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The handle http://hdl.handle.net/1887/22521 holds various files of this Leiden University dissertation.

Author: Sirichamorn, Yotsawate Title: Systematics and biogeography of , Brachypterum and () in Asia Issue Date: 2013-11-28 Chapter 1 General Introduction -like -like ) and (Wight Derris seem to be seem like taxa Brachypterum -like taxa. -like Brachypterum Brachypterum according to Geesink Brachypterum Derris- Derris -like taxa are sometimes verytaxa are sometimes -like as many of as occur usually them (not including Brachypterum Derris Lour. [including Lour. Derris -like taxa. Up to now every researcher every researcher taxa. Up to now -like ). Paraderris Derris Derris Aganope : Systematics and Biogeography: Systematics Chapter 1 - Miq., s.s. and s.s. (Miq.) Geesink. (Miq.) Derris Derris Aganope and Paraderris are lianas, which may climb over trees or other vegetation formingor trees other vegetation climb over may which are lianas, (Fig. 1-1: A). However, the tree and tree-like habit can also be found also be habit can tree-like and tree the However, A). 1-1: (Fig. Derris prefer sunlight and water. Many species growMany species exposed naturally in fully and water. sunlight prefer -like taxa” contain members of-like characterized (Fabaceae), tribe the -like taxa are common weeds in forest plantations (Fig. 1-1: E). in forest plantations (Fig. taxa are common weeds -like

Aganope, Brachypterum Brachypterum Aganope,

Derris in dry deciduous forest, along the roads or in dry, very areas (Sirichamorn, disturbed in dryroads along the or forest, in dry, deciduous TheD). growspecies on a range C, 1-1: of personal observation; Fig. soil types ofSome species (Hamid, 1999). varying clay from coarse sand to heavy the Asian Derris supply such as limestone mountains (Fig. 1-1: B). Species ofSpecies B). 1-1: mountains (Fig. as limestone supply such than more drought tolerant Paraderris A), generally(not higher 1-1: altitudes at low (Fig. or along waterways areas nearby in dryerare specialized species A few m). than 500 water a limited habitats with a thick cover cover a thick placed in previously were which or (2 3 species and ofa species (1984), single Asian indehiscent pods. The have a pantropical distribution. According to the most the to According distribution. a pantropical The have plants pods. indehiscent genericthe Asian recent Adema (2000), circumscription proposed by oftaxa consist genera the & Arn.) and Benth.] ofsimilar because also show Some species morphological features. their overlapping This the distribution. and a wide in morphological makes characters a high variation plants sometimes difficult to identify correctly to the species level. Mostof species Asian General morphology, ecology and utility of the Asian morphology, General “ by their imparipinnate with leaves opposite leaflets and typical flat, usually winged, various various levels and from different viewpoints. The and morphologicalon molecular based their phylogeny be data will which after species will be defined first, inferred. Thewill form phylogeny resulting for the basis a new and subjective less classification. theFinally, biogeographic history of the taxa willbe analysed. This generalintroduction provides informationof palaeotropic Leguminosae (Fabaceae) concerns the (Fabaceae) Leguminosae others divided genus, all taxa into a single unite would some solutions, had different genera. Theinto several them purpose of problems the at tackle is to this thesis General IntroductionGeneral One of the major problems left in the classification of tribe Millettieae of the

General Introduction

2 1 General Introduction 1 3 Derris . B (Gagnep.) Craib, a Craib, (Gagnep.) -like taxa which usually taxa which -like Benth., a common weed a common weed Benth., (Roxb.) Benth., a species Benth., (Roxb.) Derris Derris thorelii . D Derris amoena . Derrisscandens E . C tree. Hevea (Wall.) Adema, typical for most offor most typical Adema, (Wall.) General Introduction General Paraderris elliptica Paraderris . , grows also in a disturbed area, roadside. , growsarea, roadside. also in a disturbed A species inhabiting limestone mountains. mountains. inhabiting limestone species , grows in a very area. grass disturbed, covered Derris -like taxa: -like Brachypterum Brachypterum Derris Habitat of Habitat Fig. 1-1. Fig. and forming other trees cover. climbing over a thick or along waterways, grownearby exposed areas in fully ofan example sp., few the previously placed in in possibly placed species the ground and climbing up a in para-rubber covering plantations,

, . and C and . Reddish . Reddish B Aganope

Paraderris Derris scandens Derris . to be transfered be to D Brachypterum Hemsl., an example Hemsl., or no residues. or no residues. and . The flowering Theflowering . Derris Paraderris. Lour. Lour. . Theyare usually or Derris Derris Derrisalboruba Paraderris . (Roxb.) Benth. have a true have panicle, Benth. (Roxb.) . Species ofSpecies . Aganope G A (Fig. 1-2: C, D): flowers in flowers D): C, 1-2: (Fig. Derris trifoliata . C and . -like taxa: -like Derris Aganope Derris marginata Paraderris . is a pseudoraceme/pseudopanicle type, characterized characterized type, a pseudoraceme/pseudopanicle is Derris F and . Up to 41 leaflets per leaf in per found be leaflets can 41 to Up . (Fig. 1-2: A). Reddish or golden-brownish Reddish A). 1-2: (Fig. sp. and sp. Paraderris Paraderris Derris , but absent in absent , but . Leafletsvary in shape and size, but all have and Derris have this type oftype this inflorescence. have . : Systematics and Biogeography: Systematics Chapter 1 - F Aganope E and Derris Paraderris E Aganope Derris Brachypterum or but monadelphous with basal fenestrae in basal fenestrae monadelphous with but s.s. and s.s. (Miq.) B.D. (Miq.) Jacks., B.D. usually have more leafletsthan and Derris Aganope Derris -like taxa are well known as an importantknown ofsource taxa are well -like rotenone D. microphylla D. Derris e.g., e.g., , only two species shown in shown species only two , . Pods are indehiscent, typically thin and leather-like (derris, Greek: skin, Greek: (derris, thin and typically leather-like are indehiscent, . Pods Derris . Some examples ofexamples Some . found in morphological characters

Brachypterum, Aganope, Brachypterum Brachypterum Aganope,

), on which the fascicles of flowers are born. (Roxb.) Benth. (Roxb.) The inflorescence typical for ofpresence the by arrowsred in by (indicated “brachyblasts” called axes lateral wartreduced, the or knob-like D the absence of by characterized are born This and thus the flowers solitarily on elongated lateral branches. brachyblasts rare in is of an intermediate inflorescence which basally elongated lateral resembling branches, a true panicle, but higher up the and resembles a pseudoraceme. bears only brachyblasts rachis Fig. 1-2 Fig. to in present are often leaves young range of Rotenone or as dust, dips or baits. in sprays, forms, i.e. crops in various little rapidly biodegradesand leaves air and sunlight to exposed when toxin, which occurs in every occurs part, which in stems but in higher amounts especially toxin, and insecticide organic and unselective as an effective, is used Rotenone and roots. piscicide (fish poison). It can thus be appliedto control a large array of pests on a in some species (Fig. 1-3: G to I), which were previously included in in included previously were which G to I), 1-3: (Fig. species in some The wings of or pods can be found along upper suture along both sutures; the the hilum are found in eccentric a distinctly with Seeds pods are rare. wingless hilum are found in or eccentric a central slightly prefer others while season, rainy the in flower species Some species. per varies time drier According conditions. to field observations, thespecies are mostly pollinated bees. by ofSpecies are diadelphous in Paraderris The leather coat). shape of are found chambers Seed A to I). 1-3: (Fig. pods varies the an entire margin. Stipellae are generally absent but usually present in species which are generallyStipellae an margin. entire which in species present usually absent but true in are panicles Inflorescences one-wingedpods. have in or pseudopanicles pseudoracemes 1-2: (Fig. rare. are panicles on veryfascicles “brachyblasts”; shortcalled axes lateral intermediateE, F); true between G) 1-2: forms and pseudoracemes/ panicles (Fig. Theuncommon. but are also found, pseudopanicles or reddish, calyx is usually greenishsometimes or pinkish rarely Thewhite, are typically petals short with lobes. Thedeep stamens purplish, with or basal on without callosities the standard petals. species previously placed in placed previously species of species in many found be can B) 1-2: (Fig. leaves young absent in but this is usually As mentioned they earlier, all imparipinnate with have leaves opposite The leaflets. number of leaflets per leaf varies ofspecies found in from threeto nine, which is most frequently

General Introduction

4 1 General Introduction 1 5 General Introduction General : Systematics and Biogeography: Systematics Chapter 1 - Derris and

Aganope, Brachypterum Brachypterum Aganope,

General Introduction

6 1 General Introduction 1 7 . G D. Aganope Aganope (Miq.) . C (Roxb.) (Roxb.) Paraderris Paraderris (Roxb.) Benth. Benth. (Roxb.) itself are used D. scandens D. (Benth.) Polhill. Polhill. (Benth.) D. microphylla D. F.C. How. Seed chambers can be can chambers Seed How. F.C. Derris ferruginea P. elliptica P. . is the main species that species main the is F Graham ex Benth. is used to used Grahamis ex Benth. Craib. Craib. Aganope Aganope thyrsiflora . Derriseriocarpa B P. elliptica P. . I Lour. Lour. D. elegans D. Derris reticulata have also been reportedalso been and as piscicide have . E (Hamid, 1999). Derris Derristrifoliata . -like taxa, many species are also used traditionally are also used taxa, many species -like A (Roxb. ex DC.) Benth. DC.) ex (Roxb. General Introduction General Lour. (Fig. 1-2: C and 1-4: C) is used in India and used is C) C and 1-4: 1-2: (Fig. Lour. and Smith-Vaniz, 2008). Traditionally, the tuba root the Traditionally, 2008). and Smith-Vaniz, (Roxb.) Benth., Benth., (Roxb.) Derris -like taxa: -like Derris robusta . Derris -like taxa are also reported. For example, species which are which species example, taxa are also reported.-like For D. trifoliata D. H (Roxb. ex DC.) Benth. (Fig. 1-4: E, F) and E, F) 1-4: (Fig. Benth. ex DC.) (Roxb. Derris marginata . (Wall.) Adema, see Fig. 1-1: A and 1-4: A] and tuba merah [ A] and tuba A and 1-4: 1-1: Fig. see Adema, (Wall.) Derris D . Scale bar = 1 cm. (Roxb.) Benth. (Roxb.) D. robusta D. I , . Craib (Fig. 1-4: D) contain some sweet chemical substance and are used as and are used substance chemical sweet contain some D) 1-4: Craib (Fig. (Benth.) Adema] have been used as a fish poison throughoutSE Asia and (L.) Polhill. Polhill. (L.) to Variation in pods of Variation G Paraderris ellipica Paraderris Fig. 1-3. Fig. heptaphylla Other uses ofOther uses i.e trees, for and sometimes and shade trees as timbers are cultivated A), 1-2: (Fig. Jacks. B.D. Thailand.of A solution of crushedleaves Guinea. New bites in Papua snake wash Benth. (Fig. 1-1: C, 1-2: D and 1-4: B) are used as a diuretic, laxative, expectorant, laxative, a diuretic, as are used D and B) 1-2: 1-4: C, 1-1: (Fig. Benth. emmenagogueofand treatment the in common cold and backache. osteoarthritis, ofRoots after childbirth.in India milk secretion are also used to increase this species Theofplant whole and counter-irritant, and against antispasmodic, rheumatism, Indonesia as stimulant, and dysmenorrhea. Thai paralysis, roots or traditional doctors used stems chronic and anti-arthritis carminative, ofStems 1999). (Hamid, treatment as a laxative, reticulata in as an are also used expectorant or they a laxative in local medicine; a sweetener Because ofBecause of toxicity the In Thailand roots ofthe in many countries. as medicines of Stems as emmenagogueas a blood tonic. and stems the [ montana other Several species ofthe Pacific. However, insecticide. an as used are also locally ofa major source as cultivated widely is ofcultivars Several rotenone. species this of been selected have and “Ngawi” are Creeping”, “Changi No.3” “Sarawak which reported superior to be commercially used by ichthyologists for collecting fish in their studies of studies their in fish collecting for ichthyologists by used biodiversity, fish marine to minor effects side only are necessary small quantities only because and have (Robertson environment the The chemical is extremely toxic to insects and fish becauseonly slightly but respiration, cellular with interferes or and gills trachea through the it is easily absorbed of use Although the (Hamid, 1999). to mammals toxic as an rotenone insecticide Netherlands the e.g. countries in some banned been and has out being phased is introduction of to the due about 1980 since ofand lack the insecticides synthetic determinationdata about convincing of continuously rotenone has been toxicity, its Derrisscandens seen in

. to -like -like Solori . The Derris Derris Aganope Brachypterum Paraderris (Roxb. ex DC.) Benth. ex DC.) (Roxb. (Hooker, 1849) (Hooker, Adans. and Adans. and Salken Lour., as medicine and occasionally as medicine Lour., Derrisrobusta Hook.f. was described was Hook.f. . -like genera,-like together with F (Wall.) Adema, as toxin and medicine. and medicine. Adema, as toxin (Wall.) s.l., which was divided into divided was which s.l., , E (Taub.) Mendonça & Sousa] (Taub.) Dipteroderris Derris ), and other related genera Derris

Derris trifoliata . can also be grown as green manure . As the genus has indehiscent pods, pods, genus indehiscent . As the has C Ostryocarpus ther leguminous plants which usually which plants leguminous ther Paraderris elliptica Paraderris but the name was later transferredname was but the to . which is full of is full which blossoms white-pinkish Derris Euderris A -like tree species found in arid areas of species tree -like ( Derris in the same year. Bentham (1860), in his (1860), Bentham year. same in the X. stuhlmannii D. trifoliata D. Derris Derris Dunn (1911) for African species of species for African Dunn (1911) : Systematics and Biogeography: Systematics Chapter 1 - . Later, Roberty (1954) described the monotypic the described (1954) Roberty . Later, Pterocarpus , D. scandens D. Paraderris Derris Derris Fig. 1-4: B). Like o Like B). 1-4: Fig. -like -like species which have true paniculate inflorescences. sect. sect. Lour. and Lour. Ostryoderris Dunn was established in 1910. One year later Dunn also later year One in 1910. established Dunn was and species have several characters that deviate from the other other from the deviate that characters several have species L.f. Three raised to generic years later the subgenus was L.f. -like taxa with various uses: taxa with various -like Derris Brachypterum , Derris Derris (Dunn) Roberty [≡ Derris was established by Joao de Loureiro in 1790 to accommodate two two accommodate to in 1790 Loureiro de Joao by established was Craib, as medicine and sweetener in medicine recipes. recipes. in medicine and sweetener as medicine Craib, to accommodate a accommodate to Leptoderris Benth., as toxin and stems ocessionally as rope. Benth., as toxin was established by Aublet, later used by Taubert (1891) for (1891) Taubert by later used Aublet, by established was Aganope Derris pinnata (Roxb.) Benth., as medicine and ornamentalas medicine Benth., plant. (Roxb.) Derris X. chevalieri Xeroderris Derris reticulata Deguelia . . Species of. Species Asian Derris amoena D .

G Derris scandens Aganope, Brachypterum Brachypterum Aganope,

. B toxin. and Fig. 1-4 Fig. “Synopsis of all old-world the united Dalbergieae”, genusinto a single called American species, some sections: five in a wide sense (s.l.). Wight and Wight Arnottsubgenusdescribed (1834) (s.l.). in sense a wide of genus the genusthe In 1849, 1837). (Bentham, level genusthe who established (1855), Miquel by followed 1849), (Hooker, several accommodate He also recognized upper filaments uniting with others. Before being describedby Loureiro in 1790, it the name gave Linnaeus (1747) another group ofdescribed Adanson (1763) Leguminosae. The Taxonomic History ofThe Taxonomic the genus Thegenus species: new already he However, (1860). Bentham by Dalbergieae tribe the in placed firstly was it the that remarked Dalbergieae, for example, the inflorescence, adhesion ofwings andkeel petals and contribute to nitrogen fixation, species of species fixation, nitrogen to contribute 1997). soil (Hamid, 1999; Thothathri Rugayah, and to improve ornamentalpurposes (Thothathri, Hamid, 1997; Thothathri 1982; Rugayah, and as such 2002) al., et Puy Du 1999; ( season flowering its during African genus another genus described plants, which have inflorescences similar to those of of to those pods similar but genus Africa, Adans., both based on plates in Hortus Malabaricus (van Rheede, 1686; 1688). In 1688). 1686; Rheede, in Hortus on plates both based Malabaricus (van Adans., 1775,

General Introduction

8 1 General Introduction 1 9 General Introduction General

; as -like -like . The under Harms and the together Wight & Leptoderris Derris Aganope Philenoptera Derris should be (the new- (the Lonchocarpus and section Derris Xeroderris (Bentham, 1860 (Bentham, Derris are distinct from are distinct is to be preferred Deguelia Ostryoderris Euderris had to be reunited and Benth. of Benth. genus the Margaritolobium and s.l. to generic level, i.e., i.e., to genericlevel, s.l. Deguelia Deguelia Philenoptera ). He also reinstated ). Brachypterum . In contrast to Polhill, Adema . In contrast to Polhill, Fasciculati Malme, Malme, Derris to genus as proposed by Polhill Polhill to genus as proposed by Aganope Brachypterum apart. Later Polhill (1981) and (1981) apart.Later Polhill and because of because typical its also shows morphological overlap morphological also overlap shows L.f. (Geesink, 1984) and 1984) (Geesink, L.f. . In also addition, Adema concluded Wight & Arn.), also shows a history also shows of synonymisation Paraderris Derris Aganope Aganope Deguelia Dahlstedtia in a taxonomic broad ( sense in a taxonomic Xeroderris Ostryocarpus Muellera is possibly correct. Aganope to generic level and also transferred to generic level species Ostryoderris, Ostryocarpus , and similarly with Lonchocarpus s and : Systematics and Biogeography: Systematics Chapter 1 - ” was not closely related to Asian related not closely ” was with , including section , including section Lonchocarpus Brachypterum Salken Aganope Lonchocarpus . According to Geesink the name the Geesink According to . Derris Lonchocarpus Leptoderris s.s. (old-world species of species (old-world s.s. section Deguelia (formerly section Euderris Xeroderris “ and Ostryocarpu (Macbride, 1943), (Macbride, -like genus -like Derris Derris Ostryocarpus . Accepting. Paraderris Derris Derris , from tribe Dalbergieae to tribe Millettieae, because they showed a showed they because Millettieae, tribe to Dalbergieae from tribe , , but kept , but kept and this generic circumscription is still widely accepted nowadays. generic and this acceptedwidely is still circumscription nowadays. Kunth), (previously section (previously ) and . He partially accepted Polhill’s (1971) concept by considering concept(1971) by He partially. accepted Polhill’s . Adema (2000) carried. Adema (2000) on analysis based morphological out a cladistic Derris Millettia with panicles to it. Polhill also combined also Polhill to it. panicles with Solori Derris Aganope

Derris Aganope, Brachypterum Brachypterum Aganope,

inclusion of genus this with The neotropical genera. Thevarious mergedwith genuseither was with 1943), Macbride, a distinct genusto a distinct related closely though proposed to synonymise American that the the oldest name oldest the older name the conservedover over Thesuggestedresults characters. genusthe that with with some other generawith of e.g., Millettieae, and even genera,a problem causes comparable with several including thus s.l.), is difficult result will be a morphologically heterogeneous genus that, consequently, (1984), Geesink to According diagnose. to Lonchocarpus of phylogenies and classification the the not are neotropical genera However, these main focus of this thesis. Fenzl. ex A. Rich. (Bentham, 1860). (Bentham, ex A. Rich. Fenzl. (1971), Geesink synonymized it with synonymized Geesink (1971), close morphological, anatomical, and chemical resemblance with resemblance morphological,close anatomical,and chemical ofstudy Arn. genera. and related his intensive After Geesink tribe Millettieae, the many ofreinstated (1984) of sections Bentham’s Brachypterum of species world section Lonchocarpus Dipteroderris of Instead to generic level. raising section into Geesink (1981, 1984) officially transferred many generaincluding with indehiscent pods, Hutchinson (1964) united united (1964) Hutchinson reinstated Polhill In 1971 pods. of

General Introduction

10 1 General Introduction 1 11

-like -like and Paraderris Derris -like taxa -like s.s. also show also show s.s. Ostryocarpus , are introduced, are introduced, , -like taxa because taxa -like , s.s. and s.s. Derris Derris

were also not closely also not were Derris Derris , P. elliptica P. Ostryoderris (Roxb. ex DC.) Benth.], was was Benth.], DC.) ex (Roxb. Philenoptera in those analyses) appeared to appeared analyses) in those and allies. and allies. Brachypterum Derris in a narrow sense next to various genera, in a narrow next to various sense , for example , (Hu et al., 2000 ) and nuclear ITS/5.8S and nuclear al., 2000 ) et (Hu Derris robusta Ostryocarpus in a wide taxonomic sense including many sense taxonomic in a wide Derris s.l. (sensu Bentham, 1860) was a polyphyletic a polyphyletic was 1860) Bentham, (sensu s.l. -like taxa -like and the African African and the Paraderris Derris trnK/matK General Introduction General Derris were not included in those studies. Apparently, the Apparently, studies. in those included not were Derris (s.s.) in all phylogenies. Up to now, all phylogenies could all phylogenies now, Up to in all phylogenies. (s.s.)

(treated as (treated Derris . However, even in the sense of sense in the even . However, and Adema (1984a) Geesink (Roxb.) Geesink [= Geesink (Roxb.) Lonchocarpus Derris Aganope Derris and its closely related African generaAfrican related closely and its B. B. robusta s.s. was probably also not monophyletic, because a representative of a representative because monophyletic, also not probably was s.s. -like taxa. A few species (not more than three species for each genus) for each species three than more (not species A few taxa. -like , are also still obscure and make it impossible to draw final conclusions Derris Aganope Derris -like -like taxa. The first one is conclusion, there are two generic/infragenericcompeting there are conclusion, two of circumscriptions sources of phylogenetically informative data with sufficiently sampled species can an exact circumscription ofprovide Asian remains far from solved. In addition, the relationships between the Asian-African Asian-African the relationships between addition, the In remains far from solved. genus Xeroderris of species Many regardswith generic to their circumscriptions. morphological similarities among them, which sometimes oftaxa. Species the distinguish makes it difficult to widely cultivated and then finally or natural morphological formscultivars leads to a possible consequently which naturalised. The species consistsinfraspecificclassification. of several More detailed phylogenetic studies based on various molecular phylogenies. The is second molecular phylogenies. of phylogeny the preferred is the which However, option. phylogenetically three the and morphologicallygenerarelated closely not equivocally solve the generic the among the circumscriptions solve equivocally not of insufficient sampling. Thusthe problem of generic analysis. phylogenetic delimitationimproved still needs an Problems of a generic/infrageneric circumscription of In Derris ofresults the by genera already refuted and infrageneric was taxa, which the (2000) (2000) Brachypterum outside placed always were included in the phylogenetic studies of studies phylogenetic in the included were al. (1998) et Lavin by Millettieae tribe the representing taxa different small samples, the Besides 2002). al. (2000, and Hu et infrageneric groups of on based phylogenies resulting that revealed al., 2002) et (Hu The genus taxon. far separate other from the phylogeny, Millettieae in the be a basal clade Thetaxa. neotropical to Asian related reflect gene-levelchanges, which are believedto reflect truephylogenies.However, of species the to related directly studies phylogenetic molecular only few are there Asian Molecular phylogeny of phylogeny Molecular ofSystematics as these data on DNA sequence based organisms living nowadays are - Derris -like taxa -like Derris sensu Geesink sensu Adema, Derris and its related genera, Derris -like taxa. Within the Indo- taxa. Within the -like , Derris -like taxa occur in Thailand,taxa occur 17 -like -like taxa occur in Thailand?-like Derris Derris Derris Derris -like taxa have been collected, but the but collected, been taxa have -like Derris s.l. According to the most recent preliminaryrecent most According to the s.l. : Systematics and Biogeography: Systematics Chapter 1 - Derris -like taxa in Thailand -like Derris and Derris 1) How many and which species of many and which 1) How do they differ morphologically and ecologically? WhatHow are their diagnostic morphological characters? 2) Is the most recent generic concept of Ifmonophyletic? of about the monophyly not, how or other previous generic concepts?

Aganope, Brachypterum Brachypterum Aganope,

Research Questions, aims and outline of Questions, aims and outline Research the thesis questions are addressed: The research following especially the rare, endemic or endangered species. especially the rare, observationof ofspecies 16 (2000), Adema more and goes As time by, are found in Malesia and nine are common to both areas. more unidentified Thai specimens of identification is still dependent on the literature of neighbouring countries such as Flore du Cambodgethe du Viêtnam (Phan and and du Laos et Vidal, 2001) Flora ofThe al., 2008). et of study China (Wei more substantial provide plants will these and for offundamental data the study the to directly It will contribute the Fabaceae. Flora of Thailandnecessarythe and Flora Malesiana and also provide information for sustainable managementand conservation of Thai or not completely finished. The first taxonomic account of Asian of species 18 in ThailandAt that time prepared Craib (1928). by was Asian under recognised taxa were like the Indochinese region in the north region in the (Mackinnon Indochinese south the region in the Sundaic and the ThailandBecause 1997). crossroads at the lies Mackinnon, 1986; and Mackinnon, regions ofthese between the composition of Realm, the lndo-Malayan the Thai genus or species at However, of that countries. resembles neighbouring closely flora countrythe in Thailand.endemics are some there Thus, as considered be could level ofcenter collective a perfect of botanical diversity Indo- 3 major regional elements: study the However, al., 2001). et Burmese, Welzen Indo-Chinese and Malesian (van ofof especially carried slowly out this area is lacking, in plantdiversity, Fabaceae, zone where a variety of variety a zone where are able to found and hence, can be tropical ecosystems supportof a large variety including plant species, majorbiogeographical countrythe realm, two Malayan regions, between situated is Species Diversity ofDiversity Species ThailandSE Asia mainland, a hot and the humid climate within located is centrally

General Introduction

12 1 General Introduction 1 13 , and like like , related trnK-matK . Vegetative . Vegetative -like taxa and -like Derris- Paraderris Derris Brachypterum Paraderris , and ) and Aganope ). Sequence data ofSequence ). other genera in Brachypterum -like taxa? -like Derris s.s. (Adema’s genericconcept) using is tested (Adema’s s.s. , Brachypterum, Derris, Brachypterum, s.s. Derris -like taxa in Thailand is carried out, using Adema’s taxa in Thailand-like is carriedusing Adema’s out, Derris spacer, and one nuclear marker, ITS/5.8S, were were ITS/5.8S, marker, and one nuclear spacer, General Introduction General s.s. (including (including s.s. Derris -like taxon is reported, taxon of -like together with the non-monophyly Derris , like taxa is highlighted. Three chloroplast markers, Three taxa is highlighted. markers, like chloroplast Aganope, Deguelia, Leptoderris, Deguelia, Lonchocarpus, Aganope, Ostryocarpus and ? Derris trnL-F intergenic other and to other generaother and ofparticularly Millettieae,

Derris- -like taxa? How can we explain the paleotropical intercontinental explain can we taxa? How -like ? Aganope the monophyly of monophyly the , a revision of , and -like clades. clades. -like Derris species ofdisjunctions (PIDs) found in some Derris taxa such as taxa such Philenoptera are the infrageneric4) How groups proposed for the genus previously other? Are those groups and does the related to each monophyletic corroboratemolecular phylogeny these or other infrageneric groups? be classified as 5) Can the clades genera or infrageneric they taxa? Are explain the trends in recognizable morphologically can we how and ofmorphological evolution occur in the events 6) Where major diversification and when did 3) How are the three genera are the 3) How to each Derris chapter 2 chapter 3

sequenced for all generasequenced of palaeotropic (Asian and African) generaneotropic also some (American in analyses in the are also included obtained from GenBank tribe Millettieae the of all taxa. Monophyly relationship between order phylogenetic to understand the palaeotropic each and field observations. All literature related toand are described species new this currentlyTwo are enumerated. species recognised genus reviewed is again. All generatogether to the descriptions are provided, with Keys and species illustrated. and notes for all taxa. In of phylogeny the especially chapter, In this approach. phylogenetic a molecular old world the psbA-trnH descriptions and a data matrix for future phylogenetic analyses. Geographical and analyses. descriptions and a matrix data for phylogenetic future ecological data, such as floweringperiod, are obtained from herbarium specimens The basis for all research are the ofand space in time evolution no idea about the have still would we those Without herbarium collections and field observations. the In concepts of in the examined and used morphological are carefully characters and reproductive , . The. Aganope Solori against s.s, by reinstating the genusthe reinstating by s.s, are not yet validly described, validly are not yet

. Characters supporting each 4 -like taxa. More gainedinsight is -like Brachypterum Derris Derris according to the results ofresults according to the the Chapter Derris with Paraderris : Systematics and Biogeography: Systematics Chapter 1 - Paraderris Derris and s.s (including s.s Derris and by synonymising synonymising and by and is the proposal to conserve the is name the presents the integrationpresents of molecular and morphological data as total

deals with the historical biogeographythe deals with of genera, three namely 3 and 4). Molecular dating were performed in a Bayesian framework with performed Molecular dating were 3 and 4). framework in a Bayesian parsimony-based S-DIVA and likelihood based analyses under the dispersal- the under analyses based and likelihood S-DIVA parsimony-based s.s. and the infrageneric classification previously proposed. Characters

Aganope, Brachypterum Brachypterum Aganope,

the aid of and ancestral area reconstruction (AAR) analyses program the BEAST, using extinction-cladogenesisprogramsthe implemented model, and Lagrange RASP combinations proposed As the in respectively. and quotation marks “...” in this Chapter are in between names used species new the without author name. Chapter 6 Brachypterum Chapter proposed in order to maintain the monophyly of monophyly the maintain to proposed in order Brachypterum and treatment nomenclatural changes are made genus are presented in a taxonomic where necessary. Chapter 5 forgottenlong is it but used, be and should name and available oldest the is latter described in it. species was a single used, not even never Chapter 4 reconstruction for a phylogenetic evidence of in the evolution of specific morphological A traits. new generic circumscription is Derris supporting briefly discussed. clade are each

General Introduction

14 1