Taxonomy, distribution and diversity of Ficus palmata Forssk. subsp. virgata (Roxb.) Browicz (Moraceae) in India

1,3,4 Plant Diversity, Systematics and Herbarium Division, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow, Uttar Pradesh 226001, India 2,5 Botanical Survey of India, Southern Regional Centre, Coimbatore, Tamil Nadu 641003, India 6 M.B. (P.G.) College, Haldwani, Nainital, Uttarakhand 263139, India 1 rinkeynbri@gmail.com, 2 jvsbsi@yahoo.com, 3 aanshul_84@rediffmail.com, 4 dr_lbchaudhary@rediffmail.com (corresponding author), 5 gvs.23.murthy@gmail.com, 6 anjala.69@gmail.com

different areas or within an area appear to be different due to variable features.Sometimes the form with entire leaves is described as F. palmata whereas the form with lobed leaves is described as F. carica, while all variations can be seen on the same plant in different areas.Since these variations are continuous with numerous intermediate forms, delimiting infraspecific taxa is not possible.The dioecious nature of plants, which are similar in their gross morphology, makes it even more difficult to delineate.
Sometimes Ficus palmata subsp.virgata is also confused with its closely allied species F. carica (Image 1B), known only in cultivation in some parts of India for its fig/fruit commonly known as 'anjeer' (Hindi).F. carica is a native from the Mediterranean region to Afghanistan and differs from F. palmata (including F. palmata subsp.virgata) in degree of pubescence and thickness of young stems, colour and pubescence of tepals and size of figs.
In F. carica, the twigs are glabrous or glabrescent and stouter, figs more than 2cm in dia.and tepals often glabrous or rarely glabrescent and white to pink while in F. palmata the twigs are tomentose and slender, figs up to 2cm in dia.and tepals puberuolous and white (Corner 1965).In addition, the figs of F. palmata are hard and unpalatable, whereas the figs of F. carica are sweet, tasty and edible when ripe (Roxburgh 1832).Although these two species are treated separately till date, doubts have been raised by many other workers about the separate identity of the two species.King (1887-88), for example, has also pointed out that F. palmata may be a form of F. carica.In 'Figweb' (2004'Figweb' ( -2013)), it has also been stated that both the species are probably conspecific.Molecular and chemical studies of this complex group may help resolve the confusion.
In view of the above, the main purpose of the present work is to (i) reevaluate the status of F. palmata subsp.virgata in India, (ii) record the range of variations, and (iii) provide its updated distribution status in India.
The raw figs of F. palmata subsp.virgata are used as vegetable and ripened figs as fruit generally by tribals and local people in hilly and forest areas (Parmar & Kaushal 1982).Figs are demulcent, laxative and used in diseases of the lungs and bladder (Kirtikar & Basu 2001).The twigs are used as fodder.The latex is said to be employed in curdling of milk (Maheswari 1963).

INTRODUCTION
The genus Ficus L. (commonly known as 'figs') with ca.735 species in the world is divided into six subgenera, 19 sections and 27 subsections (Berg & Corner 2005).Although the genus has been studied at regional and global levels (Corner 1958(Corner , 1965(Corner , 1981;;Berg 1989Berg , 2003Berg , 2004 a,b; a,b;Kochummen 1998;Dixon 2003;Wu et al. 2003;Burrows & Burrows 2003;Berg & Corner 2005), the Indian figs have not been revised after King (1887King ( -88, 1888) ) till date while a large number of specimens have been added from different parts of India during the last 100 odd years.Recently, Chaudhary et al. (2012) published a checklist of Indian Ficus which consisted of 115 taxa (89 species and 26 infraspecific taxa).The present work is also an outcome of the revisionary study of the genus in India.
Ficus virgata Roxb.was described by W. Roxburgh in his Flora Indica in 1832 from Rohilkhand of Uttar Pradesh in India.He also described another species, F. caricoides Roxb., in the same work from Lucknow (Uttar Pradesh) and differentiated it from F. virgata chiefly based on the size of the plants.Later on, it was reduced as a synonym under quite variable species F. virgata by Brandis (1874).Subsequently, King (1888) merged these two species under F. palmata Forssk.However, there are disagreements among various workers regarding the correct identity and distribution of F. palmata and F. virgata since long.Browicz (1982) has pointed out that typical F. palmata (Image 1A) occurs only in eastern Africa and the southern Arabian Peninsula, and differs from F. virgata Roxb. in having leaves more elongated, acute or acuminate at apex and glabrescent or completely glabrous, whereas in F. virgata Roxb., the leaves are ovate, broadly ovate, elliptic or almost rounded, acute to rounded or truncate at apex and puberulous to densely tomentose on both the surfaces.He also observed that F. virgata Roxb.was distributed from Afghanistan to Nepal and treated it as a subspecies under F. palmata Forssk.

Morphological Diversity
Habit: Ficus palmata subsp.virgata (Figs.1a-m & Images 2a-f) occurs in two distinct forms: bushy shrubs and small or medium-sized trees.Sometimes the plants have a single weak stem, giving an appearance of a large herb.The bushy form of the plant is sometimes found hanging on the mountain slopes.Both shrubby and tree forms branch profusely.In the tree form, the leaves are comparatively smaller than that of the shrubs.The girth of the tree trunk ranges from 30-60 cm in diam.
Indumentum: The hairs distinctly differ from plant to plant and population to population within the species.They are quite dense on young twigs, stipules and leaves, but become sparse at maturity.Due to the presence of puberulous or scabrous types of hairs, the upper surfaces of the leaves and receptacles are rough to touch while the lower surfaces are soft, cottony and hairy.
Leaves: The leaves show tremendous variations in lamina shape, size, base, margins and lobation between plants or on the same plant.The leaves of bushy or shrubby forms are larger (up to 27x24 cm) than the leaves of the tree form.The leaves show distinct dimorphism in the taxon.Generally, the leaves are unlobed, but distinctly 3-5-lobed leaves are also not rare.In bushy and shrubby forms, the leaves are usually entire, but sometimes, in the lower portion of plants, the lobed and entire leaves are intermixed.On trees, both types of leaves are either found on the same branch or occasionally on separate branches.The leaves are ovate to broadly ovate with acute tip or sometimes almost rounded with truncate or rounded apex.The base of the leaves also vary from cuneate to obtuse or cordate.The leaf-margin, which is considered one of the important taxonomic characters, also differs from plant to plant from subdentate to dentate to distinctly serrate Fig. 2.
Stipules: The stipules are not much variable.They are caducous and leave scars on twigs.Generally, they are greenish throughout, but sometimes the tip portion becomes reddish (Image 2B).
Bracts Flower: In this taxon five types of flowers are seen: male, female, gall, pseudo-hermaphrodite and neuter.All these types of flowers do not occur within the same syconium as the taxon is dioecious.The female and neuter flowers are present in a single syconium known as a female syconium which grows on female plants.The male, gall and pseudo-hermaphrodite flowers occur within a same syconium known as male syconium which grows on male plants Fig. 1.
Male flowers in syconium are arranged in a bunch near ostiole as well as scattered inside the whole syconium along with gall flowers.In male flowers the number of tepals generally varies from 4-6 which are almost free except basal portion.Occasionally, the tepals are three, but they are almost divided nearly to the base or above the base so that they look like six in number.Irregular lobbing of tepals is also frequently seen.Similarly, the number of stamen also varies from 3-6.The gall flowers which occur throughout the syconium except the ostiolar region have tepals which are same as that of male flowers but has one ovary with a short style.The wasp completes its life cycle in the ovary of the gall flower so seeds are not formed.A pseudo-hermaphrodite flower contains three tepals, Female flowers are present throughout the inside of the female syconia.They are mostly sessile but sometimes pedicellate.In this case the number of tepals varies from 4-5.Like the male flowers a very irregular lobing in tepals has been noticed.Style is subterminal to lateral and comparatively longer than gall flowers.Stigma is usually bifid, however, linear, trifid or indistinctly bifid conditions have also been observed.

Illustration by Rinkey Tiwari
The length of the arms of stigma also varies considerably among different flowers.They may be either equal to each other or unequal.The color of the stigma varies from light to dark orange.Neuter flowers have reduced tepals without pistil and stamen, or sometimes with a staminode.

Distribution Pattern
Ficus palmata subsp.virgata is commonly distributed in northwestern India from the plains to an altitude of 2200m in the Himalayas with a maximum concentration in Himachal Pradesh, Punjab, Rajasthan, Uttarakhand and Uttar Pradesh.Singh (1986) has observed that the taxon is very common in Patna (Bihar).Mooney (1950) has mentioned that the taxon is seemingly rare in Odisha.The report from West Bengal is only based on garden grown plants as the taxon has not been reported earlier (Prain 1903).So far the species has not been reported from the north-east region, which is considered to be one of the richest regions for Ficus in India (Chaudhary et al. 2012), probably due to the high humidity and rain fall prevailing in the region.The frequency of occurrence of the species from the north to south is distinctly diminishing.In the southern region, the species is so far known only from Visakhapatnam District of Andhra Pradesh.Matthew (1999) reported the species from the Palni Hills in Tamil Nadu based on a cultivated plant, which on critical examination turned out to be F. carica.Similarly, its report from Kerala by Kader (2009) is erroneous as it is based on misidentified specimens of Broussonetia papyrifera (L.) L'H'er.ex Vent.

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Cultivated specimen sent to Botanical Garden, Calcutta sent by General Martin from Lucknow.
Phenology: Formation of figs almost throughout the year.
: The bracts are associated with syconium or fig.The number of basal bracts varies from 2-4.The basal bracts are present at the base of the fig body when figs are without stipe or they may occur at the base of the stipe in stipitate figs.The margins of basal bracts are entire or slightly undulate.The apical bracts are found in many rows around the ostiole.The outermost row of 3-6 bracts is compactly arranged whereas the bracts in inner rows are loosely arranged.These bracts show variations in the colour as well as their opening nature.Since F. palmata subsp.virgata is dioecious the male and female figs are located on different plants which are almost indistinguishable through vegetative characters.However, in the male figs the apical bracts protrude outwardly during the time of pollination to allow the insects to enter into the fig.Sometimes the outward projection of the bracts is so pronounced that it makes a clear ostiolar opening through which the stamens present near the ostiole also come outside the fig which may help in segregation of male and female plants and figs.While in the female figs the apical bracts do not protrude outwardly and remain flat.The colour of the apical bracts distinctly varies from faintly to prominently red to pink or sometimes only green with a tinge of red.Image 3. Figs: Fig is one of the important taxonomic characters in the genus Ficus.Traits like their position, number per axil, the presence or absence of peduncle and stipe, shape and size, colour, the presence or absence of hairs and bracts on fig body etc., are frequently used to distinguish the taxa at different levels.However, in the present case the figs are considerably variable in their number per axil, peduncle and stipe length, size and shape, colour etc.In the candidate taxon the figs are axillary and generally solitary.But paired figs are also seen frequently on different plants or on the same plant.As far as the length of stipe and peduncle of fig is concerned, they drastically vary from plant to plant and sometimes they become so reduced that the figs appear almost sessile and give the false appearance of a different taxon.The figs are initially green which turn dark purple to almost black after ripening (Image 2E-F).