A new Heteropterys () from semideciduous forest, with notes on wood anatomy Author(s): André M. Amorim, Lucas C. Marinho, Cleiton Pessoa and Marcelo R. Pace Source: Systematics and Evolution, Vol. 303, No. 2 (February 2017), pp. 177-185 Published by: Springer Stable URL: https://www.jstor.org/stable/44854165 Accessed: 21-06-2021 11:33 UTC

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This content downloaded from 86.59.13.237 on Mon, 21 Jun 2021 11:33:21 UTC All use subject to https://about.jstor.org/terms Plant Syst Evol (2017) 303: 177-185 (JM J CrossMark DOI 10.1007/S00606-0 16- 1360-0

ORIGINAL ARTICLE

A new Heteropterys (Malpighiaceae) from semideciduous forest, with notes on wood anatomy

André M. Amorim1,2,3 • Lucas C. Marinho2,3 • Cleiton Pessoa2 • Marcelo R. Pace4,5

Received: 11 May 201 6 /Accepted: 12 October 2016 /Published online: 4 November 2016 © Springer- Verlag Wien 2016

Abstract This paper describes and illustrates Keywords Heterop- Lianas • Inselbergs • • terys serrata , a new species endemic to semideciduous Metallophyllis informal group • South America forests associated with inselbergs in the state of Espírito Santo, Brazil. Morphological and wood anatomical traits of the new species are compared to those of other Introduction species of the Heteropterys Metallophyllis informal group. Based on wood anatomy, H. serrata and H. nitida (the Heteropterys most mor- Kunth is the largest of Malpighiaceae phologically similar species) have different with axial ca. 151par- species (Amorim 2003; Anderson 2013; enchyma, which is scarce paratracheal in H. Pessoa nitida and (andAmorim 2016). The genus as circumscribed by some other species of the Metallophyllis informal Anderson group) and Davis (2007), with Clonodia W.R.Anderson and aliform confluent in H. serrata. The most as a synonym, is monophyletic with fairly strong bootstrap notable morphological and anatomical characters support that (Davis dis- and Anderson 2010) and includes several tinguish the new species are the young hexagonal infrageneric stems, clades (with modest support) as proposed by the unusual widely serrate leaf margins and Niedenzuthe aliform (1928) (i.e. Heteropterys ser. Metallophyllis confluent axial parenchyma in the wood. Nied., Heteropterys subsect. Aptychia Nied, and sub- sect. Stenophyllarion Nied.). Despite advances based on phylogenetic studies and nomenclatural adjustments over the last decades, the of Heteropterys is still Handling editor: Ricarda Riina. poorly resolved and its biodiversity poorly understood, as attested by the discovery of several recently described Kl André M. Amorim amorim.uesc @ gmail.com species (Sebastiani and Mamede 2010; Anderson 2014; Pessoa and Amorim 2016). 1 Departamento de Ciências Biológicas, Universidade Estadual New species of Malpighiaceae have been consistently de Santa Cruz, Km 25, Ilhéus-Itabuna, Ilhéus, reported from forests along the eastern coast of South Bahia 45662-900, Brazil America, where at least 22 species have been described in 2 Herbario Centro de Pesquisa do Cacau, CEPEC, Km 22, the last 16 years (i.e. Amorimia W.R.Anderson - 2 spp., Rodovia Ilhéus-Itabuna, Itabuna, Bahia 45600-970, Brazil Heteropterys - 10 spp., Diplopterys A.Juss. - 1 sp., Hiraea 3 Programa de Pós-Graduação em Botânica, Universidade Jacq. - 4 spp., Lophopterys W.R.Anderson & C.C.Davis - Estadual de Feira de Santana, Avenida Transnordestina, Novo Horizonte, Feira de Santana, Bahia 44036-900, Brazil 1 sp., Mcvaughia W.R.Anderson - 1 sp. and Stigmaphyllon A.Juss. - 3 spp.). The high number of taxonomie novelties 4 Departamento de Botânica, Universidade de São Paulo, Rua do Matão, 277, Cidade Universitária, São Paulo, of angiosperms from these forests, which has been high- São Paulo 05508-090, Brazil lighted by several authors (Sobral and Stehmann 2009; Goldenberg et al. 2016), can be partly attributed to recent 5 Department of Botany, National Museum of Natural History, Smithsonian Institution, PO Box 37012, Washington, botanical expeditions in previously inaccessible or unex- DC 20013-7012, USA plored areas.

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Deciduous Fig. 1 Heteropterys and serrata - a, c-e and H.semideciduous nitida - b, f-h wood^ forest patches in eastern South America anatomy: a trans verse section with general vieware of secondary xylem, good examples of unexplored areas that growth rings delimited by a narrow to broad band of axial lack botanical parenchyma ( arrowheads ), gelatinous fibresrecords. present (asterisk)-, Of the ten new species of Het- eropterys b transverse described section with general view of the secondary xylem, from eastern Brazil over the last dec- ades, three diffuse porous, gelatinousare fibres present (asterisk)-,restricted c detail of to deciduous or semideciduous forests (e.g. secondary xylem, showing growth H. rings delimited by marginalandersonii (vs. Amorim (Amorim 2004), H. axial) parenchyma (arrowhead), wide vessels solitary or in radial or brunnea tangentialSebastiani multiples of 2, narrow vessels forming long radial chains & Mamede (Sebastiani and Mamede 2010) and H. arcuata C.Pessoa & Amorim (Pessoa and (arrow), parenchyma vasicentric to aliform; d longitudinal tangential Amorim 2016)). Studies of herbarium collections and section, showing non-storied rays, 3-4 cells wide, prismatic crystals in chambered axial parenchyma cells present (arrowhead)', e longitu- extensive fieldwork in the state of Espírito Santo, Brazil, dinal radial section; ray heterocellular, mixed, with procumbent, have been carried out to resolve the taxonomy of prob- square and upright cells across the ray, fibres septate (arrow), lematic taxa within Heteropterys . The present paper crystalliferous parenchyma abundant (arrowhead)-, f detail of the describes a new species of Heteropterys , known only from secondary xylem, showing growth rings marked by marginal axial semideciduous forests in Espírito Santo, and provides parenchyma in narrow bands (arrowheads), wide vessels associated with narrow ones, narrow vessels forming long radial chains (arrows), information on wood anatomy of the genus that will be paratracheal parenchyma scanty; g longitudinal tangential section, useful in future taxonomie and phylogenetic analyses. showing non-storied rays, 3-4 cells wide, prismatic crystals in chambered axial parenchyma cells present (arrowhead)', h longitudi- nal radial section, ray heterocellular, mixed, with procumbent, square Materials and methods and upright cells across the ray, fibres septate (arrow). Scale bars : a, b = 1 mm, c-f = 500 ļim, g-h = 200 pm. Photographs a-h by Marcelo R. Pace The new species was observed in situ and vegetative samples were dried and fixed in FAA 70 for laboratory analyses (Johansen 1940). The morphological description Results and the illustration are based on specimens from CEPEC, MBML, NY, RB and SPF (abbreviations follow Thiers The new species of Heteropterys is described based on 2016). The distribution map was made using the website several collections made in semideciduous forest in the SimpleMappr (Shorthouse 2010), and the conservation state of Espírito Santo, Brazil. Based on its morphology, status was assessed based on IUCN (2012) criteria. this species is considered part of the Metallophyllis infor- The wood anatomy was studied using samples of stems of mal group (= Heteropterys ser. Metallophyllis , sensu Nie- the new Heteropterys and H. nitida (Lam.) DC., which were denzu (1928)) because of the presence of abaxially golden- collected from the basal portion of the plant (ca. 2 cm diam- sericeous leaf lamina, well-developed peduncles, subequal eter), immediately fixed in FAA 70 (Johansen 1940) and basally sericeous filaments and straight or slightly diver- stored in a 70% ethanol solution. Prior to sectioning, the wood gent (distally) styles. The most notable characters that samples were softened in boiling water and glycerine for up to distinguish the new species from other species of the group a week (8 h/day) and later embedded in polyethylene glycol are the young hexagonal stems and leaves with wide, ser- 1500. The samples were then sectioned using a slide micro- rate leaf margins. tome with the aid of a polystyrene resin, which serves to The wood anatomy of the new species is characterized preserve the wood anatomy when soft tissues are combined by the following: distinct growth rings, marked by a narrow with hard tissues (Barbosa et al. 2010), a phenomenon com- to broad marginal band of axial parenchyma (1-5 cells mon in lianas. The anatomical sections were double-stained wide; Fig. la, c) and wide vessels (Fig. la); porosity with safranine and Astra blue (Bukatsch 1972). semidiffuse to diffuse (Fig. la); vessels with no particular Wood descriptions follow the recommendations of the arrangement (Fig. la, c); vessel dimorphism with wide IAWA Committee (1989). Measurements were obtained vessels associated with narrow vessels (Fig. lc); wide using the free software ImageJ 1.45s (http://imagej.nih. vessels solitary or in radial or tangential multiples of 2-3 gov/ij), with a minimum of 30 repetitions per specimen. (Fig. la, c); narrow vessels usually in radial chains of 5-7 Vouchers of anatomical study: Heteropterys sp. nov. - cells (Fig. lc); perforation plates simple; intervessel pits Brazil, Espírito Santo, Mun. Vila Pavão, estrada Vila alternate, small (5 ± 0.6 pm), vestured; vessel-ray pitting Pavão para Cristalina, margem do Rio Cricaré, 18°42'17"S similar to intervascular pits in size and shape; vessel 40o34'33"W, 9 Jun 2015 (ste), A.M. Amorim et al. 9578 diameter 135 ± 34 pm in wide vessels and 23 ± 5 pm in (CEPEC!). Heteropterys nitida - Brazil, Rio de Janeiro, narrow vessels; vessel element length 340 ± 44 pm; wide Mun. Silva Jardim, Reserva Biologica Poço das Antas, vessel frequency 6 ± 3; tyloses absent; fibres thin to thick estrada da Barragem, 22°32'43"S 42°18'01"W, 11 Sep walled (Fig. la, c), septate (Fig. le), occasionally gelati- 2012 (bud), M.R. Pace 226 , 233 (CEPEC !, SPF!). nous (Fig. la); axial parenchyma vasicentric to aliform

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(Fig. la, c), in marginal heterocellular, narrow mixed, with procumbent,bands square on and uprightthe limits of growth rings (Fig. la, c); cells axialmixed throughout parenchyma (Fig. le); prismatic crystals with large 4-5 cells per strand; perforated andray abundant, cells present inpresent; chambered upright rays ray cells and non-storied, in 3-4(-5) cells wide, less than 1 mm (Fig. Id); rays chambered axial parenchyma cells (Fig. Id, e).

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Heteropterys 2015). From what we know, nitida the presence of narrow vessels in , which is the species most morpho- logically similar radial chains is taxonomically informative to at the family the and new taxon, has wood anatomy char- acterized by even the genus the level. following: distinct growth rings, marked by a narrow marginal Regarding features that distinguish both species, the mostband of axial parenchyma (Fig. lb, f); porosity remarkablediffuse are the parenchyma type and crystal abundance.(Fig. lb); vessels with no particular arrangement Heteropterys nitida has the same(Fig. type described for all other lb, f); vessel dimorphism with wide vessels associated closely related species of the Heteropterys Metallophylliswith narrow vessels (Fig. If); wide ves- sels solitary informal group, withor the presence in of scarce paratrachealradial or tangential multiples of 2-3(^1) (Fig. lb, f); parenchyma narrow and occasional crystals in rays and chambered vessels usually in radial chains of 7-10 cells (Fig. lb, axial parenchyma f; cells (Dominguessometimes 2008). Heteropterys in clusters), with the first narrow vessel of the serrata , on the otherchain hand, differs by the presence in of contact with the wide vessel (Fig. If), or less often abundant vasicentric toindependent aliform confluent parenchyma and of the wide vessels; perforation plates simple; considerably more abundant intervessel crystals. Parenchyma distribu- pits alternate, small (6 ± 0.4 pm), vestured; tionvessel-ray is one of the most informative characters in the wood ofpitting similar to intervascular pits in size and shape; Malpighiaceae, with species thatvessel have very scarce par- diameter 166 =b 25 pm in wide ves- sels and 29 enchyma ±(e.g. Byrsonima 7 Rich.pm ex. Kunth (Record andin Hess narrow vessels; vessel element length 234 ± 66 1943;pm; Sonsin et al. 2014)) to aliformwide confluent parenchyma vessel frequency 12 =b 1; tyloses absent; fibres forming bands (e.g. Bunchosiathin Kunth (Metcalf and toChalk thick walled, septate (Fig. lb, f), some gelatinous 1950)) or(Fig. occasionally apotracheal diffuselb); parenchyma (e.g.axial paratracheal parenchyma scanty (Fig. If), Burdachiain A.Juss. narrow and Ptilochaeta Nees (Metcalf and Chalk marginal bands on the limits of the growth rings 1950)). To date, Heteropterys (Fig. serrata is the only knownlb, f); axial parenchyma with 4-5 cells per strand; species withinperforated the Metallophyllis informal group that has ray cells present, rays non-storied, rays 3 cells wide, vasicentric to aliform less parenchyma. than 1 mm (Fig. lg); rays heterocellular, mixed, with Besides scarce paratrachealprocumbent, parenchyma, other species square and upright cells mixed throughout of Heteropterys (Fig. , such as those in the Aptychialh); informal elongate crystals infrequent and in single ray group (cells, Heteropterys subset. Aptychia prismatic , sensu Niedenzu crystals in chambered axial par- enchyma (1928)),cells have abundant non-lignified (Fig. parenchyma in bands lg). (Domingues 2008; Pace 2015) that may proliferate and give rise to additional vascular tissue within the original Discussion stem (Pace 2015). Heteropterys serrata and H. nitida lack non-lignified parenchyma, but have marginal parenchyma Wood anatomy marking the limits of the growth rings. These bands are very narrow in H. nitida and range from narrow to broad in From a wood anatomical point of view, both Heteropterys H. serrata. In addition, although both species have large serrata and H. nitida share a number of features typical of prismatic all crystals in chambers, the crystals are more Malpighiaceae, namely vessels with simple perforation plates, abundant in H. serrata. Crystal abundance may reflect alternate vestured pits, paratracheal parenchyma, gelatinous seasonal and other environmental conditions and may not fibres, heterocellular rays, perforated ray cells and abundance be a good taxonomie character (Franceschi and Nakata of prismatic crystals (Solereder 1908; Metcalfe and Chalk 2005; Marcati and Angyalossy 2005). Nevertheless, par- 1950; León 2009). These two species also share with other enchyma type has great taxonomie value as it is the main species of Heteropterys the presence of narrow vessels in featurelong that differentiates H. serrata from H. nitida. radial chains associated with wide vessels (Domingues 2008), a feature also recorded for many other genera of Malpighi- Taxonomie Treatment aceae (Pace 2015). The presence of narrow vessels associated with wide vessels, named vessel dimorphism by Carlquist Heteropterys serrata Amorim, sp. nov. - HOLOT YPE: (1981), is a typical feature of many lianas (Angyalossy etBrazil, al. Espírito Santo, Mun. Vila Pavão, estrada Vila 2015) due to convergent evolution (Pace and Angyalossy Pavão para Cristalina, margem do Rio Cricaré, 18°42/17//S 2013). However, the arrangement of the small vessels is 40°34/33//W, dis- ca. 101 m a. s. 1., 16 Feb 2014 (fi.), R.C. tinctive because it varies among families, but usually notForzza, H. Medeiros and K. V. Hmeljevski 7826 (holotype: within them. For instance, in Malpighiaceae and Sapindaceae RB! Barcode 00833151; isotypes: CEPEC! NY!) (Figs, la, they appear in radial chains (Klaassen 1999; Pace 2015; c- e, 2, 3, 4). Bastos et al. 2016; Cabanillas et al. in press), but in Bignon- The unique combination of hexagonal stems in cross iaceae they are in clusters or surround large vessels (Pace etsection, al. leaf laminae with a golden-sericeous abaxial

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This content downloaded from 86.59.13.237 on Mon, 21 Jun 2021 11:33:21 UTC All use subject to https://about.jstor.org/terms New Heteropterys and notes on anatomy 181 surface and widely serrate reticulum more margins, visible below than above. stamens Inflorescences a subequal, fil- aments sericeous at base, much-branched and vasicentric panicle, terminal or axillary, aliform, 8-21 cm confluent axial parenchyma in the long, suberect,wood with clearlycorymbs of 4-7 flowers;differentiates inflorescence Het- eropterys serrata Amorim bracts similar fromto the leaves, abruptlyall other reduced to species of the genus. 5-7 x 0.5-2 mm, lanceolate to oblong, the margins serrate Liana, climbing, 5-6(-10) m tall; basal stems 3-5 cm or entire in smaller bracts; peduncle well-developed, diam, cylindrical in cross section, bark irregularly rugose, 3.5-7.3 x 0.5-1 mm, uniformly slender, subterete, young stems flattened longitudinally, hexagonal in cross straight, densely golden-sericeous; bracts section, developing scattered lenticels. Leaves opposite, 1.2-2.2 x 1-1.5 mm, ovate-orbiculate, eglandular, glab- plane; petiole (0.5-)1.3-1.6(-L9) cm long, initially seric- rous adaxially, densely sericeous abaxially; bracteoles eous, later glabrescent, eglandular; stipules not seen; lam- similar to the bracts but smaller; pedicel ina of larger leaves (4.9-)10-15.2(-19) x (2.7-)4.5-7.3(- 2-4 x 0.7-0.9 mm, straight, densely golden-sericeous, 8.7) cm, membranous to subchartaceous, oblong to uniformly slender distally. Sepals 1.3-2.5 x 0.9-1.7 mm, oblong-lanceolate, the base obtuse or slightly cuneate, the acute at apex, appressed against filaments at anthesis, margins widely serrate, teeth usually with a small gland at adaxially glabrous, abaxially densely golden-sericeous; the apex, the apex obtuse or subacuminate; lamina-bearing lateral 4 sepals biglandular, the glands green, 2 large glands abaxially at base, one on each side of 0.8-3 x 0.7-1.8 mm, free at apex, elliptic-ovate. Petals midvein; adaxial surface metallic-sericeous, glabrescent exposed in the enlarging bud, pale yellow, glabrous, claw with age; abaxial surface with densely and persistently and limb thick but not carínate, narrowly flat; lateral 4 golden-sericeous indûment, sessile, straight and appressed petals spreading, margins erose, the claw ca. hairs, the lateral veins more prominent abaxially, the 1.8 x 0.5 mm, the limb 1.7-3.1 x 1.3-2 mm, obtuse at

Fig. 2 Distribution map of Heteropterys mulgurae { white squares ), H. nitida {black triangles ) and H. serrata {white circles). Grey areas in the detail represent the Barreiras Formation along the Brazilian coast

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Fig. 3 General abaxial morphology view: the of stamen Heteropterys second fromserrata right - a floweringopposite the posterior branch; b leaf petal; lamina the shapestamen (abaxial fourth view from ) withleft opposite enlargement the anterior of leaf sepal; f detail margin and base of oneto show stamen, basal adaxial glands; view; c apical g gynoecium, stem node theshowing anterior the style to right, petiole base and its detail apex ofenlarged cross section (a-g fromof stem R.C. showing Forzza the et hexagonalal. 7826). Drawing by shape; d corymb Diana Carneiro with buds and open flowers; e androecium, laid out,

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Fig. 4 General morphology of margin Heteropterys and apex; serrata g overview and habitatof the semideciduous- forest associated a basal stem wood; b apical stemwith nodeinselbergs showing in the the Pedra petiole do base;Elefante Environmental Protection c abaxial surface of the leaf lamina Area, showing where H.the serrata base with grows glands (a-f and from A.M. Amorim et al. 9578). petiole apex; d vegetative branch Photographs in adaxial a,view; d, e eby vegetative André M. branch Amorim; b, c, f by Cleiton Pessoa; in abaxial view; f abaxial surface g by ofRenato the leafGoldenberg lamina showing the

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This content downloaded from 86.59.13.237 on Mon, 21 Jun 2021 11:33:21 UTC All use subject to https://about.jstor.org/terms 184 A. M. Amorim et al. base; posterior Misiones, Argentina, and adjacentpetal areas in the state of suberect, glandular-dentate on the proximal Paraná,1/2-2/3, Brazil (Fig. 2). the claw 1.7-2.2 x 0.4-0.6 mm, the limb 2-3.1 x 1.5-2 mm, truncate at base. Filaments Additional specimens examined (paratypes): Brazil, 1.5-2.2 x 0.4-0.8 mm, abaxially sparsely sericeous atEspírito Santo, Mun. Águia Branca, Assentamento 16 de base, connate ca. 1/3 their length, subequal, distally curved Abril, 18°54'25"S 40°44'05"W, 15 Mar 2006 (senescent sideways or backward; anthers 0.6-1.2 mm long, slightly fi.), V. Demuner et al. 1995 (CEPEC!, MBML!); Mun. reflexed at anthesis, the connective uniformly yellow. Nova Venecia, Área de Proteção Ambiental Pedra do Ovary 0.8-1.6 mm long, densely golden-sericeous; styles Elefante, trilha da Mata da Fazenda Santa Rita, 18°46'57"S 3, 0.9-1.2 mm long, as long as the anthers, glabrous, 40°25/58"W, 16 Jul 2008 (ste), A.M. Amorim et al. 7514 straight or slightly divergent distally, dorsally rounded (CEPEC!);or Mun. Vila Pavão, estrada Vila Pavão para truncate at apex, stigma lateral. Samaras unknown. Cristalina, margem do Rio Cricaré, 18°42'17"S Etymology : The specific epithet refers to the serrate leaf40o34'33"W, 9 Jun 2015 (ste), A.M. Amorim et al. 9578 margins, an unusual feature in Malpighiaceae. (CEPEC!). Phenology: found in flower during February and March. Acknowledgements The authors thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for awarding a Distribution, habitat and conservation status : Heterop- Research Productivity Fellowship to AMA (Grant # 310717/2015-9), terys serrata is endemic to the state of Espírito Santo, Ph.D Fellowship to LCM (Grant # 141561/2015-7) and for providing Brazil (Figs. 2, 4g), and occurs in fragments of semide- financial support for fieldwork (Universal Grant # 486079/2013-9 and ciduous forest associated with inselbergs (i.e. rock out- PPBIO Grant # 457483/2012-1), and Fundação de Amparo à Pesquisa do Estado de São Paulo (F APES P) for awarding a Ph.D. fellowship to crops, which mainly consist of granite or gneiss, rising MRP (Grant # 2012/01099-8) and a BIOTA project (Grant # abruptly above surrounding plains (de Paula et al. 2016)). 2013/10679-0). We also thank Dr. Rafaela C. Forzza (Jardim Botâ- Although this species occurs in a protected area (Pedra do nico do Rio de Janeiro Research Institute), who was the first to point Elefante Environmental Protection Area), according to out that a specimen of Heteropterys serrata was an unusual Malpighiaceae and for kindly providing the loan from RB, Diana IUCN (2012) criteria H . serrata should be classified as Carneiro for the drawings, Nathan Smith for the English revision and endangered (EN [B2 + bi,ii,iii,iv]) due to the following: an the Malpecos Lab staff for assistance with the herbarium collections area of occupancy less than 12 km2 that is in an advanced (http://malpecos.wix.com/malpecoslab). stage of degradation due to human activities (i.e. agricul- Compliance with ethical standards ture, cattle farming and granite quarries); and the original habitat is mostly reduced to small isolated fragments. Conflict of interest The authors declare that the manuscript does not Notes: Heteropterys serrata is easily separated from all present any kind of conflict of interests. The purpose and content of other species of the Heteropterys Metallophyllis informal the work are original and not previously published. group by its young hexagonal stems in cross section (Fig. 3c). This characteristic is common in Lophopterys References species but has never been reported for Heteropterys. Another noteworthy characteristic of the new species is the Amorim AM (2003) The anomalous-stemmed species of Heterop- widely serrate leaf margins (Figs. 3b, 4f), which is an terys subsect. Aptychia (Malpighiaceae). Brittonia 55:127-145. unusual feature in Neotropical Malpighiaceae. doi: 10. 1663/0007- 196X(2003)055[0127:TASSOH]2.0.CO;2 Some aspects of the leaves and inflorescence shape of Amorim AM (2004) A new species of Heteropterys (Malpighiaceae) from the semideciduous forests of Bahia, Brazil. Brittonia Heteropterys serrata resemble H. nitida , a rather common 56:143-146. doi : 10.1 663/0007- 196X(2004)056[0 143 : ANSO and variable species that occurs along the coast from HM]2.0.CO;2 southern Bahia to Parana, and inland as far as eastern Anderson WR (1998) Two new species of Heteropterys (Malpighi- Minas Gerais, Brazil (Anderson 1998). However, in H. aceae) from Southern South America. No von 8:215-217 Anderson WR (2013) Origins of Mexican Malpighiaceae. Acta Bot serrata the glands on the leaf margin are at the apex of the Mex 104:107-156 teeth (vs. mostly 5 mm or more from the margin in H. Anderson WR (2014) Seven new species of neotropical Malpighi- nitida ), the indûment on the abaxial surface of lamina is aceae. Acta Bot Mex 109:23-43 golden-sericeous (vs. metallic- or silvery-sericeous in H. Anderson WR, Davis CV (2007) Generic adjustments in neotropical Malpighiaceae. Contr Univ Michigan Herb 25:137-166 nitida ), the filaments are 1.5-2.2 mm long (vs. 0.3-0.6 mm Angyalossy V, Pace MR, Lima AC (2015) Liana anatomy: a broad long in H. nitida) and the styles have a rounded apex (vs. perspective on structural evolution of the vascular system. In: usually with dorsally obtuse-apiculate apex in H. nitida). Schnitzer SA, Bongers F, Burnham R, Putz FE (eds) Ecology of Another species in this complex that is strongly associated Lianas. Wiley, Oxford, pp 253-287 Barbosa ACF, Pace MR, Witovisk L, Angyalossy V (2010) A new with H. nitida (Anderson 1998) is H. mulgurae W.R.An- method to obtain good anatomical slides of heterogeneous plant derson, which is distributed between the province of parts. IAWA J 31:373-383

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