JOURNAL OF THE ARNOLD ARBORETUM

HARVARD UNIVERSITY VOLUME 60 US ISSN 0004-2625 Journal of the Arnold Arboretum

Subscription price $25.00 per year. Subscriptions and remittances should be sent to Ms. E. B. Schmidt, Arnold Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Claims will not be accepted after six months from the date of issue. Volumes 1-51, reprinted, and some back numbers of volumes 52-56 are available from the Kraus Reprint Corporation, Route 100, Millwood, New York 10546, U.S.A. EDITORIAL COMMITTEE S. A. Spongberg, Editor E. B. Schmidt, Managing Editor P. S. Ashton K. S. Bawa P. F. Stevens C. E. Wood, Jr. Printed at the Harvard University Printing Office, Boston, Massachusetts

America, Europe, Asia Minor, and in eastern Asia where the center of species diversity occurs. The only North American representative of the genus, C. caroliniana grows along the edges of streams and in wet, rich soils in forested areas from Nova Scotia and Quebec southward to Florida and westward into Minnesota, Iowa, Missouri, eastern Texas, and Oklahoma. Disjunct popula- • - - Mexico and in Central

The stems of Carpinus caroliniana, a small i bluish-gray, sinuate bark and an ; _ _ trees (Fagus spp.). The wood, which i

Second-class postage paid at Boston, Massachusetts JOURNAL OF THE ARNOLD ARBORETUM

REVISION OF MARGARITARIA (EUPIIORBIACEAE)

GRADY L. WEBSTER

AM ONC 1 THE SMALLER GENK aiphorbiaceae subfamily Phyllanthoi- deae 1 Pax , Margaritaria has a larly broad distribution (MAP 1) in •he New and Old World (except for the Pacific islands). The name' (in alluding to the 'I e C\ n I.i< p i b white cndocarp of the fruit, v . used by Hermann (1689), and the plan. 3 illustrated by Pluke >92). Although the genus was estab- lished hv Linnaeus films in 1 early two centuries passed before it achie\ general acceptance taxonomic literature. Because the :ger Linnaeus included two different genera of plants in the original iption .1 taxonornh understanding ol the genu; \v;is dow in develop- Antoine de Jussieu (1789) and Adrien de Jussieu (1824) both re- ed Mariiaritaria as a gcnu> of uncertain position, allliough the latter- abed a West Indian form of M. nobilis as Cicca antillana A. Juss. on the basis of an observation by Choisy, reported a possible rela- hip of Mariiaritaria with Cicca L. Baillon (1858) treated Margari- in a rather con toed ni.mm i pi i« in< \ uious species in four different on , oi Cicca 3 \\( II i in lli ep it. t< < ( nu \ asp > m u B nllt n

"w^.^X

orld distribution oi \1< ai aria generalized). Base maps from

t and Fellows of Harvard College, 1979. 404 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

Mueller Argovicnsis (1866) continued this association between Cicca and Margaritaria but clarified the relationships to some extent by removing Cicca distkha (-- Phvllanthus acidus (L.) Skeels) to a separate subsec- tion of Phyllunthus sect. Cicca (L.) Mueller-Arg. Baillon and Mueller saw that the Old World taxa described under Pro- sorus Dalz. by Dalzell (1852) and Thwaites (1856) were related to the New World Margaritaria. However, they failed to realize that the taxa Margaritaria/Prosorus do not belong in close juxtaposition with Cicca (scn.su stricto). Bentham (1880), who rejected Margaritaria as being founded on. a mixtme of taxa., did point out that Prosorus was discordant with Cicca because of its tricarpellate fruit. Hooker (1887) judiciously maintained Cicca and Prosorus as separate sections of Indian Phvllanthus. However, this felicitous disposition of Hooker was ignored by Tax (1890) and Pax and Hoffmann (1931). who lumped Cicca and Margaritaria to- gether in Phyllanthus sect. Cicca. Although occasional writers of floras (e.g., Britton & Millspaugh, 1920; Gamble, 1924) maintained Margari- taria or Prosorus as distinct genera, most conservative botanists have fol- lowed the usage of Mueller and of Pax, at least until the rehabilitation of Margaritaria by Webster (1957) and Airy Shaw (1966). As I have pointed out earlier (Webster, 1957, 1968), Margaritaria is really quite distinct from Phvllanthus (including Cicca) and indeed is closer to Flucggca Willd. in Phyllantheae subtribe klueggeinae Mueller- Arg. (Webster, 1975). The unique seeds, with fleshy exotesta and thick, bony endotesta separati Ma ritar'u from II otiiei kinllantheae. The fruits of most species of Margaritaria. with a brittle, irregularly shatter- ing, papery endocarp (FIGURKS 5, 6), are equalh distinctive. Staminate material of Margaritaria max he distinguished from that of Flucggca by the lack of a pistillode, and from thai of Phyllanthus by the annular disc subtending a tetramerous androeciuni. The resemblance with Phyllanthus sect. Cicca that impressed 19lh century botanists is very superficial in- deed, sine.' Cicca (liiivtc markedly in its drupaceous fruits with thin- walled, dry seeds and in its very different vegetative structure (with typical phyllanthoid branching, the ultimate axes being short lived and deciduous, in contrast to the persistent mi specialized branches of Mar-

Morphologically, the species of Margaritaria aie all rather similar, and the dioecious character of the plants makes it rather difficult to construct workable keys to imlividu I p aniens. Most taxa ire deciduous shrubs or small trees that grow in easonal for< t oi bu h .< fetation. The habit of Margaritaria agrees with that of many other unspeoialized Euphorbia- ceae; Halle ct al. (1978) cite M. discoidca as an example of the architec- tural model of Roux. in which the apline axi ha | u al phyllotaxy but the leaves on the branchlets are distichous. To judge from my own ob- servations of M. nobilis i well i from indi< it ion in the literature, all of the species in the genus would seem io belong to the model of Roux. The entire leaves, with diort petioles and distinct stipules, vary relatively slightly in morphology. Flowers, often produced at the proximal nodes VEBSTER. MARGARITARIA of leaves is mature. Opler. Frankie. and baker (1(>7(>) have reported that in the seasonal forest in Guanacaste. Costa Rica. .1/. nohilis flowers about two weeks after the first spring rain in April or May, and only remains in flower about three days. Other Maryjiritana populations in African and Asian monsoon forests may show similar flowering behavior. The tetra- meroii caly: i rathei imilai in ill no i, md th< annular floral disc is so constant that it provides a frond generic character. Tn all normal staminate flowers (FIGURE 12), the stamens are consistently 4, and the filaments are nearly always free. The tricolporate, semitectate pollen grains of M ar^aritaria are typical of many unspecialized Phyllantheae and in en imilai to tho ( oi in li Kohler, 1965). The pistillate flowers of Margaritaria ( FIGURES 11. 13) have calyx and disc similar to that of the staminate. The main differential characters are found in the ovary, which has from 2 to 6 carpels. Mueller (1866) used carpel number as a diagnostic character in defining subsections of his sect. Circa, but its value in indicating phvlewenetic relationships is not very great Carpel number has keen independently reduced to 2 in the West Indies (.1/. hottcami). the Indian Ocean islands (.1/. aiwma'a). and .Australia (J/, dubiuni-traccyi) ; within the Arnencan .'!/. nobilis it fluc- Uiat iiom 1 to o in mli in tin ia!>l< tnann i thai it cannot be used to define subspecific taxa. However, despite some overlap, the pattern of carpel number distribution is distinctively different in M. nobilis and the closely related M. discoidea (TABLES 1,2). The fruit oi mo ! [U^M- ot Mar^aritaria an different from those of all other Phyllantheae. The endocarp is thin, papery, and hyaline, and easily shatters to expose the fleshy seeds (FIGURES 5, 6). The seeds are

(exotesta), bony sclerotesta (endotesta I. and large hilum. These seeds would appear to be morphologically adapted for bird dispersal: van der Fiji (1969), in fact, discusses .10 nobilis as an example of "deceitful" ornithochorous seeds (although the sarcotesta is sufficiently thick that the notion of deceit is perhaps here misplaced). The scletole-m-i of the seed furnishes a useful systematic character, since it varies considerably in size and sculpturing. Tt is distinctively rugose in 1/. indka and M. rhom- boidalis, but smooth in most other taxa. Cytologically, Mur^aritaria appears to be diploid with a base number of Id (2// '() i hk' m,MO mi peciali/cd ta\a of subfamily Phyllan- thoideae (Webster & Ellis. 1962; Mangenot in Bancilhon, 1971). Al- though there are only two published counts (for ,1/. discoidea and M. no- bilis), the uniformity of the gemis makes it appear unlikrlv that nddi tional count would c\ ! n\ is niti< it a >l<> i< il h rsi Ecologically, species of Manjar/iaria might fairly be characterized as planl ol econdan ucce a'ons in tropical easonal climates, although they may also be found in < limax ot eve rgi '< n fore u Vdjanohoun (1964) reported that M. discoidea is common in secondary stages of moist forest JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

. 2. Carpel number variation in American and African taxa of Margaritaria.

herbarium specimens, M. nobilis displays .

Analysis of specimens from the great areas inhabited by M. nobilis, HI. discoidca, and .1/. ind'na. in America. Africa, and Asia, respectively, shows a considerable amount of ill-defined fluctuating morphological variability, as one might expect to find in extensive panmiclic populations. Only in Africa is there more than one distinct mainland continental species. Spe- ciation has occurred mainly in the Caribbean and Indian Ocean islands and thus appears to reflect a "peripheral'' effect; i.e., the definitive geo- 1979] WEBSTER, MARGARITARIA 407 graphic segregation of islands -reins to have been prerequisite to specia- tion, at least outside Africa. Reconstruction of phylogenetic relationships in M < neurit aria must be based mainly on comparison of living species, in view of the paucity and questionableness of the fossil evidence. Madel (1962) has noted the re- semblance between the wood of the Upper Cretaceous Paraphyllanthoxylou and the living Margaritaria discoidca. Bailey (1924) established the ge- nus Paraphyllanthoxylon on the basis of similar Cretaceous wood from Arizona; more recently Cahoon (1972) has described /'. alabamcnse from Cenomanian/Turonian rocks in central Alabama. The large diameter (1 m. or more) of some of the Alabama logs is so uncharacteristic of living species ol 1// iritaria that th< em ignment ippears dubious; in fact, even if these various Cretaceous '-species" of Paraphyllanthoxylon are euphorbiaceous it is doubtful if one could defend ;i closer systematic assignment than simply the tribe Phyllantheae. Since the pollen of Mar- Madagascar indicates that this region m. y b d ignatei i th mi I hi h homeland for Mar- garitaria, as was suggested earlier for another genus of Phyllantheae, Meineckia Baillon (Webster, 1965). The ornithochorous dispersal of liar gar it aria would seem to allow a faith high | mbabilih foi ucce dul transoceanic long-distance migration; however, the failure of the genus to invade the Pacific i Cud u<", . , i[-iar ;, ]i |1(1 i 1 >il11^ may be somewhat limited. As far as the intercontinental th nd.mCi i Margaritaria is con- cerned, there does not appear to be enough evidence foi a decision to be made between the alternative models of transoceanic dispersal or migra- tion over rafting tectonic plates. However, when the distribution pat- terns of other Phyllantheae such as Meineckia and Flueggea are taken into account, one ma}/ perhaps best explain the distribution of Margaritaria by postulating both expansion and contraction from i once more nearly ontinuoti rang* (prob bl\ achi ved without exl nsive long-distance dispersal).

SYSTEMATIC TREATMENT Margaritaria Linnaeus f. Suppl. PI. 66. 1781. Cicca sect. Pseudo-Cicca Baillon, Etude Gen. Euphorb. 618. 1858. Phyllanthus sect. Cicca subsect. Paracicca Mueller-Arg. Linnaea 32: 50. 1863. Phyllanthus sect. Cicca subsect. Margaritaria (L. f.) Mueller-Arg. in DC. Prodr. 15(2): 414. 1866. TYPE: \I,I, > a. n«> „/ ./<••>///' 1 . i. Prosorus Dalz. Jour. Bot. Kew. Misc. 4: 345. 1852. Cicca sect. Prosorus (Dalz.) Baillon, Etude Gen. Euphorb. 619. 1858. Phyllanthus sect. Cicca subsect. Prosorus (Dalz.) Mueller-Arg. in DC. Prodr. I5f2): 416. 1866,, i'hxdo-nhH ,, /,,»,-/, f I'll . HookcM I 11 Brit India 5: 305. 1887. 1* i Prosorus oiduus Dalz ' Margaritaria nana < )alz i \in Shaw) JUl RN Vh Ul H] \ I Ul !) \KI1URI 1 I \l |vui (>(.)

FIGURES 1-4. Leal bases of Margaritnrhr. I. .1/. satmlnts (Correll 43522). showing ulaxialh < li nneled petiole ' 1/ his/> i, • \ppcrt 107), showing nonehanneled pet ink' S '/ rhomlu d, ' -> (/ .

i „ inn u < i iillMII 1 mde Uen. Kuphorb. 620. 1S5S. TYPE: Zygospermum zcxlanicum Thwahes ex Baillon ( Mar^aritaria cyanosper- •ma (Gaertner) Airy Shaw). 1979] WEBSTER, MARGARITARIA 409

Wurtzia Baillon. Adansonia I. 1: 186. t. 7, figs. 5, 6. 1861. TYPE: Wurtzia tctracocca Baillon i l/« i int, , ,, t r, core, i [billon i Webster i Ore,/ sees. Ciccoidcs Thouars ex Baillon Ktucle (den. Kuphorh. 618. 1SVS. i'hxlU'ih'n s .(i (,( rvi'i . (_7rr

Dioecious shrubs or trees (rarely scandent) : foliage usually deciduous. new leaves appearing with flowers (or sometimes evergreen ) : phyllotaxy on branches distichou branch* pei i tent bar! u ually lenticellate in age. Leaves mostly chartaceous. pinnately veined: veins brochidodro- mous; margins entire; petioles short; stipules entire or denticulate, deciduous or subpersistent. Flowers in clusters at proximal axils of expanding leafy branches (or on short shoots): pistillate (occasionally) and staminate (raieb) Ibmei oliiai\ stamina! flower: pedicel elon- gated, often capillary, not. articulated; sepals 4. biseriate, often unequal (outer pair usually narrower), membranous or chartaceous, entire or den- ticulate, midvein u ualh pu b b inched di innulai entir< oi h lowly lobed, adnate to base of calyx (reduced oi absent in a few taxa) ; stamens 4, the filaments free (rarely basally connate), the anthers ex- trorse in bud, dehiscing longitudinally, muticous; pollen grains subglobose, 3-colporate, semitectate; pistillode absent. Pistillate flower: pedicel terete oi flattened not irticulatod calyx and disi imilai to staminate; ovary with 2 to 6 carpel;, the styles free or basally connate, bifid to bipartite, often somewhat dilated ovule 7 p< i loeuh hemitropous. Fruit capsular, more or less irregularly dehiscent, the preen exocarp separating from the thin, papery endocarp (in most taxa): seeds normally paired in each lo- cule, the outer coat (sarcotesta) fleshy, bluish, the inner coat (sclerotesta) dm I wood oi bom nooth < tu o im.i unit i t the chalazal end, the (tide perm copinu , whiti h llu embryo sir; i"1!! oi lightb ur\ "d with < on ! doo thin and il it t li lonci i Ih in th • i idi< U As here interpreted, Ma/yarifana is a relatively- homogeneous genus of 14 species that are all rather closely related. The most distinctive group is Baillon's section Ciccoidcs, with Tcarpellate fruits that do not have a thin, papery endocarp (FIGURE 8). Although M. dubium-traceyi of Aus- tralia has similar fruits, it does not appear to be immediately related to M. anomala of Madagascar and the M; caren 1 In lype of sect. Cic- coidcs. It do 101 i m i pedient i!u ofoi to i< o m/e any subgeneric taxa within Margaritaria.

glabrous, or if hirtellous then not deeply lobed.

Fruits irregularly dehiscent, the endocarp thin, hyaline, papery. JOURNAL OF THE ARNOLD ARBORETUM

Endotesta of seed smooth or nearb /so. 5. Branch tips not spinose. (). liranches glabrous or hirs utulou s but not finely scabridu-

ot basallv C( 7. Petioles adaxiallv char ; leaf blades rounded to acuminate; VOUIIK twigsi glab) • hirsutulous. 8. Carpels mostly 4 o 3 or 6); styles free or nearly so; stipule mm. 1 ong; seeds 2-4 mm. long; leaves usually 1. M. nobilis. S. Carpels mostly 3 (rarely ; ! or 4^ 1; stipules 5-10 mm. long or if shorter then styles leaves rounded to a . . . 2. M. discoidea. 7. Petioles not adaxiallv channeled; leaf blades obtuse or rounded at tip; voting twigs ieddi-L in- 11 uliiu Pels 3 6. liranches usually finely seaibridul tipules lanceolate or- ovate, basally cordate, cuspidate- a.umi nate; carpels 3 or 4.

beneath, stipules pale 3. M. scandens.

pels mostly 4; leave- not glaucous beneath, stipules dark

5. Branch tips slender, spinescent. scabridulous when young; stipules lanceolate, pale; carpels 2. 5. M. hottcana. Endotesta of seed distinctly rugose or ribbed: carpels 3; plants glabrous.

; pistillate [lowers often 2 or 3 per axil 7. M. indica. ninate calyx lobes 3.3-3.7 mm. long; anthers 1.8-2 long; pistillate (lowers solitary

10. Staminate disc obsolete or absent; veinlet reticulum distinct- ly prominulous; carpels 2 01 3 u. 1/ r/iombonhilis. 3. Fruits tardily dehiscent, the endococci not extremely thin and hyaline; leaves mostly rounded to emammate; carpels 2. 12. Leaves and fruits glabrous. 13. Kndotesta ot seed smooth: lea.l venation not prominulous; ant hers 0.S-1.-I mm. long 10. M. aiiomala. 13. Endotesta of seed rugose; leaf venation prominulous; anthers 0.3-0.5 mm. long 11. M. dubium trncrvi. I.' Leaves and I mils hispiduloiis or hirsutulous; leaf venation prom- inulous 12. M. hispidula. 2. Filaments distinctly connate: leaves not over 2 cm. long, rounded to emarginate; carpels 2 or 3, glabrous 13. M. drcaryana. Ovary hutellous ! rarely glabrous), deeply lohed, I or 2 of the 3 carpels often abortive; petioles not adaxiallv channeled; endococci hyaline-papery 14. M. obovata. 6,M.nobilis (Gtu.l, "• " • " I-UI|HHII, nnt < u.. im p., In- 1. n I i„ ex broken away to show paired seed- inside: both >< 5. 7. .1/. discoid^a su it •• 5. S. M. hispid !,i i.lppert 107), longitudinal section of fruit showin a (Mwasumbi LM '• /' i > carp llat< fruit 1 > fruit with third carpel i 412 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

1. Margaritaria nobilis Linnaeus f. Suppl. PL 428. 1781. Phyllanthus nobilis (L. f.) Mueller-Arg. in DC. Prodr. 15(2): 414. 1866. TYPE: Surinam, Dahlbcrg s.n. (LINN). FIGURES 5, 6.

\ia > i '(,<<> '/< I hi ( i I'h pin i /. 7775, ,/?Y 4. 1692. Marzuritaria altcrntiolia L. in Aim. PI. Surinam. 16. 1755, w;mra /7/r^. Cicca antHlana A. juss. Euphorb. Gen. Tent. 10S. /. 4, fig- 13B. 1824. Phyll- anthus ant Minus (A. Juss J Mueller-Arg. Linnaea 32: 51. 1865. Phyllan- thus antillanits ,l pcdic idlaris Mucllei \x\ ibid i'hviianl luts nobilis ailtil- lanus (A. Juss.) Mueller-Arg. in DC. Prodr. 15(2): 415. 1866. Margari- \ ii . / //,;/' i \ Ju^ i ^tchit \ Outntm t atal Fl. Guad. 2: 47. 1937. TYPE: St. Thomas. Ri-dlc s.n. ir-jri. Cicca sia uaincns \iuj Linnaea I '," ! I Chvli has antillanus a con- color Mueller-Arg. Linnaea 32: 51. 1863. TYPE: Surinam. Host maim 622

Cicca pavoniana Baillon, Etude Gen. Euphorb. 618. 1858. Plivllanthus no- bilis s pavoiiiamts i Baillon I Mueller-Arg. in DC. Prodr. 15(2): 415. 1866. TYPE: Peru, Pavon s.n, (p, holotype. n.v.; G, isotype). Cicca ant/tlana vai , pcdiccllaris Cri eb \I in \m Mad. 11. 8: 158. 1860. 1 T',. .' Cuba Faialloncs. \\44yjit 5S4a (GOET, holotype; G, GH, S, isotypes). Phyllanthus nobilis 4 guyaucusis Mueller-Arg. in DC. Prodr. 15(2): 414. 1866. TYPE: French Guyana, Sagot 11:>4 (c.. holotype; r, isotype). Phyllanthus nobilis ~, perirenal us Mueller-Arg. in ibid. TYPE: Peru. Spruce

Phyllanthus nobilis * brasilicnsis Mueller-Arg. in ibid. TYPE: Brazil, Minas Gerais, Clausscn s.n. t c. holotype; A. isotype). / ^ i ' i Vluellei \rg in //;/)/ TYPI Brazil J?/edc/

Plivllanthus nobilis var. fl -w/rtrr/7 Mueller-Arg. Fl. Brasil. 11(2): 70. 1873. TYPE: Brazil. Martius 281.3 < M). Phyllanthus nobilis var. paiiaiucusis Mueller-Arg. //>/)/. TYPE: Panama, £«£- /<»/ //a.vr.v /MP (M, lectotype). Phyllanthus hctcromorpha Rusby. Descr. S. Am. PI. 42. 1920. TYPE: Colom- bia, II. II. A/»//// 1716 I NY, holotype; MO, U, isotypes). Phyllanthus nobilis var. hypouialacus Stamllev, Cartie-ie Inst. Publ. 461: 68. 1955. /'. antillanus var. liv bo inula,:us i Standley > Lmulell. 1'hytologia 1: 337. 1939. TYPE: Mexico. Campeche. Lundcll 897 (F). Celastrus tctraments Standley, Contr. C. S. Natl. Herb. 23: 679. 1923. TYPE: Mexico, Iguala Canyon, Pringle 10319 (us).

Deciduous shrub or tree to 20 meters high; branches glabrous or rare- ly hirtellous, subtercte or angled, becoming prominently lenticellate; fo- liage deciduous (perhaps sometimes evergreen). Leal blades chartaceous, elliptic or oblong to oblanccolate, mostly <) 15 can. long. 2.5-5 cm. broad, acuminate, obtuse to cuneatc at base decurrent on petiole, glabrous or sometimes hirsuu.T << I •- -• . ii along the veins, sometimes glaucous be- neath; major veins mostly 7 to 12 on a side, veinlet reticulum prominent beneath but scarcely prominulous; margins plane; petiole ca. 2-5 mm. long, distinctly channeled adaxially; stipules caducous to more or less 1979] WEBSTER, MARGARITARIA 413 persistent, pale and scarious. elliptic- to oblong-lanceolate, acuminate, 1.5- 3 (-4.5) mm. long. Slaminate (lowers several per axillary cluster; pedicels ca. 2.5-8 nun. long, glabrous; sej)als biseriate, glabrous. ().') 1.7 mm long. 0.7-1.5 mm. broad (outer pair usually narrower); disc 1-2 mm. across; stamens 4, free, hi micm becoming l-; mm Lonj thi anthers elliptic or oblong, 0.5-1.1 mm. long. Pistillate ilowers 1 to several per axil; pedi- cels aibl ret( glabrou becomim? i-12( Id) mm. long; sepals 4, bi- seriate, the inner ones 1.5-2.5 mm. long, 1.5-2.5 mm. broad (outer ones usually narrower); disc 0.8-2.5 mm. across; ovary usually of 4 (65%) or 5 (28%) (rarely 3 or 6) carpels, not deeply lobed. the styles free or basally connate. 1.5-3 mm. long, bifid. Fruits subglobose to oblate, moderately lobed, 7.5-11 mm. in diametei dehis inr irregularly, endo- carps papery; seeds with thick, bluish v:i •, i ai otesia die endoti La smooth, planoconvex, (2-)2.7-4 mm. long, 1.9-3.7 mm. broad.

DISTRIBUTION. Seasonal, rain, or riparian forests, mnstlv at lower ele-

4 mi. X. ol ' La Cuesl .a. McYangh ct a!. 21190 (DAV, MICH). SAN Luis Po- Lw idell & Lundell 7259 America: Belize. C01 s). Guatemala. Pet en, L (Gin. El Salvador. Chalal- Allen US). Honduras. "! Mora 2000 WAN Is.: Grea t Swan, Proctor • 125U ( DAV) . Nicaragua. Volcan San- us). Costa Rica. , Shank & Molina 4199 (F. Panama. Uarn: > Co 0 Is., Foster 1192 MO) . West Indies: Cuba. LAS

^sSv_^te> 414 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

VILLAS: Buenos Aires, Jack 7401 (A). ORIENTE: RIO Yao, Webster 4150 (DAV, GH); Farallones, Wright 584a (GH); Charco de Toro, Wright 1955 (GOET). I'IXAU DEL Rio: Mendoza, Kkmaii 18080 i XY I. Haiti. Morne Birgonne, Ekman 112627 (A). Dominican Republic. Moncion, Ekman H12997 (DAV, S). Puerto Rico. Fa- jardo to Luquillo, Sintenis 1555 (GH). Jamaica. Trov, Harris 8698 (GH). Virgin Islands. ST. THOMAS. Eggers 175 (cm. ST. JOHN, Raunkiaer s.n. (c). Lesser Antilles. ANTIGUA: Blubber Valley, Box 1495 (A). GUADELOUPE: Vieux-Fovt, Webster, Ellis, & Miller 9157 (DAV, MICH, U). DOMINICA: Carholm Estate, Ernst 1949 (DAV, US). MARTINIOUE: Ste.-Luce, Ilahn 9o4 (GH). SABA: Boldingh 2198 (u). ST. LUCIA: Morne la Sorciere, Proctor 18169 (DAV). ST. VINCENT: Lctchwood Valley, Beard 615 (A. NV. UK GRENADA: Union. /Voc/or 17.774 (A, DAV, u). GRENADINES: Mustique. (,. W. Smith 128 ICII). Tobago. Scarborough, Broadway 4045 (on. s, u). Trinidad. Ceclros. Broadway TRIN 7411 (TRIN). South America: Colombia. Puerto lierrio, IIaught 2180 (A, us). Venezuela. Isla Margarita. Johnston 252 (GH); Delta Amacuro. Wurdack & Monachino 59689 (us). Guyana. Kanuku Mts.. ,1. C. .Sw/7// ,?5r). Brazil. SANTA CATAKIXA: Itajai, A'/em 959 (us).

This widespread species is rarely abundant in any one locality, since it appears to belong to successional stages of vegetation. It displays a considerable amount of morphological wiriation, and Mueller in his final treatment (1873) recognized no less than nine varieties, of which seven were confined to various regions of South America. After considerable study of the available specimens, 1 have concluded that it is not profitable to attempt to demarcate any subspecific taxa. since the variation in morphological characters appears to be clinal, without any prominent dis- continuities. Pubescent forms, designated l>\ Standby as /'// vllaiit/ms no- h'tlis v,i f/vponiabieus, occur promi MUHD l\ mixed with glabrous ones. There is a tendency for 3-carpellate fruits to occur more commonly in Mexico and Central America, but they also occur in Paraguay and sporadically elsewhere; 5- and even o carpellale Iruits are found in the Amazon basin and Colombia, but many West Indian specimens are also largely 5-carpellate (TABLES 1, 2). ddie nomenclature and typilicaiion of I bis widespread American species of Margaritaria presents several problems. Linnaeus (in Aim, 1775) de- scribed two species, M. oppositijolia and .1/. allcniijoHa, that are illegiti- mate because the generic name Margaritaria was not validly published at that time. In 17S1 Linnaeus filius assigned a different name, M. nobilis, probably because he combined the two taxa of 1775 (even though he ex- pressed doubt about their conspecificity). It is not evident which speci- men of those in the Linnaean Herbarium was collected by Dahlberg (as cited in the protologue), but I believe that the sheet labeled M. allcnii- jolia can reasonably be taken to be the holotype. Wheeler (1939, a, b) suggested that both the generic name and the specific epithet of Margari- taria nobilis should be rejected because ol tin original inclusion of dis- cordant elements; but under current rules of nomenclature, it appears WEBSTER, MARGARITARIA

Margaritaria discoidea (Baillon) Webster, Jour. Arnold Arb. 48: 311. 1967. Cicca discoidea Baillon, Adansonia I. 1: 85. 1860. Phyll- anthus discoideus (Baillon) Mueller-Arg. Linnaea 32: 51. 1863; in DC. Prodr. 15(2): 416. 1866. TYPE: Gambia, Albreda, Heudelot 102 ype). Shrubs or trees to 20 (rarely 30) meters high; branches subterete or angled, glabrous or hirsutulous when young, becoming prominently len- ticellate; foliage usually deciduous. Leaf blades chartaceous, elliptic or oblong, mostly acute to acuminate at apex, acute to obtuse at base, mostly 3-15 cm. long, 2-4(-5.5) cm. broad, glabrous beneath or sometimes hir- 416 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 sutulous (especially along veins): major veins usually 5 to 12 on a side, raised beneath, veinlet reticulum usually prominent beneath and some- times prominulous, margins plane; petiole ca. 3-7 mm. long, usually dis- tinctly channeled adaxially; stipules caducous (sometimes subpersistent), blunt to acuminate or acute-apiculate, mostly 4-8(-10) mm. long (oc- casionally only 2.5 mm. at first nodes of annual growth). Staminate flow- ers several per cluster; pedicels glabrous or hirsutulous, 2 5( 7) mm long; sepals 4. subequal, the larger (inner) ones O.N 1.5(2) mm. long; disc thin, entire or minutely crenulaie. (0.5 ) 1.0-1.8 mm. across; sta- mens 4, the filaments free or slightly united at base. 0.8-2 mm. long, the anthers ellipsoid, 0.4-1.4 mm. long. Pistillate flowers mostly 1 to 3 per axil; pedicels rather stout, sublerete or angled, becoming (2 )S 8( 14) mm. long; sepals \ subetpial obloii" ti )• ate the outer ones ca. 0.8-1.2 mm. long and broad, the inner ones 1.0-8.3 mm. long; disc rather thick. 1.5-2.7 mm. across; ovary usually of 8 carpels (90'; ). rarely of 2 (\.S% ) or 4 (8.5%), the styles'connate' basalU into a column 0.2-2 mm high, with tips spreading, 0.8 1.5 nun. long, bifid. Capsules subglobose to oblate, scarcely to distinctly (but not deeply) 3-lobed. (5.5-)6-9(-l 1) mm. in diameter, endocoeci thin and papery: seeds with thin to thick sarcotesta, the endotesta smooth, planoconvex. (."' >2.8-3.2(-3.9) mm. long.

DISTRIBUTION'. Decitluous forests or woodlands, sometimes in rainfor- ests, most often in succcssional communities, from sea level to ca. 1000 meters elevation; western to eastern and southeastern Africa (MAP 4).

As noted by Radcliffe-Smith (1976), this widespread African species h o MI ii (f ill (,i - III. m1h\ Mow •'••! in fonti ist to Margaritaria nobilis, it exhibit fahh . ieai differentiation into western and eastern populations that may be conveniently distinguished as subspecies. Al- though there is some oveilap ol characters, most specimens can be rather confidently assign* da cord in;; to th* ha racier: let ailed in the folio wins.1 key:

1.5-6 mm. long; stylar column 0.2 (OSf-l.ll mm. hi-h; endotesta of seed mostly 2.5-8.5 mm Inn;:; leal blade:, mo-ale acute to acuminate at tip. subsp. 2a. discoidea. Stipules mostly 2-> mm. loiiu: (at. distal nodes): larger staminate sepals 1.5- 3 mm. long; anthers 0.8-1.4 nun. IUIIL; liialing nedieob mosth •>' ] 3 mm. long; stylar column 0.7-2 mm. high; endotesta of seed mostly 3.5-4.8 mm. long; leaf blade- mostly obtuse (<> rounded at tip. subsp. 2b. nitida.

Leaf blades mostly acute to a in.be it tip (occasionally obtuse); stipules usually caducous, generally acute to acuminate. (8-) 5-8(-10) mm. long. Staminate flowers with sepals unequal innei (larger) ones 1.1-1.5 1979] WEBSTER, MARGARITARIA

MAP 4. Distribution of Margaritaria in \fric;j olid doli- If discoidca tubsp discoidca hali-hlled Mjiun1., ,1/ discoidca ,ulw. ,////r/\7 In.ci 17 .:,>&• o»

DISTRIBUTION. Seasonal to evergreen rainforests, mostly below 1000 meters; Senegal to Angola. Zaire, and along the Rift Valley in eastern Africa (MAP 4); mainly in secondary vegetation (see Aubreville, 1959). Only a small portion of the specimens >cvn from West Africa are cited

REPRESENTATIVE SPECIMENS. Angola: Cuanza X.. Cazengo, Gossweiler 4692 !•"], \v. i'i Ciimnoon: Ihnimh. /.caper 2sco < p. rsi. Central African Empire: Boukoko, Tisserant 501 (P). Ivory Coast: Rasso, Anbrerille 5o0 (A. I-I. Benin: hoi. ( /« a WMII ( iboti Eibrevilk Klaint 1031 (A P) Gambia: A1- hreda. Hcitdelot 102 (>). Ghana: Asokroa. Kumasi. Darko 595 (MO). Guinea: cntrc Konkoure et Timbo. Chevalier 124^9

. Guinea-Bissau: Fulacunda, Es- pirito Santo 2162 (coi). Liberia: barCavville. Imldwin 11125 (NY). Mali: cercle de Rita, Vuillet 247 n>). Nigeria: [jebu. Uluasogo. Ross 62 (MO, NY). Senegal: Thics-Diourbel, Raolak. Berhaitt 1091 (p). Sierra Leone: Njala. Dcighton 1109 (MO). Sao Tome & Principe: Principe. Espinto Santo 221 (cm); Sao Tome. £/r/7o Santo 3S50 (cop. Sudan: Equatoria. Torit Rirt. Andrews 1740 (K). Togo: Tamberma. Kvrslinx 545 (A). Uganda: Ripayo. Bummer 567 (MO, IS); Teso, Serere, Chandler 1065 (PRE). Zaire. EQPATEUR: Lukolela, C//fl/>/« 595 (us). Rive: Ralehe. /We/ 729 (PR). Troupin 12520. 12595 (HR); Lueki, Z?o;w'e 20

2b. Margaritaria discoidea subsp. nitida (Pax) Webster, comb, et stat. nov. Fluei^ca nitida Pax. Hot. Jahrb. 19: 76. 1894. TYPE: Mozambique. Ouilimane, Stuhtinann 550 (n.v.; the original speci- men at B presumably having been destroyed. I hereby designate as neotype the following collection: Mozambique, entre Xampevo e Mugeba, Grandvaux Barbosa & Carvalho 2950 (K)). 1979] WEBSTER, MARGARITARIA 419

I h-t ca fa folia Pax IMlanzcnw Ost.-Afr. C, 236. 1895. TYPE: Tanzania, Kilimanjaro, Yolkens 1737 (K. lectotype). PhyllantJius ? amapondmsis Sim Fores! Forest Fl Colony Cape Good Hope, 325. t. 141, fig. 2. 1907. TYPE: South Africa, Pondoland. Egossa Forest. Sim 2608 (PRE, holotype). Leaf blades mostly obtuse or Milnuite at tip. occasionally rounded or emarginate, rare!) acuminatt Lipule caducou oi pet istent ! » >( 6) mm. long; inner (larg r) taminate epah (1.5—) 1.8—2.5(—3) mm. long; staminate disc 0.5-1.1 (-1.5) mm. across; anthers 0.8-1.4 mm. long; pistillate flowers with pt-di 4s becoming (5 )6— 10(—13) mm. long; ovary nearly always of 3 carpels (97G ), very rarely of 2 or 4 (1.5% each), styles usually connate ! . to :;, their lengths, the column 0.8-2 mm. high; capsules (9-) 10-13 mm. in diameter; endotesta of seeds (3.5-)3.8-4.8 mm. long.

DISTRIBUTION. Burundi and southwestern Ethiopia south to Rhodesia and South Africa (MAP 4); a large number of additional specimens are cited by Radcliffe-Smith (1976).

SPECIMENS EXAMINED. Burundi: Bururi. Kigwena. Reekmans 3193 (PRE). Ethiopia: S. Sidamo. Chaffex 1 MK) Kenya. CEXTRAT Nyeri. Porter 933 (PRE). COAST: Kilili, Sokoke, Gisan SOK6 (PRE). NAIROBI: Nairobi. National Museum grounds. Mwangangi 17 63 (Cm); Coryndon Museum grounds. Will'nuns 127 (K, PRE); Chirorno ! e , S / i \i > Gillcll 1650 (MO) 'A I'F.R> Kakamega 1 ; o i i /'/ hiu in oiid < //(/»s/n ' (i ki i Mil. . ( i IR\L Salima, Rob- son & Steele 1613 (PRI > NORTHERN Chendo Rivei between For! Till mi ( In i i / I i ill 6' (PRI hil pa >ist Kaseye Pawel I 159 (D\\ > Karonga Disti mi \\ oi I ime i Pa 1 i 11 (mvv) Mzimba Disti 1F4 mi. SE of Mzamba2 \h ion /'. u h 11960 (DAV) Nkhata Bay Distr., road to Nkhata Bay from Alzuzu, Paivrk 130-16 {DAY), Mozambique. GAZA: Mana ; Martinhn do bilen* Grandvaux Barbosa S33S (coi); entre Chissano c Lieilo, Grandvaux Barbosa & Lemos 7070 (coj. K). IXHAMBAXE: Sul do Save, Xhacoongo Maccdo & tialsnilias 1101 (PRI I; Bazaruto E . Font Estone. Mogg 28639 (PRE). MAPUTO; Muntanhane, Balsinhas 237 (PRE), Eourengo Marques,

Borle 174 (PRE) Ilombx js \o < u L i l'< h \ M i Floo la do Licauti, Lemos & Palsinhas _ > (PRI > ZAMBEZIA end Nampevo e Mugeba, Grandvaux Barbosa & Carvalho 2300 (K); Ouelimane, Fugel M ruba i lagoa Plantation / a nil i IJ < i Miu.l"-i! 'I i \i xn Chipinga Distr.: Makosa bids, /7/e/».v //n / /'A '", / ( i i \l \SIION4LAND SOUTH. Mrewa Disti lia\vnov< Rivei bridge We// W> (XY) MATABELELAND. Se- bungwe Distr.: Kariangwe Hill, Lovemore 175 (NY, PRE) Suainga, Lovemorc 420 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

487 (PRE); Victoria Falls, Wilson s.n. CO. Matopos Distr.: Moth Shrine Koppie, Plowe 1320 (NY); Pesna Kohila, Miller 1901 (MO. PRE); Ouaringa, Miller 2522, 3295 (PRE). VICTORIA. Chibi Distr.: Mad/hare Dip. Moll 385 (NY). South Africa. CAPE OF GOOD HOPE: Pondoland. Eim^i ! nn -\ s »; 2oOS\ 10618 (PRE). NATAL: Hlabisa. Ward 2677. 2S10 (PRE); Hluhluwe Came Reserve, Codd 9627 (PRE); Fakude 20 (PRE). /V-e/ /02 ( PRE I ; Mbizatsheni. Ward 6058 (PRE); Wall's farm, Gerstner 500o i I'KEI. ln-wax uma PiOr.: Maputa. de Winter & Vahrmei- jer S601 (PRE), 1(> mi horn Xdumu toward Ingwavuma, .\loil '1353 IPKEI; con- fluence Pongola I MI '' oi v i ii dumu urn • er\ /VV I !/<

The status of the eastern African populations of Macgaritaria discoidea remains controversial Radclifte Smith ( PCo). while recognizing JM. obo- vata at varietal rank, grouped all other forms of eastern African Margari- taria in M. discoidea. However, although there are overlaps of measure- ment in most characters in llu eastern and western African populations, I believe that they are modally distinctive enormh to merit separation at the subspecific level Specimens somewhat intermediate between subsps. discoidea and nitida are known from Sudan, Luanda, and Zaire. How- ever, the sampling of collections from these areas is still not very good, so the position of the boundary line or zone between the two subspecies is not yet well defined.

3. Margaritaria scandens (Wrighl ex Griseb.) Webster, Jour. Arnold Arb. 38: 66. 1957. Cicca antillana var. a glauccscens Griseb. Mem.

Am. Acad. II. 8: 157. 1860. Phyllanthus antillanus t glauccscens (Griseb.) Mueller-Arg. Linnaea 32: 51. 1863. Cicca scandens Wright ex Griseb. Nachr. Ges. Wiss. Gottingen 1865: 165. 1865. Phyllanthus scandens (Griseb.) Mueller-Arg. in DC. Prodr. 15(2): IPS I am his I? , s vi indt is (Griseb.) O. Kuntze, Rev. Gen. PI. 2: 601. 1891. TYPE: Cuba, Oriente. Monteverde. Wright 1437 (GOET, holotype). FIGURES 1, 11. JSTKR. MARGARITARJ a

IMCI-KKS 11-13. Flowers of Mar^aritar tillatc; 12, M. hotteana (Ekman IIS909) H8919). pistillate; all X 7.5.

I'hylhintSiiis baJiamensis I'rluu. Symb. Antill. v 27'!. piO.'g Margaritaria ba- hamensis (Urban) Britton & Millsp. Bahama Fl. 220. 1920. LECTOTYPE: Andros. Red Bays, Northrop s.n. (GH, lectotype!

Straggling or somewhat sca.ndcnt. shrub or small tree up to 5 meters high; l>ranchc cabridulou when counii reddi h md becoming more or less smooth in age; foliage evergreen, or partly deciduous. Leaf blades firmly chartaceous Nipti< m >b< tt< or lanceo] u m th / cm loim 1.5-2.5 cm. broad, obtuse or bluntly pointed al tip. cuneate at base, de- current on petiole, often glaucous beneath; major veins ca. 7 to 12 on a side, not conspicuous above, beneath raised, the veinlet reticulum often prominent but not proininulou margins pi in." m recurved; petioles 3-7 mm. long, distinc I laxial hanneled ail i lanceolate scari- ous, becoming more or less indurate and subpersistent, cordate, acuminate, (1—) 1.5—2.5(—3.5) mm. long. Staminate flowers few to several per axil (rarely solitary): pedicels stout, 2-5 mm. long; sepals biseriate, ovate to elliptic, the outer pair 1—1.7 mm. long. 1—1.4 mm. broad, the inner pair 1.4-2.1 mm. long, 1.2-1.9 mm. broad; disc thickish. 1.2-2.1 mm. across; stamens 4, free, the filaments becoming 0.5-1.3 (-2) mm. long, the anthers elliptic, 0.5—0.7(—1) mm. long. Pistillate flowers 1 to 3 per axil; pedicels subterete, (1.5-)3-7(-9) mm. long; sepals 4, biseriate, subequal, 1.5- 2.3 mm. long and broad: disc thick, 2.3-3 mm. across; ovary of 3 (less commonly 4) carpels, the styles free, spreading, 1-1.5 mm. long, bifid. Fruits oblate, shallowly 3-lobed, 7 10 mm. in diameter, irregularly de- hiscent, endococci i p< eed with thicl blu gi n ireotesta, the endo- testa smooth, plano-convex. 2.7 4.2 mm. long, 2.4-3.5 mm. broad

DISTRIBUTION Ham mod pine forests mid evergreen forests; Baha- mas and Cuba (MAP 2).

SPE< [MEXS EXAMINED. Bahamas. ANDROS. N. Andros: Mangrove Cay, Brace

4Sn, (N\i - JI fioll town (one!! 77 r.vaas 43929 C\Yl; Condi Sound, Strachan Hill, Brace 6924 (NY); road to Conch Sound, Brace 67S4 (NY); road to Louisa Coppice, Brace 6270 (A); 2 mi. NVY, of Love Hill, ///// 7027. 7035 (NY) ; Mastic 422 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

Point. Brace 6983 (NY); Crow Hill. Small & Carter 8720 (NY); Smith Hill, Small & Carter 8678 (NY); 1 mi. N. of Smith's Hill. Correll 43522 (NY). ELEUTHERA: Spanish Wells, Coker 328 (NY); The lUutl, Russell 1 IDA\ 0. GREAT ABACO: road to Hole-in-the-wall, 6 mi. below intersection with Sandy Point Rd., Correll & Meyer 44559 (NY); California Road, Brace 2037 (NY). GREAT EX- UMA: Stuart Manor, Correll 44107 (NY). NEW PROVIDENCE: Nassau, Brace 3906, (NY), Curtiss 183 (A, NY); Farringdon Road, E. G. Britton 3043 (NY); Fort Charlotte Rd., Brace 7147 (NY); Maiden-hair Coppice. Wilson 8394 (NY), E. G. Britton 3435 (NY); be! ween Love and Gambier. Degener 19140 (A, NY); Water- loo, Britton & Brace 741 (NY), Degener 19139 (A, NY); Blue Hills, between Lake Cunningham and Lake Killarney, Webster, Samuel., 14 Williams 10514 (DAY). Cuba. LA HAUANA; Isle of fines. Jennings t\'s (NY). ORIENTE: Mt. Libano, 1S60, Wright s.n. (GOET); La Pcrla. Montcverde. If'r/'gA/ J4J7 (GOET, holotype; BR, F, G, GH, NY, S, W, isotypes).

Several barren specimens from Oriente Province. Cuba, are distinctive in their thick, levolute leaves with a thick, waxy, glaucous bloom beneath: Lomas de Duaba, near Baracoa, Ekman 4236 (s); Moa. La Brena, Leon & Clemente 23339 (MICH); Toa, charrascos de Pena Prieta, Alain 3481 (DAY): Pico Galano, Toa, Alain 3761 (DAV). These may represent a form ol C • u '« ia > w/ /.\ oi [irrha) I a idesciibed species. There is still some uncertainty as to the application of the name Mar- garitaria scandens because the species has not been re-collected in Cuba since Wright's original gatherings. From these incomplete specimens, it is difficult to be absolutely certain that the Bahama plants are conspecific. However, until the Cuban population can be restudied, it seems best to treat M. bahamensis as a synonym of M. scandens. Unfortunately, I had earlier (Jour. Arnold Arb. 39: 208. 1958) mistakenly treated M. baha- mensis as a synonym of M. tctracocca rather than of M. scandens.

4. Margaritaria tetracocca (Baillon) Webster, Jour. Arnold Arb. 38: 66. 1957. Wurtzia tctracocca Bailh.n, Aclansonia T. 1: 187. 1861. TYPE: Cuba, Oriente, Wright 583 (G, holotype; GOET, isotype).

Cicca antillana var. p virens Griseb. Mem. Am. Acad. II. 8: 158. 1S60. Phyll- anthus antillanus p virens (Griseb.) Mueller-Arg. Limiaea 32: 51. 1863. Cicca virens (Gn I i fright. Griseb Nachi Ges Wiss Gottingen 1865: 166. 1865 Phylh •!> < n-vs ( Gri eb ) Mueller-Arg. in DC. Prodr. 15(2): 'i i lid / sp >us nfis (( I.IM (.). Kuntze. Rev, Gen. PI. 2: 601. 1891. TYPE: Cuba, Oriente. Wright 584 (GOET, lecfolype; <;. cap s, isotypes).

Shrub to 3 meters high; branches subterete, scabridulous at least when young; foliage evergreen. Leaf blades firmly chartaceous, elliptic or ob- long, 3-9 cm. long, 1.5-3.5 cm. broad, obtuse or bluntly pointed at tip, cuneate at base, decurrent on petiole, shining above, paler but not glau- cous beneath; major veins ca. 7 to 12 on a side, inconspicuous above, raised beneath, veinlet reticulum rather conspicuous beneath but not very prominulous; margins more or less revolute; petioles 4-7 mm. long, adaxially channeled; stipules ovate, basally auriculate, acuminate, hardly scarious, becoming dark and indurate, 1.5-2.7 mm. long. Staminate flow- 1979] WEBSTER, MARGARITARIA 423 ers several per cluster; pedicels stout, 2-3 mm. long; sepals biseriate, 1.8-2.5 mm. long, the inner ones ca. 1.7-2.5 mm. broad (outer ones nar- rower) ; disc obsolete; stamens 4, free, the filaments 0.5-0.8 mm. long, the anthers oblong, 1-1.3 mm. long. Pistillate flowers solitary; pedicels subterete, scabridulous, 2.5-5.5 mm. long; sepals 4, biseriate, subequal, ovate, 1.5-2.3 mm. long, 1.2-2.1 mm. broad; disc rather thin, 1.5-1.8 mm. across; ovary of 4 (rarely 3) carpels, apically indented, the styles 1.5-2 mm. long, connate basally into a column 0.4-0.7 mm. high, bifid. Fruits oblate, shallowly to rather deeply 4-lobed, apically depressed, 9-10 mm. in diameter; seeds apparently with a thin sarcotesta, the endotesta smooth, plano-convex, 2.7-3.2 mm. long, 2.7-2.9 mm. broad. DISTRIBUTION. Endemic to montane forests; Cuba and Hispaniola (MAP 2). SPECIMENS EXAMINED. Cuba. ORIENTE: "Cuba Orientali," 1856-57, Wright 583 (G, GOET); Monteverde, 1859 60 II right 584 ((.. Gi-i GOET); Pinal de Santa Ana, Eggers 5026 (A). PINAR DEL RIO: Pan de Guajaibon, Acuna & Alain 18549, 18550 (HAC) [lain & icuna '.999 (us) Haiti. SUD [assif d< 1 Hotti central group, St. Louis du Sud. Bonnet-Carre, ca. 1150 m. alt., Ekman 119234 (s). Margaritaria tetracocca is still a poorly known species, but it appears to be quite distinct from the much more common and more widely dis- tributed M. scandens. The absence of a glaucous bloom on the underside of the leaves in M. tetracocca is characteristic, although some forms of M. scandens also lack this character. However, the stipules of M. tetracocca are consistently different in their auriculate base and indurate texture, so even barren specimens can be distinguished from M. scandens. The specimens from Pinar del Rio and from Haiti are somewhat di- vergent from the type collection, and the Haitian plant in particular may — when better known — prove to be subspecifically or even spe- cifically distinct.

5. Margaritaria hotteana (Urban & Ekman) Webster, Jour. Arnold Arb. 38: 66. 1957. Phyllanthus hotteanus Urban & Ekman, Arkiv Bot. 22A(8) : 61. 1929. TYPE: Haiti, Ekman H8909 (s, holotype; G, isotype). FIGURES 12, 13. Low, straggling shrub; twigs slender, scabridulous when young, lenti- cellate in age, some twig tips becoming spinescent. Leaf blades thinly chartaceous, elliptic or oblong to obovate, 1-2.5 cm. long, 0.4-0.8 cm. broad, obtuse or rounded (and sometimes minutely apiculate) at tip, cuneate at base, decurrent on petiole, slightly shiny above, green and paler beneath; veins obscure above, midrib prominently raised beneath, major veins 5 to 10 on a side, veinlet reticulum fine and not prominulous; margins plane or slightly recurved; petioles 1-3 mm. long, adaxially channeled; stipules lanceolate, scarious, pale, mostly deciduous, 0.7-2 mm. long. Staminate flowers 1 to 3 per axil; pedicels slender, 3-6 mm. long; 424 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 sepals biseriate, oblong to obovate. 1.3-2 mm. long, 1.0-1.7 mm. broad across; stamens 4, free, the filaments 1 1.5 mm. long, the anthers elliptic, 0.5 0.6 mm. long. Pistillate flowers solitary; pedicels subterete, 2-3 mm. long; sepals 4, biseriate, the outer ones ovate, the inner ones oblong-elliptic, 1.7-1.8 mm. long, 1.3-1.7 mm. broad; disc 4-lobed. ca. 1.5 mm. across; ovary of 2 carpels, the steles nearlv free, spreading, somewhat dilated, bifid, 0.7-1 mm. long. Fruits not seen. DISTRIBUTION. Endemic to forests on limestone: southern Haiti (MAP

SPECIMENS EXAMINED. Haiti. Sen: Massif de la Untie Morne Rochelois, Miragoane, towards Charlier, 14 Aug. 192 7. Ekman IlSQOQ (G, s); Grande- Cai'mite, Source-Fantasque. 20 Aug. 192 7, Ekman HS010 (S); St.-Michel de l'Atalaye. Morne la Cidre, limestone crags, ca. 750 m„ Ekman H8364 is). As noted by Ekman (ins.), the habit of this species is very similar to that of Fluci^ca aridoton; however, despite the lack of fruits, the floral disc clearly indicates that the plant belongs to Margaritaria. The sca- bridulous twigs suggest an affinity to M. scandens and .17. tctracocca. but it differs from those species in its slender, spinescent branch tips, smaller leaves, and bicarpellate ovary.

6. Margaritaria luzoniensis (Merr.) Airy Shaw, Kew Bull. 20: 387. 1966. Phyllanthus luzoniensis Merr. Philip. Jour. Sci. Bot. 7: 404. ( I ) 12. /' ion in >nii isis ( Men". I 'in h * KCM »nll h ! 1963. TYPE: Luzon, Pampanga Prov., Cunan FB 17698 (A, lecto- type). FIGURE 4. Tree to ca. 10 meters high; foliage deciduous; twigs subterete, reddish- hirsutulous when young, glabrate in age. leiilicellate. Leaf blades thinly chartaceous, elliptic to obovate, (2-)2.5-5 cm. long, 1-2.5 cm. broad, obtuse to rounded at apex, cuneate at base, slightly decurrent on the petiole above, beneath paler and hirsutulous when young (glabrate in age); veins (5 to 8 on a side) and veinlets anastomosing in a prominent reticulum, prominulous especially beneath; margins plane; petiole most- ly 2-4 mm. long u htl\ latei ll\ Hanged but not idaxially channeled; stipules caducous, browni h carious, lanceolate, hirtellous, ca. 1-1.5 mm. long. Staminatc flowers several pei a ill; « duster; pedicels 4-6(-8) mm. long, hirsutulous; sepals biseriate, hirsutulous, the outer ones ovate, 1.3- 1.6 mm. long, 0.8-1 mm. broad the ion a ones obovate. 1.6-2 mm. long, 1.1-1.5 mm. broad; disc thin, 0.5-0.9 mm. across; stamens 4, the fila- ments free, becoming 0.8-1.5 mm. long, the anthers elliptic, 0.6-0.7 mm. long. Pistillate flo\.,< i . olu in pediu i ulueiete lin utulous, 8-17 mm. long; calyx and ovary not seen in flower. Fruits subglobose, 6-10 mm. in ii meh carped, inell; more or less persistent; seeds with dis- tinct dark sarcotesta, the endotesta smooth, plano-convex, 4.1-5.1 mm. long, 2.6-3.1 mm. broad. 1979] WEBSTER, MARGARITARIA 425

DISTRIBUTION. Dry forests, lower elevations: endemic to Luzon.

SPECIMENS EXAMINED. Philippines. Luzon. BATAAN : Corregidor Is.. Curran FB 13225 (A, US). XCEVA ECITA: Mt. l.'mingan. Ramos & Edano BS 26282 (us). PAMPANGA: Mt. Arayat. Curran FB 17698 (A. lectotype). PANGASINAN: Umingan, Otancs BS 17834 (A, US).

Airy Shaw (1971) has questioned the distinctness of Margaritaria lu- zonicnsis from M. indie a on the basis of certain specimens of the latter that have small fruits. Actually, in the one collection (Ramos & Edano BS 2628.1) of .1/. iuzoninisis with mature fruit, the seeds are larger than those of M. indica. A better seed character is provided by the surface of the endotesta, which is smooth in .1/. luzoniensis and at least somewhat (often pronouncedly) rugose in M. indica. The small, blunt leaves with non- channeled petiole (FIGURE 4) and reddish pubescence of M. luzoniensis make it easily recognizable even in the barren condition, and there ap- pears to be no reason nor to give it specific rank.

7. Margaritaria indica (Dalz.) Airy Shaw, Kew Bull. 20: 387. 1966. Prosorus indicus Dalz. Jour. Bot. Kew Misc. 4: 345. 1852. Phyll- anfhus indicus (Dalz ) Mueller-Arg. Linnaea 32 : 52. 1863; in DC. Prodr. 15(2): 417. 1866. TYPE: India, Deccan, Dalzell s.n. (K).

I'liyllanthus stocksii Mueller-Arp. Linnaea 32: 151. 1863. TYPE India, Stocks & Law s.n. (G). Ciilococais siuidaicns Kurz ex Tensive & Binn. \at. Tijdschr. Nederl. Ind. 27: is l e,4 I'livUautluis MI,HU « I m kieller-A.-' m DC Prodr. 15(2): 1272 IS66 D spams smuhih iKurzi 0. Kuntzc. Rev. Gen. PI. 2: 599. 1891. TYPE: "in freto undaico I sm nn u (holotype, presumably BO, n.v.; G. isotype). Phyllanthus indicus f. vestita J. J. Sm. Bijdr. Booms. Java 12: 87. 1910. TYPE: Java. G. Watoedoclol. Koordars s.n. (BO, n.v.).

Glabrous trees up to 25 meters high; foliage deciduous; twigs brown- ish, more o le- prominenth lentieell, » ntjcompre t 1 Lent hPdi thinly chartaceous, elliptic to oblong-lanceolate, ca. 5-13 cm. long, 3-6 cm. broad, distinctly and ometime abruptly acuminate at tip, cuneate to rounded at base and more or less adaxially decurrent on the petiole, usually paler beneath; veins (mostly 8 to 12 on a side) and veinlets prominulous beneath; margins plane or incurved; petiole mostly 5-8 mm. long, adaxially channeled; stipules caducous, brownish-scarious, lanceo- late, more or less acuminate, 2.5-4.5 mm. long. Staminate flowers several per axillary cluster; pedicels ca. 4-6 mm. long; sepals 4, biseriate, the outer ones ovate, 1-1.5 mm. long, 0.6-1 mm. broad, the inner ones obo- vate, 1.3-1.8 mm. long, 1-1.5 mm. broad: disc adnate. smooth or some- what rugulose, 0.6-1.5 mm. across; stamens 4, the filaments free, be- coming 0.7-1.5 mm. long, the anthers elliptic or oblong, 0.6-0.9 mm. long. Pistillate flowers 1 to 3 per axil; pedicels subterete, 8-21 mm. long; sepals 4, biseriate, ovate to oblong, entire, ca. 1.5-2 mm. long; disc entire, ad- nate, 1.8-2.8 mm. across; ovary almost always of 3 (very rarely 4) car- JOURNAL OF THE ARNOLD ARBORETUM pels, the styles free or basalb cmnuu'. spifading, ca. 1.5-2 mm. Ions, bifid. Fruits sub-globose, scarcely lobed, irregularly dehiscent, (7—)9—12 mm. in diameter, endococci papery; seeds with rather thickish, bluish sarcotesta, the endotesta usually distinctly rugose or furrowed on back, plano-convex, (3.5-)3.9-4.S mm. long. (2.7-)3-3.7(-4.2) mm. broad.

DISTRIBUTION. Mostly in lowland (below 1000 meters) deciduous or semi-evergreen monsoon forests; India and Ceylon to Thailand, Formo- sa, Indochina, the Philippines. Indonesia, and \ustralia (.MAP 5). Addi- tional localities in Java are cited by J. J. Smith (1910). and other oriental localities are given by Airy Shaw (1963, 1976).

SPECIMENS EXAMINED. India. Kanara, Talbot 1173 (K). MYSORE: 20 mi. S. of Belgaum, Ritchie 1343 (on); Hassan Distr.. Vettinahalla, Saldanha 13263 (K), Nicolson et al. HFP SO (MO); Shiradi ghat, Saldanha 13140 (MO). Sri Lanka: Nannapurawa. Moneragala Distr.. Unas 003 (MO). Burma: Bassein Distr., Chaungtha Is., Lace 3034 (K). Thailand: Chaiyaphum Distr., van Beu- sekom et al. 4207 (K. MO); Lampang. Wans; Fen. II mil 1057 (K); Saraburi. Phu Kae, Chamroerngsri 2 (K). Vietnam. TONKIN: Dam-ha. Tsang 30219 (A, S); Ha-coi, Tsang 295S3 (A, S). Malaysia. PAHANG: I'ulau C'hibeh. P. Tioman. Corner SFN 29S29 (A). Indonesia. SUMATRA: without locality. Forbes 2733 (GH). JAVA: Semarang Prov., hoarders 2n47d (,\i; Semarang, Kedoeng djati, Koorders 25521 (WAG). MOLUCCAS: Soela, P. Mmgoii, Koorders 29901 (A); Tanimbar, Otimmei Kooida , < ' I'h lr.pi I i < > < Prov., Abulug, Alejandro FB 2743S (A. US); Prov. Oamarines. Alvarez FB 22655 (A, US). MINDANAO; Agiisan suhprov.. Miranda /•/>' 22943 (A, US); Butuan sub- prov., Weber 1195 (us). PALAWAN: Palawan Is.. Cenabre et al. FB 28004 (A), Elmer 13132 (GH), Fenix BS 155S9 (A. US), Natividad 25361 (us); Puerto Princesa, Cenabre FB 20209, FB 20210 (A); Inahit, Yidal 1729 (A). SULU: Ta- wilawi. iOunos Or F.dauo I'S h!5i>3 (A).

mm

MAP 5 Disti ihul ion ui l/8) reported the species from China (Kwantung), Borneo (Sarawak), and the Kangean Islands; and more recently (1971) he has extended the range to Xew Guinea. Mar^arilana iinliea is very closely related to the Vfrican M. discoidca, from which it cannot be surely distinguished excepl by virtue of the ru- gose endotesta of the seed. Airy Shaw (1971) notes that some Philippine specimens suggest possible intergradation with .17. lir.onicnsis; however, the available sampling of specimens is still far too scanty, and field in- vestigations will In in to come to am conclusions about changes

Margaritaria cyanosperma (Gaertner) Airy Shaw. Kew Bull. 20: 387. 1966. Croton ? cyanospermus Gaertner. Fruct. Semin. PI. 2: 120. pi. 107. 1791. Prosorus ^aertneri Thwaites. London Jour. Bot. 8: 272. 1856. Prosorus cyanospermus (Gaertner) Thwaites. Enum. PI. Zeyl. 281. 1861. PhyUanthus ( vanospermus (Gaertner) Mueller- Arg. Linnaea 32: 51. 1863; in DC. ITodr. 15(2) : 416. 1866. TYPE, n.v.; neotype: Ceylon, Thwaites CP 2601 (PDA, holotype, n.v.). /\ m/> ,,„/'/// zeyla•nirtun Thwaite i \ fiaillon dc Gen. Luphorb. 620. pi. 27. fig. 11. TYPE : Ceylon. Twites (r Glabrous tree: foliage deciduous; twigs brownish. Leaf blades thinly chartaceous, elliptic, ca. 5-9 cm. long, 2-4 cm. broad, cuspidate-acuminate at tip, cuneate at base; veins (ca. 5 to 10 on a side) and veinlets prominulom margins plane pliol I ; mm Inn id ixially channeled; stipules deciduous, lanceolate, scarinus. acuminate, ca. 3-4 mm. long. Staminate flowers several per cluster; pedicels mostly 8-10 mm. long; sepals 4, biseriate. reticulate veined, the outer ones oblong, ca.. 8.5-i- nun. long. 1.4-1.8 mm. broad, the inner ones obovate, 3.5-4 mm. long, 2.2- 2.7 mm. broad; disc rather tenuous. 0.8-1 mm. across; stamens 4. the filaments free, 0.5-0.7 mm. long, the anthers linear-oblong, 1.8-2 mm. long. Pistillate flowers solitary; pedicels slender, 15-17 mm. long. 2,2-8 mm. but id

DISTRIBUTION. Endemic to Sri Lanka. SPECIMENS EXAMINED. Sri Lanka: without specific locality, Thwaites CP 2155 428 JOURNAL OK THE ARNOLD ARBORETUM [VOL. 60

CO. CP 2601 (GII, LI. Tluvaites i IS'di cited AmUmammva and Ralnapour dis- tricts, but the species seems not to have been re-collected in recent times. Although the rugose endotesta of the seed suggests an affinity with Margaritaria hidica, the large slaminate flowers set .1/. cyanospenna apart from all of the other species in the genus. Unfortunately, the species re- mains poorly known, and the typification of its name remains somewhat doubtful. It is impossible to tell from Gacrtner's description, based on the fruit alone, which of the two specie:; of Marian/ana on Sri Lanka should take the epithet cyanospenna. According to Dr. \V. Frey, of Universitat Tubingen (pers. comm.), the "type material" of Gaertncr's is missing and presumed lost. However, since Thwaiies (ISM,) cleailv distinguished the two species, 1 am (in the absence of any contrary evidence) following his usage in assignment of names and am designating the specimen cited by him as neotype.

9. Margaritaria rhomboidalis (Baillon) Webster, comb. nov. Cicca ? rhomboidalis baillon. Adansonia 1. 2: 51. 1861. Phyllantlms / rhomboidalis (Baillon) Muellcr-Arg. Linnaea 32: 52. 1863, in DC. Prodr. 15(2): 417. 1866; Leandri. Not. Syst. Paris 6: 199. fig. V. PUS. [d. Madagascar 111(1): 97. fig. XV, 6-11. 1958. TYPE: Madagascar. Ambongo. Pcrvillc oo / (p, lectotype). FIGURE 3. Shrub or small tree ca. 3-8 meters high; twigs pale, subterete; foliage deciduous. Leaf bladi chartaceou to ubc< iaceou illiptic to obovate, ca. 2—5(—7) cm. long. 1.5-3 cm. broad, mostly obtuse to acute at tip, cuneate and often inequilateral at base, usually greenish on both sides (sometimes more or less glaucous beneath) ; veins (ca. 7 to 12 on a side) and veinlets conspicuous, pruminulnus especially beneath; margins plane or slightly reflexed; petioles 3- 7(-13) mm. long, plane to adaxially chan- neled; stipules caducous, brownish, scarinus, oblong-lanceolate, 1.2-4 mm. long. Staminate flowers several per axillary cluster; pedicels 3-5 mm. long; sepals 4, biseriaie, ovate, entire, 1.8-2.5 mm. long, 1.3-1.7 mm. broad; disc obscure or absent; stamens 4. the filaments free, 0.5-1.2 mm. long, the anthers oblong, (0.8- ) 1-1.4 mm. long. Pistillate flowers solitary; pedicels compressed-angled, becoming 4-12(-20) mm. long; sepals 4, biseriate, ovate, entire, 2-3.2 mm. long; disc obscure or absent; ovary of 2 or 3 carpels, the styles usually basally connate, column 0.8-1.2 mm. high, the tips bifid and recurving. 0.5 1 mm. long. Capsules subglobose, irregularly dehiscent, mostly 8.5—10(—12) mm. in diameter, the endococci thin and papery, the columella more oi less pemistent. ca. 4-5 mm. long; seeds with thickish in ite ta (whiti h on drying), th« endotesta more or less ribbed or rugose, 2.8-4.5 mm. lonm i:m hilum •-. arying in position from adaxial to abaxial, ca 0.5-0.8 mm. across. DlSTI UBUTION. Endemic to Madagascar, mostly in decidi : 300 meters or below (MAP 6). SPECIIJENS EXAMINED. Madagascar. 1 IWARAX TSOA: bassindel 1979] WEBSTER,A MARGARITARIA 429 / >' ! / ). /

a Test d'lhosy aux environs d'Ambinda, Capuron < 11 " 1 11623 (p); Ankirihitra, pres du mont Tsitondroina, Perrier de la Bathie 1346 (p); foret de Belambo, nord de Marohala, 5F K>2J,V (P). AIAJU.M.A IOK-I d< L umpihy [Tsiempihy], Leandri, Capuron, & Razufuidrakoto 2195 (i >; lAntsingy, vers Ambodiriana, r Leandri, Capuron, fr Ra:>> >i >,!,„!,,>, <> -> • v i i> > / „,>di < r „ ' , ,,/;/ T 'M ,'

10. Margaritaria anomala (Baillon) Fosberg, Kew Bull. 33: 185. 1978. Cicca anomala Baillon, £tude Gen. Euphorb. 619. 1858. Phyllanthus anomalus (Baillon) Mueller-Arg. Linnaea 32: 52. 1863; in DC. Prodr. 15(2): 418. 1866. TYPE: "Madagascar," du Petit-Thouars s.n. (p, holotype). 430 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

Flneggca major Baillon. Etude Gen. Euphorb 593 IS.vS \dansonia I. 2: 42. 186L Phylhv hiis hys, ranlhtis Uuellei \i Linn ii '" ' IS(n i - <> men superjl). TYPE: .Mauritius. Bouton s.n. (p. holotype). Flneggca •' eglandulosa Baillon. Etude Gen. Euphorb. 593. 1S5S. Pliyllanthus xihn vloid LuelU'r-Arg in I)i IVodr ISI'I 41S 1S66 Leandri, Id. Madagascar 111(1): 94. fig. AT, 1-5. 195,3. Fhyllanlhns cglandulosus ( liail- p lon) Leandri ( ml PI M;i(i i|i 1 /i\'ll ithu IO mains subsp. cry , o.wloid i v] m-ll.-r-Arg l Leandri. Not. Syst. Paris 6: 191. 19 I i I nl i' i < \inl>un,enj Lew Bull. 33: 1S5. 1978. TYPE: Aldabra, Abbott s.n. (K, lectotype; us. isotype).

Glabrous shrub or small tree I -6 meters high; bark scaly or in strips; foliage partly oi \ lioll id nm in I . » UH ii m< ill ul i i t< m compressed. Leaf blades thinly chartaceous, elliptic to obovate or spathu- late, (2—)3—6(—10) cm. long, 1.5-1 ( -(>) cm. broad, rounded or emarginate to obtuse or bluntly cuspidate at tip, cuneate at base (more or less decur- rent on petiole), u IUIK palei ( urmetimes glau ou ) heneath; veins and veinlets obscure to prominent but veinlct reticulum usually not prominu- lous beneath; leaf margins more or less incurved to revolute; petiole 2-6 (-10) mm. long, plane or adaxially somewhat channeled; stipules ca- ducous, brownish-scarious, oblong-lanceolate, acuminate or blunt and apiculate, 2.5-5 mm. long. Staminaie (lowers several per axillary cluster; pedicels mostly 3-5 mm. long; sepals 4. biseriate, usually the outer ones 1.2-2 mm. long, 1-1.3 mm. broad, ovate, entire, the inner ones larger, 1.5- 2.3 mm. long, 1.3-2 mm. broad; disc adnate, smooth or rugulose, mostly 0.5-1 mm. across, I inn n i thi nl i m a\ ree, becoming 1-1.5 mm. long, the anthers 0.8-1.4 mm. long. Pistillate flowers solitary or less often paired a proximal i il of arum 1 mrremenl I >,rowtli; ])edicels more or less compressed and angled, 5-15(-20) mm. long; sepals 4, biseriate, ovate to oblong, entire, 1.5-2.5 mm. long; disc entire, adnate, 1-2.2 mm. across; ovary of 2 (very rarely 3) carpels, the styles free or basally co- herent, ca. 0.8-1.5 mm. long, bifid, with the branches more or less sharply reflexed. Fruits subglobo.se or laterally compressed, indehiscent or irregu- larly dehiscent, 6-8 mm. in diameter, endococci not developed (wall lay- ers not separating), columella not persistent; seed with thin, bluish sarco- testa, the endotesta smooth (i,neb iomewhal rugose) more or less plano- convex or trigonous-lenticular, (2.7-)3.5-5.5 mm. long, the hilum adaxial, circular, sub-basal, 0.9-1.2 mm. across.

DISTRIBUTION. Scrub or seasonal forests below 1000 meters; Madagas- car, Aldabra, Astove, and Mauritius (MAPS 5, 6). Reported from Cos- moledo by Fosberg (1978).

SPECIMENS EXAMINED. Aldabra Islands. Without further locality, 1892, Ab- bott s.n. (us). ILE MICHEL: scrub on samb GiampL.non Fosberg 49357 (DAV). MIDDLE ISLAND: scrub, modei.uob mii-li Inn, mm bobcrg 49582, 49666, 49671 (DAV). SOUTH ISLAND: scrub on flat limestone, Fosberg 48848, 48961, 49029, 49309, 49401 (DAV). Astove Island: Grand Anse, scrub on moderately 1979] WEBSTER, MARGARITARIA 431

rough limestone, Fosberg 49715, 19736 (DAA ). Comoro Islands. Without further locality. Boivin s.n. (K, P). GRANDE CISMURK: Moroni, d'Alleizette s.n. (L). Mada-

gascar. DIEGO SUAREZ foret en!n \i.11 »<,r,, 1 -.>,. 1( \, liodimagoro, plateaux calcaires, 250 m,, Humbert iV^, , , [•,<>. IMU \ dU- dThosy, 800-1000 m., rochers siliceux /M//M ; j s >M < M\m \ Berizoka, Perrier de la Bathie 339 (P) ; collines seches, Maevatanana, Perrier de la Bdthie 339bis 1 (P); vallee de Menn\ va Perrier dt hi hi!, / \VJter (p); bois sees, Bemarivo, Perrier de la Bat hie MM (P.I; vallee tie la Betsiboka a Marovoay, 50 m., bois du terrains arenaces Humbert '•> Pernei dt !<>, 'iet/ne M> ' (p); Ambongo, Per- ville r<62 (P.) \nkarafant ika T iramaiidroso Ranuunoujisoa 2029 (p), An- karaobato, Distr. Mitsinjo. SF 5386 (PI; station forestiere de Marohogo, Ma- junga, SF 7967 (p); foret de Morarano. Marohogo. iT MM (P); foret Tsimem- ; bo,, Antsalova. .S7 8248 (P). TAMATAVE: cote littoral est, Baie d'Antongil, d'Alleizette s.n. (L) ; Andevoranto, Ambila-Lemaitso, Ramasokoto 1524 (p); Tam- pina, foret littorale, /"em'er de /a £&/«'r/; /Mn' G(/M v.s f , „ hi, mousse 139 (K) ;

Plaine des Roches. /«/*e» M/lcV 1-^4 (MAU); Ferme Cascade, T'a«g/iflw MAU 1611 (MAU).

Hooker (1887) treated Margaritaria anomala as a synonym of M. in- dica, apparently on the basis of some Wallich specimens misidentified by Mueller (1866). However, although staminate specimens are difficult to distinguish, the indehiscent. 2-carpellate fruits of M. anomala are very different from the dehiscent, 3-carpellate ones of M. indica. Leandri (1958) has attempted to separate Phyllanthus anomalus from P. erythroxyloides on the basis of fruit and leaf shape. However, as noted by Coode (1978), the type specimen of Petit-Thouars was probably col- lected on Mauritius rather than on Madagascar. There is unfortunately some uncertainty in applying the name Phyllanthus erythroxyloides, since Baillon's allusion to the obsolete floral disc in his brief description (of Flueggea eglandulosa) would sugge I th; I hi might I ive had material of Margaritaria rhomboidalis. However, the leaves of the type collection (Perville 662) lack the prominulous venation characteristic of M. rhom- boidalis, and the pistillate flower does have a disc. At any rate, the Mauritius and Madagascar populations clearly seem to be conspecific, and the name M. anomala applicable to both. Fosberg (1978) has separated the populations from Aldabra and ad- jacent islands as var. cheloniphorbe. However, he does not cite any diagnostic characters, and although the Aldabra specimens do tend to have thicker and more rounded leaves, I can find no significant and consistent differences in leaves, flowers, or fruits between the plants of Aldabra and those of Mauritius and Madagascar. JOIRNAL 01' TIIF, ARNOLD ARIJOK K'l L'Al

n.v.). Glabrous shrub or tree to 5 meters high; twigs terete, grayish, with scattered lenticels. Leaf blades chartaceous to subcoriaceous, ovate to ellip- tic-oblong, 4-6(-9) cm. long. 2-3.5 (-5.5) cm. broad, emarginate, rounded or obtuse (rarely minutely apiculate) at tip, cuneate at base (distinctly de- i! u ui on ()« lid! i oli uceou veins (ca. 6 to 9 on a side) and veinlets prominulous on both sides in a promin ni retii ilum; margins plane; petiole distinctly laterally carinate or adaxially channeled; stipules cadu- cous. Staminate flowers several ; ixillai duster; pedicels capillary, 10-13 mm. long; sepals 4. hi * rial.. * llipth lo obovate, subequal, the outer ones ca. 1.2-1.3 mm. long, 0.8 mm. broad, the inner ones 1.5-1.8 mm. long, 0.8-1 mm. broad; disc angular, ca. 1 mm. across; stamens 4, the filaments united in pairs at base, 1.5-2.5 mm. long, the anthers elliptic, 0.3-0.5 mm. long. Pistillate flowers (not seen, descr. ex Airy Shaw) 1 or 2 per axil; pedicels slender, subterete or angled, 4-8 mm. long; sepals 4. biseriate, ca. 1 mm. long; ovary of 2 carpels, tlu- styles basally connate, ca. 1 mm. long, bifid, branches spreading. Fruits subglobose or elliptic, more or less indehiscent (tardily splitting from apex), 5-7 mm. in di- ameter, endococci corneous (not tenuous and hyaline); seeds with bluish sarcotesta very tenuous except at base, the endotesta more or less ridged or grooved on back, wedge shaped, ca. 3.1-3.7 mm. long and 2.1-2.9 mm. across the back tin luhin ha il, i ' 0 . mm across.

Australia. Queensland. BURKE DISTR. : Haydon Creek, i Road, alt. 60 m.. Ilyland 5121 ('DAV>. COOK DISTR.: Spring- mount Road, 17°10'S, 145°15'E. alt. 550 in., Ilyhwd 55'J-f (K); betw. ChillaRoe and Munpana. alt 3<)0 m.. Ilyland 5S5S (DAY); S. Palmer River cachement, 16°15'S, 144°40'E all '180 m.. Uxhrad sW (HAY) Additional Queensland lo- calities, cited by Airy Shaw (19761. have been indicated on MAP 5. The relationships i Ma / * in ,i bit i-trth \ n uneeil n flu rugose endotesta of the seed 'i • . . • ihi. affinity to M. indica, al- though the fruit is qui! > dilT< n nt nd urn • mul n lo (hat of M. anomala. Possibly M. dubium irac< vi i . -'.peri ili ed !• rivativ. of .1/. indica, com- parable to the vegetatively somewhat similar 71/. luzoniensis, and the re- semblances in fruits with .1/. anomala are (\uc to convergence.

12. Margaritaria hispidula Webster, sp. nov. FIGURES 2. 8. Ab aliis speciebus Madagascariensibus difietl folds venis prominulis, infra hispidulis. capsulis mox dehiscentibus. Tree 8 meters high; twigs subterete, whitish-hispidulous at tips, glabrate below, not prominently lenticellate. Leaf blades subcoriaceous, 1979] WEBSTER, MARGARITARIA 433 shiny and subglabrous above, copiously hispidulous-hirsutulous beneath, ovate or elliptic to obovate, obtuse or subacute at tip, cuneate to rounded at base, ca. 2.5-4.5 cm. long, 1-2.5 cm. broad; major veins (6 to 8 on a side) and veinlet reticulum distinctly prominulous on both sides; margins plane or recurved; petioles 3-6 mm. long, copiously hispidulous-hirsutu- lous, not adaxially channeled; stipules oblong-lanceolate, scarious, pale brownish, 2.5-3.5 mm. long. Flowers not seen. Fruiting pedicels sub- terete, 9-11 mm. long, hispidulous; carpi , th tyles (on immature fruit) ca. 2 mm. long, united about halfway into a column; fruits sub- globose, 6-7 mm. in diametei li pi lulous tardily dehiscent, the endo- carp tough, 0.2 mm. thick, not fragile and papery; mature seeds not seen. DISTRIBUTION. Known only from the type collection (MAP 6). teich, ISO m. alt., P. Otto Appet It is regrettable that this species has to be proposed as new on the basis of such imperfect material. Vegetatively, il is similar to Margaritaria rhomboidalis, but its very different fruits suggest a closer relationship to M. anomala or M. dubium-traceyi. Although we cannot be certain of its affinities until better material is collected. T feel sure that the Appert collection represents a species distinct from the known Madagascan taxa.

13. Margaritaria decaryana (Leandri) Webster, comb. nov. Phyll- anthus decaryanus Leandri, Not. Syst. Paris 6: 198. 1938; Fl. Madagascar 111(1): 98. fig. IS. 1-5. 195S. TYPE: Madagascar, Imangory, Decary 8937 (p).

>' hytla , ' \ in vanjis vai mavambia Leandri Urm in MI Ma da i < u 8B: 228. fig. 6, 1-5. 1957. TYPE: Madagascar, between Tsivory and Ima- nombo, Capuron 8543 (v. lectotypej. Glabrous shrub or small tree; twigs subterete, smooth, grayish or brown- ish; foliage deciduous. Leaf blades membranous, broadly elliptic or sub- orbicular to spathulate or obcordate, 1.0-1.8 cm. long, rounded to emar- ginate at tip, cuneate at base, green on both sides (somewhat paler be- neath); major veins mostly 3 or 4 on each side, veinlet reticulum some- what prominulous beneath; margins plane; petioles ca. 3-5 mm. long, adaxially channeled; stipules persistent or caducous, oblong, scarious, acute and more or less denticulate-fimbriate distally, 1.0-2.5 mm. long. Staminate flowers several per cluster; pedicels slender, 4-6 mm. long; sepals subequal, becoming reflexed, elliptic or ovate, obtuse, entire or obscurely denticulate at apex, 1.6-2.1 mm. long, 1.1-1.8 mm. broad, disc entire, thin, ca. 0.7-1 mm. across; stamens 4, the filaments connate be- low, mostly 1.5-2.2 mm. long (staminal column 0.8-1.5 mm. long), the anthers oblong, 0.8-1.1 mm. long. Pistillate flowers solitary or paired; pedicels angled-compressed, ca. 3-4 mm. long; sepals 4, subequal, 2-2.5 mm. long, 1.5-2.5 mm. broad; disc thin, entire, ca. 1.5 mm. across; ovary 434 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 of 2 or 3 carpels, the styles more or less erect, 1.5-2 mm. Ion,!?, basally connate, distally bifid. Fruits not seen. DISTRIBUTION. Xeric scrub; southern Madagascar (MAP 6).

SIT.i i \ir;\s i-\ \MI\KII. Madagascar. TULE et Imanombo, Capuron 8543, 8543bis (p); Bekily et Tsivory. Seyrig 242B (P). If I am correct in associating Leandns var. manambia with his P/zy//- anthus decaryanus. then the species is well characterized by its monadel- phous androecium and small suborbicular leaves. Leandri refers to the species and variety as each having 2docular ovaries, but actually they appear to be 2-locular in Cap/iron S;>-Uhis and 8 lo< ular in Seyrig 242B.

Mueller-.\rg; IKhh). Snurincuji Iwilloniami M ucller-Arg. in DCS. Prodr. 15(2) : 451. 1S66. Flucggca bailloniana (SMueller-Arg.) Fax, Bot. Jahrb. 19: 76. 1894. TYPE: Zanzibar, floivin s.n. (p, holo- type). FIGURES 9, 10.

type. n.v.). Shrub or tree 2-10 meters high; branches subterete, reddish- or brow ish-hirtcllous at growing tips, glabrate and grayish in age; foliage usua deciduous. Leaves rhai taceous; l)iades mostly obovate or spathulate, (1 2—4(—5) cm. long, 1-3 cm. broad, usually rounded or very obtuse at t cuneate at base, above smooth, beneath more or less glaucous and h lellou,, especially along miiii ib: major veins mostly 7 to 12 on a side, beneath, veinlet reticulm or less pr curved1 or narrowly levolute; petioles i isually hi. rsutulous, ii( )t adaxially channeled. 2 4(-5.5) m ; caducom ;, lanceolate, ca. 2-. 3.S mm. long. Staminate tlowers s everal per axil; pedicels glabrous. 3-5 rr im. long; sepals 1 )iseriate. the out er ones o blong, 1.5-2 mm. long, 0.7-1 mm. broad, the ir s broadly obovate, (1.5-) 2-2.1 -; mm. long, 1.2-1.15 mm. broad: dis*c 0.5-1 mm. acn oss; stain. ?ns 4. the filaments be coming 1.3-1.S mm. Ion:;; the .anthers linear-oblong. SW> lO.S-1 mm. long. Pistillate (lowers solitary or paired; pedicels often hirsutulous, mostly 2.5- o mm. long in fruit; : epals 4. sube([iial, oblong, hirsutulous on back, 1.6- 2.5 mm. long. 1.3-2 mm. broad; disc thin. 1.7-2.0 mm. across; ovary deeply trilobed. hirsutulous; carpels 2 (22 percent) or 3 (78 percent), very rarely 4, the styles free or nearly so, spreading, bifid to bipartite, 1.2-1.5 mm. long, (apsides deeply ?.- or 3 lobed (most often only 2 car- pels well developed), mostly 10-12 mm. in diameter, the cocci subspheroid- al, usually hirsutulous. the eiidocarp hyaline-papery: endotesta of seerl , smooth or grooved on back, 3.8-4.5 mm. long. 1979] WEBSTER, MARGARITARIA 435

DISTRIBUTION. Scrubland, deciduous forest, or savanna woodland, sea level to ca. 1000 meters; Burundi and Kenya south to Mozambique. Rad- cliffe-Smith (1976) cites a large number of additional localities, includ- ing records from Zaire and Zambia (MAP 4).

SPECIMENS EXAMINED. Burundi. BUBANZA: colline Muzinda, 900 m., Le- walle 1240 (MO); Randa, savane boi e< 900 m , Lewalle '.635 (NY). BURURI: Mufamb.n foief il.un 1000 m iinkinai^ 10, , lum Rumone foret claire sur rochcrs, 900 m.. Leiealle 1604 (not. Kenya. COAST: Kilifi. Sokoke. Gisau SOK6 i PRI . SOK11 (h PREI Malawi XORTI.F.RX ; Karons Distr.. 2 mi. N. of Chilumba in Vinthukhutu For< i R< erv< Brachysi a woodland, 1800 ft., Pawek 12064 (DAV). Tanzania. COAST: Tande Hill. 30 km. WNW. of Dar-es- Salaam, 250 ft., wooded grassland, Harris, Tadros, & Mwasumbi BJH 3616 (M\I 10477 (MO); Kisarawe Distr.. Sej;wei 3652 (PRE); 21 mi. S. of Dar-es-Salaam on Kilwa Rd., 150 m.,

Harris & Rodgers BJH 3070 (MO). MOROC;ORO : Kilosa. Semsei 3332 (PRE). NORTHERN: probably from Ukerewe Is., Father Courads 6035 (PRE). SOUTHERN HIGHLANDS: Kyimbila Distr.. Stoh 1744 (ex p.) (A, MO, P), 1771 (A, GH, MO, NY, P, PRE). TANGA: Korogwe Distr.. Kwamarukanga, Shabani 57 (PRE); Mkono, 30 km. E. of Handcni. 518 m, Archhold 172S (MO). ZANZIBAR: Sacleux (P) ; Kidichi, 300 ft., Faulkner 2751 (NY).

This distinctive species, which is widespread in east Africa, is sympatric with Margaritarta diunid a o\ i : inn lh all of its rans;e and has often been confounded with i! although Pax clear!} I ;tiiiL'ui hi I tin two in 1895 (under the names Flucggca hailloniami and F. nitida). Radcliffe- Smith (1976) provide thoroimh do lamentation of its distribution and discusses its relationship with .1/. disro/dea bui conservatively treats it as a variety. However, ..!/. obovata is clearly distinguished by the deeply lobed fruit (FIGURE 9), the usually distinctive pubescence (on fruits as well as leaves), and leaves with blades that are not decurrent on the petiole as adaxial flanges (i \en u iul ibauUei pointed out In Radeb'ffe Smith). In this yenu.s where distiriyuisluny characters are difficult to find, M. obovata is nun h iiuin c I irl\ in.nliil than must i i nul 1 believ< that it deserves full specific status.

DOUBTFUL AND EXCLUDED TAXA 1/ < >p/>ositifo! L in Urn PI Surinam. 16. 1775, nomen Ulegit. = (.'oriihretai ie (Mueller-Arg in DC Prodr 15(2): 414 1S66). Wiythintlms anhmui, Leandri Bull Soc Bot France 81: 452. 1934. TYPE: M nl ILM i \nl ii / i in Hat hie 1174 ( - \lthu i h i • mclri referred thi peci to /V nth/i etmn / tin t imiri iti di < ol separate segments is not correct for Margaritaria. In the later treatment by Leandri (Fl. Madagascar 111(1): 102. fig. XVI, 4, 5. 1958), it is clear that the plant is a true Phvllantlnts. neither of sect. Cicca nor of Margari-

Phyllanthus ftacourtioides Hutch. Kew Bull. 1915: 48. 1915. SYNTYPES : Mo- zambique, Lourenco Marques, Schlechter 115QS, 11634 (K, n.v7). Hutchin- son i I LIS 11 In spe u o Mai > i discoid mil h ch ri] tie u gests that it may be a form of M. discoidea Mabsp. nitida. However, if JOURNAL OF THE ARNOLD ARBORETUM

ACKNOWLEDGMENTS

The curators of a number of institutions have generously provided loans or access to specimens: A, BR, C, COI, DAY, V. G, GII, GOKT. HAC, K, L, M, MO, NY, p, PRE, s, TRIN. u, uc. us, WAG. I wish to thank Dr. F. R. Fosberg (Smithsonian Institution, Washington, D. C.) and Mr. M. J. E. Coode (Kew) for valuable specimens of and information on Mascarene taxa, Mr. A. Radcliffe-Smith (Kew) for comments on the African taxa, and Miss E. Retief (Pretoria) for help with localities in South Africa. The floral drawings were provided by Miss Kilian Garvey. I am especially indebted to Mrs. Mary Sigourney for her assistance with the maps and photographs.

LITERATl'RE CITED

ADJANOHOUN, E. 1964. Vegetation ties savanes et des rochers decouverts en Cote d'lvoire centrale. 178 pp. O.R.S.T.O.M., Paris. AIRY SHAW, H. K. 1963. Notes on Malaysian and other Asiatic Euphorbia- ceae NXI. Prosorus iudicus in Borneo and other regions. Kew Bull. 16:

. 1966. Notes mi Malaysian and other Asiatic Euphorbiaceae LXIX. New combinations in Margaritaria L. f. Ibid. 20: 386, 387. lo'/I. s'oh on Malax ian mtl olhei ' i i [ihorbiaceae CXXIV. A key to the Securineginae and some deceptively similar Phyllanthinae. Ibid. 25: 491-493. . I'Of). Now or noteworthy uniiiin u[>lioi'liiaeea( Ibid. 31: 341-398. ALM, J. 1775. Plantae surinamenscs. 18 pp. Dissertation, Uppsala. AUBREVILLE, A. 1959. La flore forestiere de la Cote d'lvoire. ed. 2. Vol. 2. 341 pp. Publ. 15, Centre Technique Furestier Tropical, Nogent-sur-Marne. BAILEY, I. W. 1924. The problem of identilying the wood of Cretaceous and later dicotyledons: Paraphyllnnlhoxylon anzonrnsc. Ann. Bot. 38: 439-451. BAILLON, H. 1858 Etude generah do group* d. 1 m1111uo maces 08-1 pp VT tor Masson, Paris. BANCILHON, L. 1071. Coiili iliulmm a I i tiuK M\oii i m pecics oi i'o sil dicotyledonous wood. Am. Jour. Bot. 59: 5-11. COODE, M. J. E. 1978. Margaritaria in Mauritius (unpublished manuscript). DALZEI.L, N. A. 1852. Contribution to the botany of western India [continua- tion]. Jour. Bot. Kew Misc. 4: 341-347. FOSBERG, F. R. 1978. Miscellaneous notes on the llora of Aklabra and neigh- boring islands: XL Critical notes on Euphorbiaceae. Kew Bull. 33: 181- 1979] WEBSTER, MARGARITARIA

GA.MIUU. J. S. DDF Flora of (he presidency of Madras. Vol. 2, P 963-1160. West Newman & Co., London. HALLE, F., R. A. A. OLDEMAN. & P. B. TOMLIXSOX. 197S. Tropical i forest in rehitectural anah I >p pun er-Vt lierli) HER.MAW, P. 1689. Paradisi Uatavi prodromus. Pp. 301-386 /'•;/ S. ed., Schola botanica. Amsterdam. HOOKER, J. D. 1887. Euphorbiaceae. Fl. Brit. India 5: 239-477. JUSSIEU, A. DE. 1824. D( Kuphorb id num gencribus medici viribus tentarnen. 118 pp. Dido!, S'ans. JUSSIEU, A. L. DE. 1789. Genera plantarurn secundum ordines naturales disposita. 498 pp. Herissant. Paris. KEXG, H. 1951. New or critical Euphorbiaceae from eastern Asia. Jour. Wash. Acad. 41: 200-205. . 1955. The Euphorbiaceae of Taiwan. Taiwania 6: 27-66. KOHLER, E. 1965. Die Pollenmorphologie der biovulaten Euphorbiaceae und ihre Bedeutung fur die Taxonomie. Grana Palynol. 6: 26-120. LEAXDRI, J. 1958. Euphorbiacecs. Flure de Madagascar et des Comores 111(1): 1-209. LIXNAECS. G. 1781. Supplementum plantarurn. 46S pp. Braunschweig. Im- pensis Orphanotrophei. MADEL, H. 1062. Die I'ossileu Euphorbiaceen-U<">I/er nlit besonderer Berrick- sichtigung neuei Funde aus dei Gbcrkreidt Shd-AfriDi enckenben; Lethaea 43: 283-321. MUELLER, J. 1866. Euphorbiaceae. DC. Prodr. 15(2): 189-1261, 1269-1273. OPLER, P. A., G. W. FRAXKIE, & H. G. BAKER. 1976. Rainfall as a factor in the release, timing, and synchronization of antliesis by tropical trees and shrubs. Jour. Biogeogr. 3: 231-236. PAX, F. 1800. Euphorbiaceae. Nat. Pfianzenfam. III. 3(5): 1-119. . I'd Dnplunbuu n I'p '^-2-D in A IACI EK ed , Die Pflanzen- welt Ost-Afrikas und der Nachbargebieft I heil G \ erzeichniss der bis jetzt aus Ost-Afrika bekaunt gewordenen Pfkmzen. Dietrich Keimcr, Ber- lin. • & K. HOFF.MAXX. 1931. Euphorbiaceae. Xat. Pflanzenfam. ed. 2. 19c: 11-233. PiJL, L. VAN DER. 1969. Principles of dispersal in higher plants. 153 pp. Springer-Verlag, Berlin. PLUKEXET, L. 1692. Phytographia. Pars tertia. London. Pu\T, W 1962 Pollen morpD ilo-n o (hi i uphorbiact it i I pedal relei

RADCLIFFE-SMITK, A. 1976. Notes on African Euphorbiaceae: \'I. Kew Full. 30: 675-687. SMITH, J. J. 1910. Euphorbiaceae. Pp. 9-637 in S. H. KOORDERS & T. VALE- TON, eds., Bijdrage 12 tot de Kennis der Boomsoorten op Java. G. Kolff & Co., Batavia. THWAITES, G. H. K. 1856. Note on the genus Prosorus. Jour. Bot. Kew Misc. 8: 271, 272. WEBSTER, G. L. 1957. A monographic study of the West Indian species of Phyll- anthus. Jour. Arnold Arb. 38: 51-80. — — •. 1965 ' revision ol ilk enu IP n < I (I uphorbiaceae). Acta Bot. Neerl. 14: 323-365. 438 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60

1'iyy Conspectus (it ;i new cla ssitua t ion of tin- I uphorbiaceae. Taxon 24: 593-601. • & D. BURCII. 1968. Margaritaria. Pp. 217-219 in R. E. WOODSON. JR.. R. W. SCHERY, <•/ a!., Flora of Panama. Part VI. Family 97. Euphorbiaceae. Ann. Missouri Bot. Card. 54. & J. ELLIS. 1962. Cyiuiaxononiic si ml irs in the Kuphorbiaceae. sub- tribe Phyllanthinae. Am. Jour. Bot. 49: 14 18. WHEELER. L. C. 1939a. A miscellany of New World Euphorbiaceae. Contr. Gray Herb. 124: 35-42. . 1939b. A miscellany of New World Euphorbiao/aw Ibid. 127: 48-78. WEBSTER, MARGARITARIA

INDEX TO NUMBERED EXSICCATAE The numbers in parentheses refer to the corresponding species and subspecies in the text. Not all exsiccatae are cited under REPRESENTATIVE SPECIMENS in the text. Acuiia, J. B., & Fr. Alain 18549, 18550 Berti, L. M., 202, 610, 776 (1). (4). Beusekom, C. F. van, et al 4207 (7). Adam, J.-G., 27458 (2a). Binuyo, A., FH 141435 (2a). Agnew, A., 7784 (2b). Black, G. A., 47-1539, 48-2943 (1). Akpaba, G. K., 1 (2a). Blanco., C. A., 830, 851, 899 (1). Alain, E. L., A257, A1593 (1); 3481, Boldingh, I., 2198, 2735a (1). 3761 (3?). Bond, F. E., T. S. Gillin, & S. Brown Alain, E. L, & J. Acuha 2999 (4). 150 (1). Alejandro, B., FB 27438 (7). Borle. J. M.. 174 (2b). Allen, P., 5018, 7040 (1). Box, H. E., 1493 (1). Alvarez, R. J., FB 22655 (7). Brace, L. J. K, 168, 3906, 4867, 6270, Andoh, J. E.. 5694 (2a). 6784, 6924, 6983, 7147 (3). Andrews, F. W., 1740 (2a). Breteler, F. J., 1214 (2); 3797 (1). Angus, A., 687 (14). Britton, E. G., 3403 (3). Appert, P. 0, 107 (12). Britton, E. G., & D. W. Marble 1305 Archbold, M. E., 1728 (14). Aristeguieta, L., A. Liogier, & L . Car- Britton, N, L, & L. J. K. Brace 741 denas de Guevara 7207 (1). (3). Armitage, F. B., 158 (2b). Britton, N. L., & E. G. Britton 9998 Asplund, E., 14574, 14631, 1970 1 (1). Aubreville, M., 475, 537, 560 793, Britton, N. L„ & W. E. Hess 2706 (1). 1026, 1448, 1522, 1628, 2328 237,9, Britton, N. L., & J. A. Shafer 792, 2330 (2a). 1532 (1). G, 1572 Broadway, W. E.; 345, , 4043, 4515 Bailey, L. H., & E. Z. Bailey 263 (2b) TRIN7411 (1). Bainbridge, W. R., 705 (2b). Brown, W. T. S., 2162, 2319 (2a). Baldwin, J. T., Jr. 6239, 11125 (2a) Bunting, G. S., & B. Trujillo 2227 (1). Balsinhas, A., 237 (2b). Calderon, S„ 707, 1668, 1726 (1). Bangham, W. N., 319 (1). Capuron, R., 8543, 8543bis (13); Barbosa, L. A. G., 7970, 8338 (2b). 11.621 SF, 11.623 SF (9). Barbosa, L. A. G., & M. Carvalho Carvalho, de, 703 (2b). 2950 (2b). Cenabre, FB 29209, FB 29210 Barbosa, & F. de Lemos (7). 4907 j) . Cenabre, A. L, J. Baldemor, & P. Adu- Barter, 1498 (2a). viso FB 28004 (7). ;., 483, 613 (1). Chaffey, O. R., 132 (2b). P., 1421 (1). Chamroerngsri 2 (7). P., & M. Magalhaes 612 Chandler, P., 1063 (2a). Chapin, J. P., 598 (2a). Belem, P., & R. S. Pinheiro 2902 Chase, N. C, 5150 (2b). Chevalier, A., 12499, 12757, 13342, Berhaut, R. P, 1091 (2a). 16179, 18066, 18212, 20348, 20775, Bernard!, 2999 (1). 20944, 21302, 22348, 23780 (2a). Bernoulli, C. G., & R. Cario 3398 (1). Claussen, M., 743, 744 (1). J()l RXAL OK THE ARNOLD ARROR17I I M

Cochet, M., 21 (2a). Kndlidi. R.. 4.V.? i Mr I. Codd. L. E., 9627 (2b). Enti, A. A., i?695 (2a). Codd, L. E„ & R. A. Dyer 4545, 45W Ernst, W. R., 7572, 7949 (1). (2b). Fspirito Santo, J. V. G., 26, 221. 524, Coker. W. C, 328 (3). 2002. 2162, 507S, 5S52. 5860 (2a). Collins, D. J., 933 (7). Fakude, E. N., 20 (2b). Conrads, I-., 6035 (14). Fanshawe. D. 17, 6364 (1); 7^757 (2b). Contreras, E.. 5706 (1). Farrell, J. A., 765 (2b). Coode, M. J. E., ct al. 5021, 5022, Faulkner. II. (7. 574 (2b), 2757 (14). 5053 (10). Fenix, E.. BS 15589 (7). Cooper. G. P., 436 (2a). Fiebrig. K„ 4729 (1). H„ SFN 29829 (7). Finlay, K., TRIN 2470 (1). Corrcll, ., 43522, 44107 (3). Fishlock, W. C. 57, 423 (1). Correll, .. & Evans -A?92y (3). Fleischmann, F.. 480 (1). (Anvil. V ]•'. ( vl r 4-1559 (3).

Onatn 74P5, Curran, H. M., 7060 (1); F73 Jj; Z?3 776P5 (6). Curtiss. A. II., 183 (3). Foster, R.. 7 792 (1). Dalziel. f. M., 35 (2a). Friedmann, F., 2620 (10). Darko, K. 0., 595 (2a). Froes, R. L., 7776, 77547, 55727 (1). Davidse, G., & F. Llanos 5456 (1). Froes. R. L„ & G. A. Black 27168, Davy, J. B., 601 (2b). Debeaux, 0., 316 (2a). Decary, R., 1118, 1122 (9). Degener, 0, 19139, 19140 (3). 24103, 24220 (1). Deighton, F. C, -7109, 4761, 4502 (: Gehrt, A., 31733 (1). Donie. C. 2095 (2a). Gentle, P. PL. 503. 28S2, 3557, 471 Drummond, R. B., 5049 (2b). 7651 (1). Gentry. A., & J. Revilla 16305 (1). Gerstner. J„ 2/7V5, 5006 (2b). Gillett, J. B., 16593 (2b). 11018, 11104 Gilliland, IP B., 7260 (2b). Gines, PL, 3486 (1). Gisau, SOK6 (2b); 5W77 (14). Glassman, S. F., 7795 (1). Glaziou, A., 6752 (1). A., TO Gonzalez, G. J., 725 (1). s. H. F 175, 255f), //7.A. Goodier. R., 459, 664 (2b). 75574, 15744 (1); 5026 (4). Gosswviler, J., 45 79, 4677, 4692, 478 Ekman, E. L.. 7460, 2577, 2627, 4206, 9948. 15606 (2a). 4570, 4602, 4641, 5137, 6083, 9602, Grey, P. 1858 (10). 1S0S6. 18151, 18269, H4219, Hahn, P.. 964 (1). f! 13250, 1112997 (l); 4236 (3?); Haines. H. 11., 4705 (7). /A/TV 14); 118364, II8909, H8919 Hansen 909 (2a). Harris, B. J., 6775 (2b). Harris, B. J., & A. Rodgers BJH 39 WEBSTER, MARGARITARIA

Harris, B. J., Tadros, & l- Klug, G., 2013, 2338, 2686, 2695, sumbi BJH 3616 (14). Harris, W., 8698 (1). Hassler, E., 10906 (1). Hatschbach, G 0). Haught Kostermans, A. J . G. H, e* c/. KK- Have S.. 109 i SS 4L ? (7). Hedin, L., 1618 (2 Krukoff, >Q, 5663, 5685, 6281, Heller. A. A., 1266 6564, 5257 (1) Heudel. Kuasa, D., 34 (2 Hill, S. R., 3027, 3035 (4). Lace, J. H.. 50. >4 (7). Hinton, G. B., 1991, 3911, 3975, 9555, Lag(\ C, 80707 ( 10264, 10623. 12285, 15167 (1). Langlasst i, E., J5t ' (1). Hodge, W. H., 3202 (1). Sobro, J. P., Holdridge, L. R., 4 (1). Lasser, T., & C. D. Adams 4570 (1). Holm, T. H„ 312 (1). Lasser, T., & V. Vareschi 4087 (l). Hornby, A. J. W.. 2880 (2b). Lanjouw, J., & J. C. Lindeman 1199, Hornby, R. M., 2580 (2b). 1357, 1556, 1869, 3167 (1). Hostmann, F. W. R., 622 (1). Leandri, J., 948, 954 (9). Howard, R. A.. 12078 (1). Leandri. J.. R. Capuron, & A. Raza- Humbert, II., 7185 (2b); 2P95, fmdrakoto 2195, 2338 (9). 2W8ter. 18993 (10). Leandri, J., & P. Saboureau 2720, Humbert, H., & H. Perrier de la Bathie 3069 (9). 2353 (10). Leeuwenberg, A. J. M., 2919, 4211, Hyland, B., 5121, 5594, 5838, 5895 (11); (5639 (7). Lemos. F. de, Ik A. Balsinhas 259 Irvine, J., 2522 (2a). Jack, J. G.; 7401, 8086 (1). Leon, H, 11033, 11738 (;i). Jack, J. G., & H. Leon 13918 (1). Leon. H„ & H. ClementtJ 2JJ3P (3). Jacobsen. W., 2657, 2948, 2971 (2b). Leonard, J., 1391 (2a). Jacques-Felix, II., 302, 3215, 3351, Le Testu, M. G., 3843, 4555, 7275 4813, 5510 (2a). (2a). Jennings, 0. E., 678 (3). Lctouzev. R.. SRFK 1292 , 5i?F,fi: ^295 Johnston, I. M., 954, 1251, 1297, (2a). Lewalle, J., 1240, 1604, 2635 (14). J. R., 332, 333 (1). Liben. L., 3734 (2b). Julien. AA4D' 242 M (10). Lima. F.. ,lfG J0745 (1) Junocl, A., 141 (2b). Lindeman, J. C., 5589, 5625, 6627, ^.., 1567, 1625 (1). 6634 (1). Kennedy. H.. 791 (I). Linder, D. H., 125 (2a). Kennedy. J. D.. 2209, 2244 (2a). Liogier, A. H., 12247 (1) Kerr, A. F. G„ 5858, 20331 (7). Little, E. L, Jr. 6354, 6672, 13460, Kersting. A., 216, 343, 577 (2a). 13490 (1). Killip, E. P., 8346 (1). Little, E. L„ Jr., & R. R . Little 5272, Killip. E. P., & J. Cuatrecasas 39152 8292. 8393 (1). Killip, F, A. C. Smith 2dPl5, Little, E. L., Jr., & R .. Woodbury

Klaine, R. P., 154, 124, 364, 585, D. F, 138, 311, 487 (2b). 589, 1031 (2a). Klein, R., 959, 161. JOURNAL OF THE ARNOLD ARBORETUM

Funded. C. L., 897, 3136, 3432, 6974 Otanes. F., BS 17834 (6). Pamplemousse (Herb.) 139 (1C Lundell. C. L.. & A. A. Lundell 7259, Pawek, J., 1/960, 12046. 12112, 1J559 7352 (1). (2b); 5072 (14). Luteyn. J. L.. & R. Foster 949, 1353 Pegel, K„ 102 (2b). Pena, B. S„ 609 (1). Maas. P. J. M., & J. A. Tawjoeran Perrier de la Bathie, H., 33/ ', 339, 339bis, 339ter, 9527, 13318 '(10);' Macedo. A.. & A. Balsinhas 1101 (2b). 1346 (9). Maguire, B., 40717 (1). Pervillc. N., 664 (9); 662 (1C Mangoen, M., £>1J (7). Phelps, R. J., 197 (2b). Mann. C. 882 (2a). I'hipps. J. B., 2358 (2b). Martius, C. F. P. von, 2813 (l). 1'ickel, lb, 4214 (1). Matuda. E., 2d02, J09(5, 4760 (1). Pierlot, R„ 729 (2a). Maxon. W. R., 76S0 (1). Pierre, M. L., 559 (2a). Maxon, W. R., & A. D. Harvey 6531 Pires. J. M, N. T. Silva. & R. Souza 9017 ( 1 ). Maxon, W. R„ & A. T. Valentine Pittier. H„ 3468. 4518, 6603, 6(550, 6950 (1). 7059, 7809. 8925. 12399 (1). McDaniel, S., 1610(5 (1). Pittier, H., & M. Nakichenovich 15555 McYaugh. R., C. Kcddema. & R. W. Pippen 21190 (1). Plowe, D. C, 1320 (2b). Medina. E„ S24 (1). Pobeguin. M., 171, 681, 1178 , 1506 Mcikle, R. D„ 1124 (2a). Mexia, Y., 6140. 6296 (1). Porter, R. V. H.. 993 (2b). Mgaza, C. D., 5(52 (14). Powell. D., 1702 (1). Miller, 0. B„ 1991, 2017, 2522, 3295 Prance. G. T.. J. F. Ramos, & L. G. (2b). Farias 7361 (1). Miranda, D. P., FB 22943 (7). Prance, G. T., et al. 14477A (F Mogg, A. 0. D., 28639 (2b). Prestoe. R. A. R.. TRIN 1642 (1). Molina, A., 129. 755, 5//, 898, 968. Pringle, C. G, 10319 (1). 2000, 2891, 3966. 5429 (1). Procter, J.. 2984 (14). Moll, E. J., 385, 592, 3231, 3374, Proctor, G. R., 17714, 18169, J2505 4353, 4456, 4514 (2b). Moll, E. J.. & Ncl 5573. 5575 (2b). Purpus, C. A., 5092, 8454, 8789 , 8798, Moller, A. F„ 405 (2a). 17162 (1). Moore, S., 1025 (1). Put 1230, 1734 (7). Mobray, R. N., 69417 (1). Quintas, F„ 916 (2a). Muller, T., & T. A. D. Gordon 1396 Ramasokoto 1524 (10). (2b). r , a.., , „n ! Mullin, L. J., 89/51 (2b). Ramos, M., & G. Kdafio US 26282 Mwangansi. O. M., 1768 (2b). (6); BS 44303 (7). Mwasumbi, L. B„ LBM 10477 (14). Raunkiaer, (*., 2782. 2873 (1). Nash, G. V, & N. Taylor 1525 (1). Reckmans, M., 3192 (2b); 1077, 4075, Natividad, P., 23361 (7). 4224 (14). Xicolson. 1). H., ct al. 11FP SO (7). Reijenga, T. W., 934 (1). Northrop. J. T., & A. R. Northrop krnz. IF. ,vv IF M. Klein, 9575 (1). (3). Ritchie 1343 (7). Robinson. E. A, 2506 (2b). Robson. N. K. lb, & I). B. Fanshawe Ortega, J. G., 7060 556 (2b). 1979] WEBSTER, MARGARITARIA 443

Robson, N. K. B., & M. Steele «» Stern, W. L., R. L. Chambers, J. D. Dwyer. & J. E. Ebinger 76S, 919 Roig. J. T., 3243 (3). Roig, J. T, & M. Cremata 2148 (1). Sterringa, J. T., 12441, 12529 (1). Stewart, E. C, 14 (1). Rose, H.. 181 (2). Steyermark. J., 45 SOT. 47781, 48053, Rose, J. N., J. H. Painter, & J. S. ROM- 51076, 51413, 51416, 51495, 62825, 87008, 8

Webster. C. E. W. !\ Adam, K. M ler, & L. Miller J2000 (1). Webster, G. L.. J. Ellis. & K. I. M 12705, 14107 ( R. ()., 12014 i Webster. G. L.. E. Samuel. & T. W \. 8394 (3).

Welwitsch. I'.. 340 (2a). Winter. B. de. & J. Yahrmeijer 8601 Wed.'t-mann. E.. 2549 (1). Wgoni, J. F„ 65 (2b). Winzerling. 11. \\\. 11-10 ( 1 ). While, C. E., 993 (1). Wright. C. 5S4a, 1951, 1932, 1935 (I); White. F.. J4P0, J<500, 6326 (2b). 1450 (3); 1437 (3, 4); 583, 584 Wight, IE 2599 (7). Wild. EI.. 4 MO (2b). Wurdack. J. J.. & J. V. Monachino W.Mr. I. K„ & J. T. Baldwin. Jr. 39689, 41411 (1). Yuncker, T. (7, }. M. Koeppcn, a K. .Id,, W J. J. O. de. & B. E. h\ de A. Wagner 5/54 (1). Wilde-Duyfyes 2374 (2a). Zenker, G., 927, 2053, 2055, 2550, illiams. L., 565, 2481, 5404, 4050, 2S90, 5158, 4203 (2a). 8806, 10255, 10272, 11845, 13304