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PhytoKeys 61: 73-80 (2016)

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Circumscription and synopsis of Eugenia section
Speciosae Bunger & Mazine (Myrtaceae)

Mariana de Oliveira Biinger', Fiorella Fernanda Mazine’,

Eve J. Lucas*, Jodo Renato Stehmann!

| Laboratério de Sistemdtica Vegetal, Departamento de Botanica, Instituto de Ciéncias Biolégicas, Universi-
dade Federal de Minas Gerais, CEP 31270-901, Belo Horizonte, MG, Brazil 2. Departamento de Ciéncias
Ambientais, Universidade Federal de Séo Carlos — campus Sorocaba, Jodo Leme dos Santos, Km 110 - SP-264,
CEP 18052-780, Sorocaba, SP. Brazil 3 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

Corresponding author: Mariana de Oliveira Biinger (maribunger@gmail.com)

Academic editor: P de Lange | Received 25 January 2016 | Accepted 4 February 2016 | Published 25 February 2016

Citation: Biinger MO, Mazine FE, Lucas EJ, Stehmann JR (2016) Circumscription and synopsis of Eugenia section
Speciosae Biinger & Mazine (Myrtaceae). PhytoKeys 61: 73-80. doi: 10.3897/phytokeys.61.7904

Abstract

A new section of Eugenia (Myrtaceae) is described, segregate from Eugenia sect. Phyllocalyx. Phylogenetic
studies suggest that Eugenia sect. Phyllocalyx as traditionally delimited is paraphyletic. To maintain the
monophyly of each of the sections in Eugenia s./., we herein opt to circumscribe a new section and recog-
nize six taxa in sect. Speciosae, which has a distribution mostly in southeastern Brazil and northern South

America. Nomenclatural notes are made and a taxonomic key is provided for the species of the section.

Keywords

Amazon Forest, Atlantic Forest, Neotropics, Myrteae

Introduction

Eugenia Linnaeus is a widespread tropical genus with about 385 species in Brazil (Go-
vaerts et al. 2014, Sobral et al. 2015), most of which grow along the Brazilian Atlantic
rainforest biome (Oliveira-Filho and Fontes 2000). Eugenia is unique among Myrta-
ceae in having a native distribution that spans nearly the entire geographic range of the
family (Snow 2011).

Copyright Mariana de Oliveira Biinger et al. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

74 Mariana de Oliveira Biinger et al. / PhytoKeys 61: 73-80 (2016)

The genus Phyllocalyx was segregated from Eugenia by Otto Berg (1856) being
characterized especially by peduncles with leaf-like bracts and showy sepals, propor-
tionally larger than the flowers. The name Phyllocalyx O. Berg (1856) is also illegiti-
mate, being a later homonym of Phyllocalyx A. Rich. (1847). When Niedenzu, in
1893, transfered Phyllocalyx O. Berg to Eugenia, he named it Eugenia sect. Phyllocalyx.
This name is treated as nomen novum and has the same type as the illegitimate name.
It has priority from 1893 and must be cited as Eugenia sect. Phyllocalyx Nied., not as
Eugenia sect. Phyllocalyx (O. Berg) Nied. (McNeill et al. 2012 — Article 58.1).

Recently, based on a molecular (nuclear and plastid markers) phylogenetic analy-
sis, Mazine et al. (2014) recognized nine clades in Eugenia s. |. They also confirmed
the inclusion of Calycorectes, Hexachlamys, and Phyllocalyx in Eugenia. The “Phylloca-
lyx clade” or “clade 6” sensu Mazine et al. (2014) refers to Eugenia sect. Phyllocalyx
Nied. comprising c. 15 species widely distributed in the Atlantic Forest, from eastern
Brazil to Paraguay. ‘The section is characterized by peduncles with leaf-like bracts and
showy sepals, proportionally larger than the flowers (Berg 1856, under Phyllocalyx),
and is currently being monographed (Biinger et al. unpubl. res.). A remarkable result
of Mazine et al. (2014) is the placement of Eugenia wentii — traditionally included in
Eugenia sect. Phyllocalyx (Mc Vaugh 1969) — in “clade 9” althoug this clade does not
have any support.

After broad sampling of Eugenia sect. Phyllocalyx within a molecular framework
(using five markers, one nuclear and four plastid) (Biinger et al. unpubl. res.), results
show that Eugenia sect. Phyllocalyx sensu Berg emerges as a paraphyletic group. The
clade containing most species previously placed in section Phyllocalyx and also con-
taining the type-species of the section (Eugenia involucrata DC.) emerges as a well-
supported monophyletic group (PP Bayes: 0.99; PP Beast: 0.97; ML: 75). A second,
also well-supported clade (PP Bayes: 1; PP Beast: 1; ML: 100) includes species previ-
ously included in Eugenia sect. Phyllocalyx (Eugenia bunchosiifolia Nied., E. hermesiana
Mattos, E. longipetiolata Mattos, E. macedoi Mattos, E. speciosa Cambess and E£. wentii
Amshoff) but emerges with high support (PP Bayes: 0.99; PP Beast: 0.99; ML: 72) as
sister to clade 9 sensu Mazine et al. (2014). Now, the clade 9 also emerges with high
support (PP Bayes: 0.99; PP Beast: 0.99; ML: 86).

Biinger et al. (unpubl. res.) also have optimised morphological characters across
the molecular tree, presenting useful results with which to distinguish the sections.
Results indicated that these characters are uncommon in Eugenia s./. and can therefore
be used to support placement of species inside a genus/subgenus/section (e.g. Berg
1857, Niedenzu 1893, McVaugh 1969, Mattos 1989). Although these two clades do
not emerge in a monophyletic group, they share the floral characters of showy sepals
and bracteoles that could be homoplastic characters in Eugenia s.1.

To avoid continued recognition of a paraphyletic taxon we herein recognize a new
section called Eugenia sect. Speciosae and provide the new circumscription of Eugenia
sect. Speciosae, an identification key and a synopsis of the known species of this new
section.

Circumscription and synopsis of Eugenia section Speciosae Biinger & Mazine 72

Taxonomic synopsis

The section name “Speciosae” was chosen based on the fact that Eugenia speciosa is the
most geographically widespread species in this group. The specific epithet “speciosa” is
also the oldest within the section (Cambessédes 1832)

Eugenia sect. Speciosae Biinger & Mazine, sect. nov.

urn:lsid:ipni.org:names:77 153396-1

Notes. Trees or shrubs; hairs simple. Indeterminate inflorescence which produces a
floral region that, for instance, produces monads, dyads or triads and vegetative in-
novative shoots, as an auxotelic inflorescence (Briggs and Johnson 1989); bracteoles
linear or narrowly elliptic persistent at anthesis but caducous in mature fruits; flowers
showy always 4—merous; sepals showy, free, foliaceous, sepals and petals concealing
the apex of the bud; ovary 2—locular; ovules 2—many, placenta axile. Fruit crowned
by the calyx lobes. Seeds 1—2; seed coat membranous or cartilaginous; embryo with
fused cotyledons.

Type. Eugenia speciosa Cambess. Fl. Bras. Merid. 2 (19): 351. 1832.

Eugenia sect. Speciosae contains six species with three occurring in the Atlantic
Forest of Brazil, and one distributed in northern South America, in the Amazon. The
Atlantic Forest-Amazon disjunction distribution represents a classic biogeographic

pattern of the Southern Hemisphere (McVaugh 1968).

1. Eugenia bunchosiifolia Nied., Nat. Pflanzenfam. 3, Abt. 7: 82. 1893.

Basionym: Phyllocalyx grandifolius O. Berg, Fl. Bras. 14(1): 333. 1857.
Type: Brazil. Habitat ad urbem Santos in prov. S.Pauli, fructificat Majo: Sellow s.n.
(holotype: B, destroyed; lectotype here designated: K[000170006]!)
Phyllocalyx grandifolius var. pyriformis O. Berg, Fl. Bras. 14(1): 591. 1859.
Type: Brazil. Habitat in silvis prope urbem Rio de Janeiro, e.g. ad Tejuca, florebat
Novembri, fructificabat Septembri: Riedel s.n. (holotype: LE! [photo])
Eugenia santensis Kiaerskou, Enum. Myrt. Bras. 163. 1893, nom. superfl.
Type: Based on Phyllocalyx grandifolius O. Berg
Eugenia littoralis Mattos, Loefgrenia 42:1. 1970, nom. illeg.
Type: Brazil. Sao Paulo: Peruibe, Prainha, 25 Jul 1969, Mattos 15599 (holotype: HB!)
Eugenia brunoi Mattos, Loefgrenia 99:2. 1990, syn. nov.
Type: Based on Eugenia littoralis Mattos

Notes. Eugenia bunchosiifolia is a tree 3—15m alt. from the coastal Atlantic Forest of
Brazil, growing in rainforests from Parana, Rio de Janeiro and Sao Paulo states. ‘This

76 Mariana de Oliveira Biinger et al. / PhytoKeys 61: 73-80 (2016)

species has glabrous leaves with obscure glandular dots visible on both faces, leaf api-
ces are acuminate, without cartilaginous margins, the hypanthium is velutinous. The
lectotype of Eugenia bunchosiifolia was chosen because the holotype was destroyed in
the Second World War. The specimem found at K was a isotype and now considered
the lectotype of this name.

The protologue and the examinated holotype of Eugenia brunoi matches with tho-
se of E. bunchosiifolia, hence this species is here synonymized with E. bunchosiifolia

2. Eugenia hermesiana Mattos, Loefgrenia 94: 1. 1989

Type. Brazil. Sao Paulo: Salesdpolis, na Estagao Bioldgica de Boracéia, 15 Jan. 1968,
Rabello, E. s/n. (holotype: HAS, not found).

Notes. ‘This species has glabrous leaves without cartilaginous margins, dots visible
mostly abaxially, leaf apices are acute or obtuse, the hypanthium is velutinous. Eugenia
hermesiana is a shrub up to 3 m high from Sao Paulo State (Brazil), growing in the
coastal Atlantic Forest. There are few specimens located in BHCB, IAC, NY, SP and
SPSF. It is a threatened species classified as Endangered in the Brazilian Official List of
Flora Threatened Species (MMA 2014).

3. Eugenia longipetiolata Mattos, Dusenia 8: 162. 1968.
Fig. 1D

Basionym: Stenocalyx mutabilis O. Berg, Fl. Bras. 14(1): 347. 1857.
Type: Brazil. Tingua, Schott 5854 (lectotype here designated M [M-0170971]}!;
isolectotype W! [photo])
Eugenia mutabilis Nied., Nat. Pflanzenfam. 3, Abt. 7: 81. 1893, nom. illeg.
Type: Based on Stenocalyx mutabilis O. Berg
Eugenia tinguana Mattos, Loefgrenia 123: 1. 2006, nom. superfl.
Type: Based on Stenocalyx mutabilis O. Berg

Notes. Eugenia longipetiolata is a tree up to 15 m high from coastal Atlantic Forest
of Brazil, growing in ombrophilous forests from Rio de Janeiro and Sao Paulo states.
This species has leaves with visible, flat gland dots on both faces, black-floccose simple
trichomes on abaxial faces, caudate apices, non-cartilaginous margins and a ferrugi-
nous-pubescent hypanthium.

The lectotype was chosen for Stenocalyx mutabilis because Berg did not indicate
a single specimen and Mattos did not designate a ectotype when he published the
nom. nov.. The specimen from M was seen and here considered the lectotype for
the name.

Circumscription and synopsis of Eugenia section Speciosae Biinger & Mazine i

Figure |. A, B Auxotelic inflorescence and foliaceous calyx, Eugenia wentii (photo by B. Holst)

C, E, F Eugenia speciosa, mature fruit with decidous bracteoles, stem bark and position and format of the

bracteoles (photos by M.O.Biinger); D Eugenia longipetiolata (photo by M.O.Biinger).

4. Eugenia macedoi Mattos & D. Legrand, Loefgrenia 67: 24. 1975.

Type. Brasil, Minas Gerais: Ituiutaba, San Vicente, 12 Sep. 1950, Macedo, A. 2574
(holotype: MVM, not seen; isotype US! [00603977])

Notes. Eugenia macedoi is known only by two specimens colected in Minas Gerais
and Goias States (Brazil). This species is a shrub growing in the Cerrado biome (like
savannas). Apparently it is the only species of the section that occurs in dry areas. ‘This
species has glabrescent leaves without cartilaginous margins, dots visible on both faces,
leaf apices are acute, and the hypanthium is velutinous.

5. Eugenia speciosa Cambess., Fl. Bras. Merid. (quarto ed.) 2(19): 351. 1832.
Fics, Bor

Phyllocalyx speciosus (Cambess.) O. Berg, Fl. Bras. 27(2—3): 307. 1856.
Type: Based on Eugenia speciosa Cambess.

78 Mariana de Oliveira Biinger et al. / PhytoKeys 61: 73-80 (2016)

Phyllocalyx retusus O. Berg, Fl. Bras. 14(1): 331. 1857.
Type: Brazil. Habitat ad ripas flaminis Rio Pardo in Montevideo: Sellow s.n. (holo-
type B, probably destroyed; isotypes: K! [000276590], BR! [(0000005261277]).
Phyllocalyx limbatus O. Berg, Fl. Bras. 14(1): 332. 1857.
Type: Brazil. Habitat ad Angra dos Reys in prov. Rio de Janeiro: Pohl 264, 5760., loco
incerto ajusdem prov., Sellow s.n. (lectotype here designated BR! [526061-Sellow]};
isolectotype: B (fl.), probably destroyed; W (fr.) [photo]! ).
Phyllocalyx macrosepalus O. Berg, Fl. Bras. 14(1): 332. 1857.
Type: Brazil. Habitat ad Alegres et Manoel Jesu praedia in prov. Minarum: Mikan
s.n., Pohl s.n. (lectotype here designated: BR! [526984]; isolectotypes: M! [M-
0171010], W [photo]}).
Phyllocalyx marginatus O. Berg, Fl. Bras. 14(1): 332. 1857.
Type: Brazil. Habitat in prov. Rio de Janeiro: Martius s.n. (holotype: BR! [526094].
Eugenia retusa (O.Berg) Nied., Nat. Pflanzenfam. 7: 82. 1893.
Type: Based on Phyllocalyx retusus O. Berg
Eugenia caldensis Kiaerskou, Enum. Myrt. Bras. 162. 1893.
Type: Based on Phyllocalyx marginatus O. Berg
Eugenia macrocalyx Mart. ex B.D Jacks, Index Kew. 1: 908. 1893.
Type: Based on Phyllocalyx macrosepalus Berg

Type. Brazil. In sabulosis prope praedium vulgo Fazenda d ‘Araucaria in prov. S. Pauli, flo-
ret Octobri: Saint-Hilaire s.n. (lectotype: P [P01902768]}!; isolectotype: MPU! [photo])

Notes. Eugenia speciosa is a tree 5—12 m high from Atlantic Forest in southern and
southeastern Brazil. It is common in rainforests and “restingas”. This species also occurs
in montane Atlantic Forest in Minas Gerais State (Brazil) and also occurs in Paraguay,
Argentina, Uruguay and Bolivia. This species has leaves with visible, salient dots on both
faces, glabrous, obtuse apices with cartilaginous margins and a glabrous hypanthium.

The lectotypes chosen for Phyllocalyx limbatus and Phyllocalyx macrosepalus are
from BR; they were seen and we consider that the specimens that well represent the
names. The lectotype that was chosen for Phyllocalyx limbatus is a specimen that is a
duplicate (isotype) of the specimen that was in B which was destroyed in the Second
World War. For thus, we consider it as the lectotype for this species.

6. Eugenia wentii Amshoff, Recueil Trav. Bot. Néerl. 39: 160, f. 4. 1942.
Fig. 1A, B

Phyllocalyx wentii Amshoff, Recueil Trav. Bot. Néerl. 39: 158, f. 4. 1942
Type: Based on Eugenia wentii Amshoff nomen alternativ.

Calycorectes macrocalyx Rusby, Mem. New York Bot. Gard. 7: 313. 1927.
Type: Bolivia. Bopi River Valley. Rusby 666 (holotype: NY! [00386736]; isotypes:
BKL! [photo], MICH! [photo], US! [photo])

Eugenia macrocalyx (Rusby) McVaugh, Fieldiana, Bot. 29(3): 212. 1956, nom. illeg.
Type: Based on Calycorectes macrocalyx Rusby

Circumscription and synopsis of Eugenia section Speciosae Biinger & Mazine Ji)

Type. Suriname. Fluv. Coppename inf., Went FAFC 142 (holotype: U! [0005034])
Notes. Eugenia wentii is a treelet or tree 3-6 m high from the Amazon forest; it is found
in Amaz6nia and Paréa States (Brazil), French Guyana, Suriname, Venezuela, Bolivia, Co-
lombia, Ecuador and Peru. ‘This species has glabrous leaves with flat, visible gland dots on
both faces, caudate apices without cartilaginous margins and a velutinous hypanthium.

Key to species of Eugenia sect. Speciosae

1 Fiypanthiwin: clabrows:s acdsntictcnce essa cetetedacbsssqeacttecdeeataccah tains E. speciosa
— Hypanthium with trichomes
2 Leaves with caudate apices and black-floccose indument on mature leaves...
Dc anee ee tema tee eres team once Senda nemeestaeeees cement eeean E. laneiperiolees
= eae with acuminate apices, acute, obtuse or rostrate; glabrous or without
black-floccose hairs

3 Leaves usually with cartilaginous margins «0.0... eee E. bunchosiifolia
- Leaves always without cartilaginous margins

4 Leaves withaciiminate or rostrate apices’. vive veer eeka weenie E. wentii
- Leaves with acute or obtuse apices

5 Calyx lobes acuminate 50 to 70 mm long. ...... eee eee E. hermesiana
- Calyx lobesyacute- B.0 tO-d MIM ONG. a tssceihnnitnnna dled shaaie de ricetat E. macedoi
Acknowledgements

We thank all curators of the visited herbaria for allow us to study the specimens and
to Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) and Con-
selho Nacional de Desenvolvimento Cientifico e Tecnolégico (CNPq) for the financial
support received by the first and the last authors.

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