Indopiptadenia paper

PHYTOTAXA

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2014 Magnolia Press

Phytotaxa 164 (2): 061–078

www.mapress.com/phytotaxa/Article

http://dx.doi.org/10.11646/phytotaxa.164.2.1

Taxonomy of a monotypic genus Indopiptadenia (Leguminosae-Mimosoideae)

OMESH BAJPAI1, AWADHESH K. SRIVASTAVA2, ARUN K. KUSHWAHA3 & LAL BABU CHAUDHARY4*1 Plant Diversity, Systematics and Herbarium Division, CSIR-National Botanical Research Institute, Lucknow-226 001, India.

E-mail: [email protected] 2Plant Diversity, Systematics and Herbarium Division, CSIR-National Botanical Research Institute, Lucknow-226 001, India.

E-mail: [email protected] Plant Diversity, Systematics and Herbarium Division, CSIR-National Botanical Research Institute, Lucknow-226 001, India.

E-mail: [email protected] 4 Plant Diversity, Systematics and Herbarium Division, CSIR-National Botanical Research Institute, Lucknow-226 001, India.

E-mail: [email protected]

* Corresponding author

Abstract

Indopiptadenia, a monospecific genus with the species I. oudhensis, is confined to the Indo-Nepal border area in

scattered populations along the Himalayan foothills between 156–908 m elevation. I. oudhensis has been little studied

since its discovery in 1874. The taxonomy of the genus is reexamined here with extensive notes on diversity, distribution,

phenology, ecology and conservation status based on exhaustive field survey and examination of herbarium specimens.

A full description including data on many new characters and encompassing all morphological variations is provided to

better characterize the species so that its correct systematic position can be ascertained and provide the basis for proper

conservation strategies. Placed in the tribe Mimoseae, the species shows more affinity towards the Newtonia group than

the Piptadenia group. However, it differs from the former in having unarmed to armed stem, uni- or bijugate leaflets,

absence of stemonozone and pods that dehisce generally by rupturing of the pod valves over the seed chambers to leave

a persistent replum. The IUCN category Near Threatened (NT) is assigned to the taxon.

Key words: South Asia, endemic, Mimoseae, Indopiptadenia oudhensis, Near Threatened

Introduction

Indopiptadenia Brenan (1955: 178), is a monotypic genus with the species Indopiptadenia oudhensis (Brandis

[1874: 168]) Brenan (1955: 178) and is placed in the subfamily Mimosoideae, family Leguminosae (Brenan 1955;

Hutchinson 1964; Lewis & Elias 1981; Sanjappa 1992; Kumar & Sane 2003; Luckow 2005). I. oudhensis is

endemic to the Terai region of the Indo-Nepal border area in the Bhabar zone along the Himalayan foothills in

tropical moist deciduous forest (Champion & Seth 1968). Biswas & Chandra (1997) and Prakash et al. (2009) have

incorrectly suggested that the genus Indopiptadenia is found in America and Africa.

I. oudhensis is a striking evergreen small to medium sized tree with upright branches and profuse pendulous

branchlets and leaves. Recent surveys (Kashyap 2009; Prakash et al. 2009; Singh 2010) in India claim that during

the last more than 100 years the population of the trees has substantially decreased due to exploitation by local

people, habitat destruction and low regeneration ability of the species in natural habitats, and that as a result the

species has become rare and threatened.

During the course of study over the last 5 years for the tree flora of Uttar Pradesh, India, the authors have

regularly visited different forest areas where they discovered populations of I. oudhensis in the forest of Balrampur

district adjacent to the Nepal border. The plants were also discovered in the adjacent area within Nepal in Dang

district. Subsequently, the species was found in additional localities in Champawat district of Kumaun region in

Uttarakhand State in India. After studying the available information (Duthie 1903, 1906; Biswas & Chandra 1997;

Prakash et al. 2009) it was realized that although the species was discovered more than 100 years ago, it has been

Accepted by Ruth Clark: 14 Mar. 2014; published: 4 Apr. 2014 61

mailto:[email protected]




little studied since its original description, and remains poorly known. During recent years additional collections

have been made, and more areas have been surveyed, whilst the overall composition of the forest has substantially

changed due to various anthropological and ecological factors. Since it is poorly known, detailed investigation is

required to understand the species before conservation strategies for its protection can be formulated (Rana &

Ranade 2009). In view of this, the present study revisits the taxonomy, diversity, phenology, ecology and

conservation status of the species to properly characterize it, describing many new features for the first time.

Methods

The area of occurrence of the species was visited 9 times and plants were collected in all seasons to prepare

voucher specimens and record the phenological data. The earlier herbarium collections housed at BSD, DD,

K-Photos, LWG (herbarium acronyms according to Thiers, 2012) were also critically examined. Images of the

type specimens were studied from the website of RBG Kew (http://apps.kew.org/herbcat/navigator.do) to

authenticate the identity of the species. Fresh flowers, fruits and seeds were observed and photographed to

enable new description to be made. A preliminary conservation assessment of the taxon based on IUCN Red List

categories and criteria, version 10.1 (IUCN Standards and Petitions Subcommittee 2013) was produced using the

GeoCAT tool (http://geocat.kew.org/) to find out Extent of Occurrence (EOO) and Area of Occupancy (AOO)

values (Bachman et al. 2011), and then analysing this result in the context of further information available.

Results

A detailed description of the species is provided here. This includes many morphological features not described in

the original publication, such as the type of prickles on the stem, variation in leaves, pubescence of young leaves,

type of extrafloral nectaries on petiole and leaf rachis, attachment of flowers on the floral axis, arrangement of

stamens on annular cup, characters of bracts, colour of different stages of leaflets, petals and pods, new dehiscence

pattern of pods and variations in the shape and size of seeds. The genus shows close affinity to genera of the

informal Newtonia group (sens Luckow 2005) and exhibits tremendous variations in thorns, bark, leaves,

extrafloral nectaries on the rachis, size of pods and shape and size of seeds. Detailed comments on the current

conservation status, distribution, ecological associations and population size have been provided which explain that

although the distribution of species is patchy, it occurs frequently over a wide area, and may be considered as Near

Threatened.

Morphological diversity

Habitat and habit:—The species grows chiefly in gravelly-sandy soil on foothills of the Himalayas along the

edges of rivers and near streams. It is also found on flat lands in dry and hard rocky soils and on dry mountain

slopes along roadside. The plants are small to medium sized erect trees, either multi stemmed from the base or with

short main trunk with upright branches and profuse pendulous branchlets on which numerous downwardly oriented

leaves are found (Figure 1).

Stem:—In I. oudhensis the stems can be either densely prickly or without prickles, and both forms can occur in

the same population (Figure 2A, B). This is the first report of armature in Indopiptadenia, as Hutchinson (1964)

and Luckow (2005) previously have not considered Indopiptadenia to be armed. The prickles are either found only

on the main trunks of trees or distributed throughout, even on young branches. With age, the prickles degenerate to

leave scars on the stem. The bark shows wide range of variation on different plants within the same population. It

may be smooth or wrinkled but in older trees it becomes rough due to formation of exfoliating woody scales.

(Figure 2)

Leaves:—Young flush leaves are dark red, gradually becoming dark green at maturity (Figure 3). The leaves

are quite variable on the same plant (Figure 4). They are bipinnate, commonly with two pairs of opposite pinnae

(Figure 4 D). However, leaves with one pair of pinnae are also seen frequently (Figure 4 A). Sometimes one

additional pinna has also been observed on the primary rachis (Figure 4 L). Rarely the lower pair of pinnae are

subopposite (Figure 4 M). Occasionally ternate leaves are also observed either with one pair or two pairs of leaflets

(Figure 4 C, J). Generally, each pinna consists of only one pair of leaflets at its apex, which ends in a short pointed

BAJPAI ET AL.62 • Phytotaxa 164 (2) © 2014 Magnolia Press

mucro (Figure 4 A, C–E, M); however, the number of leaflets and their position on the pinna rachis varies between

leaves (Figure 4 B, F–K, N, O). In place of two leaflets, there may be four leaflets on any or all of the pinnae

(Figure 4 B, F–K, M, N) and in this way the number of leaflets per leaf may vary between 4 to 16. An aberration

with only three leaflets has also been seen rarely on some pinnae (Figure 4 K). Occasionally, the primary rachis is

absent or quite reduced so that the leaf appears palmatipinnate (Figure 4 E). Variations have also been noticed in

the shape and size of leaflets. The extrafloral nectaries found on the petiole and primary and secondary rachis show

variation in shape and size. Either they are circular in outline or may have two to three indistinct lobes (Figure 5 A–

C). Generally they are found at the junction of each or any one of the pinnae either on the petiole or primary rachis,

but rarely occur on the secondary rachis either in the middle portion or at the apex (Figure 5 D–F). The extrafloral

nectaries are smaller in size on the secondary rachis than other extrafloral nectaries found on the petiole and

primary rachis.

FIGURE 1. Indopiptadenia oudhensis: A–F, Habit and habitats; G, Close up of habit.

Phytotaxa 164 (2) © 2014 Magnolia Press • 63TAXONOMY OF INDOPIPTADENIA (LEGUMINOSAE-MIMOSOIDEAE)

FIGURE 2. Indopiptadenia oudhensis: Portion of stem showing variation in bark: A, Smooth bark without prickles; B, Bark with

prickles; C, Young branch with prickles; D, Mature branch with prickles; E–H, Degeneration of prickles with age; I, Bark after

prickles have fallen; J, Bark with exfoliating woody scales in fully matured tree; K, Blaze.

Inflorescence:—Axillary and terminal spike or raceme of spikes. When the inflorescence is axillary, it is

simple (unbranched), but when terminal it becomes branched with 3–7 spikes. In branched inflorescences generally


the terminal spike matures before the lower ones (Figure 6 D), although lower spikes have also been observed

maturing before terminal ones (Figure 6 C). The flower petals are dark red in bud but gradually turn yellowish

green and finally brown in colour (Figure 6 A–D). The petals are free from each other and similar in shape and size.

FIGURE 3. Indopiptadenia oudhensis: Different stages of leaf formation.

Pods:—The size of the pods varies significantly within the same plant. The deciduous, pendulous pods are

usually straight but falcate pods are also not uncommon (Figure 7 B). The young pods are dark red, gradually

turning green at maturity and finally brown when ripe (Figure 7 A). The dehiscence of pods is a point of interest.

Earlier it was thought that the pods dehisce through both margins (Figure 7 G). However, here for the first time it

was observed that generally the dehiscence of the pods takes place by the bursting of the fruit wall at the seed

position when they are on the plants (Figures 7 C, E). After dehiscence they usually fall on the ground (Figure 7 D).

However, when the undehisced pods fall on the ground or sometimes when they remain on the tree, they

downwardly (from apex to base) dehisce along single or sometimes both margins (Figure 7 F, G).

Seeds:—Seeds are placed singly or in pairs at a distance of 7–10 mm or they are overlapping to each other

(Figure 8 A–C). The shape and size of the seeds varies considerably between pods (Figure 8 D, E). When the seeds

are oblong they are arranged parallel to the pod. The circular and oblong seed forms never occur together within the

same pod. The seeds are dispersed by wind as they are light weight and winged.


FIGURE 4. Indopiptadenia oudhensis: Variations in leaves.

Taxonomic treatment

Indopiptadenia Brenan, Kew Bull. 1955: 178. 1955; Hutch., Gen. Fl. Pl. 286.1964; Lewis & Elias in Polhill &

Raven, Adv. Legum. Syst. 1:161.1981; Sanjappa, Legum. India 66. 1992; Kumar & Sane, Legum. S. Asia: Checkl.

118. 2003; Luckow in Lewis et al., Legum. World 170 (Fig.) & 171. 2005.

Type species:— Indopiptadenia oudhensis (Brandis) Brenan.

Small to medium sized tree, unarmed or armed with prickles. Stipules caducous. Leaves alternate, bipinnate; rachis

glandular; pinnae opposite, generally bifoliolate or 4-foliolate; leaflets large, subsessile, pinnately nerved.

Inflorescence a pedunculate spike or raceme of spikes. Bracts minute, deciduous. Calyx petaloid, cup shaped,


minutely 5-teethed, glabrous. Petals 5, valvate, completely free. Annular disk present. Stamens 10, free, just

exserted, anthers with terminal globose gland. Ovary stipitate, glabrous; style longer than the stamens; stigma

tubular. Pods deciduous, usually straight or sometimes falcate, flat, glabrous, unilocular, dehiscing by rupturing of

the fruit wall at the seed position or by one or both margins. Seeds compressed, thin, winged on both sides.

FIGURE 5. Indopiptadenia oudhensis: Variations in position and structure of extrafloral nectaries.

FIGURE 6. Indopiptadenia oudhensis: Developmental stages of inflorescence.

Indopiptadenia oudhensis (Brandis) Brenan, Kew Bull. 1955: 179. 1955; Henry & Roy, Bull. Bot. Surv. India 10:

275. 1968; Ohashi in Hara & William, Enum. Fl. Pl. Nepal 2: 123. 1979; Sanjappa, Legum. India 66. 1992; Biswas

& Chandra, Indian Forest. 123 (5): 419. 1997; Kumar & Sane, Legum. S. Asia: Checkl. 118. 2003; Prakash et al.,

Biodiv. News 1(1): 2. 2009.

Piptadenia oudhensis Brandis, For. Fl. N. W. & C. India 168. 1874; Benth., Trans. Linn. Soc. 30: 366. 1875; Baker in Hook. f.,

Fl. Brit. India 2: 289. 1879; Duthie, Fl. Upp. Gang. Pl. 1: 308. 1903; Duthie, Ann. Roy. Bot Gard. Cal. 9: 33, t. 43. 1906;

Brandis, Indian Trees 261. 1906; Kanjilal, For. Fl. Pilibhit, Oudh, Gorakhpur & Bundelkhand 161. 1933.


Holotype:— India: Ind. Or., Oudh (‘Oude’), R. Thompson 135 (communicated by Dr. Stewart July 1871) (K, photo!).

Adenanthera oudhensis J. L. Stewart, MSS; sensu Kanjilal, For. Fl. Pl. Uttar Pradesh 2: 285. 1966 (misapplied name).

Vernacular name:—Genti or Gainti (Hindi); Katrar (Kumauni); Hathipaula, Kaphali Tree, (Nepali).

(Figure 1–11)

FIGURE 7. Indopiptadenia oudhensis: Pods and their dehiscence: A, Mature pods on the tree; B, Variations in the shape and size of

pods; C, Dehisced pods on the tree; D, Fallen dehisced pods on the soil; E, Dehiscence by rupturing of fruit wall at seed position; F,

Dehiscence by single suture; G, Dehiscence by both sutures.

FIGURE 8. Indopiptadenia oudhensis: Seeds: A–C, Variations in the arrangement of seeds in the pod; D, Variation of shape and size

of the seed; E, Mature seed.

Tree, 5–15 m tall, evergreen. Main stem 1–2 m long, 17–75 cm in girth, unarmed or armed with prickles; branches

upright with profuse pendulous branchlets; branches and branchlets with or without prickles, smooth or wrinkled;

prickles large, conical, hard, sharp pointed, generally degenerating at maturity and leaving scars on old stem; bark of

stem thick, grey, brown or dusky red, smooth, wrinkled or rough with flat exfoliating woody scales; blaze red,

fibrous; wood light red, close grained, hard. Stipules 1–2 mm long, caducous, triangular, subacute at apex, glabrous

on both side or pubescent outside, glabrous inside, always densely villous along margins. Leaves 11–22 cm long,


quite variable, alternate, bipinnate, generally with two pairs of pinnae or sometimes with single or two pairs of

pinnae plus a single or pinna occasionally ternate; petiole 2.2–5.5 cm, terete, pulvinus, minutely pubescent only

when young, otherwise glabrous, bearing one or rarely up to three circular extrafloral nectaries at the junction of

first pair of pinnae; primary rachis 2–10 cm, terete, glabrous, minutely pubescent when young, otherwise glabrous,

with or without extrafloral nectaries at the junction of the pinnae; pinnae 1.5–6.0 cm, (1–) 2 (–3) pairs, opposite,

each bearing one or two pairs of opposite leaflets, lower pair smaller than upper pair; secondary rachis 1.3–5.0 cm

long, with pulvinus, in other details similar to main rachis, sometimes bearing small extrafloral nectary at the apex

(at junction of leaflets) or in the middle; leaflets 4–16, 5–10 × 2.0–8.7 cm, variable, bright red when young, dark

green at maturity, minutely petiolulate, coriaceous to subcoriaceous, smooth above, glaucous below, obovate,

broadly obovate, obliquely broadly obovate, oblong-elliptic, broadly elliptic or ovate-rhomboid, cuneate or oblique

at base, margins entire and generally undulate, obtuse, truncate or rarely retuse at apex, glabrous on both sides,

pubescent on lower side only when quite young, veins prominent on both sides, brochidodromous; petiolules up to 4

mm long, thick, wrinkled. Inflorescence an axillary spike or terminal raceme of spikes, pendulous, shorter than the

subtending leaf; peduncle c. 10 mm long, minutely adpressed pubescent; floral axis minutely adpressed pubescent;

FIGURE 9. Indopiptadenia oudhensis: Holotype.


spikes 2.5–4.0 × c. 0.8 cm, cylindrical, many-flowered, the flowers densely arranged. Bracts c. 1 mm long,

caducous, subulate, acute or obtuse at apex, entire and densely hairy along margins, almost glabrous to densely

hairy outside, glabrous inside. Flowers 5–6 mm long, sessile, jointed on floral axis with a circular structure at base,

after falling leaving basal part on floral axis, basal part falling later. Calyx c. 1 mm long, petaloid, persistent, cup

shaped, minutely 5-teethed, glabrous. Petals 5, 4.0–4.5 × 0.9–1.0 mm, valvate, equal, completely free to base,

fleshy, greenish-yellow, becoming brownish at maturity, dark red in bud, lanceolate, acute at apex, entire along

margins, glabrous, faintly veined. Stamens 10, free, almost equal, just exerted from corolla; filaments 4.5–5.0 mm

long, weakly marcescent on developing pods, filiform, glabrous, attached to the outer edge of annular disk in two

rows, five per row; anthers 1 × 0.8 mm, versatile, yellowish-green, broadly oval with a sessile terminal globose

gland, dehiscing longitudinally. Ovary c. 2 mm long, cylindrical, stalked, glabrous, unilocular, 15–25-ovuled, stipe

c. 0.5 mm long, ovules minute, reniform, arranged in two rows on dorsal suture; style c. 3 mm, off-white with slight

tinge of pink, filiform, longer than the stamens, straight or slightly curved towards apex, glabrous; stigma terminal,

capitate, surrounded by a tubular hyaline structure, glabrous. Pods 13–34 × 1.0–1.3 cm (including stipe),

pendulous, linear-oblong, subobtuse or acute at tip, cuneately narrowing towards base into a 1.5–3.0 cm long stipe,

usually straight or sometimes weakly falcate, flat, faintly raised around seeds, thickened along both sutures, red

when young, green with age, reddish-brown after ripening, glabrous, unilocular, dehiscing by the rupturing of the

fruit wall at the seed position or by longitudinal splitting along single or both margins from apex to base. Seeds 11–

22 per pod, 4–20 × 4–10 mm, compressed, thin, light weight, winged on both sides, broadly ovate, oblong-elliptic,

or reniform, attached on dorsal suture with a funicle, placed either in pairs or singly, 7–10 mm apart, parallel with

the pod sutures and overlapping when oblong, green when young, brown at maturity, glabrous.

FIGURE 10. Indopiptadenia oudhensis: A, Close up of mature flower; B, Calyx; C, Petal; D, Basal part of flower showing attachment

of filaments; E, Stamen; F, Anther; G, Anther after dehiscence; H, Pistil; I, Stigma.

Distribution:—India (Uttar Pradesh, Uttarakhand), Nepal (Central and Western Nepal: Kapilbastu to

Kanchanpur). (Figure 12)

I. oudhensis was first collected from the Gonda division in the forest of Oudh, a region of the British Indian

State ‘United Provinces of Agra and Oudh’ (now known as Uttar Pradesh) adjacent to the hills of Nepal by Richard

Thompson in 1871 (Brandis 1874, 1906; Duthie 1903, 1906). Subsequently, it was also found in Eastern Kumaun

(now in Uttrakhand) by A. F. Brown in 1883. The Gonda division presently constitutes three districts, Balrampur,

Shrawasti (partially) and Gonda s. s. The survey of the species in India in the current years (Kashyap 2009; Prakash

et al. 2009; Singh 2010) as well as in our own investigation reveals that the species occurs only in the Balrampur

district of Uttar Pradesh and E. Kumaun (Tanakpur, Champawat dist.) of Uttarakhand. All the previous collections

of specimens made from Gonda division fall now within Balrampur district. The herbarium records show that the


species has also been collected in the past from Bahraich district (adjacent to Balrampur district), however, in the

recent surveys (Panigrahi 1969; Saini 2005a, b; Bajpai et al. 2012) it has not been seen in the area, nor in the

adjacent border area of Lakhimpur Kheri (Singh 1997). Kanjilal (1933) has reported it from Gorakhpur district (on

the banks of Gandak river near Nepal Frontier), but it has not been included in the district flora by Srivastava

(1976), and no specimens have been seen from this region.

FIGURE 11. Indopiptadenia oudhensis: A, Habit; B, Twig with inflorescence; C, Flower; D, Calyx (opened); E, Opened flower

(calyx removed); F, Anther (front view); G, Anther (back view); H, Pistil; I, Twig with pods. (Reproduced from Duthie 1906).


FIGURE 12. Distribution of Indopiptadenia oudhensis.

From Nepal the species was first reported by Duthie (1903, 1906) based on Inayat Khan’s collections made in

1900. Since then, only 6 collections have been made in this country. Our recent surveys and personal

communication with the botanists of Nepal (Prof. Mohan Siwakoti, Central Dept. of Botany, Tribhuvan University,

Kathmandu, Nepal) reveal that the species grows in the border districts from Kanchanpur to Kapilbastu in scattered

populations.

Phenology:—Flowering: March - April, Fruiting: April–May (Table 1).

TABLE 1. Different phenophases in Indopiptadenia oudhensis at Balrampur, Uttar Pradesh, India.

Ecology:—I. oudhensis grows in gravelly sandy soils along Himalayan foothills between 156–908 m elevation

in forests near streams, along rivers banks, on mountain slopes and along roadsides as well as on flat lands. The

species most commonly found in association are Colebrookea oppositifolia Smith (1806: 111), Holoptelea

integrifolia (Roxburgh (1795: 56)) Planchon (1848: 259), Woodfordia fruticosa (Linnaeus [1759: 1045]) Kurz

(1871: 56), Acacia catechu (Linnaeus f. [1782: 439]) Willdenow (1806: 1079), Murraya koenigii (Linnaeus [1771:

555, 563]) Sprengel (1817: 315), Mallotus philippensis (Lamarck [1786: 206]) Müller Argoviensis (1865: 196),

Azadirachta indica Adrien Jussieu (1830: 221) and Ficus semicordata Buchanan-Hamilton ex Smith (1810: 71).

Phenophases Time Period

Vegetative phenology

1 Initiation of leaf bud bursting Early February

2 Emergence of young leaves March

3 Time required for leaf maturation 1–2 months

4 Average duration of the leaf 9–11 months

5 Nature of the tree species Evergreen

Reproductive phenology

6 Flower bud initiation Mid March

7 Appearance of mature flower Early April

8 Duration of flowering 15–20 days

9 First appearance of pods Mid April

10 Last appearance of pods Mid May

11 Duration of fruiting 15–20 days

12 Dehiscence of pods Early May

13 Length of reproductive stage April-May


Affinities:—I. oudhensis was first described by D. Brandis in 1874 in the genus Piptadenia Bentham (1840:

135) and until 1955 it was considered part of this genus. Piptadenia is a tropical South American element with c.

15 species (Lewis & Elias 1981). Brenan in 1955 observed that the Indian materials identified as Piptadenia

oudhensis Brandis (1874: 168) do not fit into Piptadenia because they have leaves that consist of 1 or 2 pairs of

pinnae, the pinnae being uni- or bijugate, eglandular petiole, glandular leaf rachis, free corolla lobes and elongated

seeds; he thus erected a new genus Indopiptadenia to accommodate the Indian plants. However, he (Brenan 1955)

mentioned that Indopiptadenia has greater morphological affinity with Piptadenia as both have pods that dehisce

along both the sutures. Recently, Luckow (2005) has mentioned that Indopiptadenia has a superficial affinity with

Piptadenia and following Lewis & Elias (1981) placed the genus in the Newtonia group of tribe Mimoseae Bronn

(1822: 130). Lewis & Elias (1981) separated this species from the Piptadenia group chiefly based on: pedicels

jointed with generally persistent basal part, petals free, anther gland present, pollen grains single, intrastaminal disc

present, style tip tubular, fruits long, flat, coriaceous, opening along one margin or both and seeds winged (Table

2). Indopiptadenia is undoubtedly closer to the Newtonia group, but it differs in having unarmed to armed stem,

uni- or bijugate leaflets, absence of stemonozone and dehiscence of pods by rupturing of the fruit wall at the seed

position. It suggests that Indopiptadenia requires a separate identity within the tribe Mimoseae. Indopiptadenia is

also sometimes confused with the genus Adenanthera Linnaeus (1753: 384) which chiefly differs in having a pod

with valves that often contort after opening and seeds which are thick, with a hard red or bicoloured testa. In

general appearance it resembles with Hardwickia binata Roxburgh (1819: 209) and Bauhinia variegata Linnaeus

(1753: 375) (Leguminosae-Caesalpinioideae).

TABLE 2. Comparison between Piptadenia group, Newtonia group and Indopiptadenia.

Piptadenia group Newtonia group Indopiptadenia

Habit Herb, shrub or tree Tree Tree (small to medium)

Stem Armed or unarmed Unarmed Unarmed or armed

Leaflets Opposite or alternate, multijugate Opposite or alternate,

multijugate

Opposite, uni- or bijugate

Extrafloral nectaries Present, diverse or absent Present except

Piptadeniastrum

Present and diverse

Inflorescence Capitate or spicate Racemes or spiciform racemes Spicate or raceme of spikes

Pedicel Sessile or subsessile Jointed or not, basal part

usually persistent

Jointed or not, basal part usually

persistent

Petals Joined or free Free Free

Stemonozone Sometimes present Often present Absent

Anther-gland Present or absent Present except some African

Newtonia

Present

Pollen grains 4–28-grained Single Single

Intrastaminal disc Present or absent Generally present Present

Style apex and stigma Tapering to a narrow porate

stigma

Funnel-shaped to tubular Tubular

Fruits Papery to pulpy, sometimes

constricted between the seeds,

straight or coiled

Long, flat Mostly straight, sometimes

falcate, long, flat

Dehiscent Indehiscent or dehiscent along one

or both margins or breaking up to

leave a ± persistent replum

Along one margin Rupturing the fruit wall at the

place of seed present or by single

or sometimes both margins

Seeds Winged or not Winged, usually on long

funicle

Winged, funicle short or long


FIGURE 13. Seedlings of Indopiptadenia oudhensis in natural habitat.

Regeneration of plants in nature and threat:—In comparison to the large number of seeds produced on each

tree, few seedlings have been observed in the area of the occurrence of the species. However, the seedlings are

more commonly found growing away from the parent trees, especially in gravelly sandy soils and among boulders

(Figure 13 A–E). The pods dehisce when they are on the plants and the winged and light weight seeds are dispersed

by air away from the parent trees. The seeds taken from the fallen undehisced pods do not germinate in the garden/

nursery, perhaps because they are immature. No trace of germination of seeds has been seen in the fields from the

fallen pods near the trees while the fallen pods remain there till the rainy season. Since the seedlings and young

plants are grazed by animals, their chances of survival are very low in natural conditions. This is perhaps one of the

major reasons why the species does not regenerate strongly in natural conditions in spite of a large number of

seedlings. Secondly, the species grows on hill slopes and foothills near streams and river, which can result in the


seeds being washed out from the original habitats to unsuitable areas. The cutting of river banks every year and

destruction due to floods in these areas also causes severe damage to the trees, especially young ones and those

growing along the riversides.

Sometimes the sprouting of young plants has been noticed from the buried old branches in the soil or remnants

of the older trees (Figure 13 F). This indicates that the propagation of the species may also be possible vegetatively

through stem cuttings. Since the seedlings often do not grow into mature trees in the natural habitats due to grazing

by animals, they must be taken out from the fields to the nurseries/gardens for their protection and propagation.

Conservation status:—More than 100 years ago the species was considered common especially in the Nepal

region (Brandis 1874; Duthie 1903, 1906). It has a fairly wide distribution from Kanchanpur to Kapilbastu districts

along the forest margins (Figure 12). However, sometimes the species has been treated as endangered to critically

endangered, at least in the Indian region (Biswas & Chandra 1997; Prakash et al. 2009; Catalogue of Life: 25th

June 2013). Recently, the Government of India in consultation with the Provincial Government of Uttar Pradesh

(vide notification no. S. O. 998 (E) dated 15 April 2009) has recorded the species as ‘on the verge of extinction’

(endangered) and prohibited its collection in order to conserve it (Goyal 2009). Inside the Indian Territory the

species is confined to 2–3 populations with 10–15 individuals at each location in Balrampur district; however they

are also seen frequently in E. Kumaun region up to 908 m elevation. Although, the species is not dominant in any

locality, it can be easily seen in many common habitats. The seedlings are also seen in an apparently healthy

condition in many places. A preliminary conservation assessment based on EOO (8,410.53 km2) and AOO (68.00

km2) values (Figure 14) suggests the status of the taxon to be Vulnerable (VU) and Endangered (EN) respectively

according to B1 and B2 of the IUCN criteria. A full IUCN conservation was not carried out for the taxon, but it was

evaluated according to the conditions of Criterion B. It does not meet the requirements for B (a), (b) or (c) of

population fragmentation, decline, or fluctuation, so does not qualify as VU or EN under Criterion B as a whole.

We therefore infer that the taxon should be assigned the category Near Threatened (NT).

FIGURE 14. Extent of occurrence of Indopiptadenia oudhensis.

Uses:—The leaves are used for fodder and the stem as fuel and timber by local inhabitants of the border area of

both countries. The wood is very hard, strong and durable (Brandis 1874). Due to profuse and dense branching and

evergreen nature the plant may be grown in gardens or as avenue tree for shade.

Specimens Examined

INDIA. Uttar Pradesh: Balrampur Dist. (‘Gonda Dist.’): Pipra, 07 May 1900, Inayat s. n. (DD); Pipra, 07 May

1900, Inayat 23638 (DD); Hasanpur, 17 May 1900, Inayat 23635(a) & 23638(a) (DD, K photo); Sunghara, 14 May


1900, Inayat 23635(b) & 23638 (DD); Foot of Nepal Hills, 20 July 1911, R. S. Trouffs 3076 (DD); Barawa, 13

April 1916, Shees Ram 889 (DD); Sohelwa Wildlife Sanctuary: Jarwa forest, 6 km before Nepal border, Near

Nalah, 162 m, N 27º 40.365' E 82º 31.795', 06 November 2011, L. B. Chaudhary & R. Tiwari 263775 (LWG); 167

m, N 27º 40.378' E 82º 31.787', 06 November 2011, L. B. Chaudhary & R. Tiwari 263776 (LWG); 1 km before

Koylabas from Jarwa, 171 m, N 27º 40.814' E 82º 31.580', 10 May 2012, L. B. Chaudhary & O. Bajpai 252716 &

252717 (LWG); 156 m, N 27º 40.354' E 82º 31.810', 08 February 2013, O. Bajpai, A. K. Srivastava & A. Kumar

264426 (LWG); 176 m, N 27º 40.831' E 82º 31.577', 10 February 2013, O. Bajpai, A. K. Srivastava & A. Kumar

264432 (LWG); 1.5 km before Koylabas from Jarwa, 160 m, N 27º 40.362' E 82º 31.802', 19 March 2013, O.

Bajpai & L. B. Chaudhary 264439 (LWG); 1.5 km before Nepal from Tulsipur, inside the river, 176 m, N 27º

40.831' E 82º 31.577', 20 March 2013, O. Bajpai & L. B. Chaudhary 264442 (LWG); 1 km before Koylabas from

Jarwa, 160 m, N 27º 40.362' E 82º 31.802', 07 April 2013, O. Bajpai & L. B. Chaudhary 264481 & 264482 (LWG);

165 m, N 27º 40.371' E 82º 31.774', 01 May 2013, O. Bajpai & L. B. Chaudhary 264497 & 264498 (LWG); 165 m,

N 27º 40.371' E 82º 31.774', 30 July 2013, O. Bajpai, A. K. Srivastava & A. Kumar 263925, 263927 & 263928

(LWG); Bahraich Dist., Bhachkae, 20 May 1920, Sri Ram s. n. (DD); Saharanpur, Botanic Garden, 02 May 1914,

R. N. Parker 6758- from cultivated plant (DD). Uttarakhand: Kumaun: E. Kumaun, 1200 ft., May 1883, A. F.

Brown s. n. (DD); Sarda Valley, 12 miles above Tanakpur, R. H. Blutter 7235 (DD); Champawat Dist.: on way to

Purnagiri, 800 m., March 1990, K. S. Negi 303 (BSD); about 15 km away from Tanakpur towards Purnagiri, 343 m,

N 29º 07.545' E 80º 09.550', 18 August 2013, L. B. Chaudhary, O. Bajpai & A. Kumar 263952 (LWG); 406 m, N

29º 07.739' E 80º 09.375', 18 August 2013, L. B. Chaudhary, O. Bajpai & A. Kumar 263953 & 263954 (LWG); 5

km before Sukhi Dang from Tanakpur, 908 m, N 29º 08.514' E 80º 04.767', 18 August 2013, L. B. Chaudhary, O.

Bajpai & A. Kumar 263956 (LWG); Dehra Dun: New Forest, 23 May 1935, M. B. Raizada 69269- from cultivated

plant (DD); 23 May 1935, M. B. Raizada 69270- from cultivated plant (DD); 28 May 1937, M. B. Raizada 74553-

from cultivated plant (DD); May 1939, M. B. Raizada 80086- from cultivated plant (DD); FRI, Arboratum

compound 4, Gasthigarni 8110- from cultivated plant (DD).

NEPAL. Western Nepal: Jalesain nala, 29 April 1900, Inayat 23636 (DD); Sunpathri nala, 04 May 1900, Inayat

23637 (DD); Mandaura nala, 2 May 1900, G. H. Inayat 23637(a) (DD, K photo); Barahmdeo, Sarda Gorga, 2000

ft., 2 March 1914, J. H. Lyall 10208 (DD); Bhaunala to Garbha Doti Dist., 13 April 1929, Shees Ram 236 (DD);

Bardia Dist., Babai-Deurrali, 351 m, N 28º 20.934' E 81º 42.226', 23 January 2001, K. K. Shreshtha et al. 666

(TUCH); Kanchanpur Dist., 250 m, 9 August 1980, K. R. Rajbhandari, P.M. Regmi & K. J. Malla 5153 (KATH);

Dang Dist., 179 m, N 27º 41.219' E 82º 31.651', 31 July 2013, O. Bajpai, A. K. Srivastava & A. Kumar 263935

(LWG); Dang Dist., 186 m, N 27º 41.112' E 82º 31.622', 31 July 2013, O. Bajpai, A. K. Srivastava & A. Kumar

263936 (LWG).

Acknowledgements

The authors are thankful to Dr. C. S. Nautiyal, Director, CSIR-National Botanical Research Institute, Lucknow,

India for facilities and the Department of Science & Technology, Government of India, New Delhi for financial

support. We are also grateful to Dr. V. Sampath Kumar, IBLO, Herbarium, Roy. Bot. Gard. Kew, England, UK, Dr.

P. Lakshminarasimhan and Dr. Subir Bandopadhyay, Botanical Survey of India, Kolkata, India, Prof. Mohan

Siwakoti, Central Dept. of Botany, Tribhuvan University, Kathmandu, Nepal and Mr. Ganga D. Bhatt, National

Herbarium and Plant Laboratories, Lalitpur, Kathmandu, Nepal for providing necessary literature and information

on the species. The curators of herbaria mentioned in the work are also duly acknowledged for granting permission

for herbarium consultation. We thank the editor of the paper Ruth P. Clark, Herbarium, Royal Botanic Gardens,

Kew and unknown reviewers for the critical revision of the manuscript and useful comments and suggestions.

References

Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott B. (2011) Supporting Red List threat assessments with GeoCAT:

geospatial conservation assessment tool. In: Smith V, Penev L (Eds) e-Infrastructures for data publishing in biodiversity

science. ZooKeys 150: 117–126. (Version BETA). GeoCAT tool available from: http://geocat.kew.org/ (accessed 1 April

2014)


http://www.pensoft.net/J_FILES/1/articles/2109/2109-G-3-layout.pdf

http://www.pensoft.net/J_FILES/1/articles/2109/2109-G-3-layout.pdf

http://dx.doi.org/10.3897/zookeys.150.2109

Bajpai, O., Kumar, A., Mishra, A.K., Sahu, N., Pandey, J., Behera, S.K. & Chaudhary, L.B. (2012) Recongregation of Tree

Species of Katerniaghat Wildlife Sanctuary, Uttar Pradesh, India. Journal of Biodiversity and Environmental Sciences

2(12): 24–40.

Baker, J.G. (1879) Leguminosae. In: Hooker, J.D. ed. Flora of British India. L. Reeve & Co., London, U.K., 2: 289.

Bentham, G. (1840) Contribution towards a Flora of South America.- Enumeration of Plants collected by Mr Schomburgh in

British Guina. Journal of Botany 2: 127–146.

Biswas, S. & Chandra, S. (1997) Indopiptadenia oudhensis (Brandis) Brenan – An Endangered Tree Legume of Uttar Pradesh

and Nepal. Indian Forester 123(5): 419–421.

Brandis, D. (1874) The Forest Flora of North West & Central India. W.H. Allen and Co., London, U.K., pp. 168–169.

Brandis, D. (1906) Indian Trees. Archibald Constable & Co. Ltd., London, U.K., pp. 261.

Brenan, J.P.M. (1955) Notes on Mimosoideae. Kew Bulletin 10(2): 178–179.

http://dx.doi.org/10.2307/4108864

Bronn, H.G. (1822) De formis plantarum leguminosarum: primitivis et derivates. Heidelberg, 140 pp.

Catalogue of Life: 25th June 2013. Available from: <http://www.catalogueoflife.org/col/details/species/id/11483003>

(Accessed: 25 June 2013).

Champion, H.G. & Seth, S.K. (1968) A Revised Survey of the Forest Types of India. Publication Division, Govt. of India, New

Delhi, 404 pp.

Duthie, J.F. (1903) Flora of the Upper Gangetic Plain and of the Adjacent Siwalik and Sub-Himalayan tracts. Superintendent

of Government Printing, Calcutta, India, 1: 308.

http://dx.doi.org/10.5962/bhl.title.10981

Duthie, J.F. (1906) Piptadenia oudhensis Brandis. In King, G., Duthie, J.F. & Prain, D. eds. A second century of new and rare

Indian plants. Annals of Royal Botanical Garden Calcutta 9: 33, t. 43.

Goyal, A.K. (2009) Collection of Indopiptadenia oudhensis prohibited. Biodiversity News 1(1): 7.

Henry, A.N. & Roy, B. (1968) Nomenclatural Notes on Indian Flowering Plants. Bulletin of Botanical Survey of India 10: 275.

Herbarium Catalogue, Royal Botanic Gardens, Kew (2006). Published on the internet: http://apps.kew.org/herbcat/

gotoHomePage.do (Accessed: 25 September 2013).

Hutchinson, J. (1964) The genera of flowering plants. Clarendon Press, Oxford, U.K., 1: 154.

IUCN Standards and Petitions Subcommittee. 2013. Guidelines for Using the IUCN Red List Categories and Criteria. Version

10.1. Prepared by the Standards and Petitions Subcommittee. Downloadable from: http://www.iucnredlist.org/documents/

RedListGuidelines.pdf. (Accessed: on 29 January 2014).

Jussieu, A.H.L. (1830) Memoire sur Le Group des Meliacees. Mémoires du Muséum d'Histoire Naturelle 19: 153–304.

Kanjilal, P.C. (1933) A Forest Flora of Pilibhit, Oudh, Gorakhpur & Bundelkhand. Narendra Publishing House, Delhi, India,

pp. 161.

Kashyap, A.K. (2009) Hathipaula ki Talash. Biodiversity News 1(1): 15–16 (in Hindi).

Kumar, S. & Sane, P.V. (2003) Legumes of South Asia - A checklist. R.B.G., Kew, U.K., pp. 118.

Kurz, W.S. (1871) On Some New or Imperfectly Known Indian Plants. Journal of the Asiatic Society of Bengal. Natural

History 40(2): 45–78.

Lamarck, J. B. A. P. de Monnet de (1786) Encyclopédie Méthodique, Botanique 2. Paris, 774 pp.

Lewis, G.P. & Elias, T.S. (1981) Mimoseae. In: Polhill, R.M. & Raven, P.H. eds. Advances in Legume Systematics. Royal

Botanic Gardens, Kew, U.K., 1: 155–168.

Linneaus, C. (1753) Species Plantarum 1 & 2. Stockholm, 1200 pp.

Linnaeus, C. (1759) Systema Naturae, Ed. 10, 2. Stockholm, pp. 825-1384.

Linnaeus, C. (1771) Mantissa Plantarum 2. Stockholm, 587 pp.

Linnaeus, C. f. (1782) Supplementum Plantarum. Brunsvigae, 468 pp.

Luckow, M. (2005) Tribe Mimoseae. In: Lewis, G., Schrire, B., Mackinder, B. & Lock, M. eds. Legumes of the World. Royal

Botanic Gardens, Kew, U.K., pp. 163–183.

Müller, A.J. (1865) Euphorbiaceae. Linnaea 34(1): 1–224.

Ohashi, H. (1979) Mimosaceae. In: Hara, H., William, T.S. & William, L.H.J. eds. Enumeration of Flowering Plants of Nepal.

British Museum, U.K., 2: 123.

Panigrahi, G., Singh, A.N. & Misra, O.P. (1969) Contribution to the Botany of the Tarai Forests of the Bahraich District of Uttar

Pradesh. Bulletin of Botanical Survey of India 11(1&2): 89–114.

Planchon, J.E. (1848) Sur Les Ulmacees. Annales des Sciences Naturelles; Botanique (sér. 3) 10: 244–341.

Prakash, A., Rawat, K.K. & Verma, P.C. (2009) Indopiptadenia oudhensis (Brandis) Brenan: monotypic, endemic and highly

endangered taxa need conservation in Uttar Pradesh. Biodiversity News 1(1): 2–5.

Rana, T.S. & Ranade, S.A. (2009) The enigma of monotypic taxa and their taxonomic implications. Current Science 96 (2):

219–229.

Roxburgh, W. (1795) Plants of the coast of Coromandel :selected from drawings and descriptions presented to the hon. court of

directors of the East India Company, W. Bulmer and Co. for G. Nicol, Londan, 68 pp. + 100 pls.

http://dx.doi.org/10.5962/bhl.title.467

Roxburgh, W. (1819) Plants of the Coast of Coromandel 3. London, 96 pp. + 300 pls.


http://dx.doi.org/10.3897/zookeys.150.2109

http://www.catalogueoflife.org/col/details/species/id/11483003

http://apps.kew.org/herbcat/gotoHomePage.do

http://apps.kew.org/herbcat/gotoHomePage.do

Saini, D.C. (2005a) Flora of Bahraich District, Uttar Pradesh I-IV. Journal of Economic and Taxonomic Botany 29(3): 528–636.

Saini, D.C. (2005b) Flora of Bahraich District, Uttar Pradesh V-VI. Journal of Economic and Taxonomic Botany 29(4): 843–

920.

Sanjappa, M. (1992) Legumes of India. Bishen Singh Mahendra Pal Singh, Dehra Dun, pp. 66.

Singh, G. (2010) Indopiptadenia oudhensis: A plant of Sohelwa Wildlife Forest Division, Balrampur, Uttar Pradesh. National

Conference on Biodiversity, Development and Poverty Allevation. Uttar Pradesh Biodiversity Board, Lucknow, pp. 145–

146. <www.upsbdb.org/pdf/Souvenir2010/35.pdf>.

Singh, K.K. (1997) Flora of Dudhwa National Park. Bishen Singh Mahendra Pal Singh, Dehradun. India, 516 pp.

Smith, J.E. (1806) Exotic Botany : R. Taylor and Co., Londan, U.K., 2: 122.

Smith, J.E. (1810) Ficus. The cyclopædia; or, Universal dictionary of arts, sciences, and literature 14, London. Ficus no. 1–105.

Sprengel, C. (1817) Systema Vegetabilium 2. Stuttgardtiae, 964 pp.

Srivastava, T.N. (1976) Flora Gorakhpurensis. Today & Tomorrow's Printers and Publishers, Delhi, India, 411 pp.

Thiers, B. (2012) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Gardens’

Virtual Herbarium. <http://sweetgum.nybg.org/ih/>.

Willdenow, C.L. (1806) Species Plantarum 4 (2). Berlin, pp. 633–1157.


http://www.upsbdb.org/pdf/Souvenir2010/35.pdf

http://sweetgum.nybg.org/ih/

http://www.tropicos.org/Person/244

Indopiptadenia paper

Documents

Transcript of Indopiptadenia paper