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1
2Sapindaceae
3Sapindaceae Juss., Gen.: 246 (1789), nom. cons., “Sapindi”.
4Aceraceae Juss. (1789), nom. cons.
5Hippocastanaceae A. Rich. (1823), nom. cons.
6Dodonaeaceae Link (1831).
7Koelreuteriaceae J. Agardh (1858).
8P. ACEVEDO-RODRI
´GUEZ,P.VAN WELZEN,F.ADEMA,AND R.W.J.M. VAN DER HAM
9Trees, treelets, shrubs, lianas or herbaceous clim-
10 bers; cork superficial; stems of climbing species
11 (i.e., Serjania,Paullinia,Urvillea,Houssayanthus,
12 and Thinouia) usually with multiple vascular
13 cylinders. Leaves pinnately or ternately compound
14 or palmate, or rarely simple, alternate, rarely
15 opposite; proximal leaflets seldom reduced, re-
16 flexed, and covering the stem to resemble a pair
17 of stipules (pseudostipules), distal leaflet in most
18 arboreal and shrubby species rudimentary; sti-
19 pules present only in climbing species, minute
20 to large. Inflorescences axillary, terminal, pseudo-
21 terminal, cauliflorous or ramiflorous, thyrso-
22 paniculate, racemose, spicate, or fasciculate, or
23 flowers solitary. Flowers 5-merous, regular, or
24 less often 4-merous and obliquely zygomorphic,
25 bisexual or more often functionally unisexual by
26 reduction (plants monoecious or rarely dioe-
27 cious); sepals distinct or connate at base; petals
28 usually white or light yellow, rarely 0, usually
29 ornamented by an adaxial appendage; appen-
30 dages variously shaped, mostly petaloid, simple,
31 bifurcate, or hood-shaped, basally adnate to the
32 petal or just a prolongation of petal margins,
33 concealing the nectary; disk extrastaminal, annu-
34 lar or unilateral, often lobed, cup-shaped or dish-
35 shaped, very rarely on both sides of the stamens
36 or intrastaminal; stamens (3–)5–8(–30); filaments
37 distinct or connate at base, equal or unequal in
38 length; anthers dorsifixed or basifixed, introrse,
39 opening by longitudinal slits; sterile stamens
40 present in pistillate flowers; gynoecia syncarpous,
41 (1–)3(–8)-carpellate; carpels with 1, 2, or excep-
42 tionally many (7–8 in Xanthoceras,8inMagonia)
43 ovules; style terminal or exceptionally gynobasic
44 (Deinbollia), 2–3-branched, or with simple, 2–3-
45 lobed, capitate stigma, sometimes (Acer) the style
46branches elongate and the style nearly 0; pistil
47usually rudimentary in staminate flowers. Fruit
48a septifragal or loculicidal capsule, a schizocarp
49with winged or non-winged mericarps, baccate or
50rarely a drupe. Seeds sessile or exceptionally (Dis-
51tichostemon) subtended by a funiculus, variously
52shaped, exalate or rarely winged, naked, with a
53partial to complete sarcotesta, or an arillode
54(arising from the integuments); embryo oily or
55starchy, lacking endosperm, notorhizal or loma-
56torhizal with straight, curved or plicate, fleshy
57cotyledons, the radicle often separated by a deep
58fold in the testa that forms a radicular pocket.
59Mostly tropical or subtropical, with a few
60genera extending to sub-temperate zones; 141
61genera and about 1,900 species.
62VEGETATIVE MORPHOLOGY. Most genera of
63Sapindaceae are predominantly medium-sized to
64large emergent trees or erect shrubs, less often
65they are tendrilled lianas or understory palm-
66like treelets, exceptionally sub-shrubs or scandent
67shrubs. The arboreal and fruticose habits are
68widespread throughout the distributional range
69of Sapindaceae, while treelets, lianas, and sub-
70shrubs are restricted to the tropics. There are
71about 500 species of lianas, all of which are exclu-
72sive to the Neotropics (with the exception of
73several species found in the Paleotropics), parti-
74cularly to tribe Paullinieae (Cardiospermum,
75Houssayanthus,Lophostigma,Paullinia,Serjania,
76Urvillea, and Thinouia), accounting for ca. 60% of
77the Sapindaceae species in the Neotropics. Sub-
78shrubs are rare and are known to occur in dry
79vegetation subject to periodic fires. In the Paleo-
80tropics, a few species of Allophylus,Laccodiscus,
81and Lepisanthes are known to be scandent shrubs,
K. Kubitzhi (ed.), Flowering Plants, Eudicots: Sapindales, Cucurbitales, Myrtaceae,
The Families and Genera of Vascular Plants X, DOI 10.1007/978-3-642-14397-7_17,
#Springer-Verlag Berlin Heidelberg 2010
371
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82 not showing any active mechanism for ascending
83 the host plants.
84 Exudates for the most part are odorless and
85 colorless. However, exudates are red in Pometia,
86 or white in numerous species of Paullinia and
87 Serjania.
88 Stems are for the most part terete and
89 smooth. However, many of the climbing species
90 have stems that are deeply furrowed, or sharply to
91 obtusely angled, and a few have thorny stems.
92 Bud scales are present in a few taxa, being
93 predominant in subfamily Hippocastanoideae.
94 These are simple and accrescent with the growing
95 buds. Bud scales have also been reported for a few
96 tropical genera, including Exothea,Melicoccus,
97 Talisia, and Sapindus that bear minute scales, or
98 Talisia (Fig. 84B) and Pseudopteris that bear large
99 leaf-like cataphylls. Cataphylls are clustered at the
100 ends of branches, on axillary buds, and at the
101 base of inflorescences. Their size and form vary
102 considerably. In some species they can be up to
103 25 cm long, while in others they are pinnatifid
104 and a few cm in length.
105 Leaves are predominantly spirally arranged,
106 and variously compound, less often opposite,
107 simple or digitate. Compound leaves include
108 the following types: palmate, pinnate, bipinnate,
109 tripinnate, trifoliolate, biternate, triternate, or
110 a combination of these. A peculiar character
111 of numerous Sapindaceae with pinnately com-
112 pound leaves is the presence of a single terminal
113 rudimentary leaflet or process (Fig. 85A). At
114 first glance, this structure may be confused with
115 an undifferentiated leaflet primordium present in
116 other families such as the Meliaceae. However, in
117 Meliaceae, this structure is produced in pairs, and
118 slowly developed into new leaflets. Leaflets are
119 predominantly entire, but there is variation,
120 with some genera having entire, crenate or serrate
121 leaflets. Venation is variable and represented by
122 brochidodromous, cladodromous, craspedodro-
123 mous, mixed-craspedodromous, semi-craspedo-
124 dromous, and palinactinodromous types (Hickey
125 1979). Stipules are largely absent in Sapindaceae,
126 and restricted to the genera of the Paullinieae.
127 Pseudostipules are present in a few species of
128 the paleotropical genera Alectryon,Blighiopsis,
129 Chouxia,Cupaniopsis,Eriocoelum,Glenniea,
130 Haplocoelopsis,Laccodiscus,Lepisanthes,Mac-
131 phersonia,Otonephelium,Placodiscus, and Pome-
132 tia (Weberling 1976). Although superficially
133similar to stipules, these are in reality proximal
134leaflets reduced in size, which often clasp the
135stem. Petioles and leaf rhachises are terete,
136angled, carinate, sulcate, and sometimes narrowly
137to broadly winged. The base of petioles and
138the petiolules are more often enlarged, with the
139adaxial portion very often depressed or furrowed,
140or only very rarely nearly cylindrical.
141Tendrils are found only in genera of the Paul-
142linieae. They are opposite and coiled, and seem to
143be homologous to the proximal pair of cincinni
144(or drepania) of their thyrsoid inflorescences.
145Very often, the portion of the inflorescence
146above the tendrils is aborted, resulting in a short
147axillary branch that produces a pair of opposite
148tendrils in its distal portion.
149The indumentum in the family is quite
150variable. Plants are either glabrous or exhibit
151several different kinds of indumentum, which
152are predominantly composed of simple, erect or
153curly, non-glandular trichomes. Less frequently,
154the indumentum may include multicellular-
155glandular, papilliform, fasciculate, stellate tri-
156chomes, or peltate scales. Trichomes occur as a
157pure stand or as a mixture of different types, and
158are often classified as puberulent, appressed-
159pubescent, tomentose, tomentulose, sericeous,
160velutinous, setiferous, hirsute, woolly, pilose,
161pilosulous, or furfuraceous. Stiff, irritating hairs
162occur on the outer surface of fruits of Cnesmo-
163carpon and Jagera species.
164VEGETATIVE ANATOMY. A detailed anatomical
165survey of Dodonaeoideae and Sapindoideae,
166which covers both vegetative and reproductive
167organs, was presented by Radlkofer (1890) and
168used in explaining his classification. Among
169other things, Radlkofer described carefully the
170location and structure of secretory cells that con-
171tain saponins and mucilaginous cells, which
172occur in the leaf epidermis. An important trait
173discovered by Radlkofer is the constant presence
174of a cylinder of sclerenchyma in the pericycle,
175which consists of phloem fibers and stone
176cells. This structure was observed in all genera
177of Sapindaceae as circumscribed by Radlkofer
178(1890) (¼Dodonaeoideae þSapindoideae), with
179the notable exception of Xanthoceras and Valen-
180zuelia (the latter renamed as Guindilia), in which
181the ring of sclerenchyma is not continuous;
182incidentally, Guindilia has opposite leaves. The
372 P. Acevedo-Rodrı
´guez et al.
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183 sclerenchyma cylinder is also constantly
184 present in tribe Hippocastaneae, whereas in
185 tribe Acereae sometimes (e.g., Acer negundo)it
186 is well developed but sometimes (A. pseudoplata-
187 nus,Dipteronia) it is represented only by isolated
188 strands of fibers.
189 Nodes of Sapindaceae are trilacunar, rarely
190 also 5-lacunar in Hippocastaneae. The vessel seg-
191 ments usually have simple perforation; scalari-
192 form perforation is rarely seen in Sapindoideae,
193 and more often in Hippocastaneae. Rays are
194 mostly 1-seriate, but in Acereae mixed 1- and
195 pluriseriate rays are found. A comprehensive
196study of wood anatomy of Dodonaeoideae and
197Sapindoideae was published by Klaassen (1999).
198Numerous woody vines of the Sapindoideae
199present anomalous secondary thickening of their
200stems. The most salient feature of this anomalous
201thickening can be described as multistelate,
202where the stem has a single central stele sur-
203rounded by three, five, or up to ten peripheral
204steles. Anomalous secondary thickening is pre-
205dominant in Serjania (Fig. 76A, D) and Paullinia
206(Fig. 76B), where 58% and 12%, respectively of
207the species show some kind of anomaly
208(Acevedo-Rodrı
´guez 1993).
Fig. 76. Sapindaceae. Stem cross sections of Sapinda-
ceae vines. ASerjania sp., stem with a central stele
and two smaller peripheral steles. BPaullinia alata,
stem with a central stele and three smaller peripheral
steles. CPaullinia ingifolia, stems with single stele
and white latex. DSerjania grandifolia Radlk., vas-
cular tissue produced in alternate concentric layers
alternating with connective tissue; note the white
latex. (orig.)
Sapindaceae 373
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209 INFLORESCENCES. The inflorescences in Sapin-
210 daceae are variously shaped thyrses with lateral
211 dichasia, cincinni, drepania, or reductive forms
212 thereof (for details, see Radlkofer 1890: 178 seq.).
213 They are axillary, distal, cauliflorous, or supra-
214 axillary, solitary or fasciculate. Some genera have
215 racemes that seem to be derived from thyrsoid
216 ancestors. The flowers within the inflorescences
217 are predominantly unisexual or bisexual, but with
218 a strong tendency for one sex to predominate.
219 FLOWERS. Pedicels are usually conspicuous,
220 and have an abscission zone or articulation any-
221 where from base to near the apex, less often they
222 are inconspicuous or non-articulated. The peri-
223 anth is more often 5-merous, but there is consid-
224 erable variation in the number of parts. Sepals are
225 distinct (Figs. 77, 78) to completely connate
226 (Fig. 86A, D), and may be as few as three or as
227 many as ten. They are usually of similar size and
228 shape, or less often dimorphic. The petals are
229 distinct, with imbricate aestivation, inserted
230 on the base of an extrastaminal nectary disk.
231 Their number varies from four to six, in addition
232 to the 5-merous corolla. However, there are
233 numerous genera or species that completely lack
234 a corolla. Petals are erect (Fig. 83C) or reflexed
235 (Fig. 78), those of Sapindoideae with an adnate
236 adaxial petaloid appendage (Figs. 83, 84), or
237 with extended inrolled basal margins lacking
238 appendages, or lacking appendages and in-
239 rolled margins altogether (Fig. 86E). The petaloid
240 appendages are simple (Fig. 84D, G), bifid, hood-
241 shaped (Fig. 83D), or corniform, and very often
242 sericeous or tomentose. Leinfellner (1958) ana-
243 lyzed these appendages, emphasizing their peltate
244 nature. The disk is extrastaminal, annular
245 (Fig. 77), more rarely amphistaminal (Fig. 80A, B)
246 or intrastaminal, cup-shaped, 2-, 4-, 5(–8)-lobed
247 (Fig. 83C), of elongated corniform lobes, unilat-
248 eral and semi-annular, or rudimentary, with vari-
249 ous indumenta, or glabrous altogether. In many
250 genera, nectar is produced as a reward for polli-
251 nators. The number of stamens is more often
252 eight but there is considerable variation, with
253 genera containing from five to eight or from
254 eight to ten. Stamens are sometimes as few as
255 four in Cupania or Dictyoneura, or as many
256 as 20 in Hornea,30inDeinbollia, and 74 in
257 Distichostemon. Filaments are glabrous or vari-
258 ously pubescent, of equal or unequal lengths,
259erect, spreading, or sigmoid. Anthers are basi-
260fixed or dorsifixed, oblong to linear or elliptic to
261ovate with an obtuse, apiculate or retuse apex,
262opening along longitudinal slits. The ovary is
263syncarpous and usually 3-carpellate. However,
264there are numerous genera possessing 2-carpellate
265ovaries in addition to the 3-carpellate ones. Blo-
266mia and sometimes Alectryon and Nephelium
267have unicarpellate gynoecia. Chytranthus and
268Radlkofera have gynoecia with up to eight
269carpels. The septae are complete, with the ovary
270containing the same number of locules as carpels,
271except in Melicoccus (Fig. 77) and Zollingeria,
272where the septae are partially developed, result-
273ing in a unilocular ovary. Ovules are anatropous,
274hemitropous, or campylotropous, one per carpel
275in subfam. Sapindoideae, or two per carpel in
276subfam. Dodonaeoideae, and 7–8 in Magonia
277(Dodonaeoideae) and Xanthoceras (Xanthoceroi-
278deae); placentation is axile, commonly in the
279middle or less often basal, or apical. The style is
280terminal and simple in all Sapindaceae except
281for Deinbollia, which has a gynobasic or a sub-
282terminal style; in Dipteronia and some Acer, the
283style can be very short, or lacking, and has two
284prominent stigmatic style branches (Fig. 80C, H).
285The stigmatic surface is variable, very often re-
286presented by a line along the stigmatic branches,
287or by as many lines as carpels along the distal
288portion of a simple style. Stigmas are sometimes
289capitate, elongated-cylindrical, or lobed.
Fig. 77. Sapindaceae. Melicoccus bijugatus Jacq.
APistillate flower. BDitto, longitudinal section. (Ace-
vedo-Rodrı
´guez 1996; reproduced with permission of
the artist Bobbi Angell)
374 P. Acevedo-Rodrı
´guez et al.
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290 For the aspect of oblique floral monosymme-
291 try, see the ontogenetic studies of Koelreuteria by
292 Ronse Decraene et al. (2000).
293 SEX DISTRIBUTION. Functionally bisexual flow-
294 ers are rare among Sapindaceae; they are known
295 from Acer,Aesculus,Bizonula,Dodonaea,Ex-
296 othea,andHandeliodendron. Most Sapindaceae
297 are usually labeled as polygamous, and there is
298 evidence for a wide distribution within the family
299 of a particular sexual system, duodichogamy,
300 which has been unveiled in such distantly related
301 taxa as Cupania (Bawa 1976), Acer,Dipteronia,
302 Hippocastanum,Deinbollia, and Koelreuteria (de
303 Jong 1976), Sapindus (Subba Reddi et al. 1983),
304 Serjania (Acevedo-Rodrı
´guez 1993), Talisia (Ace-
305 vedo-Rodrı
´guez 2003), and Paullinia (Somner,
306 unpubl. data).
307 Duodichogamy implies a sequence of three
308 distinct phases of flowering in which all flowers
309 of a given individual are in the same phase. Dur-
310 ing the first phase, male flowers release functional
311 pollen; they have a reduced pistil. When they are
312 dropped, female flowers appear on the same
313 inflorescence, which have a well-developed pistil
314 and short stamens with indehiscent anthers.
315 These phases may overlap for a few days. After
316 fertilization of the female flowers (from flowers of
317 an individual that is in a different phase), in a
318 third phase hermaphrodite flowers with well-
319 developed stamens and pistils appear but usually
320 are effectively only male. Sometimes, as in Acer,
321 the third phase may be female when the first
322female phase is repressed for some reason. In
323many Sapindaceae, the first or usually the last
324phase may be missing, so that the reproductive
325system is dichogamous. Since self-incompatibility
326seems to be frequent in Sapindaceae, (duo)
327dichogamy acts as a barrier against selfing. Most
328genera and species of the family share the posses-
329sion of male and apparently hermaphrodite but
330functionally female flowers, and it is likely that
331they are dichogamous; in the absence of experi-
332mental data, however, this remains unproven. In
333the generic descriptions, these cases are labeled
334“falsely polygamous”. True dioecy is more rarely
335documented in the family, as for instance for
336some Acer,Xerospermum noronhianum, and
337some Nephelia (van Welzen 1989). It is likely
338that duodichogamy is basal in the family
339(de Jong 1976), and van Welzen (1989) supposes
340that it is symplesiomorphic.
341POLLINATION. The open, white, fragrant, and
342nectar-secreting flowers of tribe Hippocastaneae
343appear primarily bee-pollinated, such as several
344Aesculus, among which the colored spot on the
345petals functions as a nectar guide. Nevertheless,
346such flowers can also be attractive to other kinds
347of pollinators including Lepidoptera, and species
348of Aesculus sect. Pavia, particularly the red-flowered
349Ae.pavia, are pollinated by humming birds. Sim-
350ilarly, in Billia the white-flowered B.columbiana
351is probably bee-pollinated, whereas the red flow-
352ers of B.hippocastanum suggest bird pollination
353(Forest et al. 2001). In Acer, the transition from
354entomogamy to anemophily is accompanied by
355one from dichogamy to dioecy (de Jong 1976;
356Hesse 1979). In Sapindoideae, apart from nectar
357also pollen appears to be an important reward,
358because male flowers are not only much more
359abundant than functionally female flowers, but
360they also have a longer duration and are much
361more visited than the females (van Welzen 1989),
362which implies that the anthers in the female flow-
363ers act as mimics.
364POLLEN MORPHOLOGY. The literature on pollen
365morphology of Sapindaceae is quite extensive.
366The most complete work at the family level is
367that of Muller and Leenhouts (1976; including
36878% of genera), where pollen types were evalu-
369ated in regard to their systematic significance.
370More recent progress has been made by several
Fig. 78. Sapindaceae. Vouarana anomala.A,BMale flow-
ers. CPetals, ventral view. DPistillode with subtaining
annular disk. EPistillode. (Acevedo-Rodrı
´guez 1997;
reproduced with permission of the artist Bobbi Angell)
Sapindaceae 375
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371 workers (van der Ham 1990; Acevedo-Rodrı
´guez
372 1993; Ferrucci and Anzo
´tegui 1993), but their
373 analyses apply only to some of the genera.
374 The present overview is based on pollen data for
375 137 of the 141 genera currently recognized in
376 Sapindaceae.
377 Sapindaceae pollen grains are usually iso-
378 polar or subisopolar monads. Tetrads occur
379 only in Magonia (Fig. 79F). Pollen grain size is
380 usually between 20 and 30 mm, and the grains are
381 oblate to prolate in shape. Colporate pollen is
382 usually suboblate to prolate (Fig. 79A–E),
383 whereas pollen with small apertures (porate, bre-
384 vicolporate) or with connected apertures (syncol-
385 porate, parasyncolporate) has a more oblate
386 shape (Fig. 79G–L). The equatorial outline is
387 almost circular (Fig. 79C, D) to bluntly triangular
388 (Fig. 79G–L); the meridional outline is almost
389 circular to more or less elliptic (Fig. 79B, E, H).
390 Generally, Sapindaceae pollen is 3-aperturate
391 (Fig. 79A–L), but often small percentages of
392 2-and 4-aperturate grains co-occur. Colporate
393 pollen (Fig. 79A–C, E, F) is the commonest,
394 being known from many genera in all four sub-
395 families. It is a relatively basic type found in many
396 other angiosperm families. Several other types
397 are more restricted. Syncolporate (Fig. 79J) and
398 parasyncolporate pollen (Fig. 79K), with and
399 without apocolpial fields, respectively, are
400 known only in subfamily Sapindoideae, being
401 present in most Cupanieae, Alectryon,Schlei-
402 chera,Castanospora,Tristira, and Tristiropsis.
403 Parasyncolporate and syncolporate are not clear-
404 cut character states. Moreover, several genera (e.g.,
405 Alectryon,Arytera,andCupania) possess both
406 colporate and (para)syncolporate pollen, and
407 often intermediates as well (van der Ham 1990;
408 van Bergen et al. 1995). Small ectoapertures occur
409 in brevicolporate and porate pollen found in a
410 few species of Allophylus (Fig. 79H), Lepisanthes,
411 Pometia (Fig. 79L), Talisia, and tribe Paullinieae.
412 Pollen of several Paullinieae is heteropolar with a
413 demisyncolporate aperture system, e.g., species of
414 Serjania,Cardiospermum,Houssayanthus,and
415 Urvillea (Fig. 79G, I), which have pollen grains
416 that are proximally syncolporate, and distally
417 have a short demicolpus (Ferrucci and Anzo
´tegui
418 1993). Distichostemon pollen has indistinct ectoa-
419 pertures, and a few species of Harpullia have
420 pollen without recognizable ectoapertures (crypto-
421 aperturate; Fig. 79D). Sapindaceae pollen grains
422nearly always have lalongate, elliptic to subcircular
423endoapertures, though usually hidden by the
424ectoaperture margins. Acer pollen may have aper-
425tures without or with indistinct endoapertures.
426The exine is usually clearly stratified, showing
427a tectum, a columellate infratectal layer, and a
428nexine. In Diplopeltis,Distichostemon, and Dodo-
429naea, the infratectal layer is granular/columellate,
430which might relate to wind-pollination. Mostly,
431the nexine consists of a distinctly delimited foot
432layer and endexine. The endexine is thin in non-
433apertural parts, thickens toward the apertures,
434and is maximal along and under the apertures.
435The ornamentation of the exine shows much
436variation. Striate (Fig. 79A, E), rugulate (Fig. 79H, J),
437perforate (Fig. 79G), and intermediate types are
438most common; verrucate (Fig. 79C), scabrate
439(Fig. 79B), and reticulate (Fig. 79D, L) types are
440less frequent. In subfamilies Xanthoceroideae,
441Hippocastanoideae, and Dodonaeoideae, rugu-
442late pollen is rare and psilate pollen entirely
443absent. In subfamilies Hippocastanoideae and
444Sapindoideae, verrucate and scabrate ornamenta-
445tion are absent or rare (present only in some Acer
446species, and in the Malesian genera Cubilia,
447Dimocarpus,Jagera, and Trigonachras), but it
448is common in Xanthoceras and Dodonaeoideae
449(e.g., Dodonaea,Exothea,Filicium,Ganophyllum,
450and Zanha). The tectum is usually perforate, but
451less densely and less distinctly so toward the
452apertures. Perforate types are linked with more
453or less reticulate types. Simple reticulate orna-
454mentation is rare (e.g., Pometia; Fig. 79L). Striate
455ornamentation is often associated with a colporate
456aperture system (Fig. 79A, E), and rugulate with a
457(para)syncolporate aperture system (Fig. 79J, K),
458which probably reflects a functional, harmome-
459gathic relation. Ornamentation often varies with-
460in genera, while, e.g., Dimocarpus longan shows
461remarkable infraspecific variation: striate, perfo-
462rate, scabrate, and intermediates thereof (van der
463Ham 1990).
464Muller and Leenhouts (1976) recognized 12
465pollen types, which were based largely on ectoa-
466perture features and pollen grain shape. The col-
467porate spheroidal type A (Fig. 79A–C, E), found
468in most genera, was considered basic. Type B1
469and type B2 (Fig. 79J, K; many genera) include
470spheroidal to oblate pollen with a (para)syncol-
471porate aperture system. These types intergrade
472and are combined here (type B1/2). This applies
376 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
Sapindaceae 377
Uncorrected Proof
473 also to the brevicolporate oblate types A1 and C1
474 (Fig. 79L; 8 genera; type A1/C1). Type C2
475 (Fig. 79G, I; 4 genera) includes oblate pollen
476 with a heteropolar aperture system, and type C3
477 (Fig. 79H; 2 genera) oblate pollen with porate
478 apertures. Type D was described as 2-porate.
479 However, of the two genera mentioned, Lophos-
480 tigma has 4-aperturate pollen, while the very rare
481 2-porate Allophylus pollen is probably a variant
482 of the normal 3-porate type (Fig. 79H). Type E
483 (Fig. 79F) represents the tetrad pollen of Mago-
484 nia. Type F includes colporate (per)prolate, equa-
485 torially constricted pollen, and is found only in
486 Diplopeltis huegelii. Type G was described as
487 spheroidal pollen with protruding endoapertures
488 without ectoapertures, occurring in Distichoste-
489 mon. However, indistinct ectoapertures appeared
490 to be present; therefore, this pollen type is con-
491 sidered here as a variant of the basic colporate
492 type. Type H (Fig. 79D) includes the spheroidal
493 cryptoaperturate pollen occurring in a few
494 Harpullia species. As far as possible, the pollen
495 type(s) occurring in each genus is/are indicated
496 in the generic descriptions below.
497 Comparison with the molecular phylogenetic
498 trees of 64 genera by Harrington et al. (2005)
499 suggests that the colporate spheroidal type A is
500 indeed basal in the Sapindaceae, subfamilies
501 Xanthoceroideae, Hippocastanoideae, and Dodo-
502 naeoideae being characterized or heavily dominated
503 by this pollen type. Pollen types E, F, and H are
504 each restricted to one or a few species of Dodo-
505 naeoideae. In subfamily Sapindoideae, the com-
506 monest pollen types A and B1/2 occur largely
507 clustered—for example, type A in clade A (Lepi-
508 santheae-Sapindeae) and clade B (Nephelieae),
509 and type B1/2 in clade C (Cupanieae), whereas
510 type A1/C1 is found in a few isolated taxa. Pollen
511types C2 and C3 are restricted to clade D (Paulli-
512nieae-Thouinieae).
513EMBRYOLOGY. Pollen grains are 2-celled when
514shed. The ovule is anatropous to hemitropous
515or campylotropous (at least after fertilization),
516bitegmic, crassinucellate, and the micropyle is
517formed by both integuments or rarely only
518by 7the inner one. The chalazal megaspore devel-
519ops into a Polygonum type embryo sac. The
520endosperm is nuclear, and the tissue is digested
521before it becomes cellular (Davis 1966; Johri et al.
5221992).
523FRUIT AND SEED. The vast majority of Sapin-
524daceae genera have trilocular fruits, although
525many also have bilocular, unilocular, or some-
526times quadrilocular fruits. In addition, fruits of
527several genera are predominantly unilocular due
528to the abortion of carpels, but sometimes they
529may develop more than one locule. Partly trilo-
530cular fruits are produced in Koelreuteria and
531Euphorianthus because of their distally incom-
532plete septa. Fruits of Chytranthus and Radlkofera
533contain up to 8 locules. Fruits are either indehis-
534cent or dehiscent. Indehiscent fruits are baccate
535with a leathery to crustose pericarp, or exception-
536ally a drupe with a stony (Tristiropsis) or woody
537(Hypelate) endocarp. Indehiscent fruits are either
538unlobed, deeply lobed, or made of monocarps,
539these of the same number as the carpels or less,
540due to abortion. Dehiscent fruits are capsular
541or schizocarpic. The former are predominantly
542loculicidal and early dehiscent, or less often
543septicidal, septifragal, calyptrate, tardily dehis-
544cent, or incompletely dehiscent. Their texture
545varies from woody, coriaceous to papery or
546membranous. Schizocarpic fruits are
Fig. 79. Sapindaceae. Pollen micrographs. A–FSubfam.
Dodonaeoideae. G–LSubfam. Sapindoideae. All micro-
graphs in polar view, except for B,E, and H, which are
in equatorial view. AErythrophysopsis aesculina, colpo-
rate, striate pollen grain, 2,100. BDodonaea truncatialis,
colporate, scabrate pollen grain (prolate equatorial view)
with mesocolpial scabrae in a coarser pattern, 1,500. C
Exothea copalillo, colporate, loosely scabrate-verrucate
pollen grain, 3,150. DHarpullia crustacea, cryptoaper-
turate, reticulate (muri with scabrae) pollen grain, 2,500.
EHandeliodendron bodinieri, colporate, striate pollen
grain (prolate equatorial view) showing verrucate colpus
membrane, 2,000. FMagonia pubescens, pollen tetrad
(upper pollen grain in polar view) with striate-gemmate
ornamentation, 900. G–IUrvillea ulmacea, heteropolar,
demisyncolporate, perforate-reticulate pollen grains, with
proximal view showing long, fused demicolpi (G), equato-
rial view (H), and distal view showing short demicolpi (I),
1,800. JCupania belizensis, syncolporate, rugulate pol-
len grain, 1,900. KMatayba apetala, parasyncolporate,
perforate-indistinctly rugulate pollen grain, 2,250. L
Pometia pinnata, brevicolporate, reticulate pollen grain,
1,500. (Photos R. van der Ham)
378 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
547 predominantly made of samaroid mericarps, and
548 less often of sub-globose, unwinged mericarps.
549 Seeds are ellipsoid to sub-globose or less
550 often lenticular or flattened, exalate or excep-
551 tionally winged (Diplokeleba and Magonia), gla-
552 brous or less often pubescent, the testa totally
553 dry (scleroidal), with fleshy portions, or
554 completely fleshy (sarcotestal), the scleroidal
555 testa totally naked or partially to completely cov-
556 ered by an arillode (of integumental origin),
557 which sometimes bears a funiculus-like basal
558 projection (e.g., Guioa,Mischocarpus,andSar-
559 copteryx). Scleroidal seeds sometimes bear a
560 small to large hilum. The embryo is usually
561 fleshy, oily or starchy, lacking endosperm, notor-
562 hizal or lomatorhizal with straight, curved or
563 plicate, fleshy cotyledons, the radicle often sepa-
564 rated by a deep fold in the testa that forms a
565 radicular pocket.
566 PHYTOCHEMISTRY. (based on Hegnauer 1973,
567 1990) A large amount of quebrachitol, a mono-
568 methyl ether of 1-inositol, occurs in leaves, barks,
569 flowers, and fruits of the whole family. Cyano-
570 genic glucosides are widely distributed in the
571 family (but not in Hippocastanoideae); they are
572 distributed throughout the plant body, including
573 ripe seeds. There they are often replaced by
574 cyanolipids, which in part also are very toxic.
575 Polyphenolics are found in Sapindaceae mainly
576 as coumarins, flavonoids, and proanthocyani-
577 dins. Condensed tannins, based on proanthocya-
578 nidins, can occur in large quantities, amounting
579 to 15–20% of the dry bark in Sapindoideae.
580 Gallotannins and their derivatives (bergenin) are
581 also present. The family is also rich in saponins,
582 which are contained in idioblasts within the
583 vegetative tissues and very often in the seeds.
584 These compounds include mono- and bidesmo-
585 sidic saponins with triterpene sapogenins, and
586 some have prominent ichthyotoxic and detergent
587 properties. The resinous exudates of Dodonoi-
588 deae are based on diterpenes and triterpenes.
589 Non-proteinogenic amino acids of the hypoglycin
590 type with branched carbon chains of 6 or 7
591 C-atoms occur free or as glutamyl peptides in
592 the seeds, and are known from the whole family.
593 Unspecific alkaloid tests have been positive for a
594 large number of species of the family, but specific
595 compounds rarely have been isolated and char-
596 acterized. An exception are the purine bases
597(caffeine, theobromine, theophylline), which
598are accumulated in large amounts in the seeds
599and other plant parts of Paullinia cupana and
600related species, from which in Amazonia a stimu-
601lating drink (“guarana
´”) is prepared since time
602immemorial.
603DISPERSAL. Wind-dispersed fruits consist of
604samaroid mericarps (Figs. 82, 83F) that are straw-
605colored at maturity. The genera Diplokeleba and
606Magonia have capsules with winged seeds that
607may also be dispersed by wind. Capsular fruits
608have green, red, orange, or yellow pericarps, with
609seeds that are either sarcotestal or arillate, and
610presumably dispersed by birds. Indehiscent bac-
611cate fruits have green, yellow, or red pericarps,
612and usually contain sarcotestal seeds very likely
613dispersed by mammals such as monkeys or bats.
614Several genera with inflated capsules seem to lack
615dispersal mechanisms; in these, fruit walls usually
616disintegrate, and the seeds are ultimately released
617by the effect of gravity.
618FOSSILS.Acer fruits first occur in the late Paleo-
619cene of North America, and in the Eocene and later
620Tertiary the genus is well represented there but
621also in Europe and Asia (Manchester 1999). The
622early occurrence of Acer in North America with
623the coeval appearance of Dipteronia (McClain and
624Manchester 2001) and the related extinct Eocene
625fruit, Deviacer, may point to a North American
626origin of the group. Leaves and fruits of Aesculus
627are on record from the Maastrichtian to early
628Eocene of North America (Manchester 2001), and
629also from the Paleogene of Spitsbergen and
630Kamtchatka (Budantsev 1983), whereas in Europe
631the genus does not appear before the Miocene (see
632also Harris et al. 2009). Fossil seeds attributable to
633Sapindoideae (from genus Sapindospermum Reid
634& Chandler) are known from the Cretaceous of
635North America, Greenland, Europe, and Siberia,
636and fossil wood (Sapindoxylum Kr€
ausel) has been
637recovered from Cretaceous beds in Egypt
638(Knobloch and Mai 1986). Sapindaceous flowers
639are reported from the middle Eocene of British
640Columbia (Erwin and Stockey 1990). Koelreuteria
641has an ample fossil record in the northern hemi-
642sphere that extends back to the middle Eocene
643(Manchester 1999).
644The fossil pollen record of the family begins in
645the Cretaceous, represented mainly by the pollen
Sapindaceae 379
Uncorrected Proof
646 genus Cupanieidites, which corresponds to the
647 extant genus Cupaniopsis and several related
648 genera (see Cookson and Pike 1954). Cupaniei-
649 dites appeared first in the Coniacian and Santo-
650 nian of Gabon, and in the Senonian of India and
651 Brazil, became more frequent during the Maas-
652 trichtian in North America, and appeared during
653 the Paleocene in Australia and was abundant
654 there in the Eocene (see Muller 1981).
655 FAMILY CLASSIFICATION. Sapindaceae were
656 first proposed by Jussieu in 1789 in his Genera
657 Plantarum as a family distinct from Aceraceae
658 (including Aesculus). This concept of Sapinda-
659 ceae has been followed by numerous workers
660 such as de Candolle (1824, who proposed Hippo-
661 castanaceae as a separate family), Cambesse
`des
662 (1829), and Don (1831). Later workers, such as
663 Reichenbach (1834), Lindley (1862), Bentham
664 and Hooker (1862), Baillon (1874), and Blume
665 (1878), included Aceraceae, Hippocastanaceae,
666 and various genera currently recognized in
667 other families such as Staphyleaceae, Sabiaceae,
668 and Melianthaceae within Sapindaceae.
669 The concept of Sapindaceae held by the end
670 of the 19th century and through most of the 20th
671 century as a family distinct from Aceraceae
672 and Hippocastanaceae is the result of the monu-
673 mental work of Radlkofer (1888, 1890, 1931). He
674 provided the first worldwide system of classifica-
675 tion for the family, recognizing two subfamilies
676 and 14 tribes, and although outdated, his work is
677 still a useful framework for the identification
678 of taxa and the understanding of phylogenetic
679 relationships within Sapindaceae. His system
680 considered the uniovulate Sapindaceae to be
681 the most basic members within the family. In
682 1964, Scholz provided modern names for the
683 two recognized subfamilies (i.e., Dodonaeoideae
684 and Sapindoideae), and inverted their sequence,
685 i.e., considering the multiovulate Dodoneaiodeae
686 as basal within the family. This rearrangement
687 was followed by Capuron (1969), who in addition,
688 modified some of the generic concepts. Muller
689 and Leenhouts (1976) independently proposed
690 a rearrangement to Radlkofer’s system that
691 agreed with Scholz’s general rearrangement but
692 proposed more substantial changes. They
693 reduced tribe Aphanieae into Lepisantheae, and
694 suggested the inclusion of families Aceraceae and
695 Hippocastanaceae into subfamily Dodonaeoideae,
696Aceraceae in its own tribe, and Hippocastanaceae
697as part of tribe Harpullieae.
698The inclusion of Aceraceae and Hippocasta-
699naceae by Muller and Leenhouts (1976) was
700slowly to be adopted. Thorne (1976), Cronquist
701(1981), and Takhtajan (1997) continued recogniz-
702ing Aceraceae and Hippocastanaceae as distinct
703families. At the same time, floristic works were
704not confronted with this dilemma, because Sapin-
705daceae s.s. are predominately tropical, while
706Aceraceae and Hippocastanaceae are largely tem-
707perate. More recently, researchers are following
708the suggestions of Muller and Leenhouts to
709include Aceraceae and Hippocastanaceae within
710Sapindaceae. Judd et al. (1994, 1999), who did
711a cladistic analysis based on morphological char-
712acters, supported the inclusion of both families
713within Sapindaceae. Savolainen et al. (2000), Soltis
714et al. (2000), and Johnson and Chase (in Klaassen
7151999), from results of studies based on DNA
716sequence data for a small number of taxa, found
717Hippocastanaceae and Aceraceae to be sister
718groups and to be nested within Sapindaceae.
719In a later study, Harrington et al. (2005),
720using sequencing data from two plastid genes
721(matK and rbcL) for 64 genera of Sapindaceae,
722Aceraceae, and Hippocastanaceae, found strong
723support for the recognition of four major clades.
724Their analyses show weak support for a Hippo-
725castaniodeae clade (Aceraceae and Hippocasta-
726naceae) being sister or basal to the remaining
727Sapindaceae, which could be interpreted as
728support for the recognition of Aceraceae and
729Hippocastanaceae as distinct families. They,
730however, opted for the recognition of four sub-
731families within Sapindaceae (i.e., Xanthoceroi-
732deae, Hippocastanoideae, Dodonaeoideae, and
733Sapindoideae), instead of four or five closely
734related families. The acceptance of four sub-
735families within Sapindaceae seems to be gaining
736support even in the absence of strong cladistic
737evidence. Thorne and Reveal (2007) modified
738their concept of Sapindaceae to recognize four
739subfamilies as suggested by Harrington et al. In
740a more recent study, Buerki et al. (2009), using
741sequence data from eight plastid and nuclear
742genes for 104 genera of Sapindaceae s.l., found
743ample support for the recognition of four clades
744within Sapindaceae, as proposed by Harrington
745et al., but like these authors, they could not place
746the Xanthoceroid clade with certainty among the
380 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
747 remaining clades, but left room for the interpreta-
748 tion of Aceraceae and Hippocastanaceae as a clade
749 closely related to Sapindaceae s.str. The current
750 treatment follows the approach of Harrington
751 et al. (2005) in recognizing four subfamilies within
752 Sapindaceae, instead of multiple families.
753 TRIBAL CLASSIFICATION. Radlkofer (1931) re-
754 cognized 14 tribes within Sapindaceae based
755 mostly on flower symmetry and fruit type.
756 His system remained largely unchanged until
757 Muller and Leenhouts (1976) cast doubts on the
758 monophyly of the Melicocceae, and the validity of
759 the Cupanieae as distinct from the Schleichereae
760 and Nephelieae. They ultimately suggested the
761 possibility of reducing the latter three tribes into
762 one. In a more recent cladistic analysis based on
763 morphological data, Acevedo-Rodrı
´guez (2002)
764 cast further doubts on the validity of the tribe
765 Melicocceae, suggesting the genera Castanospora,
766 Tristira, and Tristiropsis not to belong there.
767 Leenhouts (1978) evaluated the tribe Nephelieae,
768 but did not address the limits of the closely
769 related Cupanieae and Schleichereae. These
770 studies were followed by those of Johnson and
771 Chase (in Klaassen 1999) who analyzed many
772 more genera (40) than did previous authors.
773 Their studies, although including few genera,
774 suggest clades that conflict with the current
775 tribal classification, except for the Paullinieae-
776 Thouinieae clade. Their analysis also suggests
777 Paullinieae and Thouinieae to be sister groups
778 of a larger clade. These results contrast with
779 those of Acevedo-Rodrı
´guez (1993), Harrington
780 et al. (2005), and Buerki et al. (2009) where
781 the genera of Paullinieae are nested within the
782 Thouinieae, therefore calling for the merging of
783 the Paullinieae and Thouinieae tribes.
784 The work of Harrington et al. (2005) found
785 support for various lineages within their pro-
786 posed four subfamilies. Some of these lineages
787 are strongly supported, while others are only
788 weakly so. In summary, they found support for
789 the Acereae and Hippocastaneae clades within
790 subfamily Hippocastanoideae; a dehiscent fruit
791 clade and an indehiscent fruit clade within
792 subfamily Dodonaeoidea; clade Lepiantheae-
793 Sapindeae, clade Nephelieae, clade Cupanieae,
794 and clade Paullinieae-Thouinieae within sub-
795 family Sapindoideae. Buerki et al. (2009) found
796 strong support for the topologies obtained by
797Harrington et al. (2005) for the first three sub-
798families, but differ by proposing 10 groups within
799subfamily Sapindoideae. Buerki et al.’s results
800provide strong support for the Paullinia,Melicoc-
801cus,Tristiropsis,Macphersonia,Sapindus,Schlei-
802chera, and Delavaya groups, and only weak
803support for the Blomia,Cupania, and Koelreu-
804teria groups. However, the relationship among
805these clades is not well supported in their ana-
806lyses. Some of the proposed topologies seem
807inconclusive, since some of the morphologically
808well-defined genera are suggested to be either
809paraphyletic or polyphyletic, and many topolo-
810gies are only weakly supported. With exception of
811Paullinieae (expanded to include Thouinieae),
812which has support from various analyses, the
813tribal classification in Sapindaceae is still largely
814unresolved.
815In the present treatment, we follow Harrin-
816gton et al. (2005) in recognizing four subfamilies
817and four tribes within subfamilies Hippocasta-
818noideae and Dodonaeoideae. With regards to
819subfamily Sapindoideae, we will not make an
820attempt to divide it into tribes, as most relation-
821ships suggested by previous workers are either
822not supported by molecular data or are inconclu-
823sive (Harrington et al. 2005; Buerki et al. 2009).
824Instead, we will recognize only the tribes Paulli-
825nieae (including Thouinieae) and Melicocceae
826(excluding Tristira,Tristiropsis,Castanospora,
827and Dilodendron) for which there seems to be
828strong support from either morphology or DNA
829sequencing data; the remaining genera will be
830treated alphabetically as incertae sedis within
831subfamily Sapindoideae. We also follow Harrin-
832gton et al. (2005) in including Koeleuteria and
833Ungnadia within subfamily Sapindoideae.
834AFFINITIES. An early, very extensive study on
835the relationship of Sapindaceae with other
836families, involving molecular data, was that of
837Gadek et al. (1996). They analyzed rbcL sequence
838data for representatives of all putative sapinda-
839lean families. Their results identified a sapinda-
840lean clade sister to representatives of Malvales.
841This premise supports a broader concept of
842Sapindales similar to that of Cronquist (1981)
843but with the exclusion of some families. More
844recent work has fully confirmed the work of
845Gadek et al. (1996); see INTRODUCTION TO SAPINDALES
846in this volume.
Sapindaceae 381
Uncorrected Proof
847 ECONOMIC IMPORTANCE. Sapindaceae are the
848 source of numerous products, some of which
849 are economically important, either globally or
850 locally. Among the most important ones, fruit
851 crops are high on the list. These include edible
852 fruits such as litchi (Litchi sinensis), longan
853 (Dimocarpus longan), rambutan (Nephelium
854 lappaceum), and pulasan (N. ramboutan-ake)at
855 a global level, and the mamoncillo or keneep
856 (Melicoccus bijugatus), pitomba (Talisia escu-
857 lenta), and cutupliı
´o guaya (T. olivaeformis)at
858 a local level. The arillodes of Blighia sapida are
859 the source of the nutritious ackee, widely con-
860 sumed in Jamaica, but highly toxic when eaten
861 unripe (Rashford 2001). Numerous species of
862 Paullinia have been reported to be useful in the
863 preparation of medicines, caffeine-rich beverages,
864 binding and weaving material, and for fish,
865 human and arrow poisoning (Beck 1990). The
866 seeds of Paullinia cupana are the source of the
867 important Brazilian crop guarana
´, a source of
868 caffeine and flavoring of soft drinks. Almost all
869 Sapindaceae are used around the tropics for fish
870 poisoning (Acevedo-Rodrı
´guez 1990). The wood
871 of some species of Euphorianthus, Harpullia, and
872 Schleichera are used in the construction of houses.
873 Numerous genera are grown as ornamentals—
874 e.g., Acer,Aesculus,Arfeuillea,Allophylus,Cardi-
875 ospermum,Filicium,Harpullia,Koelreuteria,
876 Sapindus, and Xanthoceras. Minor products
877 include oils from Pappea and Schleichera, and
878 arrow poison from Paullinia pinnata.
879 KEY TO THE GENERA
880 1. Carpels multiovulate 2
881 – Carpels uniovulate 41
882 2. Leaves simple or unifoliolate 3
883 – Leaves compound or palmately lobed 8
884 3. Leaves opposite; carpels 2; fruits of 2 indehiscent,
885 distally winged mericarps 2.Acer (in part)
886 – Leaves alternate; carpels 3 or more numerous,
887 sometimes 2 but then fruits capsular 4
888 4. Petals absent 5
889 – Petals present 7
890 5. Floral disk unilateral 17.Llagunoa (in part)
891 – Floral disk obsolete or absent 6
892 6. Plant often viscid; sepals 3–7; stamens 5–15, in one
893 row 13.Dodonaea (in part)
894– Plant not viscid; sepals 5–8; stamens 8–74, usually
895in two or more rows 12.Distichostemon
8967. Plant with glandular and simple trichomes; inflores-
897cence a thyrse, with numerous flowers; sepals of
898similar size; fruit a schizocarp of 3 indehiscent
899obovoid cocci or a crustose capsule 11.Diplopeltis
900– Plants without glandular trichomes; inflorescence
901racemose, 1–3-flowered; outer two sepals smaller;
902fruit an inflated capsule 124.Stocksia
9038. Leaves opposite or subopposite 9
904– Leaves alternate 16
9059. Leaves digitate or trifoliolate 10
906– Leaves pinnate 14
90710. Leaves trifoliolate 11
908– Leaves digitate 12
90911. Corolla zygomorphic, usually of only 4 petals;
910petals clawed, crested above the claw; carpels (2)3(4);
911stigma unlobed; fruit a loculicidal, unwinged cap-
912sule 5.Billia
913– Corolla actinomorphic, of 5 petals; petals not
914clawed nor crested; carpels 2; stigmas two; fruit of
915two indehiscent, distally or dorsally winged
916mericarps 2.Acer (in part)
91712. Fruit of two indehiscent, distally or dorsally
918winged mericarps 2.Acer (in part)
919– Fruit a loculicidal, unwinged capsule 13
92013. Sepals connate half or more of their length
9214.Aesculus
922– Sepals distinct to the base 6.Handeliodendron
92314. Ovary 3(5)-carpellate; fruit capsular
92413.Dodonaea (in part)
925– Ovary 2(3)-carpellate; fruit schizocarpic, of 2 win-
926ged mericarps 15
92715. Mericarps completely surrounded by a wing
9283.Dipteronia
929– Mericarps with a distal wing 2.Acer (in part)
93016. Leaves trifoliolate 17
931– Leaves pinnate 20
93217. Plant bearing stellate
933hairs 9.Cossinia (in part)
934– Plant glabrous or with simple hairs 18
93518. Petals 0; floral disk unilateral
93617.Llagunoa (in part)
937– Petals 5; floral disk annular 19
93819. Fruit indehiscent, 1-locular, baccate; petals lacking
939appendages 27.Hypelate
940– Fruit a dehiscent, 2–3-coccate, loculicidal capsule;
941petals with marginal appendages 64.Delavaya
94220. Plant bearing stellate hairs 21
943– Plant glabrous or with simple hairs, lacking stellate
944hairs 25
382 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
945 21. Floral disk with 5 elongated, horn-like lobes; leaflet
946 margins serrate; fruit thick woody, with corky
947 endocarp 1.Xanthoceras
948 – Floral disk unilateral, semi-annular or annular,
949 without horn-like lobes; leaflets entire or crenate;
950 fruit crustose or papery to woody but not corky
951 22
952 22. Seeds with white or red sarcotesta on lower half or
953 base 16.Harpullia (in part)
954 – Seeds without sarcotesta (seed coat black or dark
955 brown) 23
956 23. Abaxial surface of leaflets densely stellate pubes-
957 cent; seeds glabrous 9.Cossinia (in part)
958 – Abaxial surface of leaflets sparsely stellate pubes-
959 cent; seeds hispidulose 24
960 24. Disk semi-annular, or nearly complete, formed by
961 two semi-annular halves; fruits papery
962 7.Arfeuillea
963 – Disk annular or semi-annular, dish-shaped to pen-
964 tagonous; capsules crustose 20.Majidea
965 25. Corolla 0 or rudimentary 26
966 – Corolla present 29
967 26. Fruit a dehiscent, loculicidal capsule
968 8.Averrhoidium
969 – Fruit indehiscent, baccate 27
970 27. Fruit 2-locular 25.Ganophyllum
971 – Fruit 1-locular 28
972 28. Leaf rachis winged; inflorescences of axillary,
973 glomerate cymes; stamens not coiled in bud
974 21.Doratoxylon
975 – Leaf rachis naked; inflorescences of thyrses;
976 stamens coiled in bud 28.Zanha
977 29. Petals lacking appendages 30
978 – Petals bearing appendages 36
979 30. Pollen dispersed as tetrads; ovules 7–8 per locule;
980 seeds winged 19.Magonia
981 – Pollen dispersed as monads; ovules 2(3) per locule;
982 seeds unwinged 31
983 31. Leaf rachis winged 26.Hippobromus
984 – Leaf rachis unwinged 32
985 32. Petals suborbiculate, sessile 22.Euchorium
986 – Petals ovate or clawed 33
987 33. Carpels 2; fruit baccate, indehiscent 23.Exothea
988 – Carpels 3(4); fruit capsular, loculicidal 34
989 34. Floral disk unilateral, double, the inner lobe concave,
990 4-dentate; sepals fimbriate-glandular; ovary with
991 stipitate glands 18.Loxodiscus
992 – Floral disk annular, lobed; sepals not fimbriate nor
993 glandular; ovary lacking stipitate glands 35
994 35. Floral disk simple; capsule fleshy, 1(3)-coccate
995 15.Eurycorymbus
996– Floral disk double, the central rim tubular
997sometimes elongated into a gynophore; capsule
998membranous, with more or less compressed
999locules 59.Conchopetalum
100036. Fruits indehiscent 72.Erythrophysopsis
1001– Fruits dehiscent 37
100237. Fruits coriaceous or woody 38
1003– Fruits membranaceous, or chartaceous 39
100438. Seeds pubescent 71.Erythrophysa (in part)
1005– Seeds glabrous 138.Ungnadia
100639. Sepals imbricate; disk semi-annular; seeds
1007pubescent 71.Erythrophysa (in part)
1008– Sepals valvate; disk annular; seeds glabrous 40
100940. Petals yellow; appendages formed by the fim-
1010briate margins of the sagitate blade base; floral
1011disk elevated on a short androgynophore; stamens
1012straight in bud; distal leaflet fully developed; cap-
1013sules with incomplete septa 82.Koelreuteria
1014– Petals white; appendages formed by a narrow mar-
1015ginal flap; floral disk sessile; stamens geniculate
1016in bud; distal leaflet rudimentary; capsule with
1017complete septa 120.Sinoradlkofera
101841. Leaves unifoliolate or simple 42
1019– Leaves compound 50
102042. Calyx and corolla actinomorphic 43
1021– Calyx and corolla zygomorphic 46
102243. Fruit indehiscent 44
1023– Fruit dehiscent 45
102444. Corolla of 4 or 5 petals, anthers dorsifixed;
1025sarcotesta 0 115.Sapindus (in part)
1026– Corolla 0; anthers basifixed, sarcotesta present
102773.Glenniea (in part)
102845. Fruit a 1-locular (by abortion), loculicidal capsule;
1029seed with lobed arillode 100.Pappea
1030– Fruit 1–3-coccate, septicidal or loculicidal-
1031calyptrate; seed with granular sarcotesta
103244.Alectryon (in part)
103346. Fruit indehiscent 47
1034– Fruits schizocarpic 48
103547. Calyx 4-merous; fruit of 1–2 rounded cocci
103629.Allophylus (in part)
1037– Calyx 5-merous; fruit of 3, ellipsoid, apically keeled
1038cocci 95.Namataea
103948. Mericarps sub-globose, exalate 34.Guindilia
1040– Mericarps winged 49
104149. Calyx 5-merous; mericarps with a dorsal wing
104231.Bridgesia
1043– Calyx 4-merous; mericarps with a distal wing
104440.Thouinia (in part)
104550. Distal leaflet well-developed 51
1046– Distal leaflet rudimentary 61
Sapindaceae 383
Uncorrected Proof
1047 51. Trees or shrubs, exceptionally climbing shrubs;
1048 stipules absent 52
1049 – Vines, lianas or climbing shrubs, sometimes not
1050 climbing in early stages; stipules minute, or excep-
1051 tionally large and early deciduous 55
1052 52. Leaves pinnate 53
1053 – Leaves trifoliolate 54
1054 53. Fruit schizocarpic; mericarps winged; leaves once
1055 pinnate 30.Athyana
1056 – Fruit a loculicidal capsule; leaves bipinnate
1057 66.Dilodendron (in part)
1058 54. Fruit schizocarpic, of 3 winged mericarps
1059 40.Thouinia (in part)
1060 – Fruits indehiscent, of 1(2) unwinged monocarps
1061 29.Allophylus (in part)
1062 55. Fruits schizocarpic, splitting into 3 1-winged
1063 mericarps 56
1064 – Fruit capsular, winged or not winged 59
1065 56. Flowers actinomorphic 39.Thinouia
1066 – Flowers zygomorphic 57
1067 57. Mericarps with a dorsal wing surrounding the
1068 coccus 35.Houssayanthus
1069 – Mericarps with a proximal wing 58
1070 58. Stamens with filaments of similar length; pollen
1071 cylindrical 36.Lophostigma
1072 – Stamens with filaments of unequal lengths; pollen
1073 triangular 38.Serjania
1074 59. Capsules woody, coriaceous or crustaceous, not
1075 inflated; pollen 3-porate 37.Paullinia
1076 – Capsules papery, inflated or nearly so; pollen
1077 3-(demi)(syn)colporate 60
1078 60. Capsules not winged or only narrowly so
1079 32.Cardiospermum
1080 – Capsule dorsally winged 41.Urvillea
1081 61. Leaves bipinnate or tripinnate 62
1082 – Leaves once pinnate 66
1083 62. Leaves tripinnate 133.Tripterodendron
1084 – Leaves bipinnate 63
1085 63. Sepals valvate 50.Bizonula
1086 – Sepals imbricate 64
1087 64. Anthers basifixed; placentation axile; fruit a
1088 loculicidal capsule 66.Dilodendron (in part)
1089 – Anthers dorsifixed; placentation basal; fruit inde-
1090 hiscent, baccate or nearly so 65
1091 65. Stigma capitate, sessile; aril and sarcotesta present
1092 90.Macphersonia (in part)
1093 – Stigma of 3 elongated branches; aril or sarcotesta
1094 absent 135.Tristiropsis
1095 66. Calyx zygomorphic 67
1096 – Calyx actinomorphic (slightly zygomorphic in
1097 Tinopsis)96
109867. Sepals connate to form a tubular or urceolate calyx
109968
1100– Sepals distinct 81
110168. Calyx bilabiate 111.Pseudopancovia
1102– Calyx of (4)5(–7) lobes, not bilabiate 69
110369. Calyx urceolate 70
1104– Calyx tubular 71
110570. Fruit 5–8-locular, not ribbed; inflorescence of
1106densely packed flowers; bracts longer than the
1107flowers 113.Radlkofera
1108– Fruit 3–8-locular, longitudinally ribbed; inflores-
1109cence with loosely spaced flowers; bracts shorter
1110than the flowers 57.Chytranthus
111171. Petals 4; disk unilateral; stamens (6)8; style
1112terminal 99.Pancovia
1113– Petals 5; disk annular; stamens (8)12–30; style
1114gynobasic or sub-terminal 63.Deinbollia
111572. Fruit indehiscent 73
1116– Fruit dehiscent 76
111773. Fruit deeply lobed or lobed-coccate
111887.Lepisanthes (in part)
1119– Fruit ovoid, obovoid or ellipsoid, not lobed 74
112074. Fruit locules unwinged 87.Lepisanthes (in part)
1121– Fruit with 3 dorsal wings 75
112275. Corolla 0; disk annular; fruit trilocular
1123134.Tristira
1124– Corolla of 4–5 petals; disk unilateral, flattened;
1125fruit unilocular 140.Zollingeria
112676. Fruit schizocarpic, splitting into indehiscent mer-
1127icarps 77
1128– Fruit capsular 83
112977. Mericarps unwinged 78
1130– Mericarps winged 80
113178. Seeds covered with sticky, saponiferous pulp
1132115.Sapindus (in part)
1133– Seeds not surrounded by a sticky or saponiferous
1134pulp 79
113579. Mericarp membranaceous, bladdery 109.Porocystis
1136– Mericarps woody, not bladdery
1137119.Scyphonychium
113880. Mericarps with a proximal wing 131.Toulicia
1139– Mericarps with a distal or dorsal wing 81
114081. Calyx 4-merous; corolla zygomorphic
114133 Diatenopteryx
1142– Calyx 5-merous; corolla actinomorphic 82
114382. Leaves with 10–12 leaflets; disk annular or semi-
1144annular; stamens 8; ovary 3-carpellate; fruits of 3
1145mericarps 48.Atalaya
1146– Leaves with 2–4 leaflets; disk 5-lobed; stamens
114718–24; ovary bicarpellate; fruits of 2 mericarps
114880.Hornea
384 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
1149 83. Corolla absent 65.Dictyoneura (in part)
1150 – Corolla present 84
1151 84. Corolla zygomorphic; capsule echinate or muricate
1152 85
1153 – Corolla actinomorphic; capsules smooth 86
1154 85. Capsules echinate; petal appendages marginal
1155 121.Sisyrolepis
1156 – Capsules muricate; petal appendage basal, simple
1157 104.Phyllotrichum
1158 86. Capsules 1-locular 96.Neotina
1159 – Capsules 2- or 3-locular 87
1160 87. Seeds winged, non-arillate 10.Diplokeleba
1161 – Seeds, ellipsoid, sub-globose, unwinged, arillate 88
1162 88. Gynoecium 2-carpellate 89
1163 – Gynoecium (2)3(4)-carpellate 90
1164 89. Leaflets entire; anthers retuse at apex
1165 138.Vouarana (in part)
1166 – Leaflets serrate or crenate-serrate; anthers apicu-
1167 late at apex 128.Tina (in part)
1168 90. Arillode basal or lateral 91
1169 – Arillode partially to nearly completely covering the
1170 seed 92
1171 91. Arillode basal, surrounding the hilum; disk
1172 annular 114.Rhysotoechia
1173 – Arillode at base of seed but not covering the hilum;
1174 disk pentagonal 110.Pseudima
1175 92. Petal appendages absent 93
1176 – Petal appendages present or sometimes
1177 rudimentary 94
1178 93. Petals more or less cuneate at base; inflorescence
1179 axillary; arillode fimbriate at apex
1180 94.Molinaea (in part)
1181 – Petals clawed at base; inflorescence cauliflorous;
1182 arillode open but not fimbriate at apex
1183 74.Gloeocarpus
1184 94. Arillode with a basal funiculus-like appendage
1185 (rudimentary in one species) 77.Guioa
1186 – Arillode lacking a basal funiculus-like appendage
1187 95
1188 95. Petals with one or two basal appendages
1189 62.Cupaniopsis
1190 – Petals with marginal appendages or without
1191 appendages 94.Molinaea (in part)
1192 96. Petals absent 97
1193 – Petals present 118
1194 97. Fruits indehiscent or tardily dehiscent
1195 (pseudodehiscent) 98
1196 – Fruits dehiscent 107
1197 98. Sepals connate into a cupular or urceolate calyx
1198 99
1199 – Sepals distinct or connate only at base 101
120099. Fruit tardily dehiscent (pseudodehiscent), usually
12011-coccate, the pericarp smooth
1202123.Stadmania (in part)
1203– Fruit indehiscent, 3-locular, or if 1-coccate, then
1204with muricate pericarp 100
1205100. Gynoecium 3-carpellate; fruit usually 3-locular, 3-
1206sulcate or 3-lobed 105.Placodiscus
1207– Gynoecium bicarpellate; fruit usually 1-coccate (1
1208coccus rudimentary), indehiscent, baccate; muricate
120988.Litchi
1210101. Seeds exarillate 73.Glenniea (in part)
1211– Seeds arillate 102
1212102. Gynoecium 2-carpellate; arillode basal 103
1213– Gynoecium 3(4)-carpellate; arillode nearly cover-
1214ing the entire seed 104
1215103. Pseudostipules present; floral disk glabrous
121698.Otonephelium
1217– Pseudostipules 0; floral disk pubescent
121867.Dimocarpus (in part)
1219104. Arillode with a dorsal split 79.Haplocoelum
1220– Arillode covering the seed completely (no dorsal
1221split) 105
1222105. Sepals connate at base (1/4 of their length); floral
1223disk unlobed; anthers dorsifixed
1224118.Schleichera
1225– Sepals distinct to base; floral disk lobed; anthers
1226basifixed 106
1227106. Sepals 5; stamens 8; cotyledons smooth
122884.Lecaniodiscus
1229– Sepals (5)6 or 7(8); stamens 6–8(10); cotyledons
1230with brain-like appearance (cerebriform)
123149.Beguea
1232107. Calyx with imbricate aestivation 108
1233– Calyx with valvate aestivation 111
1234108. Pseudostipules present; anthers basifixed; fruits
12351-locular 52.Blighiopsis
1236– Pseudostipules 0; anthers dorsifixed; fruits
1237(1)3(4)-locular 109
1238109. Leaflets serrate 65.Dictyoneura (in part)
1239– Leaflets entire 110
1240110. Aril with a basal funiculus-like appendage; inflor-
1241escences of thyrses or panicles
124293.Mischocarpus (in part)
1243– Aril lacking a basal funiculus-like appendage;
1244inflorescences of racemes 51.Blighia
1245111. Fruit with circumscissile dehiscence 112
1246– Fruit with loculicidal dehiscence 113
1247112. Seed with a dorsal, white arillode
1248107.Podonephelium
1249– Seeds with red sarcotesta on lower half
125044.Alectryon (in part)
Sapindaceae 385
Uncorrected Proof
1251 113. Seeds naked (without arillode or sarcotesta);
1252 endotesta ruminately grown together with embryo
1253 76.Gongrospermum
1254 – Seeds arillate or sarcotestal 114
1255 114. Seeds arillate 136.Tsingya
1256 – Seeds sarcotestal 115
1257 115. Sepals distinct to the base 116
1258 – Sepals connate at least half way to form cupular or
1259 acetabuliform calyx 117
1260 116. Gynoecium unicarpellate; stigma capitate
1261 53.Blomia (in part)
1262 – Gynoecium (1)2(–4)-carpellate; stigmata 2(3),
1263 elongated, and usually coiled
1264 97.Nephelium (in part)
1265 117. Fruit warty, spiny or echinate
1266 97.Nephelium (in part)
1267 – Fruit smooth 44.Alectryon (in part)
1268 118. Fruits indehiscent 119
1269 – Fruits dehiscent 133
1270 119. Seeds laterally flattened, with a longitudinal ventral
1271 hilum 106.Plagioscyphus
1272 – Seeds variously shaped, hilum if present restricted
1273 to the basal-ventral part 120
1274 120. Gynoecium 2(3)-carpellate 121
1275 – Gynoecium 3-carpellate 128
1276 121. Stamens 5(–7) 122
1277 – Stamens (6–)8(–10) 125
1278 122. Seeds exarillate 24.Filicium
1279 – Seeds completely covered
1280 by an arillode 123
1281 123. Sepals connate half of their length 108.Pometia
1282 – Sepals distinct 124
1283 124. Cataphylls present; petals lacking appendages
1284 112.Pseudopteris
1285 – Cataphylls absent; petals bearing basal or marginal
1286 appendages 129.Tinopsis
1287 125. Testa fleshy (sarcotestal) 126
1288 – Testa not fleshy 127
1289 126. Fruit ellipsoid or globose, smooth; carpels with
1290 incomplete septa 42.Melicoccus
1291 – Fruit 1–2-coccate, warty or spiny; carpels with
1292 complete septa 139.Xerospermum
1293 127. Seed exarillate; calyx aestivation valvate; fruit
1294 smooth 55.Castanospora
1295 – Seed arillate; calyx aestivation imbricate; fruit
1296 smooth to echinate 67.Dimocarpus (in part)
1297 128. Sepals connate into a cup-shaped calyx
1298 123.Stadmania (in part)
1299 – Sepals distinct, or if connate, then the calyx
1300 tubular 129
1301 129. Filaments coiled in bud 130
1302– Filaments short, not coiled in bud 131
1303130. Petal appendages marginal
130490.Macphersonia (in part)
1305– Petal appendages basal 56.Chouxia
1306131. Seeds naked (not arillate nor sarcotestal)
1307115.Sapindus (in part)
1308– Seeds arillate or sarcotestal 132
1309132. Seeds sarcotestal 43.Talisia
1310– Seeds completely covered by a translucent arillode
131154.Camptolepis
1312133. Fruit schizocarpic, splitting into winged mericarps
1313127.Thouinidium (in part)
1314– Fruit capsular 134
1315134. Fruit echinate, warty or with hispid or setaceous
1316hairs 135
1317– Fruit smooth or scrobiculate, glabrous or variously
1318pubescent 138
1319135. Fruit a 1- or 2-coccate, warty or echinate capsule;
1320petals lacking appendages 60.Cubilia
1321– Fruit a 3-locular, trigonous or trilobed, hispid cap-
1322sule; petals with appendages 136
1323136. Disk cupular; sepals valvate; seeds arillate at base;
1324fruits without irritating hairs 70.Eriocoelum
1325– Disk annular; sepals imbricate; seeds non-
1326arillate, with hilar sarcotesta; fruit with irritating
1327hairs 137
1328137. Fruit locules not winged 81.Jagera
1329– Fruit locules winged on lower dorsal portion
133058.Cnesmocarpon
1331138. Capsules 1-locular 139
1332– Capsules 2- or 3-locular 143
1333139. Seed naked (not arillate nor sarcotestal), with large
1334white hilum 101.Paranephelium
1335– Seed arillate or sarcotestal, with small hilum 140
1336140. Seed arillate, the arillode with dorsal split
133775.Gongrodiscus
1338– Seed sarcotestal 141
1339141. Fruit smooth 142
1340– Fruits tuberculate to echinate
134197.Nephelium (in part)
1342142. Seed black with partial red sarcotesta
134344.Alectryon (in part)
1344– Seed completely covered by a brownish sarcotesta
134553.Blomia (in part)
1346143. Petals lacking appendages 144
1347– Petals bearing appendages 148
1348144. Sepals connate; seeds exarillate
1349122.Smelophyllum
1350– Sepals distinct; seeds arillate 145
1351145. Capsule obovoid or trigonous 146
1352– Capsule deeply 2- or 3-lobed 146
386 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
1353 146. Seeds sarcotestal at its base or lower half, not
1354 fimbriate at apex 16.Harpullia (in part)
1355 – Seeds arillate, arillode covering at least lower half,
1356 fimbriate at apex 94.Molinaea (in part)
1357 147. Arillode ventrally split and fimbriate
1358 125.Storthocalyx
1359 – Arillode ventrally attached, covering whole seed,
1360 not fimbriate 85.Lepiderema
1361 148. Calyx with valvate aestivation 149
1362 – Calyx with imbricate aestivation 155
1363 149. Sepals distinct 150
1364 – Sepals connate at least at base 153
1365 150. Petals with a single ventral appendage 151
1366 – Petals with marginal appendages 152
1367 151. Gynoecium 3-carpellate; capsule (1–2)3-coccate,
1368 wider than long 45.Amesiodendron
1369 – Gynoecium 2-carpellate; capsule 2-lobed or ellip-
1370 tic, longer than wide 86.Lepidopetalum
1371 152. Seeds sarcotestal; gynoecium bicarpellate
1372 46.Aporrhiza
1373 – Seed arillate; gynoecium tricarpellate
1374 91.Matayba
1375 153. Sepals connate at base 154
1376 – Sepals connate at least half of their length
1377 44.Alectryon (in part)
1378 154. Petals with marginal appendages; fruit locules not
1379 winged dorsally 92.Mischarytera
1380 – Petals with ventral, bifid appendage; fruit locules
1381 with a narrow dorsal wing 116.Sarcopteryx
1382 155. Petals with marginal appendages 156
1383 – Petals with ventral appendages 166
1384 156. Seeds sarcotestal 157
1385 – Seeds arillate 159
1386 157. Pubescence of stellate hairs
1387 16.Harpullia (in part)
1388 – Pubescence of simple hairs 158
1389 158. Pseudostipules 0; sarcotesta cupular
1390 117.Sarcotoechia
1391 – Pseudostipules present; sarcotesta completely cov-
1392 ering the seed 83.Laccodiscus
1393 159. Arillode basal 160
1394 – Arillode covering at least lower half of the seed 163
1395 160. Petals longer than the sepals, with crested
1396 appendages 130.Toechima
1397 – Petals shorter than the sepals, with non-crested
1398 appendages 161
1399 161. Petals clawed at base 69.Elattostachys
1400 – Petals not clawed at base 162
1401 162. Pericarp woody; seeds arillate at base, lacking a
1402 funiculus-like appendage
1403 138.Vouarana (in part)
1404– Pericarp coriaceous; seeds completely covered by
1405an arillode, with a funiculus-like appendage
140693.Mischocarpus (in part)
1407163. Capsule dehiscent by a loculicidal calyptra or
1408septicidal 47.Arytera
1409– Capsules loculicidal 164
1410164. Seeds lenticular 68.Diploglottis
1411– Seeds obovoid or ellipsoid 165
1412165. Gynoecium bicarpellate; stigma an invaginate
1413prolongation of the style 128.Tina (in part)
1414– Gynoecium tricarpellate; stigma simple, with three
1415stigmatic lines 61.Cupania
1416166. Petal bearing a single appendage 167
1417– Petals bearing 2 appendages 171
1418167. Sepals connate at base 168
1419– Sepals distinct to base 169
1420168. Seed exarillate 102.Pavieasia
1421– Seed sarcotestal 89.Lychnodiscus
1422169. Petals clawed at base 132.Trigonachras (in part)
1423– Petals cuneate at base 170
1424170. Gynoecium 3-carpellate; disk 7- or 8-lobed; ovary
1425hirsute 103.Pentascyphus
1426– Gynoecium 2-carpellate; disk annular; ovary gla-
1427brous 78.Haplocoelopsis
1428171. Petals not clawed; appendage crested
1429126.Synima
1430– Petals clawed, appendage not crested 172
1431172. Seed with a 2- or 3-lobed sarcotesta around the
1432hilum 14.Euphorianthus
1433– Seed not sarcotestal (naked)
1434132.Trigonachras (in part)
1435
1436
SUBFAMILIES,TRIBES,AND GENERA
1437OF SAPINDACEAE
1438I. SUBFAM. XANTHOCEROIDEAE Thorne & Reveal
1439(2007).
1440Leaves alternate; petals without appendages;
1441ovules 7–8 per locule; disk with orange horn-
1442like appendages.
14431.Xanthoceras Bunge
1444Xanthoceras Bunge, Mem. Sav. Etr. Acad. Petersb. 2: 85
1445(1834).
1446Falsely polygamous trees. Leaves alternate,
1447imparipinnate, with stellate pubescence; leaflets
1448serrate; distal leaflet rudimentary; stipules 0.
Sapindaceae 387
Uncorrected Proof
1449 Inflorescences terminal racemose thyrses. Flow-
1450 ers actinomorphic, bisexual or functionally uni-
1451 sexual; sepals 5, distinct; petals 5, with darker
1452 coloration at base, clawed; disk with 5 erect corni-
1453 form lobes, alternating with the petals; stamens 8;
1454 pollen colporate, loosely verrucate; ovary 3-
1455 carpellate, with 7–8 ovules per carpel; style fili-
1456 form with capitate, 3-sulcate stigma. Fruit a 3-
1457 locular, woody loculicidal capsule, with corky
1458 endocarp. Seeds exarillate. 2n¼30.
1459 A single species, X. sorbifolia Bunge, endemic
1460 to China, and commonly cultivated as an orna-
1461 mental tree.
1462 II. SUBFAM. HIPPOCASTANOIDEAE Burnett (1835).
1463 Leaves opposite; ovules 2 per locule.
1464 1.Tribe Acereae (Durande) Dumort. (1827).
1465 Flowers actinomorphic; petals without appen-
1466 dages; disk annular.
1467 2.Acer L. Fig. 80A–I
Acer L., Sp. Pl. 1054 (1753); de Jong, Meded. Landbou.
1468 Wageningen Nederl. 72: 1–201 (1976).
1469Duodichogamous or dioecious trees or shrubs.
1470Leaves opposite, simple, unlobed, palmately
1471lobed to deeply dissected, or trifoliolate, palmati-
1472compound or imparipinnate, margins entire, den-
1473tate, serrate or lobed; petioles usually long; stipules
14740 or rarely present. Inflorescences terminal or axil-
1475lary, corymbose-thyrsoid, racemose or fasciculate.
1476Flowers unisexual or bisexual, actinomorphic;
1477sepals (4)5, distinct or less often connate, imbri-
1478cate; petals 4–5 or 0, white or greenish, not clawed;
1479disk extrastaminal or less often intrastaminal,
1480amphistaminal or 0, annular or annular-lobed,
1481glabrous or pubescent; stamens (5)8 (10, 12), the
1482filaments equal or unequal; pollen colpate or col-
1483por(oid)ate, striate, rugulate or reticulate, some-
1484times scabrate; ovary 2(3, 5, 8)-carpellate, with
14852 ovules per carpel; style branches stigmatic, 2;
1486style sometimes very short. Fruits schizocarpic,
1487splitting into 2 samaroid, 1-seeded, mericarps,
1488each with a long, dorsal or distal wing. Seed later-
1489ally compressed, with papery testa.
1490Sixteen sections and about 126 species,
1491northern Asia, Japan, Europe, and North
1492America.
14933.Dipteronia Oliver Fig. 80J
1494Dipteronia Oliver, Hooker’s Icon. Pl. t. 1898. (Oct 1889).
Fig. 80. Sapindaceae. A–CAcer platanoides.ALeaf. B
Male flower. CFemale flower. DAcer laurinum, part of
infructescence. EAcer carpinifolium, part of infructes-
cence. F–JAcer negundo.FLeaf. GMale flower. HFemale
flower. IFruit. JDipteronia sinensis, fruit. (Takhtajan
1981; drawn by A. Schischtkin)
388 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
1495 Duodichogamous trees. Leaves opposite, impar-
1496 ipinnate; leaflets 9–17, with serrate margins;
1497 terminal leaflet fully developed; stipules 0. Inflor-
1498 escences terminal, thyrsoidate. Flowers function-
1499 ally unisexual, actinomorphic; sepals 5, distinct;
1500 petals 5, cream or greenish, clawed; disk extra-
1501 staminal, annular-lobed, stamens 6–8, the fila-
1502 ments of equal length; pollen colporate, striate;
1503 ovary 2-carpellate, with 2 ovules per carpel; stig-
1504 mata 2, connate at base. Fruits seemingly schizo-
1505 carpic, of 2 suborbicular, samaroid mericarps,
1506 each with a wing completely surrounding the
1507 seed locule. Seeds exarillate.
1508 A single species, D. sinensis Oliver, endemic
1509 to China.
1510 2.Tribe Hippocastaneae (DC.) Dumort. (1827).
1511 Flowers zygomorphic; petals usually with appen-
1512 dages; disk unilateral.
1513 4.Aesculus L.
1514 Aesculus L., Sp. Pl.: 344. (1753).
1515 Hippocastanum Miller (1754).
1516 Trees or shrubs. Leaves opposite, palmati-
1517 compound; petioles usually long; stipules 0.
1518 Inflorescences terminal thyrses or racemes. Flow-
1519 ers bisexual or functionally staminate, zygomor-
1520 phic; sepals 5, imbricate, connate to form a
1521 tubular or campanulate calyx; petals 4(5), equal
1522 or unequal, clawed; appendages 0 or minute
1523 and placed above the claw; disk unilateral, 4-
1524 lobed; stamens (5)6–8, the filaments of unequal
1525 length; pollen colporate, finely striate, sometimes
1526 scabrate, usually with heavily verrucate colpus
1527 membranes; ovary (2)3(4)-carpellate, with
1528 2 ovules per carpel; style with punctiform or
1529 obscurely 3-lobed stigma. Fruit usually a 1-
1530 seeded loculicidal capsule, smooth or echinate.
1531 Seeds exarillate, with a large, pale hilum.
1532 Thirteen species, southeastern Europe, Asia,
1533 Japan, and North America.
1534 5.Billia Peyritsch
1535 Billia Peyritsch, Bot. Zeit. 16: 153 (1858).
1536 Trees. Leaves opposite, trifoliolate; margins
1537 entire; petioles long; stipules 0. Inflorescences of
1538 terminal panicles bearing bisexual and unisexual
1539 flowers. Flowers zygomorphic; sepals 5, imbri-
1540 cate, distinct, unequal; petals 4(5), red or white,
1541unequal, clawed; appendages minute, two or a
1542single bifid or dissected, ventral on the claw;
1543disk unilateral, 4-lobed; stamens 6–8, the fila-
1544ments of unequal length; pollen colporate, finely
1545striate, with heavily verrucate colpus membranes;
1546ovary (2)3(4)-carpellate, with 2 ovules per carpel;
1547style with punctiform stigma. Fruit usually a
15481-seeded, loculicidal capsule, smooth. Seeds exar-
1549illate, with a small, pale hilum.
1550Two species, from southern Mexico to north-
1551ern South America.
15526.Handeliodendron Rehder
1553Handeliodendron Rehder, J. Arnold Arb. 16: 65 (1935).
1554Trees. Leaves opposite, digitate. Inflorescence
1555of terminal panicles. Flowers zygomorphic, bi-
1556sexual; sepals 5, distinct, imbricate; petals 4–5,
1557clawed, with 2 marginal appendages above the
1558claw; disk unilateral, lobed; stamens (7)8,
1559unequal; pollen colporate, striate, with heavily
1560verrucate colpus membranes (Fig. 79E); ovary
15613-carpellate, stipitate, with 2 ovules per carpel;
1562style short, stigma capitate. Fruit a (1–)3-locular,
1563stipitate, coriaceous, loculicidal capsule. Seeds
1564with a double arillode at base.
1565A single species, H. bodinieri (Le
´v.) Rehder,
1566endemic to China.
1567III.SUBFAM.DODONAEOIDEAE Burnett (1835).
1568Leaves alternate; petals usually without
1569appendages.
15703. Tribe Dodonaeaeae Kunth ex DC. (1824)
1571Disk (semi)annular; ovules (1)2(3 or 8) per loc-
1572ule; fruits dehiscent.
15737.Arfeuillea Pierre ex Radlk.
1574Arfeuillea Pierre ex Radlk. in Engl. & Prantl., Nat. Pflan-
1575zenfam. III, 5: 362 (1895); Radlk. in Engl., Pflanzenr. 98:
15761467 (1933); Welzen in Santisuk & Larsen, Fl. Thailand 7:
1577185, 186 (1999).
1578Falsely polygamous trees with simple and stellate
1579hairs. Leaves alternate, paripinnate; leaflets
1580crenate or entire; distal leaflet rudimentary. Inflo-
1581rescences axillary or terminal thyrses, with folia-
1582ceous bracts. Flowers zygomorphic, pistillate
1583or staminate on same inflorescence; sepals 5,
Sapindaceae 389
Uncorrected Proof
1584 petal-like, distinct, imbricate; petals 4, without
1585 appendages; disk semi-annular, or nearly com-
1586 plete, formed by two semi-annular halves; stamens
1587 (6)7–8(9); pollen colporate, striate; ovary 3-carpel-
1588 late, with 2 ovules per carpel. Fruit a 3-locular,
1589 loculicidal, papery capsule. Seed one per carpel,
1590 obovoid, exarillate.
1591 A single species, A. arborescens Pierre ex
1592 Radlk., from Thailand and Laos, cultivated
1593 throughout the tropics as an ornamental. Doubt-
1594 fully distinct from Majidea.
1595 8.Averrhoidium Baill. Fig. 81
1596 Averrhoidium Baill., Adansonia 11: 244 (1874).
1597 Dioecious trees. Leaves alternate, paripinnate;
1598 leaflets serrate or entire; distal leaflet rudimen-
1599 tary; stipules 0. Inflorescences axillary thyrses.
1600Flowers actinomorphic, functionally unisexual;
1601sepals 5, distinct, imbricate; petals 0 or rudimen-
1602tary, 1–4, without appendages; disk annular-
1603lobed; stamens (7)8; pollen colporate, striate;
1604ovary 3-carpellate, with 2 ovules per carpel; style
1605filiform, with 3 stigmatic groves. Fruit a 1-locular,
1606loculicidal, crustose capsule. Seeds 1–2 per fruit,
1607with sarcotesta.
1608Four species, one in Mexico, the remaining
1609from tropical South America.
16109.Cossinia Commers. ex Lam.
1611Cossinia Commers. ex Lam., Encycl. 2: 132 (1786). [also
1612spelled Cossignia]
1613Falsely polygamous trees or shrubs, with
1614stellate pubescence. Leaves alternate, trifoliolate
1615or imparipinnate, terminal leaflet well-developed.
1616Inflorescences terminal paniculate or corymbose
1617thyrses. Flowers actinomorphic or zygomorphic,
1618bisexual or functionally unisexual; sepals 5,
1619imbricate; petals 4–5, without appendages; disk
1620annular or semi-annular; stamens 5–6(–8); pollen
1621colporate, with clearly scabrate mesocolpia and
1622indistinctly scabrate to perforate apocolpia; ovary
16233-carpellate, with 2 ovules per carpel; stigma sub-
1624capitate. Fruit a septicidal, 3-locular, crustose
1625capsule. Seeds globose, exarillate.
1626Three species, two from Mauritius and one
1627from New Caledonia.
162810.Diplokeleba N.E. Brown
1629Diplokeleba N.E. Brown, Trans. & Proc. Bot. Soc. Edin-
1630burgh 20: 50 (1894).
1631Falsely polygamous trees. Leaves paripinnate;
1632leaflets entire or undulate; distal leaflet rudimen-
1633tary. Inflorescences terminal, corymbose thyrses.
1634Flowers functionally unisexual; calyx zygomor-
1635phic, the sepals 5, imbricate, outer sepals smaller;
1636petals 5, without appendages; disk annular, cup-
1637shaped, crenulate; stamens 8; pollen colporate,
1638irregularly striate to rugulate; ovary 3-carpellate,
1639with a single ovule per carpel; style filiform, with
16403 stigmatic lobes. Fruit a 3-locular, woody loculi-
1641cidal capsule. Seeds winged.
1642Two species, southern South America.
164311.Diplopeltis Endl.
1644Diplopeltis Endl. in Endl. et al., Enum. Pl. Hugel.: 13
1645(1837).
Fig. 81. Sapindaceae. Averrhoidium dalyi.AFruiting
branch. BFruits showing apical (left) and basal (right)
dehiscence. CSeed, lateral view (left) and ventral view
(right). DEmbryo, lateral view. (Drawn by A. Tangerini)
390 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
1646 Monoecious sub-shrubs. Leaves alternate, simple;
1647 stipules 0. Inflorescences terminal thyrses. Flow-
1648 ers zygomorphic; sepals 5, imbricate; petals 4(5),
1649 clawed, without appendages; disk unilateral,
1650 lobed, erect; stamens 8; pollen colporate, loosely
1651 to densely scabrate to finely striate; ovary 3-
1652 carpellate, with 2 ovules per carpel; style filiform.
1653 Fruit a 3-locular, loculicidal, crustose capsule or a
1654 schizocarp of 3 indehiscent cocci. Seeds arillate,
1655 ovoid, 1 or 2 per locule.
1656 Five species, Australia.
1657 12.Distichostemon F. Muell.
1658 Distichostemon F. Muell., Hooker’s J. Bot. Kew Gard.
1659 Misc. 9: 306 (1857).
1660 Falsely polygamous trees. Leaves alternate, simple,
1661 entire, serrate or crenate; stipules 0. Inflorescences
1662 axillary or terminal. Flowers actinomorphic,
1663 functionally unisexual; sepals 5–8, imbricate;
1664 petals 0; disk rudimentary; stamens 8–74; pollen
1665 colporate or brevicolporate, finely scabrate; ovary
1666 3(–6)-carpellate, with 2 ovules per carpel; style
1667 filiform, with 3 stigmatic lobes. Fruit a 3(–6)-
1668 locular, septifragal, marginicidal, capsule, the
1669 locules dorsally or distally winged, inflated or
1670 flattened. Seeds exarillate, subtended by a con-
1671 spicuous funiculus, with a conspicuous annular
1672 ring around the hilum, 2 per locule.
1673 Six species, Australia.
1674 13.Dodonaea Miller Fig. 82
1675 Dodonaea Miller, Gard. Dict. Abr. ed.: 4 (1754); Leenh-
1676 outs, Blumea 28: 271 (1983), reg. rev.; West, Brunonia 7:
1677 18 (1984), reg. rev.
1678 Dioecious or falsely polygamous-dioecious
1679 shrubs or trees, with viscous glandular hairs.
1680 Leaves alternate or rarely opposite, simple or
1681 paripinnate; distal leaflet fully developed or rudi-
1682 mentary. Inflorescences axillary or terminal
1683 racemes or thyrses. Flowers actinomorphic, uni-
1684 sexual or bisexual; sepals (3–)5(–7), imbricate
1685 or valvate; petals 0; disk 0 or rudimentary in
1686 pistillate flowers; stamens 5–15; pollen colporate,
1687 finely to coarsely scabrate, with scabrae often
1688 in coarse patterns (Fig. 79B); ovary (2)3–5(6)-
1689 carpellate, with 2 ovules per carpel; style filiform;
1690 stigma grooved or divided. Fruit a 2–6-locular,
1691 septifragal or septicidal capsule, the locules
1692dorsally winged. Seed exarillate, on enlarged
1693placenta. 2n¼28, 30, 32.
1694Primarily Australian with 59 endemic species
1695and 8 pantropical species.
169614.Euphorianthus Radlk.
1697Euphorianthus Radlk., Sitzungsber. Math.-Phys. Cl.
1698K€
onigl. Bayer. Akad. Wiss. M€
unchen 9: 673 (1879); Leenh-
1699outs, Blumea 33: 198 (1988), rev.
1700Falsely polygamous trees. Leaves alternate, pari-
1701pinnate; distal leaflet rudimentary. Inflorescences
1702axillary to pseudo-terminal thyrses. Flowers acti-
1703nomorphic, bisexual or functionally unisexual;
1704sepals 5, distinct, imbricate; petals 5, clawed,
1705with a pair or woolly appendages above the
1706claw; disk annular; stamens (6)7–8; pollen para-
1707syncolporate, rugulate; ovary 3-carpellate, with a
1708single ovule per carpel; style filiform; stigma
1709grooved to slightly lobed. Fruit an incompletely
17103-locular, sub-globose, velvety, fleshy, loculicidal
Fig. 82. Sapindaceae. Dodonaea viscosa Jacq. AFruiting
branch. BCapsules. CD. angustifolia, fruit. DD. polyan-
dra, fruit. (Adema et al. 1994; drawn by J. Wessendorp)
Sapindaceae 391
Uncorrected Proof
1711 capsule, the endocarp sericeous. Seeds with a
1712 2(3)-lobed sarcotesta around the hilum.
1713 A single species, E. euneurus (Miq.) Leenh.,
1714 found in eastern Malesia, from the Philippines
1715 and Celebes to Vanuatu. The wood is used in
1716 the construction of houses.
1717 15.Eurycorymbus Hand.-Mazz.
1718 Eurycorymbus Hand.-Mazz., Akad. Wiss. Wien. Math.
1719 Naturwiss. Kl. Anz. 59: 104 (1922).
1720 Dioecious trees. Leaves alternate, paripinnate;
1721 leaflets serrate; distal leaflet rudimentary. Inflor-
1722 escences axillary, corymbose thyrses. Flowers
1723 actinomorphic, unisexual; sepals 5; petals 5,
1724 spatulate, without appendages; disk annular,
1725 dish-shaped-crenate; stamens 8; pollen colporate,
1726 striate; ovary 3(4)-carpellate, with 2 ovules per
1727 carpel; style filiform, with 3 stigmatic branches.
1728 Fruit a 1(3)-coccate, fleshy, loculicidal capsule.
1729 Seeds exarillate, sub-globose, puberulent, one
1730 per locule, persistent on receptacle.
1731 A single species, E. austrosinensis Hand.-
1732 Mazz., from China.
1733 16.Harpullia Roxb.
1734 Harpullia Roxb., Fl. Ind. 2: 441 (1824); Leenhouts &
1735 Vente, Blumea 28: 1 (1982), rev.
1736 Falsely polygamous shrubs or trees. Indumentum
1737 of stellate hairs. Leaves alternate, paripinnate;
1738 distal leaflet rudimentary; petiole and rachis
1739 winged or unwinged. Inflorescences axillary, ter-
1740 minal, ramiflorous, or cauliflorous thyrses. Flow-
1741 ers actinomorphic, bisexual or functionally
1742 unisexual; sepals 5, distinct, imbricate; petals 5,
1743 clawed, with 2 marginal appendages above the
1744 claw or the petals sessile without appendages;
1745 disk annular; stamens 5–8; pollen colporate,
1746 sometimes cryptoaperturate, striate, striate-
1747 rugulate, rugulate, reticulate with scabrae or
1748 coarsely reticulate with finely striate to scabrate
1749 muri (Fig. 79D); ovary 2–3(4)-carpellate, with 1
1750 or 2 ovules per carpel; style with 2 or 3 stigmatic
1751 lines. Fruit a 2–3-locular, chartaceous to woody
1752 loculicidal capsule. Seeds with an orange sarco-
1753 testal ring around the hilum or completely
1754 covered by an aril that is basally adnate to the
1755 testa. 2n¼30.
1756 About 26 species from India, Sri Lanka,
1757 southeastern China, Malesia, Australia to New
1758Caledonia and Tonga. Several species are
1759cultivated for their wood or as ornamentals.
176017.Llagunoa Ruiz & Pavo
´n
1761Llagunoa Ruiz & Pavo
´n, Prodr.: 126 (1794).
1762Falsely polygamous shrubs or trees. Leaves alter-
1763nate, simple or trifoliolate. Flowers solitary or in
1764axillary cymes. Flowers zygomorphic, bisexual
1765or functionally unisexual; sepals 5, imbricate;
1766petals 0; disk unilateral, semi-annular; stamens
17678; pollen colporate, striate; ovary 3-carpellate,
1768with 2 ovules per carpel; style subulate; stigma
1769subcapitate. Fruit a 3-locular, 3-lobed, crustose,
1770loculicidal capsule. Seeds exarillate. 2n¼20.
1771Three or four species from the Andean high-
1772lands of South America.
177318.Loxodiscus Hook. f.
1774Loxodiscus Hook. f., Hooker’s J. Bot. Kew Gard. Misc. 9:
1775200 (1857).
1776Falsely polygamous shrubs or trees. Leaves alter-
1777nate, imparipinnate; leaflets serrate; distal leaflet
1778rudimentary. Inflorescences terminal thyrses.
1779Flowers zygomorphic, bisexual or functionally
1780unisexual; sepals 5, imbricate, with fimbriate-
1781glandular margins; petals 4(5), clawed, without
1782appendages; disk semi-annular, double, the
1783inner lobe concave, 4-dentate; stamens 7–8;
1784pollen colporate, striate; ovary 3-carpellate, with
1785stipitate glands and 2 ovules per carpel; style
1786subulate; stigma obtuse. Fruit a 3-locular, loculi-
1787cidal, membranous capsule. Seeds exarillate.
1788A single species, L. coriaceus Hook. f., from
1789New Caledonia.
179019.Magonia St. Hil.
1791Magonia St. Hil., Bull. Sci. Soc. Philom. Paris 1824: 78
1792(1824); Joly et al., Brittonia 32: 380–386 (1980).
1793Falsely polygamous trees. Leaves alternate, pari-
1794pinnate; leaflets entire; distal leaflet rudimentary.
1795Inflorescences axillary or terminal thyrses. Flowers
1796zygomorphic, bisexual or functionally unisexual;
1797sepals 5, distinct, imbricate; petals 5, without
1798appendages; disk complete, half of one side con-
1799sisting of 4 short glands, the other half of two
1800concentric erect, fleshy laminae, the outer one
1801taller; stamens 8; pollen grains in tetrads,
392 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
1802 colporate (6 pairs of colpi per tetrad), striate-
1803 gemmate (Fig. 79F); ovary 3-carpellate, with
1804 8 ovules per carpel; style short, stigma capitate,
1805 sub-3-lobed. Fruit a large, 3-locular, trigonous,
1806 woody loculicidal capsule. Seeds flattened, sur-
1807 rounded by a wing. 2n¼30.
1808 One species, M. pubescens St.-Hil., from
1809 Brazil, Bolivia, and Paraguay.
1810 20.Majidea Kirk ex D. Oliver
1811 Majidea Kirk ex D. Oliver, Hooker’s Icon. Pl. tab. 78, 1097
1812 (1871).
1813 Falsely polygamous trees. Indument of fasciculate
1814 stellate hairs. Leaves alternate, paripinnate; distal
1815 leaflet rudimentary. Inflorescences axillary or
1816 terminal thyrses. Flowers actinomorphic or
1817 zygomorphic, bisexual or functionally unisexual;
1818 sepals 5, imbricate; petals 0 or only 1–2 or 4; disk
1819 annular or semi-annular, dish-shaped to penta-
1820 gonous; stamens 7–8; pollen colporate, striate;
1821 ovary 3-carpellate; with 2 ovules per carpel; sty-
1822 le elongated with punctiform stigma. Fruit a
1823 3-locular, deeply lobed, loculicidal capsule, with
1824 crustose pericarp. Seeds exarillate. 2n¼24.
1825 Four or five species from Africa and
1826 Madagascar.
1827 4. Tribe Doratoxyleae Radlk. (1890).
1828 Disk annular; ovules (1)2(3) per locule; fruits
1829 indehiscent.
1830 21.Doratoxylon Thouars ex Hook. f.
1831 Doratoxylon Thouars ex Hook. f. in Benth. & Hook. f.,
1832 Gen. 1: 408 (1862).
1833 Falsely polygamous-dioecious shrubs or treelets.
1834 Leaves alternate, paripinnate or imparipinnate;
1835 leaflets entire or crenate; distal leaflet rudimen-
1836 tary; rachis narrowly winged; stipules 0. Inflores-
1837 cences axillary, glomerate cymes. Flowers
1838 actinomorphic, functionally unisexual or bisex-
1839 ual; sepals 5, imbricate; petals 0; disk annular-
1840 5-lobed; stamens 5(6–7), the anthers dorsifixed;
1841 pollen colporate, scabrate to coarsely verrucate
1842 with striate to rugulate verrucae; ovary 2-
1843 carpellate, with 2 ovules per carpel; style short;
1844 stigma capitate. Fruits baccate, indehiscent, 1-
1845 locular. Seed ellipsoid, exarillate.
1846 Six species, one from Mauritius and five from
1847 Madagascar.
184822. Euchorium Ekm. & Radlk.
1849Euchorium Ekm. & Radlk., Repert. Spec. Nov. Regni Veg.
185021: 230 (1925).
1851Dioecious trees. Leaves alternate, paripinnate;
1852distal leaflet rudimentary; stipules 0. Inflore-
1853scences of axillary racemes. Flowers unisexual;
1854calyx zygomorphic, sepals 5, imbricate, the
1855outer two smaller; petals 5, without appendages;
1856disk annular; stamens 8; pollen unknown;
1857ovary 3-carpellate, with 2 ovules per carpel; style
1858filiform, obtuse. Fruits unknown.
1859A single species, E. cubense Ekm. & Radlk.,
1860endemic to western Cuba.
1861In the absence of information on fruits, this
1862genus is included in tribe Dodonaeaeae because
1863of its resemblance to Exothea.
186423.Exothea Macfad.
1865Exothea Macfad., Fl. Jamaica 1: 232 (1837).
1866Falsely polygamous dioecious trees. Leaves alter-
1867nate, paripinnate; distal leaflet rudimentary. In-
1868florescences axillary or sub-terminal corymbose
1869thyrses. Flowers bisexual or functionally unisex-
1870ual; calyx zygomorphic, sepals 5, imbricate, the
1871outer two smaller; petals 5, without appendages;
1872disk annular, dish-shaped-crenate; stamens (7)8
1873(–10); pollen colporate, loosely scabrate to verru-
1874cate (Fig. 79C); ovary 2-carpellate, with 2 ovules
1875per carpel; style elongated; stigma capitate. Fruits
18761(2)-locular, globose, baccate, indehiscent. Seeds
1877with coriaceous-fleshy testa.
1878Three species, distributed throughout the
1879West Indies, Mexico, Guatemala, Colombia, and
1880Ecuador.
188124.Filicium Thwaites ex Hook. f.
1882Filicium Thwaites ex Hook. f. in Benth. & Hook. f., Gen.
1883Pl. 1: 325. (1862); Thwaites, Enum. Pl. Zeyl.: 58, 408
1884(1864); Welzen in Fl. Males. I, 11: 754 (1994); Welzen in
1885Fl. Thailand 7: 198, 199 (1999).
1886Falsely polygamous trees. Leaves alternate, paripin-
1887nate; distal leaflet rudimentary; rachis (broadly)
1888winged. Inflorescences axillary or pseudo-termi-
1889nal thyrses. Flowers actinomorphic, functionally
1890unisexual; sepals 5, distinct, valvate; petals 5,
1891without appendages; disk annular, 5-lobed,
1892woolly; stamens 5; pollen colporate, loosely
Sapindaceae 393
Uncorrected Proof
1893 scabrate; ovary 2-carpellate, with a single pendant
1894 ovule per carpel; style short; stigma lobed. Fruits
1895 1–2-locular, baccate, indehiscent, smooth, gla-
1896 brous, the pericarp fleshy. Seeds exarillate.
1897 2n¼32.
1898 Three or four species, mainly in east Africa
1899 and Madagascar, one extending to India and Sri
1900 Lanka, and widely cultivated as an ornamental
1901 throughout the tropics.
1902 The taxonomic placement of Filicium has
1903 been much debated over decades. It was origi-
1904 nally described as an Anacardiacea, and later
1905 transferred into Sapindaceae. In 1862, it was
1906 transferred into Burseraceae by Hooker f.,
1907 followed by a transfer back into Sapindaceae
1908 by Baillon in 1874. In 1890, Filicium was placed
1909 by Radlkofer into the tribe Doratoxyleae of
1910 the subfamily Dodonaeoideae, in spite of its
1911 uniovular carpels; this position is supported by
1912 Harrington et al. (2005).
1913 25.Ganophyllum Blume
1914 Ganophyllum Blume, Mus. Bot. Lugd.-Bat. 1: 230 (1850)
1915 [1851]; Radlk. in Engl., Pflanzenreich 98: 1423 (1933);
1916 Leenhouts in Fl. Males. I, 11: 538 (1994).
1917 Dioecious trees. Indument of glandular scales.
1918 Leaves alternate, paripinnate; distal leaflet rudi-
1919 mentary. Inflorescences axillary thyrses. Flowers
1920 actinomorphic; calyx cup-shaped, (4)5(–7)-
1921 lobed, valvate; petals 0; disk annular-lobed;
1922 stamens 5(–7); pollen colporate, irregularly ver-
1923 rucate; ovary 2(3)-carpellate, with 2 ovules per
1924 carpel; style short; stigma indistinctly lobed.
1925 Fruits indehiscent, 2-locular, ovoid, baccate,
1926 with papery endocarp and leathery mesocarp.
1927 Seeds exarillate.
1928 One or two species, from western and central
1929 Africa, Andaman and Nicobar Islands, through-
1930 out Malesia to northeastern Australia, New
1931 Guinea, and the Solomons.
1932 26.Hippobromus Ecklon & Zeyher
1933 Hippobromus Ecklon & Zeyher, Enum.: 151 (1836).
1934 Falsely polygamous dioecious trees. Leaves
1935 alternate, paripinnate; leaflets serrate; distal
1936 leaflet rudimentary; rachis winged. Inflores-
1937 cences axillary thyrses. Flowers zygomorphic,
1938 bisexual or functionally unisexual; sepals 5,
1939connate at base, imbricate; petals 5(6), shorter
1940than the sepals, without appendages; disk annu-
1941lar, pentagonous; stamens 8; pollen colporate,
1942rugulate; ovary 3-carpellate, with 2 or less often
19433 ovules per carpel; style short; stigma decur-
1944rent. Fruits 1-locular, indehiscent, baccate.
1945Seeds exarillate.
1946A single species, H. alatus Ecklon & Zeyher,
1947Africa.
194827.Hypelate P. Browne
1949Hypelate P. Browne, Civ. Nat. Hist. Jamaica: 208 (1756).
1950Falsely polygamous shrubs or trees. Leaves
1951trifoliolate; stipules 0. Inflorescences axillary or
1952terminal thyrses. Flowers bisexual or functionally
1953unisexual; calyx zygomorphic, sepals (4)5, imbri-
1954cate, the outer two smaller; petals (4)5, without
1955appendages; disk annular, obsolete 5-lobed; sta-
1956mens (7)8; pollen colporate, loosely verrucate;
1957ovary 3-carpellate, with 2 ovules per carpel; style-
1958short; stigma capitate. Fruit a 1-locular, indehis-
1959cent drupe with woody endocarp and fleshy
1960mesocarp and exocarp. Seeds exarillate, with
1961papery testa.
1962A single species, H. trifoliata Sw., endemic to
1963the West Indies and the southeastern United
1964States (Florida).
196528.Zanha Hiern
1966Zanha Hiern, Cat. African Pl. 1: 128 (1896).
1967Dialiopsis Radlk. (1902) [nomen nudum] and in Engl. &
1968Prantl., Nat. Pflanzenfam., Nachtr. 3: 207 (1907).
1969Dioecious trees. Leaves alternate, paripinnate;
1970leaflets 3–7 pairs, entire or serrulate; distal leaflet
1971rudimentary; stipules 0. Inflorescences distal,
1972short thyrses. Flowers actinomorphic, function-
1973ally unisexual; sepals 4–5(6), imbricate, connate
1974on lower 1/3–1/2; petals 0; disk annular or funnel-
1975shaped; stamens (3)4–5, coiled in bud; pollen
1976colporate, loosely scabrate; ovary 2-carpellate,
1977with 2 ovules per carpel; style short, filiform,
1978stigmas 2-lobed. Fruit 1-locular, indehiscent,
1979with fleshy mesocarp and coriaceous endocarp.
1980Seed 1 per fruit, with coriaceous or slightly fleshy
1981testa.
1982Three species from southern Africa and
1983Madagascar. Fruits of Z. suaveolens Capuron are
1984said to be edible.
394 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
1985 IV. SUBFAM.SAPINDOIDEAE
1986 Leaves alternate; petal appendages usually pres-
1987 ent; disk annular or unilateral; carpels uniovulate.
1988 5. Tribe Paullinieae Kunth ex DC. (1824)
1989 Tribe Thouinieae Blume (1847).
1990 Herbaceous vines, shrubs or small trees,
1991 often stipulate; leaves imparipinnate; flowers
1992 zygomorphic.
1993 29.Allophylus L
1994 Allophylus L., Sp. Pl. 348 (1753); Leenhouts, Blumea 15:
1995 301–358 (1967); Fritsch, Kulturpfl. 18: 194 (1970);
1996 Ferrucci, Bol. Soc. Argentina Bot. 24: 200–202 (1985).
1997 Duodichogamous or dioecious shrubs or trees,
1998 or less often scandent shrubs. Leaves alternate,
1999 trifoliolate or less often unifoliolate or digitate (in
2000 a few Asian collections); leaflets serrate, dentate
2001 or entire; petioles elongated; stipules 0. Inflore-
2002 scences axillary racemes, panicles or thyrses.
2003 Flowers zygomorphic, 4-merous, functionally
2004 pistillate or staminate; sepals distinct, imbricate,
2005 the outer distinctly smaller than the inner; petals
2006 with a single, 2-lobed appendage or 2 marginal
2007 appendages; disk unilateral; stamens 8; pollen
2008 brevicolporate to porate, rugulate, sometimes
2009 striate, reticulate or psilate; ovary 2(3)-carpellate,
2010 with a single ovule per carpel; style with 2–3
2011 stigmatic branches. Fruit an indehiscent drupe,
2012 1–2-coccate, with crustose endocarp and fleshy
2013 exocarp. Seeds exarillate, with papery testa.
2014 2n¼28.
2015 Depending on circumscription, a single
2016 species or about 250 species with circumtropical
2017 distribution. Several species are planted as
2018 ornamentals.
2019 The notorious difficulties in species delimita-
2020 tion in Allophylus led Leenhouts (1967) to revise
2021 the status of the genus, which he found to be
2022 dominated by an enormous degree of clinal
2023 variation and intergradation among regional
2024 populations. As a consequence, he suggested to
2025 treat Allophylus as a single polymorphic species
2026 with the possibility of adding existing species
2027 names for informally characterizing local races.
2028 Since then, however, no progress in an under-
2029 standing of the systematic structure of Allophylus
2030has been made, and most botanists still continue
2031recognizing regional populations as species.
203230.Athyana (Griseb.) Radlk.
2033Athyana (Griseb.) Radlk. in T. Durand, Index Gen. Phan.:
203473 (1887) [1888].
2035Monoecious trees. Leaves alternate, imparipin-
2036nate; leaflets serrate; distal leaflet fully developed;
2037rachis winged; stipules 0. Inflorescences axillary
2038or pseudo-terminal thyrses. Flowers zygomor-
2039phic, functionally unisexual; sepals 5, of similar
2040size, valvate; petals 4, of same length, with a single
2041petaloid hood-shaped appendage; disk semi-
2042annular-lobed; stamens (7)8; pollen colporate,
2043finely perforate; ovary 3-carpellate, with a single
2044ovule per carpel; stigma capitate. Fruit schizocar-
2045pic, splitting into 3 samaroid mericarps with a
2046distal wing. Seed exarillate.
2047A single species, A. weinmannifolia Radlk.,
2048southern South America (Peru, Bolivia,
2049Argentina).
205031.Bridgesia Bertero ex Cambess., nom. cons.
2051Bridgesia Bertero ex Cambess., Nouv. Ann. Mus. Hist.
2052Nat. 3: 234 (1834), non Hook. (1831).
2053Falsely polygamous shrubs. Leaves alternate,
2054simple, lobed or serrate; stipules 0. Inflorescence
2055a short, axillary cyme or flowers solitary. Flowers
2056zygomorphic, unisexual or bisexual; sepals 5,
2057imbricate; petals 4, with a single cucullate, crested
2058appendage; disk semi-annular, 4-lobed; stamens 8;
2059pollen colporate, striate; ovary 3-carpellate, with
2060a single ovule per carpel; style filiform with 3
2061stigmatic branches. Fruit schizocarpic, splitting
2062into 3 samaroid, papery mericarps with a short
2063dorsal wing. Seed exarillate.
2064A single species, B. incisifolia Bertero &
2065Cambess., from Andean Chile.
206632.Cardiospermum L.
2067Cardiospermum L., Sp. Pl.: 366 (1753).
2068Monoecious herbaceous vines. Leaves alternate,
2069ternately compound or biternate; stipules minute.
2070Inflorescences axillary thyrses bearing tendrils at
2071base of rachis. Flowers zygomorphic, functionally
2072staminate or pistillate; sepals 4–5, unequal, imbri-
2073cate; petals 4, with a basal, hood-shaped append-
2074age; disk unilateral, 2-lobed; stamens 8; pollen
Sapindaceae 395
Uncorrected ProofUncorrected Proof
2075 heteropolar, demi(syn)colporate proximally, per-
2076 forate to reticulate; ovary 3-carpellate, with
2077 a single ovule per carpel; style with 3 stigmatic
2078 branches. Fruit a septifragal, marginicidal, in-
2079 flated, capsule with thin-membranous walls.
2080 Seeds black, with a small, heart-shaped arillode
2081 surrounding the micropyle. x¼10, 11.
2082 About 15 species native to the Neotropics,
2083 3 of which are widely distributed throughout
2084 the tropics. Two species are cultivated as
2085 ornamentals.
2086 33.Diatenopteryx Radlk.
2087 Diatenopteryx Radlk., Sitzungsber. Math.-Phys. Cl.
2088 K€
onigl. Bayer. Akad. Wiss. M€
unch. 8: 284 (1878);
2089 Ferrucci, Sapindaceae in Spichiger & Ramella, Flora del
2090 Paraguay (1991).
2091 Falsely polygamous shrubs or trees. Indumentum
2092 of simple hairs and scales. Leaves alternate,
2093 imparipinnate; leaflets dentate or serrate; distal
2094 leaflet rudimentary or fully developed; stipules 0.
2095 Inflorescences axillary thyrses. Flowers zygomor-
2096 phic, functionally unisexual; sepals 4, distinct,
2097 imbricate; petals 4, with a basal hood-shaped,
2098 crested appendage; disk semi-annular, lobed;
2099 stamens (6–)8; pollen colporate, rugulate; ovary
2100 2-carpellate, with a single ovule per carpel; style
2101 filiform; stigma subcapitate. Fruits schizocarpic,
2102 splitting into 2 samaroid mericarps with a long
2103 distal wing. Seed exarillate.
2104 Two species from southern South America
2105 (Brazil, Bolivia, Paraguay, and Argentina).
2106 34.Guindilia Gilles ex Hook. & Arn.
2107 Guindilia Gilles ex Hook. & Arn., Hooker’s Bot. Misc. 3:
2108 170 (1833).
2109 Valenzuelia Bert. ex Cambess. (1834), non Mutis (1810).
2110 Falsely polygamous trees. Leaves alternate, sim-
2111 ple, opposite, entire or tridentate at apex; stipules
2112 0. Inflorescences axillary cymes. Flowers zygo-
2113 morphic, bisexual or functionally unisexual;
2114 sepals 5, imbricate; petals 4(5), with a hood-
2115 shaped, crested, ventral appendage; disk unilat-
2116 eral, modified into a 2-lobed mount; stamens 8;
2117 pollen colporate, striate; ovary 3-carpellate, with
2118 a single ovule per carpel; style filiform; stigma
2119 3-lobed. Fruit schizocarpic, splitting into (1–)3
2120 sub-globose, crustose mericarps. Seed exarillate.
2121 Three species from southern South America.
212235.Houssayanthus Hunziker
2123Houssayanthus Hunziker, Kurtziana 11: 17 (1978);
2124Ferrucci, Bonplandia 5 (19): 164–174 (1981); Ferrucci,
2125Candollea 41: 218 (1986) and ibid. 42: 805–807 (1987);
2126Rzedowski & Caldero
´n de Rzedowski, Acta Bot. Mexicana
212776: 89–98 (2006).
2128Woody vines. Cross section of stem simple or
2129with a central and three peripheral vascular cylin-
2130ders. Leaves trifoliolate, 5-pinnately compound,
2131or biternate; stipules minute. Inflorescences
2132axillary thyrses with a pair of tendrils at base of
2133rachis. Flowers zygomorphic, functionally unisex-
2134ual; sepals 4 or 5, unequal, imbricate; petals 4, with
2135a basally adnate hood-shaped appendage; disk
2136unilateral, 4-lobed; stamens 8; pollen heteropolar,
2137demisyncolporate, perforate; ovary 3-carpellate,
2138with a single ovule per carpel; style with 3 stig-
2139matic branches. Fruit a schizocarp splitting into 3
2140mericarps with a short dorsal wing. Seeds lentic-
2141ular, exarillate. 2n¼24.
2142Five species, two from Mexico, one from
2143Venezuela, and two from Brazil, Bolivia,
2144Paraguay, and northeastern Argentina.
214536.Lophostigma Radlk.
2146Lophostigma Radlk. in Engl. & Prantl, Nat. Pflanzenfam.,
2147Nachtr. 1: 228 (1897); Acevedo-Rodrı
´guez, Syst. Bot. 18:
2148379–388 (1993), rev.
2149Woody vines. Cross section of stem with a single
2150vascular cylinder. Leaves alternate, trifoliolate; sti-
2151pules minute. Inflorescences axillary thyrses with a
2152pair of tendrils at base of rachis. Flowers zygomor-
2153phic, functionally unisexual; sepals 5, unequal,
2154imbricate; petals 4, much shorter than the sepals,
2155with 2 basally adnate digitate appendages; disk
2156unilateral, 4-lobed; stamens 8, with short fila-
2157ments; pollen cylindric-ellipsoid, 4-aperturate,
2158possibly a lengthy variant of heteropolar 3-demi-
2159syncolporate Paullinieae pollen (e.g., Serjania);
2160ovary 3-carpellate, with a single ovule per carpel;
2161style with 3 stigmatic branches. Fruit a schizocarp
2162splitting into 3 mericarps with an elongated proxi-
2163mal wing. Seeds lenticular, exarillate.
2164Two species, one from Ecuador and Peru, the
2165other from Bolivia.
216637.Paullinia L.
2167Paullinia L., Sp. Pl.: 365 (1753); Radlkofer, Monogr.
2168Paullinia (1895); Simpson, Fieldeana Bot. 36: 125–164
2169(1976), reg. rev.
396 P. Acevedo-Rodrı
´guez et al.
Uncorrected ProofUncorrected Proof
2170 Woody vines, usually producing milky sap. Cross
2171 section of stem with a central and 3–5 peripheral
2172 vascular cylinders, or with a single vascular
2173 cylinder. Leaves alternate, trifoliolate, pinnate
2174 (5-pinnate), bipinnate, biternate, or variously
2175 dissected; stipules minute to large. Inflorescences
2176 axillary thyrses with a pair of tendrils at base of
2177 rachis, or cauliflorous thyrses without tendrils.
2178 Flowers zygomorphic, functionally unisexual;
2179 sepals 4–5, unequal, imbricate; petals 4, with
2180 a basally adnate hood-shaped appendage; disk
2181 unilateral, 4-lobed; stamens 8; pollen porate,
2182 perforate; ovary 3-carpellate, with a single ovule
2183 per carpel; style with 3 stigmatic branches. Fruit a
2184 1–3-locular, septifragal-marginicidal capsule, the
2185 locules dorsally winged or exalate, exceptionally
2186 spiny. Seeds globose, oblong or ellipsoid, with a
2187 partial to nearly complete sarcotesta. 2n¼24.
2188 About 190 species native to the Neotropics,
2189 one species extending into Africa. Numerous
2190 species are used as fish poisons; P. cupana
2191 Kunth is the source of the economically impor-
2192 tant “guarana
´” used in the confection of a stimu-
2193 lating drink or soft drinks in Brazil.
2194 38.Serjania Plum. ex Miller Fig. 83
2195 Serjania Plum. ex Miller, Gard. Dict. Abr. ed. 4 (1754);
2196 Acevedo-Rodrı
´guez, Mem. New York Bot. Gard. 67: 1–93
2197 (1993).
2198 Chimborazoa H.T. Beck (1994); Acevedo-Rodrı
´guez,
2199 Novon 8: 106 (1998).
2200 Duodichogamous woody or herbaceous vines,
2201 often producing milky sap. Cross section of
2202 stem with a single or multiple vascular cylinders,
2203 usually 3, but also 8 or 10. Leaves alternate, ternately
2204 compound (ternate, biternate, or triternate) or 5-
2205 pinnately compound; stipules small and early
2206 deciduous. Inflorescences axillary or terminal
2207 thyrses. Flowers zygomorphic, functionally uni-
2208 sexual; sepals 4–5, unequal, imbricate; petals 4,
2209 with a basally adnate hood-shaped appendage;
2210 disk unilateral, 2- or 4-lobed; stamens 8; pollen
2211 heteropolar, demi(syn)colporate proximally,
2212 sometimes with short distal demicolpi as well,
2213 perforate, sometimes reticulate, psilate or faintly
2214 rugulate; ovary 3-carpellate, with a single ovule
2215 per carpel; style with 3 stigmatic branches. Fruit
2216 schizocarpic, splitting into three samaroid meri-
2217 carps, each with a proximal wing. Seeds lenticular
2218 to globose, exarillate. 2n¼24.
2219About 230 species native to tropical and sub-
2220tropical America.
222139.Thinouia Triana & Planchon
2222Thinouia Triana & Planchon, Ann. Sci. Nat. Bot. IV, 18:
2223368 (1862).
2224Allosanthus Radlk. (1933).
2225Lianas. Secondary growth of stems with numer-
2226ous cortical steles. Leaves alternate, trifoliolate;
2227stipules minute. Inflorescences umbelliform
2228thyrses, seldom bearing tendrils, axillary or
2229aggregate into terminal thyrsoid inflorescences.
2230Flowers actinomorphic, bisexual, 5-merous; calyx
2231cup-shaped, the sepals valvate; petals obovate
2232to spatulate, with a pair of short, marginal appen-
2233dages; disk annular; stamens 6–8; pollen colpo-
2234rate, striate; ovary 3-carpellate, with a single
2235ovule per carpel; style elongated with 3 stig-
2236matic branches. Fruit schizocarpic, splitting into
Fig. 83. Sapindaceae. Serjania polyphylla.AFertile
branch. BBiternate leaf with tendril. CMale flower.
DPetal with adnate hood-shaped appendage. ELongitu-
dinal section of male flower. FWinged mericarp attached
to floral axis. GEmbryo. HCross section of stem showing
numerous vascular cylinders. (Acevedo-Rodrı
´guez 1996;
reproduced with permission of the artist Bobbi Angell)
Sapindaceae 397
Uncorrected Proof
2237 3 mericarps, each with a distal wing. Seeds nearly
2238 spherical, exarillate.
2239 About 12 species from Central and South
2240 America.
2241 40.Thouinia Poit., nom. cons.
2242 Thouinia Poit., Ann. Mus. Natl. Hist. Nat. 3: 70 (1804),
2243 non L. f. (1782), nom. rej.; Votava, Taxonomic revision of
2244 the genus Thouinia (Sapindaceae). Ph.D. dissertation,
2245 Columbia University, New York, 235 pp. (1973), rev.
2246 Falsely polygamous shrubs or trees. Leaves alter-
2247 nate, trifoliolate or unifoliolate; stipules 0. Inflor-
2248 escences axillary, racemose thyrses. Flowers
2249 zygomorphic, functionally unisexual; sepals 4,
2250 imbricate; petals 4, with a ventral, bifid append-
2251 age; disk unilateral, lobed; stamens 8; pollen
2252 brevicolporate, rugulate; ovary 3-carpellate,
2253 with a single ovule per carpel; style filiform,
2254 with 3 stigmatic branches. Fruit schizocarpic,
2255 splitting into 3 samaroid mericarps, each bearing
2256 a distal wing. Seeds exarillate.
2257 About 30 species from Central America and
2258 the West Indies.
2259 41.Urvillea Kunth
2260 Urvillea Kunth in H.B.K., Nov. Gen. Spec. 5: 105 (1821).
2261 Herbaceous to woody vines. Stems terete and
2262 lenticellate, becoming trilobate at age, producing
2263 milky sap. Leaves alternate, trifoliolate or biter-
2264 nate; stipules minute, deciduous. Inflorescences
2265 axillary thyrses. Flowers zygomorphic, bisexual;
2266 sepals 5, unequal, imbricate; petals 4, spatulate,
2267 with an adnate hood-shaped appendage on adax-
2268 ial surface; disk unilateral, 4-lobed, receptacle
2269 enlarged into a short androgynophore; stamens
2270 8; pollen heteropolar, demisyncolporate proxi-
2271 mally, often with short distal demicolpi as
2272 well, perforate, indistinctly rugulate or reticulate
2273 (Fig. 79G–I); ovary 3-carpellate, with a single ovule
2274 per carpel; style with 3 stigmatic branches. Fruit a
2275 3-locular, thin, papery, semi-inflated capsule,
2276 with a narrow, marginal (septal) wing. Seeds
2277 sub-globose, with a heart-shaped or reniform,
2278 white arillode around the hilum. 2n¼22.
2279 About 15 species from Central and South
2280 America.
2281 6. Tribe Melicocceae Blume (1847).
2282 Flowers actinomorphic; fruits indehiscent, usu-
2283 ally single seeded; seeds with sarcotesta.
228442.Melicoccus P. Browne Fig. 77
2285Melicoccus P. Browne, Hist. Jamaica: 210 (1756); Acevedo-
2286Rodrı
´guez, Moscosoa 9: 58–61 (1997); Acevedo-
2287Rodrı
´guez, Fl. Neotrop. 87 (2002), rev.
2288Melicocca L. (1762), nom. illeg.
2289Dioecious or monoecious trees. Leaves alternate,
2290paripinnate; leaflets 1–2 pairs; distal leaflet rudi-
2291mentary, 0 or exceptionally present; rachis usu-
2292ally winged; stipules 0. Inflorescences terminal
2293panicles or racemes. Flowers actinomorphic,
2294functionally pistillate or staminate. Calyx cup-
2295shaped, sepals 4(5), equal, imbricate; petals 4(5)
2296erect or reflexed; appendages 0 or rudimentary;
2297disk annular, slightly lobed; stamens 8; pollen
2298colporate, striate; ovary 2-carpellate, unilocular,
2299with a single ovule per carpel; style obsolete;
2300stigmatic surface lobed, subcapitate. Fruit inde-
2301hiscent, with leathery pericarp, sub-globose or
2302ellipsoid. Seeds 1(2) with edible sarcotesta.
23032n¼32.
2304Ten species, native to South America and
2305Dominican Republic. Melicoccus bijugatus Jacq.
2306is widely cultivated in the Neotropics for its
2307edible, tasty fruits.
230843.Talisia Aublet Fig. 84
2309Talisia Aubl., Hist. Pl. Guiane 1: 349 (1775); Mennega,
2310Acta Bot. Neerl. 21: 578–586 (1972); Acevedo-Rodrı
´guez,
2311Fl. Neotrop. 87: 1–179 (2003), rev.
2312Tapirocarpus Sagot (1882).
2313Duodichogamous unbranched shrubs or small
2314to large trees. Leaves alternate, paripinnate
2315or imparipinnate; leaflets entire; distal leaflet
2316rudimentary. Inflorescences axillary, terminal or
2317cauliflorous thyrses. Flowers 5-merous, actino-
2318morphic, staminate or pistillate; calyx usually
2319cup-shaped, sepals imbricate or valvate, equal or
2320unequal in size; petals reflexed or ascending, with
2321marginal appendages or with a basal, petaloid,
2322sericeous appendage; disk annular, 5–8-lobed;
2323stamens 5–8; pollen either colporate, striate, or
2324brevicolporate, psilate; ovary 3-carpellate, with a
2325single ovule per carpel; style subulate, crowned
2326by a capitate to cylindrical stigma. Fruit 1(–3)-
2327seeded, indehiscent, with leathery or less often
2328woody pericarp. Seeds entirely covered with a
2329sarcotesta.
398 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
2330 Fifty two species predominantly from South
2331 America, some occurring in Central America and
2332 southern Mexico.
2333 Genera 44–140: Incertae Sedis
2334 44.Alectryon Gaertn
2335 Alectryon Gaertn., Fruct. Sem. Pl. 1: 216, pl. 46 (1788);
2336 Reynolds, Austrobaileya 2: 332–338 (1987); Leenhouts,
2337 Blumea 33: 313–327 (1988), reg. rev.
2338 Falsely polygamous shrubs or trees. Leaves alter-
2339 nate, paripinnate or less often unifoliolate; leaf-
2340 lets serrate or entire; distal leaflet rudimentary;
2341lower pair of leaflets (pseudostipules) sometimes
2342clasping the stem. Inflorescences axillary thyrses
2343or panicles. Flowers actinomorphic; calyx aceta-
2344buliform, sepals 4–5(6), valvate or sub-imbricate;
2345petals 0 or 4–5, shortly clawed, the appendages
2346apparently of marginal origin, connate to form
2347a funnel-shaped structure with the petal; disk
2348annular-lobed, glabrous; stamens (5–)8, the
2349filaments subequal, inserted on the disk; pollen
2350colporate to parasyncolporate, striate to rugulate;
2351ovary (1)2–4(5)-carpellate, with a single ovule per
2352carpel; stigma grooved or lobed. Fruits capsular,
23531–3-coccate, lobed or unlobed, circumscissile or
2354dehiscent septifragally along septum. Seed with a
2355partial granular, red sarcotesta. 2n¼32.
2356About 25 species in eastern Malesia, Australia,
2357New Zealand, New Caledonia, and extending into
2358the Pacific to Samoa and the Sandwich Islands.
2359Two subgenera: Synalectryon and Alectryon.
236045.Amesiodendron Hu
2361Amesiodendron Hu, Bull. Fan. Mem. Inst. Biol. 7, Bot.: 207
2362(1936); Lo, Acta Phytotax. Sin. 17: 36, f. 3 (1979); Leenh-
2363outs in Fl. Males. I, 11: 465 (1994).
2364Polygamous monoecious trees. Leaves alternate,
2365once pinnate, paripinnate; leaflets serrate; distal
2366leaflet rudimentary. Inflorescences axillary or ter-
2367minal panicles. Flowers actinomorphic; sepals 5,
2368distinct, valvate; petals 5, with a single basal
2369appendage; disk annular, bowl-shaped, forming
2370a ring on upper portion; stamens (6)7–8(9); pollen
2371syncolporate, rugulate; ovary 3-carpellate, with a
2372single ovule per carpel; style with two stigmatic
2373lines. Fruits loculicidal capsules, 3-coccate,
2374sometimes 1 or 2 cocci rudimentary. Seeds with
2375sarcotestal ring around hilum.
2376One to three species, from Indochina, southern
2377China, peninsular Malaysia, and Sumatra.
237846.Aporrhiza Radlk.
2379Aporrhiza Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
2380Bayer Akad. Wiss. M€
unchen 8: 338 (1878).
2381Falsely polygamous trees or shrubs. Leaves alter-
2382nate, once pinnate, paripinnate; terminal leaflet
2383rudimentary. Inflorescences terminal, axillary or
2384cauliflorous thyrses. Flowers actinomorphic,
2385functionally unisexual; sepals 5, distinct, valvate;
2386petals 5, clawed, with a pair of appendages
2387formed by the inflexed margin above the claw;
Fig. 84. Sapindaceae. Talisia princeps Oliver. ADistal
portion of pinnate leaf and inflorescence. BCataphylls.
CFlower bud. DPistillate flower. EDitto, perianth
removed to show disk, sterile stamens and pistil. F
Stamen. GAbaxial and adaxial views of petal with adnate
appendage. HLongitudinal section of fruit showing
mesocarp and seed. IFruit and ventral view of embryo.
(Acevedo-Rodrı
´guez 2003; drawn by A. Tangerini)
Sapindaceae 399
Uncorrected Proof
2388 disk annular, sub-lobed; stamens 6–8; pollen
2389 colporate, perforate; ovary 2-carpellate, with a
2390 single ovule per carpel; style short; stigma
2391 2-lobed. Fruit a 2-locular, loculicidal capsule.
2392 Seeds with a sarcotesta on lower half. 2n¼28.
2393 Four to six species from tropical Africa.
2394 47.Arytera Blume Fig. 85
2395 Arytera Blume, Rumphia 3: 169 (1847); van der Ham,
2396 Blumea 23: 289–300 (1977); Turner, Blumea 38: 137–144
2397 (1993); van Bergen et al., Blumea 40: 195–209 (1995).
2398 Trees or shrubs. Leaves alternate, once pinnate,
2399 paripinnate; leaflets usually with domatia; distal
2400 leaflet rudimentary. Inflorescences axillary or
2401 pseudo-terminal thyrses. Flowers actinomorphic,
2402 functionally unisexual; sepals 5, connate to nearly
2403 distinct, valvate to imbricate; petals (2–)5(6), with
2404 2 marginal appendages, sometimes clawed; disk
2405 annular, lobed; stamens (5–)7–8(–10); pollen
2406colporate to parasyncolporate, rarely syncolpo-
2407rate, rugulate to striate-rugulate; ovary (1)2–3-
2408carpellate, with a single ovule per carpel; style
2409filiform, with 2–3 stigmatic lines or 2–3-lobed.
2410Fruit a 1–3-coccate, loculicidal capsule. Seeds
2411with basal arillode, apically open, covering half
2412to nearly the entire seed.
2413About 28 species in northeast India and
2414southeast Asia, throughout Malesia, and Australia,
2415Solomon Islands and Pacific Islands.
2416Section Arytera has simple hairs, sepals exter-
2417nally pubescent, and 2–3-lobed ovaries; Section
2418Azarytera has glandular-scaly indument, glabrous
2419sepals, and 2-lobed ovaries.
242048.Atalaya Span. ex Blume
2421Atalaya Span. ex Blume, Rumphia 3: 186 (1847); Radlk. in
2422Pflanzenreich 98: 605 (1932); Reynolds, Austrobaileya 1:
2423398–406 (1981), reg. rev.; Leenhouts in Fl. Males. I, 11: 479
2424(1994), reg. rev.
2425Falsely polygamous trees or shrubs. Leaves
2426alternate, once pinnate, paripinnate or less often
2427imparipinnate, up to 6-jugate; distal leaflet
2428rudimentary; rachis sometimes winged. Inflores-
2429cences terminal or axillary thyrses. Flowers uni-
2430sexual or bisexual; calyx slightly zygomorphic,
2431sepals 5, distinct, imbricate, the outer 2 smaller;
2432petals (4)5, clawed, with a pair of appendages
2433formed by the inflexed margins above the claw;
2434disk annular or semi-annular; stamens 8; pollen
2435colporate, rugulate; ovary 3-carpellate, with a
2436single ovule per carpel; placentation axile; style
2437conical with 3 stigmatic lines. Fruits schizocarpic,
2438splitting into (1–)3 divaricate, samaroid meri-
2439carps, each with a long, dorsal wing. Seed
2440exarillate, laterally compressed, with papery testa.
2441Twelve species, mostly from northern and
2442eastern Australia, also in the Lesser Sunda
2443Islands, southeastern New Guinea, and South
2444Africa.
244549.Beguea Capuron
2446Beguea Capuron, Mem. Mus. Natl. Hist. Nat., B, Bot. II,
244719: 105 (1969).
2448Dioecious trees. Leaves alternate, paripinnate;
2449leaflets 3–7 pairs, opposite or alternate, entire;
2450distal leaflet rudimentary; stipules 0. Inflore-
2451scences axillary racemes or thyrses. Flowers acti-
2452nomorphic, unisexual; sepals (5)6–7(8), distinct,
Fig. 85. Sapindaceae. Arytera multijuga.AFlowering
branch. BMale flower. CPetal. (Adema et al. 1994)
400 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
2453 valvate; petals 0; disk annular-lobed; stamens 6–8
2454 (–10); pollen colporate, striate; ovary 3-carpellate,
2455 with a single ovule per carpel; style filiform with 3
2456 stigmatic branches. Fruit indehiscent, crustose,
2457 1-seeded. Seeds arillate, cotyledons cerebriform
2458 (brain-like).
2459 A single species, B. apetala Capuron, endemic
2460 to Madagascar.
2461 50.Bizonula Pellegrin
2462 Bizonula Pellegrin, Bull. Soc. Bot. France 71: 299 (1924).
2463 Hermaphroditic trees. Leaves bipinnate; distal
2464 leaflet rudimentary; stipules 0. Inflorescences
2465 terminal thyrses. Flowers actinomorphic, bisexual;
2466 sepals 5, distinct, valvate; petals 5, with a basal
2467 appendage forming a pocket; disk annular, double;
2468 stamens 12–13; pollen colporate, striate; ovary
2469 3-carpellate, with a single ovule per carpel; style
2470 elongated, weakly lobed. Fruit not known.
2471 A single species, B. le-testui Pellegrin, from
2472 Gabon, Africa.
2473 51.Blighia Koenig
2474 Blighia Koenig, Ann. Bot. 2: 571 (1806).
2475 Falsely polygamous-dioecious, usually trees or
2476 shrubs. Leaves alternate, paripinnate; leaflets
2477 1–5 pairs, entire; distal leaflet rudimentary; sti-
2478 pules 0. Inflorescences axillary racemose thyrses.
2479 Flowers actinomorphic, functionally unisexual;
2480 sepals 5, valvate; petals 5, connate along append-
2481 age margins to form a pouch; disk annular,
2482 8-lobed; stamens 8–10; pollen colporate, striate;
2483 ovary 3(4)-carpellate, with a single ovule per car-
2484 pel; style elongated-conical. Fruit a 3(4)-locular,
2485 loculicidal, capsule. Seeds with a basal arillode.
2486 2n¼32.
2487 Three species from tropical Africa. Blighia
2488 sapida Koenig is widely cultivated for its edible
2489 arillodes in Jamaica and as an ornamental
2490 in Africa, the West Indies, and areas of the
2491 Neotropics.
2492 52.Blighiopsis van der Veken
2493 Blighiopsis van der Veken, Bull. Jard. Bot. E
´tat. 30: 413
2494 (1960).
2495 Dioecious trees. Leaves alternate, paripinnate;
2496 leaflets opposite or alternate, entire; distal leaflet
2497 rudimentary; pseudostipules small. Inflore-
2498scences thyrsoid. Flowers actinomorphic, uni-
2499sexual; sepals (4)5(7), slightly imbricate, distinct
2500to base; petals 0; disk annular; stamens 5(7),
2501anthers basifixed; pollen colporate, striate; ovary
25023-carpellate, with a single ovule per carpel; style
2503short, crowned by a trigonous stigma. Fruit a
2504tardily loculicidal, 1-locular, coriaceous capsule.
2505Seed solitary, with basal arillode.
2506A single species, B. pseudostipularis van der
2507Veken, Central Africa.
250853.Blomia Miranda.
2509Blomia Miranda, Annales Inst. Biol. Univ. Nac. Mexico 24:
251082 (1953).
2511Tikalia Lundell (1961).
2512Falsely polygamous-dioecious trees. Leaves alter-
2513nate, paripinnate; leaflets crenate, 1-4 pairs; distal
2514leaflet rudimentary; stipules 0. Inflorescences
2515axillary thyrses. Flowers actinomorphic, uni-
2516sexual or bisexual; sepals 5, distinct, valvate;
2517petals 0 or vestigial, with a pair of minute mar-
2518ginal appendages; disk annular-lobed; stamens
25195–6; pollen colporate, striate; ovary 1-carpellate,
2520with a single ovule per carpel; style short; stigma
2521capitate. Fruit a 1-locular, tardily loculicidally
2522dehiscent, coriaceous, red capsule. Seeds with
2523thin sarcotesta.
2524A single species, B. cupanioides Miranda, in
2525Mexico, Guatemala, and Belize.
252654.Camptolepis Radlk.
2527Camptolepis Radlk. in Engl. & Prantl., Nat. Pflanzenfam.,
2528Nachtr. 2–4, 3: 207 (1907); Capuron, Mem. Mus. Natl.
2529Hist. Nat. B, Bot. II, 19: 1–189 (1969).
2530Hypseloderma Radlk. (1932) [1933].
2531Dioecious trees. Leaves alternate, paripinnate;
2532leaflets entire; distal leaflet rudimentary; stipules
25330. Inflorescences ramiflorous, short thyrses.
2534Flowers actinomorphic, unisexual; sepals 5, dis-
2535tinct, imbricate; petals 5, with a single basal, short
2536appendage; disk annular-lobed; stamens (10–)12;
2537pollen colporate or brevicolporate, perforate;
2538ovary 3-carpellate, with a single ovule per carpel;
2539style elongated; stigma 3, ellipsoid. Fruit indehis-
2540cent, (1–)3-locular. Seeds completely covered by
2541a translucent arillode.
2542Four species from tropical east Africa and
2543Madagascar.
Sapindaceae 401
Uncorrected Proof
2544 55.Castanospora F. Muell.
2545 Castanospora F. M€
uller, Fragm. 9: 92 (1875); Reynolds,
2546 Austrobaileya 2: 34–35 (1984).
2547 Falsely polygamous trees. Leaves alternate, pari-
2548 pinnate; leaflets entire, alternate or subopposite;
2549 distal leaflet rudimentary; stipules 0. Inflore-
2550 scences axillary racemes or thyrses, with lateral
2551 dichasia. Flowers actinomorphic, functionally
2552 staminate or pistillate; calyx cup-shaped, sepals 5,
2553 ovate, sub-valvate; petals 5, with 2 minute basal
2554 appendages; disk annular-lobed; stamens 8;
2555 pollen parasyncolporate, striate-rugulate; ovary
2556 2-carpellate, with a single ovule per carpel;
2557 style short, with 2 reflexed stigmatic branches.
2558 Fruit indehiscent, crustose-fleshy, bicoccate,
2559 sometimes with a thin septa. Seeds large,
2560 chestnut-like, brown, exarillate.
2561 A single species, C. alphandi F. Muell., eastern
2562 Australia.
2563 56.Chouxia Capuron
2564 Chouxia Capuron, Mem. Mus. Natl. Hist. Nat., B, Bot. II,
2565 19: 130 (1969); Schatz et al., Adansonia III, 21: 51–62
2566 (1999), rev.
2567 Dioecious or monoecious trees. Leaves alternate,
2568 paripinnate; leaflets opposite or subopposite,
2569 entire; distal leaflet rudimentary; stipules 0.
2570 Inflorescences cauliflorous racemes or thyrses.
2571 Flowers actinomorphic, staminate or pistillate;
2572 sepals (4)5(6), distinct, imbricate; petals 5, with
2573 a simple basal appendage; disk annular; stamens
2574 (7)8(–10); pollen colporate, striate-reticulate;
2575 ovary (2)3-carpellate, with a single ovule per
2576 carpel; style elongated; stigma 3. Fruit 1–3-locular,
2577 indehiscent, baccate. Seeds arillate.
2578 Six species endemic to Madagascar.
2579 57.Chytranthus Hook. f.
2580 Chytranthus Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 403
2581 (1862).
2582 Glossolepis Gilg (1897).
2583 Falsely polygamous dioecious treelets. Leaves
2584 alternate, paripinnate, terminal leaflet rudimen-
2585 tary; stipules 0. Inflorescences cauliflorous, fasci-
2586 culate, racemose thyrses. Flowers zygomorphic,
2587 functionally staminate or pistillate; calyx urceolate,
2588sepals 5, connate at base, imbricate; petals 4(5–7),
2589with a basal, simple or hood-shaped and some-
2590times crested appendage; disk semi-annular or
2591reniform; stamens 7–8(9–11); pollen colporate
2592of brevicolporate, striate; ovary 3–8-carpellate,
2593with a single ovule per carpel; style subulate.
2594Fruit indehiscent, 3–8-locular, deeply ribbed,
2595fleshy. Seeds with sarcotesta. 2n¼32.
2596Twenty-six to 29 species from west tropical
2597Africa.
259858.Cnesmocarpon Adema
2599Cnesmocarpon Adema, Blumea 38: 195–201 (1993).
2600Trees. Leaves alternate, paripinnate; leaflets
2601papillate beneath; distal leaflet rudimentary.
2602Inflorescences axillary or ramiflorous thyrses.
2603Flowers actinomorphic; sepals 5, imbricate,
2604slightly unequal; petals 5, with 2 marginal appen-
2605dages, or less often lacking appendages; disk
2606annular or semi-annular; stamens 8; pollen
2607syncolporate or parasyncolporate, psilate or
2608indistinctly rugulate; ovary 3-carpellate, with a
2609single ovule per carpel, style with 3 stigmatic
2610lines. Fruit a 3-locular, fleshy, loculicidal capsule,
2611the locules basally winged, covered with irritating
2612hairs. Seeds obovoid, testa shiny black, with
2613carunculoid sarcotesta at the base.
2614Four species in Australia and Papua
2615New Guinea, occurring in primary lowland to
2616montane forest.
261759.Conchopetalum Radlk.
2618Conchopetalum Radlk. in T. Durand, Index Gen. Phan.: 81
2619(1887) [1888].
2620Shrubs or trees. Leaves alternate, paripinnate;
2621leaflets entire; distal leaflet rudimentary; stipules 0.
2622Inflorescences axillary or ramiflorous, fasciculate,
2623pseudo-umbelliform cymes. Flowers actinomor-
2624phic, functionally unisexual; sepals 5, connate at
2625base, imbricate; petals 5, red, without appen-
2626dages; disk annular, double, the central rim
2627sometimes tubular resembling an androgyno-
2628phore; stamens 8, exserted; pollen colporate,
2629striate-rugulate; ovary 3-carpellate, with 2 ovules
2630per carpel; style elongated, the stigma puncti-
2631form. Fruit a 3-locular, chartaceous capsule.
402 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
2632 Seeds with brown sarcotestal ring around the
2633 hilum.
2634 Two species endemic to Madagascar.
2635 60.Cubilia Blume Fig. 86
2636 Cubilia Blume, Rumphia 3: 100 (1847); Leenhouts,
2637 Blumea 24: 297 (1978).
2638 Falsely polygamous trees. Leaves alternate, pari-
2639 pinnate; terminal leaflet rudimentary. Inflores-
2640 cences terminal or pseudo-terminal, thyrsoid or
2641 corymbiform. Flowers actinomorphic; calyx urce-
2642 olate, with a narrow opening and 5 minute lobes;
2643 petals 5, shorter than the calyx, without appen-
2644 dages; disk annular, fleshy; stamens 5 (or 6); pollen
2645 small (1213 mm), colporate, scabrate; ovary
2646 2-carpellate, with a single ovule per carpel; stigma
2647 short, with 2 stigmatic lobes. Fruit a 1–2-coccate,
2648 loculicidal, echinate, woody capsule. Seeds with a
2649 large round hilum, arillate on lower half or third.
2650 A single species, C. cubili (Blanco) Adelb.,
2651 from eastern Borneo, the Philippines, Celebes,
2652 and the western Moluccas Islands.
265361.Cupania L.
2654Cupania L., Sp. Pl.: 200 (1753).
2655Duodichogamous or dioecious trees. Leaves
2656alternate, paripinnate or imparipinnate; leaflets
2657mostly serrate; stipules 0. Inflorescences axillary
2658or terminal thyrses or racemes. Flowers actino-
2659morphic, functionally unisexual; sepals 5, short,
2660imbricate; petals 5, with a pair of marginal
2661tomentose appendages; disk annular-lobed;
2662stamens (4, 6) 8; pollen syncolporate (Fig. 79J)
2663or parasyncolporate, rugulate; ovary 3-carpellate,
2664with a single ovule per carpel; style elongated;
2665stigma conical. Fruit a 2- or 3-locular, woody or
2666leathery, loculicidal capsule. Seeds with a cupular
2667arillode at base. 2n¼32.
2668About 50 species from tropical and subtropical
2669America.
267062.Cupaniopsis Radlk.
2671Cupaniopsis Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
2672Bayer. Akad. Wiss. M€
unch. 9: 483, 498, 584 (1879);
2673Adema, Leiden Bot. Series 15: 75 (1991), rev.
Fig. 86. Sapindaceae. Cubilia cubili.AMale flower. B
Longitudinal section of male flower. CAnthers, ventral
and dorsal views. DFemale flower. EPetal. FCapsule. G
Dehisced capsule showing seed with arillate base. H
Longitudinal section of seed. (Adema et al. 1994)
Sapindaceae 403
Uncorrected Proof
2674 Shrubs or treelets, pubescence of simple or
2675 glandular trichomes, or scaly. Leaves alternate,
2676 paripinnate; leaflets entire or less often dentate
2677 or serrate; distal leaflet rudimentary; stipules 0.
2678 Inflorescences axillary or cauliflorous thyrses. Flow-
2679 ers unisexual; calyx zygomorphic, sepals (4)5(–7),
2680 distinct, imbricate, the outer two distinctly
2681 smaller; petals (4)5, with 1 or 2 basal appendages;
2682 disk annular-lobed; stamens (5–)8–14; pollen
2683 syncolporate or parasyncolporate, sometimes
2684 colporate, rugulate, striate-reticulate, reticulate
2685 or perforate, sometimes verrucate; ovary (2)3
2686 (4)-carpellate, with a single ovule per carpel.
2687 Fruit a (1)2–3-locular, dehiscent, fleshy capsule.
2688 Seeds with basal arillode, covering half to
2689 nearly the entire seed, or exceptionally naked
2690 and sarcotestal.
2691 Sixty species from eastern Malesia, Caroline
2692 Islands, northern and eastern Australia, and
2693 from Solomon Islands to Samoa including New
2694 Caledonia.
2695 63.Deinbollia Schumach.
2696 Deinbollia Schumach., Beskr. Guin. Pl.: 242 (1827).
2697 Falsely polygamous or dioecious, shrubs or
2698 trees. Leaves alternate, paripinnate; leaflets
2699 entire; distal leaflet rudimentary; stipules 0.
2700 Inflorescences axillary or terminal thyrses. Flow-
2701 ers functionally unisexual; calyx zygomorphic,
2702 sepals 5, imbricate, the outer two smaller; petals
2703 5, with a basal, deeply bilobed appendage; disk
2704 annular, cup-shaped or vase-shaped; stamens
2705 (8–)12–30; pollen colporate, perforate; ovary
2706 (2)3(5)-carpellate, with a single ovule per carpel;
2707 style gynobasic or sub-terminal, filiform. Fruits
2708 of (1)2–3(–5) indehiscent, fleshy cocci. Seeds
2709 exarillate. 2n¼30.
2710 About 38 species from southern Africa,
2711 Madagascar, and Mascarene Islands.
2712 64.Delavaya Franch.
2713 Delavaya Franch., Bull. Soc. Bot. France 33: 462 (1886).
2714 Falsely polygamous shrubs or trees. Leaves alter-
2715 nate, trifoliolate; leaflets serrate-denticulate;
2716 stipules 0. Inflorescences terminal thyrses. Flow-
2717 ers functionally unisexual; calyx zygomorphic,
2718 sepals 5, imbricate, the outer two smaller; petals
2719 5, with a pair of marginal appendages; disk
2720pulvinate, nearly shortly tubular; stamens 8; pol-
2721len colporate, striate; ovary 2–3-carpellate, with
27222 ovules per carpel; style subulate. Fruits 2–3-
2723coccate, loculicidal capsules. Seeds exarillate.
2724A single species, D. yunnanensis Franch.,
2725China.
272665.Dictyoneura Blume
2727Dictyoneura Blume, Rumphia 3: 163 (1847); van Dijk,
2728Blumea 31: 437–449 (1986).
2729Falsely polygamous shrubs or trees. Leaves alter-
2730nate, paripinnate; leaflets serrate to lobed; distal
2731leaflet rudimentary; stipules 0. Inflorescences
2732axillary, spicate or racemose thyrses. Flowers
2733unisexual; sepals (5)6, imbricate, of same length
2734in actinomorphic calyx or the inner and/or the
2735outermost sepals smaller in zygomorphic calyx;
2736petals 0; disk annular; stamens (4)5(6); pollen
2737colporate, rugulate; ovary 2(3)-carpellate, with
2738a single ovule per carpel; style short, with 2(3)
2739stigmatic grooves. Fruit a 2(3)-locular, sub-
2740fleshy capsule, endocarp granular. Seeds with a
2741partial cupular, ventral or nearly complete sar-
2742cotesta.
2743Two or three species from eastern Borneo,
2744the Philippines, Celebes, Moluccas, and New
2745Guinea.
274666.Dilodendron Radlk.
2747Dilodendron Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
2748Bayer. Akad. Wiss. M€
unchen 8: 355 (1878); Gentry &
2749Steyermark, Ann. Missouri Bot. Gard. 74: 533–538
2750(1987), rev.
2751Falsely polygamous-dioecious trees. Leaves alter-
2752nate, bipinnate or sub-tripinnately compound;
2753distal leaflets on rachises fully developed or rudi-
2754mentary; secondary rachises marginate, alternate;
2755stipules 0. Inflorescences terminal thyrses. Flow-
2756ers 5-merous, functionally staminate or pistillate;
2757calyx slightly zygomorphic with one sepal larger
2758than the others, aestivation imbricate; petals
2759much shorter than sepals, with narrow, marginal
2760appendages; disk annular and flattened; stamens
27616–8(9); pollen colporate, striate; ovary 3-carpellate,
2762with a single ovule per carpel; style short with 3
2763stigmatic lobes. Capsule loculicidal, 2–3-locular,
2764woody. Seeds exarillate.
2765Three species from tropical continental
2766America.
404 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
2767 67.Dimocarpus Lour.
2768 Dimocarpus Lour., Fl. Coch.: 233 (1790); Leenhouts,
2769 Blumea 19: 113–131 (1971), rev.; van der Ham, Palyno-
2770 sciences 2: 239–254 (1993).
2771 Falsely polygamous trees or shrubs; indumen-
2772 tum of simple and stellate hairs. Leaves alter-
2773 nate, paripinnate, rarely unifoliolate; leaflets
2774 serrate or entire; distal leaflet rudimentary.
2775 Inflorescences terminal or less often axillary
2776 thyrses. Flowers actinomorphic, functionally
2777 unisexual; sepals 5–6, imbricate, connate at
2778 base or distinct; petals 5(6) or 0, without appen-
2779 dages; disk annular, 5-lobed, pubescent; stamens
2780 (6–)8(–10); pollen colporate, striate to perforate,
2781 sometimes scabrate or rugulate; ovary 2(3)-car-
2782 pellate, with a single ovule per carpel; style fili-
2783 form, with 2–3 stigmatic spreading lobes. Fruits
2784 of 1(2) indehiscent, or tardily dehiscent, warty,
2785 smooth or spiny mericarps. Seeds with a thin,
2786 translucent-white, fleshy arillode around the
2787 hilum. 2n¼30.
2788 Six species in southern and southeastern
2789 Asia from Sri Lanka and India to eastern Malesia
2790 and Australia. Dimocarpus longan Lour. is widely
2791 cultivated as the source of the tropical fruit
2792 Longan.
2793 68.Diploglottis Hook. f.
2794 Diploglottis Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 395
2795 (1862); Reynolds, Austrobaileya 1: 390 (1981); Reynolds,
2796 Austrobaileya 2: 328 (1987), reg. rev.; Leenhouts in Fl.
2797 Males. I, 11: 520 (1994), reg. rev.
2798 Falsely polygamous trees. Leaves alternate, pari-
2799 pinnate; distal leaflet rudimentary. Inflorescences
2800 axillary or pseudo-terminal thyrses. Flowers acti-
2801 nomorphic, functionally unisexual; sepals 5,
2802 imbricate; petals 4–5, clawed, with a pair of
2803 appendages formed by the inflexed margin
2804 above the claw, theses sometimes crested; disk
2805 annular or semi-annular, lobed; stamens 6–9;
2806 pollen parasyncolporate, rugulate or psilate;
2807 ovary 2–3-carpellate, with a single ovule per car-
2808 pel; style filiform, with 3 stigmatic grooves. Fruit
2809 a 2–3-locular, loculicidal capsule, coccate, deeply
2810 lobed or ovoid. Seeds with 2-lobed arillode cov-
2811 ering most of the seed.
2812 Twelve species from northeastern Australia
2813 and Papua New Guinea.
281469.Elattostachys (Blume) Radlk.
2815Elattostachys (Blume) Radlk., Actes Congr. Bot. Amster-
2816dam 1877: 101 (1879); Adema in Fl. Males. I, 11: 527
2817(1994).
2818Falsely polygamous trees or shrubs. Leaves alter-
2819nate, paripinnate; distal leaflet rudimentary.
2820Inflorescences (supra)axillary thyrses. Flowers
2821actinomorphic, bisexual or functionally unisexual;
2822sepals 5, valvate to slightly imbricate; petals 5,
2823clawed with 2 marginal appendages; disk annular,
2824dish- or cup-shaped; stamens 8; pollen parasyn-
2825colporate or colporate, striate to rugulate; ovary
28263-carpellate, with a single ovule per carpel; style
2827with 3 stigmatic lines. Fruit a 3-locular, woody,
2828loculicidal capsule. Seeds completely arillate or
2829only so at base.
2830About 20 species, from Malesia, Australia,
2831Solomon Islands, New Hebrides, New Caledonia,
2832Fiji, Samoa, and Tonga.
283370.Eriocoelum Hook. f.
2834Eriocoelum Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 400
2835(1862).
2836Monoecious or falsely polygamous trees or
2837shrubs. Leaves alternate, paripinnate; leaflets
28382–5 pairs; distal leaflet rudimentary; pseudosti-
2839pules usually present. Inflorescences axillary,
2840spicate, racemose or thyrsoid. Flowers actino-
2841morphic, functionally unisexual; sepals 5, dis-
2842tinct, valvate; petals 5, with a short, pubescent
2843ventral appendage; disk annular, cupular,
28448–10-lobed-crenate; stamens 8–10; pollen col-
2845porate, striate; ovary 3-carpellate, with a single
2846ovule per carpel; style filiform, with 3 stigmatic
2847lobes. Fruit a 3-locular, woody, loculicidal capsule,
2848sometimes with setaceous-hispid pubescence.
2849Seeds with an arillode at base.
2850About 10 species from tropical Africa.
285171.Erythrophysa E. Meyer ex Arnold
2852Erythrophysa E. Meyer ex Arnold, J. Bot. (Hooker) 3: 258
2853(1841) (as Erythrophila, corr. Sonder in Harvey & Sonder,
2854Fl. Cap. 1: 237 (1860).
2855Falsely polygamous shrubs or trees. Leaves alter-
2856nate, imparipinnate; distal leaflet fully developed.
2857Inflorescences axillary or terminal thyrses. Flowers
2858zygomorphic, bisexual or functionally unisexual;
2859sepals 5, imbricate; petals 4(5), with a pair or
Sapindaceae 405
Uncorrected Proof
2860 digitiform, simple or dissected appendages above
2861 the claw; disk semi-annular; stamens 8; pollen
2862 colporate, striate; ovary 3-carpellate, with 2 ovules
2863 per carpel; style elongated, filiform; stigma punc-
2864 tiform. Fruit a 1–3-locular, membranous,
2865 inflated, loculicidal capsule. Seeds globose, exar-
2866 illate, sparsely pubescent, the testa delineating the
2867 contour of cotyledons.
2868 Five species from South Africa and Madagascar.
2869 72.Erythrophysopsis Verdc.
2870 Erythrophysopsis Verdc., J. Linn. Soc. Bot. 58: 202 (1962).
2871 Polygamous shrubs or trees. Leaves alternate,
2872 imparipinnate; distal leaflet fully developed. In-
2873 florescences axillary or terminal thyrses. Flowers
2874 zygomorphic, functionally unisexual; sepals
2875 5, distinct, imbricate; petals 4, with a pair or
2876 digitiform, simple or dissected appendages
2877 above the claw; disk unilateral, 4-lobed; stamens
2878 8; pollen colporate, striate (Fig. 79A); ovary
2879 3-carpellate, with 2 ovules per carpel; style
2880 elongated, filiform; stigma punctiform. Fruits
2881 indehiscent, 1–3-locular, woody or crustose.
2882 Seeds 1–3 per fruit, sub-globose, exarillate, woolly
2883 pubescent, the testa delineating the contour of
2884 cotyledons.
2885 A single species, E. aesculina (Baill.) Verdc.,
2886 Madagascar.
2887 73.Glenniea Hook. f.
2888 Glenniea Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 404
2889 (1862); Leenhouts, Blumea 21: 91–103 (1973) and 22:
2890 411–414 (1975).
2891 Melanodiscus Radlk. (1887) [1888].
2892 Hedyachras Radlk. (1920).
2893 Falsely polygamous-dioecious or monoecious
2894 trees or shrubs; indumentum of simple or stellate
2895 hairs. Leaves alternate, unifoliolate or paripin-
2896 nate; leaflets entire; distal leaflet rudimentary;
2897 pseudostipules sometimes present. Inflores-
2898 cences terminal, thyrsoid. Flowers actinomor-
2899 phic, functionally unisexual; sepals (3)4–5,
2900 imbricate or valvate, distinct; petals 0; disk patel-
2901 liform or annular-lobed; stamens 4–8; pollen
2902 colporate, striate to striate-reticulate; ovary 2(3)-
2903 carpellate, with a single ovule per carpel; stigma
2904 lobed. Fruits indehiscent, crustose or baccate.
2905 Seed exarillate.
2906Eight species, 3 in tropical Africa, 1 in Mada-
2907gascar, 1 in Sri Lanka, and 3 in Indochina and
2908Malesia.
290974.Gloeocarpus Radlk.
2910Gloeocarpus Radlk., Philipp. J. Sci. 8, Bot.: 464 (1914);
2911Welzen, Blumea 35: 389 (1991), rev.
2912Trees; indument of glandular hairs. Leaves
2913alternate, paripinnate; leaflets serrate; distal
2914leaflet rudimentary. Inflorescences ramiflorous
2915thyrses. Flowers functionally unisexual; calyx
2916zygomorphic, the sepals 5, distinct, imbricate,
2917outer 2 smaller; petals 5, with 2 marginal appen-
2918dages; disk annular; stamens 7; pollen syncolpo-
2919rate or parasyncolporate, rugulate; ovary
29203-carpellate, with a single ovule per carpel; style
2921short, filiform; stigma not lobed. Fruit a 3-locular,
2922woody, deeply lobed, loculicidal capsule. Seeds
2923completely covered with a thin arillode.
2924A single species, G. patentivalvis (Radlk.)
2925Radlk., endemic to the Philippines.
292675.Gongrodiscus Radlk.
2927Gongrodiscus Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
2928Bayer. Akad. Wiss. M€
unchen 9: 503, 607 (1879); Turner &
2929van der Ham, Bull. Mus. Natl. Hist. Nat. Paris IV, 18:
2930339–349 (1996).
2931Falsely polygamous shrubs or trees. Leaves alter-
2932nate, paripinnate; distal leaflet rudimentary.
2933Inflorescences axillary, thyrsoid. Flowers actino-
2934morphic, bisexual or functionally unisexual;
2935sepals 5, distinct, valvate; petals 5, with 2 marginal
2936appendages; disk annular, 5-lobed; stamens (7)
29378; pollen parasyncolporate, rugulate; ovary
29383-carpellate, with a single ovule per carpel;
2939style short; stigma sub-trilobed. Fruit a 1-locular
2940(incomplete septa), fleshy, tardily dehiscent
2941capsule. Seeds arillate along ventral portion.
2942Three species endemic to New Caledonia.
294376.Gongrospermum Radlk.
2944Gongrospermum Radlk., Philipp. J. Sci. 8, Bot.: 469 (1914);
2945Welzen, Rheedea 1: 60 (1991), rev.
2946Trees. Leaves alternate, paripinnate; leaflets
2947papillate on lower surface; distal leaflet rudimen-
2948tary. Inflorescences axillary, simple or thyrses.
2949Flowers actinomorphic, functionally unisexual;
406 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
2950 sepals 5, valvate, distinct; petals 0; disk sub-cup-
2951 ular, 5-lobed; stamens 8; pollen syncolporate,
2952 rugulate; ovary 3-carpellate, with a single ovule
2953 per carpel; style pyramidal. Fruit a 1-locular,
2954 leathery, loculicidal capsule. Seeds exarillate,
2955 endotesta ruminately grown together with
2956 embryo.
2957 A single species, G. philippinense Radlk.,
2958 Philippines.
2959 77.Guioa Cav.
2960 Guioa Cav., Icon. 4: 49, t. 373 (1798); Welzen, Leiden Bot.
2961 Ser. 12: 146 (1989), rev.
2962 Shrubs, trees, or treelets. Leaves alternate, pari-
2963 pinnate; leaflets entire or less often crenate or
2964 serrate; distal leaflet rudimentary; rachis terete
2965 or winged. Inflorescences axillary or terminal
2966 thyrses. Flowers functionally unisexual; calyx
2967 zygomorphic, sepals 5(6), imbricate, petaloid,
2968 outer 2 smaller; petals 5(6), usually clawed, with
2969 two marginal or ventral appendages that are usually
2970 crested; disk annular or semi-annular; stamens
2971 (7)8; pollen syncolporate or parasyncolporate,
2972 rarely colporate, rugulate to perforate or psilate;
2973 ovary 3-carpellate, with a single ovule per carpel;
2974 style apical, pyramidal, with 3 stigmatic lines.
2975 Fruit a 3-locular, obcordate, deeply 3-lobed to
2976 coccate, leathery, loculicidal capsule. Seeds
2977 almost entirely covered by an arillode that has a
2978 basal projection.
2979 About 64 species ranging from southeastern
2980 Asia, throughout Malesia to Australia, New
2981 Caledonia, and Samoa.
2982 78.Haplocoelopsis Davis
2983 Haplocoelopsis Davis, Kew Bull. 52: 231 (1997).
2984 Monoecious shrubs or small trees. Leaves
2985 alternate, paripinnate or imparipinnate; leaflets
2986 entire; distal leaflet rudimentary; pseudosti-
2987 pules present. Inflorescences axillary, racemose
2988 thyrses. Flowers actinomorphic, unisexual; sepals 5,
2989 distinct, imbricate; petals 5, with a short, bilobed
2990 ventral appendage; disk annular; stamens 8–9;
2991 pollen unknown; ovary bilobed, 2-carpellate,
2992 with a single ovule per carpel; style with 2 stig-
2993 matic lobes. Fruit a 2-locular, circular, laterally
2994 compressed, loculicidal capsule. Seeds not
2995 known.
2996A single species, H. africana Davies, from
2997central and east Africa.
299879.Haplocoelum Radlk.
2999Haplocoelum Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3000Bayer. Akad. Wiss. M€
unchen 8: 336 (1878).
3001Falsely polygamous-dioecious trees. Leaves alter-
3002nate, paripinnate; distal leaflet rudimentary.
3003Inflorescence a congested axillary polychasium.
3004Flowers actinomorphic, functionally unisexual;
3005sepals 5–6, distinct, narrowly imbricate; petals 0 or
3006less often present; disk annular; stamens 5–6,
3007inserted on the disk; pollen colporate, rugulate;
3008ovary (2)3-carpellate, with a single ovule percar-
3009pel; style with 3 stigmatic lobes. Fruit 1–2-locular,
3010indehiscent, baccate. Seeds ellipsoid or laterally
3011compressed, with a dorsally or distally split aril-
3012lode.
3013About 7 species from tropical Africa and
3014Madagascar.
301580.Hornea Baker
3016Hornea Baker, Fl. Mauritius: 59 (1877).
3017Falsely polygamous shrubs or trees. Leaves alter-
3018nate, paripinnate; leaflets 2 or 4, entire; distal
3019leaflet rudimentary. Inflorescences terminal,
3020corymbose or thyrsoid. Flowers bisexual or
3021functionally unisexual; calyx zygomorphic, sepals
30225, orbicular, concave, distinct, imbricate, the
30232 outer sepals smaller; petals 5, clawed, with 2 ven-
3024tral appendages above the claw; disk 5-lobed;
3025stamens 18–24; pollen colporate, rugulate; ovary
30262-carpellate, with a single ovule per carpel; style
3027short; stigma punctiform. Fruits of 2 mericarps
3028with a dorsal wing. Seeds exarillate.
3029A single species, H. mauritiana Baker,
3030endemic to Mauritius.
303181.Jagera Blume
3032Jagera Blume, Rumphia 3: 155 (1847); Leenhouts in Fl.
3033Males. I, 11: 614 (1994).
3034Falsely polygamous, often pachycaulous trees or
3035shrubs. Leaves verticillate, sometimes opposite or
3036spirally arranged, paripinnate; leaflets serrate;
3037distal leaflet rudimentary. Inflorescences axillary
3038thyrses. Flowers actinomorphic, bisexual or func-
3039tionally unisexual; sepals 5, slightly connate at
3040base, imbricate; petals 5, with marginal or ventral
3041appendages; disk annular; stamens (7)8(–10);
Sapindaceae 407
Uncorrected Proof
3042 pollen syncolporate or parasyncolporate, per-
3043 forate or reticulate with often finely tapering
3044 scabrae; ovary 3-carpellate, with a single ovule
3045 per carpel; style filiform with 3 stigmatic lines.
3046 Fruit a 3-locular, woody, loculicidal capsule, with
3047 stiff, irritating hairs. Seeds with a small sarcotesta
3048 around the hilum.
3049 Two species in the Moluccas, New Guinea,
3050 and eastern Australia.
3051 82.Koelreuteria Laxm.
3052 Koelreuteria Laxm., Nov. Comm. Acad. Petrop. 16: 562, t.
3053 18 (1772); Meyer, J. Arnold Arb. 57: 129–166 (1976);
3054 Adema in Fl. Males. I, 11: 755 (1994).
3055 Duodichogamous trees. Leaves alternate, once
3056 or twice pinnate, imparipinnate; leaflets entire,
3057 serrate or crenate; terminal leaflet well-devel-
3058 oped. Inflorescences terminal thyrses. Flowers
3059 zygomorphic; sepals 5, valvate; petals 4, yellow,
3060 clawed, with fimbriate involute base of lamina
3061 forming an appendage; disk annular, undulate,
3062 on a short stipe; stamens (5–)8; pollen colporate,
3063 striate; ovary 3-carpellate, with 2 ovules per
3064 carpel, the septa incomplete on distal portion;
3065 stigma entire or trifid. Fruit a 3-locular, papery,
3066 inflated, loculicidal capsule with incomplete
3067 septa; seeds 2 per locule, exarillate. 2n¼22, 30, 32.
3068 About four species native to Japan, southern
3069 China, Taiwan, and perhaps indigenous to Fiji.
3070 Koelreuteria paniculata Laxm. and K. bipinnata
3071 Franchet are cultivated worldwide in temperate
3072 areas as ornamentals.
3073 83.Laccodiscus Radlk.
3074 Laccodiscus Radlk., Sitzungsber. Math-Phys. Cl. K€
onigl.
3075 Bayer. Akad. Wiss. M€
unchen 9: 496 (1879).
3076 Falsely polygamous trees or scandent shrubs.
3077 Leaves alternate, paripinnate; distal leaflet rudi-
3078 mentary; pseudostipules present. Inflorescences
3079 axillary or terminal thyrses. Flowers actinomor-
3080 phic, bisexual or functionally unisexual; sepals 5,
3081 connate at base, imbricate; petals 5, with short
3082 marginal appendages; disk annular-lobed or
3083 5-lobed; stamens 8–10; pollen colporate, per-
3084 forate; ovary 3-carpellate, with a single ovule per
3085 carpel; style curved; stigma papillose. Fruit a
3086 3-locular, woody, 3-lobed, loculicidal capsule.
3087 Seeds arillate (fide Fouilloy and Halle
´1973a).
3088 About six species native to West Africa.
308984.Lecaniodiscus Planch. ex Benth.
3090Lecaniodiscus Planch. ex Benth. in Hook., Niger Fl.: 250
3091(1849).
3092Chiarinia Chiov. (1932).
3093Falsely polygamous-dioecious shrubs or trees.
3094Leaves alternate, paripinnate; leaflets 3–7 pairs;
3095distal leaflet rudimentary; stipules 0. Inflore-
3096scences axillary, racemose or thyrsoid. Flowers
3097actinomorphic, functionally unisexual; sepals 5,
3098imbricate, distinct; petals 0 or 5, with 2 minute
3099basal appendages; disk annular, lobed; stamens 8;
3100pollen colporate, striate; ovary (2)3-carpellate,
3101with a single ovule per carpel; style short, stigma
3102subsessile and 3-lobed or with 2 reflexed stig-
3103matic branches. Fruits crustose, 1-locular or
3104bicoccate, indehiscent or tardily splitting from
3105the base. Seeds nearly entirely covered by an
3106arillode.
3107Two species native to tropical Africa.
310885.Lepiderema Radlk.
3109Lepiderema Radlk., Actes Congr. Bot. Amsterdam 1877:
3110250 (1879); Reynolds, Austrobaileya 1: 488 (1982), reg.
3111rev.; Schot, Blumea 36: 235 (1991), reg. rev.
3112Falsely polygamous shrubs or trees; indument
3113of lepidote scales. Leaves alternate, paripin-
3114nate; distal leaflet rudimentary. Inflorescences
3115axillary, ramiflorous, or pseudo-terminal race-
3116mose thyrses. Flowers actinomorphic, bisexual
3117or functionally unisexual; sepals 5, petaloid,
3118distinct, imbricate; petals 5, shortly clawed, with-
3119out appendages; disk annular; stamens (6–)8;
3120pollen syncolporate, parasyncolporate or colpo-
3121rate, rugulate-reticulate; ovary 3-carpellate, with
3122a single ovule per carpel; style spirally twisted;
3123stigma lobed. Fruit a 3-locular, trigonous, stipi-
3124tate, woody, loculicidal capsule. Seed basally
3125arillate.
3126Eight species, 6 in Australia and 2 in New
3127Guinea.
312886.Lepidopetalum Blume
3129Lepidopetalum Blume, Rumphia 3: 171 (1847); Welzen,
3130Piskaut & Windadri, Blumea 36: 452 (1992), rev.
3131Falsely polygamous trees. Leaves alternate, pari-
3132pinnate; distal leaflet rudimentary. Inflorescences
3133ramiflorous, axillary or pseudo-terminal thyrses.
3134Flowers actinomorphic, functionally unisexual;
408 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
3135 calyx shortly cupular, sepals 5(6), distinct, val-
3136 vate; petals 5(–7), shorter than the sepals, with a
3137 single basal appendage bigger than the blade; disk
3138 annular or semi-annular; stamens (7)8(–10), the
3139 filaments nearly equal, the anthers dorsifixed;
3140 pollen colporate, striate to striate-reticulate;
3141 ovary 2-carpellate, with a single ovule per carpel;
3142 stigma sessile, fleshy, of two elongate lobes adnate
3143 to the distal portion of ovary. Fruit a 2-locular,
3144 leathery or woody, glabrous, loculicidal capsule.
3145 Seed with a basal to complete, orange sarcotesta.
3146 Six species throughout Malesia and north-
3147 eastern Australia.
3148 87.Lepisanthes Blume
3149 Lepisanthes Blume, Bijdr. Fl. Nederl. Ind. 5: 237 (1825);
3150 Leenhouts, Blumea 17: 33–91 (1969).
3151 Aphania Blume (1825).
3152 Erioglossum Blume (1825).
3153 Otophora Blume (1847).
3154 Hebecoccus Radlk. (Radlk. (1878).
3155 Thraulococcus Radlk. (1878).
3156 Aphanococcus Radlk. (1887) [1888].
3157 Manongarivea Choux. (1927).
3158 Sapindopsis How & Ho (1955), non Fontaine (1889).
3159 Howethoa Rauschert (1982).
3160 Monoecious trees, shrubs, or climbing shrubs.
3161 Leaves alternate, (im)paripinnate, sometimes
3162 unifoliolate; terminal leaflet rudimentary; pseu-
3163 dostipules sometimes present. Inflorescences ter-
3164 minal, axillary, ramiflorous or cauliflorous thyrses.
3165 Flowers bisexual or functionally unisexual; calyx
3166 zygomorphic, the sepals 5, imbricate, the outer
3167 2 distinctly smaller; petals (2–)4(5), with mar-
3168 ginal or ventral appendages; disk semi-annular
3169 or annular, 5-lobed or crenate; stamens (4–)8
3170 (–18); pollen colporate or brevicolporate, rarely
3171 syncolporate, rugulate to reticulate, rarely psilate;
3172 ovary 2–3(4)-carpellate, sessile or stipitate, with a
3173 single ovule per carpel; style short, apical; stigma
3174 capitate, sometimes sessile. Fruits indehiscent,
3175 2–3-lobed, sub-fleshy, sometimes deeply lobed
3176 or coccate with distinct monocarps. Seeds ellip-
3177 soid, obovoid to sub-globose, exarillate. 2n¼26,
3178 28, 30.
3179 About 24 species in tropical Africa, Madagascar,
3180 southern and southeastern Asia from Sri Lanka to
3181 Hainan,Malesia,andnorthwesternAustralia.
3182 Four subgenera: subgen. Lepisanthes: Leaves
3183 paripinnate, without pseudostipules; petiole and
3184rachis not winged; outer sepals sericeous outside;
3185petals longer than sepals; fruits usually only
3186slightly lobed, septa continuous; subgen. Oto-
3187phora: leaves pari- or imparipinnate, without
3188pseudostipules; petiole winged or not; outer
3189sepals glabrous or hairy outside; petals shorter
3190than the sepals; fruits not or slightly lobed, septa
3191often interrupted; subgen. Erioglossum: Leaves
3192paripinnate, without pseudostipules; petiole and
3193rachis not winged; outer sepals glabrous outside;
3194petals longer than the sepals; fruits lobed, septa
3195complete; subgen. Aphania: Leaves paripinnate,
3196sometimes simple, sometimes with pseudosti-
3197pules (petiole and rachis winged); outer sepals
3198glabrous; petals as long as sepals; fruits lobed,
3199septa complete.
320088.Litchi Sonn
3201Litchi Sonn., Voy. Ind. Or. Chine 2: 230, t. 129 (1782);
3202Leenhouts, Blumea 24: 398 (1978).
3203Euphoria Commers. ex Juss. (1789).
3204Duodichogamous trees; indument of 2-branched
3205hairs. Leaves alternate, paripinnate; distal leaflet
3206rudimentary. Inflorescences terminal and axillary
3207thyrses. Flowers actinomorphic bisexual or func-
3208tionally unisexual; calyx cup-shaped, with 4–5
3209equal, minute lobes; petals 0; disk annular;
3210stamens (6)7(–11); pollen colporate, striate;
3211ovary 2-carpellate, with a single ovule per carpel;
3212stigma of 2 elongated, spreading or coiled lobes.
3213Fruit 1-coccate (1 coccus rudimentary), indehis-
3214cent, with leathery, muricate pericarp. Seeds
3215partly or completely covered by a translucent,
3216fleshy, convolute arillode. 2n¼28, 30.
3217A single species, L. chinensis Sonn., from
3218southeastern China, Indochina, Malay Peninsula,
3219Java, Borneo, and the Philippines. Widely
3220cultivated in subtropical regions for its edible
3221fruits, commonly known as Litchi.
322289.Lychnodiscus Radlk.
3223Lychnodiscus Radlk., Sitzungsber. Math-Phys. Cl. K€
onigl.
3224Bayer. Akad. Wiss. M€
unchen 8: 332 (1878).
3225Falsely polygamous shrubs or trees. Leaves
3226alternate, paripinnate; distal leaflet rudimentary.
3227Inflorescences axillary or terminal thyrses or pa-
3228nicles. Flowers actinomorphic, bisexual or func-
3229tionally unis exual; sepals 5, connate at base,
3230imbricate; petals 5, with a single ventral
Sapindaceae 409
Uncorrected Proof
3231 appendage; disk annular; stamens 10–12; pollen
3232 colporate, striate; ovary 3-carpellate, with a
3233 single ovule per carpel; stigma clavate. Fruit a
3234 (1–)3-locular, 3-lobed, loculicidal capsule; seeds
3235 sarcotestal.
3236 About seven species from tropical Africa.
3237 90.Macphersonia Blume
3238 Macphersonia Blume, Rumphia 3: 156 (1847).
3239 Dioecious trees. Leaves alternate, bipinnate, or
3240 once pinnate; distal leaflet rudimentary. Inflores-
3241 cences axillary, racemose or spicate, or rarely
3242 thyrsoid. Flowers actinomorphic, bisexual or
3243 functionally unisexual; sepals 5, distinct, imbri-
3244 cate; petals 5, clawed, with appendages formed by
3245 the inflexed margins above the claw; disk annular,
3246 dish-shaped; stamens 8; pollen colporate, striate;
3247 ovary (2)3-carpellate, with a single ovule per car-
3248 pel; stigma sessile, capitate or trigonous. Fruit
3249 1–2-locular, indehiscent or tardily dehiscent,
3250 sub-fleshy. Seeds completely covered by a trans-
3251 lucent arillode.
3252 About eight species from Aldabra, Madagascar,
3253 and west tropical Africa.
3254 91.Matayba Aublet Fig. 79K
3255 Matayba Aublet, Hist. Pl. Guiane 1: 331 (1775).
3256 Falsely polygamous-dioecious, large or small
3257 trees. Leaves alternate, paripinnate or imparipin-
3258 nate; leaflets entire; distal leaflet rudimentary.
3259 Inflorescences axillary or terminal thyrses. Flow-
3260 ers 5-merous, actinomorphic, bisexual or unisex-
3261 ual; sepals short (less than 2 mm long), distinct,
3262 valvate; petals as long as the sepals or longer, with
3263 a pair of marginal tomentose appendages; disk
3264 annular, usually lobed; stamens (4–6)8; pollen
3265 syncolporate or parasyncolporate, sometimes
3266 colporate, rugulate; ovary 3-carpellate, with a sin-
3267 gle ovule per carpel; stigma trilobed or trifid.
3268 Fruit a 2- or 3-locular, trigonous or lobed,
3269 woody or leathery, loculicidal capsule. Seeds
3270 nearly globose or ellipsoid, arillate at base or
3271 seldom nearly entire.
3272 About 50 species from tropical and sub-
3273 tropical America.
3274 92.Mischarytera H. Turner
3275 Mischarytera H. Turner, Blumea Suppl. 9: 210 (1995), rev.
3276Trees. Leaves paripinnate; distal leaflet rudimen-
3277tary. Inflorescences axillary to pseudo-terminal
3278thyrses. Flowers actinomorphic, functionally uni-
3279sexual; sepals 5, connate at base, valvate; petals 5,
3280clawed, with appendages formed by the inflexed
3281margins above the claw, or these 0; disk annular,
3282unlobed or lobed; stamens 7–8; pollen parasyn-
3283colporate, sometimes colporate, rugulate; ovary
32843-carpellate, with a single ovule per carpel; stigma
3285shortly 3-lobed. Fruit a 3-locular, loculicidal or
3286loculifragal capsule. Seeds arillate.
3287Three species in eastern Australia and Papua
3288New Guinea.
328993.Mischocarpus Blume, nom. cons.
3290Mischocarpus Blume, Bijdr. 1825: 238 (1825); van der
3291Ham, Blumea 23: 251 (1977).
3292Pedicellia Loureiro (1790), nom. rej.
3293Mischocodon Radlk. (1913).
3294Falsely polygamous shrubs or trees. Leaves
3295alternate, paripinnate; distal leaflet rudimentary.
3296Inflorescences pseudo-terminal, axillary or rami-
3297florous thyrsoids. Flowers actinomorphic, bisex-
3298ual or functionally unisexual; sepals imbricate,
3299distinct or connate into a crateriform calyx,
3300crowned by 5 subequal lobes; petals 0, reduced
3301or slightly longer than the sepals, with marginal
3302appendages; disk annular or semi-annular;
3303stamens (5–)8(9); pollen syncolporate or para-
3304syncolporate, sometimes colporate, rugulate to
3305rugulate-reticulate; ovary (2)3(4)-carpellate, with
3306a single ovule per carpel; stigma 3-lobed. Fruit a
3307(1)2-locular, coriaceous, loculicidal capsule.
3308Seeds completely covered by an arillode with a
3309basal, funiculus-like appendage.
3310About 15 species from southeastern Asia,
3311throughout Malesia to Australia.
331294.Molinaea Commers. ex Juss.
3313Molinaea Commers. ex Juss., Gen. Pl.: 248 (1789).
3314Falsely polygamous shrubs or trees. Leaves alter-
3315nate, paripinnate; leaflets entire or serrate, distal
3316leaflet rudimentary. Inflorescences axillary,
3317simple or paniculate thyrses. Flowers bisexual or
3318functionally unisexual; calyx zygomorphic, sepals
33195, distinct, imbricate, the outer two smaller; petals
33205, with marginal appendages or without appen-
3321dages; disk annular or semi-annular; stamens 8;
3322pollen parasyncolporate, rarely syncolporate
410 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
3323 or colporate, rugulate; ovary 3-carpellate, with
3324 a single ovule per carpel; style short; stigma
3325 3-gonous, grooved. Fruit a 3-locular, loculicidal
3326 capsule. Seeds arillate.
3327 About ten species from Madagascar,
3328 Mauritius, and Mascarene Islands.
3329 95.Namataea D.W. Thomas & D.J. Harris
3330 Namataea D.W. Thomas & D.J. Harris, Kew Bull. 54: 951
3331 (1999).
3332 Seemingly dioecious shrub. Leaves alternate, sim-
3333 ple; stipules 0. Inflorescences cauliflorous, rami-
3334 florous or less often axillary, racemes. Flowers
3335 unisexual; calyx zygomorphic, urceolate, the
3336 sepals 5, connate at lower quarter or third,
3337 slightly imbricate, third and fifth sepals slightly
3338 longer than remaining sepals; petals 4, a basal
3339 hood-shaped, crested appendage (the crest infun-
3340 dibuliform); disk reniform; stamens 7; pollen not
3341 known; ovary 3-carpellate, with a single ovule per
3342 carpel; style stout. Fruit fleshy, indehiscent, of 3
3343 ellipsoid, cocci, each with an apical beak.
3344 A single species, N. simplicifolia D.W.
3345 Thomas & D.J. Harris, Cameroon.
3346 96.Neotina Capuron
3347 Neotina Capuron, Mem. Mus. Natl. Hist. Nat. B, Bot., II,
3348 19: 174 (1969).
3349 Monoecious or dioecious trees. Leaves alternate
3350 or subopposite, paripinnate; distal leaflet rudi-
3351 mentary. Inflorescences axillary thyrses. Flowers
3352 unisexual; calyx zygomorphic, the sepals 5, im-
3353 bricate, outer two sepals smaller; petals 4–5, with
3354 2 marginal or ventral appendages; disk annular;
3355 stamens 5(6–8); pollen parasyncolporate or col-
3356 porate, rugulate to rugulate-reticulate; ovary
3357 2-carpellate, with a single ovule per carpel; style
3358 elongated, with 2 stigmatic lines. Fruit a 1-locular,
3359 sub-fleshy, loculicidal capsule. Seeds with red or
3360 orange arillode for 2/3 of their length.
3361 Two species from Madagascar.
3362 97.Nephelium L.
3363 Nephelium L., Syst. Nat. ed. 12, 2: 623 (1767); L., Mantissa
3364 Pl.: 18 (1767); Leenhouts, Blumea 31: 373–436 (1986).
3365 Falsely polygamous or dioecious trees or less
3366 often shrubs. Leaves alternate, paripinnate; leaf-
3367lets distinctly glaucous beneath; distal leaflet
3368rudimentary. Inflorescences axillary, pseudo-
3369terminal or terminal (in N. cuspidatum Blume
3370also rami- and cauliflorous) thyrses. Flowers
3371actinomorphic; calyx cup-shaped, crowned by
33724–6 subequal, valvate lobes; petals 0 or 4–6,
3373clawed, with a bilobed appendage; disk annular;
3374stamens 4–10; pollen colporate, striate; ovary (1)2
3375(–4)-carpellate, with a single ovule per carpel;
3376style elongated; stigma usually 2-lobed. Fruits
33771(2)-coccate, tardily loculicidally dehiscent, the
3378pericarp warty to spiny, coriaceous or less often
3379woody or corky. Seeds completely covered by
3380edible sarcotesta. 2n¼22.
3381About 16 species from southeastern Asia in
3382Yunnan and Assam to Hainan and Malesia.
3383Nephelium lappaceum L. (the Rambutan) and
3384N. ramboutan-ake (Labill.) Leenh. (the Pulasan)
3385are widely cultivated for their edible fruits.
338698.Otonephelium Radlk.
3387Otonephelium Radlk, Sitzungsber. Math-Phys. Cl. K€
onigl.
3388Bayer. Akad. Wiss. M€
unchen 20: 253, 288 (1890).
3389Falsely polygamous trees. Leaves alternate, pari-
3390pinnate; distal leaflet rudimentary; pseudosti-
3391pules present. Inflorescences terminal or axillary
3392thyrses. Flowers actinomorphic bisexual or func-
3393tionally unisexual; sepals 5, imbricate, distinct;
3394petals 0; disk annular, glabrous; stamens 5–9;
3395pollen colporate, striate, sometimes irregularly
3396striate or rugulate; ovary 2-carpellate, with a
3397single ovule per carpel; stigma bifid. Fruit
33981-coccate, with rudimentary cocci at base,
3399indehiscent, baccate-crustose, with soft spines.
3400Seeds arillate.
3401A single species, O. stipulaceum Radlk.,
3402western India. Doubtfully different from Dimo-
3403carpus by its glabrous disk and the presence of
3404pseudostipules.
340599.Pancovia Willd., nom. cons.
3406Pancovia Willd., Sp. Pl. 2: 285 (1799); non Fabricius
3407(1759), nom. rej.
3408Falsely polygamous or dioecious, trees or shrubs.
3409Leaves alternate, paripinnate; leaflets 2–12 pairs;
3410distal leaflet rudimentary. Inflorescences axillary
3411or cauliflorous, fasciculate, racemose or thyrsoid.
3412Flowers zygomorphic, functionally unisexual;
3413sepals 4–5, imbricate or sub-valvate; petals 3–4,
Sapindaceae 411
Uncorrected Proof
3414 clawed, with 2 inflexed or dissected appendages
3415 above the claw; disk semi-annular, unilateral;
3416 stamens (6–)8; pollen colporate, rugulate; ovary
3417 3-carpellate, with a single ovule per carpel; style
3418 subulate; stigma sub-clavate. Fruit indehiscent,
3419 3-locular, with fleshy to woody pericarp. Seeds
3420 exarillate, laterally compressed. 2n¼32.
3421 Ten to 12 species native to west tropical and
3422 subtropical Africa.
3423 100.Pappea Eckl. & Zeyh.
3424 Pappea Eckl. & Zeyh., Enum. Pl. Afr. austr. extratrop. 1: 53
3425 (1834–1835).
3426 Dioecious trees or shrubs. Leaves alternate,
3427 simple. Inflorescences axillary, racemose or
3428 thyrsoid. Flowers actinomorphic, functionally
3429 unisexual; sepals 5, valvate; petals (4)5(6), with
3430 a pair of marginal, hairy appendages; disk annu-
3431 lar; stamens 8(–10); pollen colporate, striate;
3432 ovary 3-carpellate, with a single ovule per carpel;
3433 style short; stigma sub-lobed. Fruit a 1-locular,
3434 fleshy, loculicidal capsule. Seeds with a lobed
3435 arillode.
3436 One to four species native to southern Africa.
3437 The fruit of P. capensis (Spreng.) Eckl. & Zeyh. is
3438 said to be edible, a bland oil is expressed from the
3439 seeds (Harvey and Sonder 1894).
3440 101.Paranephelium Miq
3441 Paranephelium Miq., Fl. Ind. Bat. Suppl. 509 (1861)
3442 [1860]; Davids, Blumea 29: 425 (1984), rev.
3443 Mildea Miquel (1867), non Griseb. (1866).
3444 Scyphopetalum Hiern (1875).
3445 Falsely polygamous trees. Leaves alternate,
3446 mainly imparipinnate; distal leaflets rudimentary
3447 or well-developed. Inflorescences ramiflorous or
3448 terminal thyrses. Flowers actinomorphic; calyx
3449 shortly cupular, the sepals (4)5(–7), mainly
3450 equal, distinct or connate at base, valvate; petals
3451 (4)5(–7), often clawed, with a single appendage;
3452 disk annular, 5-lobed, cup-shaped; stamens 5–9;
3453 pollen syncolporate or parasyncolporate, rugu-
3454 late; ovary mainly 3-carpellate; with a single
3455 ovule per carpel; stigma usually lobed. Fruits 1
3456 (–3)-locular, globular, smooth to densely spiny,
3457 woody, loculicidal capsules or dehiscing ran-
3458 domly. Seeds exarillate, with an enlarged white,
3459 round hilum.
3460Four species in southeast Asia from Yunnan,
3461Myanmar, and Indochina to Sumatra, Borneo,
3462and the Philippines.
3463102.Pavieasia Pierre
3464Pavieasia Pierre, Fl. Forest. Cochinch.: t. 317 (1894).
3465Falsely polygamous trees or shrubs. Leaves alter-
3466nate, paripinnate; distal leaflet rudimentary.
3467Inflorescences terminal thyrses. Flowers actino-
3468morphic, bisexual or functionally unisexual;
3469calyx shortly cupular, sepals 5, connate at base,
3470imbricate; petals 5, with a single basal appendage;
3471disk annular; stamens 8; pollen syncolporate, stri-
3472ate; ovary 3-carpellate, pilose, with a single ovule
3473per carpel; style filiform; stigma obscurely
34743-lobed at apex. Fruit a 3-locular, loculicidal
3475capsule. Seeds exarillate.
3476One to three species endemic to China.
3477103.Pentascyphus Radlk.
3478Pentascyphus Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3479Bayer. Akad. Wiss. M€
unchen 9: 472, 495, 539 (1879).
3480Falsely polygamous trees or shrubs. Leaves alter-
3481nate, paripinnate; leaflets alternate, entire; distal
3482leaflet rudimentary; stipules 0. Inflorescences ter-
3483minal or axillary thyrses. Flowers actinomorphic,
3484bisexual or functionally unisexual; sepals 5, dis-
3485tinct, imbricate; petals 5, obovate to cuneate,
3486with a single short, basal appendage connate to
3487petals’ margins; disk annular, 7–8-lobed; stamens
34888; pollen syncolporate or parasyncolporate, rugu-
3489late; ovary 3-carpellate, hirsute, with a single
3490ovule per carpel. Fruit unknown.
3491A single species, P. thyrsiflorus Radlk., in
3492French Guiana, Surinam, and Brazil (Amazonas).
3493104.Phyllotrichum Thorel ex Lecomte
3494Phyllotrichum Thorel ex Lecomte, Notul. Syst. (Paris) 2:
34958 (1911).
3496Falsely polygamous trees. Leaves alternate, pari-
3497pinnate, with 5–6 pairs of leaflets; distal leaflets
3498rudimentary. Inflorescences ramiflorous, race-
3499mose thyrses. Flowers zygomorphic; sepals 5, dis-
3500tinct, imbricate; petals 4, with a single basal
3501appendage; disk unilateral, semi-annular;
3502stamens 8(9); pollen syncolporate or parasyn-
3503colporate, striate; ovary 3-carpellate, densely
3504pubescent, with a single ovule per carpel; style
412 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
3505 nearly trigonous; stigma nearly 3-lobed. Fruit a
3506 3-locular, loculicidal, muricate capsule. Seeds
3507 ovoid, exarillate.
3508 A single species, P. mekongense Thorel ex
3509 Lecomte, endemic to Laos.
3510 105.Placodiscus Radlk.
3511 Placodiscus Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3512 Bayer. Akad. Wiss. M€
unchen 8: 332 (1878).
3513 Falsely polygamous trees or shrubs. Leaves alter-
3514 nate, paripinnate, with 2–9 pairs of leaflets;
3515 distal leaflets rudimentary; pseudostipules some-
3516 times present. Inflorescences cauliflorous, spicate
3517 thyrses. Flowers actinomorphic; calyx urceolate
3518 or turbinate, crowned by 5 valvate or narrowly
3519 imbricate sepals; petals 0; disk annular, dish-
3520 shaped; stamens 8; pollen colporate, striate;
3521 ovary 3-carpellate, tomentose, with a single
3522 ovule per carpel; style short; stigma minute.
3523 Fruits baccate, 1–3-locular, 3-sulcate or 3-lobate,
3524 indehiscent. Seeds exarillate.
3525 About ten species native to tropical Africa.
3526 106.Plagioscyphus Radlk.
3527 Plagioscyphus Radlk., Sitzungsber. Math.-Phys. Cl.
3528 K€
onigl. Bayer. Akad. Wiss. M€
unchen 8: 335 (1878).
3529 Cotylodiscus Radlk. ibid. 8: 334 (1878).
3530 Strophiodiscus Choux (1926).
3531 Poculodiscus Danguy & Choux (1927).
3532 Falsely polygamous shrubs. Leaves alternate,
3533 paripinnate; leaflets 1–5 pairs, serrate; distal
3534 leaflets rudimentary. Inflorescences axillary, ra-
3535 cemose thyrses. Flowers actinomorphic; sepals 5,
3536 distinct or connate at base, imbricate; petals 4–5,
3537 the appendages 2 and marginal or single and
3538 basal; disk annular, vase-shaped, or unilateral,
3539 semi-annular or semi-vase-shaped; stamens (7)8;
3540 pollen colporate or brevicolporate, perforate to
3541 reticulate; ovary 2–3-carpellate, tomentose, with a
3542 single ovule per carpel; stigma apiculate. Fruits
3543 baccate, 1–3-locular, indehiscent. Seeds arillate,
3544 with a longitudinal ventral hilum.
3545 About ten species native to Madagascar.
3546 107.Podonephelium Baill.
3547 Podonephelium Baill., Adansonia 11: 245 (1874).
3548 Falsely polygamous-dioecious shrubs. Leaves
3549 alternate, paripinnate, with 3–6 pairs of leaflets;
3550distal leaflets rudimentary. Inflorescences axillary
3551thyrses. Flowers actinomorphic; calyx crateriform,
3552with 4–7, valvate lobes; petals 0; disk cup-shaped,
35538-crenate; stamens (5–)8; pollen colporate, stri-
3554ate; ovary 3-carpellate, tomentose, with a single
3555ovule per carpel; stigma 3-lobed. Fruit a 1(2)-
3556coccate, crustose, circumscissile dehiscent cap-
3557sule. Seeds with white arillode mostly along
3558dorsal portion.
3559Four species native to New Caledonia.
3560108.Pometia Forst. & Forst.
3561Pometia Forst. & Forst., Char. Gen. Pl.: 55, t. 55 (1775);
3562Jacobs, Reinwardtia 6: 109–144 (1962).
3563Falsely polygamous trees, producing red exu-
3564dates. Leaves alternate, paripinnate; leaflets entire
3565or serrate, often with large orbicular glands
3566beneath; distal leaflet rudimentary; pseudosti-
3567pules present. Inflorescences terminal or axillary
3568thyrses. Flowers actinomorphic, functionally uni-
3569sexual; sepals 5, connate at base to half of their
3570length, valvate; petals 5 without appendages;
3571disk annular or semi-annular; stamens 5(6); pol-
3572len brevicolporate, reticulate (Fig. 79L); ovary
35732(3)-carpellate, with a single ovule per carpel;
3574style filiform; stigma obtuse, emarginate. Fruits
35751(2)-locular, indehiscent, with fleshy mesocarp.
3576Seeds fully covered by an arillode.
3577Two species found in Sri Lanka, Andaman
3578and Nicobar Islands, Indochina, Taiwan, Malesia,
3579Fiji, Samoa, and Tonga. The wood of P. pinnata
3580Forst. is used as firewood in the Pacific Islands.
3581109.Porocystis Radlk.
3582Porocystis Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3583Bayer. Akad. Wiss. M€
unchen 8: 353 (1878).
3584Falsely polygamous trees. Leaves alternate,
3585paripinnate or imparipinnate; leaflets entire; dis-
3586tal leaflet rudimentary or exceptionally fully
3587developed. Inflorescences terminal or axillary
3588thyrses. Flowers zygomorphic, bisexual or func-
3589tionally unisexual; sepals 5, distinct, imbricate;
3590petals 4, clawed, with a bifid appendage above
3591the claw; disk unilateral, semi-annular to
35924-lobed; stamens 8; pollen colporate, rugulate-
3593reticulate; ovary 3-carpellate, with a single ovule
3594per carpel; style short, with 2 recurved, stigmatic
3595branches. Fruit a 3-locular schizocarp, splitting
Sapindaceae 413
Uncorrected Proof
3596 into 3 membranous, inflated, wrinkled mericarps.
3597 Seeds exarillate.
3598 Three species, 2 from Brazil (Amazonas) and
3599 1 from French Guiana.
3600 110.Pseudima Radlk.
3601 Pseudima Radlk., Nouv. Giornale Bot. Ital. 10: 108 (1878).
3602 Falsely polygamous-dioecious, small to large
3603 trees. Leaves alternate, pinnately compound;
3604 distal leaflet rudimentary; stipules 0. Inflores-
3605 cences axillary or terminal thyrses. Flowers bi-
3606 sexual or unisexual; calyx zygomorphic, sepals
3607 5, imbricate, concave, the outer 2 sepals shorter;
3608 petals 5, longer than the sepals, lacking appen-
3609 dages; disk annular, 5-lobed; stamens 8 or 10,
3610 shorter than the petals; pollen colporate, per-
3611 forate; ovary 2(3)-carpellate, with a single ovule
3612 per carpel; style filiform; stigma obtuse. Fruit a
3613 2(3)-coccate, loculicidal, leathery capsule, the
3614 cocci equally developed or one of them rudimen-
3615 tary. Seeds large, arillate.
3616 One species from the lowlands of tropical
3617 continental America.
3618 111.Pseudopancovia Pellegr.
3619 Pseudopancovia Pellegr., Bull. Soc. Bot. France 102: 228
3620 (1955).
3621 Falsely polygamous-dioecious, shrubs. Leaves
3622 alternate, paripinnate; distal leaflet rudimentary;
3623 stipules 0. Inflorescences axillary, spicate thyrses.
3624 Flowers zygomorphic, bisexual or unisexual;
3625 calyx 2-lipped, 4–5-lobed; petals 3–4, clawed,
3626 with a basal appendage forming a pocket; disk
3627 unilateral; stamens 7; pollen colporate, rugulate;
3628 ovary 3-carpellate, with a single ovule per carpel.
3629 Fruit unknown.
3630 A single species, P. heterophylla Pellegr.,
3631 endemic to west equatorial Africa.
3632 112.Pseudopteris Baill.
3633 Pseudopteris Baill., Adansonia 11: 243 (1874).
3634 Falsely polygamous trees. Leaves alternate, pari-
3635 pinnate; leaflets opposite or alternate, entire
3636 or crenate-serrate; distal leaflet rudimentary;
3637 stipules 0. Inflorescences of axillary racemose
3638 thyrses. Flowers actinomorphic, bisexual or uni-
3639 sexual; sepals 5, distinct, imbricate; petals 5,
3640 smaller than the sepals, cucullate, without appen-
3641dages; disk 5-lobed ; stamens 5; pollen colporate,
3642striate; ovary 2–3-carpellate, with a single ovule
3643per carpel. Fruit baccate, 1–3-locular, indehiscent.
3644Seeds sub-globose, completely covered by an
3645arillode.
3646Three species endemic to Madagascar.
3647113.Radlkofera Gilg
3648Radlkofera Gilg, Bot. Jahrb. Syst. 24: 300 (1897).
3649Falsely polygamous unbranched trees. Leaves
3650alternate, paripinnate, with 13–20 pairs of leaflets;
3651distal leaflet rudimentary; stipules 0. Inflores-
3652cences axillary, racemose thyrses, with elongated
3653bracteoles. Flowers zygomorphic, bisexual or uni-
3654sexual; calyx urceolate, sepals 5, connate at base,
3655imbricate; petals 4, with a ventral appendage;
3656disk unilateral; stamens 7–8; pollen colporate,
3657indistinctly rugulate; ovary 5–7(8)-carpellate,
3658tomentose, with a single ovule per carpel; style
3659elongated. Fruit indehiscent, 5–8-locular, fusi-
3660form, with fleshy mesocarp. Seeds exarillate,
3661orange.
3662A single species, R. calodendron Gilg, western
3663Africa.
3664114.Rhysotoechia Radlk.
3665Rhysotoechia Radlk., Actes Congr. Bot. Amsterdam 1877:
3666131 (1879); Etman, Blumea 39: 41 (1994).
3667Falsely polygamous trees or shrubs. Leaves alter-
3668nate, paripinnate. Flowers seemingly bisexual;
3669calyx zygomorphic, the sepals 5, distinct, imbri-
3670cate, 2 outer sepals smaller; petals 5, clawed, with
3671marginal appendages or these 0; disk annular;
3672stamens (7)8; pollen syncolporate or parasyn-
3673colporate, rugulate to perforate, sometimes psi-
3674late; ovary (2)3-carpellate, with a single ovule
3675per carpel; style with 3 stigmatic lines. Fruit a
3676(2)3-locular, sub-globose, obovoid, or subcor-
3677date, loculicidal capsule. Seeds with a cup-shaped
3678arillode at base.
3679About 14 species from Australia, Borneo, Phi-
3680lippines, Sulawesi, Moluccas, and New Guinea.
3681115.Sapindus Plum. ex L.
3682Sapindus Plum. ex L., Sp. Pl.: 367 (1753); Radlkofer in
3683Pflanzenreich 98: 630 (1932); Leenhouts in Fl. Males. I, 11:
3684713 (1994).
414 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
3685 Duodichogamous trees. Leaves alternate, paripin-
3686 nate or unifoliolate; leaflets 2–8 pairs, often
3687 falcate; distal leaflet rudimentary. Inflore-
3688 scences terminal, thyrses. Flowers actinomorphic
3689 or partly zygomorphic, functionally pistillate or
3690 staminate; sepals 5, distinct, imbricate, the outer
3691 2 smaller; petals 4 or 5, with a single large
3692 appendage, a transverse ridge, or a pair reduced
3693 marginal appendages; disk annular or cup-
3694 shaped; stamens 8; pollen colporate, rugulate;
3695 ovary 3-carpellate, with a single ovule per carpel;
3696 style short, stigma capitate or with 3 conivent
3697 stigmatic branches. Fruit schizocarpic, 1(2)-
3698 coccate, with 2(1) rudimentary cocci, separating
3699 into indehiscent globose mericarps, with fleshy
3700 pericarp containing much saponin. Seeds
3701 globose, exarillate. n¼11, 15, 18.
3702 About ten species with tropical to sub-
3703 temperate distribution. Several species used as
3704 ornamentals.
3705 116.Sarcopteryx Radlk.
3706 Sarcopteryx Radlk. Actes Congr. Bot. Amsterdam 1877:
3707 127 (1879); Reynolds, Austrobaileya 2: 53 (1984), reg. rev.;
3708 Welzen, Blumea 36: 91 (1991), reg. rev.
3709 Falsely polygamous shrubs or trees. Indumentum
3710 of simple hairs and red glandular hairs. Leaves
3711 alternate, paripinnate; leaflets 1–5 pairs, entire;
3712 distal leaflet rudimentary. Inflorescences axillary
3713 or terminal, simple or thyrses. Flowers actino-
3714 morphic, functionally unisexual; sepals 5, con-
3715 nate at base, valvate; petals 5, clawed, with
3716 2 dissected or crenate appendages above the
3717 claw, these sometimes crested; disk annular;
3718 stamens 8; pollen syncolporate or parasyn-
3719 colporate, rugulate to rugulate-reticulate; ovary
3720 3-carpellate, with a single ovule per carpel; style
3721 filiform. Fruit a 3-locular, loculicidal, leathery-
3722 woody, capsule, each locule with a narrow, dorsal
3723 wing. Seeds completely covered by dorsally
3724 opened arillode with a basal funiculus-like
3725 appendage, the testa papery.
3726 Twelve to 13 species, from eastern Australia,
3727 Moluccas, and New Guinea.
3728 117.Sarcotoechia Radlk.
3729 Sarcotoechia Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3730 Bayer. Akad. Wiss. M€
unchen 9: 501 (1879); Leenhouts,
3731 Blumea 33: 198 (1988).
3732Falsely polygamous trees. Leaves alternate, pari-
3733pinnate or unifoliolate; leaflets serrate or entire;
3734distal leaflets rudimentary. Inflorescences axil-
3735lary, or ramiflorous thyrses. Flowers actinomor-
3736phic; sepals 5, distinct, slightly imbricate; petals 5,
3737shorter than the sepals, with a pair of seemingly
3738marginal appendages; stamens (5–)7(8); pollen
3739parasyncolporate, rugulate; ovary 2–3-carpellate,
3740with a single ovule per carpel; style apical; stigma
3741slightly lobed. Fruit a 2–3-locular, 2–3-lobed,
3742fleshy, loculicidal capsule. Seeds with a cupular
3743or reduced sarcotesta.
3744About 11 species from Australia (northern
3745Queensland), Papua New Guinea, and Moluccas.
3746118.Schleichera Willd., nom. cons
3747Schleichera Willd., Sp. Pl. 4, 2: 1096 (1806); Leenhouts in
3748Fl. Males. I, 11: 727 (1994).
3749Falsely polygamous trees, with glandular indu-
3750ment. Leaves alternate, paripinnate; leaflets
3751entire; distal leaflet rudimentary. Inflorescences
3752axillary, racemose or thyrsoid. Flowers actino-
3753morphic, functionally unisexual; sepals 4–6,
3754equal, valvate, connate at base; petals 0; disk
3755annular, dish-shaped; stamens 5–9; pollen para-
3756syncolporate, striate; ovary 2–4-carpellate, with a
3757single ovule per carpel; stigma lobed. Fruits
37581-locular, indehiscent, coriaceous, usually with a
3759spiny pericarp. Seed completely covered by an
3760arillode. 2n¼30, 32.
3761A single species, S. oleosa (Lour.) Oken,
3762distributed from Sri Lanka and India to Indo-
3763china, Malesia east to the Moluccas and Lesser
3764Sunda Islands. The seeds are the source of macas-
3765sar oil widely used as hair ointment; the wood is
3766used as construction material.
3767119.Scyphonychium Radlk.
3768Scyphonychium Radlk., Sitzungsber. Math.-Phys. Cl.
3769K€
onigl. Bayer. Akad. Wiss. M€
unchen 9: 473, 495, 519
3770(1879); Ferrucci, Bonplandia 6: 117–124 (1989).
3771Falsely polygamous trees. Leaves alternate, pari-
3772pinnate; leaflets entire; distal leaflet rudimentary.
3773Inflorescences terminal thyrsoids. Flowers func-
3774tionally unisexual; calyx zygomorphic, sepals 5,
3775distinct, imbricate, the outer 2 smaller; petals 5,
3776clawed, twice as long as the sepals, with a bilobed
3777minute appendage forming a pocket above
3778the claw; disk annular, cup-shaped, 5-lobed;
Sapindaceae 415
Uncorrected Proof
3779 stamens 8; pollen colporate, perforate; ovary 2-
3780 carpellate, with a single ovule per carpel; stigma
3781 shortly bifid. Fruit, (1)2-coccate, woody, schizo-
3782 carpic. Seeds exarillate.
3783 A single species, S. multiflorum Radlk., native
3784 to northern and eastern Brazil, and French
3785 Guiana.
3786 120.Sinoradlkofera F. Meyer
3787 Sinoradlkofera F. Meyer, J. Arnold Arb. 58: 183 (1977).
3788 Boniodendron Gagnep., nom. inval.; Leenhouts, Blumea
3789 28: 45 (1982).
3790 Falsely polygamous trees. Leaves paripinnate;
3791 leaflets serrate; distal leaflet rudimentary. Inflor-
3792 escences terminal thyrses. Flowers sub-actino-
3793 morphic; sepals 5, valvate; petals 5, white,
3794 clawed, appendages marginal or 0; disk annular;
3795 stamens 8, geniculate in bud; pollen colporate,
3796 striate; ovary 3-carpellate, with 2 ovules per
3797 carpel; style subulate. Fruits 3-locular, inflated,
3798 loculicidal capsules. Seeds exarillate.
3799 A single species, S. minor (Hemsley) F. Meyer,
3800 native to China.
3801 121.Sisyrolepis Radlk.
3802 Sisyrolepis Radlk. in F. N. Williams, Bull. Herb. Boiss. II,
3803 5: 222 (1905); Leenhouts, Blumea 23: 336 (1977); Welzen
3804 in Santisuk & Larsen, Fl. Thailand 7: 243 (1999).
3805 Delpya Pierre ex Radlk. (1910).
3806 Falsely polygamous shrubs or trees. Leaves alter-
3807 nate, paripinnate; leaflets crenate; distal leaflet
3808 rudimentary. Inflorescences axillary thyrses.
3809 Flowers zygomorphic; sepals 5, distinct, imbri-
3810 cate; petals 4(5), with a pair of marginal appen-
3811 dages; disk semi-annular, lobulate; stamens 8(9);
3812 pollen syncolporate, finely striate; ovary 3-carpel-
3813 late, with a single ovule per carpel; stigma
3814 not lobed. Fruit a 3-locular, echinate, loculicidal
3815 capsule. Seeds exarillate.
3816 A single species, S. muricata (Pierre) Leenh.,
3817 Thailand and Cambodia.
3818 122.Smelophyllum Radlk.
3819 Smelophyllum Radlk., Sitzungsber. Math.-Phys. Cl.
3820 K€
onigl. Bayer. Akad. Wiss. M€
unchen 8: 330 (1878).
3821 Falsely polygamous shrubs. Leaves alternate,
3822 paripinnate; leaflets 3–4 pairs; distal leaflet
3823rudimentary. Inflorescences axillary thyrses.
3824Flowers actinomorphic; calyx cupular, sepals
38255, connate, imbricate; petals 5, without appen-
3826dages; disk annular, 5-lobed; stamens 8; pollen
3827colporate, striate; ovary (2)3-carpellate, with
3828a single ovule per carpel; style thickened. Fruit
3829a (1)2(3)-coccate, chartaceous capsule. Seeds
3830exarillate.
3831A single species endemic to South Africa.
3832123.Stadmania Lam.
3833Stadmania Lam., Tabl. Encycl. 2: 443 (1793); reg. rev.;
3834Capuron, Mem. Mus. Natl. Hist. Nat. B, Bot. II, 19:
3835151–160 (1969).
3836Pseudolitchi Dand. & Choux (1926).
3837Dioecious or monoecious trees. Leaves alternate,
3838paripinnate; leaflets (1)3–5 pairs, serrate; distal
3839leaflet rudimentary. Inflorescences terminal or
3840axillary, racemose thyrses. Flowers actinomor-
3841phic, functionally unisexual; calyx cupular,
3842sepals 5, valvate; petals 5, sometimes clawed,
38430 in one species; appendages simple, crested or
3844dissected, basal or above the claw; disk annular
3845to 5-crenate; stamens (6–)8; pollen colporate,
3846striate or perforate, sometimes irregularly striate
3847or rugulate; ovary 3-carpellate, with a single
3848ovule per carpel; style short. Fruit a 1(3)-coccate,
3849indehiscent. Seeds covered with a translucent
3850arillode.
3851Six species from Mauritius, Madagascar, and
3852eastern tropical Africa.
3853124.Stocksia Benth.
3854Stocksia Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 304
3855(1853).
3856Polygamous spiny shrubs. Leaves simple, fasci-
3857culate on short branches axillary to the spines;
3858stipules 0. Inflorescences axillary, racemose thy-
3859rses. Flowers zygomorphic, functionally unisexual;
3860sepals 5, imbricate, distinct, the outer smaller;
3861petals 5, clawed, without appendages; disk annu-
3862lar-lobed; stamens 7–8; pollen colporate, striate;
3863ovary 3-carpellate, with 2 ovules per carpel; style
3864distal; stigma minute. Fruit a 3-locular, membra-
3865nous, inflated, loculicidal capsule. Seeds exaril-
3866late.
3867A single species, S. brahuica Benth., from
3868Persia and Afghanistan.
416 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
3869 125.Storthocalyx Radlk.
3870 Storthocalyx Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3871 Bayer. Akad. Wiss. M€
unchen 9: 499, 660 (1879).
3872 Falsely polygamous shrubs or trees. Leaves alter-
3873 nate, paripinnate, with 2–10 leaflets; distal leaflet
3874 rudimentary; stipules 0. Inflorescences axillary,
3875 spicate or thyrsoid. Flowers actinomorphic, func-
3876 tionally unisexual; sepals 5, distinct, narrowly
3877 imbricate or valvate; petals 5, without appen-
3878 dages; disk annular; stamens 8; pollen syncol-
3879 porate, perforate; ovary 3-carpellate, with a
3880 single ovule per carpel; style subulate; stigma
3881 minute. Fruit a 3-locular, trigonous or pyriform,
3882 corticose-woody loculicidal capsule. Seeds with
3883 a ventrally split, fimbriate arillode.
3884 Four species from New Caledonia.
3885 126.Synima Radlk.
3886 Synima Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3887 Bayer. Akad. Wiss. M€
unch. 9: 501 (1879); Leenhouts &
3888 Adema in Fl. Males. I, 11: 730 (1994).
3889 Monoecious trees. Leaves alternate, paripinnate;
3890 leaflets crenate-denticulate, without domatia or
3891 glands; distal leaflet rudimentary. Inflorescences
3892 axillary, together sometimes pseudo-terminal.
3893 Flowers actinomorphic; calyx shortly cupular,
3894 sepals 5, short, distinct, narrowly imbricate;
3895 petals 5, rhomboidal, with a pair of recurved,
3896 woolly or ciliate appendages, usually distinctly
3897 crested; disk annular; stamens 8; pollen parasyn-
3898 colporate, rugulate-reticulate; ovary 3-carpellate,
3899 with a single ovule per carpel; style apical;
3900 stigma slightly lobed. Fruit a 3-locular, trigonous,
3901 loculicidal capsule. Seeds with basal or dorsal
3902 sarcotesta.
3903 Two species; Australia (N Queensland) and
3904 southeast New Guinea. Lowland and montane
3905 rain forest, mossy oak forest, up to 900 m.
3906 127.Thouinidium Radlk.
3907 Thouinidium Radlk., Sitzungsber. Math.-Phys. Cl. K€
onigl.
3908 Bayer. Akad. Wiss. M€
unchen 8: 284 (1878).
3909 Falsely polygamous trees. Leaves alternate, pari-
3910 pinnate; leaflets 2–8 pairs, serrate or entire; distal
3911 leaflet rudimentary; stipules 0. Inflorescences
3912 terminal thyrses. Flowers functionally unisexual;
3913 calyx actinomorphic or slightly zygomorphic,
3914 sepals 5, distinct, imbricate; petals (4)5, clawed,
3915with an appendage above the claw forming
3916a pocket; disk cup-shaped; stamens 6–8(–10);
3917pollen colporate, perforate; ovary 3-carpellate,
3918with a single ovule per carpel; style short, subu-
3919late. Fruit schizocarpic, splitting into 3, distally
3920winged, samaroid mericarps. Seeds exarillate.
3921Six species from Mexico, Central America,
3922and some islands in the Greater Antilles.
3923128.Tina Schult., nom. cons. prop.
3924Tina Schult. in Roem. & Schult., Syst. Veg. 5: XXXII, 414
3925(1819–1820) (‘1819’).
3926Gelonium Gaertn., Fruct. Sem. Pl. 2: 271, fig. 139 n. 8 (1791),
3927nom. rej. prop.
3928Falsely polygamous shrubs or trees. Leaves alter-
3929nate, paripinnate, with 2–6 pairs of leaflets; leaf-
3930lets serrate or crenate-serrate; distal leaflet
3931rudimentary; stipules 0. Inflorescences axillary
3932thyrses. Flowers functionally unisexual; calyx
3933actinomorphic or zygomorphic, sepals (3–)5,
3934distinct, imbricate, in two series; petals 5, with
39352 marginal appendages; disk annular; stamens
3936(6–)8; pollen syncolporate or parasyncolporate,
3937rugulate; ovary 2(3)-carpellate, with a single
3938ovule per carpel; style subulate; stigma a invagi-
3939nate prolongation of the style. Fruit a 2-locular,
3940loculicidal capsule. Seeds arillate.
3941Six species from Madagascar.
3942129.Tinopsis Radlk.
3943Tinopsis Radlk. in T. Durand, Index Gen. Phan.: 78 (1887)
3944[1888].
3945Bemarivea Choux (1927).
3946Falsely polygamous-dioecious trees. Leaves
3947alternate, paripinnate, with 2–4 pairs of leaflets;
3948distal leaflet rudimentary; stipules 0. Inflo-
3949rescences axillary or terminal thyrses. Flowers
3950actinomorphic, functionally unisexual; calyx
3951slightly zygomorphic, the sepals 5, imbricate,
3952distinct; petals 5, with a single appendage form-
3953ing a pocket or with 2 marginal appendages; disk
3954annular, slightly sulcate; stamens 5(7); pollen
3955colporate, rugulate to rugulate-reticulate; ovary
39562-carpellate, with a single ovule per carpel. Fruit
3957a 2-locular, indehiscent or less often tardily
3958and incompletely dehiscent capsule, usually
39591-seeded. Seed arillate.
3960Eleven species from Madagascar.
Sapindaceae 417
Uncorrected Proof
3961 130.Toechima Radlk.
3962 Toechima Radlk., Actes Congr. Bot. Amsterdam 1877: 130
3963 (1879); Reynolds, Austrobaileya 2: 176 (1985); Leenhouts,
3964 Blumea 33: 203 (1988); Leenhouts in Fl. Males. I, 11: 732
3965 (1994).
3966 Falsely polygamous trees. Leaves alternate, pari-
3967 pinnate; leaflets entire to serrate. Inflorescences
3968 axillary thyrses. Flowers actinomorphic; sepals 5,
3969 equal, valvate to narrowly imbricate; petals 5,
3970 clawed, with a single crested appendage; disk
3971 annular; stamens 8; pollen parasyncolporate, rug-
3972 ulate to rugulate-reticulate; ovary 2–3-carpellate,
3973 with a single ovule per carpel; stigma lobed. Fruit
3974 a 2–3-locular, fleshy, loculicidal capsule. Seed
3975 with a basal placental arillode.
3976 About eight species distributed in Australia
3977 and New Guinea.
3978 131.Toulicia Aublet
3979 Toulicia Aubl., Hist. Pl. Guiane 1: 359 (1775).
3980 Falsely polygamous-dioecious, small,
3981 unbranched trees. Leaves imparipinnate; leaflets
3982 usually falcate, opposite or alternate; distal leaflet
3983 rudimentary. Inflorescences terminal or axillary
3984 thyrses. Flowers zygomorphic, staminate or
3985 pistillate; sepals 5, unequal, imbricate; petals 4,
3986 with a petaloid, bifid ventral appendage or with
3987 2 marginal appendages; disk unilateral, semi-
3988 annular; stamens 8; pollen colporate, perforate;
3989 ovary 3-carpellate, with a single ovule per carpel;
3990 style with 3 stigmatic branches. Fruits schizo-
3991 carpic, splitting into three, samaroid mericarps,
3992 each with a proximal wing and a papery, inflated
3993 locule. Seeds exarillate.
3994 About 12 species from the lowlands of South
3995 America.
3996 132.Trigonachras Radlk.
3997 Trigonachras Radlk., Actes Congr. Bot. Amsterdam 1877:
3998 116 (1879); Leenhouts, Blumea 33: 204 (1988).
3999 Falsely polygamous trees. Leaves alternate, pari-
4000 pinnate; leaflets entire, often with glands below;
4001 distal leaflet rudimentary. Inflorescences axillary
4002 or terminal thyrses or panicles. Flowers actino-
4003 morphic, bisexual or unisexual; sepals 5, distinct,
4004 narrowly imbricate (sub)equal; petals 5, clawed,
4005 with (1)2 appendages above the claw; disk annu-
4006 lar; stamens (7)8(9); pollen syncolporate or para-
4007syncolporate, perforate to scabrate with often
4008finely tapering scabrae; ovary 3-carpellate, with
4009a single ovule per carpel; style with 3 stigmatic
4010lines. Fruit a 3-locular, fleshy, loculicidal capsule.
4011Seeds exarillate.
4012About eight species, occurring throughout
4013Malesia, but absent from Java and the Lesser
4014Sunda Islands.
4015133.Tripterodendron Radlk.
4016Tripterodendron Radlk., Sitzungsber. Math.-Phys. Cl.
4017K€
onigl. Bayer. Akad. Wiss. M€
unchen 20: 290 (1891).
4018Falsely polygamous-dioecious trees. Leaves
4019alternate, tripinnate; leaflets denticulate or ser-
4020rate; distal leaflet rudimentary; stipules 0. Inflor-
4021escences axillary thyrses. Flowers actinomorphic,
4022bisexual or functionally unisexual; sepals 5–6,
4023valvate; petals clawed, 5(6–8), with a pair of app-
4024endages above the claw; disk annular; stamens 8;
4025pollen colporate, striate; ovary 2-carpellate, with
4026a single ovule per carpel; style short, with a mar-
4027ginal stigmatic line. Fruit a 2-locular, fleshy, loc-
4028ulicidal capsule. Seeds arillate.
4029A single species endemic to east-central Brazil.
4030134.Tristira Radlk.
4031Tristira Radlk., Actes Congr. Bot. Amsterdam 1877: 133
4032(1879); Leenhouts in Fl. Males. I, 11: 740 (1994).
4033Falsely polygamous trees. Leaves alternate, pari-
4034pinnate; leaflets opposite or alternate, entire
4035or dentate; distal leaflets rudimentary. Inflores-
4036cences terminal or axillary thyrses. Flowers
4037bisexual or functionally unisexual; calyx zygo-
4038morphic, sepals 5, distinct, imbricate, the outer
4039two smaller; petals 0; disk annular, lobed; sta-
4040mens 8(9); pollen parasyncolporate, rugulate;
4041ovary 3-carpellate, with a single ovule per carpel;
4042style subulate, with 3 stigmatic lines. Fruits
40433-locular, indehiscent drupes, with slightly fleshy
4044exocarp and stony endocarp, each locule with a
4045dorsal wing. Seeds exarillate.
4046A single species, T. triptera (Blanco) Radlk.,
4047eastern Philippines, Celebes, and Moluccas.
4048135.Tristiropsis Radlk
4049Tristiropsis Radlk. in T. Durand, Index Gen. Phan.: 76
4050(1887)[1888]; Leenhouts in Fl. Males. I, 11: 742 (1994), rev.
4051Palaoea Kanehira (1935).
418 P. Acevedo-Rodrı
´guez et al.
Uncorrected Proof
4052 Falsely polygamous trees. Leaves alternate, bipin-
4053 nate; leaflets entire; distal leaflet rudimentary.
4054 Inflorescences axillary thyrses. Flowers bisexual
4055 or functionally unisexual; calyx zygomorphic,
4056 sepals 5, imbricate, the outer 2 smaller; petals
4057 0 or 5, the appendages either forming a pocket,
4058 or a folded marginal outgrowth; disk annular;
4059 stamens 8(–13); pollen parasyncolporate, rugu-
4060 late; ovary 3(–5)-carpellate, with a single ovule
4061 per carpel; stigma not lobed, grooved. Fruit a
4062 (2)3-locular, indehiscent drupe, with slightly
4063 fleshy exocarp and stony endocarp; seeds
4064 exarillate.
4065 Three species from Borneo, the Philippines
4066 and throughout eastern Malesia to northeastern
4067 Australia, the Solomons, Marianas, and Christmas
4068 Islands in the Pacific.
4069 136.Tsingya Capuron
4070 Tsingya Capuron, Mem. Mus. Natl. Hist. Nat., B, Bot. II,
4071 19: 104 (1969).
4072 Monoecious trees. Leaves alternate, paripinnate;
4073 leaflets entire; distal leaflet rudimentary; stipules
4074 0. Inflorescences axillary racemose thyrses. Flow-
4075 ers actinomorphic, functionally unisexual; sepals 5,
4076 valvate; petals 0; disk annular, pulvinate; stamens
4077 8–10; pollen colporate, striate; ovary 3-carpellate,
4078 with a single ovule per carpel; style with 3 stig-
4079 matic lines. Fruit (immature) unilocular by abor-
4080 tion. Seed one per locule, arillate, with long
4081 ventral hilum.
4082 A single species, T. bemarana Capuron,
4083 endemic to Madagascar.
4084 137.Ungnadia Endl.
4085 Ungnadia Endl., Atakta Bot. t. 36 (1835) [1833].
4086 Falsely polygamous shrubs or trees. Leaves alter-
4087 nate, paripinnate; leaflets serrate; distal leaflet
4088 fully developed; stipules 0. Inflorescences axillary
4089 or ramiflorous, pseudo-umbelliform. Flowers
4090 zygomorphic, bisexual or functionally unisexual;
4091 sepals 5, imbricate, 3 distinct, 2 connate; petals
4092 4–5, clawed, with a tuft of filiform appendages
4093 above the claw; disk unilateral, undulate, with
4094 androgynophore; stamens (7)8(–10); pollen col-
4095 porate, finely striate; ovary 3-carpellate, stipitate,
4096 with 2 ovules per carpel; style filiform with punc-
4097 tiform stigma. Fruit a 3-locular, 1–2-seeded,
4098loculicidal, coriaceous capsule. Seeds exarillate,
4099with large white hilum. 2n¼32.
4100A single species, U. speciosa Endl., Mexico
4101and southern United States (Texas).
4102138.Vouarana Aubl. Fig. 78
4103Vouarana Aubl., Pl. Guiane 2: (Suppl.) 12, fig. 374 (1775).
4104Falsely polygamous medium-sized trees. Leaves
4105alternate, paripinnate; leaflets entire; distal leaflet
4106rudimentary. Inflorescences axillary or terminal
4107thyrses. Flowers 4–5-merous, actinomorphic or
4108zygomorphic, bisexual or unisexual; sepals dis-
4109tinct, unequal, concave, imbricate; petals rhom-
4110bic, shorter than the sepals, with two marginal
4111appendages; disk annular and lobed; stamens
41126–8; pollen parasyncolporate, rugulate; ovary
41132-carpellate, with a single, basal ovule per carpel;
4114stigma subulate and papillose. Fruit a 1–2-seeded,
4115woody, loculicidal capsule. Seeds ellipsoid with a
4116large basal arillode.
4117Two species from Costa Rica to northern
4118Brazil.
4119139.Xerospermum Blume
4120Xerospermum Blume, Rumphia 3: 99 (1847); Leenhouts,
4121Blumea 28: 389 (1983), rev.
4122Falsely polygamous or dioecious trees. Leaves
4123alternate, paripinnate; leaflets entire, abaxially
4124with flat, orbicular glands toward proximal por-
4125tion of blade; distal leaflet rudimentary. Inflores-
4126cences axillary thyrses. Flowers actinomorphic,
4127bisexual or functionally unisexual; sepals 4–5,
equal, imbricate; petals 4–5, sessile to clawed,
4129without appendages; disk annular or semi-annular;
4130stamens 7–9; pollen colporate, striate to psilate,
4131sometimes irregularly striate or rugulate; ovary 2
4132(3)-carpellate, with a single ovule per carpel;
4133stigma lobed. Fruit indehiscent, 1–2-coccate,
4134coriaceous to woody, granular to shortly spiny.
4135Seed with complete, thin, sarcotesta. 2n¼32.
4136Two species from Bangladesh, Indochina
4137and western Malesia. X. noronhianum Blume is
4138dioecious.
4139140.Zollingeria Kurz, nom. cons.
4140Zollingeria Kurz, J. Asiat. Soc. Bengal, 41, 2: 303 (1872),
4141non Schultz.-Bip. (1854), nom. rej.; Adema, Blumea 37: 73
4142(1992); Welzen in Fl. Thailand 7: 248 (1999).
Sapindaceae 419
Uncorrected Proof
4143 Falsely polygamous trees. Leaves alternate, pari-
4144 pinnate; leaflets entire; distal leaflet rudimentary.
4145 Inflorescences axillary thyrses. Flowers actino-
4146 morphic or zygomorphic; sepals 5, distinct, slightly
4147 to distinctly unequal, usually imbricate; petals
4148 4–5, with or without appendages; disk annular
4149 or semi-annular; stamens 8; pollen colporate,
4150 irregularly striate; ovary 3-carpellate, unilocular,
4151 with a single ovule per carpel; stigma 3-lobed, or
4152 3 stigmatic lines. Fruit 1-locular, 3-winged,
4153 coriaceous, indehiscent. Seed exarillate, flattened,
4154 elongated.
4155 Three or four species from Myanmar, Laos,
4156 Thailand, and Borneo.
4157INSUFFICIENTLY KNOWN GENUS:
4158 141.Chonopetalum Radlk.
4159 Chonopetalum Radlk., Bot. Jahrb. Syst. 56: 258 (1920).
4160 Falsely polygamous trees. Leaves alternate, pari-
4161 pinnate; distal leaflet rudimentary; stipules 0.
4162 Inflorescences axillary thyrses. Flowers actino-
4163 morphic, functionally staminate or pistillate;
4164 calyx cup-shaped, sepals 5, distinct, imbricate;
4165 petals 5, with a single basal appendage as long
4166 as the petal; disk annular; stamens 8; pollen
4167 unknown; ovary 3-carpellate (as inferred from
4168 the pistillodes; pistillate flower not known).
4169 Fruit not known.
4170 One species, Ch. stenodictyum Radlk., known
4171 only from the type collection from Equatorial
4172 Guinea, Africa.
4173 Doubtful Genus:
4174 Hirania Thulin
4175 Hirania Thulin, Nord. J. Bot. 24(5): 510 (2007).
4176 Dioecious ? shrubs. Leaves simple, alternate on
4177 long shoots, or congested in short, lateral shoots;
4178 stipules 0. Inflorescences terminal thyrses. Flow-
4179 ers zygomorphic; sepals 5, partially imbricate;
4180 petals 4, clawed, without appendages, pink; disk
4181 unilateral, of two erect segments; stamens 8.
4182 A single species recently described from
4183 Somalia based on a specimen bearing only pistil-
4184 late flowers.
4185 We have not seen any material of this puta-
4186 tive Sapindaceous genus, and since the original
4187 description portrays the floral disk as intrastam-
4188inal, we wonder if it really belongs in Sapindaceae
4189or not. Additional material would be necessary to
4190place this genus in the right family.
4191Selected Bibliography
4192Acevedo-Rodrı
´guez, P. 1990. The occurrence of piscicides
4193and stupefactants in the plant kingdom. Adv. Econ.
4194Bot. 8: 1–23.
4195Acevedo-Rodrı
´guez, P. 1993. Systematics of Serjania
4196(Sapindaceae). Part I: A revision of Serjania Sect.
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