Udulutsch Et Al 2009 Adenocalymma Calcareum

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Adenocalymma calcareum sp. nov. (Bignoniaceae) from BrazilianAmazonia and a key to the Amazonian species of the genus

Renata Giassi Udulutsch, Marco Antonio de Assis and Pedro Dias

R. G. Udulutsch ([email protected]) and P. Dias, PNPD/CAPES and Programa de Pos-graduacao em Ciencias Ambientais, Univ. doEstado de Mato Grosso, BR78.200-000 Caceres, MT, Brazil. M. A. de Assis, Depto de Botanica, Inst. de Biociencias, Univ. EstadualPaulista, Caixa Postal 199, BR13506-900 Rio Claro, SP, Brazil.

A new species of Adenocalymma (Bignoniaceae, Bignonieae) is described and illustrated. Adenocalymma calcareum

Udulutsch & P. Dias sp. nov. is known only from the Rondo nia State of western Brazil. Its main diagnostic features aredark yellow corolla provided with nectaries, lepidote ovary, glabrous capsule with prominently thick and calcareous wingsand wingless seeds. A discussion on the relationships of the new taxon and a key to all species ofAdenocalymmareportedfrom Brazilian Amazonia are provided.

The Bignoniaceae Juss. comprise about 860 speciesincluded in 104 genera (Fischer et al. 2004). The familyhas a pantropical distribution, with most of its speciesdiversity concentrated (ca 78% of the species) to theNeotropics (Gentry 1980). According to Gentry (1980),333 species and 56 genera occur in Brazil, which thus has amajor part of the diversity of the family.

Adenocalymma Mart. ex Meisn. nom. cons. (McNeillet al. 2006), is a Neotropical genus and one of the largestgenera of the tribe Bignonieae (47 species). It occurs fromMexico to northern Argentina (Udulutsch 2008). In Brazil,

Adenocalymma is found in 25 out of 26 states, but mostspecies occur in the northeastern and southeastern parts ofthe country. It is represented in all vegetational types,including rain forests, seasonal semideciduous forests, andsavanna formations.

Adenocalymma is morphologically distinguished fromother genera of the tribe Bignonieae by leaves with 2 or 3leaflets, simple (for the climbing species) tendrils, axillaryshoots with woody and often nectariferous prophylls [citedas pseudostipules by Gentry (1973, 1993); according to

Macbride (1961, p. 3), the term pseudostipule should beavoided] (following the terminology of Blaser 1944), andracemes. Moreover, most species have bracts, bracteoles,and calyx bearing pateliform (dark brown to black onexsiccate) nectaries.

Being lianas, most Adenocalymma species may presentsome particular difficulties for collectors (Putz 1984,Udulutsch et al. 2004). According to Gentry (1973), thosedifficulties may lead to: rarity of collections, lackof ecological/geographical information, and taxonomicproblems (e.g. new species being described based on

fragmentary material such as A. prancei A. H. Gentry,Gentry 1978).

A recent phylogenetic study of the whole tribeBignonieae by Lohmann (2006) included five species of

Adenocalymma, which, together with Gardnerodoxa Sand-with, Memora Miers, Neojobertia Baill. and Sampaiella

J. C. Gomes, emerged as a well supported clade (SMANG

clade, with 100% of parsimony and maximum likelihoodnon-parametric bootstrap and 100% of Bayesian posteriorprobability), sister to the unsupported Core Bignonieaeclade. Within the SMANG clade, there is a clade contain-ing Adenocalymma, Memora, and Sampaiella, whose sup-port varies from 91100%. However, the relationshipbetween Adenocalymma and Memora is polytomic, thus

Adenocalymmas status as a monophyletic group is stillunknown.

The first author recently prepared a taxonomic revisionof Adenocalymma, including descriptions of all species(Udulutsch 2008). The discovery of this new species is aresult from these studies and collecting work in Brazil.

Material and methods

Only fully developed structures were used for the morpho-logical description. Flowers were fixed in 70% alcohol inthe field for later measurements and illustrations. Termsused to describe two-dimensional shapes follow Hickey(1973), those used to describe indumentum are accordingto Payne (1978). Inflorescence type is according to

Weberling (1989).

Nordic Journal of Botany 27: 449453, 2009doi: 10.1111/j.1756-1051.2009.00490.x,

# The Authors. Journal compilation # Nordic Journal of Botany 2009

Subject Editor: Bertil Stahl. Accepted 6 May 2009

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Adenocalymma calcareum Udulutsch &P. Dias sp. nov. (Fig. 1)Species nova ex affinitate Adenocalymmatis gracielzae A. H.Gentry, a hac specie corolla aurantiaca, ovario dense minutelepidoto sine nectariis, valvis fructus glabris albostriatis cumala crasso indurato (calce incrustata similiter!) bene differt.

Type: Brazil, Rondonia, Presidente Medice, BR-364 high-

way, 6.5 km of road to Alvorada do Oeste, from PresidenteMedice to Vilhena, left side of the highway, roadto Embratels hill, then 14 km on the road, right side.Secondary forest. Clay soil. 11815?58.8??S, 61852?11.7??W,222 m a.s.l., 17 Nov 2005, fl. & fr., R. G. Udulutsch &P. Dias 2668 (holotype: HRCB, isotypes: MG, SPF).

Liana; branchlets drying brown, inconspicuously striate, withlenticels when older, minutely puberulous, glabrescent;axillary shoot prophylls persistent, 3.04.1)1.22.5 mm,lanceolate, apex acute, midvein prominent, puberulous,glabrescent, with 28 conspicuous dark brown to blacknectar glands. Leaves petiolate; petiole 1.35.8 cm long,semiterete, minutely puberulous, glabrescent at apex; lateralpetiolules 0.72.4 cm long, the terminal petiolule 1.53.8 cm long, semiterete, swollen at apex, puberulous,glabrescent; tendril puberulous at base; leaflet blades dryingolive and opaque above, chartaceous, margin flat andnon-cartilaginous, glabrous except for short trichomes onmidvein, venation prominent on both sides, lateral leaflets

6.817.9)2.39.4 cm, ovate to elliptic, the apex acuminateto occasionally obtuse, apiculate, base symmetric, acute toobtuse, the terminal leaflet 9.215.1)5.39.1 cm, ovate tosometimes obovate, the apex obtuse or rounded to occasion-ally retuse, apiculate, base symmetric, acute to rounded.Inflorescence a lateral raceme, pauciflorous, congested,drying olive to brown, peduncle 0.40.6 mm long, rachis(2.1() 3.712.4 cm long, puberulous, trichomes greyish,nectaries lacking, with conspicuous scares from falling

flowers; bracts 2, persistent, 1.01.7)0.61.2 mm, ovate,puberulous, lacking nectar glands or with a single dark brownto black gland, apex acute; first order bracteoles caducous,1.92.0)1.21.4 mm, symmetric, ovate, puberulous, with47 dark brown to black glands at apex, apex acuminate;second order bracteoles caducous, 2.02.5)1.11.6 mm,symmetric, obovate, puberulous, with 14 irregularly dis-tributed dark brown to black glands, apex acute. Flowerspedicelate, pedicel 0.41.0 cm long, puberulous; calyx0.81.0)0.70.9 cm, campanulate, sub-truncate, outsidepuberulous, inside papillose, lobes apiculate, with 25 nectarglands per lobe at apex; corolla dark yellow, infundibuliform,with conspicuous greenish to dark brown glands at the base

of lobes and mouth of tube, lobes 0.8

1.5)

0.8

1.3 cm,obovate, puberulous on both sides, apex retuse, tube3.16.1 cm long, 1.12.0 cm wide at the mouth, thecylindric base of tube 1.21.5 ((1.8))0.30.4 cm, tubeoutside puberulous, with dendritic trichomes, except for thebase, inside glabrous but glandulartomentose at the level ofstamen insertion; stamens as long as or longer (exserted) than

Figure 1. Adenocalymma calcareumsp. nov. (a) flowering shoot, (b) axillary shoot prophyll, (c) portion of inflorescence with flowers, (d)first order bracteole, (e) second order bracteole, (f) flower bud showing nectaries at the base of the corolla lobes, (g) disc and gynoecium,(h) ovary, longitudinal section showing two series of ovules, (i) fruit, (j) seed. (a) from Udulutsch & Dias 2780, (b) (i) from Udulutsch &Dias 2668, (j) from Udulutsch & Dias 2671. Line illustrations by R. G. Udulutsch.

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the corolla throat, the longer filaments 2.43.8 cm, shorterfilaments 1.52.4 cm, glabrous, anthers 4.55.5 mm long,staminode 4.05.5 mm long, apex ovate, membranaceous;disc 1.62.0)3.63.7 mm; ovary 3.25.8)1.61.7 mm,tetragonal in cross section, lepidote, lacking glands, ovulesbiseriate, 1112 per series, style 2.85.8 cm long, glabrous,stigma foliaceous 2.12.5)1.32.4 mm, elliptic, glabrous.Fruit drying brown, ellipsoid to seldom obovoid, with onepair of thick and hard wings along each valve; valves woody,

8.915.7 cm long, 4.46.2 cm wide (including wings),glabrous, white-striate, lacking nectaries, apex obtuse, wings0.51.6 cm wide, 3.04.5 mm thick, greyish white at thecorners, calcareous; seeds wingless, drying light brown, 2.33.6 cm long, 2.22.9 cm wide, 0.71.4 cm thick, hilumirregularly shaped, pale brown, 0.61.9)1.72.2 cm.

Distribution and habitat

This species is known from a few collections made at thetype locality, and another three from adjacent counties insoutheastern Rondonia State, western Brazil (Fig. 2). It wasfound in a secondary vegetation capoeira with some

remains of Amazonian forest (terra firme forest, Dalyand Mitchell 2000). Flowering specimens were collected inOctober, November and January, fruiting in October andNovember.

Notes

Adenocalymma calcareum is most similar to A. gracielzae A.H. Gentry. Like A. calcareum, A. gracielzae has nectary-bearing corolla, winged fruits (a character also shared by A.bracteolatum DC. and A. divaricatumMiers, Fig. 3c, 3d),and wingless seeds. Adenocalymma calcareum differs byhaving dark yellow petals (not greenish yellow as in

A. gracielzae, Fig. 4), lepidote, non-nectariferous ovary(not puberulous, nectariferous ovary as in A. gracielzae),and, importantly, white-striate fruit with prominently thickand hard wings (not inconspicuously veined fruit withmembranaceous wings as in A. gracielzae, Fig. 3b). For

more details on characters for distinguishing A. calcareum

and A. gracielzae see Table 1. In addition to the abovementioned differences, A. calcareumand A. gracielzaediffersin distribution and habitat. Adenocalymma calcareumgrowsin terra firme forests in Rondonia, and A. gracielzae isrestricted to the varzea forests of the Xingu and Itacaiunasrivers (Para State, Brazil, Fig. 2). The other two winged-fruit species are also allopatric: A. bracteolatum occurs inVenezuela, Colombia, Brazil (Roraima, Para, Mato Grosso,and Acre States), Peru, and Bolivia; whereas A. divaricatumis restricted to eastern Brazil (Fig. 2).

Besides A. calcareum, another two species are foundin Rondonia State: A. impressum (Rusby) Sandwith and

A. uleanum Kraenzl. Those species can be distinghuishedfrom A. calcareum by having included stamens, wingless

fruits, and winged seeds (not stamens as long as or longerthan the corolla throat, winged fruits, and wingless seeds asin A. calcareum). Moreover, A. impressum has glabrouscorolla and non-nectariferous calyx (not puberulouscorolla and nectariferous calyx as in A. calcareum); and

Figure 2. Known distribution of Adenocalymma bracteolatum(stars), A. calcareum (circles), A. divaricatum (squares), andA. gracielzae (triangles).

Figure 3. Fruits ofAdenocalymma. (a) A. calcareum, arrow indicates one of the calcareous wings, (b) A. gracielzae, arrow indicates one ofthe membranaceous wings, (c) A. bracteolatum, (d) A. divaricatum. (a) from Udulutsch & Dias 2771, (b) from Silva 3614 (MG), (c) fromUdulutsch & Dias 2776 (HRCB), (d) from Udulutsch & Dias 2818 (HRCB). Photos by R. G. Udulutsch.

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A. uleanum has branched inflorescences (not non-branchedinflorescences as in A. calcareum.)

The epithet calcareum makes allusion to the fruit of thisspecies that has valves with wings like limestone (Fig. 3a).

Additional specimens examined (paratypes)

Brazil. Rondonia: Ariquemes, BR-364 highway, fromCuiaba to Porto Velho, km 495, road on the right, ca2.2 km of the perimeter of Ariquemes, 12 km from thehighway, 10804?13.6??S, 62857?41.2??W, 175 m a.s.l., 19Nov 2005, fl., R. G. Udulutsch & P. Dias 2673 (HRCB).Campo Novo, Campo Novo mining, ca 100 km southwestof Ariquemes, 10834?S, 63837?W, 14 Oct 1979, fr., J. L.

Zarucchi 2690 (INPA, MO, NY). Ji-Parana, BR-364highway, 25 km of Ji-Parana, going to Porto Velho, terrafirme forest, Oct 1983, fl., W. A. Rodrigues 10440 (UEC).Presidente Medice, road to Embrapa and Embratels hill,6.5 km of road to Alvorada do Oeste, from PresidenteMedice to Vilhena on BR-364 highway, road on left side ofthe highway, then 2.25 km on the road, left side, over afence, secondary forest, clay soil, 11815?34.2??S,61852?54.3??W, 207 m a.s.l., 5 Jan 2007, fl., R. G.Udulutsch & P. Dias 2780 (HRCB, NY); clay soil withlots of gravel, 11815?49.0??S, 61851?51.2??W, 234 m a.s.l.,17 Nov 2005, fr., R. G. Udulutsch & P. Dias2671 (HRCB, SPF); 11815?49.0?? S, 61851?51.2??W,234 m a.s.l., 17 Nov 2005, fl. & fr., R. G. Udulutsch &P. Dias 2672 (HRCB, SPF).

Key to species of Adenocalymmareportedfrom Brazilian Amazonia

1. Corolla glabrous; calyx and prophylls of the axillaryshoot lacking or rarely with 13 inconspicuous nectarglands . . . . . . . . . . A. impressum (Rusby) Sandwith Corolla papillose or puberulous to villous; calyxand prophylls of the axillary shoot with nectarglands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Stamens as long as or longer (exserted) than the corolla

throat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Stamens shorter (included) than the corollathroat. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Corolla dark yellow, cylindric base of tube 1.21.5((1.8) cm long; ovary lepidote, lacking nectar glands;fruit glabrous and white-striated, wings prominentlythick and calcareous in hardness and appearance (likelimestone). . . . . . A. calcareumUdulustch & P. Dias Corolla greenish yellow, cylindric base of tube2.53.0 cm long; ovary puberulous, with nectarglands; fruit puberulous, wings membranaceous . . . .. . . . . . . . . . . . . . . . . . . A. gracielzae A. H. Gentry

4. Corolla with nectar glands on the abaxial surface to the

throat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Corolla lacking nectar glands . . . . . . . . . . . . . . 85. Leaflet margins whitish, cartilaginous; connective leaf-

like . . . . . . . . . . . . . . .A. inundatum Mart. ex DC. Leaflet margins concolored, non-cartilaginous; connec-tive apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Axillary shoot prophylls falcate. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . A. subincanum HuberAxillary shoot prophylls symmetric, never falcate . 7

7. Axillary shoot prophylls 3.5)1.2 mm, with a few(34) irregularly distributed nectar glands; inflores-cence lepidote with orange scales . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . A. prancei A. H. Gentry

Figure 4. Flowering shoots and flowers of Adenocalymma. (a)(b)A. calcareum, (c)(d) A. gracielzae. (a) from Udulutsch & Dias2780, (b) from Udulutsch & Dias 2668, (c)(d) from Udulutsch &Dias 2706 (HRCB). Photos by R. G. Udulutsch.

Table 1. Distinguishing characters of Adenocalymma calcareumand A. gracielzae.

Character A. calcareum A. gracielzae

Axillary shootprophylls

28 nectaries 12 nectaries

LeafletAdaxial surface(on exsiccate)

opaque lustrous

Abaxial surface short trichomeson midvein only

puberulous

Terminal leaflet ovate to obovate elliptic to obovate

First orderbracteoles

47 nectaries 12 nectaries

Second orderbracteoles

14 nectaries lacking nectaries

Calyx (mm) 810 5.45.7

CorollaLength of tubebase (cm)

1.21.5 (1.8) 2.53.0

Color of petals dark yellow greenish yellow

OvaryIndument lepidote densely puberulousNectaries absent presentOvules(per series)

1112 1415

Fruit valvesWing (mm) thick (3.04.5) membranaceous (0.51.5)Indument glabrous densely puberulousSurface white-striate olivebrown,

inconspicuously veined

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Axillary shoot prophylls 79)2.54.0 mm, withseveral (1420) longitudinally and linearly arrangednectaries; inflorescence lacking scales . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . A. uleanum Kraenzl.

8. Leaflet margin concolored, non-cartilaginous; fruitwinged. . . . . . . . . . . . . . . . . . A. bracteolatum DC. Leaflet margin whitish, cartilaginous; fruitwingless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Axillary shoot prophylls oblong; connective leaf-like;

ovary densely lepidote. . . .A. calderonii (Standl.) Seibert Axillary shoot prophylls spathulate; connective apicu-late; ovary lacking scales . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . A. marginatum (Cham.) DC.

AcknowledgementsThis paper is part of the PhD thesis of the firstauthor. We thank J. Prado for kindly providing the original Latindiagnosis and IBAMA for the collecting licenses no. 024/2005 &055/2006 COMON. We also thank J. Prado and L. G.Lohmann for reading an earlier draft of this manuscript. We aredeeply indebted to B. Stahl, Subject Editor, for providing manyhelpful suggestions and to E. Ljungstrand for providing the finalLatin diagnosis. RGU was supported by CNPq (proc. 140945/

2004-0) and IAPT Research Grants 2005, and PD was supportedby FAPESP (procs. 02/09762-6 and 04/15141-0).

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Udulutsch Et Al 2009 Adenocalymma Calcareum