REINWARDTIA
Vol. 20. No. 1. pp: 27‒36
DOI: 10.14203/reinwardtia.v20i1.4031
27
DIVERSITY AND PHENETIC STUDY ON SYCONIUM OF FICUS L.
(MORACEAE) FROM KERALA, INDIA REVEALING NATURAL
CLASSIFICATION ALONG WITH AN IDENTIFICATION KEY
Received January 31, 2021; accepted May 3, 2021
SREEHARI S. NAIR
Research and P. G. Department of Botany, MES Asmabi College, P. Vemballur, Thrissur, Kerala, India.
Email: harificus@gmail.com
K. H. AMITHA BACHAN
Research and P. G. Department of Botany, MES Asmabi College, P. Vemballur, Thrissur, Kerala, India.
Email: amithabmes@gmail.com
P. J. EBIN
Department of Botany, Sacred Heart College, Thevara, Cochin, Kerala, India.
Email: ebinpj@shcollege.ac.in
ABSTRACT
NAIR, S. S., BACHAN, K. H. A. & EBIN, P. J. 2021. Diversity and phenetic study on syconium of Ficus L.
(Moraceae) from Kerala, India revealing natural classification along with an identification key. Reinwardtia 20(1):
27–36. — Ficus L. commonly called ‘figs’ is one of the most complex genera among the angiosperms with its
specialised inflorescence called syconium that looks like a fruit. Syconium of 33 species of Ficus reported from Kerala
were observed here to develop a novel key, solely based on syconium morphology. Numerical taxonomic methodo
logy for syconium morphological characters were standardised, considering 22 characters with 104 character states
and analysed using similarity clustering. The floral features of the genus are very much complex and all the existing
keys for the species identification relays on both vegetative as well as floral features. Hence, the present key will be
practical in use when syconium is the only available part. The numerical analysis of the syconium features well
clustered and separated the trees with cauliflorous inflorescence, hemi epiphytic-epiphytic life forms and independent
trees similar to the natural classification of the figs as “Atthi, Itthi and Aal”, indicating that phenetic analysis using the
syconium characters alone provided a grouping similar to the natural grouping based on the habit. Preliminary
phylogenetic analysis of figs also provided a similar clustering. This gives an insight into the fact that the separation of
figs into these natural groups is reflecting phylogenetic trait. Detailed studies including more morphological traits and
molecular analysis could establish the phylogenetic relation of figs in relation to the evolutionary history of climate
and vegetation.
Key words: Diversity, Ficus, identification key, natural classification, phenetics, syconium.
ABSTRAK
NAIR, S. S., BACHAN, K. H. A. & EBIN, P. J. 2021. Penelitian keanekaragaman dan fenetik bentuk sikonium Ficus
L. (Moraceae) dari Kerala, India, mengungkap klasifikasi alami dan kunci identifikasinya. Reinwardtia 20(1): 27–36.
— Ficus L. biasa disebut 'buah ara' adalah salah satu marga paling kompleks di antara angiospermae dengan
perbungaan khusus yang disebut sikonium, yang terlihat seperti buah. Sikonium dari 33 jenis Ficus yang dilaporkan
dari Kerala diamati di sini untuk mengembangkan kunci baru, hanya berdasarkan morfologi sikonium. Metodologi
taksonomi numerik untuk karakter morfologi sikonium distandarisasi, dengan mempertimbangkan 22 karakter dengan
104 status karakter dan dianalisis menggunakan penggugusan keserupaan. Ciri-ciri bunga dari marga sangat kompleks
dan semua kunci yang ada untuk identifikasi jenis bergantung pada ciri vegetatif dan juga ciri-ciri bunga. Karenanya,
kunci sekarang akan praktis digunakan jika sikonium adalah satu-satunya bagian yang tersedia. Analisis numerik fitur
sikonium mengelompok dengan baik dan memisahkan pohon dengan perbungaan kauliflorus, bentuk kehidupan hemi
epifitik-epifitik dan pohon independen yang mirip dengan klasifikasi alami buah ara sebagai "Atthi, Itthi dan Aal",
menunjukkan bahwa analisis fenetik menggunakan karakter sikonium sendiri memberikan pengelompokan yang mirip
dengan pengelompokan alami berdasarkan perawakan. Analisis filogenetik awal dari buah ara juga memberikan
pengelompokan yang serupa. Ini memberikan wawasan tentang fakta bahwa pemisahan buah ara ke dalam kelompok
alami ini mencerminkan sifat filogenetik. Studi terperinci termasuk lebih banyak sifat morfologi dan analisis
molekuler dapat menetapkan hubungan filogenetik buah ara dalam kaitannya dengan sejarah evolusi iklim dan
vegetasi.
Kata kunci: Fenetik, Ficus, keanekaragaman, klasifikasi alami, kunci identifikasi, sikonium.
mailto:harificus@gmail.com
REINWARDTIA 28 [VOL.20
INTRODUCTION
The genus Ficus L. is the most advanced
member in the family Moraceae with a worldwide
distribution of about 750 species (Berg & Corner,
2005; Chaudhary et al., 2012). They are common-
ly called ‘fig’ plants. The plant is characterised by
the typical hypanthodium inflorescence and the
infructescence, a syconium generally referred as
‘fig’ or ‘receptacle’. Fig plants are considered as a
keystone species (Vanitharani et al., 2009; Kumar
et al., 2011) in many ecosystems, with nutrient
rich figs being eaten by reptiles, birds and
mammals the year around. In India, the genus has
about 115 species including 24 infraspecific taxa
(Chaudhary et al., 2012). Thirty one species of
figs have been so far reported from Kerala
(Sasidharan, 2004). In general, there is a natural
classification of the figs (Matthew, 1995) as
“Atthi” (trees with cauliflorous inflorescence),
“Itthi” (hemi epiphytic and epiphytic trees,
stranglers) and “Aal” (independent trees), which
were the vernacular names used by Hendrik van
Rheede in Hortus Malabariucs (Rheede, 1678).
The presence of minute microscopic flowers
within the hypanthodium make the identification
of the taxa highly problematic. The keys currently
in practice are not solemnly based on syconium
characters but rather based on vegetative and flo-
ral morphology (Gamble, 1925; Corner, 1965).
Development of a key based on syconium features
with very detailed diagnosis can make the identifi-
cation of figs much easier, since the syconium
being a very characteristic feature and could be
easily collected.
In addition to molecular data, morphological
characters can also be used for establishing
phylogenetic relationship (Scotland et al., 2003;
Wiens, 2004). Consideration of more morpho-
logical characters with detailed character states
can provide a clear segregation of a genetically
related group (Henderson & Ferreira, 2002;
Sonibare et al., 2004; Bolourian & Pakravan,
2011; Rahman et al., 2013) and this could be clos-
er towards phylogenetic segregation when com-
pared to the traditional morpho taxonomic meth-
od.
The present study focuses to provide the diver-
sity of syconium and a key based on the syconium
morphology alone for the identification of Ficus
species. Also, to understand whether the clustering
of figs based on syconium morphology have any
similarity with the habit based natural classifica-
tion of figs into three groups, ‘Atthi’, ‘Itthi’ and
‘Aal’. Thus, this paper uses the numerical taxo-
nomic techniques for phenetic segregation of figs
based on syconium features for developing a
strong foundation of syconium based key as well
as for testing the similarity with natural classifica-
tion.
If the natural classification is reflected in the
phenetic syconium character segregation, this
could be taken as first level confirmation towards
the hypothesis that the natural grouping in figs are
reflecting the phylogenetic affinity.
MATERIALS AND METHODS
Extensive field studies were conducted for the
collection of specimens. Syconium of the figs were
collected and were preserved in FAA solution
(Bowles, 2004). Morphological characters of syco-
nium were observed including the vestiture and the
images were taken using Coolpix Digital Micro-
scope. The species were identified using regional
floras (Gamble, 1925; Sasidharan, 2004) and the
identified species were confirmed by herbarium
consultation in the herbariums of CALI, KFRI,
TBGT and MH. Herbarium specimens and floras
were consulted for the analysis of syconium of the
species whose fresh materials were unavailable.
The syconium characters were enumerated in a
data sheet prepared specifically for syconium
features following the terminologies used by
Simpson (1953). Multiple samples were analysed
for the conformity of the characters. Numerical
taxonomic methodology for syconium characters
were developed and standardised (Sneath & Sokal,
1973). Thirty three taxa were diagnosed and
entered into the data base with 22 characters cover-
ing 104 character states (Table 1). The develop-
ment of the different character state is based on the
range of each character and suitable for entry into a
numerical matrix representing 0 and 1 for absence
and presence respectively (Fasila et al., 2020). The
data was analysed using PAST Software for simi-
larity-based clustering, giving equal weight age for
all the character states studied.
A novel key was prepared from the observations
of syconium characters and also based on the clus-
tering provided by dendrogram. The habit specifi
city of the species was confirmed according to
Berg (2003). The similarity clustering dendrogram
obtained by analysing 22 morphological characters
of syconium was used to test the relatedness of
habit based natural grouping in figs with the clus-
tering of figs based on syconium morphology and
thereby to understand whether they have any
phylogenetic relation.
RESULTS
Syconium of 33 species of figs studied shows
significant character dissimilarity, that are enough
to differentiate them into separate species (Fig. 2).
Out of the 33 species studied, 27 species had axil-
lary inflorescence and 6 species had cauliflorous
inflorescence. Syconium of 12 species were sessile
NAIR et al.: Diversity and phenetic study on syconium of Ficus L.
2020] 29
Fig. 1. Study area, Kerala State, India.
and 21 were pedunculate. Among the 12 species
with sessile syconium, 10 species were having out-
er bracts, while two species were devoid of outer
bracts. Within the 21 pedunculate species, 15 spe-
cies had outer bracts and 6 species lack outer
bracts (Fig. 3). Vestiture on the surface of the sy-
conium were characteristic and were consistent in
all the species studied. Glabrous or hairy vestiture
of the syconium were also used to separate the spe-
cies.
Numerical taxonomic analysis of the syconium
characters also showed species level separation
with significant similarity and dissimilarity co-
efficient. The maximum dissimilarity was observed
for F. auriculata Lour (44%). Ficus rigida var.
bracteata (Corner) Bennet & F. superba Miq and
F. microcarpa L.f & F. binnendijkii (Miq.) Miq.
were the most similar species with a 92% similari-
ty.
The dendrogram developed through similarity
clustering using PAST software is given (Fig. 4).
The species were clustered into three major groups
based on the similarity of syconium morphology.
Habit preference of the species plotted on the den-
drogram and it showed a similar grouping pattern
with the syconium morphology-based clustering.
DISCUSSION
The morphological features of the syconium
were well segregated providing a syconium based
identification key for 33 species. This could be
very useful in practise, since the syconium is the
most available part of a fig during any kind of
field study. This can help field botanists as well as
ecologists and people from other disciplines. The
numerical analysis provided more clarity for
preparing such a practical key based on syconium
features.
Natural grouping in figs based on habit divides
them into three groups as ‘Atthi, Itthi and Aal’ in
Malayalam, which include the trees with cauliflo-
rous inflorescence, hemi epiphytic-epiphytic life
forms and independent trees respectively. Similar-
ity clustering based on 104 character states of the
syconium morphology alone separated the species
significantly and also clustered them into three
groups similar to the natural grouping based on
habit, strengthening relatedness between cluster-
ing based on syconium morphology and natural
grouping. The only exceptions in the clustering
were F. superba and F. hispida, two cauliflorous
species ware grouped along with the independent
REINWARDTIA 30 [VOL.20
Table 1. Character and character states of Ficus species studied.
Sl. No. Character Character States
1 Inflorescence position Axillary, cauliflorous
2 Inflorescence clustering Solitary, paired, clustered
3 Basal bracts Present, absent
4 Syconium length maximum
1–10 mm, 11–20 mm, 21–30 mm, 31–40 mm, 41–50
mm, 51–60 mm, 61–70 mm, 71–80 mm, 81–90 mm, 91–
100 mm
5 Syconium length minimum
1–10 mm, 11–20 mm, 21–30 mm, 31–40 mm, 41–50
mm, 51–60 mm
6 Syconium width maximum
1–10 mm, 11–20 mm, 21–30 mm, 31–40 mm, 41–50
mm, 51–60 mm, 61–70 mm
7 Syconium width minimum
1–10 mm, 11–20 mm, 21–30 mm, 31–40 mm, 41–50
mm
8 Syconium shape Obovoid, globose, turbinate, oblong, ovoid
9 Syconium pubescence Glabrous, hairy, hairy near ostiole
10 Syconium texture
Glabrous, pubescent, pilose, scars, scabrid, strigose,
puberulent, tomentose
11 Peduncle Present, absent
12 Peduncle length maximum 1–10 mm, 11–20 mm, 21–30 mm, 31–40 mm
13 Peduncle length minimum 1–10 mm, 11–20 mm, 21–30 mm, 31–40 mm
14 Peduncle width maximum 1–5 mm, 6–10 mm, 11–15 mm
15 Peduncle width minimum 0–1.0 mm, 1.1–2.0 mm, 2.1–3.0 mm, 3.1–4.0 mm
16 Peduncle shape Terete, angular
17 Peduncle pubescence Glabrous, hairy
18 Peduncle texture
Glabrous, pubescent, pilose, scars, scabrid, strigose,
puberulent, villous
19 Syconium colour at young stage Green, pale green
20 Syconium colour on maturity
Green, dark green, pale brown, yellow, pale yellow,
orange, pale red, purple, white, grey
21 Syconium colour on ripening
Green, dark green, pale brown, yellow, pale yellow,
orange, pale red, purple, brown
22 Lenticels Present, absent
NAIR et al.: Diversity and phenetic study on syconium of Ficus L.
2020] 31
Fig. 2. Syconium of figs. A. F. amplissima. B. F. anamalayana. C. F. arnottiana. D. F. auriculata. E. F.
beddomei. F. F. benjamina. G. F. benghalensis. H. F. callosa. I. F. drupacea. J. F. asperata. K. F. hetero-
phylla. L. F. hispida. M. F. microcarpa. N. F. nervosa. O. F. pumila. P. F. racemosa. Q. F. religiosa. R. F.
talbotii. S. F. tinctoria ssp. gibbosa. T. F. tinctoria ssp. parasitica. U. F. tsjahela. V. F. virens. W. F.
amplocarpa. X. F. binnendijkii. Y. F. caulocarpa. Z. F. costata. A1. F. dalhousiae. A2. F. elastica. A3. F.
guttata. A4. F. krishnae. A5. F. mollis. A6. F. rigida var. bracteata. A7. F. superba.
REINWARDTIA 32 [VOL.20
Fig. 3. Diversity of the syconium.
Syconium Based Key for the Identification of Ficus species
1a. Syconium sessile, peduncle absent ……………...……………………………......……………….….... 2
1b. Syconium not sessile, peduncle present ….…...…………………...……………………....................... 3
2a. Surface of the syconium hairy ….……………….…………………………………….……….……..... 4
2b. Surface of the syconium glabrous ……………………………………………...............................…… 5
4a. Receptacles paired, surface with puberulent hairs, basal bracts absent ..........................… F. krishnae
4b. Receptacles clustered, surface with pubescent or tomentose hairs, basal bracts present …................. 6
6a. Figs with pubescent hairs, 1.5–2.5 cm in size, red coloured on ripening ………….. F. benghalensis
6b. Figs with tomentose hairs, 0.8–1.0 cm in size, brown coloured on ripening .……….....….. F. mollis
5a. Surface of the syconium with scars, syconium solitary in arrangement .……………...….... F. talbotii
5b. Surface of the syconium glabrous without scars, syconium not solitary in arrangement,
clustered ……………………………………………………………………………………...…...…. 7
7a. Basal bracts present ……………………...………………………………………….………...…… 8
7b. Basal bracts absent ……………….………………….…….…………………….…… F. benjamina
8a. Figs arranged solitary or in pairs …….…………………………………………………………... 9
8b. Figs arranged in clusters …….…………………………...…………………..……………..….. 10
9a. Syconium obovoid in shape, arranged solitary …………..……………..……….. F. amplissima
9b. Syconium not obovoid in shape, arranged in pairs ………………….………...…….. ……… 11
11a. Shape of the receptacle oblong ……………………………..…..……………....…………. 12
11b. Shape of the receptacle not oblong, globose or ovoid ………………………..………...… 13
12a. Figs up to 4 cm, red coloured on ripening ………………………………....…. F. drupacea
12b. Figs up to 1.5 cm, yellow coloured on ripening …………................………...… F. elastica
13a. Receptacle globose in shape, 0.8–1.0 cm in size, orange coloured on
ripening ………………………….…………………………………...…….. F. microcarpa
13b. Receptacle ovoid in shape, 0.3–0.5 cm in size, red coloured on ripening …………..
…...………………….………………………………………….………….... F. binnendijkii
NAIR et al.: Diversity and phenetic study on syconium of Ficus L.
2020] 33
10a. Syconium 0.4–1.0 cm in size, red coloured on ripening ……………..………..….. F. religiosa
10b. Syconium 0.5– 0.6 cm in size, pale brown coloured on ripening …………..…...…. F. tsjakela
3a. Figs cauliflorous ……………………………………..………………………….…………….……… 14
3b. Figs axillary ……………………………………………………………..………………..………...… 15
14a. Receptacles in pairs, arising from long hanging racemes, rarely axillary ……………..…. F. hispida
14b. Receptacles in clusters, arising from the main trunk only ……..…………………...…………….. 16
16a. Turbinate shaped syconium, surface of the syconium hairy…….…………………… F. auriculata
16b. Globose shaped syconium, surface of the syconium glabrous …………..……………...…….… 17
17a. Figs less than 1.5 cm across, purple coloured in ripening ………………………...... F. superba
17b. Figs above 2.0 cm across, red coloured in ripening ……….……….……………………...….. 18
18a. Receptacles in clusters of many, on long racemes (up to 15 cm) ……..………..... F. racemosa
18b. Receptacles in clusters of few, not on long racemes …………………………………...…… 19
19a. Figs large, 3.0–6.5 cm in size, peduncle up to 1.0 cm in length, mature figs without
ooze of resin …………………………………………................................…. F. amplocarpa
19b. Figs small, up to 2.5–3.5 cm in size, peduncle above 1.0 cm in length, mature figs
with ooze of resin ………………………….…………………………………...…. F. guttata
15a. Syconium arranged as clusters on the axis ………………….………………..………………...…. 20
15b. Syconium arranged in pairs or solitary on the axis ……………………….………………………. 21
20a. Peduncle and figs hairy, peduncle up to 0.2–0.4 cm long …………...…………...… F. caulocarpa
20b. Peduncle and figs glabrous, peduncle up to 0.1–0.2 cm long ………………….…… F. arnottiana
21a. Figs paired ……………………………………………………………………………...……….. 22
21b. Figs solitary …………………………………………………………………………………...… 23
22a. Basal bracts present ………………………………………………….………………...……… 24
22b. Basal bracts absent ………………………………………………...…….……………………. 25
24a. Globose shaped receptacles …………………………………………….....……………….... 26
24b. Obovoid shaped receptacles …………………………………………………..….………..... 27
26a. Peduncle long, 0.5–1.0 cm, figs less than 1.8 cm across …………………...…………...… 28
26b. Peduncle short, 0.1–0.3 cm, figs 2.0 cm and above ………..……………...….....…. F. virens
28a. Receptacles 1.2–1.6 cm in size, peduncle 0.5–0.8 cm long, yellow–red coloured on
ripening …………………………………………………………………...…….. F. costata
28b. Receptacles 0.8–1.2 cm in size, peduncle 0.5–1.0 cm long, pale brown–purple
coloured on ripening …………………………………….…...….... F. rigida var. bracteata
27a. Syconium above 3.5 cm, surface glabrous with scars, peduncle angular in shape ……
……………………………………………………………………………...….... F. beddomei
27b. Syconium below 2.5 cm, surface hairy without scars, peduncle terete in shape ………..... 29
29a. Figs and peduncle hairy, figs 1.0–1.6 cm across, peduncle 0.8–1.0 cm ……. F. dalhousiae
29b. Figs hairy near ostiole and peduncle glabrous, figs 1.6–2.3 cm across, peduncle
0.5–1.2 cm ………………………...…………………………………........ F. anamalayana
25a. Length of the peduncle above 1.0 cm, receptacles red coloured on ripening … F. exasperata
25b. Length of the peduncle below 0.8 cm, receptacles yellow coloured on ripening ………… 30
30a. Surface of the syconium hairy, below 0.8 cm in size, peduncle 0.4–0.6 cm in length
…..……………………………………………………...………… F. tinctoria ssp. gibbosa
30b. Surface of the syconium glabrous, above 1.0 cm in size, peduncle 0.5–1.0 cm in
length ………..…………….……………..………………...….. F. tinctoria ssp. parasitica
23a. Receptacle obovoid in shape, hairy, with basal bracts ……………..………….……… F. pumila
23b. Receptacle globose in shape, glabrous, without basal bracts ……………...……...…….…….. 31
31a. Syconium with scars on the surface, 2.8–3.5 cm across, peduncle hairy …...… F. heterophylla
31b. Syconium without scars on the surface, glabrous, below 3.0 cm across, peduncle glabrous.. 32
32a. Receptacles 2.5 cm and above, peduncle length above 1.0 cm …………………. F. callosa
32b. Receptacles below 2.0 cm, peduncle length 0.8–1.0 cm …………………….…. F. nervosa
REINWARDTIA 34 [VOL.20
Fig. 4. Dendrogram showing the clustering of figs based on syconium morphology and habit
preference.
NAIR et al.: Diversity and phenetic study on syconium of Ficus L.
2020] 35
trees. Ficus superba might have grouped among
them due to the close similarity in the syconium
morphology with other species in the group. Ficus
hispida shows cauliflorous inflorescence as well as
axillary inflorescence. Along with this, the close
similarity in syconium morphological characters
could be the reason for the exceptional clustering
of this species with other members. Phylogenetic
studies on figs also provided a similar grouping
(Sreehari & Bachan, 2020).
Morphometric studies were used to establish the
genetic relatedness in many genera (Rahman et al.,
2013). Since the phenetic analysis of the syconium
and preliminary phylogenetic analysis shows a
clustering, similar to the natural grouping of figs,
this could be taken as first level confirmation
towards the hypothesis that the natural grouping in
figs is reflecting the phylogenetic affinity.
CONCLUSION
The syconium based numerical analysis
showed that the morphological variations in syco-
nium characters alone is significant to separate the
species. Hence, the syconium morphology-based
identification key will be practical in use, even
when the syconium is the only available part. The
syconium morphology-based clustering was much
similar to the natural grouping of figs into trees
with cauliflorous inflorescence, hemi epiphytic-
epiphytic life forms and independent trees, indi-
cating phylogenetic affinity. Detailed phenetic
analysis incorporating more morphological charac-
ters, molecular studies including a greater number
of taxa and their ecological association within a
geographical enclosure can provide more light on
the phylogeny of figs in relation to the evolution-
ary history of climate and vegetation.
ACKNOWLEDGEMENTS
We are grateful to the Department of Botany,
MES Asmabi College, P. Vemballur and Sacred
Herat College, Thevara, for providing the facilities
to carry out the work. The authors are thankful to
the curators of CALI, KFRI, TBGT and MH for
the services provided for the completion of the
study.
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