IAWA Journal, Vol. 21 (2), 2000: 213–235

WOOD ANATOMY OF () by W. John Hayden & Sheila M. Hayden Department of Biology, University of Richmond, Richmond, VA, 23173, USA

SUMMARY Via LM and SEM, we studied wood structure of 51 genera representing 19 tribes of Acalyphoideae, the largest subfamily of Euphorbiaceae. Many acalyphoid woods possess the following features: growth rings indistinct or weakly defined; pores evenly distributed; simple perfora- tion plates (but admixture of irregular scalariform plates common); al- ternate intervessel pits; vessel-ray pits larger than intervessel pits, circu- lar to elongate and alternate to irregular; thin to moderately thick-walled non-septate fibre-tracheids or libriform wood fibres; parenchyma dis- tribution diffuse, diffuse-in-aggregates, and scanty paratracheal, some- times in thin-tangential bands; heterocellular rays seldom more than 3 cells wide; and prismatic crystals in parenchyma and/or ray cells. Within this syndrome, a number of other wood characters also occur but at lower frequency. For the most part, the unusual features have not proven systematically informative at the tribal level. Presence of lysi- genous radial canals, however, supports recognition of Alchorneae. Wood data do not support the segregation of and from subfamily Acalyphoideae. Key words: Acalyphoideae, Euphorbiaceae, Pandaceae, Peraceae, wood anatomy.

INTRODUCTION

Acalyphoideae is the largest and most complex of the five subfamilies of the Euphorbiaceae. Its diversity can be summarized succinctly via statistics from Websterʼs (1994) classification: 20 tribes, 116 genera, and c. 2,000 that are found through- out the world but are especially abundant in the tropics. Within the , Acalyph- oideae consists of those that have uniovulate locules but lack the characteristic pollen, latex, and indumentum features that define Crotonoideae and Euphorbioideae (Webster 1994). Thus, Acalyphoideae appears to be a paraphyletic assemblage. In- cluded within the ranks of Acalyphoideae are several genera of uncertain taxonomic placement. For example, has been segregated as family Peraceae (Airy Shaw 1965; Radcliffe-Smith 1987), the genera of tribe Galearieae have been placed in Pandaceae (Cronquist 1981; Radcliffe-Smith 1987; Takhtajan 1997), and has been included in Pandaceae (Webster 1987) or, sometimes, in Euphorbiaceae sub- family Phyllanthoideae (e.g., Pax & Hoffmann 1931; Mennega 1987).

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Approximately 86 percent of acalyphoid genera can be characterized as woody. Although a large number of publications contain some information on wood structure of various members of Acalyphoideae (see Appendix), most of these consider a small number of genera and no general summary of wood anatomical diversity for the sub- family exists. This paper, therefore, attempts to fill this gap based on readily available materials from collections at the Smithsonian Institution (USw) and University of Utrecht (Uw). This paper joins papers on Phyllanthoideae (Mennega 1987), Oldfield- ioideae (Hayden 1994), and Euphorbioideae (Mennega, in preparation) as a contribu- tion towards the elucidation of the anatomy and systematics of Euphorbiaceae.

MATERIALS AND METHODS

Wood specimens of Acalyphoideae examined here are listed in the Appendix. This also includes a guide to previous wood anatomical literature for each . We pre- pared microscope slides from 110 specimens and examined 46 more via slides loaned from USw and Uw. Materials studied thus represent a total of 19 tribes, 51 genera, and 110 species. Wood blocks were rehydrated by boiling in tap water with a few drops of Aerosol OT solution, rinsed, and sectioned on a sledge microtome. Sections were stained in Harrisʼ hematoxylin and counterstained in safranin. Macerations were prepared by treatment in equal parts of 10 percent nitric and 10 percent chromic acids, followed by several rinses in water, staining in safranin, dehydration in tert- butyl alcohol, and clearing in toluene. Photomicrographs were prepared from Kodak Technical Pan film developed in Kodak HC110 developer at dilution F. For study with SEM, wood sections approximately 1 mm thick were cut from air-dried wood blocks, affixed to specimen stubs, sputter-coated with a gold-palladium mixture, and studied with a Hitachi S-2300 SEM. Scanning electron micrographs were prepared from Kodak Tri-X film developed in Kodak HC110 developer at dilution B.

RESULTS AND DISCUSSION

Wood anatomy of Acalyphoideae Growth rings — Growth rings, when detectable, are weakly defined by variation in fibre wall thickness or, sometimes, by the presence of boundary parenchyma. Vessels — Vessels are evenly distributed radially and, hence, diffuse-porous when growth rings are present. There are two cases of unusual large-scale vessel distribu- tion: in Necepsia vessels are organized in alternating radial files of wood with and wood without vessels (Fig. 1) and, in , vessels are organized into radial files separated by broad aggregate rays (Fig. 2). At small scales, vessels are most fre- quently grouped in a mixture of solitary cells or small radial multiples and/or clusters of 2–6 cells (Fig. 1–10, 12). Predominantly solitary vessels were noted in (Fig. 3), malayana, guianensis, C. simulata, Megistostigma, , and (Fig. 4). Pore outlines are routinely circular to elliptical but distinctly angular in Dicoelia (Fig. 5) and somewhat so in Microdesmis. Perfora- tions on vessel elements are generally simple (Fig. 13). Perforations are exclusively

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Fig. 1–3. Transverse sections of acalyphoid woods. – 1: Necepsia afzelii, Cooper Y. 13726, vessels grouped in radial files. – 2:Podadenia sp., Krukoff 4074, aggregate ray. – 3: Botryophora geniculata, Boeea 5494, pith flecks. — Scale bars = 500μ m.

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Fig. 4–12. Transverse sections of acalyphoid woods. – 4: Megistostigma malaccense, Boeea 1389, lianous stem with large, solitary vessels and thin-walled fibres. – 5: Dicoelia sp., Boeea 7794, vessels with angular outline. – 6: grandifolium, Krukoff 5553, paren-

Downloaded from Brill.com10/07/2021 08:57:34AM via free access Hayden & Hayden — Wood anatomy of Acalyphoideae 217 scalariform in Microdesmis and Panda. In some genera or particular species, simple perforations occur mixed with scalariform and/or irregular perforations; such mixed perforations occur in some species of , Adenophaedra, , Apar- isthmium (Williams 1936), tamanduana (Fig. 15), Botryophora, Caryoden- dron grandifolium, (Fig. 16), , (Metcalfe & Chalk 1950), Coccoceras (Fig. 14), Conceveiba krukoffii, C. guianensis, Dicoelia, Discoclaoxylon (Normand 1955), , zenkeri, some species of (Metcalfe & Chalk 1950), , (Metcalfe & Chalk 1950), (Metcalfe & Chalk 1950), and (Metcalfe & Chalk 1950). The frequency of per- forated ray cells in acalyphoid woods and their propensity for perforations of vari- able and irregular form (see below) demand great care in distinguishing perforation features for ordinary vessel elements as opposed to perforated ray cells. Intervessel pits are circular to elliptical or sometimes angular (polygonal) if crowded, alternate (Fig. 18), and most frequently 5–12 μm in vertical diameter; Dechamps (1979) re- ported intervessel pit diameters of 15 μm for , but our material has a range of 7–10 μm. Vestured pits have been reported for vessels of Galearia and Panda (Forman et al. 1966) and initial observations via light microscopy suggested vesture- like structures in intervessel pits of Bernardia, , and ; SEM studies, how- ever, have failed to reveal true vestured pits in any of these woods. In Bernardia and Clutia, spherical wart-like structures located on the surface of the pit cavity are re- sponsible for the pseudovestures observed via LM. Confluent inner apertures were noted on intervessel pits of ricinella, , Conceveiba, , and Octospermum. Intervessel pits are notably small (c. 2 μm) in Microdesmis and also in Claoxylon africanum and , but pits of other species of the latter two genera are much larger. Vessel-ray pits are circular to elongate and alter- nate to irregular; elongate pits may be oriented horizontally, vertically, or diagonally and they are characteristically larger than the intervessel pits (Fig. 13, 17, 18). Ves- sel elements are notably long in Coccoceras (c. 2 mm), notably short in Clutia (230– 340 μm), and notably wide in the liana, Megistostigma malaccense (up to 390 μm) (Fig. 4). Fibriform vessel elements were observed grouped with larger vessel ele- ments in Dicoelia and Clutia; in C. abyssinica, some fibriform vessel elements bear few-barred scalariform perforations as opposed to the simple perforations in ordinary vessels; in C. kilimandsharica, fibriform vessel elements possess helical thickenings. Otherwise, we observed helical thickenings only in vessels of Mallotus polyadenus and Microdesmis but they have also been reported for Cleidion, Pogonophora, and Trewia (Metcalfe & Chalk 1950); elongate inner apertures of intervessel pits form grooves on the vessel walls of Bernardia tamanduana (Fig. 15). Tyloses are frequent chyma in thin, irregular tangential bands. – 7: Aparisthmium cordatum, Maguire et al. 51902, vessels multiple with aliform vasicentric parenchyma, fibres moderately thick-walled. – 8: Homonoia riparia, Stern 2104, a rheophyte, thin-walled fibres difficult to distinguish from axial parenchyma. – 9: Ricinus communis, USw 8879, vessel groupings. – 10: , USw 18341, vessels in radial multiples, thick-walled fibres, and diffuse axial parenchyma. – 11: Galearia celebica, FPAw 29249, vessels in radial multiples, and thick-walled fibres. – 12: schomburgkianus, USw 52702, vessels with sclerified tyloses. — Scale bars of 4 & 9 = 100 μm; all others = 50 μm.

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Fig. 13–21. SEM images of acalyphoid woods, all radial sections except 15. – 13: Botryophora geniculata, Boeea 5494, simple perforation plate and vessel-ray pits. – 14: Coccoceras sp., Boeea 5448, scalariform perforation plate. – 15: Bernardia tamanduana, Bartlett 10356,

Downloaded from Brill.com10/07/2021 08:57:34AM via free access Hayden & Hayden — Wood anatomy of Acalyphoideae 219 in heartwood and relatively thin-walled (Fig. 34). However, sporadic, thick-walled sclerotic tyloses occur in tokbrai, , Chaetocarpus (Fig. 12, 22), Homonoia, hainanense, Pera, and Pogonophora (Metcalfe & Chalk 1950); in Cephalomappa, Homonoia and Koilodepas hainanense (Fig. 23), sclerotic tyloses enclose prismatic crystals, as do the lightly sclerified tyloses of dioscoreifolia. Thin-walled tyloses of Macaranga spinosa enclose druses.

Fibres — Fibres are mostly non-septate and most commonly fibre-tracheids. Fibre- to-fibre pits range from relatively large (c. 7 μm in vertical diameter) in Cheilosa, to very small (c. 1–2 μm in vertical diameter) in Cleidion spp., Mareya, and . Libriform wood fibres (non-septate) are found in Adelia, Bernardia, Caryodendron, Cleidion spp., Clutia kilimandsharica, Coccoceras, Dalechampia dioscoreifolia, Koilo- depas, , Moultonianthus, Necepsia, Neoscortechinia, Pera bumeliaefolia, and Pogonophora. Septate fibres occur in Acalypha (Fig. 24, 27), Clutia abyssinica, Dicoelia, and Mareya. Fibres in Acalypha are mostly septate libriform fibres, although some species have septate fibre-tracheids. Contrary to the usual condition of several septa per cell, each short septate fibre-tracheid of Clutia abyssinica bears a single septum dividing the lumen into two approximately equal-sized chambers. In Dicoelia, small groups of septate libriform fibres occur intermixed with non-septate cells. We cannot confirm the presence of septate fibres in Mallotus, as reported by Kanehira (1921a, b) nor in Necepsia, as reported by Normand (1955). Average fibre length ranges from c. 0.5 mm for Clutia to c. 2.8 mm for Coccoceras. Wall thickness varies most frequently from thin to moderately thick; however, very thick-walled fibres with lumina nearly closed were observed in Caryodendron orinocoense, Chaetocarpus, Coccoceras, Galearia (Fig. 11), Microdesmis, and Panda (Fig. 10). Groups of fibres with gelatinous walls are frequent.

Axial parenchyma — Axial parenchyma distribution is characterized by diffuse and diffuse-in-aggregate patterns (Fig. 4, 6, 9–12) with some scanty paratracheal cells. Parenchyma organized in thin tangential bands, mostly 1–2 cells wide, is found in Acalypha repanda var. denudata, Agrostistachys, Aparisthmium, Caryodendron, Chaetocarpus, Dalechampia, Discoclaoxylon (Normand 1955), Homonoia (as bound- ary parenchyma), Macaranga reineckii (Burgerstein 1909), Pera bicolor, P. ferruginea, P. glabrata, Pogonophora, , (as boundary parenchyma, ac- cording to Metcalfe & Chalk 1950), and Syndyophyllum; Normand (1955) reported thick bands of parenchyma in . Apotracheal parenchyma is absent in scalariform perforation plate and vessel wall sculpture, tangential section. – 16: Claoxylon purpureum, Stern 2307, irregular perforation plate. – 17: Claoxylon echinospermum, USw 30421, vessel-to-ray pits. – 18: Coccoceras sp., Boeea 5448, vessel-to-ray pits (left) and intervessel pits (right). – 19: Mareya micrantha, USw 24337, perforated ray cell with simple perforations. – 20: Mareya micrantha, USw 24337, perforated ray cell with vertical bars on scalariform perforation. – 21: Botryophora geniculata, Boeea 5494, perforated ray cell with irregular perforation plate incorporating some bordered pits. — Scale bars of 13, 14, & 17–19 = 50 μm; 15 & 21 = 25 μm; 16 = 20 μm; 20 = 100 μm.

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Dicoelia which does, however, possess some scanty paratracheal cells. Paratracheal parenchyma is more extensive in Omphalea than in most other genera and may be characterized as vasicentric. Sometimes the banded parenchyma in Aparisthmium (Fig. 7), Dalechampia and Pogonophora approach aliform to confluent patterns. Axial parenchyma is absent or sparse in Acalypha and Clutia; when present, parenchyma in these genera is usually restricted to a few scanty paratracheal cells. Parenchyma can be difficult to discern from the fibres in transverse section in certain woods with thin- walled fibres (Fig. 8). Subdivision of paratracheal parenchyma strands ofOctospermum is unique; strands are irregularly two cells wide at the middle and taper to single cells at the tips of the strand; evidently, during development in the cambial zone, in addi- tion to the usual transverse divisions that subdivide the strand, these cells were stretched laterally in concert with the expansion of the relatively wide adjacent vessels, result- ing in additional vertical or oblique planes of division. Pith flecks (Fig. 3) are rela- tively frequent, especially in woods with thin-walled fibres. Parenchyma cells of the pith flecks of cordifolia contain prismatic crystals. Most frequently, ordi- nary cells of axial parenchyma are devoid of crystalline inclusions. However, rhom- boidal/prismatic crystals in chambered axial parenchyma were found in a sizeable minority of genera: Alchorneopsis, Blumeodendron, Caryodendron (Fig. 25), Cephalo- mappa, Chaetocarpus, Cheilosa, Galearia, Gavarretia, Koilodepas, Macaranga spp., Mareya, Neoscortechinia, Octospermum, Pera, Podadenia, Ptychopyxis, Trigono- pleura, and . Frequently, crystal-bearing cells are more or less sclerified rela- tive to the ordinary parenchyma cells. Rarely encountered crystal features of axial parenchyma include: prismatic crystals in ordinary (not subdivided) cells in Alchornea and Neoscortechinia; elongate prismatic crystals in species of Claoxylon; and druses in Macaranga kilimandsharica. In Caryodendron (Fig. 25), chambered crystal-bear- ing parenchyma cells are notably wider, both radially and tangentially, than ordinary axial parenchyma. Silica in the form of irregular spheroids up to 12 μm in diameter were observed in axial parenchyma of Neotrewia and, rarely, in Aparisthmium.

Rays — A wide range of ray features exist within Acalyphoideae. A great many woods, however, have numerous closely-spaced narrow rays, most often only one or two, sometimes three, cells wide (Fig. 29, 30). Rays are most often heterocellular, with erect cells located at the margins or between multiseriate portions of polymerous (or vertically fused or interconnected) rays. There are two notable exceptions to the usual heterocellular condition: there is a tendency towards homocellular procumbent in Leu- cocroton and, conversely, towards homocellular square to erect in Acalypha. Wide rays (with maximum cell width indicated parenthetically) were found in Acalypha (5) (Fig. 27), Blumeodendron tokbrai (4), Botryophora (7), Cheilosa (4), Old World spp. of Cleidion (4), Coccoceras (5), Discoglypremna (4) (Normand 1955), Galearia

Fig. 22–25. Radial sections of acalyphoid woods. – 22: Chaetocarpus schomburgkianus, USw 52702, sclerified tyloses. – 23: Koilodepas hainanensis, Uw 21400, crystal-bearing sclerified tyloses. – 24: Acalypha obovata, Woytkowski 5896, septate fibres. – 25: Caryodendron grandifolium, Krukoff 5553, axial xylem parenchyma, ordinary cells and chambered cells each bearing a prismatic crystal. — Scale bars of 22, 23, & 25 = 50 μm; 24 = 25 μm.

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(9) (Fig. 26), Macaranga fimbriata (5), Mareya (5), Microdesmis (5), Moultonianthus (4), Panda (5), Podadenia (5), Pogonophora (4), Ricinus (4) (Fig. 28), and Syndyo- phyllum (4). Predominantly or exclusively uniseriate rays were found in Aparisthmium (Fig. 30), Bernardia, Caryodendron, Conceveiba, Dicoelia, Leucocroton, Melanolepis, Pera bicolor, P. glomerata, and P. oppositifolia. Aggregate rays occur in the one speci- men of Podadenia studied (Fig. 2); the two aggregate rays observed occur at transi- tion regions in grain orientation in the wood and whereas one aggregate ray consists of closely spaced rays of ordinary morphology for the genus, component rays of the other example seen are highly variable, much wider than usual, and bounded or tra- versed by convoluted fibres. Metcalfe & Chalk (1950) report aggregate rays inNecepsia and Pseudagrostistachys. As noted above, vessels are grouped in distinct radial files in Necepsia (Fig. 1), but since ray frequency is nearly the same in vesselled and vesselless portions of this wood, we interpret the distinctive large-scale pattern strictly as a vessel distribution feature. Disjunctive ray cells were prominent in Bernardia and Leucocroton. Polymerous (vertically fused) rays are especially frequent in Neo- scortechinia. Multiseriate rays of Acalypha, Galearia, Mareya, and Microdesmis have weakly or inconsistently developed sheath cells. Most acalyphoid woods possess crystaliferous ray cells. Most commonly, the crystals are rhomboidal/prismatic and occur in ordinary ray cells and/or bijugate pairs, i.e., adjacent crystal-bearing cells evidently formed by subdivision of an ordinary ray cell (Fig. 31, 33). Occasionally, erect ray cells are subdivided to form four crystal-bearing cells (Fig. 31). Rarely two prismatic crystals occur in the same cell. Moultonianthus is unique in possessing bi- jugate crystal-bearing cells that are also sclerified. Druses were observed in ray cells of Acalypha amentacea, Claoxylon purpureum (Fig. 32), Koilodepas, Macaranga kilimandsharica (bijugate pairs), and Ricinus; similarly, druses in ray cells are re- ported for Müll.Arg. (Chattaway 1955). In Acalypha elliptica, A. repanda var. denudata, and Claoxylon sandwichense, prismatic crystals were fre- quent but some druses were also observed. In Koilodepas, druses occur in procum- bent ray cells while prismatic crystals are common in erect ray cells. No crystals were observed in rays of some Acalypha spp., Agrostistachys, Bernardia, Chaetocarpus, Claoxylon africanum, C. longifolium, and Dicoelia; Espinoza de Pernía (1987), how- ever, reports prismatic crystals in ray cells of Chaetocarpus schomburgkianus. Silica in the form of irregular spheroids up to 12 μm in diameter is present in ray cells of Agrostistachys, Aparisthmium, Cephalomappa, Dalechampia, Neotrewia, and Pogon- ophora. In Aparisthmium, silica and prismatic crystals sometimes co-occur in the same cell. Perforated ray cells were seen in Acalypha, Adenophaedra, Botryophora (Fig. 21), Claoxylon, Cleidion, Coccoceras, Coelebogyne, Conceveiba, Galearia,

Fig. 26–30. Tangential sections of acalyphoid woods. – 26: Galearia celebica, FPAw 29249, very tall multiseriate ray and several uniseriate rays. – 27: Acalypha obovata, Woytkowski 5896, portions of one wide multiseriate ray and several uniseriate rays, septate fibres. – 28: Ricinus communis, USw 8879, abundant multiseriate rays and few uniseriate rays. – 29: Neo- scortechinia nicobarica, USw 34315, uni- and biseriate rays. – 30: Aparisthmium cordatum, Maguire et al. 51902, uniseriate rays, tangential section. — Scale bars of 26, 28, & 29 = 100 μm; 27 & 30 = 50 μm.

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Fig. 31–34. Crystals in ray cells and lysigenous canal in acalyphoid woods. – 31: Koilodepas hainanensis, Uw 21400, bijugate and double-bijugate ray cells each with a prismatic crystal, radial section. – 32: Claoxylon purpureum, Stern 2307, druse in erect ray cell, radial section. –

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Fig. 35 & 36. Nonarticulated laticifers in acalyphoid woods. – 35: Dalechampia dioscoreifolia, Wetmore & Abbe 11, laticifers in multiseriate ray, tangential section. – 36: Dicoelia sp., Boeea 7794, laticifer in multiseriate ray, tangential section. — Scale bars = 50 μm.

Mareya (Fig. 19, 20), Melanolepis, Microdesmis, Moultonianthus, Neoscortechinia, Neotrewia, and Panda and they are reported for Pera glabrata (Giraud 1983). Perfo- rations of perforated ray cells range from simple (Fig. 19) to multiple, in which case the bars may be horizontal, vertical (Fig. 20), or irregular, sometimes extremely so (Fig. 21). Two different kinds of secretory structures occur in rays of acalyphoid woods, laticifers and large lysigenous canals. Non-articulated laticifers are present in some rays of Dalechampia (Fig. 35) and Dicoelia (Fig. 36); our report confirms the obser- vations of Heimsch (1942) in regard to Dalechampia. Laticifers in rays of Dalechampia are continuous with the laticiferous system of the pith and our inference from very limited material is that the same is true for Dicoelia. We also observed a single laticifer in a fragment of pith tissue of Acalypha stachyura, but saw no evidence that the lati- cifer system extended into the wood.

33: Galearia celebica, FPAw 29249, bijugate ray cells each with a prismatic crystal, radial sec- tion. – 34: Conceveiba krukoffii, Krukoff 8396, vessel with tyloses (left) and lysigenous radial canal (right), tangential section. — Scale bar of 31 = 50 μm; 32 & 33 = 25 μm; 34 = 500 μm.

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We observed lysigenous canals in anomalously large rays of Adenophaedra prealta, Alchornea, and Conceveiba. Whereas ordinary rays in these woods are one to two cells wide, those containing lysigenous canals in Adenophaedra prealta and Alchornea range up to nine cells wide. Dimensions of the lysigenous canals of Adenophaedra prealta and Alchornea range from 350–700 μm vertically and 160–325 μm tangen- tially. The lysigenous canals of Conceveiba (Fig. 34) are larger, ranging from 2–6 mm vertically and 280–460 μm tangentially; otherwise they are structurally similar to those seen in Adenophaedra prealta and Alchornea. Although lysigenous canals gen- erally possess secretory functions, the canals appear devoid of contents in our prepa- rations so we cannot speculate on the nature of their secretory activity. Others have reported similar radial structures for the three genera discussed above and, also, in Aparisthmium, Gavarretia, and Pera (Williams 1936; Record & Hess 1943; Metcalfe & Chalk 1950; Pérez-Mogollón 1973; Détienne & Jacquet 1983; Dechamps 1979, 1985). However, lysigenous canals were not evident in the specimens of Aparisthmium, Gavarretia, and Pera available to us. In some cases, these radial secretory canals have been described as laticiferous (Pérez-Mogollón 1973; Détienne & Jacquet 1983). On the other hand, Record and Hess (1943) remarked that these canals suggested “remnant traces,” a characterization repeated by Metcalfe and Chalk (1950). Careful study of appropriate-aged specimens would be required to ascertain whether these secretory canals bear any connection with leaf traces. Inconsistencies between our observations and previous reports of lysigenous canals could be explained by differing ages of the samples studied or because these features occur at such wide intervals in the wood that they are included only sporadically in standard-sized sec- tioning blocks. Some observations of the rays of Coelebogyne and Wetria insignis might be pertinent in regard to the lysigenous canals discussed above. We observed frequent irregular wide rays incorporating contorted fibres scattered among the ordi- nary rays of Coelebogyne and Wetria. Dimensions of these anomalous rays in Coel- ebogyne and Wetria are consistent with the lysigenous canals described above but they differ in that all cells are intact. We note further that Record and Hess (1943) and Metcalfe and Chalk report rays up to 8 cells wide in Aparisthmium, but that rays in our material were mostly uniseriate with a few biseriates. These anomalous wide rays reported in Aparisthmium, and observed in Coelebogyne, and Wetria might thus rep- resent a pre-lysigenous condition of the sorts of canals that we observed in Adeno- phaedra prealta, Alchornea, and Conceveiba. We note further that, in the case of Wetria, the frequency and close, irregular spacing of anomalously large rays does not easily support a developmental connection with leaf traces.

Relationships with other subfamilies Compared with euphorbiaceous woods in general, those of subfamily Acalyphoideae constitute a reasonably cohesive group. In contrast, woods of subfamily Phyllanthoi- deae are highly diverse (Mennega 1987). According to either the classical Baileyan concepts of evolutionary trends in woody tissue of dicots (summarized in Baas 1986, Carlquist 1988, Stern 1978) or more recently, as patterns discerned in the fossil record (Wheeler & Baas 1991), Phyllanthoideae includes woods that range from strikingly primitive structure to those that are more highly advanced. For example, Blotia pos-

Downloaded from Brill.com10/07/2021 08:57:34AM via free access Hayden & Hayden — Wood anatomy of Acalyphoideae 227 sesses very long vessel elements with scalariform perforations, non-septate fibres, abundant apotracheal parenchyma, and rays that are tall, wide, and highly heterocel- lular (Mennega 1987). Blotia thus serves as an appropriate model for primitive wood structure in Euphorbiaceae. However, as one example of more extreme specializa- tion, Phyllanthoideae also includes woods with short vessel elements bearing simple perforations, prosenchyma consisting of septate-fibres, parenchyma essentially ab- sent, and smaller, less extremely heterocellular rays; woods with this syndrome con- stitute the so-called “Glochidion-type” (Metcalfe & Chalk 1950). Between these two extremes, phyllanthoid woods exhibit a wide range of vessel, fibre, parenchyma, and ray features. In comparison, woods of Acalyphoideae are much less diverse. Most commonly, woods of Acalyphoideae possess vessel elements with simple perforations, large and irregular vessel-ray pits, thin- to moderately thick-walled non- septate fibres, abundant apotracheal parenchyma, and relatively narrow heterocellu- lar rays that are frequently polymerous (vertically fused) and bear crystals in the pat- tern we described as bijugate. Mennega (1987) commented that woods of the other two uniovulate subfamilies, Crotonoideae and Euphorbioideae, are similar, except that laticiferous elements in the ray system are more frequently encountered in Crotonoideae and Euphorbioideae than in Acalyphoideae (see also Rudall 1987). Perusal of wood descriptions for several crotonoid woods studied by Stern (1967) re- veals that the general similarity with acalyphoid woods described here even extends to a similar range of crystal forms and structural details of crystal-bearing paren- chyma and ray cells. Evidence for the monophyly of uniovulate euphorbs (Acalyphoideae + Crotonoideae + Euphorbioideae) is emerging from molecular sequence data (Wurdack & Chase 1999) and it is evident that, by and large, woods of these plants form a mutually co- hesive group. Laticifers, however, are not uniformly present throughout the uniovulate euphorbs; they are present and widespread in Crotonoideae and Euphorbioideae but occur in just a few genera of Acalyphoideae, suggesting that relationships of Dale- champia and Dicoelia might lie outside of the subfamily to which they are presently assigned (see below). In addition, the distribution of laticiferous systems in uniovulate woods suggests that Acalyphoideae is a paraphyletic assemblage, a conclusion also reached by molecular analyses (Wurdack & Chase 1999).

Relationships within Acalyphoideae Acalyphoid woods are not completely uniform. For the most part, however, we have been unable to draw meaningful conclusions concerning relationships among acalyphoid genera based on wood features alone. Unusual wood features, i.e., char- acter states with restricted taxonomic distribution within the subfamily fail to be informative because they are either plesiomorphic, autapomorphic, or occur in seem- ingly random patterns, i.e., not correlated with other wood structures or not correlated with the existing taxonomic system (Webster 1994). In general, we have not found wood characters of Acalyphoideae to occur in nested hierarchical clusters. One char- acter, however, occurrence of lysigenous radial canals is exceptional and supports recognition of Alchorneae as a distinct tribe (see below).

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Exclusively scalariform perforations are found in Microdesmis and Panda of tribe Galearieae, and rays within this tribe are remarkably tall and wide; Galearia has mixed simple and scalariform perforations; arguably, the genera of tribe Galearieae possess some of the most primitive woods within Acalyphoideae. By their plesio- morphic nature, these vessel and ray features offer little insight in determining whether the genera of Galearieae should be retained within Acalyphoideae or segregated as family Pandaceae (Cronquist 1981, 1988; Takhtajan 1997). Molecular data, however, suggests that Pandaceae do form a distinct lineage (Wurdack & Chase 1999). In addi- tion to tribe Galearieae, plesiomorphic multiple perforations and wide rays occur spo- radically in the rest of the subfamily; aside from this tribe, however, plesiomorphic vessel and ray features show no obvious clustering or grouping with the tribes defined in Websterʼs (1994) classification. Another probable symplesiomorphy is the pres- ence of angular vessels in Dicoelia and Microdesmis. Based on present knowledge, the following features appear to be autapomorphic within Acalyphoideae: occurrence of vessels in radial files, wide vessels as an adap- tation to lianous growth habit, druses in tyloses, prismatic crystals in thin-walled tyloses, aggregate rays, bijugate cells with crystals that are also sclerified. It is con- ceivable that as additional acalyphoid woods are studied, these apparent autapo- morphies may prove to be synapomorphous and, thus, systematically informative. Except as noted immediately below, all other wood features that diverge from the usual syndrome (and thus merit special mention in the general description, above) occur without obvious correlation to either Websterʼs (1994) classification or to other wood features. The occurrence of laticifers in Acalyphoideae offers a striking example of the difficulty in the taxonomic interpretation of wood data for the group. Non-articulated laticifers in secondary xylem were observed only in Dalechampia and Dicoelia. These genera are classified in different tribes and they possess scarcely any other morpho- logical or anatomical characters that could convincingly link them together. Ana- tomically, the presence of laticifers in these woods suggest that classification in Cro- tonoideae or Euphorbioideae might be preferable to Acalyphoideae, yet the biovulate locules of Dicoelia would be just as discordant in any of these three subfamilies. There seems little recourse beyond repeating the assertion that Dicoelia represents an ancient isolated relic (Airy Shaw 1972). In contrast, the presence of large, lysigenous radial canals (or their anomalous wide ray precursors), appears to be phylogenetically informative. Based on presently available data for five of the nine constituent genera, lysigenous radial canals are likely to be synapomorphous for tribe Alchorneae. Secco (1999) found tribe Alchorneae to be monophyletic; monophyly of Alchorneae could be further tested by examina- tion of woods of the remaining genera of Alchorneae for occurrence of lysigenous radial canals. In addition to Alchorneae, lysigenous canals occur in Adenophaedra prealta, classified by Webster (1994) in tribe Bernardieae. As far as we know, no other member of Bernardieae possesses lysigenous radial canals. Given that the pol- len of Adenophaedra prealta also differs from other members of the genus but con- forms with the Alchornea-type (Punt 1962), we suggest that this species may be mis-

Downloaded from Brill.com10/07/2021 08:57:34AM via free access Hayden & Hayden — Wood anatomy of Acalyphoideae 229 placed within Bernardieae and may be better assigned to tribe Alchorneae. Similarly, we note that Wetria of tribe possesses anomalously large rays, possibly a pre- lysigenous condition and, therefore, a feature that could support Airy Shawʼs (1975) alignment of this genus with Alchornea (Webster 1994). Based on the foregoing, we observe that wood structure does not convincingly support the segregation of certain genera from Acalyphoideae as distinct families. The description above makes special note of Pera (Peraceae) and the genera of tribe Galeariae (Galearia, Microdesmis, and Panda) for possession of a number of fea- tures that are unusual for Acalyphoideae; however, they possess no wood feature that cannot be found in other acalyphoid woods. In the case of Pera, long considered to occupy an isolated position within the family (Webster 1994), we note that reports of lysigenous radial canals (Record & Hess 1943; Metcalfe & Chalk 1950; Pérez- Mogollón 1973; Détienne & Jacquet 1983; Dechamps 1985) suggest a potential, novel, taxonomic association for this poorly understood genus with tribe Alchorneae.

ACKNOWLEDGMENTS

We are grateful to Alberta M.W. Mennega (Uw) and Rusty Russel (USw) for their assistance in providing sectioning blocks and loaning prepared slides. This study was supported by Kuyk Chair funds from the University of Richmond.

REFERENCES

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Appendix — Guide to Previous Literature and List of Specimens Cited

Acalypha L. (Ilic 1991; Kanehira 1921a; Metcalfe & Chalk 1950; Record & Hess 1943; Williams 1936). – A. amentacea Roxb.: , Stern 2091 (CAHU, CLP, ILL, MICH, US), USw 31752. – A. brasiliensis Müll.Arg.: Brazil, Irwin 2060 (US, TEX), USw 15187. – A. diversifolia Jacq.: Peru, Williams 4391, USw 10230. – A. elliptica Sw.: Jamaica, Miller 1481, USw 5998. – A. macrophylla Kunth ex Poepp.: Peru, Williams 5185, USw 10298. – A. macrostachya Jacq.: Panama, Stern et al. 340 (MO), USw 16212. – A. mortoniana Lundell: Guatemala, Bartlett 12740 (MICH), USw 29790. – A. obovata Benth.: Peru, Williams 5134, USw 10290; Woytkowski 5896 (MO), USw 18282. – A. repanda Müll.Arg. var. denudata Müll.Arg.: Fiji, Smith 5005 (A, US), USw 30393. – A. stenoloba Müll.Arg.: Peru, Woytkowski 5856 (MO), USw 18275. – A. villosa Jacq.: Guatemala, Bartlett 12151 (MICH), USw 29682. Adelia L. (Metcalfe & Chalk 1950). – A. barbinervis Schltdl. & Cham.: , King 3938 (US), USw 27199; King 4376 (US), USw 27342. – A. ricinella L.: Puerto Rico, Britton & Kramer Y577 (Y), USw 4410. Adenophaedra (Müll.Arg.) Müll.Arg. (as Cleidion prealtum: Détienne & Jacquet 1983; Giraud 1983). – A. prealta (Croizat) Croizat: Brazil, Amazonas, Municipality Humayta, Krukoff 6391 (US), USw 7686; Krukoff 6570 (US), USw 7780; Krukoff 6649 (US), USw 7846. Agrostistachys Dalzell (Metcalfe & Chalk 1950). – A. borneensis Becc.: ʻMalaya,ʼ Uw 21397 (slide) = KEPw 5725. Alchornea Swartz (Détienne & Jacquet 1983; Giraud 1983; Manieri & Chimelo 1989; Metcalfe & Chalk 1950; Record & Hess 1943; Tortorelli 1956; Williams 1936). – A. brachygyne Pax & Hoffm.: Brazil, Amazonas, Municipality Humayta, Krukoff 6906 (US), USw 8035 = Uw 8035 (slide). – A. cordata Müll.Arg.: Costa Rica, Iica cco-23 Uw 20676 (slide). – A. cordifolia Müll.Arg.: Liberia, Jensen 1690, Uw 34214 (slide). – A. costaricensis Pax & Hoffm.: Panama, Canal Zone, Christopherson 198 (US), USw 127 (slide). – A. grandis Benth.: Colombia, van Rooden et al. 695, Uw 25667 (slide). – A. schomburgkii Klotzsch: Brazil, Matto Grosso, Krukoff 1414, Uw 19345 (slide). – A. triplinervia (Spreng.) Müll.Arg. var. laevigata Müll.Arg.: Suriname, Lanjouw & Lindeman 1216, Uw 1425 (slide). Alchorneopsis Müll.Arg. (Dechamps 1979; Détienne et al. 1982; Détienne & Jacquet 1983; Metcalfe & Chalk 1950). – A. floribunda (Benth.) Müll.Arg.: Brazil, Amazonas, Krukoff 6386 (US), USw 7682; Krukoff 6467 (US), USw 7749. – A. portoricensis Urb.: Domini- can Republic, Liali, Abbott 1603, USw 1908; Abbott 2591, USw 2021 (slide). – A. trimera Lanj.: Brazil, Maguire et al. 51802 (NY), USw 39804; Suriname, Stahel 356, USw 13143. Aparisthmium Endlicher (Dechamps 1979; Détienne et al. 1982; Détienne & Jacquet 1983; Giraud 1983; Metcalfe & Chalk 1950; Record & Hess 1943; Williams 1936). – A. cordatum (A. Juss.) Baill.: Brazil, Krukoff 8726 (NY), USw 38471; Maguire et al. 51902, USw 39321. Argythamnia P. Brown (Rickman & Hayden 1990). – A. blodgettii (Torr.) Chapm.: USA, Florida, Big Pine Key, Hayden 719 (URV); Hayden 1429 (URV); Hayden 2036 (URV). Bernardia Houstoun ex Miller. – B. dichotoma (Willd.) Müll.Arg.: Haiti, Gonaive I., Abbott & Leonard 3349, USw 1774. – B. tamanduana (Baillon) Müll.Arg.: Brazil, Amazonas, Munici- pality Humayta, Krukoff 6095 (US), USw 7477. Blumeodendron (Müll.Arg.) Kurz (Metcalfe & Chalk 1950). – B. kurzii (Hook. f.) J.J. Smith: Borneo, , (SAR), USw 30791. – B. tokbrai (Blume) J.J. Smith: Indonesia, , Boeea 8080 (MICH), USw 29131; USw 3356 (slide); USw 3406 (slide); USw 3608 (slide); USw 3780 (slide); USw 3810 (slide). Botryophora Hook. f. – B. geniculata (Miq.) Beumée ex Airy Shaw: Indonesia, Sumatra, Boeea 5494 (MICH), USw 28795; Boeea 6767 (MICH), USw 28882.

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Caryodendron Karst. (Détienne & Jacquet 1983; Giraud 1983; Metcalfe & Chalk 1950; Williams 1929). – C. grandifolium (Müll.Arg.) Pax: Brazil, Krukoff 5553, USw 9625. – C. orino- coense Karst.: Peru, Schunke-Vigo 4451 (US, F), USw 41881. Cephalomappa Baill. – C. paludicola Airy Shaw: Borneo, Sarawak, (SAR), USw 30793. Chaetocarpus Thwaites (Dechamps 1979; Détienne et al. 1982; Détienne & Jacquet 1983; Espinoza de Pernía 1987; Giraud 1983; Lebacq & Dechamps 1964, 1967; Manieri & Chimelo 1989; Metcalfe & Chalk 1950; Pfeiffer 1926; Record & Hess 1943). – C. castano- carpus (Roxb.) Thwaites: Burma, FPRL 797, USw 11194; , , SAN 56916 (SAN), USw 37104. – C. schomburgkianus (Kuntze) Pax & Hoffm.: USw = ʻWS 2702ʼ (slide). Cheilosa Blume – C. malayana (Hook. f.) Corner ex Airy Shaw: Malaysia, Mohd. Yatim 8520 (KEP), Uw 21414. Claoxylon A. Juss. (Ilic 1991; Metcalfe & Chalk 1950; Versteegh 1968). – C. africanum Müll.Arg.: Cameroon, Zenker 712 (K), USw 31266. – C. echinospermum Müll.Arg.: Fiji, Smith 5403 (A, US), USw 30421. – C. fallax Müll.Arg.: Fiji, Smith 6683 (A, US), USw 30498. – C. longifolium (Blume) Endl. & Hassk.: Indonesia, Sumatra, Boeea 1485 (MICH), USw 27797. – C. polot (Burm. f.) Merr.: Indonesia, , Schodde & Craven 3930 (CANB, US), USw 41767. – C. purpureum Merr.: Philippines, Stern 2307 (CAHU, CLP, ILL, MICH, US), USw 31927. – C. sandwicense Müll.Arg.: , Kauai I., Stern & Carlquist 1310 (US), USw 26006; Stern 2947 (US), Uw 18559 (slide); Carlquist 512 (RAS, Y), USw 15294. Cleidion Blume ( Ilic 1991; Meniado et al. 1970; Metcalfe & Chalk 1950; Williams 1936). – C. amazonicum Ule: Brazil, Amazonas, Municipality Humayta, Krukoff 6102 (US), USw 7480. – C. javanicum Blume: Burma, FPRL 4544, USw 11201. – C. spiciflorum Merr.: Indonesia, Sumatra, Boeea 68 (MICH), USw 27461. – C. vieillardii Baill.: New Caledo- nia, USw 4746. Clutia L. – C. abyssinica Jaub. et Spach: Ethiopia, Schimper 383, Uw 21375. – C. kiliman- dscharica Engl.: Tanzania, Volkens 620, Uw 25942. Cnesmone Blume (Metcalfe & Chalk 1950). Coccoceras Miq. (Metcalfe & Chalk 1950). – Coccoceras sp.: Indonesia, Sumatra, Boeea 5448 (MICH), USw 28784. Coelebogyne J. Smith (Metcalfe & Chalk 1950). – Coelebogyne sp.: , Queensland, Webster & Hyland 18896 (DAV). Conceveiba Aublet (Breteler & Mennega 1994; Dechamps 1979; Détienne et al. 1982; Détienne & Jacquet 1983; Giraud 1983; Metcalfe & Chalk 1950; Pfeiffer 1926; Record & Hess 1943). – C. guianensis Aubl.: Venezuela, Breteler 3795 (MER, NY, U, US, WAG), USw 35641. – C. krukoffii Steyerm.: Brazil, Krukoff 8396 (NY), USw 38410. – C. magnifica Steyerm.: Brazil, Amazonas, São Paulo, Belem Creek, Krukoff 8698, USw 19349. – C. simu- lata Steyerm.: Brazil, Amazonas, São Paulo, Belem Creek, Krukoff 8609, USw 19348. Dalechampia Plumier ex L. (Heimsch 1942; Metcalfe & Chalk 1950). – D. dioscoreifolia Poepp. & Endl.: Panama, Barro Colorado I., Wetmore & Abbe 11, Uw 21733. Dicoelia Benth. (Metcalfe & Chalk 1950; Mennega 1987). – Dicoelia sp.: Indonesia, Sumatra, Boeea 7794 (MICH), USw 29045. Discoclaoxylon (Müll.Arg.) Pax & Hoffm. (Normand 1955). Discoglypremna Prain (Lebacq & Dechamps 1964; Normand 1955; Normand & Paquis 1976). Galearia Zoll. & Moritzi (Forman 1966; Ilic 1991). – G. aristifera Miq.: Indonesia, Sumatra, Bartlett 6981 (MICH), USw 29324. – G. celebica Koord.: Indonesia, New Guinea, CSIRO 29249 (L), USw 34813. – G. filiformis (Blume) Pax: Indonesia, Sumatra, Boeea 7927 (MICH), USw 29083.

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Gavarretia Baill. (Détienne & Jacquet 1983; Metcalfe & Chalk 1950; Record & Hess 1943). – G. terminalis Baill.: Brazil, Amazonas, Manaos, Pensador, Ducke 228 (Y), USw 8693; Brazil, Krukoff 8916 (NY), USw 38269. Homonoia Lour. (Metcalfe & Chalk 1950). – H. javensis (Blume) Müll.Arg.: Indonesia, New Guinea, (L), USw 23894. – H. riparia Lour.: Philippines, Stern 2104 (CAHU, CLP, ILL, MICH, US), USw 31765. Koilodepas Hassk. – K. bantamense Hassk.: K.I.T. 44, Uw 21367. – K. hainanense (Merr.) Croizat: ʻMalayaʼ, KEPw 7863 = Uw 21400. Griseb. (Metcalfe & Chalk 1950). Leucocroton Griseb. (Metcalfe & Chalk 1950; Vales & Carreras 1987). – L. wrightii Griseb.: Cuba, Mathews & Crosby s.n., USw 3014. Macaranga Du Petit Thouars (Burgerstein 1909; Chattaway 1955; Ilic 1991; Kanehira 1921a; Lebacq & Dechamps 1964, 1967; Lecomte 1922; Metcalfe & Chalk 1950; Normand 1955; Normand & Paquis 1976; Sebastine 1955). – M. fimbriata S. Moore: New Guinea, B.W. 528, Uw 28074 (slide). – M. fragrans Perry: New Guinea, B.W. 4225, Uw 28929 (slide). – M. kilimandscharica Pax: ʻEast Africaʼ, Schlieben 1701, Uw 15924 (slide); USw (slide) = ʻWS 2173ʼ; USw (slide) = ʻWS 2174ʼ. – M. maingayi Hook. f.: USw (slide) = ʻWS 3842ʼ. – M. spinosa Müll.Arg.: Ivory Coast, Leeuwenberg 2920, Uw 6580 (slide). – M. polyadenia Pax & Hoffm.: New Guinea, Fokkinga 4547 = BW 1487, Uw 28378 (slide). – M. roxburghii Wight: India, Bhat 5013, Uw 32713 (slide). – M. zenkeri Pax: Cameroon, Zenker 962 (K), USw 31267. Mallotus Lour. (Baker 1917; Chattaway 1955; Ilic 1991; Kanehira 1921a, b; Lecomte 1925; Metcalfe & Chalk 1950; Normand 1955; Ohtani 1983; Ohtani & Ishida 1978; Pearson & Brown 1932). – M. claoxyloides Müll.Arg.: Australia, Forestry Comm. of N.S.W. 3860, Uw 21322 (slide). – M. confusus Merr.: Philippines, F.P.R.I. 585 (CLP), Uw 10783 (slide). – M. discolor F. Muell.: Australia, Uw 33125 (slide). – M. multiglandulosus (Reinw. ex Blume) Hurus.: Philippines, F.P.R.I. 546 (CLP), Uw 33126 (slide). – M. polyadenus F. Muell.: Australia, Queensland, Webster & Hyland 18875 (DAV), Uw 31199 (slide). – M. ricinoides (Pers.) Müll.Arg.: Philippines, F.P.R.I. 522 (CLP), Uw 10758 (slide). Mareya Baill. (Breteler et al. 1997; Giraud 1983; Metcalfe & Chalk 1950; Normand 1955; Normand & Paquis 1976). – M. micrantha (Benth.) Müll.Arg.: Ivory Coast, USw 24337. Megistostigma Hook. f. – M. malaccense Hook. f.: Indonesia, Sumatra, Boeea 1389 (MICH), USw 27745; Boeea 1975 (MICH), USw 27943. Melanolepis Reichb. f. & Zoll. (Ilic 1991). – M. multiglandulosa (Reinw. ex Blume) Reichb. f. & Zoll.: Indonesia, Sumatra, Boeea 3645 (MICH), USw 28334; Mariana Islands, Dutton 113 (US), USw 39070. Mercurialis L. (Metcalfe & Chalk 1950). Microdesmis Planch. (Giraud 1983; Lebacq & Dechamps 1964, 1967; Metcalfe & Chalk 1950; Normand 1955; Stern 1967). – M. caseariifolia Planch.: Indonesia, Sumatra, Bartlett 7621 (MICH), USw 29504. – M. puberula Hook. f. ex Planch.: Liberia, Cooper Y. 15136 (Y), USw 4826. Moultonianthus Merr. – M. leembruggianus (Boerl. & Koord.) van Steenis: Malaysia, Sarawak, Omar 7884, Uw 21418; Indonesia, van Balgooy & van Setten 5429, Uw 31770 (slide). Necepsia Prain (Giraud 1983; Metcalfe & Chalk 1950; Normand 1955). – N. afzelii Prain: Liberia, Cooper Y. 13726 (Y), USw 4509; Cooper Y. 15178 (Y), USw 4856. Neoscortechinia Pax (Ilic 1991; Metcalfe & Chalk 1950). – N. kingii (Hook. f.) Pax & Hoffm.: Indonesia, Sumatra, Boeea 3422 (MICH), USw 28259. – N. nicobarica (Hook. f.) Pax & Hoffm.: Indonesia, New Guinea, FPAW 15909 (L), USw 34315; FPAW 29707 (L), USw 35271.

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Neotrewia Pax & Hoffm. (Meniado, Robillos & Zamuco 1970). – N. cumingii (Müll.Arg.) Pax & Hoffm.: Philippines, FPRI 316 (CLPW), USw 30205; Stern 2190 (CAHU, CLP, ILL, MICH, US), USw 31845. Octospermum Airy Shaw. – O. pleiogynum Airy Shaw: New Guinea, B.W. 12446, Uw 18239; B.W. 10124, Uw 29042. Omphalea L. (Metcalfe & Chalk 1950; Barajas Morales & Echenique-Manrique 1976; Record & Hess 1943). – Omphalea sp.: Suriname, Maguire et al. 54825 (NY), USw 39561. – O. triandra L.: Jamaica, Miller 1482, USw 5999. Panda Pierre (Forman 1966). – P. oleosa Pierre: Zaire, ʻBelgian Congoʼ, (BR), USw 18341. Pera Mutis (Babos & Borhidi 1978; Dechamps 1985; Détienne et al. 1982; Détienne & Jacquet 1983; Espinoza de Pernía 1987; Giraud 1983; Ilic 1991; Lebacq et al. 1973; Metcalfe & Chalk 1950; Peréz-Mogollón 1973; Record & Hess 1943; Williams 1936). – P. bicolor (Klotzsch) Müll.Arg.: Brazil, Amazonas, Municipality Humayta, Krukoff 6905 (US), USw 8034. – P. ferruginea (Schott) Müll.Arg.: Venezuela, Steyermark 86353 (NY), USw 39586. – P. glabrata (Schott) Baill.: Suriname, Stahel 273, USw 13074. – P. glomerata Urb.: Haiti, Tortue I., Leonard 12523, USw 4280. – P. obovata Baill.: Brazil, , (HBR, US), USw 25562. – P. oppositifolia Müll.Arg.: Cuba, Pinar del Rio, Fors 951, USw 21463. Podadenia Thwaites. – Podadenia sp.: Indonesia, Sumatra, Krukoff 4074, USw 6309. Pogonophora Miers ex Benth. (Dechamps 1979; Détienne et al. 1982; Détienne & Jacquet 1983; Metcalfe & Chalk 1950; Record & Hess 1943; Stern 1967). – P. schomburgkiana Miers ex Benth.: Suriname, Bosbeheer 10741, USw (slide) = Uw 11195. Pseudoagrostistachys Pax & Hoffm. (Metcalfe & Chalk 1950). Ptychopyxis Miq. (Metcalfe & Chalk 1950). – P. bacciformis Croiz.: Malaysia, Borneo, SAN 58065 (SAN), USw 37151. – P. kingii Ridley: Malaysia, Sabah, SAN 58068 (SAN), USw 37116. Pycnocoma Benth. (Metcalfe & Chalk 1950). Ricinus L. (Fahn et al. 1986; Ilic 1991; Messeri 1938; Metcalfe & Chalk 1950; Schweingruber 1990). – R. communis L.: Mexico, King 4574 (US), USw 27390; USA, (US), USw 8879 (slide). Syndyophyllum Laut. & Schum. (Ilic 1991). – S. excelsum Laut. & Schum.: New Guinea, B.W. 12477, Uw 18257; Fokkinga 4124 = B.W. 1844, Uw 28511. Tragia Plumier ex L. (Metcalfe & Chalk 1950). Trewia L. (Metcalfe & Chalk 1950; Pearson & Brown 1932). – T. nudiflora L.: India, (US), USw 8880 (slide); Burma, FPRL 897 (FPRL), USw 11352. Hook. f. (Stern 1967). – T. malayana Hook.: Indonesia, K.I.T. E-1031, Uw 21374; ʻMalayaʼ, KEPw 1788 = Uw 21401. Wetria Baill. (Metcalfe & Chalk 1950). – W. insignis (Steud.) Airy Shaw: Malaysia, Pawanchee s.n., KEPw 196 = Uw 21422.

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