Research Article |
Corresponding author: Ruth P. Clark ( r.clark@kew.org ) Academic editor: Gwilym Lewis
© 2022 Ruth P. Clark, Kai-Wen Jiang, Edeline Gagnon.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Clark RP, Jiang K-W, Gagnon E (2022) Reinstatement of Ticanto (Leguminosae-Caesalpinioideae) – the final piece in the Caesalpinia group puzzle. In: Hughes CE, de Queiroz LP, Lewis GP (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae Part 1: New generic delimitations. PhytoKeys 205: 59-98. https://doi.org/10.3897/phytokeys.205.82300
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A recent molecular phylogenetic analysis of the Caesalpinia group demonstrated that it comprises 26 genera, but the recognition of a putative 27th genus, Ticanto, remained in doubt. This study presents a phylogenetic analysis of ITS and five plastid loci revealing a robustly supported monophyletic group representing the Ticanto clade, sister to the morphologically distinct genus Pterolobium. Based upon this evidence, along with a morphological evaluation, the genus Ticanto is here reinstated. Descriptions are provided for all nine species of Ticanto, together with a key to the species, maps, and colour photographs. Nine new combinations are made: Ticanto caesia (Hand.-Mazz.) R. Clark & Gagnon, T. crista (L.) R. Clark & Gagnon, T. elliptifolia (S. J. Li, Z. Y. Chen & D. X. Zhang) R. Clark & Gagnon, T. magnifoliolata (Metcalf) R. Clark & Gagnon, T. rhombifolia R. Clark & Gagnon, T. sinensis (Hemsl.) R. Clark & Gagnon, T. szechuenensis (Craib) R. Clark & Gagnon, T. vernalis (Champion ex Benth.) R. Clark & Gagnon and T. yunnanensis (S. J. Li, D. X. Zhang & Z.Y. Chen) R. Clark & Gagnon. The final major question in the delimitation of segregate genera from within Caesalpinia sensu lato and the Caesalpinia group is thus resolved.
Biancaea, Caesalpinia crista, Caesalpinieae, China, Fabaceae, Guilandina bonduc, Mezoneuron, phylogeny, Pterolobium, South-East Asia, winged fruit
Caesalpinia s.l., and the Caesalpinia group more broadly, for a long time defied taxonomic classification, their circumscriptions and generic limits being difficult to define. This was due in part to high levels of morphological homoplasy and the consequent lack of defining characteristic synapomorphies available to delineate segregate genera. Caesalpinia s.l. has most often been treated as a single, pantropical genus with up to ca. 150 species encompassing a great diversity of morphological forms, but it has also been considered to comprise numerous smaller genera under as many as 30 generic synonyms (
Despite the dense sampling achieved by
The difficulties inherent in morphologically defining the elements of the Caesalpinia group are exemplified by Ticanto. It lacks obvious diagnostic synapomorphies and was not morphologically characterised by
The aim of our study is to test the monophyly of the putative genus Ticanto using molecular phylogenetic methods and detailed investigation of morphological characters compared with those of the most closely related genera in the Caesalpinia group, particularly Pterolobium, Mezoneuron and Biancaea (
The species descriptions were developed using herbarium specimens studied at HITBC, IBK, K, and KUN, NPH, and from online specimen images at A, AU, BM, C, CDBI, CSFI, CZH, E, FJSI, GXMG, GXMI, GZAC, GZTM, HGAS, HHBG, IBSC, IMC, IMDY, JIU, L, MO, NAS, NF, NY, P, PE, PEY, SM, SN, SYS, SZG, TAIF, TNM, UC, US, W, WAG, WUK, ZM, via the Chinese Virtual Herbarium (CVH, https://www.cvh.ac.cn/index.php), National Specimen Information Infrastructure (NSII, http://nsii.org.cn/2017/), Plant Photo Bank of China (PPBC, http://ppbc.iplant.cn/), and JSTOR https://plants.jstor.org), in combination with data from protologues and other relevant literature (
Due to the relative homogeneity of vegetative and floral characters between T. crista, T. magnifoliolata, T. sinensis and T. szechuenensis, the descriptions of these species were generated using a subset of the available specimens consisting of fruiting specimens and selected flowering or sterile specimens that could be confidently identified.
The x-ray images of fruit for study of the venation patterns were taken using a Faxitron MX101 machine with a 4-inch square digital plate.
A representative selection of specimens that were consulted, or for which the identification could be verified via a digital specimen image, contributed the primary data set used to generate the distribution maps. To encompass the full geographical range of the species, additional records were downloaded from the Global Biodiversity Information Facility (
Tools used for georeferencing were Google Earth Pro, Google Maps (https://www.google.com/maps) and online gazetteers (GEOLocate, https://geo-locate.org/; Falling Rain Global Gazetteer, http://www.fallingrain.com/world/; and Getty Thesaurus of Geographic Names, https://www.getty.edu/research/tools/vocabularies/tgn/). Preliminary mapping of point localities was carried out using GeoCAT (http://geocat.kew.org/editor). The distribution maps were created using ArcMap 10.5 (
DNA samples were taken from field-collected specimens dried in silica gel or from herbarium specimens. A total of 19 accessions were sequenced, representing six species of Ticanto, two of Pterolobium, one of Mezoneuron and one of Biancaea (Table
Accessions sequenced and used to generate the molecular based phylogeny, with GenBank numbers.
Genus | species | Collector name | Collector number | Country | Herbarium | ITS | trnL-F | matK | rps16 | trnDT |
---|---|---|---|---|---|---|---|---|---|---|
Biancaea | millettii | Zhi-Ming Zhong | ZZM003 | China | IBSC | ON922869 | ON932059 | - | ON971386 | ON971410 |
Caesalpinia | crista | Kai-Wen Jiang | KwT033 | China | NPH | ON922872 | ON932062 | ON971417 | ON971381 | ON971400 |
Caesalpinia | crista | Kai-Wen Jiang | TH101 | China | NPH | - | ON932064 | ON971418 | ON971383 | ON971407 |
Caesalpinia | crista | Zhong-Cheng Liu et al. | LXP-13-23687 | China | SYS | ON922873 | ON932063 | - | - | - |
Caesalpinia | crista | Zhu-Qiu Song | 2021057 | China | IBSC | ON922871 | ON932061 | ON971419 | ON971396 | ON971411 |
Caesalpinia | magnifoliolata | Kiyama et al. | 1233 | China | KUN | ON922868 | ON932058 | - | ON971387 | - |
Caesalpinia | sinensis | Clark | 415 | China | K, IBK | ON922875 | ON932066 | ON971423 | ON971390 | ON971399 |
Caesalpinia | sinensis | Clark | 429 | China | K, IBK | ON922876 | ON932067 | ON971413 | ON971394 | ON971405 |
Caesalpinia | sinensis | Hang Sun | 1672 | China | KUN | ON922874 | ON932065 | ON971415 | ON971388 | - |
Caesalpinia | sinensis | Yun-Hong Tan | s.n. | China | HITBC | ON922877 | ON932068 | ON971428 | ON971397 | - |
Caesalpinia | aff. szechuenensis | Clark | 422 | China | K, IBK | ON922870 | ON932060 | ON971426 | ON971392 | ON971398 |
Caesalpinia | vernalis | Shi-Jin Li | 787 | China | IBSC | ON922880 | ON932071 | ON971425 | ON971389 | ON971412 |
Caesalpinia | vernalis | Ya-Min Zhang | YS023 | China | NPH | ON922881 | ON932072 | ON971422 | ON971384 | ON971408 |
Caesalpinia | vernalis | Zhu-Qiu Song | 2021061 | China | IBSC | ON922879 | ON932070 | ON971420 | ON971382 | ON971406 |
Caesalpinia | sp. | Yong-Mei Yi | YYM05 | China | NPH | ON922878 | ON932069 | ON971421 | ON971385 | ON971409 |
Mezoneuron | scortechinii | Wieringa et al. | 4195 | Australia | WAG | ON922882 | ON932073 | ON971424 | ON971391 | ON971401 |
Pterolobium | punctatum | Clark | 424 | China | K | ON922883 | ON932074 | ON971427 | ON971393 | ON971404 |
Pterolobium | stellatum | MPU | 39 | South Africa | NGB | ON922884 | ON932075 | ON971416 | - | ON971402 |
Pterolobium | stellatum | RBGKewMSB | 145895 | Kenya | K | ON922885 | ON932076 | ON971414 | ON971395 | ON971403 |
Five genetic markers were amplified: the nuclear internal transcribed spacer (ITS) region of the 18S–5.8S–26S nuclear ribosomal cistron, and four plastid loci, namely rps16, the trnD-trnT intergenic spacer, the matK gene and flanking 3’-trnK intron, and the trnL-trnF intron-spacer region. DNA was extracted from ca. 0.1–0.2 g silica gel-dried leaves or 0.1–0.2 g leaves from herbarium sheets using either: (1) QIAGEN DNeasy Plant Mini Kit, following the manufacturer’s instructions; or (2) 2× CTAB (hexadecyltrimethylammonium bromide) method modified from
The PCR reactions were carried out in 25 μl volumes, using 2× PCR Premix ‘Dream Taq’ DNA polymerase buffer (4.0 mM MgCl2) (Thermo Fisher Scientific), 5×TBT (
Forward | Reverse | Reference | ||
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ITS | AB101 | AB102 |
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ITS nested | ITS2 | ITS3 |
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|
rps16 | rpsF | rpsR2 |
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trnD-T | trnD | trnT |
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trnD-T nested | trnD | trnE |
|
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trnD-T nested | trnY | trnT |
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matK-trnK | nested | trnK685F | matKC6-Caesalpinia |
|
nested | trnK4La | trnK2R |
|
|
trnL-F | trnL | trnF |
|
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trnL-F nested | trnLc | trnLd |
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trnL-F nested | trnLe | trnLf |
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The concentration and quality of DNA in each sample was assessed using a Nanodrop 2000 Spectrophotometer (Thermo Fisher Scientific). Amplification products were sequenced directly with modified dideoxy cycle sequencing with dye terminators (according to the manufacturer’s protocol; Thermo Fisher Scientific). Cycle sequencing reactions were run on an ABI 3730 automated sequencer (according to the manufacturer’s protocols; Thermo Fisher Scientific), using 5× Sequencing Buffer, DMSO, BigDye Premix 3.1, primers diluted 1/10, and 50–300 ng of genomic DNA, depending on quality and concentration. Sequencing was performed with 26 cycles using the standard settings: 0.10 minutes at 96 °C, 0.05 minutes at 50 °C, and 4.00 minutes at 60 °C. Automated sequence output files were edited and assembled using Geneious (version 8.1.9, Biomatters, Auckland, New Zealand).
Sequences of the same five genetic markers generated as described above (ITS, rps16, trnD-T, matK-trnK and trnL-F) from 60 accessions representing 51 Caesalpinia group species and two outgroups were downloaded from GenBank and incorporated into the analysis (Suppl. material
Sequences were aligned using MUSCLE (
The ML analyses were implemented using RaxML-HPC2 v. 8.2.10 (
Following visual comparison of the resulting phylogenies, all sequences were concatenated to create a six-locus matrix (ITS + plastid) of 8171 bp and the combined dataset was analysed using both ML and Bayesian methods as described above. In the preliminary RaxML analyses of this six-locus matrix, each accession was separate in the matrix and represented by a separate terminal in the tree. Where accessions were missing two or more loci, multiple accessions of single species were concatenated for subsequent analyses if they appeared in the same clade in the initial analyses, thus minimising missing data for each species. Accessions were concatenated in this way for six species, and these are highlighted in bold in Suppl. material
Results from the phylogenetic trees were visualised using Figtree v1.4.2 (
The concatenated five-locus matrix included 79 accessions (17 newly sequenced, and two of the accessions used by
Separate analyses of the plastid and nuclear datasets revealed the same major clades in both the ML and Bayesian analyses. Incongruences between the nuclear and plastid trees were found at the interspecific level within clades but were unsupported in the nuclear analyses by either bootstrap or posterior probability values; these discrepancies are therefore considered non-contradictory. The three major clades that are of most relevance to this study represent the genera Ticanto, Pterolobium and Mezoneuron, and these were recovered in both the ML and Bayesian analyses (Fig.
Caesalpinia group ML phylogeny from the combined dataset. Bootstrap values above 50 are shown, values > 75 are indicated with an orange dot at the node. Branches in bold indicate Posterior Probability greater than 0.95 in equivalent BI analysis. Arrows indicate nodes not recovered in BI analysis. The collector number of the corresponding voucher for each terminal is included with the species name. Where a terminal results from analysis of multiple vouchers, the collector numbers are separated by an underscore (see Suppl. material
Sequences of Biancaea millettii and Caesalpinia vernalis are incorporated into our analyses. These two species were initially included in the phylogeny of
The six (including accession Yi YYM05, determined as Ticanto sp.) sampled species of the proposed genus Ticanto are resolved as a monophyletic group in all analyses (Fig.
Partial sequences of ITS, rps16 and trnL-F were obtained from a single accession of T. magnifoliolata (Kiyama et al. 1233), which in both the ML and Bayesian analyses was resolved as part of the Ticanto clade. Because the position of this accession is poorly supported due to a high proportion of missing data (80.8%), the version of the phylogeny including this accession is presented separately (Suppl. material
Caesalpinia sect. Nugaria DC., Prodr. 2: 481, 1825.
Nugaria Prain, J. As. Soc. Beng. 66(ii): 470, 1897 nom. inval. nom. provis.
Guilandina paniculata Lam.
Despite reference in the protologue of Ticanto to the plate H.M. 6. t. 19, this did not constitute typification of the name because Adanson did not mention a previously or simultaneously published species name, nor the type of such a name (
The name Ticanto was a vernacular name used for these plants by the Brachmanes, also known as Brahmanas, Brahmans, or Brahmins, a sector of Hinduism. This was referenced by
Scandent shrubs or lianas to 15 m. Stems usually with scattered, recurved prickles. Leaves pari-bipinnate, pinnae 1–16 opposite pairs, leaflets 2–15 opposite pairs, leaf rachis with recurved prickles at base of pinnae and usually scattered in between. Stipules 0.25–3 mm long. Leaflets elliptic to ovate or obovate, oblong or rhombic. Inflorescence a terminal or axillary raceme or panicle 7–42 cm long; pedicels articulated; bracts at base of racemes, caducous, bracteoles at base of pedicels, caducous. Flowers zygomorphic, with a hypanthium, calyx lobes 5, free, the lower lobe cucullate over the others in bud; petals 5, 3.5–12 × 2–7 mm, the median petal distinct from the others in shape, usually with an approximately circular patch of hairs on the inner surface, the lateral petals glabrous or with few hairs; stamens 10, free, 4–14 mm long, the basal half tomentose; ovary 1–2-ovuled, glabrous or hairy; style 4–12 mm long; stigma funnel-shaped and more or less papillate, or truncate. Fruit coriaceous or ligneous, dehiscent or indehiscent, elliptic, lunate, or sub-circular, 1.5–7 × 1.5–5 cm, apex acute or beaked, with or without a stipe, the upper suture with or without a narrow wing 0.5–4 mm wide, or a carinate wing 5–6 mm deep, 1(–2)-seeded.
Andaman Islands, Australia, Cambodia, China (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hong Kong, Hubei, Hunan, Sichuan, Taiwan, Yunnan, Zhejiang), India, Indonesia, Japan (Ryukyu Islands), Malaysia, Mauritius, Micronesia, Myanmar, New Caledonia, Papua New Guinea, Philippines, Polynesia, Sri Lanka, Thailand, Vietnam (Maps
1 | Pinnae 8–16 pairs, fruit dehiscent, ligneous | T. vernalis |
– | Pinnae 1–8(–9) pairs; fruit usually indehiscent, coriaceous or ligneous | 2 |
2 | Leaflets 8–15 pairs; 0.8–1.5 × 0.4–0.6 cm | T. caesia |
– | Leaflets 2–7 pairs; (1.2–)1.5–13(–15) × 0.8–8 cm | 3 |
3 | Leaflets rhombic, 1.5–2 × 0.8–1.3 cm; fruit without a wing | T. rhombifolia |
– | Leaflets elliptic, ovate or obovate, 1.2–13(–15) × 0.8–8 cm; fruit with or without a wing | 4 |
4 | Fruit dehiscent, ligneous; without a wing; fruit venation not prominent | T. yunnanensis |
– | Fruit indehiscent, coriaceous; with or without a wing; fruit venation prominent | 5 |
5 | Fruit without a wing, slightly asymmetrical to sub-lunate | 6 |
– | Fruit usually with a flat or carinate wing along the upper suture, strongly asymmetrical, sub-circular to lunate or teardrop-shaped | 7 |
6 | Leaflets 7–13 × 4.5–8 cm, underside of leaflets with brown hairs | T. elliptifolia |
– | Leaflets 2.1–7.2 × 1–3.3 cm, underside of leaflets usually glabrous or occasionally with sparse ferruginous hairs | T. crista |
7 | Leaflets 3.5–10.8(–15) × 2.1–7 cm, apex usually rounded; ovary glabrous; fruit wing carinate | T. magnifoliolata |
– | Leaflets 1.2–10.7 × 0.8–5.1 cm, apex usually acute or acuminate; ovary sparsely to densely tomentose, or subglabrous; fruit wing flat or absent | 8 |
8 | Leaflets 1.2–6 × 0.8–3 cm, leaflet apex usually acute; fruit 1.5–3.4 × 1.5–3 cm, wing 1–3 mm wide, present only along part of the fruit length or absent | T. szechuenensis |
– | Leaflets 1.8–10.7 × 0.8–5.1 cm, leaflet apex usually acuminate; fruit 3–5.8 × 1.9–4.1 cm, wing 0.5–4 mm wide | T. sinensis |
Caesalpinia hypoglauca
Chun & F. C. How., Acta Phytotax. Sin. 7: 20 pl. 6. 1958. Type: China. Kwangtung, Sup Man Ta Shan [Mt. Shiwandashan], 26 Jul. 1933, H.Y. Liang 69864 (lectotype: (designated by
Caesalpinia caesia Hand.-Mazz., Oesterr. Bot. Z. 85: 215. 1936.
China. Kwangsi, Fenzel 3 (W!).
Habit a climber. Stems with sparse recurved prickles, puberulent. Stipules unknown. Leaves with 5–8(–9) pairs opposite pinnae; leaf rachis and petiole 15–20 cm, leaf rachis and pinnae rachises pilose; leaflets 8–12(–15) opposite pairs per pinna, subsessile, chartaceous, oblong, base strongly asymmetric, apex truncate or obtuse-rounded, emarginate, 0.8–1.5 × 0.4–0.6 cm, both surfaces glabrous. Inflorescence a panicle, supra-axillary or terminal, 10–15 cm, the axes brown puberulent; pedicels 4–7 mm, articulated. Flowers with a hypanthium, this glabrous, lower calyx lobe ca. 6 mm long, others 3.5–4 mm, all lobes glabrous; petals obovate-oblong, ca. 3.5–5.5 mm long, median petal with rhombic patch of dense hairs on the inner surface at base of blade, other petals pubescent, shortly clawed; stamen filaments ca. 6 mm long, ferruginous pilose at base; ovary glabrous, 2-ovuled, style ca. 4 mm long. Fruit blackish when dry, indehiscent, ligneous, elliptic, inflated at maturity, venation prominent, glabrous, ca. 4.5–5 × 2.3–5 cm, ventral suture narrowly winged. Seed 1, lenticulate, 1.5 × 2.0 cm, blackish.
Sparse forests along rivers, elevation 200–1000 m.
Flowering July-September, fruiting August.
China (Guangxi, Hainan) (Map
Only one specimen collected from Hainan was seen by the current authors (H. Fenzel s.n., see the citation below), of which the detailed locality is unknown (not recorded on the specimen). To include Hainan in the species distribution, we georeferenced this specimen in the centre of the island.
China. Guangxi: Fangcheng, Naliang, s. coll., s.n. (PEY). Fangcheng, Dongzhong, Dakeng Village, Shiwandashan Exped. 3224 (IBK). Shiwandashan, C.L. Tso 23669 (IBSC). Hainan: H. Fenzel s.n. (IBSC).
=Guilandina nuga L., Sp. Pl., 2. 1: 545. 1762. Type: [Indonesia]. East Indies, Ambon, Nugae silvarum Rumph. Herb. Am. 5. p.95, t. 50. 1750.
=Guilandina axillaris Lam., Encycl. 1(2): 435. 1785. Type: [India]. Rheede. Hort. Mal. 6: t. 20. 1686.
=Ticanto nuga (L.) Medik., Theodora 52. 1786. Type: based on Guilandina nuga L.
=Guilandina paniculata Lam., Encycl. 1(2): 435. 1785. Type: [India]. Malabar, Kaka Mullu vel Kaka Moullou (in caption Kaka Mullu) Rheede, Hort. Mal. 6: t. 19. 1686.
=Genista scandens Lour., Fl. Cochinch. 2: 428. 1790. Type: Cochinchina (n.v.).
=Guilandina parvifolia Stokes, Bot. Mat. Med. 2: 466. 1812. Type: [Indonesia]. East Indies, Ambon, Nugae silvarum Rumph. Herb. Am. 5. p.95, t.50. 1750.
=Caesalpinia nuga (L.) W.T. Aiton, Hort. Kew, ed 2, 3: 32. 1811. Type: based on Guilandina nuga L.
=Caesalpinia paniculata (Lam.) Roxb., Hort. Beng. 32. 1814. Type: based on Guilandina paniculata Lam.
=Caesalpinia scandens Heyne ex Roth, Nov. Pl. Sp. 209. 1821. Neotype: (designated by
=Caesalpinia axillaris (Lam.) DC., Prodr. 2: 481. 1825. Type: based on Guilandina axillaris Lam.
=Caesalpinia laevigata Perr., Mém. Soc. Linn. Paris 3: 104. 1825. Type: Philippines. Perrottet s.n. (n.v.).
=Caesalpinia crista var. parvistipula Urb., Symb. Antill. 2(2): 271. 1900. Type: Trinidad. Cult. Hort. Trinidad, Broadway 5589 (n.v.).
Caesalpinia crista L. Sp. Pl. 1: 380. 1753, emend Dandy & Exell in J. Bot. 76: 179. 1938. ≡ Guilandina crista (L.) Small, Fl. S.E. U.S. 591, 1331 (1903).
[Sri Lanka] Ceylon (“Ceylan”), Herb. Hermann vol. 1, fol. 68, no. 157 (lectotype (designated by Skeels in Science, n.s., 37: 922. 1913): BM [BM000621459!]) (note: the sheet bearing this specimen was previously identified by a single barcode, BM000594500, which was subsequently replaced with four barcodes representing the four separate specimens on the sheet).
Habit
a liana or scrambling shrub to 15 m. Stems with few, scattered recurved prickles to 5 mm, sometimes with spine-tipped corky tubercles on older stems, or unarmed, glabrous or occasionally sparsely tomentose. Stipules persistent, triangular, ca. 1 × 1 mm. Leaves with 3–6(–8) pairs pinnae, these opposite to occasionally slightly subopposite; petiole 1.8–5 cm; rachis 4–31 cm, usually armed with recurved prickles, these sometimes also on pinnae rachises, petiole and rachis usually glabrous, occasionally sparsely to moderately ferruginous tomentose; leaflets 2–4(–7) pairs per pinna, coriaceous, opposite, elliptic, base cuneate to obtuse, apex rounded to obtuse, less commonly acute, obtuse, or acuminate, terminal leaflets 2.1–7.2 × 1–3.3 cm, lateral leaflets 2.1–5.8 × 1–3.1 cm, upper surface glabrous, often glossy, lower surface glabrous or occasionally sparsely ferruginous tomentose, venation reticulate, anastomosing, visible on both surfaces. Inflorescence a raceme or panicle, axillary or terminal, 8–40 cm, axes glabrous or sparsely tomentose; pedicels 5–15 mm; bracts persistent, triangular or lunate, 1–1.5 × 1 mm; bracteoles caducous, broadly elliptic, apex acute, margins sometimes with small teeth, 1–2.5 × 0.5–1 mm. Flowers with a hypanthium 1–2 × 3–6 mm, glabrous or sparsely ferruginous tomentose; lower calyx lobe 6–8(–10) × 3–4 mm, other lobes 5–6(– 8) × 2–3 mm, all lobes glabrous, or margins ciliate, or sparsely ferruginous tomentose; median petal 6–9 × 3–7 mm, with dense circular patch of hairs on inner surface at base of blade; upper laterals 6–10 × 3–5 mm, including claw ca. 1 mm, glabrous or inner surface of claw sparsely hairy; lower laterals 7–11 × 3–6 mm, including claw ca. 1 mm, glabrous or inner surface of claw sparsely hairy; stamen filaments 4–12 mm, the vexillary shorter than the lower ones, orange tomentose on lower ½–¾ on inner surface; anthers 1–1.5 mm; ovary 2–4 mm long, glabrous or sparsely or partially tomentose; style 4–11 mm; stigma cupular or funnel-shaped, the rim papillate, sometimes only slightly so, 0.5–1 mm wide. Fruit indehiscent, coriaceous, elliptic to lunate, subsymmetrical to somewhat asymmetrical, slightly inflated, base cuneate, stipe 2–5 mm, apex acute to beaked, beak 1–10 mm, venation reticulate, prominent, glabrous or very sparsely tomentose, (2–)2.7–7 × 2.2–3.7 × 0.3–0.8 cm, lacking a wing. Seeds 1, rarely 2, circular to reniform in outline, flat, ca. 2–2.5 × 1.5–2 × 0.5–1 cm. (Fig.
Riverbanks, sandy beaches, in and behind sandy parts of mangroves, on chalk rocks and limestone, at low altitude, elevation rarely up to 350 m.
The flowering and fruiting time of this species varies throughout its distribution and may be correlated with latitude or biome as indicated below; however, the periodicity listed below derives in some cases from few records and thus may be incomplete.
China, Japan: Flowering February-April, fruiting April-October;
Bangladesh, India, Myanmar: Flowering August-December, fruiting December-October;
Thailand, Vietnam: Flowering January-June, fruiting January-June;
Malaysia, Indonesia, Palau Islands: Flowering February-December, fruiting February-January;
Philippines: Flowering December-March, fruiting May-December;
New Guinea: Flowering January-November, fruiting February-November;
New Hebrides, Solomon Islands: Flowering February-December, fruiting February-December;
New Caledonia: Flowering May, fruiting unknown;
Mauritius: Flowering unknown, fruiting February.
Andaman Islands, Australia, Cambodia, China (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hong Kong, Hubei, Hunan, Sichuan, Taiwan, Yunnan), India, Indonesia, Japan (Ryukyu Islands), Malaysia, Mauritius, Micronesia, Myanmar, New Caledonia, Papua New Guinea, Philippines, Solomon Islands, Sri Lanka, Thailand, Vanuatu, Vietnam (Map
The current authors have not seen any specimens or specimen records from Cambodia (other than a single record in GBIF which is not from a preserved specimen), but
Australia. Queensland: Daintree area, Noah Creek, Mangroves, J.J. Wieringa 4199 (WAG). Bangladesh. Chittagong, Flagstaff 255 (K). China. Chongqing: Jiulongpo, Jinfeng, Baihe Village, Jiulongpo Exped. 500107150402-289LY (IMC). Mt. Jinyunshan, Z.Y. Liu 182996 (IMC). Fujian: Hua’an, Wenhua, W.D. Han 20667 (NF). Pinghe, Daxi, Jiangzhai Village, H.B. Chen s.n. (FJSI). Xiamen, Yunding Cliff, G.D. Ye 1208 (IBSC). Yunxiao, Huotian, Baihuayang Reservoir, G.D. Ye 2482 (FJSI). Zhangzhou, Zhaoan, Wushan, X.F. Zeng ZXF19839 (CZH). Zhangzhou, Zhaoan, Wushan, Jinshui Village, X.F. Zeng ZXF41029 (CZH). Zhao’an, Jinshui Village, s. coll., s.n. (AU). Guangdong. Boluo, Mt. Luofushan, near Damiao, Yue78 5714 (IBSC). Dianbai, Luokeng, Mt. Shuangjiling, H.G. Ye 6379 (IBSC). Huidong, Pingshan Forest Farm, Mt. Chenshuishan, Zhulian?, P.Y. Chen, B.H. Chen & G.C. Zhang 46 (IBSC). Jiangmen, Mt. Guifengshan, J.Y. Chen 20165220 (SN). Qingyuan, Yangshan, near Qincaitang Reservoir, K.W. Jiang KwT033 (NPH). Ruyuan, Daqiao Health Center, back mountain, Yue73 1182 (CSFI). Shenzhen, Longgang, Nan’ao, Yangmeikeng, S.Z. Zhang, L.Q. Li et al. 185 (SZG). Xin’an, Ng-tung Shan, T.M. Tsui 231 (NAS). Zhaoqing, Mt. Dinghushan, S.J. Li 30 (IBSC). ibid., Z.Q. Song 2021057 (IBSC). Guangxi: Liuzhou, Longtan Park, Mt. Jiaodingshan, Longtan & Dule Exped. 242 (IBK). Yang-shoh, H. Fung 21112 (SYS). Guizhou: Tungtze, Y. Tsiang 4894 (IBSC). Hainan: Kan-en, Chim Fung Ling, near Sam Mo Watt Village, S.K. Lau 3582 (IBSC). Lin’gao, Maniao, Wende Village, Z.X. Li et al. 911 (IBSC). Qionghai, Lehui, near Shuangbang Village, Y. Zhong 4472 (IBSC). Wanning, Mt. Dongshan, the second mountain range, S.P. Kao 52115 (IBSC). Wenchang, Longlou, nera Mt. Beijianshan, G.W. Tang, Z.M. Li & J. Li TangGW2525 (IBSC). Hubei: Shennongjia, Xingshan to Yangri, D.G. Zhang ZB130226624 (JIU). Xingshan, Xiakou, Jianyangping, Lifangyan to Huangliang, D.G. Zhang zdg4185 (JIU). Hunan: Jianghua, C.J. Qi 3822 (CSFI, IBSC). Xiangxi, Yongshun, Zejia, Donglu Village, K.D. Lei ZZ40516121 (JIU). Jiangxi: Ji’an, Suichuan, Daijiapu, Xianmo, Z.C. Liu, W.J. Xiong, F. Ye, L. Deng, M. Tu, X.J. Zhang, L. Feng, Q.Y. Yin & N.N. Liu LXP-13-23687 (SYS). Taiwan: Hsinchu, Hsienchiaoshih, Z.-H. Chen 277 (TAIF). Yunnan: Hekou, Erqu, W.X. Liu 277 (HITBC). Micronesia. Caroline Islands, Yap Group, Gorror Island, Central Plateau, E.Y. Hosaka 3319 (K). India. Kaswar, R.J. Bell 7750 (K). Kuppam River, Taliparamba, C.A. Barber 8788 (K). North Kanara, W.A. Talbot 1256 (K). S. Andaman, Dr King’s Collector s.n. (K). Myanmar. Myebon, H.S. McKee 6069 (K). Rangoon, D.R. Khant 1079 (K). Mauritius. The Pouce, J. Gueko s.n. (K). New Caledonia. Yate, Touaourou, s. coll. s.n. (K). Thailand. Narithiwat: Kulok river mouth next to bridge on road from Tak Bai to Sungai Ko-lok, P.S. Herendeen & R. Pooma 1-V-1999-3 (US). Vanuatu. Aniwa Island, Isavai village, P. Curry 1447 (K). Banks Islands, Port Patterson, A. Morrison s.n. (K). New Hebrides, Erromanga, between Nouanko Camp and Ipota, about 10 km E of Ipota, P.S. Green RSNH1318 (K). New Hebrides, Port Vila, A. Morrison s.n. (K).
Caesalpinia elliptifolia S. J. Li, Z. Y. Chen & D. X. Zhang, Nordic J. Bot. 22: 349. 2003.
China Guangdong, Fengkai, Qixing, alt. 120m, 20 July 2000, Shijin Li 026 (holotype: IBSC!)
Habit a liana to 15 m. Stems occasionally with scattered, recurved prickles to 2 mm. Stipules caducous. Leaves with 1–2 pairs opposite pinnae; leaf rachis 20–30 cm, leaf rachis and pinnae rachises with recurved prickles; petiolules 2–3 mm; leaflets 2 opposite pairs per pinna, coriaceous, broadly elliptic, base cuneate to rounded, apex rounded, obtuse or acute, 7–13 × 4.5–8 cm, upper surface glabrous, glossy, lower surface with brown hairs especially on midvein; venation anastomosing, finely reticulate. Inflorescence a panicle, supra-axillary or terminal, 15–25 cm, all parts densely hairy; pedicels 8–12 mm, articulated; bracts caducous, lanceolate, 1–3 mm; bracteoles caducous, ca. 1.5 mm. Flowers with a hypanthium, this with brown hairs; calyx lobes ca. 6 × 2 mm, with brown hairs; median petal blade reflexed, claw ca. 3.5 × 1 mm, blade ca. 7 × 6–7 mm, circular patch of brown hairs at base of blade, otherwise glabrous; lateral petals 10–12 × 4–5 mm, claw ca. 1 mm, glabrous; stamen filaments 9–14 mm, the basal ca. ½ tomentose, anthers 2 mm; ovary subsessile, ca. 2 mm long, tomentose, 1- or 2-ovuled; style (2–)7–10 mm, occasionally as short as 2 mm, glabrous; stigma truncate, papillate. Fruit indehiscent, coriaceous, oblong-elliptic to sub-lunate, sub-symmetrical, compressed but slightly inflated when mature, base cuneate, stipe short, apex acute to attenuate, beak ca. 1–5 mm, veins prominent and reticulate, ca. 4.5–5 × 2.2–2.5 cm, lacking a wing. Seeds 1 or 2, brownish black, compressed, sub-circular, ca. 10–15 mm cm in diameter.
Beside ditches, elevation ca. 100 m.
Flowering April, fruiting May-June.
China (Guangdong) (Map
Caesalpinia magnifoliolata Metcalf. Lingnan Sci. J. 19: 553. 1940.
China. Kwangsi, Ling Yun Hsien, Steward, A.N. & Cheo, H.C. 583 (holotype: A [A00059894!]).
Habit
a scrambling shrub. Stems with scattered recurved prickles, ferruginous puberulent, glabrescent. Stipules not seen. Leaves with 2–3(–4) pairs opposite pinnae; petiole 3.5–9 cm; leaf rachis 3.3–18.5 cm, with paired recurved prickles at the pinna insertion points and scattered in between, or unarmed; pinnae 2–9 cm; leaflets 2(–3) opposite pairs per pinna, coriaceous, elliptic to obovate, base oblique, apex usually rounded or obtuse, retuse to emarginate, occasionally acute; terminal leaflets 3.5–10.8(–15) × 2.1–7 cm; lateral leaflets 3.5–9.3 × 2.1–4.6 cm; both leaf surfaces glabrous, or lower surface sparsely puberulent; venation reticulate, anastomosing. Inflorescence a raceme or panicle, axillary or terminal, 15–30 cm; axes and pedicels sparsely to moderately ferruginous tomentose; pedicels 5–11 mm, articulated, glabrous; bracts and bracteoles not seen. Flowers with a hypanthium ca. 1 × 2–4 mm, glabrous to sparsely orange tomentose; lower calyx lobe ca. 7 × 3 mm; other calyx lobes ca. 5–6 × 2 mm; all calyx lobes with ciliate margins; median petal inrolled, with a patch of hairs at base of blade on inner surface, ca. 7–10 × 3–5 mm; upper laterals ca. 7–10 × 3–5 mm, hairy on the claw inner surface; lower laterals ca. 7–10 × 3–5 mm, hairy on the claw inner surface. Stamen filaments ca. 5–9(–10) mm, the basal ½ tomentose; ovary ca. 3 mm long, glabrous, subsessile; style 5–10 mm, glabrous; stigma funnel-shaped, papillate, sometimes laterally placed. Fruit dark brown, indehiscent, coriaceous, lunate, stipe ca. 1 mm, beak 2–7 mm, venation prominent, glabrous, 2.8–4.2 × 2.2–3.1(–3.5) × 0.4–0.7 cm, wing on ventral suture carinate, 5–6 mm deep. Seed 1, brownish black, compressed, sub-circular, ca. 2 × 2.5 cm (Fig.
Photos of Ticanto A T. crista (i) habit (ii) leaves (iii) flowers (E.D. Liu 8629, Yunnan, China (KUN)) (iv) leaves (K.W. Jiang KwT033 (NPH)) (v) fruit and seeds (Z.Q. Song s.n. (fruit and seeds in NPH)) B T. magnifoliolata fruit and seeds (Q. Meng s.n., Guizhou, China (fruit and seeds in NPH)) .
Forests, scrubland; 400–1800 m.
Flowering February-July, fruiting May-November.
China (Guangdong, Guangxi, Guizhou, Yunnan) (Map
China. Guangdong: Luoding, Caotang, Lianshi Village, B.H. Chen et al. 1793 (IBSC). Guangxi: Donglan, Sannong, Xiangtiandong Village, Haoyantun, H.Z. Lü, Z.Z. Lan & H.F. Cen 451224180425005LY (GXMG). Fusui, Zhongdong, Luhan?, near Mt. Damingshan, S.H. Chun 12435 (IBSC). Huanjiang, Mulun Nature Reserve, Xiazhai Observation Deck, W.B. Xu, R.C. Hu & M.Q. Han ML1296 (IBK). Leye, on the way from Gantian to Daping, M. Shi s.n. (NPH). Lingle, Xinhua, Sanhe, Lingle Exped. 34420 (IBK). Longzhou, Jinlong, Jinmei, Nongqiao, W.B. Xu, Y.S. Huang et al. LZ1370 (IBK). Napo, Bing Mung, Rong La Shan, H. Akiyama, H. Kudo, J. Murata, T. Sugawara, N. Tanaka, Y. Tateishi, Y.G. Wei & S.G. Wu 1233 (KUN). Pingguo, Taiping, Chami Village, Chajiangtun, H.Z. Lü, Y. Lin, L.L. Mo & B.Z. Ban 451023150325048LY (GXMG). Pingnan, Yaoshan, Luoxiang, Mt. Lingdingshan, C. Wang 39106 (IBK). Guizhou: Anlong, Huali, Dewo, Guizhou Exped. (C.S. Chang & Y.T. Chang) 3543 (HGAS). Anlong, Longshan, Dushan, Guizhou Exped. (C.S. Chang & Y.T. Chang) 3138 (HGAS, IBSC, KUN, NAS, PE). Ceheng, Shuanghe, Jishanlin, C.Z. Dang 1684 (HGAS). Pingba, Qibo, Taohua Village, Baidong, R. Yang & L.B. Yan PB1356 (GZAC). Xingren, Nongchang, near Shanhe, Guizhou Exped. (C.S. Chang & Y.T. Chang) 7790 (HGAS, IBSC, KUN, NAS, PE, WUK). Xingyi, Qingnan, Yangping, Bajiaoping?, Anshun Exped. 744 (HGAS). Yunnan: Funing, Zhesang, Nonguo Village, J. Cai, J.D. Ya, X.Q. Yu, Y. Su & C.H. Li 14CS9136 (KUN). Mali, Xialiangshuijing, Malipo Exped. 5326240386 (IMDY). Malipo, on the way from Huilong to Mabiao, X.X. Zhu, B. Xiao, G.S. Wang & J. Wang LiuED8612 (KUN). Si-chour-hsien, Ping-chai, K.M. Feng 12522 (KUN). Simao, Simaogang, H. Wang 3842 (HITBC).
Caesalpinia rhombifolia J. E. Vidal, Adansonia, n.s., 15: 394. 1976.
Vietnam. Quang Ninh, Dam Ha, W.T. Tsang 29830 (holotype: P [P02142684!]; isotypes: C [C10011919!], E [E00313521!], G, K [K000789332!], L [L0018793!], SING).
Habit a liana. Stems with sparse recurved prickles, glabrous. Stipules caducous, not seen. Leaves with 3–6 pairs opposite pinnae; petiole 2.5–3 cm; leaf rachis ca. 10–15 cm, with paired recurved prickles at base of pinnae and scattered in between, glabrous; pinna rachis 3.5–6 cm, glabrous; petiolules ca. 1 mm; leaflets 3–5 opposite pairs per pinna; chartaceous, slightly rhombic, base broadly cuneate, apex acute, rarely slightly emarginate, 1.5–2 × 0.8–1.3 cm, both surfaces glabrous; venation reticulate, anastomosing. Inflorescence a panicle, axillary or terminal, 10–20 cm; axes glabrous; pedicels 6–9 mm, articulated, glabrous or subglabrous. Flowers with a hypanthium that is sparsely ferruginous tomentose or glabrous; calyx lobes with ciliate margins; petals ca. 7 mm long, median petal smaller than the others, rounded at apex, with a patch of hairs at base of blade, other petals hairy on inner surface of claw; stamen filaments ca. 7 mm long, pale orange tomentose on basal ca. ½ on inner surface; anthers ca. 1.5 mm long; ovary glabrous, 1- or 2-ovuled; style ca. 10 mm, glabrous. Fruit indehiscent, coriaceous, asymmetrical, sub-lunate, inflated when mature, stipe ca. 3 mm, apex slightly beaked, venation prominent, glabrous, ca. 3–4 × 2.3–2.5 cm, lacking a wing. Seed 1, broadly ellipsoid, ca. 1.8–2 × 1–1.5 × 1.1 cm, matt black.
Thickets.
Flowering May-July, fruiting October-November.
China (Guangxi), Vietnam (Map
China. Guangxi: Dongxing, Jiangping, Hezhou, Dongxing Exped. 450681180510051LY (IBK). Fangcheng, Dawangjiang Village, Y.S. Huang & L. Wu H110397 (IBK).
=Caesalpinia chinensis Roxb. in Fl. Ind. ed. 2: 361. 1832., nom. rej. Li et al. Taxon 51: 816. 2002. Type: not designated.
=Mezoneuron sinense var. parvifolium Hemsl., J. Linn. Soc., Bot. 23: 205. 1887. Type: China. Hupeh Province, Ichang, Henry, A. 2238 (lectotype, designated here: K [K000264687!]; isolectotype: P [P00751902!])
=Caesalpinia tsoongii Merr., Philipp. J. Sci. 27: 162. 1925. Type: China. Szechuen, Tsoong 4190. (holotype: UC [UC227358!]; isotype: GH [A00059897!]).
=Caesalpinia stenoptera Merr., J. Arnold Arbor. 19: 35. 1938. Type: Indo-China. Tonkin, Cao Bang, Ban Gioc, Jun. 1933, Petelot, A. 4757 (lectotype, designated here: A [A00059899!], isolectotypes: P [P02142685!, P02142686!], NY [NY00003575!, NY00003576!, NY00003577!]).
Mezoneuron sinense Hemsl., J. Linn. Soc., Bot. 23: 204. 1887. ≡ Caesalpinia sinensis (Hemsl.) J.E. Vidal in J.E. Vidal & S. Hul Thol, Bull. Mus. Natl. Hist. Nat., ser. 3, 395 (Bot. 27): 90. 1976. nom. cons. Li et al. Taxon 51: 816. 2002.
China. Hupeh, Ichang, A. Henry, (Herb. Kew) (lectotype (designated by
Habit
a scandent shrub or vine to 13 m. Stems with scattered recurved prickles to 4 mm, glabrous or sparsely whitish to pale orange tomentose. Stipules persistent, triangular, 1–3 × 1–2.5 mm. Leaves with 2–4(–5) pairs opposite pinnae; petiole (1.3–)3–7 cm; leaf rachis 2.5–24 cm, with paired recurved prickles at pinna insertion points and scattered in between, sometimes densely; pinnae 2.5–12.5 cm, sometimes with recurved prickles in pairs at the leaflet insertion points and scattered in between; leaf rachis and petiole glabrous to sparsely whitish to pale orange tomentose; pinna rachis glabrous to sparsely pale orange tomentose; leaflets 2–5 opposite pairs per pinna; elliptic, base cuneate to rounded, sometimes oblique, apex usually acuminate, or acute, occasionally rounded; terminal leaflets 1.8–10.7 × 0.9–5.1 cm; lateral leaflets 1.9–9.2 × 0.8–4.7 cm; all leaflets glabrous on both surfaces or lower surface sparsely orange tomentose at base and on midvein, sometimes at margins, glossy above; venation reticulate, anastomosing. Inflorescence a panicle, axillary, supra-axillary or terminal, 7–42 cm long, axes sparsely to densely ferruginous tomentose, axis sometimes with small, recurved prickles; pedicels (3–)4–12(–17 in fruit) mm, articulated, sparsely to moderately ferruginous tomentose; bracts caducous, triangular, 0.5–2 × 1–1.5 mm; clusters of triangular scale-like bracts sometimes below base of raceme; bracteoles caducous, broad, elliptic, acute, 2–3 × 1–1.5 mm. Flowers with a hypanthium 1–2 × 3–5 mm, sparsely to moderately ferruginous tomentose; lower calyx lobe 6–8 × 3–5 mm, other lobes 5–6 × 2–3 mm, all lobes sparsely to densely pale orange to ferruginous tomentose on inner and outer surface; median petal obovate, sometimes reflexed backwards, inrolled, 6–8 × 3–4 mm, including claw 1–2 mm long, a circular patch of orange hairs between claw and blade, hairs on margins of claw; upper laterals obovate, 6–10 × 2–6 mm, including claw ca. 1 mm long, petal glabrous or with a few hairs on inner surface of claw; lower laterals 6–10 × 2–6 mm, including claw ca. 1 mm long, glabrous or with a few hairs on inner surface of claw; stamen filaments flattened, 5–12 mm long, densely orange villous on basal ½; anthers 1–2 mm long; ovary 2–5 mm long, sparsely to densely, sometimes partially, orange tomentose, occasionally glabrous; style 6–12 mm long, sparsely hairy on basal ½; stigma funnel-shaped, not or very slightly papillate, sometimes slightly laterally placed. Fruit light green, indehiscent, coriaceous, sub-circular to lunate, base cuneate to rounded, not stipitate or stipe 0–2 mm, apex with a pronounced beak to 25 mm, venation prominent, sparsely ferruginous tomentose, the indumentum most dense at base and on margins, glabrescent or glabrous, 3–5.8 × 1.9–3.6(–4.1) cm × ca. 4–8 mm deep, wing on ventral suture 0.5–4 mm wide. Seed 1, ca. 1.8–2.5 cm diameter, matt or glossy dark brown. (Fig.
A T. sinensis (i) habit (ii) leaves (iii) flowers (photos by Xin-Xin Zhu, Guizhou, China, unvouchered) (iv) habit (v) fruit (R.P. Clark 429, Guangxi, China, (IBK, K)) (vi) leaves (R.P. Clark 415, Guangxi, China (IBK, K) B T. aff. szechuenensis (i) habit (ii) inflorescence and leaves (R.P. Clark 422, Guangxi, China (IBK, K)).
Forest and thicket, on limestone. Elevation 100–1500m.
Flowering March-May, fruiting March-October.
China (Chongqing, Guangxi, Guangdong, Hong Kong, Hubei, Sichuan, Yunnan), Laos, Myanmar, Vietnam (Map
The current authors have not seen any specimens or specimen records from Hong Kong but the area is included within the distribution listed by
China. Chongqing: Qijiang, Wansheng, Heishangu Ave., opposite of Yaqulou, S.R. Yi YSR9620 (NPH). Shimiaoxiangzhai, S.G. Tang s.n. (SM). Wushan, Guandu River, T.P. Wang 10431 (WUK). Fujian: Mengtongyang, Chengmenkan, H.Y. Zou 0931 (NF). Guangdong: Dinghu, Xinghu, Yuping Peak, K.C. Ting & G.L. Shi 1337 (WUK). Zhaoqing, Qixingyan, F.C. How 74128 (IBK). Guangxi: Bama, Xishan, Z.T. Li 601739 (KUN). Donglan, Ma’an mountain, R.P. Clark 429 (K, IBK). Debao, Longguang, Miaohuai Village, Debao Exped. 451024160516009LY (IBK). Fusui, Quli, Jidao Village, B.Y. Huang, Y.Y. Xie & H.F. Cen 451421160523025LY (GXMG). Jingxi, Longlin to Ande, R.P. Clark 415 (K, IBK). Liuzhou Longtan Park, W.E. Qun 150 (K). Long’an, Nanxu, Longxintun, Long’an Exped. 450123130506007LY (IBK). Longlin, Kechang, Haichang Village, Dankuntun, L.Y. Yu, Y.D. Peng & X.Y. Hu 451031140410083LY (GXMG). Napo, county town, Hongshui River Exped. 400 (TNM). Ningming, W Tingliang, C.C. Huang et al. 2111 (GXMI). On the slopes of the limestone mountain near the county seat of Lingle, Z.T. Li 603637 (IBK). S Nanning, Dar Shan, Seh-Feng, Me-Jon, R.C. Ching 8435 (US). Tianyang, Wucun, Dalu Village, Longlitun, Tianyang Exped. 451021150410060LY (GXMG). Guizhou: Ceheng, Rongxian, Huangtian to Maoping, Z.Y. Cao 544 (PE). Kwanlin, Kwanlinchow, Da-Swee-Tzi, S.W. Teng 1641 (IBSC). Zhenfeng, Beipanjiang, Shuiyanba Village, Y. Jia 522325190716483LY (GZTM). Hubei: Badong, T.P. Wang 10830 (IBK). Sichuan: Jiang’an, Nanyan, Hongfo Temple, K.Y. Lang 3033 (PE). Xuanhan, Dong’an, Xuanhan Exped. 1498 (SM). Yunnan: Between Szemao and Puerhfu, J.F. Rock 2849 (NY, US). Lushui, near Nujiang River, H. Sun 1672 (KUN). Xichou, Changqing, C.W. Wang 81282 (KUN). Yingjiang, 23 km milestone on X309 Road from Pingyuan to Kachang, Y.J. Guo, W.L. Zhao, P.X. Tang, X.L. Jin & X.Q. Zhang 13CS7525 (KUN). Laos. Phou Phung pres de Louang Prabang, M. Poilane 20257 (K, L). Myanmar. Bhamo District, Lapycke to Sinlum Kabo, J.H. Lace 5769 (K). Vietnam. Indo-China, Tonkin, A. Petelot 4757 (NY). Ninh Binh: Cuc Phuong National Park, N.M. Cuong 464 (MO).
=Caesalpinia kwangtungensis Merr., J. Arnold Arbor. 8: 7. 1927; Herkl. in Hong Kong Naturalist ix. 32. 1938, descr. ampl. Type: China. Kwangtung, Wilson in Canton Christ. Coll. 12838 (lectotype, designated here: (GH [A00059893!], isolectotypes: BM [BM000958803!], E [E00313522!], LU, NAS, [NAS00071304!, NAS00071305!], P [P02142689!], US [US00002578!]).
Caesalpinia szechuenensis Craib, Pl. Wilson. (Sargent) 2(1): 92. 1914.
China. Western Szechuan, Kiating Fu, May 1908, E.H. Wilson 3255. (lectotype, designated here: K [K000980490], isolectotypes: A [A00059895!, A00059896!], BM [BM000958802!], E [E00313523!], GH, NY [NY00003572!], US.)
Habit
a scandent shrub. Stems with sparse, scattered recurved prickles, glabrous. Stipules minute, ca. 0.5 × 0.25 mm, triangular, subglabrous. Leaves with 3–6 pairs opposite to strongly subopposite pinnae; petiole 1.8–7 cm, leaf rachis with prickle at the base of each pinna and scattered in between pinnae insertions, 6–22 cm; pinnae 3.8–8.5 cm, occasionally with prickle at base of petiolule; leaflets 3–6 opposite pairs per pinna, elliptic to slightly ovate, the apex usually acute, occasionally slightly rounded; terminal leaflets 2–6 × 1–3 cm, lateral leaflets 1.2–6 × 0.8–3 cm, upper surface glabrous, lower surface glabrous or with a few ferruginous tomentose hairs on midvein at base or with sparse short ferruginous hairs; venation prominent on both surfaces, reticulate, anastomosing. Inflorescence a terminal, few-branched panicle, 11–15 cm long, axes subglabrous to sparsely to moderately ferruginous tomentose; pedicels (5–)9–11 mm, articulated, glabrous to sparsely ferruginous tomentose; bracts not seen; bracteoles caducous, 1 × 0.25 mm, lanceolate-acuminate. Flowers with a hypanthium ca. 1–2 × 3–4 mm, sparsely ferruginous tomentose; lower calyx lobe 8 mm long, other lobes 5 mm long; median petal 6–9.5 × 2.5–5 mm, with a patch of hairs at base of blade and few hairs on the claw; upper laterals ca. 5–10 × 3–4 mm, glabrous or with a few hairs on inner surface of the claw; lower laterals ca. 5–10 × 3–4 mm, glabrous or with a few hairs on inner surface of claw; stamen filaments flattened, ca. 9 mm long, densely orange villous on basal ½; ovary ca. 2.5 mm long, sparsely to moderately densely pale orange tomentose; style 10–12 mm, with a few hairs at the base, otherwise glabrous, ovules 2; stigma funnel-shaped, papillate, ca. 1 mm wide. Fruit indehiscent, coriaceous, strongly asymmetrical, sub-lunate to sub-circular or teardrop-shaped, stipe 0–1 mm, beak 1–5 mm, venation prominent, glabrous, 1.5–3.4 × 1.5–3 cm × 0.4–0.6 cm, wing sometimes present along part of length of ventral suture, 1–3 mm wide. Seed 1, circular, dark brown, 1.4–1.7 cm diameter (Fig.
Mountain forest, thicket, on limestone, elevation 260–1500 m.
Flowering April-August, fruiting June-October.
China (Chongqing, Fujian, Guangdong, Guangxi, Hong Kong, Hunan, Sichuan) (Map
The current authors have seen no specimens or specimen records from Hong Kong and inclusion of the species in that area follows
China. Chongqing: Jiangjin, Mt. Simianshan, Sunzigang, Z.Y. Liu, J. Zhang et al. S-2006 (IMC). Nanchuan, Mt. Jinfoshan, Sanquan, Lengshuixi, Z.Y. Liu 960468 (IMC). Fujian: Yunxiao, Xiaban, Mt. Dachenshan, G.D. Ye 2038 (PE). Zhangzhou, Yunxiao, Mt. Liangshan, Yunliang Reservoir, X.F. Zeng ZXF36083 (CZH). Guangdong: Gaoyao, at the foot of Mt. Dinghushan, C. Huang 161752 (IBSC). Ruyuan, Daqiao, Yue71 466 (IBSC). Guangxi: Jingxi to Longbang, R.P. Clark 422 (K, IBK). Liuzhou, Rongan, Banqiao, Guban Village, Rong’an Exped. 450224170806001LY (GXMG). Hunan: Yizhang, Changle, Mt. Xinpingshan, S.K. Lau 29560 (IBSC). Yongzhou, Jiangyong, Lanxi, Shangjin Village, X.C. Jiang, G.H. Tang & X.W. Pan SCSB-HNJ-0051 (KUN). Sichuan: Changning, Xiangling, Liushuiyan, s. coll. 704 (SM). Gongxian, Luobiao, Wangjia, s. coll. 278 (SM). Hongya, Liujiang, Shuguang, Laoyingzui, Hongya Group 420 (SM). Junlian, Tuanjie, Lüzhu Temple, Sichuan Economic Plants 0281 (PE). Leibo, Zhongshanping, Xining, Sichuan Economic Plants 487 (CDBI). Mt. Emei, Heilongjiang, K.T. Fu 12134 (WUK). Pingshan [Ping-shan], F.T. Wang 22721 (PE). Tongliang, Xiquan, Xiafeng, Tongliang Exped. 267 (SM). Xuyong, Shuiwei, Guandou Village, across the Qiaogoutou River, X.F. Gao, Y.D. Gao & W.B. Ju HGX10640 (CDBI).
Caesalpinia vernalis Champion ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 4: 77. 1852.
China. Hong Kong, Champion in Herb. Bentham 502 (neotype (designated by
Habit
a liana. Stems moderately to densely ferruginous tomentose, glabrescent when old, sometimes with scattered recurved prickles. Stipules triangular, 1–2 × ca. 1 mm. Leaves with 8–16 pairs opposite to strongly subopposite pinnae; petiole 1–2.5 cm; rachis 20–43 cm long, with a recurved prickle at the base of each pinna and scattered along the rachis between the pinnae insertions, moderately to densely ferruginous tomentose; pinnae 4.5–8 cm; leaflets 5–10 opposite pairs per pinna, coriaceous, elliptic to ovate, apex acute, mucronulate, terminal leaflets 1.4–2.8 × 0.5–1.5 cm, lateral leaflets 1.2–2.5 × 0.5–1.3 cm, both surfaces glabrous, or lower surface sparsely ferruginous tomentose, or only on midvein; venation reticulate, anastomosing, obscure. Inflorescence a raceme or many-branched panicle 12–35 cm long, in axils of upper leaves or terminal, axes and pedicels densely ferruginous tomentose; bracts not seen, bracteoles ca. 1–2 × 1 mm, apex acuminate, sparsely to densely ferruginous tomentose; pedicels 6–12(–16 in fruit) mm. Flowers with a hypanthium ca. 2 × 4 mm, this moderately to densely ferruginous tomentose; lower (cucullate) lobe ca. 7–11 × 4 mm, sparsely to moderately ferruginous tomentose on centre of outer surface, becoming glabrous towards the edges, other lobes ca. 6–12 × 2 mm, (very) sparsely ferruginous tomentose inner and outer surfaces sparsely ferruginous tomentose; median petal (6–)9 × 2 mm, inrolled, with dense circular patch of hairs at base of blade, and some hairs on claw, particularly on the margins; upper laterals ca. 9 × 3 mm, sparsely tomentose on inner surface of claw; lower laterals ca. 10–11 × 3 mm, sparsely tomentose on inner surface of claw; stamen filaments flattened, ca. 9–12 mm, pale orange tomentose on lower ca. 2/3 on inner surface; anthers ca. 1.5–2 mm long; ovary ca. 2.5 mm long, densely ferruginous tomentose, stipe ca. 1 mm long, style ca. 6 mm, glabrous, ovary 2-ovuled; stigma funnel-shaped, slightly papillate, ca. 1 mm wide. Fruit dehiscent, ligneous, obliquely oblong or sub-elliptic, slightly asymmetrical, apex beaked, venation obscure, sparsely to densely ferruginous tomentose, 4–6 × 2.5–4 × 1–1.3 cm, ventral suture lacking a wing. Seeds (1–)2, lunate, ca. 2.1–2.7 × 1.3–2.1 cm, matt black (Fig.
Moist sandy soils, beside rocks along valleys, in thickets; elevation ca. 600 m.
Flowering February-April, fruiting September-December.
China (Fujian, Guangdong, Hong Kong, Zhejiang) (Map
China. Fujian: Hua’an, Eshan, W.D. Han 20542 (NF). Yunxiao, Datian, G.D. Ye s.n. (FJSI). Guangdong: Baoan, Shatian, T. Chung M185 (IBSC). Chaochow, Raoping, Fenghuangshan, N.K. Chun 42662 (IBSC). Guangzhou, Conghua, Daling, Shanshizao, S.J. Li 787 (IBSC). Haifeng, Lianhua, Lügong, Dakeng, G.X. Chen 24 (IBSC). Huidong, Z.Q. Song 2021061 (IBSC). Luofushan, Sulao Taoist Temple, on the way to Dachashan, Y. Tsiang 1751 (IBSC). Meizhou, Fengshun, Yanping, Fengbei Village, X.F. Zeng ZXF01805 (CZH). Hong Kong: Lantau, Tung Chung, S.Y. Hu 12897 (PE). N. T. Lan Nai Chung, Sai Kong, S.Y. Hu 8570 (PE). Zhejiang: Huangyan, Western part, Shidun, N.Z. Wang s.n. (NAS). Jiande, Jiande Forest Farm, J. Zhao et al. 8524205 (PE). Jingning, Wangkeng, M.L. Yu et al. 25125 (HHBG). Jiansae, Laufuyoh, K.W. Jiang, J.P. Wu, Y.F. Zhang, M.S. Zhang et al. YS022 (NPH). ibid., Y.M. Zhang YS023 (IBSC). Linhai, Yongdongkou, Dakeng, s. coll. 196 (HHBG). Ningpo, Tientungssu, H. Migo s.n. (NAS). Suichang, Daban, Yakoumen, M.L. Yu 25756 (NAS). Tiantai, Gaoming, L.S. Que 28517 (ZM). Wencheng, Shuiyanhu, J.P. Feng 499 (HHBG). Yueqing, Dajing, Dianling, Dazhuyuan, mountainside, Hangzhou Botanical Garden Herbarium 2493 (HHBG).
Caesalpinia yunnanensis S. J. Li, D. X. Zhang & Z.Y. Chen. Novon 16(1): 78–80. 2006.
China. Yunnan, Xishuangbanna, T. P. Zhu (Zhu Tai-Ping) 139 (holotype KUN: [1206956!], isotype IBSC [0162107!]).
Habit a liana. Stems with recurved prickles to ca. 2 mm long, glabrous. Stipules caducous, not seen. Leaves with 3 pairs opposite pinnae, petiole 7–8 cm, rachis 15–20 cm long, rachis with recurved prickles at the base of each pinna and scattered in between the pinnae insertions, pinnae 5–6.5 cm long; leaflets 2–3 opposite pairs per pinna, blade coriaceous, elliptic or narrowly elliptic, base obtuse to cuneate, apex bluntly acuminate, margin incurved abaxially, 6–11.5 × 2.5–4.5 cm, upper surface glossy, lower surface dull, both surfaces glabrous, 2o venation anastomosing, 3o venation finely reticulate. Inflorescence an axillary raceme, more than 20 cm long; pedicels ca. 7 mm. Flowers unknown; receptacle remnant ca. 3 mm wide. Fruit dehiscent, ligneous, oblong to elliptic, slightly asymmetrical, base widely cuneate, apex with beak ca. 2 mm long, venation reticulate, not prominent; 5–7 × 2.8–3.5 cm, ventral suture lacking a wing. Seed 1.
Thickets along riversides, sparse woodlands along roadsides, elevation ca. 600 m.
Flowering unknown, fruiting October.
China (Yunnan) (Map
Specimen Chen 0066 which is listed as a paratype of T. yunnanensis has leaflets that are asymmetrical, with an asymmetrical base and distinctly acuminate tip, and the secondary veins are at a more acute angle to the midvein when compared with typical T. yunnanensis. It is sufficiently morphologically divergent from the concept of T. yunnanensis to be considered by the current authors to represent a different taxon (not determined), and the characters of that specimen are therefore not included in the description above.
China. Yunnan: Xishuangbanna, Mengla, Yiwu, J.H. Zhang 19335 (HITBC).
Guilandina rotunda Noronha, Verh. Batav. Genootsch. Kunst. 5(Art. 4): 16 (1790), nom. inval. nom. nud.
Butea loureiroi Spreng., Syst. Veg., ed. 16(3): 186. 1826. nom. inval. nom. superfl.
Caesalpinia scandens J. Koenig ex Baker in Hook. F., Fl. Brit. India 2(5): 255 (1878), nom. inval.
Although there are no fossils definitively attributed to Ticanto, some partial leaf fossils from the Upper Tochiwara Formation of Japan, dating to the mid Miocene, have been tentatively likened to Caesalpinia crista and assigned the name Caesalpinia hokiana
The protologue of Caesalpinia crista L. lists three type elements: Fl. Zeyl. 157; Pluk. alm. 4. t. 2. f. 2.; Breyn. ic. 58. t. 28. Of these, only the Flora Zeylandica element references material (in Hermann's Herbarium) attributable to Caesalpinia crista L. whilst the Plukenet figure and Breynius plate are representative of Guilandina bonduc L. (1753) (see
The impact of the application of the name Caesalpinia crista to two widespread species, both of which have medicinal uses, is significant. A wide range of studies record the presence of numerous phytochemicals in C. crista along with a wealth of reported pharmacological benefits including antioxidant, antibacterial, antiviral (including for treatment of Covid-19 [
The implications of misidentification of samples used to test for the presence of biologically active phytochemicals and to evaluate medicinal properties are clear. Taxonomic uncertainty could lead to false assumptions of the properties of a species or inclusion of the wrong species in medical preparations, and potential harm to human health. Reiteration of the correct application of the name Caesalpinia crista L. provided here will contribute to avoidance of this issue in future studies.
Genera segregated from Caesalpinia s.l. are most often morphologically characterised by fruit and floral characters, as well as by glands and trichomes (
The morphological resemblance of species now included in Ticanto was detected as long ago as the 19th century, when
The vegetative characters of Ticanto do not distinguish it clearly from Pterolobium, Mezoneuron and Biancaea, all being lianas or scandent or trailing shrubs armed with recurved prickles, and with pari-bipinnate leaves (Fig.
The flowers of all four genera can be considered as ‘typical’ Caesalpinia group flowers consisting of a short hypanthium and five sepals, the lower of which is cucullate over the others in bud, and five oblong, obovate or spathulate (occasionally bilobed) petals that are yellow or white, sometimes with red markings on the median petal, usually with some degree of pubescence. Stamens are free and tomentose, the ovary is glabrous or hairy, and the stigma is cupular, funnel-shaped or truncate, often papillate. The flowers of all four genera appear to be adapted for a range of generalist pollinators (mostly species of bee) and do not exhibit modifications related to novel pollination syndromes (although ambophily, i.e., pollination by both wind and insects, is reported uniquely in T. crista by
Like several other genera in the Caesalpinia group, Ticanto, and the genera to which it is most closely related (Pterolobium, Mezoneuron and Biancaea) are distinguished primarily by differences in fruit morphology. The fruits of Ticanto are elliptic, circular or lunate, compressed or inflated, coriaceous or ligneous, usually indehiscent (two dehiscent exceptions), 1–2-seeded, and with or without a narrow wing up to 4 mm wide along the upper suture (Fig.
Comparative characters of Ticanto species (characters considered to be most taxonomically informative are in bold).
T. caesia | T. crista | T. elliptifolia | T. magnifoliolata | T. rhombifolia | T. sinensis | T. szechuenensis | T. vernalis | T. yunnanensis | |
---|---|---|---|---|---|---|---|---|---|
Pairs pinnae | 5–8(–9) | 3–6(–8) | 1–2 | 2–3(–4) | 3–6 | 2–4(–5) | 3–6 | 8–16 | 3 |
Position of pinnae | opposite | opposite (occasionally slightly subopposite) | opposite | opposite | opposite | opposite | opposite (rarely subopposite) | opposite to strongly subopposite | opposite |
Pairs leaflets | 8–15 | 2–4(–7) | 2 | 2–3 | 3–5 | 2–5 | 3–6 | 5–10 | 2–3 |
Leaflet size | 0.8–1.5 × 0.4–0.6 | 2.1–7.2 × 1–3.3 | 7–13 × 4.5–8 | 3.5–10.8(–15) × 2.1–7 | 1.5–2 × 0.8–1.3 | 1.8–10.7 × 0.8–5.1 | 1.2–6 × 0.8–3 | 1.2–2.8 × 0.5–1.5 | 6.0–11.5 × 2.5–4.5 |
Leaflet apex | truncate, obtuse, rounded, emarginate | obtuse or rounded to retuse, acute, or acuminate | rounded, obtuse or acute | usually obtuse, rounded, retuse or emarginate, occasionally acute | acute | usually acuminate or acute, occasionally rounded | usually acute, occasionally slightly rounded | acute | bluntly acuminate |
Inflorescence indumentum | brown puberulent | glabrous or sparsely tomentose | densely hairy | sparsely to moderately tomentose | glabrous or subglabrous | sparsely to densely ferruginous tomentose | subglabrous to moderately tomentose | densely tomentose | unknown |
Inflorescence length cm | 10–15 | 8–40 | unknown | 15–30 | 10–20 | 7–42 | 11–15 | 12–35 | >20 |
Pedicel length mm | 4–7 | 5–15 | 8–12 | 5–11 | 6–9 | 3–12(–17 in fruit) | 5–11 | 6–12(–16 in fruit) | ca. 7 |
Bracteoles mm | unknown | 1 – 2.5 × 0.5 – 1 | unknown | unknown | unknown | broad, 2–3 × 1–1.5 | caducous, 1 × 0.25 mm, lanceolate-acuminate | 1.2 × 1 | unknown |
Median petal size mm | 3.5–5.5 (L) | 6–9 × 3–7 | ca. 10.5 × 6–7 | ca. 7–10 × 3–5 | ca. 7 (L) | 6–8 × 3–4 | 6–9.5 × 2.5–5 | 6–9 × 2 | unknown |
Lateral petals size mm | 3.5–5.5 (L) | 6–11 × 3–6 | 10–12 × 4–5 | ca. 7–10 × 3–5 | ca. 7 (L) | 6–10 × 2–6 | 5–10 × 3–4 | 9–11 × 3 | unknown |
Median petal indumentum (inner surface) | rhombic patch of hairs | dense circular patch of hairs in middle | circular patch of hairs | patch of hairs | patch of hairs | circular patch of orange hairs in middle, hairs on margins of claw | patch of hairs in middle; few hairs on margins of claw | dense circular patch of hairs in middle, some hairs on claw | unknown |
Stamen length mm | ca. 6 | 4 – 12 | 9–14 | 5–10 | 7 | 5–12 | ca. 9 | 9–12 | unknown |
Style length mm | ca. 4 | ca. 4–11 | 7–10 | ca. 5–10 | 10 | 6–12 | 10–12 | 6 | unknown |
Ovary indumentum | glabrous | glabrous or sparsely or patchily hairy | tomentose | glabrous | glabrous | sparsely to densely tomentose or glabrous | sparsely to moderately tomentose | densely pilose | unknown |
Fruit dehiscence | indehiscent | indehiscent | indehiscent | indehiscent | indehiscent | indehiscent | indehiscent | dehiscent | dehiscent |
Fruit size cm | 4.5–5 × 2.3–5 | 2.7–7 × 2.2–3.7 | 4.5–5 × 2.2–2.5 | 2.8–4.2 × 2.2–3.5 | 3–4 × 2.3–2.5 | 3–5.8 × 1.9–4.1 | 1.5–3.4 × 1.5–3 | 4–6 × 2.5–4 × 1–1.3 | 5–7 × 2.8–3.5 |
Fruit wing | narrow | absent | absent | carinate, 5–6 mm | absent | 0.5–4 mm wide | sometimes present, 1–3 mm wide, along part of fruit length | absent | absent |
Fruit shape | elliptic | elliptic to lunate, sub-symmetrical to somewhat asymmetrical | oblong-elliptic, sub-symmetrical to sub-lunate | lunate | asymmetrical, sub-lunate | Sub-circular to lunate | strongly asymmetrical, sub-lunate to sub-circular or teardrop-shaped | obliquely oblong or sub-elliptic, slightly asymmetrical | oblong to elliptic, slightly asymmetrical |
Fruit stipe mm | unknown | 2–5 | short | ca. 1 | ca. 3 | 0–2 | 0–1 | unknown | unknown |
Fruit beak mm | unknown | 1–10 | 1–5 | 2–7 | slight beak | pronounced beak to 25 | 1–5 | beak present | ca. 2 |
Fruit venation | prominent | prominent | prominent | prominent | prominent | prominent | prominent | not prominent | not prominent |
Fruit indumentum | glabrous | glabrous or very sparsely tomentose | unknown | glabrous | glabrous | glabrous or glabrescent | glabrous | sparsely to densely tomentose | unknown |
Fruit texture | ligneous | coriaceous | coriaceous | coriaceous | coriaceous | coriaceous | coriaceous | ligneous | ligneous |
Ecology & elevation | forests along rivers, 200–1000 m | often coastal; limestone, up to 350 m | ca. 100 m | forests, 400–1800 m | thickets | forest, thicket, limestone, 100–1500 m | forest, thicket, limestone, 260–1500 m | Moist sandy soils, thickets, ca. 600 m | unknown |
The fruits of Pterolobium bear some structural similarities to those of Ticanto in that they comprise a 1(–2)-seeded locule that is always (Pterolobium) or sometimes (Ticanto) winged, and the wing is non-vascularised (no venation is conspicuous on the exterior surface or in X-ray, Fig.
The fruits of Biancaea differ from Ticanto in that they are usually dehiscent and wingless (except B. decapetala which may have a narrow wing or ridge along the upper suture, and B. millettii which may have a very narrow wing along the upper suture) whilst those of Ticanto are usually indehiscent (with the exceptions of T. vernalis and T. yunnanensis) and often with a narrow wing along the upper suture, although some species are wingless (T. crista, T. elliptifolia, T. rhombifolia, T. vernalis, T. yunnanensis). The ovary indumentum of Biancaea is (densely) hairy, as opposed to glabrous or sparsely to (less commonly) densely hairy in Ticanto. The fruits of Biancaea are 2–8-seeded (apart from B. millettii with 1 seed), versus 1(–2)-seeded in Ticanto. Biancaea usually has large stipules, ranging from 3 mm to 4.5 cm long (except for B. millettii in which they are 2 mm long), whilst those of Ticanto are 0.25–3 mm long. Biancaea millettii has morphological affinities with Ticanto in having small stipules and single-seeded fruits that sometimes have a narrow wing, and its distribution (Guangdong, Guangxi, Hunan, Jiangxi in China) is congruent with the centre of diversity of Ticanto. The leaves of B. millettii, which bear numerous pinnae and numerous, small, oblong leaflets, resemble those of Ticanto caesia. The morphological affinities of B. millettii with T. caesia (including certain fruit characters), and the distribution of the former in southern China (the centre of diversity of Ticanto), raised the question of whether B. millettii might belong in Ticanto; however, the molecular phylogenetic analysis here presented demonstrates it to be correctly placed in Biancaea. The fruit of B. millettii, despite some similarity with Ticanto fruits, exhibits features typical of Biancaea fruits in being dehiscent and with a puberulent indumentum. This supports the hypothesis that fruit characters are important in delineating segregate genera of Caesalpinia s.l.
The distribution of Ticanto compared with closely related genera suggests it to be a distinct evolutionary lineage. The centre of diversity of Ticanto is Southern China, where all species are present and six are endemic (T. caesia, T. elliptifolia, T. magnifoliolata, T. szechuenensis, T. vernalis, and T. yunnanensis). Of the three remaining species, T. rhombifolia occurs also in northern Vietnam, T. sinensis extends into northern Laos, Myanmar and Vietnam, and only T. crista is more widely distributed throughout South-East Asia. The centre of diversity of its sister genus Pterolobium is South-East Asia (and a single species in Africa) from India through Myanmar into Indochina, and to Indonesia, Borneo, the Philippines, and Malaysia. Only two species of Pterolobium are found in China, namely P. macropterum Kurz, in Yunnan and Hainan provinces, and P. punctatum Hemsl., which is a broadly distributed Chinese endemic extending into at least nine provinces (International Legume Database and Information Service, https://ildis.org/LegumeWeb/). The distribution of Ticanto reflects its preferred ecological niche, characterised by a drier and more seasonal climate and scrub or dry forest habitat, and that of Pterolobium likewise reflects its preference for a warmer, moister climate and lowland forest habitat. Mezoneuron occurs (like Pterolobium) predominantly in the moist tropics, but with a more widespread and disjunct distribution (across South East Asia, two species in Africa, six species endemic to New Caledonia, one endemic in Madagascar, and one endemic in Hawaii), with only two species (M. cucullatum and M. enneaphyllum) present in southern China, represented by just a few specimens collected close to the borders. Of the six species in the Asian genus Biancaea, B. millettii is endemic to a few provinces in southern China, whilst the widely distributed B. decapetala is present throughout southern China, and the remaining four species occur in moist tropical areas from India to Thailand, Cambodia, Vietnam, and Malesia. Ticanto is therefore the only genus in the Caesalpinia Group of which most species occur either partially or exclusively in China.
We thank Hua-Fei Cen and H. Tang for organising and facilitating successful fieldwork in Guangxi, China. We are grateful to curators of herbaria A, CDBI, CSFI, CZH, FJSI, GXMG, GXMI, GZAC, GZTM, HGAS, HHBG, HITBC, IBK, IBSC, IMC, IMDY, K, KUN, L, MO, NAS, NF, NPH, NY, PE, PEY, SM, SN, SYS, SZG, TAIF, US, W, WUK and ZM for facilitating access to herbarium specimens, particularly Ma-Wen Zhang (‘Dennis’) at KUN. We greatly appreciate the valuable expertise of Dr Rafael Govaerts, Helen Hartley, Melanie Thomas, Dr Nicholas Turland, and Ian Turner in helping to resolve nomenclatural questions. We thank Dr Shi-Jin Li, Dr En-De Liu, Dr Zhu-Qiu Song, Dr Yong-Mei Yi, Dr Wan-Yi Zhao, Dr Zhi-Ming Zhong, Dr Xin-Xin Zhu, Miss Shi-Ran Gu, Miss Ding-Xiang Yu, Miss Ya-Min Zhang, and Mr Jian-Ping Wu for providing specimen material or high-quality photos. We are very thankful to Dr Tie-Yao Tu for supplying sequences of several accessions for the molecular phylogenetic analyses. The first author is particularly grateful to Laszlo Csiba, who gave training in laboratory techniques and helped to resolve many problems in the lab with endless patience and good humour. Many thanks to Dr Anne Bruneau and Dr Gwilym Lewis for very helpful and thorough reviews of the draft manuscript.
Appendix 1
Data type: Occurrences.
Explanation note: Table S1. Specimens used for mapping point localities.
Appendix 2
Data type: Docx file.
Explanation note: Table S2. Sequences obtained from GenBank.
Caesalpinia group Bayesian phylogeny
Data type: Phylogeny.
Explanation note: Caesalpinia group Bayesian phylogeny based on the combined dataset.
Caesalpinia group ML phylogeny
Data type: Phylogeny.
Explanation note: Caesalpinia group ML phylogeny based on the combined dataset, including the accession Kiyama et al. 1233, Ticanto magnifoliolata.