Revisionary Insights Into The Genus Ledebouria (Asparagaceae) From India
Revisionary Insights Into The Genus Ledebouria (Asparagaceae) From India
Revisionary Insights Into The Genus Ledebouria (Asparagaceae) From India
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https://doi.org/10.11646/phytotaxa.641.1.1
Abstract
The genus Ledebouria Roth (Asparagaceae: Scilloideae) is currently known to have 3 species from India. The accepted name
for L. hyacinthina is L. revoluta. The two taxa have been segregated based on presence of bulblets, threads when leaves and
bulb scales are torn, number of inflorescence & presence of basal ovary lobes. Molecular evidence supports this segregation.
L. hyderabadensis and L. karnatakensis are reduced to variety & forma of L. hyacinthina respectively. One new variety and
a new forma of L. hyacinthina are also described here from Maharashtra.
Introduction
The genus Ledebouria Roth (1821: 195) belonging to the family Asparagaceae was described from India, based on a
single species Ledebouria hyacinthina Roth (1821: 195). The genus comprises of approximately 64 species distributed
predominantly in South Africa, South west Arabian Peninsula, Madagascar, India, and Sri Lanka (POWO, 2023).
Ledebouria is characterised by six lobed tepals, stamens fused at its base & stipitate ovary. Until the 21st century
L. hyacinthina was the only known species from India. In 1995, Stedje & Thulin, for the first time, synonymized
L. hyacinthina under L. revoluta (L.f.) Jessop (1970: 255). At present, Indian Ledebouria is represented by three
species (Deshmukh et al. 2022), namely, Ledebouria revoluta (L.f.) Jessop, Ledebouria karnatakensis Punekar &
Lakshminarasimhan (2011: 500) and Ledebouria hyderabadensis Ramana et al. (2012: 561). Ledebouria hyacinthina
shows phenotypic plasticity especially in characters such as leaf shape, size, structure, coloration, inflorescence and
number of flowers, making species identification perplexed. Literature and herbarium (Virtual: K, BM, PRE, MNHN,
UPS) studies of African L. revoluta and field studies of L. hyacinthina felt authors the necessity to revisit whether L.
hyacinthina and L. revotuta are synonymous and ascertain the taxonomic status of the recently described species of
Ledebouria from India. Further, during our studies on Indian Ledebouria we also came across two different populations
of Ledebouria with a unique set of characters. These populations are described here as new infraspecific taxa from
India.
Morphological study
Field studies were conducted to collect Ledebouria samples during the flowering season from May–August (2020–
2023) across Peninsular India (Table 1 & Figure 1). Approximately 150 individuals were observed in the field.
Total genomic DNA was extracted from fresh leaves using the NucleoSpin Plant II kit (Macherey-Nagel, Düren,
Germany). The extracted DNA was evaluatedfor its quality and quantity using an EPOCH MicroplateSpectrophotometer
(BioTEK, Winooski, Vermont, U.S.A.).Poor-quality DNA extracts were further purified using the NucleoSpin g-DNA
clean-up XS kit (Macherey-Nagel). DNAamplification and sequencing of the trn L-trn F and trnC -ycf6 region was
performed using the primer trnL-trnF (Dangi, 2015) and trnC -ycf6 (pfosser, et al. 2012) Nr DNA (Internal Transcribed
the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press •
Spacer) sequences were not amplifiable and comparative sequences are absent on NCBI nucleotide database (Ali
et al. 2012). On the contrary, cp DNA (trn L-F) regions have been shown to be useful at lower taxonomic levels
(Stedge 1998, Bohle et al. 1994, Fangan et al. 1994) and trn L-F region was also widely used by various author
for phylogenetic analysis (Manning et al. 2004, Manning et al. 2009, Pfosser & Speta 1999) another maker trn C-
ycf6 used in study of Hyacinthaceae relationships (pfosser, et al. 2012). Therefore, for the purpose of the present
phylogenetic analysis cp DNA (trnL-F and trnC-ycf6) were selected over nr DNA. Sequences were assembled and
subjected to multiple sequence alignment using MUSCLE (Edgar, 2004) implemented in MEGA X v.10.1.8 (Kumar
et al. 2018) and revised using gBlock online software. Phylogenetic analysis was conducted independently for both
regions. Phylogenetic analyses were performed using RAxML v.8.1.18 (Stamatakis et al. 2008; Stamatakis, 2014) and
MrBayes v.3.3.6-svn(r1040) x64 (Ronquist et al. 2012) on the CIPRES Science Gateway v.3.3 (Miller et al. 2010).
RAxML was run under the GTR-GAMMA model with 20 heuristic searches from distinct random stepwise addition
sequence parsimony starting trees, followed by selecting the best-scoring tree. RAxML bootstrap (BS) values were
calculated using 1000 replicates. We used Tracer v.1.6.0 (Rambaut et al. 2014) to check for convergence of the chains.
Specifically, we visually inspected the Tracer plots and further checked that all ESS values were above 200. Each DNA
region was analysed separately first, then all plastid regions combined (partitioned in coding vs. non-coding regions).
Next, we compared all well-supported clades (here defined as ≥ 75 % RAxML BS) in the plastid.
Taxonomy:—Ledebouria was described from India, with a single species Ledebouria hyacinthina Roth. The genus
Scilla Linnaeus (1753: 308) was described by Linnaeus which was identified by bulb scales without fibres, tepals erect,
blue-pink rarely white and ovary oblong. Baker (1870) in his revision of the genus Scilla treated Ledebouria within
Scilla as a section and considered L. hyacinthina as synonym of Scilla indica (Wight 1853: 2041) Baker (1870: 12)
(currently nom. illeg. POWO, 2023). These two species, namely S. indica and S. revoluta Baker (1870:6) differ in
soboliferous roundish bulb, scape 2 to 3 times as long as scape and stipitate ovary with discoid dilated base (in S.
revoluta Baker) against, ovoid, not soboliferous bulb, scape as long as leaves & sessile, deeply blunt, 3 lobed ovaries
without any dilation (S. indica Baker). Baker’s (1870) choice was followed by Hooker (1892), Cooke (1907).
Baker re-revised the genus Scilla in 1896 (Baker 1896). It is known that “most of the species he recognized were
known from single specimens or illustration. He never visited South Africa and the living plants used by him were
grown in greenhouses in Great Britain” (Venter 1993).
Jessop (1970), in his revision of bulbous Liliaceae concentrated on the generic concept of Scilla, Ledebouria, and
Drimiopsis. and re-established Ledebouria (in family Hyacinthaceae) on the basis of the presence of stipitate ovary. He
made a new combination L. revoluta (L.f. 1782:204) Jessop (1970:255) based on Hyacinthus revolutus L.f. (1782:204)
(Type: “Cap. Bonaespei”, Thunb. s.n., UPS, holo, Microfiche no. 8508!). However, Jessop did not synonymize Indian
L. hyacinthina with South African L. revoluta.
Macbride (1918) made a new combination S. hyacinthina (Roth) Macbride (1918:14), based on L. hyacinthina
and synonymised S. indica Baker. Deb & Dasgupta (1981) adopted the genus Scilla (S. hyacinthina (Roth) Macbride,
basionym L. hyacinthina Roth) in their revisionary work. Venter (1993) in his revision of South African Ledebouria
from a multidisciplinary approach including morphology, cytology & anatomy has not considered L. hyacinthina as
a synonym of L. revoluta. In 1995, Stedje & Thulin, for the first time, synonymized L. hyacinthina under L. revoluta.
This concept was followed by Venter (2008) in his synopsis of the genus Ledebouria in South Africa (2008:143) and
synonymised L. hyacinthina and L. revoluta. In India, several taxonomists used Scilla instead of Ledebouria after
Baker’s publication (Hooker 1892, Wight 1903, David 1903, Cooke 1907, Macbride 1955, Deb & Dasgupta
1975, 1981, Sharma et al. 1996, Shetty & Singh 1991, Karthikeyan & Kumar 1993). Almeida (2009) used the name
Ledebouria hyacinthina for the Indian element. From 2013 onwards, Ramana et al. (2012) started adopting Ledebouria
revoluta as accepted name for the Indian species, Ledebouria hyacinthina (Table 2.).
Morphology:—Ledebouria hyacinthina (India) is known by solitary plants; bulb hypogeal, dead bulb scales
compactly arranged, without threads when torn, outer bulb scales brown, membranous; inner bulb scales whitish;
inflorescence 1–3 per bulb, flowers up to 60 (rarely 100) per inflorescence, occasionally ca 20 abortive flowers at base
of the inflorescence; ovary tricarpellary, syncarpous, superior, stipe ca 1mm long, each carpel bilobed, ovary conical
to ellipsoid; capsule globose, fruits 1–2(3). A comparative account differentiating Indian Ledebouria (L. hyacinthina)
from S. African Ledebouria (L. revoluta) is provided in Table 2.
Phenology:—Flowering period is dependent on arrival of monsoon. Ledebouria revoluta (South African regions)
flowers from August to November and in India it starts by the end of May; later fruiting continues till end of July or
beginning of August in some areas. In some areas, bulbil forms at the tip of the leaf which bends down to the soil and
produce roots and new plant.
Phylogeny:—The trnC-ycf6 dataset comprised 107 accessions with a total length of 1155 base pairs aligned of
which 786 were conserved, 369 were variable and 272 were parsimony informative. The trnL-F dataset comprised
87 accessions with a total length of 1124 base pairs aligned; of which 587 were conserved, 632 were variable and 380
were parsimony informative. Both the cp DNA resolved all Indian accessions as a separate clade. The newly added
Ledebouria sequences were grouped into the clade having other Indian accessions of Ledebouria (supplementary Table
4, 5.). Good branch node support 100 % segregates the Indian Ledebouria from all the African species of Ledebouria.
It is important to note that all the other accessions have out grouped themselves and formed clades separated from the
Indian clade. Internally, the phylogenetic relationship within the Indian clade was poorly resolved, figure 2 & 3 &
some resolved nodes had weak support.
Description:—Bulbous herb, 10–25 cm long, solitary, bulb hypogeal, dead bulb scales compactly arranged, without
threads when torn, outer scales brown, membranous, inner fleshy scales whitish. Foliage leaves synanthous, 3–6 per
bulb, 20–30 (45) × 1–4 cm, rosulate, erect, spreading or flapping, sessile or pseudo-petiolate, lanceolate to ovate,
canaliculate, fleshy, mucilaginous, without threads when torn, dark green to purple blotches, base cuneate, apex acute
to obtuse, green occasionally spotted with purple, margins entire, green or transparent; leaf tip bulbils present or absent.
Inflorescence 1–3 per bulb, longer than leaves, erect to flaccid. Flowers more than 25 per raceme, protandrous, tepals 6,
campanulate, 5–6 × 5–6 mm, oblong, arranged in two whorls of three each, white to purple with green recurved, equal,
cucullate. abortive flowers may be observed at the base of raceme ca 20 in number. Bracts 2, well developed, glabrous,
membranous, one linear 1–1.5 mm long and other hyaline 1.5–2 mm long, lanceolate to ovate, mostly acuminate at apex
or bifid, hyaline and turn green as the flowers mature, persistent. Pedicel spreading, 4–12 mm long, maroon, Stamens
6, erect, filament as long as style, uniseriate, white at base and purple towards apex, inserted at throat of the perianth
the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press •
FIGURE 2. Best Maximum likelihood tree retrieved after the analysis of cpDNA (trnC-ycf6) sequenced 107 samples bootstrap value >75
% indicated along with branches, Names are to right of the tree.
the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press •
tube, Filament perianth tube fused epi-tepalous, base slightly swollen. Anther 1–3 mm long, pale yellow at maturity,
dehiscence longitudinal. Ovary tricarpellary, 2 ovules per locule, syncarpous, superior, stipitate, stipe ca 1mm long,
each carpel bilobed, ovary conical to ellipsoid (in lateral view), basal lobes absent. Style triangular in transverse
viewglabrous, purple. Stigma below the level of anther lobes in young flowers, gradually projecting upwards. Capsule
globose, Fruits 1–2 (3). Seed globose to ovate, surface wrinkled, brown.
Specimen examined:—INDIA. Goa: Colem, 19 June 2003, Datar, (AHMA!). Maharashtra: Pune, Bhor hill, 29
July 1957, Vartak, 9550 (AHMA!); Sinhagad, 20 July 1958, Vartak, 13480, 13481 (AHMA!); Junnar, Hirve, 07 June
2001, Nagarkar 19916, 19017 (AHMA!); 05 July 2003, Nagarkar, 22745 (AHMA!); Vetal hill, Law college, 16
June 1988, Joshi, VH538, VH538a (AHMA!); Vetal hill, 18 July 1955, Vartak, 751 (AHMA!); Mulashi, Sutarwadi,
28 June1984, Upadhye, MP039 (AHMA!); Pune University, 15 June1973, Datar. 8073, 8074, 8075, 8076, 8077
(AHMA!); 12 June 1973, Datar, 8070, 8071, 8072 (AHMA!); 28 August 1980, Gaunjadkar , 16566 (AHMA!);
Parvati, 25 June1981, Kulkarni, 431 (AHMA!); Katraj hill, 15 June 1956, Vartak, 4549 (AHMA!); Shivneri hills, 25
July 1963,Ansari, 88681(BSI!); AlandI, Chakan, 9 June1963, Rao, 85287 (BSI!); Nawargaon forest, 21 August1973,
Wadhwa, 137447a (BSI!); Shivneri fort, 24 June1964, Hemadri, 94322 (BSI!); Purandhar Road, 19 July 1963, Seshadri,
86561 (BSI!); Golf club, Yerwada, 28 July 1962, Rao, 39529 (BSI!); Shivneri hill, 17 July 1965, Ansari, 88792,
(BSI!); Alandi, 28 June 1966, Moorthy, 68572 (BSI!); Katraj, 16 June1956, Puri, 2354 (BSI!); Haveli, 16
July1964, Ansari, 97512 (BSI!); Khanapur, 11 August1964, Ansari, 97596 (BSI!); Dive Ghat, 04 July 2016,
Nandikar, 1409 (NGCPR!); 15 July 2017, Giranje, 1429 (NGCPR!); Inglun Ghat, 12 June 2018, Giranje, 1434
(NGCPR!), Junnar, 23 June 2014, Punekar, 1401 (NGCPR!); Inglun Ghat, 22 June 2017, Nandikar, 1418 (NGCPR!),
06-2015, Nandikar, 1403 (NGCPR!), Ahmadnagar, Sangamner, Dalasane, 23 June 2017, Nandikar et al., 1419
(NGCPR!); Amaravati, Belkund, 28 June 1977, Ansari, 149319 (BSI!); Umarkhed, Amaravati, 18 June 1977, Ansari,
146802 (BSI!), Dhami, 2 July 1977, Ansari, 149359 (BSI!); Burmi forest, 19 June1977, Ansari, 146816 (BSI!); Bairat,
22 June 1977, Ansari, 146885 (BSI!); Toribanda, 27 August 1976, Ansari, 144097 (BSI!); Burmi forest, 19 June
1977, Ansari, 146859 (BSI!); Silwassa, 28 June 1964, Ansari, 98505 (BSI!); Chalisgaon, 14 July1957, Mahajan,
20632 (BSI!); Nashik, Igatpuri, 8 May1962, Rane, 81331 (BSI!); Khandala, 30-06-1956, Puri, (BSI!); Akola, Adan
Dam Site, 26 June 1977, Kamble, 153869 (BSI!); Narnala fort, 24 June 1978, Kamble, 153759 (BSI!); Mahan, 22
June 1978, Kamble, 153725, 153726 (BSI!); Satara, Patan, Borirange, 8 August 1979, Karthikeyan, 160076 (BSI!);
Bhade Village, Veer Dam, 15 June 2017, Giranje et al. 1417 (NGCPR!); Kas Plateau, 12 September 2015, Nandikar,
1404 (NGCPR!); 22 June 2017, Nandikar, 1418 (NGCPR!); Pandavdara, 01 July 2017, Giranje, 1424 (NGCPR!);
Shirwal, Chheda Company, 27 June 2017, Balakrishna , 1423 (NGCPR!); Shirwal, Miraj, 15 June 2017, Giranje
et al., 1416 (NGCPR!); Palashi, 21 June 2016, Nandikar, 1406 (NGCPR!); Kas, June 1991, Bachulkar, 5137 (SUK!);
Suttagatti, 23 June 2007, Chandore, ANC320 (SUK!); Buldhana: Wasali forest, 30 June 2011, Karpure, 1256
(28253); Solapur: Gangewadi, 28 July 2011, Jayanthi, 199217 (BSI!); Kolhapur, Kandalgaon, 22 November 2002,
Das, 183895 (BSI!); Raigad, Kankeshwar, 14 June 1978, Sane, K-699 (AHMA!); Ratnagiri, Guhaghar, 11 June 2018,
Giranje, 1433 (NGCPR!); Jaitapur, 27 May 2022, Suchandra et al, SHK 46 (RDNCP!); Raigad, Kankeshwar Temple,
15 June 2017, Giranje, 1420 (NGCPR!); Kankeshwar Temple, 2 July 2023, Suchandra et al. SHK 48 (RDNCP!)
Mapgaon, 25 June 2017, Giranje, 1420 (NGCPR!); Gadchiroli, Armori, 15 June 2015, Kahalkar, 1421 (NGCPR!);
Satara, Kas Pathar, 30 July 2021, Suchandra et al, SHK26(RDNCP!); Kolhapur, Ajra , 15 July 2022, Suchandra et al.
SHK24 (RDNCP!). Karnataka: Belgaon, 8 July 2010, Chavan, 2042 (AHMA!); Mysore, 11 August1963, Raghavan,
90012 (BSI!); Birnali, 10 August1976, Singh, 142828 (BSI!); North Kanara, 5 June 1885, Talbot, (BSI!); North Kanara
10 July 1889, Talbot (BSI!); Dodda malki, 04 June 2014, Jayanthi, 195758 (BSI!); Dharwad, 20 May1904, Bhide,
16696 (BSI!); Belgaon, Suttagatti, 01 June 2017, Giranje, 1414 (NGCPR!), Gokak, 16 July 2020, Nandikar, 1410,
(NGCPR0!); Khanapur, 24 June 2016, Girnanje et al. 1407 (NGCPR!); Bagalkot, Badami, 17 July 2016, Nandikar,
1411, 1431 (NGCPR!); Bagalkot, Badami, 23 July 2023, Suchandra et al. SHK21 (RDNCP!). Andra Pradesh:
Shesachalam, 19 August 2018, Malpure, 1441 (NGCPR!); Palakona, 3 September 2014, Prasanna, 004546 (BSID!);
Thumallaballu-Rajampet, 01 September 2014, Prasanna et al. 004507 (BSID!); East Godavari, Machalipatanam, 14
September 1980, Ramamurthy, 88870 (BSID!); Prakasham, Chinnaganjam, 25 June 1988, Ravikumar, 88122 (BSID!);
Kadapa, Mabbuchintala palli, 19 August 2009, Pajaikullaiswamy, 37682 (BSID!); Adilabad, Kanchanpalli, 28
June 1987, Ravishankar, 85223 (BSID!); Mehabubnagar, 26 June 1987, Husagoulai, 84957 (BSID!). Tamilnadu:
Kunnathur lake, 20 August 2017, Kishor , 1430 (NGCPR!); Annamalai Tiger Reserve, 23 August 2016, Nandikar, 1413
(NGCPR!).
1. Leaf narrowly lanceolate; apex acute with no bulbil; inflorescence 1–3 per plant; ovary with downward projected beak in each
carpel........................................................................................................................... Ledebouria hyacinthina var. hyderabadensis
1. Leaf lanceolate to spathulate; apex acute or obtuse; inflorescence upto 1–2 bulbil may or may not be present; ovary without beak
...................................................................................................................................................................................................................2
2. Leaves lanceolate, oblanceolate or elliptic; apex acute (up to 700); bulbil may be present or not........................................................
.......................................................................................................................................... Ledebouria hyacinthina var. hyacinthina
2. Leaves spathulate; apex obtuse (between70–1300); bulbil absent............................................Ledebouria hyacinthina var. obtusata
Ledebouria hyacinthina var. hyderabadensis (M.V Ramana, Prasanna & Venu) S.R.Dutta, Chakral & H. Rodrigues
var. et. stat. nov. ≡ Ledebouria hyderabadensis M.V Ramana, Prasanna & Venu (2012: 561), basionym. (Figure 6.)
Type:—INDIA. Telangana, Hyderabad, Damaiguda hills, 27 July 2009, P.V Prasanna & V.N Rao, 1459, (Holotype: CAL0000024579!;
Isotype: BSID0002007!)
Description:—Bulb 1.5–2 cm long; leaves 5–6, 10–15 × 0.5–1 cm, linear-lanceolate, apex acute, apex angle < 90,
blotches absent, bulbil absent, margin straight; inflorescence 2–3, erect or flaccid, bearing ca 25 flowers, absence of
sterile flower, ovary top shaped with beak at base.
Distribution:—India, Telangana, Hyderabad, Damaiguda hill, Osmania university.
Habitat:—Growing on boulders where sandy soil is accumulated.
Phenology:—July to August.
Associates:—Cyperus meeboldii Kükenthal (1922:345), Ophioglossum costatum Robert Brown (1810: 163), Striga
asiatica (Linnaeus 1753:630) Kuntze (1891:466)
Specimens examined:—INDIA. Telangana, Hyderabad, Rajendranagar hill, 21 July 2016, P.T Giranje, 1412
(NGCPR00147!, NGCPR00148!), 15 June 2018, P.T Giranje, 1435 (NGCPR00149!, NGCPR00150!).Damaiguda
hill, 10 July 2022, Suchandra et al, SHK31 (RDNCP!).
Notes:—Ledebouria hyderabadensis was described on the basis of number of flowers (25 per inflorescence),
shape of the leaf, absence of bulbil and blotches on the leaf. However, our phylogenetic & morphological work reveals
that this taxon is nested within the Ledebouria hyacinthina clade along with other members of Ledebouria from India
(Figure 2, 3). Beaked carpel a unique character only observed in this taxon can be used to relegate its status as a variety
of L. hyacinthina.
Ledebouria hyacinthina var. obtusata S. Dutta, Chakral & H. Rodrigues var. nov. (Figure 5.)
Type:—INDIA. Maharashtra, Palghar, Takmak, SHK 15 (Holotype: SHK0015!; Isotype: BLAT SHK0016, RDNCP SHK0017!).
Description:—Bulb 2–4 × 1.5–3 cm, leaves broadly lanceolate—spathulate, apex obtuse (70o–130o) without bulbil;
inflorescence bearing more than 40 (100) flowers; flowers pinkish in colour; ovary obtusely deltate without beak at
the base.
Distribution:—Maharashtra: Manori (Mumbai), Takmak, Tungareshwar (Palghar), Uttan, Mama-bhanja
(Thane) Masai (Kolhapur).
Habitat:—From low level Basalt mesa to high altitude lateritic plateau of the western ghats, rocky plateau, forest
floor, sandy soil near the sea and cultivated areas.
Phenology:—Flowering and fruiting from June to August.
Etymology:—The epithet ‘obtusata’ refers to the obtuse apex of the matured leaf.
Associates:—Senna tora (Linnaeus 1753: 376) Roxb. (1832:340), Dipcadi saxorum Blatter (1928: 736),
Mesosphaerum suaveolens (L. 1759:1100) Kuntze (1891:525), Lepidagathis Willdenow. (1800: 400).
Specimen examined:—INDIA. Maharashtra: Mumbai, National Park, Borivali, 27 June 1953 Fernades, R1303
(BLAT!); Andheri, Mahakali Caves, Salsette, 07 July 1945, Santapau, 6767 (BLAT!); Malad Quary Hills, 17 June 1955,
Shah, Shah4538 (BLAT); Malad, Marve Road, 03 July 1955, Shah, Shah4589 (BLAT); Vihar Lake, 5 July 1958,
Merchant, 589 (BLAT); Mahakali Caves, Andheri, 13 June 1951, Santapau, 12823 (BLAT); Kanheri hill top, 14
August 2011, Shinde, 28078 (AHMA!); Manori, 15 July 2023, Suchandra et al. SHK35 (RDNCP!); Pune District,
14 July 1955, Vartak, 782 (AHMA!); Pune University, 15 June 1973, Datar, M.A.C.S 8077 (AHMA); Dive ghat, 15
July 2017, Giranje, 1429 (NGCPC!); 1426 (NGCPR181!); Mulshi, Rawalewadi, 04 July 2017, Giranje et al., 1426
(NGCPR!); Bovdhan hill, 22 July 1954, J.A.V, 5069 (BSI!); Chattarshungi hills, Pune, 5 June 1917, Ezekiel 30576
(BLAT!); Chattarshungi hills, Pune, 28 June 1954 , Rukmini, 153 B.R (BLAT!); Chattarshungi hills, Pune, 28 June
the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press •
FIGURE 4. L. hyacinthina f. recurvata S. Dutta, Chakral & H. Rodrigues: A Habit, B Leaf, C Bulbil, D Inflorescence, E Tepal, F & G
Flower, H Stamen, I & J Pistil, K T.S of ovary, L Fruit, M T.S. of leaf.
the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press • 11
FIGURE 6. Ledebouria hyacinthina var. hyderabadensis S. Dutta, Chakral & H. Rodrigues: A Habit, B Leaf, C Inflorescence, D Tepal, E
& F Flower, G Stamen, H Pistil, I T.S of ovary, J Fruit, K seed.
1. No blotch on leaf; leaf margin much recurved; epidermal cell margin straight; number of bulbils per leaf goes up to 5–6..................
.................................................................................................................................................................Ledebouria hyacinthina f. recurvata
1. Purple blotch on young leaf; leaf margin slightly bent downwards; epidermal cell margin wavy; number of bulbil per leaf is 1......
............................................................................................................................................Ledebouria hyacinthina f. karnatakensis
Ledebouria hyacinthina f. karnatakensis (Punekar & Lakshmin.) S. Dutta, K. Chakral & H. Rodrigues forma et. stat.
nov. ≡ Ledebouria karnatakensis Punekar & Lakshminarasimhan (2011: 500), basionym.
Type:—INDIA. Karnataka, Uttar Kanada, Anshi National Park, Ulavi Road, 6 June 2006, Sachin A Punekar 5168 (Holotype: CAL!)
Description:—Bulb 2–4 × 1.5–3 cm; leaves 2–4, 15–20 × 1–2.5 cm, lanceolate, apex acute, apex angle < 90, blotches
present during the time of flowering, bulbil present, margin straight or slightly bend; inflorescence 1–2, erect or
flaccid, bearing ca 40 flowers, sterile flower present at the base; ovary ellipsoid without beak at base.
Distribution:—Amboli road, along the roadside Ulavi-Kumbharwada road.
Habitat:—Growing on the open basalt rocky outcrops as well as on soil under the shades of trees.
Phenology:—May to July.
Associates:—Senna tora (Linnaeus 1753: 376) Roxburgh (1832:340), Cyperus nemoralis Chermezon (1921:
553) and species of Cyperus Linnaeus (1753: 44)
Specimens examined:—INDIA. Karnataka: Uttar Kanada, 17 June 2018, Nandikar, 1437, 1438 (NGCPR!);
Anshi, 17-06-2016, Nandikar, 1437 (NGCPR!). 6 June 2023, Suchandra et al., SHK 24 (RDNCP!).
Notes:—Ledebouria karnatakensis was described on the basis of presence of blotches, bulbil and number
of flowers. However, after a detailed morphological study, we conclude that it resembles Ledebouria hyacinthina
population collected from Kankavli, Maharashtra (Table 3.) Phylogenetically, L. karnatakensis is nested within L.
hyacinthina. Thus, we reduce the status to a variety level.
the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press • 13
TABLE 4 (supplementary). Gene bank voucher number, locality & accession of trnL-trnF cpDNA region.
No Accession no Taxon Locality
1 Z99148 Ledebouria urceolata Ethopia
2 AM909711 Ledebouria nossibeensis Madagascar
3 MK644154 Ledebouria revoluta India
4 Z99145 Ledebouria revoluta India
5 OQ850332 Ledebouria revoluta India
6 OQ850329 Ledebouria sp India
7 AJ507944 Ledebouria hyacinthina India
8 AJ507945 Ledebouria hyacinthina India
9 OQ850330 Ledebouria sp India
10 OQ850331 Ledebouria revoluta India
11 JX090555 Ledebouria revoluta Unverified, INDIA
12 JX090556 Ledebouria socialis South Africa
13 X74576 Ledebouria revoluta Africa
14 AJ232501 Ledebouria socialis Cultivated in Vienna
15 AJ507938 Ledebouria sp India
16 AJ507939 Ledebouria sp. India
17 AJ507937 Ledebouria floribunda Ethopia
18 Z99146 Ledebouria revoluta Africa
19 AJ507941 Ledebouria sp INDIA
20 AJ507946 Ledebouria concolor South Africa
21 AJ507951 Ledebouria socialis South Africa
22 DQ313330 Resnova lachenalioides South Africa
23 DQ313331 Resnova maxima South Africa
24 AJ507943 Resnova maxima South Africa
25 AJ507942 Resnova humifusa South Africa
26 DQ313329 Resnova sp South Africa
27 AJ508004 Schnarfia messeniaca Cultivated in Vienna
28 OP096330 Muscari macrocarpum Turkey
29 OP096365 Bellevalia speciosa Turkey
30 OP096364 Brimeura amethystina Turkey
31 FJ423214 Brimeura amethystina RBGE Cultivated
32 OP096344 Scilla monophyllos Portugal
33 AJ508000 Tractema monophyllos Portugal
34 FJ423219 Tractema monophyllos Portugal
35 AJ508001 Autonoe haemorrhoidalis Canary Island
36 FJ423213 Autonoe madeirensis Portugal
37 FJ423212 Autonoe latifolia Portugal
38 FJ423238 Hyacinthoides mauritanica Portugal
39 FJ423241 Hyacinthoides mauritanica Morocco Atlas
40 FJ423234 Hyacinthoides reverchonii Spain
41 FJ423242 Scilla flahaultiana Morocco Atlas
42 FJ423255 Hyacinthoides hispanica Spain
43 FJ423251 Hyacinthoides hispanica Portugal
44 NC046498 Hyacinthoides non-scripta north west Portugal, Europe
45 FJ423245 Hyacinthoides non-scripta Spain
46 FJ423250 Hyacinthoides hispanica Portugal
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the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press • 15
TABLE 5 (supplementary). Gene bank voucher number, locality & accession of trnL-ycf6 cpDNA region.
No Accession no Taxon Locality
1 MT954823 Squilla pancration Italy
2 MT954824 Squilla pancration Malta: Gozo
3 MT954833 Squilla undulata Spain: Valencia Province
4 MT954821 Squilla palaestina Israel: Nahal Adorayim
5 MT954829 Squilla serotina Morocco: Larache
6 MT954814 Squilla aff. hesperia Morocco: Dchar Abiad
7 MT954828 Squilla secundiflora “Morocco:
8 MT954819 Squilla maritima Spain: Madrid Province
9 MT954812 Squilla aff. hesperia “Morocco:
10 MT954817 Squilla aff. hesperia “Morocco: NE Larache
11 MT954813 Squilla aff. hesperia “Morocco: Tanger, Cabo Espartel”
12 MT954816 Squilla aff. hesperia “Morocco: Larache”
13 MT954818 Squilla aff. hesperia “Morocco: NE Larache”
14 MT954820 Squilla numidica Spain: Ibiza, close to Portinatx
15 HE575035 Rhadamanthopsis sp. SOUTH AFRICA
16 MT954754 Rhadamanthopsis monophyllus south Africa: Eastern Cape
17 HE575036 Rhadamanthopsis karooicus SOUTH AFRICA
18 MT954749 Rhadamanthopsis hyacinthoides South Africa: Eastern Cape
19 MT954750 Rhadamanthopsis hyacinthoides South Africa: Eastern Cape,
20 MT954748 Rhadamanthopsis haworthioides South Africa: Western Cape
21 HE575034 Rhadamanthopsis sp SOUTH AFRICA
22 MT954747 Rhadamanthopsis haworthioides South Africa: Eastern Cape
23 MT954755 Rhadamanthopsis monophyllus South Africa: Eastern Cape
24 MT954633 Aulostemon mzimvubuensis South Africa: Eastern Cape
25 HE575037 Rhadamanthopsis karooicus SOUTH AFRICA
26 HE575065 Litanthus pusillus SOUTH AFRICA
27 HE575066 Litanthus pusillus SOUTH AFRICA
28 MT954743 Litanthus pusillus Namibia: Pockenbank,
29 HE575064 Litanthus pusillus SOUTH AFRICA
30 MT954745 Litanthus stenocarpus South Africa: Western Cape
31 MT954901 Zulusiadelagoensis South Africa: KwaZulu-Natal
32 MT954902 Zulusia delagoensis South Africa
33 MT954903 Zulusia edwardsii South Africa: KwaZulu-Natal
34 HE575062 Drimia haworthioides South Africa
35 MT954679 Drimia haworthioides South Africa: Eastern Cape,
36 MT954676 Drimia cf. elata South Africa
37 MT954675 Drimia cf. elata South Africa: Eastern Cape
38 MT954678 Drimia haworthioides South Africa: Eastern Cape
39 MT954688 Drimia sphaerocephala South Africa: Eastern Cape,
40 MT954685 Drimia sp. South Africa: Western Cape
41 HE575063 Drimia elata South Africa
42 MT954677 Drimia elata “South Africa: Darling,
43 MT954681 Drimia cf. elata South Africa: Northern Cape
44 MT954686 Drimia sp. South Africa: Northern Cape
45 MT954757 Rhadamanthopsis namibensis Namibia: Karas
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the genus Ledebouria from India Phytotaxa 641 (1) © 2024 Magnolia Press • 17
TABLE 5 (Continued)
No Accession no Taxon Locality
90 MT954794 Sagittanthera cyanelloides South Africa: Eastern Cape
91 MT954738 Iosanthus khubusensis South Africa: Northern Cape
92 MT954739 Iosanthus khubusensis South Africa: Northern Cape
93 MT954741 Iosanthus toxicarius Namibia: Pockenbank,
94 MT954740 Iosanthus sp Namibia: Windhoek
95 HE575017 Bowiea volubilis South Africa
96 MT954674 Bowiea volubilis Ex Cult. Botanical Garden Vienna
97 MT954671 Bowiea gariepensis South Africa: Northern Cape
98 MT954673 Bowiea kilimandscharica Kenya: Massai Mara
99 HE574952 Ledebouria sp. Madagascar
100 FJ269110 Autonoe madeirensis Portugal: Madeira”
101 FJ269109 Autonoe latifolia Morocco: Anti-Atlas
102 submitted to genebank Ledebouria hyacinthina India-Manori
103 submitted to genebank Ledebouria hyacinthina India-Manori-2
104 submitted to genebank Ledebouria hyacinthina India-Anshi
105 submitted to genebank Ledebouria hyacinthina India-Badami
106 submitted to genebank Ledebouria hyacinthina India-Hyderabad
107 submitted to genebank Ledebouria hyacinthina India Kankavali
Ledebouria hyacinthina f. recurvata S. Dutta, K. Chakral & H. Rodrigues f. nov. (Figure 4.)
Type:—INDIA. Maharashtra: S. Dutta, H. Rodrigues & K. Chakral, 23 May 2022, SHK06 (Holotype CAL!, Isotype: RDNCP!)
Description:—Bulb 5–6 × 3–4 cm; Leaves elliptic, margin recurved, apex acute, with outline of epidermal cell straight,
single bulbil develops per leaf but reaches up to 5-6 when leaf tears longitudinally; inflorescence 1–2, erect and flaccid,
bearing 40-60 flowers; flowers pinkish in colour; sterile flowers up to 20 at the base of raceme; ovary transversely
elliptic; Seed brown when formed & reach maturity, at times seed does not mature but withers.
Distribution:—India, Maharashtra, Sindhudurg, Kankavali, Brahamwadi.
Habitat:—Senna tora (Linnaeus 1753: 376) Roxburgh (1832:340), Neanotis lancifolia
W.H.Lewis (1966: 39), Smithia sensitiva Aiton (1789: 496) and species of Lepidagathis.
Phenology:—Late May to July.
Etymology:—The epithet ‘recurvata’ is derived from the plants recurved leaf margin observed in the transverse
section of leaf.
Associated plants:—Senna tora (Linnaeus 1753: 376) Roxburgh (1832:340), Rhamphicarpa fistulosa
(Hochstetter 1841: 374) Bentham 1846: 504, Neanotis lancifolia (Hooker f. 1880: 73) W.H.Lewis 1966: 39, Alysicarpus
vaginalis (Linnaeus 1753: 746) de Candolle 1825: 353 and species from genera, Rotala Linnaeus (1771: 143).
Specimen examined:—INDIA. Maharashtra, Sindudurg, Kankavali, Brahamanwadi, S.R. Dutta, Chakral, H.
Rodrigues, SHK016 (RDNCP0018!)
Ledebouria revoluta (L.f) Jessop ≡ Hyacinthus revolutus L.f. in Suppl.pl (1783:204) Type:—South Africa, Cap
Bonaespei, C. P Thunberg (UPS8508!)
Leaves broadly lanceolate—spathulate, apex acute without bulbil; presence of fibres, inflorescence bearing more than 40 (100) flowers;
flowers pinkish in colour; ovary obtusely deltate without beak at the base, presence of basal lobes.
Specimen studied:—SOUTH AFRICA. KaNgwane: Songimvelo game reserve, 12 July 1992, Meyer. 2531CC (PRE!); Cape de
Bonae, Drege (MNHN-P-P!); Herbier E. Drake, (MNHN-P-P!); Transwaal, 24 September 1959, J.C. Scheepers (MNHN-P-P!);
MOZAMBIQUE. N., Cabo Delagado province, 22 November 2009, Goyder et, al. 6111 (MNHN-P-P!); Carwford 684 (MNHN-P-
P!).
Our sincere thanks to Dr. Mats Hjertson, Curator (Vascular Plants) Uppsala University, Sweden for providing the
type image of Ledebouria revoluta; to the directors and curators of CAL, BLAT, BSI, BSID, AHMA, NGCPR, SUK for
extending their help for studying voucher specimens of Ledebouria; Dr. Subir Bandhopahaya,Rtd. Scientist, Central
National Herbarium (Howrah), Botanical Survey of India for his critical comments during the course of the project; We
are grateful to Mr. Himanshu Agarwal, Assistant Librarian, R. D. & S. H. National College, Bandra (Mumbai) for his
help in procuring original literature.
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