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Summary. A new family of Asparagales, Xeronemataceae, is described. This small family of two species is native to New Zealand and was formerly included by several authors in Phormiaceae, to which they are grossly similar. However, they differ in a number of significant ways, and the results of DNA sequence analysis strongly support them as occupying an isolated position as sister to a clade composed of most of the higher Asparagales, thus making their description as a new family appropriate. Xeronema Brongn. & Gris is a poorly known genus of two species. It was initially thought to be monotypic with the only species being X. moorei Brongn. & Gris from New Caledonia. However, in 1924 another species, X. callistemon W. R. B. Oliv., was discovered on inaccessible inland cliff areas of the Poor Knights Islands, situated 15 miles off the northeastern coast of New Zealand (Oliver 1925, 1926). Xeronema callistemon was later found on the outlying New Zealand island Taranga (Cranwell 1933) but has never been recorded growing naturally in mainland New Zealand. The two species are rather similar in morphology, but in general X. callistemon is a larger plant than X. moorei, and there are differences in flower structure; for example, X. callistemon has a more persistent style and shorter, more tapering seeds (Moore 1957). Both species have leathery, unifacial equitant (ensiform) leaves and congested racemes bearing elongate, crimson flowers that all face upwards. The stamens are highly exserted and have persistent filaments. All parts of the flowers are red or orange, including the pollen; they could be bird or butterfly pollinated. The flowers have six free tepals, six free stamens with versatile anthers dehiscing by longitudinal introrse slits and a secretory tapetum. Pollen is monosulcate, with a finely reticulate exine, and normally boat-shaped with an elliptical sulcus (Fig. 1A), also reported by Cranwell (1953) and Schulze (1982). In material examined here (with standard SEM techniques), we found rare trichotomosulcate grains (Fig. iB), with a Y-shaped sulcus, among predominantly monosulcate grains (Fig. 1A). The ovary is superior, trilocular, with axile placentation and several crassinucellate ovules per locule. Prominent, branched, highly vascularised septal nectaries extend from the ovary base to openings around the style base (Rudall unpublished). Calcium oxalate raphides are present in idioblasts throughout all tissues, but styloid crystals are absent (Prychid & Rudall 1999). The fruit is purplish, with numerous black seeds with papillate surfaces (Oliver 1926).

Five Central American species of Dieffenbachia, D. beachiana Croat & Grayum, D. concinna Croat & Grayum, D. grayumiana Croat, D. hammelii Croat & Grayum, and D. tonduzii Croat & Grayum, as well as one species of Rhodospatha, R. pellucida Croat & Grayum, are described as new. Because of the impending completion of the Flora de Nicaragua, five names in Dieffenbachia and one in Rhodospatha must be published here in order to be included in that work. Revisions of both genera with additional new species (for Central America only for Dieffenbachia) are forthcoming. Dieffenbachia beachiana Croat & Grayum, sp. nov. TYPE: Panama. Bocas del Toro: along road between Chiriqui Grande and Fortuna, 13.2 mi. W of Chiriqui Grande, 8?45'N, 82?10'W, 310 m, Croat & Grayum 60130 (holotype, MO-3230653; isotypes, K, PMA, US). Planta terrestris, 40-100 cm; internodia 2-6 cm longa, 1.5-3.5 cm diam.; petiolus 17-46 cm longus, vaginatus 1/4-8/10 longitudinis; vagina decurrens ad apicem; lamina anguste elliptica vel lanceolata, 16-41 cm longa, 6.515 cm lata. Inflorescentia 1-3 in quoque axilla; pedunculus 9-13 cm longus; spatha 10-19 cm longa; spadix usque 13-15 cm longus; pistila 48-66. Slender herb 40-100 cm tall; stems briefly creeping at base; internodes 2-6 x 1.5-3.5 cm, medium green to olive-green, sometimes streaked with cream, semiglossy and obscurely roughened. Leaves clustered toward apex of stem, erect-arching; petioles 17-46 cm long (averaging 26.5 cm), broadly C-shaped in cross section, flat with acute, erect margins or sharply to bluntly sulcate adaxially, dark green to brownish, pale-green mottled, matte, usually minutely roughened and with scattered whitish raphide cells visible, drying with scattered clusters of pustular raised areas with granular-puberulent progressions; unsheathed portion (2.5-)10-30 cm long; sheaths decurrent distally, 10-22 cm long (averaging 14.2 cm), comprising 0.25-0.83% of the total length of the petiole (averaging 0.54% of petiole length); blades narrowly elliptic to lanceolate, rarely narrowly ovate, 16-41 x 6.5-15 cm, 1.8-5.3 times longer than wide, 0.71.9 times longer than petioles, inequilateral, one side 1-2 cm wider than the other, + equilaterally acuminate at apex (acumen to 5 mm long), slightly inequilateral and acute, rounded or truncate at base (rarely subcordate in Panama), thinly coriaceous to subcoriaceous, drying papyraceous, usually quilted, weakly bicolorous, crisp-undulate along the margins; upper surface semiglossy (rarely matte), plain dark green (in Costa Rica, but sometimes mottled white or cream in Panama); lower surface slightly paler, semiglossy to weakly glossy or matte below; midrib flat-raised, often striate and usually concolorous above, convex or thicker than broad beneath; primary lateral veins 23-36 pairs, sunken above, convex beneath, drying paler than surface, departing midrib at an acute angle, then straight to weakly curved to the margin at 70o-110o before sweeping prominently toward apex (to ca. 450 toward apex and sometimes at the very base, sometimes with adjacent veins alternately reflexed and ascending); interprimary veins usually present, not markedly weaker than the primary lateral veins; midrib and primary lateral veins puberulent on lower surface with thick, whitish trichomes; minor veins moderately distinct and weakly raised beneath. Inflorescences 1-3 per axil; peduncles 9-13 cm long; spathe 10-19 cm long, uniformly green; gradually long-tapered to apex, gradually contracted at middle, which corresponds to the sterile section of the spadix; tube 1-2 cm diam. when furled; blade to 3 cm wide when flattened; spadix to 13-15 cm long; staminate portion of spadix 5--6.5 cm long, drying 4 mm diam.; male flowers 1.8-2.6 mm diam., irregularly subrounded, drying smooth and light brown at apex; pistillate portion of spadix 4.5-6 cm long, drying 7 mm diam., usually separated from the staminate portion by a slender, almost completely sterile section 2.0-3.5 cm long, drying 2 mm diam. (the staminate and pistillate portions sometimes almost contiguous); female flowers 4866, 2-4 situated across the width of the spadix, pistils oblong-ellipsoid, 1.5-2.0 mm long, 1.0-1.6 mm diam.; stigma subglobular, about as broad as the ovary; staminodia narrowly clavate, usually not at all fused at base, ca. twice as long as pistil. Infructescence somewhat flattened, yellow-green NovoN 9: 491-502. 1999. This content downloaded from 157.55.39.144 on Wed, 07 Sep 2016 04:37:01 UTC All use subject to http://about.jstor.org/terms

Three new species of Manettia (Rubiaceae) are described from Panama: M. arboricola, M. microphylla, and M. longicalycina. Manettia Mutis ex L. (Rubiaceae) is a genus of herbaceous or somewhat woody, usually twining vines distributed in the West Indies, Mexico, Central America, and South America south to subtropical Argentina and Paraguay. Comprising an estimated 80-130 species, this genus was revised by Wernham (1918, 1919) and treated for Central America and Mexico in the North American Flora (Standley, 1921), Flora of Guatemala (Standley & Williams, 1975), and Flora of Panama (Dwyer, 1980). Although two papers on the classification of Manettia were published by Chung (1967, 1968), no recent comprehensive accounts of the genus exist. Traditionally, Manettia has been placed in the Cinchoneae (de Candolle, 1830; Schumann, 1889; Steyermark, 1974; Robbrecht, 1988), on account of its dry fruits with numerous ovules per locule and winged seeds. Because Manettia contains raphides, Bremekamp (1966) and Verdcourt (1958) removed it from Cinchoneae and placed it in Hedyotideae, an option also mentioned by Robbrecht (1988). In a recent cladistic study of Cinchoneae incorporating cpDNA data and other information, Andersson & Persson (1991) suggested that the presence or absence of raphides in Rubiaceae is not as important a character as previously thought. Andersson and Persson placed Manettia in Hedyotideae in the same clade as Bouvardia, Hedyotis, and Hindsia, based on shared characters of distinct placentas attached centrally by a stipe, testal outgrowth forming a concentric wing, wood having fiber tracheids and predominantly solitary vessels, and loculicidal capsules dehiscent only at the apex. During the course of preparing a treatment of Manettia for Flora Mesoamericana, three new species-all from Panama-came to light and are described below. Most interesting are the two hemiepiphytic species, M. arboricola Lorence and M. microphylla Lorence & Dwyer, which climb up tree trunks by means of adventitious roots, an unusual feature previously unreported in the genus. These species are otherwise characteristic of Manettia in terms of their flower, capsule, and seed morphology. 1. Manettia arboricola Lorence, sp. nov. TYPE: Panama. Comarca de San Bias: El Llano-Carti Road, km 16.7, 350 m, 9019'N, 78?55'W, 16 June 1985, G. de Nevers & S. Charnley 5907 (holotype, MO 3623655, photo, PTBG; isotypes, F, SCZ). Figure 1. Species non volubili hemiepiphyto habitu, radicibus adventitiis, inflorescentia terminali in ramis frondosis disposita, floribus 3-5, corolla caerulea, tubo gracili 0.50.6 mm in diametro distinguibilis. Hemiepiphytic vine ascending tree trunk, the main stem woody, 1.5-2 mm diam., the bark brownish white, peeling, adventitious roots produced at nodes, the primary branches 10-23 cm long, arising at nodes, sparsely branching again, the secondary branches 4-10 cm long, 1 mm diam., usually leafy and terminated by a single inflorescence, the internodes 1-3 mm long, each side with a longitudinal ridge flanked by 2 furrows, smooth, glabrous, whitish; leaves petiolate, the petioles 2-6 x 0.5 mm, glabrous; lamina ovate, 2.5-5 x 1-2.5 cm, the base cuneate or rounded, the apex acute, glabrous, drying dark brownish green, chartaceous or membranaceous, the 2? veins 4-6 pairs, arcuate, weakly brochidodromous; stipules narrowly deltoid, 0.5 mm long, thickened. Inflorescence terminal, glabrous, cymose, 3-5-flowered, the peduncle 10-13 mm long; flowers 4-merous, the pedicels 4-10 x 0.2-0.3 mm, bracteolate, the bracteoles subulate, 1 mm long, the hypanthium obovoid-turbinate, 1.5-2 x 1-1.2 mm, glabrous; corolla blue when fresh, salverform, the tube 8-9 mm long, 0.5-0.6 mm diam., glabrous externally and internally, the lobes narrowly lanceolate-elliptic, 7-8 x 1.5-2 mm, glabrous; stamens included, affixed 2 mm below throat, the anthers 1.5 mm long, linear-ellipsoid; style 1.5 mm long, the stigma bilobed, the lobes linear, 0.5 mm NOVON 3: 59-62. 1993. This content downloaded from 207.46.13.150 on Sat, 23 Jul 2016 05:06:44 UTC All use subject to http://about.jstor.org/terms

Caespitose palm with stem 1.5-3-5 m tall and 1*5 cm diam.; internodes 2-5 cm long. Leaf sheath tough and fibrous, 15 cm long, very densely rustytomentose; petiole 40-47 cm long; lamina 36-54 cm long with 3-5 pairs of pinnae; pinnae 21-28 x 2-5-8 cm, with lower veins and rachis densely rufous-hairy. Inflorescences borne among the leaves; prophyll as long as leaf sheath; second peduncular bract projecting 8 cm beyond the leaf sheath, densely brown-tomentose. Inflorescence to 49-57 cm long, with peduncle to 29 cm; rachillae 4-8 in number, 19-33 cm long, the lowermost rachillae sometimes with second-order branching. Staminate flower with calyx 1 mm long, corolla 3 mm long, ribbed; stamens with lobed anthers 1-5 mm long, tissue in the stomial region with conspicuous large raphides; pistillode columnar, 2-5 mm long. Pistillate flowers (immature) with calyx 1 mm long; corolla 2 mm long and ribbed; staminodes minute; ovary ovoid, 1.5 mm long. Fruit elongate, 1-5 cm long by 5-6 mm broad at the base, tapering to 3 mm at the apex, with 3 ribs on the dorsal side and 2 lateral ribs extending less than half way from the base of the fruit, apex curved towards the basal stylar remains; young fruit green, turning white and ripening red. Endosperm homogeneous, curved and smooth (without ribs).

Ludwigia anastomosans, a tree to 10 m, is studied because it is unusual in the genus in its arborescent habit. It proves to have unusually wide vessels; it also has interxylary phloem, hitherto reported for only one species of the genus. Wood anatomy of Ludwigia peduncularis and L. torulosa shows that they may be more closely related than so far indicated. Vestigial bars on perforation plates of L. torulosa are the first observed in Onagraceae and are believed to represent an instance of paedomorphosis, but also retention of a primitive feature. Other indicators of paedomorphosis in Ludwigia are abundance of erect ray cells and notably long vessel elements. The hypothesis that degree of vessel grouping is related to ecology in taxa having fiber-tracheids or libriform fibers is validated by Ludwigia, which has the lowest degree of vessel grouping for the family and is essentially aquatic. Other anatomical features reflective of ecology, combined in the Mesomorphy ratio, present a not dissimilar pattern that can be integrated with that given by vessel grouping if one takes into account probable transpiration rates and temperature regimes as well as water availability. Data on wood anatomy of eight species of Ludwigia have been presented earlier (Carlquist, 1975, 1 982a). That number seems small unless one takes into account the fact that Ludwigia is predominantly herbaceous; the most familiar species are nonwoody herbs of very wet habitats such as ponds, ditches, and streams. One of the species in the present study is a notable exception: L. anastomosans (DC.) Hara is a tree. The data on the collection studied here describe it as a tree 10 m tall with a trunk 15 cm dbh. Wood anatomy is of special interest because of this habit. In fact, the results obtained below demonstrate that the wood of L. anastomosans differs appreciably from that of other Ludwigia species. Dr. Elsa Zardini, who collected the material of L. anastomosans, also kindly supplied material of two other species because she wished to see if wood anatomy demonstrated the degree of relationship between them. Dr. Zardini has contemplated the idea that L. peduncularis (Griseb.) Gomez may be closely related to L. torulosa (Am.) Hara. The wood anatomy of Ludwigia is of considerable interest with respect to ecology because Ludwigia characteristically grows in very wet places. Ludwigia anastomosans was collected in bamboo clumps by a blackwater stream in the Parque Natural de Caraca, Minas Geraes, Brazil. The material of L. peduncularis came from ditches near Havana, Cuba. The L. torulosa specimen was collected in a natural pond 17 km south of Tumeremo, Distrito Roscia, Estado Bolivar, Venezuela. In taxa with libriform fibers such as Onagraceae, Carlquist (1 984a) hypothesized degree of vessel grouping to be in direct proportion to adaptation to dry conditions. In this case Ludwigia species ought to exhibit a low degree of vessel grouping. Although figures for vessels per group were developed in the earlier survey of woods of the family (Carlquist, 1975), no comparisons were made between those figures and ecological regimes occupied by the various species. Ludwigia is a group of interest with relation to paedomorphosis in wood anatomy. This is, in turn, related to habit and ecology. The species in the present study were examined in this context to see if woodiness in Ludwigia is primary or secondary. Although a study on wood anatomy can be expected to reveal new records for anatomical features, two in the present study proved of especial interest and worthy of discussion: occurrence of interxylary phloem and presence of vestigial bars on some perforation plates. I Grants from the National Science Foundation to Peter H. Raven made possible the collection and selection of the material for this research. 2 Rancho Santa Ana Botanic Garden, Claremont, California 91711, U.S.A., and Department of Biology, Pomona College, Claremont, California 9171 1, U.S.A. ANN. MISSOURI BOT. GARD. 74: 889-896. 1987. This content downloaded from 207.46.13.151 on Fri, 25 Mar 2016 08:49:45 UTC All use subject to http://about.jstor.org/terms 890 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74 MATERIALS AND METHODS Voucher specimens are located at the Missouri Botanical Garden. Appreciation is expressed to Dr. Elsa Zardini for providing dried wood samples suitable for study. For L. peduncularis and L. torulosa, samples represented basal portions, but diameter was small (2 and 4 mm, respectively). The material of L. anastomosans was from a branch about 1.5 cm in diameter. Although this is much less than the 15 cm diameter reported for trunks of this species, -the branch material is considered here to represent an essentially mature pattern. Woods were boiled in water, stored in 50% ethyl alcohol, and sectioned on a sliding microtome. Sections prepared in this way were, in part, satisfactory, but cell collapse on account of thinness of wood cells was excessive in some instances. Therefore an alternative method involving further softening, embedding in paraffin, and sectioning on a rotary microtome (Carlquist, 1 982b) was employed. Sections prepared by both techniques were stained in a safranin-fast green combination. Macerations were prepared with Jeffrey's Fluid and stained with safranin. Means are based upon 25 measurements (fewer if feature is scarce) except for vessel wall thickness, libriform fiber diameter, and libriform fiber wall thickness, in which a few typical cells were measured. Vessel diameter includes the wall, although lumen diameter may be preferable for some purposes and may be calculated by subtracting wall thickness from the data presented here. Mean values for vessel grouping are obtained on the following basis: a solitary vessel = 1.0, a pair of vessels in contact = 2.0, etc. Bark was not observed specifically in the present study, although sections of the stem of L. torulosa included portions of the spongy stem covering that proves to be aerenchyma like that figured for "Jussiaea repens" L. (now a Ludwigia) by Metcalfe & Chalk (1950), and studied in detail by Ellmore (1981) for L. peploides (HBK) Raven. ANATOMICAL DESCRIPTIONS Ludwigia anastomosans (DC.) Hara, Zardini & Gentry 2175 (Figs. 1-6). Growth rings inconspicuous, and probably related to water level of the riparian habitat (Fig. 1). Mean number of vessels per mm2, 51. Mean number of vessels per group, 1.24; vessels tending to be grouped into radial multiples (Figs. 1, 3). Mean-vessel diam., 109 gim. Mean vessel wall thickness, 2.5 gim. Mean vessel element length, 458 Aim. Perforation plates simple (Figs. 2, 4). Lateral wall pitting of vessels alternate, pits crowded and circular to polygonal in outline, about 12 Am diam. on vessel-vessel interfaces (Figs. 4, 5). Vessel-axial parenchyma and vesselray pitting alternate to scalariform, pit apertures long (sometimes scalariform), pit apertures wide ("gaping"). Pitting with relatively conspicuous vesturing on vessel-vessel pits (Fig. 5, upper right), vesturing less pronounced on vessel-axial parenchyma and vessel-ray pits. Occasional vessel-vessel pits in aberrant patterns (Fig. 6). Tyloses abundant (Figs. 1, 3). Imperforate tracheary elements all libriform fibers because pits apparently simple, although a few exceptional pits with small borders also observed. Many libriform fibers septate. Mean libriform fiber diam., 28 gim. Mean libriform fiber wall thickness, 2.3 ,im. Mean libriform fiber length, 588 gim. Numerous libriform fibers with gelatinous walls (Fig. 6) and therefore probably reaction wood. Axial parenchyma-vasicentric scanty. Bands of phloem-containing axial parenchyma present in marginal positions (end of-growth rings) or scattered without any relation to growth rings (Figs. 1, 3). Rays multiseriate anduniseriate, the former slightly more frequent (indicated by relative heights in Fig. 2). Ray cells predominantly erect and square (Fig. 2), a few procumbent cells present in multiseriate portions of multiseriate rays. Mean multiseriate ray height, 2,707 gim. Mean uniseriate ray height, 349 gim. Mean width multiseriate rays at widest point, 3.95 cells. Ray cell walls moderately thin, lignified. Wood nonstoried. Raphides present in phloem-containing axial parenchyma strands. Ludwigia peduncularis (Griseb.) Gomez, Ekman 13416 (Figs. 7, 8). Growth rings absent (portion studied probably less than one year's accumulation). Mean number of vessels per mm2, 62. Mean number of vessels per group, 1.30. Mean vessel diam., 50 gim. Mean vessel wall thickness, 1.8 gim. Mean vessel element length, 484 gim. Perforation plates simple. Lateral walls of vessels with crowded alternate elliptical pits (about 4 x 8 gin) with pointed ends and narrow apertures (vessel-vessel contacts). Vessel-axial parenchyma and vesselray pitting similar but with pits longer (appearing pseudoscalariform) and wider, and with wider This content downloaded from 207.46.13.151 on Fri, 25 Mar 2016 08:49:45 UTC All use subject to http://about.jstor.org/terms 19871 CARLQUIST-WOOD ANATOMY OF LUDW IGI. 891

Keywords. - Tricalysia bridsoniana sp. nov. (var bridsoniana, Kenya; var. pandensis var. nov., Tanzania); Batopedina pulvinellata subsp. glabrifolia subsp. nov. (Zaire); Tricalysia ovalifolia (Somalia); Tricalysia somaliensis sp. nov. (Somalia); Didymosalpinx; Dictyandra congolana; Argocoffeopsis eketensis comb. nov.; raphides (taxonomic value).

Stems creeping, rather stout, rooting at the nodes. Leaves chartaceous, elliptic, obtuse or rounded at the apex, rather coarsely crenate, pinnately 3-nerved, glabrous, those of each pair often unequal in size; larger leaves up to 5.5 cm. long, 3.4 cm. wide, with petioles 1.5-3 cm. long; smaller leaves 2.5-4 cm. long, I.5-2.6 cm. wide, the petioles I-2 cm. long; upper leafsurface densely covered with minute linear raphides, the lower surface papillose; pistillate inflorescence longer than the leaves, peduncled, its branches spreading; achenes ovate, about o.9 mm. long.