
21.
Stomata paracytic, subsidiaries 2 distinct, epidermal
cell wall undulate ………………… …………………………………… C. candelabrum
21.
Stomata tetracytic, subsidiaries 4 indistinct,
epidermal cell wall rounded ………………………..…………………………… C. elegans
22.
Epidermal cell wall rounded / straight, subsidiaries distinct
………………………………………………………………… ……………... 23
22.
Epidermal cell wall undulate, subsidiaries distinct / indistinct
…….………………………………………...……………………………… ……. 24
23.
Subsidiaries 2, epidermal cell wall straight ………………………………… ……………………………………………………………….
C. fantastica
23.
Subsidiaries 2–4, epidermal cell wall rounded
…………………………………………………………………………………………… …………………… 25
25.
Plants densely hirsute, SI 11.88, SD 152.73 ……………………………………………………… ………………………………………………………
C. hirsuta
25.
Plants sparsely hirsute, especially on leaves, SI 14.76, SD 230.91 ……………… ……………………………………………………………
C. oculata
24.
Subsidiaries 2 distinct …………………………………………………………………… ………………………………………………………........……….
C. evansii
24.
Subsidiaries 2–3 indistinct …………………………………………………………....... ..............................................................…………
C. vincifolia
Table 1. Plant materials
(taxa) used in the present study.
|
|
Taxa name |
Status* |
Distribution* |
Locality of collection |
Specimen number |
|
1 |
Ceropegia anantii Yadav et al. |
EN |
Endemic; MS |
Konkan, Ratnagiri, MS |
1122 |
|
2 |
C. anjanerica Malpure et al. |
CR |
Endemic; MS |
Anjaneri, Nashik, MS |
23926 |
|
3 |
C. attenuata Hook. var. attenuata
|
EN |
Endemic; MS, Goa, KA, Raj. |
Junnar, Pune, MS |
907, 1116, 21441 (AHMA) |
|
4 |
C. attenuata Hook. var. mookambikae
Diwakar & Singh |
EN |
Endemic; MS, KA |
Konkan, Ratnagiri, MS |
1112 |
|
5 |
C. bulbosa Roxb. var. bulbosa
|
Common |
Asia & Africa |
Junnar, Pune, MS |
18544, 18545 (AHMA) |
|
6 |
C. bulbosa Roxb. var. lushii
(Grah.) Hook. f. |
Common |
India, Pakistan |
Junnar, Pune, MS |
21685, 24312 (AHMA) |
|
7 |
C. candelabrum L. |
VU |
India, Sri Lanka |
Waynad, KL |
23924 |
|
8 |
C. concanensis Kamble et al.
|
EN |
Endemic; MS |
Konkan, Ratnagiri, MS |
1120, 1121 |
|
9 |
C. elegans Wall. |
Occasional |
India, Sri Lanka |
Coimbatore, TN |
23923 |
|
10 |
C. evansii McCann |
EN |
Endemic; MS |
Ambegaon, Pune, MS |
139, 1117, 1118 |
|
11 |
C. fantastica Sedgwick |
CR |
Endemic; MS, KA, Goa |
Amboli, Sindhudurg, MS |
23920 |
|
12 |
C. hirsuta Wight & Arn. |
Common |
India, Thailand |
Junnar, Pune, MS |
0466, 23908, 23909, 23910, 23911, |
|
13 |
C. jainii Ansari & Kulkarni |
EN |
Endemic; MS |
Kas, Satara, MS |
23927 |
|
14 |
C. juncea Roxb. |
Occasional |
India, Sri Lanka |
Coimbatore, TN |
23925 |
|
15 |
C. lawii Hook. f. |
EN |
Endemic; MS |
Junnar, Pune, MS |
0316 |
|
16 |
C. maccannii Ansari |
EN |
Endemic; MS |
Simhagarh, Pune, MS |
0832 |
|
17 |
C. mahabalei Hemadri & Ansari var. hemalatae
Rahangdale & Rahangdale |
CR |
Endemic; MS |
Junnar, Pune, MS |
0136, 0137 (Types), 1114 |
|
18 |
C. mahabalei Hemadri & Ansari var. mahabalei |
CR |
Endemic; MS |
Junnar, Pune, MS |
0138, 0908, 23915, 23916, 23918 |
|
19 |
C. maharashtrensis Punekar et al. |
CR |
Endemic; MS |
Junnar, Pune, MS |
23222 |
|
20 |
C. media (Huber) Ansari |
EN |
Endemic; MS |
Junnar, Pune, MS |
0774, 1119, 22735X, 23917 |
|
21 |
C. oculata Hook. |
VU |
Endemic; MS, KL, TN |
Junnar, Pune, MS |
23902, 23903, 23904, 23914 |
|
22 |
C. odorata Nimmo ex. Hook f. |
CR |
Endemic; MS, Raj, Guj. |
Junnar, Pune, MS |
22922 |
|
23 |
C. panchganiensis Blatt. & McCann |
CR |
Endemic; MS |
Junnar, Pune, MS |
18549 (AHMA) |
|
24 |
C. rollae Hemadri |
CR |
Endemic; MS |
Junnar, Pune, MS |
1123, 23209, 23334, 23912, 23913 |
|
25 |
C. sahyadrica Ansari & Kulkarni |
CR |
Endemic; MS |
Ambegaon, Pune, MS |
0188, 23907 |
|
26 |
C. vincifolia Hook. |
VU |
Endemic; MS |
Junnar, Pune, MS |
0313 |
CR—Critically Endangered | EN—Endangered |
VU—Vulnerable | Guj—Gujarat | KA—Karnataka |
KL—Kerala | MS—Maharashtra | Raj—Rajastha | TN—Tamil
Nadu | *—as per Singh et al. (2015).
Specimens are deposited at AHMA—Agharkar
Research Institute, Pune and Herbarium of Hon. Balasaheb
Jadhav College, Ale.
Table 2. Epidermal and
stomatal characters observed in the taxa under study.
|
|
Name of taxon |
Stomatal distribution |
Type of Stomata # |
Subsidiary cells |
Anticlinal cell wall pattern (Epidermal Cell
outline) $ |
|
|
Number |
Character |
|||||
|
1 |
Ceropegia anantii |
Hypostomatic |
Isotricytic & Tetracytic |
3–4 |
Indistinct |
Rounded |
|
2 |
C. anjanerica |
Hypostomatic |
Paracytic |
2–3 |
Distinct |
Undulate |
|
3 |
C. attenuata var. attenuata |
Hypostomatic |
Isotricytic & Tetracytic |
3–4 |
Indistinct |
Undulate |
|
4 |
C. attenuata var. mookambikae |
Hypostomatic |
Paracytic |
2 |
Distinct |
Undulate |
|
5 |
C. bulbosa var. bulbosa |
Amphistomatic |
Anomocytic |
3–4 |
Indistinct |
Straight |
|
6 |
C. bulbosa var. lushii |
Amphistomatic |
Anomocytic |
3–4 |
Indistinct |
Straight |
|
7 |
C. candelabrum |
Hypostomatic |
Paracytic |
2 |
Distinct |
Undulate |
|
8 |
C. concanensis |
Hypostomatic |
Paracytic |
4 |
Indistinct |
Undulate |
|
9 |
C. elegans |
Hypostomatic |
Tetracytic |
4 |
Indistinct |
Rounded |
|
10 |
C. evansii |
Hypostomatic |
Paracytic |
2 |
Distinct |
Undulate |
|
11 |
C. fantastica |
Hypostomatic |
Paracytic |
2 |
Distinct |
Straight |
|
12 |
C. hirsuta |
Hypostomatic |
Paracytic |
3–4 |
Distinct |
Rounded |
|
13 |
C. jainii |
Hypostomatic |
Paracytic |
2–3 |
Distinct |
Rounded |
|
14 |
C. juncea |
Hypostomatic |
Paracytic |
2–4 |
Distinct |
Straight |
|
15 |
C. lawii |
Hypostomatic |
Paracytic |
2 |
Distinct |
Undulate |
|
16 |
C. maccannii |
Hypostomatic |
Anomocytic |
4–5 |
Indistinct |
Straight |
|
17 |
C. mahabalei var. hemalatae |
Hypostomatic |
Paracytic |
2–4 |
Indistinct |
Rounded |
|
18 |
C. mahabalei var. mahabalei |
Hypostomatic |
Paracytic |
2 |
Distinct |
Undulate |
|
19 |
C. maharashtrensis |
Hypostomatic |
Isotricytic & Tetracytic |
3–4 |
Indistinct |
Straight |
|
20 |
C. media |
Hypostomatic |
Paracytic |
2 |
Distinct |
Straight |
|
21 |
C. oculata |
Hypostomatic |
Paracytic |
2 |
Distinct |
Undulate |
|
22 |
C. odorata |
Hypostomatic |
Paracytic |
2 |
Indistinct |
Rounded |
|
23 |
C. panchganiensis |
Hypostomatic |
Anomocytic |
4–5 |
Indistinct |
Rounded |
|
24 |
C. rollae |
Hypostomatic |
Paracytic & Anomocytic |
2–4 |
Distinct & Indistinct |
Straight |
|
25 |
C. sahyadrica |
Hypostomatic |
Isotricytic & Tetracytic |
3–4 |
Indistinct |
Rounded |
|
26 |
C. vincifolia |
Hypostomatic |
Paracytic |
2–3 |
Indistinct |
Undulate |
#—as per Prabhakar (2004) | $—as per Dilcher (1974) | Curved—Rounded | Wavy—Undulate.
Table 3. The stomatal
measurements recorded in the taxa.
|
|
Name of taxon |
Stomatal index |
Stomatal density/mm2 |
Stomatal length (µ) |
Stomatal width (µ) |
Stomatal pore length (µ) |
Size of stomata (µ2) |
|||||
|
Mean |
SE ± |
Mean |
SE ± |
Mean |
SE ± |
Mean |
SE ± |
Mean |
SE ± |
|||
|
1 |
C. anantii |
16.71 |
0.28 |
390.91 |
10.50 |
26.00 |
0.61 |
18.50 |
0.61 |
17.50 |
1.12 |
481.0 |
|
2 |
C. anjanerica |
14.69 |
0.16 |
338.18 |
9.85 |
31.00 |
0.61 |
21.00 |
1.00 |
22.00 |
0.94 |
651.0 |
|
3 |
C. attenuata var. attenuata |
21.80 |
0.47 |
300.00 |
12.03 |
28.50 |
1.00 |
20.50 |
0.50 |
16.50 |
0.61 |
584.3 |
|
4 |
C. attenuata var. mookambikae |
15.05 |
0.16 |
329.09 |
7.99 |
23.00 |
0.50 |
15.00 |
0.79 |
12.00 |
0.50 |
345.0 |
|
5 |
C. bulbosa var. bulbosa ˄ |
14.04 |
1.64 |
101.82 |
2.40 |
33.5 |
1.7 |
21.5 |
1.00 |
23.5 |
0.61 |
|
|
|
C. bulbosa var. bulbosa ˅ |
17.15 |
1.05 |
152.73 |
2.68 |
31.0 |
1.0 |
18.5 |
1.27 |
19.0 |
1.27 |
|
|
|
C. bulbosa var. bulbosa
* |
15.60 |
0.60 |
127.27 |
7.41 |
32.25 |
1.35 |
20.00 |
1.13 |
21.25 |
0.94 |
645.0 |
|
6 |
C. bulbosa var. lushii ˄ |
13.80 |
1.04 |
96.36 |
1.70 |
31.5 |
0.61 |
20.0 |
0.0 |
20.5 |
0.5 |
|
|
|
C. bulbosa var. lushii
˅ |
12.41 |
1.13 |
114.55 |
2.19 |
29.5 |
0.50 |
19.5 |
0.5 |
19.5 |
0.5 |
|
|
|
C. bulbosa var. lushii
* |
13.10 |
0.48 |
105.45 |
4.17 |
30.50 |
0.55 |
19.75 |
0.25 |
20.00 |
0.50 |
602.4 |
|
7 |
C. candelabrum |
14.39 |
0.28 |
145.45 |
3.71 |
31.50 |
0.61 |
20.50 |
0.50 |
20.00 |
0.00 |
645.8 |
|
8 |
C. concanensis |
12.11 |
0.30 |
258.18 |
9.36 |
26.00 |
0.61 |
15.50 |
0.50 |
13.00 |
0.50 |
403.0 |
|
9 |
C. elegans |
12.39 |
0.23 |
198.18 |
5.80 |
30.00 |
0.79 |
21.00 |
0.61 |
20.50 |
0.50 |
630.0 |
|
10 |
C. evansii |
15.04 |
0.52 |
209.09 |
12.03 |
33.00 |
0.50 |
19.50 |
0.50 |
19.50 |
0.50 |
643.5 |
|
11 |
C. fantastica |
16.94 |
0.46 |
269.09 |
7.84 |
28.50 |
1.27 |
23.00 |
1.22 |
16.00 |
0.61 |
655.5 |
|
12 |
C. hirsuta |
11.88 |
0.40 |
152.73 |
5.17 |
30.00 |
0.00 |
22.00 |
1.46 |
19.50 |
0.50 |
660.0 |
|
13 |
C. jainii |
12.26 |
0.30 |
370.91 |
8.58 |
31.50 |
1.00 |
20.00 |
0.00 |
20.50 |
0.50 |
630.0 |
|
14 |
C. juncea |
8.18 |
0.37 |
98.18 |
3.40 |
39.00 |
1.50 |
26.50 |
1.70 |
27.00 |
0.94 |
1033.5 |
|
15 |
C. lawii |
14.49 |
0.46 |
263.64 |
11.13 |
29.00 |
1.00 |
15.50 |
0.94 |
18.50 |
0.61 |
449.5 |
|
16 |
C. maccannii |
17.36 |
0.41 |
254.55 |
8.30 |
28.00 |
0.50 |
23.00 |
0.50 |
18.50 |
0.61 |
644.0 |
|
17 |
C. mahabalei var. hemalatae |
20.32 |
0.33 |
341.82 |
10.54 |
30.50 |
0.50 |
20.00 |
0.00 |
24.00 |
1.00 |
610.0 |
|
18 |
C. mahabalei var. mahabalei |
16.50 |
0.12 |
325.45 |
10.05 |
29.50 |
0.50 |
19.50 |
0.50 |
24.50 |
0.94 |
575.3 |
|
19 |
C. maharashtrensis |
15.48 |
0.18 |
403.64 |
10.60 |
28.00 |
1.46 |
21.00 |
0.61 |
15.50 |
0.50 |
588.0 |
|
20 |
C. media |
14.85 |
0.46 |
314.55 |
17.39 |
28.00 |
1.66 |
16.25 |
1.12 |
16.25 |
1.12 |
455.0 |
|
21 |
C. oculata |
14.76 |
0.52 |
230.91 |
8.27 |
32.00 |
0.94 |
16.75 |
0.50 |
20.00 |
1.37 |
536.0 |
|
22 |
C. odorata |
13.51 |
0.43 |
250.91 |
8.35 |
28.00 |
0.94 |
21.50 |
0.61 |
21.00 |
0.61 |
602.0 |
|
23 |
C. panchganiensis |
11.24 |
0.28 |
205.45 |
5.33 |
29.50 |
0.50 |
21.00 |
0.61 |
19.50 |
0.50 |
619.5 |
|
24 |
C. rollae |
12.99 |
0.22 |
201.82 |
4.61 |
30.00 |
0.00 |
18.50 |
0.61 |
19.00 |
0.61 |
555.0 |
|
25 |
C. sahyadrica |
19.94 |
0.60 |
285.45 |
15.19 |
30.00 |
0.00 |
16.00 |
0.61 |
20.00 |
0.00 |
480.0 |
|
26 |
C. vincifolia |
11.81 |
0.20 |
189.09 |
3.40 |
32.50 |
0.00 |
20.00 |
0.00 |
22.50 |
0.00 |
650.0 |
|
|
Experimental mean @ 25df |
14.75 |
0.35 |
252.31 |
8.50 |
29.84 |
0.73 |
19.68 |
0.67 |
19.40 |
0.64 |
587.3 |
*—the data of amphistomatic
taxa are taken from mean of both the surfaces and then the experimental mean is
calculated | ˄—upper surface of leaf | ˅—lower surface of leaf.
Table 4. List of
characters and character state used for phylogenetic analysis.
|
|
Character with character state |
|
1 |
Tuber: Absent
(0), Present (1) |
|
2 |
Stem erect: Absent (0), Present (1) |
|
3 |
Stem twining: Absent (0), Present (1) |
|
4 |
Stem scandant: Absent (0),
Present (1) |
|
5 |
Stem: Non succulent (0), Succulent(1) |
|
6 |
Leaves: Large (0), Minute (1) |
|
7 |
Plant habit:
Small (0), Robust (1) |
|
8 |
Leaves ovate: Absent (0), Present (1) |
|
9 |
Leaves lanceolate: Absent (0), Present (1) |
|
10 |
Leaves linear: Absent (0), Present (1) |
|
11 |
Leaves: Non succulent (0), Succulent (1) |
|
12 |
Leaves: Non hairy (0), Hairy (1) |
|
13 |
Leaf margin: Straight (0), Undulate (1) |
|
14 |
Leaf veins winged on lower surface: Not winged (0),
Winged (1) |
|
15 |
Leaf: Amphi-stomatic (0),
Hypo-stomatic (1) |
|
16 |
Epidermal cell anticlinal wall, Straight: Absent
(0), Present (1) |
|
17 |
Epidermal cell anticlinal wall, Rounded: Absent (0),
Present (1) |
|
18 |
Epidermal cell anticlinal wall, Undulate: Absent
(0), Present (1) |
|
19 |
Stoma: Mixed types (0), Single type (1) |
|
20 |
Stoma paracytic: Absent
(0), Present (1) |
|
21 |
Stoma anomocytic: Absent
(0), Present (1) |
|
22 |
Stoma tetracytic: Absent
(0), Present (1) |
|
23 |
Stoma isotricytic & tetracytic: Absent (0), Present (1) |
|
24 |
Stoma paracytic & anomocytic: Absent (0), Present (1) |
|
25 |
Subsidiary cells: Indistinct (0), Distinct (1) |
|
26 |
Subsidiary cell number two: Absent (0), Present (1) |
|
27 |
Subsidiary cell number four: Absent (0), Present (1) |
|
28 |
Subsidiary cell number 2–3: Absent (0), Present (1) |
|
29 |
Subsidiary cell number 3–4: Absent (0), Present (1) |
|
30 |
Subsidiary cell number 4–5: Absent (0), Present (1) |
|
31 |
Size of stoma 300–450 μ: Out of range (0),
Within range (1) |
|
32 |
Size of stoma 450–600 μ: Out of range (0),
Within range (1) |
|
33 |
Size of stoma 600–750 μ: Out of range (0),
Within range (1) |
|
34 |
Size of stoma 750–1100 μ: Out of range (0),
Within range (1) |
|
35 |
Stomatal index (SI) range value 8–11 : Out of range
(0), Within range (1) |
|
36 |
Stomatal index (SI) range value 11–14: Out of range
(0), Within range (1) |
|
37 |
Stomatal index (SI) range value 14–17: Out of range
(0), Within range (1) |
|
38 |
Stomatal index (SI) range value 17–22: Out of range
(0), Within range (1) |
|
39 |
Stomatal density (SD) range value <100: Out of range (0), Within range (1) |
|
40 |
Stomatal density (SD) range value 100-200: Out of
range (0), Within range (1) |
|
41 |
Stomatal density (SD) range value 200-300: Out of
range (0), Within range (1) |
|
42 |
Stomatal density (SD) range value 300-400: Out of
range (0), Within range (1) |
Table 5. Character state
of taxa under study (Character no. as per Table 4).
|
Sno |
Character No. |
1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
12 |
13 |
14 |
15 |
16 |
17 |
18 |
19 |
20 |
21 |
22 |
23 |
24 |
25 |
26 |
27 |
28 |
29 |
30 |
31 |
32 |
33 |
34 |
35 |
36 |
37 |
38 |
39 |
40 |
41 |
42 |
|
Taxa |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1 |
C. anantii |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
|
2 |
C. anjanerica |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
|
3 |
C. attenuata var. attenuata |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
|
4 |
C. attenuata var. mookambikae |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
|
5 |
C. bulbosa var. bulbosa |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
|
6 |
C. bulbosa var. lushii |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
|
7 |
C. candelabrum |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
|
8 |
C. concanensis |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
|
9 |
C. elegans |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
|
10 |
C. evansii |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
|
11 |
C. fantastica |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
|
12 |
C. hirsuta |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
|
13 |
C. jainii |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
|
14 |
C. juncea |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
|
15 |
C. lawii |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
|
16 |
C. maccannii |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
|
17 |
C. mahabalei var. hemalatae |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
|
18 |
C. mahabalei var. mahabalei |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
|
19 |
C. maharashtrensis |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
|
20 |
C. media |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
|
21 |
C. oculata |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
|
22 |
C. odorata |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
|
23 |
C. panchganiensis |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
|
24 |
C. rollae |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
|
25 |
C. sahyadrica |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
|
26 |
C. vincifolia |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
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