ABSTRACT
A preliminary checklist of Papaveraceae s.l. in Indian subcontinent has been prepared on the basis of primary observations of different taxa belonging to this family in wild habitats and on secondary observations based on examining herbarium specimens and taxonomic literature. On the Indian subcontinent (comprising Bangladesh, Bhutan, Myanmar, Nepal, Pakistan, Sri Lanka and India), the family Papaveraceae is poorly represented (22.46%). For better understanding of the species dynamics of Papaveraceae in Indian subcontinent, the present status of phytoendemism has been compared to the data of previous investigations done in nineteenth century. On the basis of phytogeographical analysis, economic potential and survival threat, fossil ancestry of Papaveraceae in Indian subcontinent, its mode of phytoendemism has been categorized under Holoendemics as well as Island endemics. The present studies on dynamics of Papaveraceae in Indian subcontinent substantiate its present mode of panendemism or diversification.
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DOI: 10.3923/ijb.2008.249.259
URL: https://scialert.net/abstract/?doi=ijb.2008.249.259
INTRODUCTION
Papaveraceae s.l. consists of Papaveraceae S.Str. (ca. 23 genera, 240 species); Fumariaceae DC. (16 genera, 500 species); Hypecoaceae L. (1 genus and 18 species) and monogeneric taxon Pteridophyllum Sieb. and Zucc. and it is characterized by its paracarpous gynoecium of two or more carpels (Chase, 1998; APG II, 2003; Kadereit et al., 1994, 1995, 1997). The possession of idioblasts or laticifers with watery or milky sap or latex (Friedel, 1938) and calcium oxalate crystals in the inner epidermis of the outer integument of the seeds (Brueckner, 1985) support Fumariaceae/Hypecoaceae and Papaveraceae S.Str. as monophyletic. Though Pteridophyllum is devoid of these two characters, yet its association with Hypecoaceae has been established by the presence of four unspurred petals and four stamens in these groups (Fedde, 1909, 1936). Hypothesis of Kadereit et al. (1994, 1997) have circumscribed Pteridophyllum as basal clad of Papaveraceae s.l. for two characteristics: possession of two sepals and bicarpellate gynoecium.
From the review of geographical distribution, Papaveraceae s.l. has been considered to be originated and diversified in Laurasia in geological past (Raven and Axelrod, 1974, 1978; Kadereit, 1988a, 1990, 1993a). Papaveraceae s.l. with the members of herbs and shrubs, distributed mostly in temperate and sub-tropical region of N. America, Europe, Asia, Australia and South Africa, having 42 genera and about 690 species (Debnath and Nayar, 1993; Ellis and Balakrishanan, 1993; Grey Wilson, 1993; Hoot et al., 1997; Kadereit, 1886a, b, 1987, 1988b, 1993a, b, 1996; Liden, 1989, 1991, 1995; Ryberg, 1960; Zhuang, 1993; Zuloaga and Morrone, 1996; Liden, 1998; Wu, 1999; Carolan, 2004). Most of the times Papaveraceae s.l. has been studied under four subfamilies: Chelidonioideae (Old world/New World); Eschscholzioideae (New World); Papaveroideae (Old World/New World) and Platystemonoideae (New World) (Schwarzbach and Kadereit, 1994; Hoot and Crane, 1995; Jork and Kadereit, 1995; Kadereit et al., 1995, 1997; Carolan et al., 2006). To have an idea about relative age of extant taxa, ecological analysis of a family is necessary. It is interesting to notice that Pteridophyllum, parts of Fumariadeae tribe Corydaleae p.p. and most members of Papaveraceae S.Str. subfam. Chelidonioideae occur in forest floor and ecologically related habitat; whereas rest of the family: Hypecoum, Fumariaceae tribe Corydaleae p.p., tribe Fumarieae, Papaveraceae subfam. Eschscholzioideae and Papaveroideae incl. Platystemonoideae and Glaucium/Dicranostigma are distributed in arid land, open and disturbed habitats and this transition from forest floor to open habitat has been taken place few times where stimulus like aridification, morphological divergence played the key roles (Schwarzbach and Kadereit, 1994; Kadereit et al., 1995). According to the hypothesis of Stebbins (1952), arid surroundings might have three major effects on evolution of plants. (i) when moisture is limited, local diversity in soil and other environmental factors has a great effect on the vegetation than in the regions where moisture is adequate; (ii) The division of larger sized populations into smaller sized units which are isolated from each other is favored (iii) Selection for specialized structures which helps plants to withstand severe droughts is strong. A modified version of these hypothesis have been put forwarded by Stebbins (1952, 1972, 1975) and Axelrod (1967, 1972) who envisaged that high degree of morphological diversion in arid and semi-arid region might be due to rapid morphological evolution in those areas like diversification and fragmentation of populations leads to geographic isolation followed by natural selection and adaptive radiations push further quantum-evolution. The high degree of morphological divergence in generic level has been explained under fast radiation hypothesis by Macnair (1989), who explained that rapid speciation could occur in response to adaptive challenge when colonizers are faced with vacant adaptive niches and speciate to fill them. It is interesting to notice that ecological divergence has been noticed in the generic level of the family Papaveraceae s.l.; 16 of 23 genera (220 of 240 species) of Papaveraceae S.Str. grows in arid and open habitat whereas 250-300 species of Corydalis of Fumariaceae DC. prefer to grow on forest floor (Liden, 1986; Liden et al., 1995) and those taxa extant in open habitats would have been considered younger than the groups growing on forest floor (Raven and Axelord, 1974, 1978). However, in Old World and New World, the horticultural as well as economic potential of the members of Papaveraceae s.l. should be considered as genetic resource from regional economic aspects; naturally members of this family deserve proper conservation. Apart from ethno botanical importance, the underexploited genetic resources belonging to this family deserve conservation which is on the verge of depletion of wild habitats. The lack of contemporary phytogeographical analysis of Papaveraceae in South Asian region, which is pre-requisite criteria to adopt conservation strategies on a national level as well as regional level, lead to undertake the present studies. The last phytogeographical review of this family on the Indian subcontinent was done in 1939-40 (Chatterjee, 1939, 1962). Thus an appraisal is necessary to monitor the species dynamics of Papaveraceae in the same geographical location after a period of time. Species dynamics is a key indicator in predicting whether a particular group of plants is in a mode of expansion or extinction. Apparently, it is an academic study where economic potential of an under-exploited group of plants is not taken into consideration. Hence, species dynamics should be considered as an important parameter in determining sustainable utilization pattern as well as planning of conservation strategies. For a developing country, underexploited plant resource should be considered as potential natural resource, which could play an important role in socio-economic development. But, most of the time it has been found the conservation initiative, strategies, ethics in developed countries is much more pro-active than developing countries, where underexploited plant is being considered as a potential resource, whereas it is just weed-like plant in other world.
MATERIALS AND METHODS
To prepare a preliminary checklist of Papaveraceae in Indian subcontinent, available floristic works of Indian subcontinent and other major regions have been consulted, starting with Index Kewensis (Hooker and Jackson, 1895) and its supplements. The list of endemic taxa has been prepared from the literature and confirmed from herbaria, viz. ASSAM (BSI, Eastern Circle, Meghalaya), BSIS (Industrial Section of Indian Museum, Botanical Survey of India), CAL (Central National Herbarium, Botanical Survey of India), DD (Herbarium Divn. Forest Research Institute, DehraDun), BSD (Botanical Survey of India, Northern Circle, DehraDun), CDRI (Botany Divn. Central Drug Research Institute, Lucknow), CIMAP (Herbarium, Central Institute of Medicinal and Aromatic Plants, Lucknow), RRCBL (RRL, Bangalore), RRLB (RRL, Bhubaneswar) and K (Royal Bot. Gard., Kew). The microfiches of C. Linnaeus`s collection from two European herbaria, have also been studied viz. Herbarium, London (LINN) and Herbarium, Stockholm (S). In search of endemic and threatened taxa, field surveys have been undertaken between 1999-2005 in parts of North-East India, North-West, Western and Central Himalayas in India. The required data on threatened taxa of Papaveraceae were partially accumulated from World Conservation Monitoring Centre, Cambridge; UK. The assortment of endemic taxa of Papaveraceae in different phytocorias has been made following the phytocorial classification of Kundu (2001).
RESULTS AND DISCUSSION
Distribution of Papaveraceae in Indian subcontinent: In India, the family is represented by ca. 9 genera and ca. 93 species (Debnath and Nayar, 1993; Ellis and Balakrishnan, 1993; Rawat, 1997). During present investigation it has been found that there are ca. 97 taxa (87 species and 10 sub-species) under 9 genera belonging to Papaveraceae distributed in India. The checklist of Papaveraceae of Indian subcontinent (Jaffri and Qaiser, 1974a, b; Whitmore, 1979; Long, 1984a, b; Dahl, 1990; Kadereit, 1993a, b; Kadereit et al., 1994, 1995, 1997; Wu, 1999; Ohba et al., 2006) is given in Table 1.
So, from Table 1, it has been found that there are 155 taxa under 11 genera belonging to Papaveraceae are spatially distributed over Indian subcontinent and other parts of the world.
Table 1: | Checklist of Papaveraceae in Indian subcontinent and other places |
Table 2: | The checklist of endemic taxa of Papaveraceae in India |
+: Presence, -: Absence, Phytocorial Divisions: North-West Himalayas (P-1), Western Himalayas (P-2), Central Himalayas (P-3), Eastern Himalayas (P-4), North East India (P-5), Indian Desert (P-6), Semi Arid Region (P-7), Gangetic plain (P-8), Central Deccan plateau (P-9), Northern Eastern Ghat (P-10), Southern Eastern Ghat (P-11), North Western Ghat (P-12), Central Western Ghat (P-13), Southern Western Ghat (P-14), Coromandel Coast (P-15), Malabar Coast (P-16), Andaman and Nicobar Islands (P-17), Laccadive and Minicoy Is. (P-18) (Kundu, 2001) |
Table 3: | Phytogeographical distribution of endemic taxa of Papaveraceae in India |
Table 4: | Species composition and representation of endemic taxa of Papaveraceae in India |
Endemism of Papaveraceae in India: Analysis on species distribution of Papaveraceae revealed that there are six taxa restricted in distribution in Indian territory (Table 2). From Table 2, it has been found that out of six taxa, there are three taxa belonging to Corydalis sp. two taxa belong to Meconopsis sp. and one taxon belongs to Dicentra sp. Out of these six taxa, one is confined in N. W. Himalayas (P-1) and two taxa are restricted to Western Himalayan (P-2) regions, 2 taxa are confined in (P-3) Central Himalayas and the rest one is extended From N.W. to W. Himalayas (P-1, P-2). Distribution of six endemic taxa of Papaveraceae in different phytographical region is shown in Table 3.
From the Table 3, it is very difficult to designate any particular matrix as endemic matrix of Papaveraceae in India. But, it is clear that the endemic taxa, belonging to Papaveraceae, are confined in Himalayan region, only. However, species composition and representation of endemic taxa, belonging to Papaveraceae in India, is presented in Table 4.
Among the three genera, Corydalis is represented with higher number of 3 endemic taxa with 4.76% endemism, followed by Meconopsis with 11.11% having 2 taxa, whereas Dicentra shows highest degree of endemism i.e., 25% with ca. 1 taxon only. It has been found that C. cavei, D. Long. flowers for a long time from May-October, but other two Corydalis spp. flushes for a short span (August-September) but rest of the three endemic taxa belonging to Dicentra and Meconopsis flowers from July- October. It is also found that Dicentra ventii Khanh occurs in the altitude of 1500-2700 m altitudinal range but Corydalis boweri Hemsley, C. changuensis D. Long, C. duthiei Maxim, occur in greater altitudinal range from 2700-4300 m and two species of Meconopsis genus: Meconopsis latifolia (Prain) Prain and M. robusta Hook.f and Thoms. occur in the intermediate altitudinal range between 2400-4000 m.
Endemism of Papaveraceae in Indian subcontinent: In the Broader aspect, extent of endemism of Papaveraceae in Indian subcontinent is tabulated in Table 5.
From Table 5, it has been found that there are ca.44 taxa, confined in Indian subcontinent. Out of ca.44 taxa, eight taxa are confined in Indo-Pakistan region (i.e., N. W. Himalayan regions), two taxa are confined in Indo-Pakistan-Nepal (Northwestern-Western Himalayan regions), nine taxa are confined in Indo-Bhutan region (Central Himalayan regions), fourteen taxa are confined in Indo-Nepal region (Western Himalayan regions), six taxa are confined in Indo-Nepal-Bhutan (Central-Western Himalayan regions) region, two taxa are spatially distributed over Indo-Nepal-Myanmar region (Western Himalayas-gateway of South East Asian regions) and rest of the two taxa are confined in Indo-Mayanmar region (i.e., Gateway of Southeast Asian regions). In the broader aspect, those taxa, which are found to be restricted distribution in Indian subcontinent, should be treated as Broad Range Endemics or B.R.E. (Kundu, 2005). From the distributional pattern of B.R.E., it has been found that Western and Central Himalayan region is the congenial abode for endemic taxa of Papaveraceae in Indian subcontinent (Fig. 1), on the other hand the strictly confined six taxa in Indian territory, should normally be treated as Narrow Range Endemics or N.R.E. (Kundu, 2005), distributed in Northwestern Himalayas and Western Himalayas, respectively. Comprising N.R.E.s and B.R.E.s there are 50 taxa restricted (Endemic) in India subcontinent (E.I.S.) and out of 50 taxa, 33 are restricted in Central and Western Himalayas, which helps to identify the are as Ideal resort for endemic taxa belonging to Papaveraceae s.l.
Table 5: | The checklist of endemic taxa of Papaveraceae in Indian subcontinent |
+: Presence, -: Absence, #: Data is not available |
It has been found that the total number of taxa belonging to Papaveraceae on the Indian subcontinent is 155; whereas the total number of taxa in India is 103 (66.45%). The number of endemic taxa in Indian subcontinent is not so impressive: 50 (32.25%). The number of Narrow Range Endemics is 6 (3.87%) and the number of Broad Range Endemics is 44 (28.38%). From the retrospective viewpoint of species dynamics, the percentage of endemism of Papaveraceae on Indian subcontinent in 1939-1940 (Chatterjee, 1939) was 65.11%, whereas in 2004-2006, it has become 32.25%. The negative index of phytoendemism could be defined as either decreasing number of endemic taxa or further expansion of distributional area of endemic taxa or combination of both.
Threatened taxa, belonging to the family Papaveraceae in India: From the viewpoint of fiscal evaluation of natural resources, the increasing rate of phytoendemism is an important index for national economy of any developing state but shrinkage of a particular germplasm due to habitat destruction and anthropogenic interference is a key negative gradient from conservation as well as economic viewpoint. If the potential resource is underexploited the loss is irreversible the nation as the diminishing resource disappears forever with its unknown potential without further contributing to human being. Though Argemone mexicana var. mexicana is neither threatened nor endemic but it would be considered as underexploited plant resource which has recently been found having antibacterial properties against Gram positive (Staphylococcus aureus, Bacillus subtilis) and Gram negative (Escherichia coli, Pseudomonas aeruginosa) bacteria (Bhattacharjee et al., 2006). The endemics of Papaveraceae and locally confined pan endemics with isolated small patches with less than minimal viable population size are the outfall of habitat disturbance and anthropogenic interference like ethno botanical utilization, conversion of wild habitat to cultivation field, urbanization etc. and over-exploitation of wild germplasm (e.g., Meconopsis smilacifolia, M. grandis, M. betonicifolia found first time in wild habitat between 1880-1886 and germplasm have been successfully introduced in cultivation since end of eighteenth century as ornamental plants) (Chittendon, 1956; Huxley, 1992; Polunin and Stainton, 1984). It is ascertained that there are five taxa of Indian Papaveraceae facing various degree of threats (Table 6), mainly due to anthropogenic exploitation (over-exploited by nurserymen) for its horticultural importance and medicinal potentialities (e.g., Meconopsis napaulensis), chiefly used as folklore and ethno medicine for its narcotic properties and poisonous plants (e.g., Papaver dubium, P. nudicaule, P. orientale etc.) (Chittendon, 1956; Chopra et al., 1986; Envis Centre, 2007; Facciola, 1990; Kunkel, 1984; Tanaka, 1976).
Fig. 1: | An outline of endemism of Papaveraceae in Indian subcontinent |
Table 6: | Plants facing survival threat belonging to the family Papaveraceae in India |
#: Data is not available, I: Indeterminate, E: Endangered, I.S.P.: Isolated Small Population, Ec: Economic, Hort.: Horticultural (Ornamental), V: Vulnerable, Fl: Flower |
Though Papaver somniferum var. somniferum (widely cultivated species) is not threatened plant belonging to this family in India but economy of Indian subcontinent has been highly influenced by opium trade (psychoactive chemical derivative of this species) since Sixteenth century. There are two major centers of opium production in two opposite side of Indian subcontinent: Golden Triangle (comprising of Myanmar, Laos and Thailand) and Golden Crescent (comprising of Pakistan, Afghanistan and Iran). The US Department of States estimates that the Golden Triangle led world production until 1991, with Myanmar ranking at top and afterwards Golden Crescent took the lead with Afghanistan breaking the previous record in 1999.
Possible fossil evidence in relation to endemics of Papaveraceae in India: The phylogeny of Papaveraceae s.l. has suggested that the presence of multistaminate androecium helped to recognize it as primitive family (Murbeck, 1912; Karrer, 1991). The absence of nectariferous stamens and presence of multistaminate stamens indicates towards the pollination pattern done by pollen collecting insects and this feature further strengthen the hypothesis of primitive origin. It is interesting to focus on phylogenetic relationship by studying the fossil remains of the ancestors of the modern taxa, which are referable to the endemic taxa of the family Papaveraceae s.l. now-a-days. The geographical distribution of Papaver S.Str. and Papaver sect. Horrida and Fumariadeae tribe Fumarieae in both Mediterranean to Central Asian region and in eastern; South Africa suggests that these taxa supposed to be existed in the middle to late Tertiary when drastic aridification allowed an expansion of the distributional range across the equator (Kadereit, 1988a, b; Kadereit et al., 1995). The fossil remnant of Papaveraceae in India has been represented by Papaveraceaepites thalmanii Biswas, of Lower Eocene period, Tertiary era, found in N. E. India, particularly in Garo Hills, Meghalaya (Biswas, 1962). The fossil taxon has little affinity with the living Papaver spp. except Papaver somniferum L., which is a panendemic taxon, whereas other Papapver species are concentrated in N. W. Himalayan region. Though lack of fossil remnants of Meconopsis in Papaveraceae has helped little to establish the relationship between fossil ancestors and living, endemic representative but the existing fossil representatives of other genera belonging to Papaveraceae during Tertiary era doesn`t help to predict it as Palaeoendemics whereas the morphological data particularly floral morphology indicates towards primitiveness; which feature refrained us to categorize the endemics as Neoendemics; so categorically it might be considered under Holoendemics as well as Island endemics, confined on Himalayan mountain ranges, the physically island like habitats.
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