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Jablonskia, a New Genus of Euphorbiaceae from South America

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Jablonskia, a New Genus of Euphorbiaceae from South America SystematicBotany (1984), 9(2): pp. 229-235 ? Copyright1984 by the American Society of Plant Taxonomists Jablonskia,a New Genus of Euphorbiaceae fromSouth America GRADY L. WEBSTER Botany,University of California,Davis, California 95616 ABSTRACT.A new monotypicgenus, Jablonskia, based on the South American Securinegacon- gesta (Euphorbiaceae, Phyllanthoideae), resembles genera of the Antidesmeae in pollen and wood characters,but the floralstructure is more similar to that in genera of Aporuseae such as Ashtonia and Richeria.From Securinega,where it was originally placed, and the taxa of Antidesmeae and Aporuseae, Jablonskiamay be separated by a combination of characteristics:monoecious; inflores- cences axillary; flowersconspicuously bracteate,the staminatesessile; pollen grains prolate, exine tectate-perforate,germ pore lalongate; seeds with fleshy exotesta, paired in each locule of the irregularlydehiscing capsule; embryowith radicle equalling the cotyledons. Among the taxa of Euphorbiaceae that were Muell. Arg. was originallydescribed as a species never revised by Pax and Hoffmann in their of Phyllanthusby Mueller (1863) and later long series of monographictreatments is Phyl- transferredto Securinegaby Mueller (1873). lanthinae. As circumscribedby Pax and Hoff- Mueller referredthis species to his sect. Secu- mann (1931), this subtribe included five gen- rinegastrum[=sect. Securinega],which had pre- era: Zimmermannia,Securinega, Pleiostemon, viously included only the type species from Phyllanthus,and Reverchonia.Relationships in Mauritius, S. durissimaJ. F. Gmelin. Although this complex have remained rather poorly Bentham (1878, 1880) concurredwith its place- understood because of the lack of comprehen- ment in Securinega,the South American species sive study since the last, now badly outdated, is clearly anomalous because of its monoecy, treatments of Mueller (1866) and Bentham conspicuouslybracteate flowers, and seeds with (1880). In an attemptto improve the classifica- fleshyexotesta. tion of this group, I have considered problems The discordant position of S. congestais also of generic delimitation in studies on Phyllan- apparent in its pollen characters.In his survey thus(1956-58), Reverchonia(1963 with Miller), of Euphorbiaceous pollen, Punt (1962) indicat- Meineckia(1965), and Margaritaria(1979). ed that the pollen of S. congestadiffers from The mostchallenging problem of generic de- that of other species of Securinegain its prolate limitationinvolves SecurinegaJuss. Although shape and elongated colpus transversalis.Al- Pax and Hoffmann(1931) treated Securinegain though Kbhler (1965) did not reporton the pol- a broad sense, including FlueggeaWilld., Ben- len of S. congestain his pollen survey of 'bio- tham (1878) characterizedit as a "purely arti- vulate' Euphorbiaceae, it appears to fitinto his ficial genus" and maintained Flueggea as dis- 'Antidesmatype' because of the prolate grains tincton the basis of its fruitand seed structure. with long narrow colpi and tectateexine with Aftera study of this problem over the last 15 small lumina. This may be seen in the figures years,I have concluded that Bentham was cor- included here, which show that the pollen of rect in separating Flueggeaand that the artifi- S. congesta(fig. 1) is much more similar to that ciality of delimitation of Securinegashould be of Hyeronimaalchorneoides (fig. 2) and Celianella resolved by segregatingthose elements whose montana(fig. 3) in Antidesmeae than it is to the diagnostic featuresare incompatible with the pollen of Richeriagrandis (fig. 4) and Ashtonia type species, S. durissimaJ. F. Gmelin. Part of praeterita(fig. 5) in Aporuseae or Securinegadur- this task was accomplished with the reinstate- issimain Phyllantheae (fig. 6). The tectate-per- ment of Meineckia(Webster 1965), but a num- forate exine of S. congestapollen is strikingly ber of discordantgroups remain,including the similarto thatin pollen of Euphorbioideae and South American plants that are the subject of is unique in Phyllantheae; however, it appears this paper. to be derived fromthe finelyreticulate exine The South American taxon currentlyknown of genera such as Celianella(fig. 3). as Securinegacongesta (Benth. ex Muell. Arg.) The indications of affinitysuggested by pol- 229 This content downloaded from 169.237.8.35 on Mon, 16 Sep 2013 18:29:31 PM All use subject to JSTOR Terms and Conditions 230 SYSTEMATICBOTANY [Volume 9 5~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~9 6 - ';, FIGS.1-6. Pollengrains of somegenera of Euphorbiaceae-Phyllanthoideae;scales =10 sm. 1. Jablonskia congesta(Rimachi 4838, DAy). 2. Hyeronimaalchorneoides (Alves Lages RB80704,DAy). 3. Ceizanellamontana (Maguireand Maguire35071, NY). 4. Richeriagrandis (Webster 13477, DAy). 5. Ashtoniapraeterita (Soepadmo 783, K). 6. Securinegadurissima (For. Dept. 11399,DAy). This content downloaded from 169.237.8.35 on Mon, 16 Sep 2013 18:29:31 PM All use subject to JSTOR Terms and Conditions 1984] WEBSTER: JABLONSKIA 231 len morphology are consistentwith the avail- able data for wood anatomy. In the brief,and only available, previously published descrip- tion of the wood of S. congesta,Record (1938) mentioned that it differsfrom the wood of Se- curinegaacidoton by having vessel elementswith scalariformperforations, thin-walled wood fi- bers, and reticulatexylem parenchyma. In the classification of Phyllanthoidean woods by Mennega (1984; ined.), S. congestaappears to fit best into group A2, which includes Aporusaand other genera of Aporuseae (as delimited by Webster 1975). Mennega (pers. comm.) notes, however, that genera of Aporuseae such as Aporusa,Ashtonia, and Richeriadiffer from S. congesta in having diffuse parenchyma and longer non-septatefibers. In gross morphologicalcharacters, S. congesta has much in common with the South American Richeria,both in general habit (fig. 7) and in lcm floral details (figs. 8-10). In the treatmentof Phyllanthoideae by Pax and Hoffmann(1922), it keys to Richeriaif treated as a member of Antidesminae.However, S. congestadiffers from Richeriain its monoecy,flowers in axillaryclus- ters (rather than spikes), thin-walled capsules with paired seeds in each locule, and embryo with elongated radicle (about equalling the cotyledons).In some respects,such as the thin- walled capsules and fleshy seeds, S. congesta FIG. 7. Branchof Jablonskiacongesta with imma- more closely resemblesthe Old World Aporusa. ture fruits(Revilla 22, DAV). However, Aporusais dioecious; the staminate flowers usually have only two or three sta- mens, a reduced pistillode, and no disk; the pistillate flowerslack a disk and usually have S. congestaand those of Amanoa,which is also two carpels with fimbriatestyles; and the fruits monoecious and has pentamerous flowers. have at most one seed per locule with cotyle- However, Amanoa differsin a number of sig- dons much longer than the radicle. Further- nificantcharacters, particularly the flowerswith more, S. congestadiffers from both Richeriaand well-developed petals, sessile stigmas, drupa- Aporusain pollen and wood characters. ceous fruits,and seeds with endosperm scanty More similar to S. congestain some respects or absent. Furthermore,the pollen of Amanoa is the recentlydescribed Malesian Ashtonia(Airy is very different,with a coarsely reticulate or Shaw 1968, 1972, 1974),which has pentandrous bizarrely ornamented exine, as illustrated by staminateflowers and capsular fruitwith fleshy Punt (1962) and Kohler (1965). seeds. However, the flowersof Ashtonialack a Mention should also be made of Celianella, well-developed floraldisk; the staminateinflo- recentlydescribed fromthe Guyana highlands rescences are racemose; the sepals are three or by Jablonski (1965). The pollen of Celianella four,not five; the pollen grain exine (fig. 5) is montanaJabl. (fig. 3), although finelyreticulate reticulate; and the styles are undivided and ratherthan tectate-perforate,is somewhat sim- stigmatiform.None of the other genera in An- ilar to the pollen of S. congesta(fig. 1) and of tidesmeae or Aporuseae (sensu Webster 1975) Hyeronima(fig. 2) and other genera of Antides- appears to be more similar.There is some sim- meae. Furthermore,in the accompanying pa- ilaritybetween the bracteateflower clusters of per Mennega (1984) notes a resemblance be- This content downloaded from 169.237.8.35 on Mon, 16 Sep 2013 18:29:31 PM All use subject to JSTOR Terms and Conditions 232 SYSTEMATIC BOTANY [Volume 9 8 ~~~~~~~~~~~~~~~~~~10, IAp" FIGS. 8-10. Flowers of Jablonskia.8-9. Staminate flower (one sepal removed in fig. 9); Rimachi4838 (DAV). 10. Pistillateflower (2 sepals cut away; gynoecium removed to show disk); Revilla22 (DAV). tween Celianellaand Securinegacongesta in wood curinegas.str. differt foliis glanduligeris,plantis anatomical characteristics.However, Celianella monoicis,floribus bracteatis, calyce glabro,pol- differsfrom S. congestain having racemose ped- linis granulis prolatis,perforato-tectatis, semi- icellate staminate flowerswith pendulous an- nibus carnosulis. thersthat are rathertypical for Antidesmeae. It Monoecious glabrous trees or shrubs. Leaves also differsin the stronglyaccrescent pistillate alternate,distichous, simple, entire, pinnately calyx and in the seeds, which are apparently veined, short-petiolate, with small laminar solitaryin each locule of the capsule and have glands near base; stipules elongate, caducous. an embryowith the radicle much shorterthan Flowers apetalous, sessile or subsessile in axil- the cotyledons (in this respect fittingAporu- larybracteate
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