Peter C. van Welzen

Molecular phylogeny of Macaranga, Mallotus, and related genera (Euphorbiaceae s.s.): insights from plastid and nuclear DNA sequence data

Macaranga and Mallotus (Euphorbiaceae s.s.) are two closely related, large paleo(sub)tropical genera. To investigate the phylogenetic relationships between and within them and to determine the position of related genera belonging to the subtribe Rottlerinae, we sequenced one plastid (trnL-F) and three nuclear (ITS, ncpGS, phyC) markers for species representative of these genera. The analyses demonstrated the monophyly of Macaranga and the paraphyly of Mallotus and revealed three highly supported main clades. The genera Cordemoya and Deuteromallotus and the Mallotus sections Hancea and Oliganthae form a basal Cordemoya s.l. clade. The two other clades, the Macaranga clade and the Mallotus s.s. clade (the latter with Coccoceras, Neotrewia, Octospermum, and Trewia), are sister groups. In the Macaranga clade, two basal lineages (comprising mostly sect. Pseudorottlera) and a crown group with three geographically homogenous main clades were identified. The phylogeny of the Mallotus s.s. clade is less clear because of internal conflict in all four data sets. Many of the sections and informal infrageneric groups of Macaranga and Mallotus do not appear to be monophyletic. In both the Macaranga and Mallotus s.s. clades, the African and/or Madagascan taxa are nested in Asian clades, suggesting migrations or dispersals from Asia to Africa and Madagascar.

Macaranga and Mallotus species (Euphorbiaceae) as indicators for disturbance in the mixed lowland dipterocarp forest of East Kalimantan (Indonesia)

The indicator value (IV) of Macarangaand Mallotus species (Euphorbiaceae) for different types of disturbance in lowland dipterocarp forest was assessed by counting and identifying all individuals of species of these genera taller than 30 cm in 45 (10 m × 300 m) plots at nine locations. Twelve Macarangaand nine Mallotus species were found. The main forest disturbance types (primary forest, secondary forest, selectively logged forest, forest burned once, and repeatedly burned forest used for shifting-cultivation) each had their own set of indicator species. The level of disturbance in the forest types was assessed by measuring nine forest structural parameters. The occurrence of Macarangaand Mallotusspecies was closely related to the level of disturbance in a forest. Most Macarangaspecies were characteristic of high disturbance levels, while most Mallotusspecies preferred intermediate to low levels of disturbance. However, both genera had species at both disturbance extremes. Using multiple regression analysis, combinations of Macarangaand Mallotus species were formed and used to predict the separate forest structural parameters and the general level of disturbance of a forest. The Macaranga and Mallotus species could be grouped into (1) primary forest ‘remnant’ species; (2) generalist pioneer species; and (3) high disturbance pioneer species.

Historical distribution of Sundaland’s Dipterocarp rainforests at Quaternary glacial maxima

Significance The effect of glacial cycles on Southeast Asian (SEA) rainforest during the Quaternary is unresolved. Some historical evidence suggests rainforests were confined to small refugia during glacial maxima, but dynamic vegetation models suggest evergreen rainforests were widespread. Because Dipterocarpaceae dominate current SEA rainforests, their distributions closely reflect general rainforest extent. Here, we use an extensive georeferenced database of collection records for 317 Dipterocarpaceae species to model their climatic niches, based on current climatic conditions. These distribution models were then hindcast onto historical climatic conditions of the last glacial maximum. The results indicate that central Sundaland, exposed because of lower sea levels at glacial maxima, harbored suitable environmental conditions for Dipterocarpaceae and was probably covered by rainforest. The extent of Dipterocarp rainforests on the emergent Sundaland landmass in Southeast Asia during Quaternary glaciations remains a key question. A better understanding of the biogeographic history of Sundaland could help explain current patterns of biodiversity and support the development of effective forest conservation strategies. Dipterocarpaceae trees dominate the rainforests of Sundaland, and their distributions serve as a proxy for rainforest extent. We used species distribution models (SDMs) of 317 Dipterocarp species to estimate the geographic extent of appropriate climatic conditions for rainforest on Sundaland at the last glacial maximum (LGM). The SDMs suggest that the climate of central Sundaland at the LGM was suitable to sustain Dipterocarp rainforest, and that the presence of a previously suggested transequatorial savannah corridor at that time is unlikely. Our findings are supported by palynologic evidence, dynamic vegetation models, extant mammal and termite communities, vascular plant fatty acid stable isotopic compositions, and stable carbon isotopic compositions of cave guano profiles. Although Dipterocarp species richness was generally lower at the LGM, areas of high species richness were mostly found off the current islands and on the emergent Sunda Shelf, indicating substantial species migration and mixing during the transitions between the Quaternary glacial maxima and warm periods such as the present.

A PHYLOGENETIC CLASSIFICATION OF THE MALESIAN HIPPOMANEAE (EUPHORBIACEAE)

The delimitation of genera of the tribe Hippomaneae in the Euphorbiaceae has never been solved satisfactorily. The species of this tribe present in the Malay Archipelago (Malesia) were recently revised, but no satisfying generic classification was available. Therefore, a partial phylogenetic analysis of the Malesian Hippomaneae was performed. The delimitation of genera after the analysis was done as conservatively as possible, with only those monophyletic genera distinguished that possessed unique apomorphies within the whole of the Hippomaneae. The genera recognized are not likely to be altered in a more elaborate analysis of the complete Hippomaneae. For this reason the use of a partial analysis for generic delimitations seems justified and presents, when carefully executed, a procedure with which the value of larger regional flora revisions can be greatly improved. The analysis resulted in eight trees, all pointing to the same generic delimitation. Before the analysis 12 groups could be distinguished, but after the analysis it was apparent that only 11 of them constituted monophyletic groups. Two new genera will have to be described, some others will have to be united, and several old delimitations, already abolished for some time, will have to be reestablished. Most remarkable are the union of Excoecaria with Glyphostylus and Sebastiania; and the position of Gymnanthes near taxa formerly attributed to Sapium. The nature of this article is unique in that it will be an attempt to distinguish several natural subunits within Euphorbiaceae tribe Hippomaneae based on only a subset of the included taxa. A phylogenetic analysis of only a part of the tribe will be executed and, based on the monophyletic groups present in the cladogram, genera will be

A Phylogeny of Mallotus (Euphorbiaceae) Based on Morphology: Indications for a Pioneer Origin of Macaranga

A phylogenetic parsimony analysis based on 76 morphological characters of 43 Mallotus ,3 Macaranga ,1 Claoxylon ,1 Cleidion ,1 Sampantaea ,a nd 1Wetria species, resulted in 314 trees of length 602 (CI 5 0.600, RI 5 0.685), which could be summarized in a well resolved consensus cladogram. This consensus cladogram indicates that the genus Mallotus is possibly polyphyletic and that sections Hancea and Oliganthae should probably be excluded from Mallotus. It is proposed to refer to Mallotus as either Mallotus sensu lato (with the sections Hancea and Oliganthae )o rMallotus sensu stricto (excluding the sections Hancea and Oligantae). The genus Macaranga forms a monophyletic group within Mallotus s.s. The traditional section delimitations within Mallotus s.s. do not circumscribe monophyletic clades of species with the exception of section Polyadenii. It is suggested that sections Stylanthus, Rottlera, Mallotus, and the genus Macaranga should form one monophyletic clade. Sections Axenfeldia and Rottleropsis remain unresolved, but are probably closely related. The position of Macaranga within Mallotus suggests that Macaranga evolved from pioneer ancestors that lived in open, scrub-like vegetation. It is hy- pothesised that Macaranga originated relatively recently in South East Asia during the Oligocene or early Miocene (between 38 and 15 million years ago). Macaranga Thou. (ca. 250 species) and Mallotus Lour. (c. 150 species) are two large genera in the Euphorbi- aceae (Webster 1994). They mainly occur in (sub-)trop- ical Asia and the Pacific with only a few species in tropical Africa. They can be found in a large variety of habitats, ranging from the forest understory to com- pletely open scrub vegetation, from dry to swampy or even inundated terrains, and from sandy-loam to lime- stone soils (Pax and Hoffmann 1914; Airy Shaw 1975; Whitmore 1975; Slik et al. 2000). They also show a large variety of life-histories, with typical shade-pre- ferring forest understory species as well as typical shade-avoiding pioneer species (Primack and Lee 1991;A phylogenetic parsimony analysis based on 76 morphological characters of 43 Mallotus, 3 Macaranga, 1 Claoxylon, 1 Cleidion, 1 Sampantaea, and 1 Wetria species, resulted in 314 trees of length 602 (CI 5 0.600, RI 5 0.685), which could be summarized in a well resolved consensus cladogram. This consensus cladogram indicates that the genus Mallotus is possibly polyphyletic and that sections Hancea and Oliganthae should probably be excluded from Mallotus. It is proposed to refer to Mallotus as either Mallotus sensu lato (with the sections Hancea and Oliganthae) or Mallotus sensu stricto (excluding the sections Hancea and Oligantae). The genus Macaranga forms a monophyletic group within Mallotus s.s. The traditional section delimitations within Mallotus s.s. do not circumscribe monophyletic clades of species with the exception of section Polyadenii. It is suggested that sections Stylanthus, Rottlera, Mallotus, and the genus Macaranga should form one monophyletic clade. Sections Axenfeldia and Rottleropsis remain unresolved, but are probably closely related. The position of Macaranga within Mallotus suggests that Macaranga evolved from pioneer ancestors that lived in open, scrub-like vegetation. It is hypothesised that Macaranga originated relatively recently in South East Asia during the Oligocene or early Miocene (between 38 and 15 million years ago). Macaranga Thou. (ca. 250 species) and Mallotus Lour. (c. 150 species) are two large genera in the Euphorbiaceae (Webster 1994). They mainly occur in (sub-)tropical Asia and the Pacific with only a few species in tropical Africa. They can be found in a large variety of habitats, ranging from the forest understory to completely open scrub vegetation, from dry to swampy or even inundated terrains, and from sandy-loam to limestone soils (Pax and Hoffmann 1914; Airy Shaw 1975; Whitmore 1975; Slik et al. 2000). They also show a large variety of life-histories, with typical shade-preferring forest understory species as well as typical shade-avoiding pioneer species (Primack and Lee 1991; Davies et al. 1998; Slik et al. 2000). Additionally, some species are myrmecophytic while others are not (Elias and Sun 1985; Fiala et al. 1999). Both genera belong to the subfamily Acalyphoideae, tribe Acalypheae. Macaranga is the only genus in subtribe Macaranginae, while Mallotus belongs to subtribe Rottlerinae (Webster 1994). The placement in different subtribes suggests that the genera are rather distinct and well separated. However, morphologically it is often difficult to keep them apart. Many characters, like the gland dotted lower leaf surfaces, suggest a close relationship between the two genera. The phylogeny of Macaranga is currently being studied by several groups around the world, and parts of it are already available (Davies 1996; Blattner et al., unpublished data) or will be soon (Whitmore and Davies, in prep.). Here we investigate the phylogeny of Mallotus and try to clarify its relation to Macaranga. Mallotus is rather difficult to circumscribe because most characters show considerable morphological variability among and within species. It is polythetic, and many species can only be characterised by a combination of characters (Bollendorff et al., 2000; Slik et al. 2000; Slik and van Welzen, 2001). The large number of species in combination with the variability of their morphology has resulted in several different sub-generic classifications, the first of which was proposed by Müller Argoviensis (1865, 1866). He subdivided the genus into five sections: Melanolepis (Rchb.f. & Zoll.) Müll. Arg., Cordemoya (Baill.) Müll. Arg., Eumallotus Müll. Arg., Blumeodendron Müll. Arg., and Rottleropsis Müll. Arg. However, he placed some species in different sections that are now considered to be synonyms, thus illustrating the large morphological variability of some of the species and the unclear section delimitation. Sections Blumeodendron and Melanolepis were later regarded as separate genera and excluded from Mallotus (Pax and Hoffmann 1914). Pax and Hoffmann (1914) also proposed a new subdivision of the genus into ten sections: Echinocroton (F. Muell.) Pax & K.Hoffm., Plagianthera (Rchb.f. & Zoll.) Pax & K.Hoffm., Diplochlamys (Müll. Arg.) Pax, Philippinensis Pax & K.Hoffm., Pleiogyni Pax & K.Hoffm., Axenfeldia (Baill.) Pax & K.Hoffm., Polyadenii Pax & K.Hoffm., Hancea (Seem.) Pax & K.Hoffm., Echinus (Lour.) Pax & K.Hoffm., and Stylanthus (Rchb.f. & Zoll.) Pax & K.Hoffm. This subdivision was later refined by Airy Shaw (1963, 1966, 1968, 1969, 1971, 1972a & b, 1975, 1976, 1980a, b & c, 1981). Based on new nomenclatural rules and new ideas about the relationships between the species, he subdivided the genus into eight sections: Rottleropsis, Mallotus, Stylanthus, Oliganthae Airy Shaw, Rottlera ([Willd.] Rchb.f. & Zoll., corr.) Airy Shaw, Axenfeldia, Polyadenii, and Hancea. Sections Hancea, Polyadenii, and Stylanthus were subsequently revised by Bollendorff et al. (2000) and Slik and van Welzen (2001). In these revisions four species were excluded 2001] 787 SLIK & VAN WELZEN: PHYLOGENY OF MALLOTUS from section Hancea (M. brachythyrsus, M. havilandii, M. insularum, M. miquelianus), but their affiliation to the other sections in Mallotus remained unclear. In this paper we study the relationships between the species in the genus Mallotus and their relation to the genus Macaranga using a phylogenetic (parsimony) analysis with a data matrix of 76 morphological characters. Our main questions are: (1) is Macaranga derived from Mallotus or vice versa; (2) do the eight Mallotus sections of Airy Shaw, represent recognisable monophyletic groups; and (3) to which of the Mallotus sections are the four excluded species affiliated? MATERIAL AND METHODS Species selected from several genera closely related to Mallotus were included in the phylogenetic analysis to serve as outgroups (Webster 1994). These selected genera were Claoxylon A.Juss., Cleidion Blume, Sampantaea Airy Shaw, Wetria Baill., and Macaranga (Table 1). The three selected Macaranga species represent distinctly different species groups within this genus (Whitmore 1975). We included Mallotus species belonging to sections Hancea, Polyadenii, and Stylanthus as circumscribed by Bollendorff et al. (2000) and Slik and van Welzen (2001) and the four species excluded from Hancea (Slik and van Welzen 2001) (Table 1). Of the five remaining sections at least three species, including the type, were studied if

Delimitation of Sauropus (Phyllanthaceae) Based on Plastid matK and Nuclear Ribosomal ITS DNA Sequence Data

BACKGROUND AND AIMS A recent molecular phylogenetic study showed that Sauropus is deeply embedded within Phyllanthus together with its allies, Breynia and Glochidion. As relationships within Sauropus are still problematic and the relationship with Breynia has long been doubted, more molecular data are needed to test/corroborate such a broad definition of Phyllanthus. This study aims to clarify the status and delimitation of Sauropus and establish its position within Phyllanthaceae. METHODS Plastid matK and nuclear ribosomal ITS DNA sequence data for Sauropus and its allies were used to construct phylogenetic trees using maximum parsimony and Bayesian methods. KEY RESULTS Within Phyllanthus, Sauropus can be split into the mainly south-east Asian Sauropus sensu stricto (s.s.) plus Breynia and the mainly Australian Sauropus (formerly Synostemon). Sauropus s.s. plus Breynia comprise two distinct clades; one is the combination of Sauropus sections Glochidioidei, Sauropus and Schizanthi and the other is the combination of Sauropus sections Cryptogynium and Hemisauropus and the monophyletic genus Breynia. CONCLUSIONS Molecular data indicate that Synostemon should be reinstated at the same level as Sauropus s.s. and that Sauropus s.s. should be united with Breynia under the latter, older name. The molecular data corroborate only two of the five infrageneric groups of Sauropus recognized on the basis of morphological data.

Phylogeny of palaeotropic Derris-like taxa (Fabaceae) based on chloroplast and nuclear DNA sequences shows reorganization of (infra)generic classifications is needed

PREMISE OF THE STUDY Palaeotropic Derris-like taxa (family Fabaceae, tribe Millettieae) comprise 6-9 genera. They are well known as important sources of rotenone toxin, which are used as organic insecticide and fish poison. However, their phylogenetic relationships and classification are still problematic due to insufficient sampling and high morphological variability. METHODS Fifty species of palaeotropic Derris-like taxa were sampled, which is more than in former studies. Three chloroplast genes (trnK-matK, trnL-F IGS, and psbA-trnH IGS) and nuclear ribosomal ITS /5.8S were analyzed using parsimony and Bayesian methods. KEY RESULTS Parsimony and Bayesian analyses of individual and combined markers show more or less similar tree topologies (only varying in terminal branches). The old-world monophyletic genera Aganope, Brachypterum, and Leptoderris are distinct from Derris s.s., and their generic status is here confirmed. Aganope may be classified into two or three subgeneric taxa. Paraderris has to be included in Derris s.s. to form a monophyletic group. The genera Philenoptera, Deguelia, and Lonchocarpus are monophyletic and distinct from each other and clearly separate from Derris s.s. Morphologically highly similar species of Derris s.s. are shown to be unrelated. Our study shows that previous infrageneric classifications of Derris are incorrect. Paraderris elliptica may contain several cryptic lineages that need further investigation. CONCLUSIONS The concept of the genus Derris s.s. should be reorganized with a new generic circumscription by including Paraderris but excluding Brachypterum. Synapomorphic morphological features will be examined in future studies, and the status of the newly defined Derris and its closely related taxa will be formalized.

Phylogeny and proposed circumscription of Breynia, Sauropus and Synostemon (Phyllanthaceae), based on chloroplast and nuclear DNA sequences

Abstract. Previous estimates of phylogeny in the Phyllanthaceae, Phyllantheae, have been hampered by undersampling of species from morphologically distinctive groups and using too few gene regions. To increase the phylogenetic resolution, sequences of two nuclear (ITS1–5.8S–ITS2) and Phytochrome C (PHYC)) and two non-coding chloroplast (accD–psaI, trnS–trnG) DNA markers were analysed using maximum parsimony and Bayesian inference with expanded sampling in Breynia, Glochidion, Sauropus and Synostemon. Our results supported reinstatement of Synostemon, previously included in Sauropus s.str., to generic rank, and provided evidence towards its future infrageneric classification. The results also indicated expansion of Breynia to include Sauropus s.str.; this combined monophyletic group consists of two strongly supported clades. Finally, we showed monophyly for Glochidion, which is sister to Phyllanthus subg. Phyllanthodendron, both still remaining undersampled. Morphological features characteristic of Breynia, Sauropus and Synostemon are discussed, as well as the desirability of dividing Phyllanthus into smaller genera.

A phylogenetic analysis of Harpullia (Sapindaceae) with notes on historical biogeography

Abstract A phylogenetic analysis of Harpullia (two incompletely-known species excluded, Conchopetalum and Majidea as outgroups) based on morphological characters resulted in five most parsimonious cladograms. These cladograms and the strict consensus cladogram show a few polytomies. Only a few branches have relatively high Decay Indices. The strict consensus cladogram indicates that Harpullia is monophyletic (five synapomorphies) and does not support the existing subgeneric classification. Thus, it is advisable to refrain from any subgeneric classification. The historical biogeographic pattern confirms the one already found for eight other genera of Sapindaceae. Harpullia originated on the New Guinean—Australian plate and from there dispersal and speciation occurred towards New Caledonia and Southeast Asia main land. Communicating Editor: Matt Lavin

A Revision of Ficus Subsection Urostigma (Moraceae)

Abstract The present taxonomic revision of Ficus subsection Urostigma recognizes 27 species, of which three are new: F. chiangraiensis, F. middletonii, F. pseudoconcinna. Two new varieties are distinguished within F. virens, var. dispersa and var. matthewii. Ficus lecardii and L. salicifolia, formerly subspecies of F. cordata, are again reinstated to the species level. Typical characters for the subsection are monoecy, monostaminate flowers, red(-brown) colored ovaries and cystoliths on only the abaxial leaf surface. Ficus amplissima and F. rumphii (section Leucogyne) were formerly part of subsection Urostigma, and they have been added here to the key and descriptions because of their morphological resemblance with the species in F. subsection Urostigma. Molecular based phylogenetic analyses showed that at least F. rumphii is unrelated to subsection Urostigma. The two species only differ from subsect. Urostigma in their whitish ovaries and cystoliths at both sides of the leaf blade and they are pollinated by a different group of wasps, species of Eupristina subg. Parapristina.