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Taxonomy of Xylopia barbata (Annonaceae) and related species from the Amazon/Orinoco region PDF

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Preview Taxonomy of Xylopia barbata (Annonaceae) and related species from the Amazon/Orinoco region

Contr. Univ. Michigan Herb. 21-28. 22: 1999. TAXONOMY OF XYLOPIA BARB ATA (ANNONACEAE) AND RELATED FROM THE SPECIES AMAZON/ORINOCO REGION Kenneth Bagstad and David M. Johnson J. Department Botany-Microbiology of Ohio Wesleyan University Delaware, Ohio 43015 Annonaceae Xylopia, a pantropical genus of characterized by axillary inflo- rescences, long narrow petals, and dehiscent fruits, well-represented in the Neo- is tropics, where about 50 species are known to occur (Fries 1959). One of the more named distinctive Neotropical species X. barbata Hoffmanns, ex Martius, for is brown the shaggy or brownish yellow hairs that clothe the twigs and lower surfaces of the leaves; also has especially long and narrow flowers. Fries (1931), in it describing X, conjungens, noted the similarity of leaf and flower pubescence, its floral structure, and leaf venation to those of X. barbata, but observed that the X Many leaf shape was similar to that of frutescens Aublet. recent specimens with this combination of distinctive pubescence and elongate flowers are either inter- mediate between X, barbata and X, conjungens in leaf shape or in neither fit taxon. As a result of a herbarium study, we place X, conjungens in synonymy under X, barbata, and recognize two new species, X. rigidiflora and X. orinocensis. The long hairs and narrow petals of these three species are a unique combination of among character states Neotropical xylopias, suggesting that these species form a common The natural group. pollen tetrads are of a type Xylopia (Kathleen in Peterson, pers. comm.), and the seeds bear the bilobed that characterizes the aril largely Neotropical Section Xylopia. All three species are restricted to sandy perhaps group was habitats, indicating that the ancestor of this a habitat specialist. Fries (1959) did not indicate the relationship of X. barbata or X. conjungens to them means other species, choosing instead to group by of a key, with artificially, narrow and species that share petals, free sepals, elliptic to oblong-elliptic leaf blades. Intrageneric relationships of the Xylopia barbata group remain unclear. Key Group to the of the Xylopia barbata Species cm Leaves 2.3-3.8 wide, concolored or only slightly darker adaxially than abaxially; leaf venation 1. with loops outside the closed arches of the secondary veins ("festooned-brochidodromous''); mm outer petals 2.7-3.2 wide midpoint, fleshy and somewhat woody. X. at rigidiflora cm Leaves 0.8-2.3 wide, discolored, distinctly darker adaxially than abaxially; leaf venation 1. mm L3-L9 without regular loops adorning the secondary vein arches; outer petals wide at midpomt, coriaceous to fleshy. Lamina base cuneate to attenuate and minutely decurrent on petiole; corolla orange, 2. pubescence not brown. X. orinocensis 2. Lamina base cuneate to broadly cuneate but not decurrent on petiole; corolla yellow, with brown pubescence. X, barbata 21 — — VOLUME HERBARIUM CONTR. UNIVERSITY OF MICHIGAN 22 22 Xylopia barbata Hoffmannsegg ex Martius, Fl. bras. 13(1): 40. 1841. Xylopicnim barbatiun (Hoffmannsegg ex Martius) Kuntze, Revis. gen. 1891. pi. 8. 1; Type: Brazil. [Para:] "In ripa fluvii Tocantins, locis sabulosis virgultosis, MO! M, ad Camcta," Sieber [J5J] (leclotype, here designated: photos: F! NY!; BR). isolectotype: Xylopia conjungens Acta Horti Berg. 1931. Type: Peru. Loreto: Fries, 10: 333. Feb-Mar Mishuyacu, 100 m, near Iquitos, 1930, Khig 970 (holotype: B; NY!). IF-H. isotypes: F! Fig. DBH m m Shrub 2.5 to medium-sized tree 25 6-41 cm; bark pale red, tall tall, mm wrinkled (ex Benko-Tseppon). Leaf-bearing twigs 0.4-2.5 diameter, tomen- in mm-long tose-velutinous to glabrate, the hairs golden to straw-colored, dotted 1 brown Lamina cm with light circular Icnticels. of larger leaves 3.2-7.3 long, 1-2.3 cm wide, subcoriaceous, elliptic or oblong, paler abaxially, cuneatc to broadly cuneate and not decurrent on petiole at the base, acuminate, acute, obtuse, or mm acumen, retuse at the apex, the present, 1.5-14 long, glabrous adaxially, if sparsely pubescent to puberulent abaxially, midrib tomentose abaxially, margins puberulent to tomentose; midrib impressed to plane adaxially, raised and semiter- ete abaxially; secondary veins 6-12 per side, departing at 45-75*^ from midrib, mm indistinctly brochidodromous, anastomosing (1.2-) 1.6-3.9 from margin; sec- ondary and higher-order veins indistinct, raised adaxially, slightly raised abaxially. mm mm Petiole 0.6-2.5 long, 0.4-1.4 wide, tomentose-velutinous puber- terete, to ulent. Inflorescences Lflowered, occasionally 2-3-flowcrcd from a very short pedun- mm mm cle, axillary; pedicels 1.4-3.5 long, 0.5-1.4 thick, bracts 2-3, occasionally mm Buds 1.9-3.8 long, ovate, clasping, tomentose-velutinous to glabrate. linear- 4, mm mm lanceolate, acute. Calyx 3.2-5.8 long, 3.8-7.4 diameter, cup-shaped, in mm mm coriaceous, sparsely tomentose-velutinous; lobes 1.8-3.5 long, 1.8-3.7 wide, apiculate apex. Corolla yellow, brown- or maroon-pubescent, coriaceous at tomentose to fleshy, to thickly velutinous; outer petals spreading slightly anthe- at mm mm cm 1.4-2.2 long, 2.2-3.1 wide at base, 1.3-2.2 wide at midpoint, linear, sis, acute at apex, expanded and concave at base, glabrous adaxially, finely appressed mm cm pubescent abaxially; inner petals 1.3-1.9 long, 1.8-2.6 wide base, 0.6-0.8 at mm wide at midpoint, linear-filiform, acute at apex, expanded and concave at mm rhomboidal Stamens base, in cross section, puberulent. 50-80, 1-1.2 long, mm narrowly oblong, glabrous; anthers septate at anthesis; apex of connective 0.2 mm long, truncate or slightly hemispheric; filament 0.3-0.4 long; innermost sta- mens staminodial, appressed to the stigma bases. Carpels 4-5, seated in the con- mm cavity of the torus with only stigmas emergent; ovaries Ll-1.2 long, long- mm 2-3 sericeous; ovules attached laterally; stigmas 4.2-5.6 long, coherent, (-6), mm filiform, pubescent. Torus 1.8-2.5 in diameter, conical, deeply concave in mm center, glabrous. Fruit of 2-4 monocarps borne on pedicel 4.9-5.9 long, 1.4- a mm tomentose 1.7 thick, to glabrate, with sepals persistent; torus o\ fruit 3.2-4.2 mm monocarps in diameter; dehiscent, yellow or green with a red interior, 2.2-4 mm cm long, 4.6-7.5 wide, oblong, falcate to semi-torulose, surface verrucose and somewhat with the venation of the pericarp raised, puberulent to glabrate; stipe mm mm mucro 4-5.5 long, 1.8-2 wide midpoint; apex acute, with an oblique at mm mm mono- 1.2-3 long; pericarp ca. 1 thick. Seeds 3-6, at 75"" to long axis of mm mm mm carp, 9.8 long, 4.2 wide, 3.7 thick, ellipsoid, elliptic in cross section, mm black or dark brown, smooth and somewhat shining; white, bilobed, 2 aril mm long, 3.8 wide. — BAGSTAD AND JOHNSON: XYLOPIA 1999 23 FIG. Xylopia barbata, X. rigidiflora, and X. orinoceiisls. A-B, X. orinocensis. A. Habit, xl. B. 1. Close-up of pedicel, calyx, and petal bases, x5. C-E, X. rigidiflora. C. Stamen, x20. D. Staminode, F-H, x20. E. Habit, xl. X. barbata. F. Habit, xl. G. Leaf of Peruvian specimen, xl, H. Seed, lateral view, x2.5. Based on: A-B, Yanez 172 (US); C-E, Silva 221 (INPA, NY); F, Benko-Iseppon 373 (OWU); & G, Vdsqiiez Jaramillo 15259 (MO); H, Prance al 5433 (US). et Additional Specimens Examined. Brazil. Amazonas: Rio Urutii [Urubu?-sec note below], Froes margem 25487 (NY); Manaus, do Ig. dos Franceses, Mello 3177 (U); Enseada Grande, Ponta Negra, margem do rio, Rodrigiies & Lima 2205 (F); Manaos, Rio Negro, Ule 5428 (K). Para: Mpio. de Santarem, Vila de Alter do Chao, 2^31'S, 55°00'W, Albernaz 148 (K, NY, U); Mpio. Alter do Chao, km em 69 a sul de Santarem, 'ilha" do Rio Tapajos, cerca de 200 ni distante da principal rua do (OWU, Mpio. Alter do Chao, Benko^Iseppon 373 SPF, U); Mpio. de Santarem, Vila de Alter do — VOLUME HERBARIUM CONTR. UNIVERSITY OF MICHIGAN 22 24 Kuhhmmn Chao, 2''3rS, 55°()0'W, Ferreira 054 (K, NY, U); Santarem, 1736 (NY, U); Sla. Terezinha, & margcm dircita do Rio Tapajos, Maciel Cordelro 137 (NY, U); in siccis arenosis tempori pluviali Nov NY, inundalis Sanlarcm, Riedel 1566 (NY, US); vicinibus Santarem, 1849, Spruce 388 (K, in G K B fragments ex and at F; only the sheet from Herb. Benlh. at bears date and collection number); Dec Le)\ver Amazons, campos near Santarem, Trail! 8 (K-2 sheets, one dated 21 1873, the other 25 Dec 1873); Santarem, Bellerra, as proximidades do igarape Iru(janga, entre Pindobal e Porto Novo, MO, NY, Vilhena ul 162 (NY, U). Rondonia: vicinity of Mutumparana, Prance al 5433 (K, U, et et (OWU). US); margin of Mutumparana Prance 8851 Peru, Loketo: Maynas, Mishana, airstrip. et al. 3°55'S, 73^35'W, Vasquez et al. 4048 (MO); Iquilos, Allpahuayo, Estacion Experimental del Instiluto & de Investigaciones de Amazonia Peruana, 4°10'S, 73°30'W, Vasquez Jaramillo 15259 (MO), la (MO, 15268 U). Flowering specimens have been from October through March, and one collected Peruvian specimen was collected with buds in May. Fruits have been collected in July known and November. Xylopia barbata from Loreto Department Peru, and from in is Amazonas, and Rondonia the Brazilian states of Para, (Fig. occurring at elevations 2), The of 100-180 meters. usual habitats of X. barbata are praia arenosa, restinga, and savanna forest, sandy open habitats along rivers, which are widely distributed i.e., & Amazon and numerous endemic Prance within the basin contain species (Pires f985). The type oi X, conjungens, as well as subsequent collections from Loreto, Peru, X differ from most material of barbata in having distinctly acuminate leaf apices and golden spreading pubescence on the twigs and abaxial midribs of leaves. These speci- X m mens were collected from trees 20-25 In contrast, plants of barbata from all tall. m the vicinity of Santarem are typically shrubs or small trees 2.5-10 with acute to tall, obtuse or even retuse leaf apices and brown or brownish yellow pubescence. These characters seem vary independently, however: the two Prance specimens from to Rondonia have golden hairs, but the leaves are acute to obtuse, and the trees were m Amazonas 10-11 the Froes specimen from has acute leaf apices and from a tall; is m We tree 12 but hairs are not conspicuously golden. have therefore conclud- tall, its ed that arc best treated as a single species for which there seems to be clinal all variation in size, shape of the leaf apex, and color of the pubescence. The Amazo- specimen collected by Froes from the ''Rio Urutu'' in the stale of Amazon nas in fact, probably from the Rio Urubii, a river that enters the just is, downstream from Manaus: Froes number 25355 from Urubu on collected the his September 22 1949, just 13 days earher. Martius based Xylopia barbata on specimens collected by Sieber the (1841) in Cameta and by Riedel Santarem, and used Hoffmannsegg's unpub- vicinity of at hshed binomial. Hoffmannscgg was also the source of the Sieber collections seen by Martius (Lasegue 1845), and the Sieber syntype accordingly chosen as lecto- is number BR; type. Fries (1930) cited a Sieber specimen without collection at the Museum specimen in the photograph distributed by the Field attributed to the is Munich Herbarium and shows number ''353." a collection tag bearing the M B Amazo- Type: frazil. nas: Mpio. Barcelos, Rio Jauari, 00^42'N, 63°22'W, abaixo do entronca- mento com INPA!; igarape Pretinho, 2 Jul 1985 Silva 221 (holotype: (fl), IC-E. isotypes:NY! U!). Fig. ambabus Species X, barbatae X. orinocensi ab laminis foliorum et similis, mm concoloribus vel leviter discoloribus 2.3-3.8 arcubus adjectis praeter latis, venarum medio arcus secundariarum, exterioribus carnoso-lignosis 2.7- et petalis mm 3.2 latis differt. BAGSTAD AND JOHNSON; XYLOPIA 1999 25 TROPICAL AMERICA map Flora Neotropica base no 1 C 979 by thB Univarsity o( Utrsc^it Published by the Slale Univeriily of UtracKI, the Netherlands Deportment of Systomaljc Botany 1 FIG. 2. Distribution oi Xylopia barbcifa, X. rigidiflora, and X, orinocensis in South America. m mm Shrub 2-4 Leaf-bearing twigs 0.6-2.1 diameter, tomentose- in tall. mm velutinous or puberulent to glabrate, with grayish hairs ca. 1 long persistent on younger twigs, marked with occasional circular cream-colored lenticels. Lami- cm cm na of larger leaves 5.3-9.1 long, 2.3-3.8 wide, subcoriaceous coriaceous, to elliptic or occasionally oblong, concolored or slightly darker adaxially than abaxi- ally, cuneate but not decurrent on petiole at base, cuspidate to short-acuminate at mm apex, the acumen 5-15 long, glabrous adaxially, puberulent with a tomentose midrib abaxially, margins puberulent to sparsely pubescent; midrib impressed to plane adaxially, raised and semiterete abaxially; secondary veins 9-12 per side, departing 55-80° from midrib, festooned-brochidodromous, anastomosing 2.5- at mm 8.9 from margin; secondary and higher-order veins indistinct, raised adaxially, mm mm tomen- sHghtly raised abaxially. Petiole 1.2-2.0 long, 0.6-1.5 wide, terete, mm tose or puberulent. Inflorescences 1-flowered, axillary; pedicel 2.4-3.0 long, mm mm 0.8 thick, tomentose; bracts 2-3, 3-3.5 long, clasping, ovate, tomentose. mm mm Buds Hnear-lanceolate, obtuse. Calyx 4.3-6.5 long, 5.6-8.3 in diameter, mm cup-shaped, tomentose coriaceous, to puberulent; lobes 2.6-4.1 long, 2.7-3.6 mm wide, apiculate at apex. Corolla yellow, fleshy and somewhat woody; outer mm cm spreading wide petals slightly at anthesis, 1.5-2.1 long,, 3.9-4.6 at base, mm mm 2.7-3.2 wide and 1.1 thick at midpoint, linear-lanceolate, acute and slightly trigonous apex, expanded and concave puberulent at slightly at base, adaxially. — HERBARIUM VOLUME CON UNIVERSITY OF MICHIGAN 26 TR. 22 mm cm sericeous and keeled abaxially; inner petals 1.8-1.9 long, 2.8-3 slightly mm wide at base, 1.6-1.8 wide at midpoint, linear, acute at apex, expanded and concave rhomboidal puberulent on pubcr- base, cross section, keel adaxially, at in mm ulent abaxially. Stamens 1-1.2 long, narrowly oblong, glabrous; anthers sep- mm tate at anthesis; apex of connective 0.15-0.3 long, truncate or sHghtly hemi- mm spheric, papillate; filament 0.2-0.3 long; innermost stamens staminodial, appressed to stigma bases, flattened, oblong or quadrate. Carpels seated in the concavity 9, mm of the torus with stigmas and upper half of ovaries emergent; ovaries 2 long, Torus lanceolate, long-sericeous; ovules stigmas coherent, filiform, pubescent. 3; mm mm diameter, tomentose around circumference, center portion 3.6 in 1.9 in mm diameter, 0.9 high, conical, deeply concave. Fruits and seeds unknown. kms Additional Specimens Examined. Brazil. Roraima: Mpio. Sao Luiz do Aiiaua, cnlrc 350 e Aug 355 da Estrada Manaus-Caracarai (BR-174), proximo ao Equador, O^UO', 6()°45'W, 21 1987, Cid Fcnciya9064 (OV^V.V). Both collections are of flowering material, one from July and the other from August. Both were collected from lowland campina habitat on sandy northern soil in Brazil near the frontier with Venezuela (Fig. This area inhabited by a num- 2). is endemic woody Pagamea ber of other species: aracaensis (Rul^iaceae), Coiiepia amamlae and Licania nelsonli (Chrysobalanaceae), Pithecellobium prancei (Mimos- aceae), Ternstroemia canipinicola (Ternstroemiaceae), and Caraipa longisepala & (Clusiaceae) (Prance Johnson 1992). The and petals of X. rigidiflora are, as specific epithet suggests, thicker its stiffer than those of the other tw^o species treated here; the outer petals are also The wider. leaf markedly wider, with formation of strong loops outside the is arches of the secondary veins; in the type specimen the leaves are strongly cuspi- The date. leaves are concolored or only slightly darker adaxially than abaxially, whereas two the leaves of the other species are distinctly discolored. & Xylopia orinocensis K. Bagstad D. M. Johnson, nov. Type: Venezuela. sp. km SE Amazonas: Dpto. Atabapo, Alto Rio Orinoco, 25 de Esmeralda, la Aynmrd & 3"03'N, 65^25"W, 80 m, f7 Feb 990 Delgado 7921 (holo- f f (fl), MO!; NY! lA-B. type: isotypes Ul). Tig. Species X. barbatae valde affinis, a qua imprimis laminis foliorum angustiori- bus basi attenuatis et minute decurrentibus, staminibus carpelliscjue leviter brevior- ibus, el floribus aurantiacis differt. m DBH Tree 10-21 talk up to 25 cm; bark light gray. Leaf-bearing twigs 0.4- mm 1.9 in diameter, flexuous, thickly velutinous, the dense golden patent hairs 1- mm marked brown Lamina 1.5 long, persistent, with circular light lenticels. of cm cm larger leaves 3.5-6.3 long, 0.8-1.5 wide, subcoriaceous, oblong to lan- ceolate, occasionally paler abaxially, cuneate to attenuate and minutely elliptic, acumen decurrent on petiole at base, acuminate to cuspidate at apex, the 3.5-10 mm long, glabrous adaxially, sparsely pubescent to tomentose with midrib tomen- tose-w^ooUy to puberulent abaxially, margins puberulent to tomentose; midrib im- pressed to plane adaxially, raised and semitercte abaxially; secondary veins 7-11 per departing 40-70° from midrib, brochidodromous, anasto- side, at indistinctly mm mosing 1.6-3 from margin; secondary and higher-order veins indistinct, raised mm mm adaxially, shghtly raised abaxially. Petiole 0.5-1.7 long, 0.4-0.7 wide, BAGSTAD AND JOHNSON: XYLOPIA 1999 27 tomentose tomentose-woolly. terete, to Inflorescences 1-2-flowered, axillary; mm mm pedicel 1.6-4.2 long, 0.6-0.8 thick, tomentose to tomentose-woolly; bracts mm Buds 2-3, 1.9-4 long, ovate, clasping, pubescent. linear-lanceolate, acute. mm mm Calyx 2.6-4 long, 3.5-4.5 in diameter, cup-shaped, coriaceous, tomentose- mm mm woolly tomentose; to lobes 1.1-2.2 long, 1.8-2.4 wide, apiculate apex. at Corolla orange, coriaceous outer spreading to fleshy; petals shghtly anthesis, at mm mm cm 1.7-2.4 long, 1,4-3 wide base and 1.3-1.5 wide midpoint, at at linear- lanceolate, acute at apex, cuneate at base, glabrous adaxially, appressed-pubes- mm mm cm cent abaxially; inner petals 1.8-2.0 long, 1.3-1.6 wide at base, 0.5-0.6 wide at midpoint, linear-filiform, acute at apex, expanded and concave base, at rhomboidal in cross section, glabrous adaxially, sparsely pubescent abaxially. Sta- mm mens ca. 60, 0.8-1 long, narrowly oblong, glabrous; anthers septate anthe- at inner stamens staminodial. Carpels 4-5, seated concavity of the ova- sis; in torus; mm mm ries 0.7 long, long-sericeous; ovules stigmas 3.5 long, coherent, fihform, 2; mm Torus pubescent. 1.6 in diameter, conical, deeply concave, pubescent around circumference. Fruits and seeds unknown. Additional Specimens ExAiMiNED. Venezuela, Amazonas: Prov. Alto Orinoco, Cano Surimoni ca. 30 mill, downstream by boat (40 PS) from La Esmeralda, right tributary of Rfo Orinoco, 3°12'N, Nov km eS'^SS'W, 26 1995, Rainer 282 (U); Rfo Caname, a 36 de confluencia con Rio Atabapo, la el Nov 3"37'N, 67"07'W, 1989, Yanez 172 (US). Plants were collected from forests in the upper Orinoco River drainage of southern Venezuela (Fig. quartzite outcrops were noted in one locality. All 2); three specimens bear flowers two were November and one only; collected in in February. Xylopia orinocensis has distinctly long, narrow leaves with a more at- tenuate lamina base than either X. rigidiflora or X. barbata, and the lamina is minutely decurrent on the petiole. The dimensions of the flower parts are similar to those of X. barbata. This species not included in the recently pubhshed Flora is Guayana of the Venezuelan (Steyermark but the combination of spreading 1995), cm golden pubescence, small leaves, and narrow petals over long should 1,7 dis- tinguish from any other species oi Xylopia in the Flora area. it ACKNOWLEDGMENTS Funding for study of these species was provided by the Ohio Wcsleyan University Howard Hughes Program funded by a grant from the Howard Hughes Medical Institute Undergraduate We Biological Sciences Education Program. thank Nancy A. Murray comments and help with for for the Froes locality. Kathleen Peterson provided us with unpubUshed pollen data. The following her- OWU, baria made specimens available for study: F, INPA, K, MO, NY, SPF, U, US. LITERATURE CITED Fries, R. E. 1930. Revision der Arten einiger Anonaceen-Gattungen-I. Acta Horti Berg. 1-128. 10(1): 1931. Revision der Arten einiger Anonaceen-Gattungen-IT. Acta Horti Berg. 129-341. 10(2): . 1959. Annonaceae. In Die nalurUchen Pflanzenfamilien, ed. H. Melchior, 2d ed., ]7all: 1-170. . & Duncker Humblot. Berlin: Lasegue, A. 1845. Musee botanique de M. Benjamin Delessert. Paris: Librairie de Fortin, Masson et Cie. Martins, C. F. P. de. 1841. Anonaceae, In Flora brasiliensis, ed. C. Martius, 13(1): 1-64. Munich, Leipzig; F. Fleischer. Pires, M., and G. T. Prance. 1985. The vegetation types of the Brazilian Amazon, In Key environ- J. ments: Amazonia^ ed. G. T. Prance and T. E. Lovejoy. Oxford: Pergamon Press. VOLUME MICHIGAN HERBARIUM CONTR. UNIVERSITY OF 22 28 Prance, G. and D. M. Johnson. 1992. Plant collections from the plateau of Serra do Araca T., Kew (Aniazonas, Brazil) and their phytogeographic affinities. Bull. 47: 1-24. & Steyermark, A. 1995. Xylopia. In Flora of the Venezuelan Cuayana, vol 2: PteridopJiytes Spcr- J. matophytes (Acanthaceae-Araceae). ed. P. E. Berry, B, K. Hoist, and K. Yatskievych. Portland, Oregon: Timber Press.

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