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Systematics and Biodiversity
ISSN: 1477-2000 (Print) 1478-0933 (Online) Journal homepage: https://www.tandfonline.com/loi/tsab20
Lapaea (Plantaginaceae, Gratioleae), a new genus
endemic to the Espinhaço Range (Brazil) with a
remarkable red-flowered new species
André Vito Scatigna, Vinicius Castro Souza, Raquel Moura Machado & André
To cite this article: André Vito Scatigna, Vinicius Castro Souza, Raquel Moura Machado &
André Olmos Simões (2020): Lapaea (Plantaginaceae, Gratioleae), a new genus endemic to
the Espinhaço Range (Brazil) with a remarkable red-flowered new species, Systematics and
Biodiversity, DOI: 10.1080/14772000.2020.1771470
To link to this article: https://doi.org/10.1080/14772000.2020.1771470
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Published online: 24 Jun 2020.
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Lapaea (Plantaginaceae, Gratioleae), a new genus endemic to the
Espinhac¸o Range (Brazil) with a remarkable red-flowered
ANDRÉ VITO SCATIGNA
, VINICIUS CASTRO SOUZA
, RAQUEL MOURA MACHADO
ANDRÉ OLMOS SIMÕES
Departamento de Ci^
ogicas, Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de S~
ao Paulo, Avenida
adua Dias, 11, Piracicaba, 13418-900, S~
ao Paulo, Brazil
Programa de P
ao em Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas, Rua Monteiro Lobato,
255, Campinas, 13083-970, S~
ao Paulo, Brazil
(Received 24 February 2020; accepted 5 May 2020)
Stemodia sensu lato is one of the largest genera of the Gratioleae (Plantaginaceae). Recent phylogenetic studies
corroborated the paraphyly of Stemodia in its traditional circumscription and suggested the segregation of S. stellata
from this genus. This species belongs to a morphologically cohesive group along with S. cipoensis,S. harleyi and S.
lobata, all endemic to the Espinhac¸o Range, eastern Brazil. In this study, we tested the monophyly and interspecific
relationships of this group by reconstructing its phylogeny, based on sequences of three cpDNA regions (rps16 and trnL
introns and the trnL–trnF intergenic spacer), and one nrDNA region (ITS1 spacer), using Maximum parsimony and
Bayesian inference. The monophyly of the 'S. stellata group' was confirmed, and its segregation from Stemodia
corroborated. We propose Lapaea as a new genus to accommodate its species, with subsequent four new combinations:
L. cipoensis,L. harleyi,L. lobata, and L. stellata. Additionally, we describe and illustrate L. rubriflora, a remarkable
red-flowered new species, based on morphological and micromorphological evidence. Lastly, a taxonomic revision of
the species recognized in Lapaea is presented, providing an identification key, photos, descriptions, distribution map and
notes on their morphology, geographic distribution, habitat, ecology, and conservation status. Lapaea is restricted to the
Espinhac¸o Range comprising five threatened species, each of them with a narrow distribution and phylogenetic
relationships consistent with biogeographic provinces.
Key words: Campo rupestre, ITS, Lamiales, lapa, micromorphology, phylogenetics, rps16,Stemodia, taxonomy,
Stemodia L. is one of the largest genera of the tribe
Gratioleae of the Plantaginaceae, comprising 50–60 spe-
cies distributed throughout the tropics, especially in the
New World and Australia (Souza & Giulietti, 2009;
Turner & Cowan, 1993a). It is generally characterized by
the calyx with equal to subequal sepals, and four fertile
stamens with two thecae in each anther separated by a 2-
brachiate connective, often described as stalked/stipitate
thecae; otherwise being morphologically heterogeneous
(Minod, 1918; Souza & Giulietti, 2009; Turner & Cowan,
1993a). Phylogenetic studies based on plastid and nuclear
ribosomal DNA sequences suggested that Stemodia,inits
current circumscription, is not monophyletic (Estes &
Small, 2008; Fritsch et al., 2007; Scatigna et al., 2018). In
the most recent phylogenetic study of Gratioleae
(Scatigna et al., 2018), the five species of Stemodia
sampled were retrieved in three distinct clades: in the
first, Stemodia foliosa Benth. formed a grade with S. ver-
ticillata (Mill.) Hassl., related to Scoparia L.; in the
second, Stemodia microphylla J.A.Schmidt formed a
grade with Stemodia maritima L., related to Achetaria
Cham. & Schltdl.; and in the third, Stemodia stellata
B.L.Turner was recovered as sister group of Philcoxia
P.Taylor & V.C.Souza and related to Tetraulacium Turcz.
Stemodia maritima, type species of its genus, was
assigned by Souza and Giulietti (2009)toa
Correspondence to: Andr
e Vito Scatigna. E-mail:
ISSN 1477-2000 print / 1478-0933 online
#The Trustees of the Natural History Museum, London 2020. All Rights Reserved.
Systematics and Biodiversity (2020), 0(0):1–18
Published online 24 Jun 2020
morphological assemblage of species characterized by
the erect habit; the sessile, frequently clasping, leaves;
and the sessile to sub-sessile, bracteolate flowers, fre-
quently concentrated at the apex of the stems. This
group of species, which includes S. durantifolia (L.)
Sw., S. hyptoides Cham. & Schltdl., S. palustris
A.St.Hil., S. perfoliata Scatigna & V.C.Souza and S.
stricta Cham. & Schltdl. (Scatigna et al., 2018), should
be referred to as Stemodia sensu stricto; a phylogenetic
study of this group is being prepared by our research
group. Meanwhile, S. stellata has been assigned by
Scatigna et al. (2017) to a second cohesive group of
species along with S. cipoensis Scatigna, S. harleyi
B.L.Turner and S. lobata J.A.Schmidt, characterized by
the procumbent, suffruticose, and generally rupicolous
habit, with usually 3-whorled and clearly petiolate
leaves, and axillary, long-pedicelate and ebracteolate
flowers. In the course of systematic studies in Stemodia
s.l., we came across some specimens that are consistent
with this assemblage of species, herein addressed as the
‘Stemodia stellata group’, but with a remarkable
divergent trait, that is, flowers with red corolla; these
specimens clearly represent a new species.
The ‘Stemodia stellata group’is restricted to the
quartzite outcrops of the Espinhac¸o Range (ER), which is
a vast mountain chain with a range of 1200 km, in a
north-south orientation, in eastern Brazil, crossing the
states of Bahia and Minas Gerais (Scatigna et al., 2017;
Silveira et al., 2016). The ER shelters the core area of
the campo rupestre, an azonal vegetation mosaic charac-
terized by a grass-shrubby physiognomy, associated with
rocky formations of quartzite, sandstone, and ironstone,
usually interspersed with white sandy patches, occurring
above 900 m asl. (Alves & Kolbek, 2010; Alves et al.,
2014; Silveira et al., 2016). Campo rupestre is also
known for its remarkable levels of plant endemism
(Colli-Silva et al., 2019;Conceic¸~
ao & Pirani, 2005).
Given the phylogenetic placement of Stemodia stellata
(Scatigna et al., 2018) and its morphological affinities
(Scatigna et al., 2017; Souza & Giulietti, 2009), in this
study we aim to: (1) test the monophyly of the 'Stemodia
stellata group' and its interspecific relationships, and
evaluate the need to recognize it as a new genus; (2) pre-
sent a taxonomic revision of the ‘S. stellata group’,
including the description of a new species, an identifica-
tion key, photos, and notes on morphology, geographic
distribution, habitat, ecology, and conservation status.
Materials and methods
The cpDNA dataset (rps16 and trnL introns and the
trnL-trnF intergenic spacer) as well as the nrDNA ITS1
intergenic spacer, were retrieved from Scatigna et al.
(2018). This was based on 50 samples, representing: 28
species and eight genera of Gratioleae; six species and
two genera of Angelonieae, sister group of Gratioleae;
and Plantago media L. (Plantagineae) as rooting taxon.
All these sequences are available at GenBank (https://
www.ncbi.nlm.nih.gov/genbank/). The sampling of
Philcoxia was reduced to one accession for each species
of this genus, and two or more accessions of each of the
five species of the 'Stemodia stellata group' were
included. Methodology for the generation of new
sequences followed Scatigna et al. (2018). Maximum
parsimony (MP) and Bayesian inference (BI) analyses
were performed for each region, separately, and then
applied to a combined dataset with the three plastid
regions (Partially Combined (PC)). Parameters for
phylogenetic analyses followed Scatigna et al. (2018).
Threshold for support values, above which results were
considered statistically meaningful, were: Bootstrap
(BS) 75% in the MP analysis, and Posterior
Probability (PP) 0.95 in the Bayesian analysis.
The descriptions and morphological comparisons were
based primarily on field observations and examination
of herbarium specimens, complemented with data from
the literature (Scatigna et al., 2017; Schmidt, 1862;
Souza & Giulietti, 2009; Turner & Cowan, 1993b).
Morphological terminology follows J. Harris and Harris
(2001). All collections of Gratioleae housed at the fol-
lowing herbaria were analysed: ALCB, BHCB, BHZB,
CEN, CEPEC, CVRD, DIAM, EAC, ESA, HCJS, HRB,
HRCB, HUEFS, IAC, IAN, ICN, MAR, MBM, MBML,
MG, OUPR, R, RB, SLUI, SP, SPF, UB, UEC, UFG,
and UPCB, in addition to the digital images from HBG,
G, K, LE, MO, NY, and P; herbarium acronyms follow
Thiers (2020 (continuously updated)). All cited speci-
mens were consulted in person, except for those indi-
cated by '[digital image]'. Conservation status
assessments followed the IUCN Red List categories and
criteria (International Union for Conservation of Nature
(IUCN), 2012) and subsequent guidelines (IUCN
Standards & Petitions Committee, 2019). We estimated
the extent of occurrence (EOO) and area of occupancy
(AOO) with the Geospatial Conservation Assessment
Tool (GeoCAT; Bachman et al., 2011) using a cell
width of 2 km.
Leaf and stem samples of the new species were fixed in
formol-acetic acid alcohol (FAA) 50% for 24 hours and
then stored in 70% ethanol. The material was critical
2 A. V. Scatigna et al.
point dried, mounted on aluminium stubs with double-
sided adhesive tape and covered with 50 nm of gold in a
Bal-Tec SCD 050 sputter coater. Images were observed
in a Jeol JSM 5800LV Scanning Electron Microscope,
at 10 kV, and captured with SemAfore 5.21 software.
Micromorphological terminology follows Scatigna
et al. (2017).
Results and discussion
In addition to the sequences used by Scatigna et al.
(2018), our study included 45 newly generated sequen-
ces of 12 accessions of four additional species of the
'Stemodia stellata group', henceforth called Lapaea, with
the new combinations having been already applied
throughout the text and figures. List of voucher speci-
mens, GenBank accession numbers, as well as all inde-
pendent and merged datasets are available as
Supplementary material.Table 1 summarizes the parsi-
mony scores and nucleotide substitution models for each
data matrix. There were no visual incongruences
between trees generated from each of the three cpDNA
regions using either MP or BI approaches (tree files
generated by MP and BI from each cpDNA region, sep-
arately, are available as Supplementary material); hence,
the majority-rule consensus tree generated by BI from
the PC dataset is the only one presented and discussed
in detail (Fig. 1). Although both MP and BI analyses of
ITS1 provided overall poorly resolved trees (see
Supplemental material), they showed a strong incongru-
ence relative to the PC dataset within one clade
(Subclade L, in Fig. 1). Therefore, the relationships
within this clade, generated either from PC and ITS1
datasets, are presented separately in Fig. 2.
The PC dataset consisted of 50 accessions and 2301
aligned characters, of which 479 were parsimony-
informative, 1531 were constant, and 291 were variable
but uninformative. The nucleotide substitution model
selected by the Bayesian Information Criterion (BIC)
was TVM þG for rps16, TIM1 þG for trnL, and
TPM3uf þG for trnL–trnF. We recovered the
Gratioleae in a well-supported clade (PP ¼1; BS ¼98)
formed by Mercadonia procumbens and its sister clade
(PP ¼1; BS <50) composed of the same three main
clades (clades A, B, and C in Fig. 1), reinstating the
clade composition and placement already obtained in
Scatigna et al. (2018). All species of Lapaea formed a
clade (subclade L; PP ¼1; BS ¼99) sister to Philcoxia
(subclade P; PP ¼1; BS ¼100); these two subclades
form a clade (PP ¼1; BS ¼98) sister to Tetraulacium
veroniciforme, composing clade C (PP ¼0.94; BS ¼
54). Within subclade L, L. harleyi and L. rubriflora
form a subclade (subclade BA, PP ¼1; BS ¼99) sister
to another subclade (subclade MG, PP ¼1; BS ¼95)
formed by L. lobata,L. cipoensis, and L. stellata.
Within subclade BA, all accessions of L. harleyi formed
a strongly supported clade (PP ¼1; BS ¼100) sister to
the clade formed by the accessions of L. rubriflora (PP
¼1; BS ¼75). Within subclade MG, L. cipoensis and
L. stellata form a clade (PP ¼1; BS ¼93) sister to L.
lobata; the accessions of L. stellata formed a poorly
supported clade (PP ¼0.55) and the accessions of L.
cipoensis were recovered in a polytomy.
The ITS1 dataset consisted of 49 accessions and 465
aligned characters, of which 176 were parsimony-
informative, 235 were constant, and 54 were variable
but uninformative. The nucleotide substitution model
selected by the BIC was GTR þG. Trees from this data-
set are only available as Supplementary material, except
subclade L, which is described below and presented in
Fig. 2.2. Subclade L (Fig. 2.2) was composed of all spe-
cies of Lapaea forming the same subclades BA (PP ¼
0.99; BS ¼92) and MG (PP ¼1; BS ¼100) as in the
PC dataset. Subclade BA was composed of all represen-
tatives of L. harleyi forming a clade (PP ¼1; BS ¼99)
sister to the clade formed by all L. rubriflora accessions
(PP ¼1; BS ¼97). Subclade MG was formed by a
subclade composed of all accessions of Lapaea cipoen-
sis (PP ¼0.92; BS ¼81), sister to a subclade formed
by L. lobata and L. stellata (PP ¼1; BS ¼95); the
Table 1. Parsimony scores and evolutionary model for each dataset. PC ¼Partial Combined.
Dataset rps16 trnL trnL–FPC ITS1
Terminals 47 49 50 50 49
Missing taxa 6% 2% 0% —2%
Number of characters 1085 665 551 2301 465
Parsimony informative characters 254 96 129 479 176
% informative 23.4 14.4 23.4 20.8 37.8
Tree length 561 190 268 1043 740
Consistency index (CI) 0.66 0.72 0.66 0.65 0.47
Retention index (RI) 0.88 0.91 0.89 0.88 0.75
Evolution model (BIC) TVM þG TIM1 þG TPM3uf þG—GTR þG
Lapaea, a New Genus from Brazil 3
accessions of L. stellata formed a clade (PP ¼1; BS ¼
94), but the accessions of L. lobata were recovered in
The analysis of the cpDNA combined dataset (PC)
retrieved, in a broad sense, a clade composition, and
relationships among and within the clades, consistent
with those obtained by Scatigna et al. (2018), even
though we have not included the rpl16 dataset in
All species of Lapaea formed a well-supported mono-
phyletic group, sister to Philcoxia and distantly related
to Stemodia maritima. Our results corroborate the
Fig. 1. Majority-rule consensus tree generated by Bayesian inference from the Partial Combined (PC) dataset including sequences of
the rps16 and trnL introns and the trnL–trnF intergenic spacer of the cpDNA. Number next to nodes are Posterior Probability/
Bootstrap values. Clades represented by letters are discussed in the text. Dark arrow points Stemodia maritima, type species of
4 A. V. Scatigna et al.
segregation of Lapaea from Stemodia s.s. and disagree
with Souza and Giulietti (2009), in what concerns the
relationship of S. microphylla with L. stellata.Stemodia
microphylla is probably closer to S. veronicoides
J.A.Schmidt with which it shares the resupinate, bibrac-
teolate flowers (Souza & Giulietti, 2009). Further phylo-
genetic studies associated with thorough morphological
analyses may support the segregation of even more spe-
cies from Stemodia s.s.
The Espinhac¸o Range (ER) has two major centres of
endemism, the Chapada Diamantina, in Bahia, and the
Southern Espinhac¸o, in Minas Gerais, which in turn are
subdivided in several smaller centres of endemism
(Bitencourt & Rapini, 2013; Echternacht et al., 2011;
Ribeiro et al., 2014). Recently, Colli-Silva et al. (2019)
formalized the descriptions of Chapada Damantina (CD)
and Southern Espinhac¸o (SE) as biogeographic provin-
ces. Within SE, Colli-Silva et al. (2019) recognized
three districts, namely Gr~
ao-Mogol, Diamantina Plateau
and Iron Quadrangle. The Lapaea clade consists of two
subclades, the first comprising the two species restricted
to the Chapada Diamantina, namely L. harleyi and L.
rubriflora, and the second including the three species
restricted to the Southern Espinhac¸o, L. cipoensis,L.
lobata, and L. stellata. This phylogenetic pattern, of a
clade composed of species endemic to the CD as sister
to another formed by species from the SE, is consistent
with that found in other genera such as Philcoxia
(Scatigna et al., 2018), Paepalanthus Mart.
o et al., 2013) and Minaria
T.U.P.Konno (Apocynaceae; Ribeiro et al., 2014) and
supports the recognition of the two biogeographic prov-
inces described by Colli-Silva et al. (2019).
The relationships between the species from SE are still
controversial. The cpDNA markers suggest that L. stel-
lata and L. cipoensis, both from the Diamantina Plateau
province (sensu Colli-Silva et al., 2019), are more closely
related to each other than to L. lobata, from the Iron
Quadrangle province (sensu Colli-Silva et al., 2019), a
relationship that is geographically consistent with the
areas of endemism retrieved by Echternacht et al. (2011).
However, according to the ITS1 dataset, L. cipoensis is
sister to a clade formed by L. lobata and L. stellata.
Incongruence between cpDNA and nrDNA was observed
in tribe Veroniceae (Plantaginaceae) and attributed to bio-
logical causes, such as hybridization and polyploidization
(Albach & Chase, 2004). We would encourage studies at
population level and on reproductive biology in this
group to elucidate this issue.
Although in campo rupestre there is a high rate of
endemism at species level, only a few genera are exclu-
sive to these areas (Alves & Kolbek, 2010). This is the
case of Lapaea, which is restricted to the campo
rupestre of the ER, and where each of its five species
has a narrow geographic distribution consistent with cur-
rently recognized biogeographic units (Colli-Silva et al.,
2019; Echternacht et al., 2011; Ribeiro et al., 2014).
This genus is sister to Philcoxia, which, in its turn, com-
prises a clade with species also restricted to the campo
rupestre of the ER. The geographically structured diver-
sity found in the ER has been documented in unrelated
lineages (Echternacht et al., 2011; Silveira et al., 2016),
and was also observed in Lapaea and Philcoxia. This
identifies these two genera as potential model-groups
for reconstructing biogeographic scenarios, in a global
hotspot with the highest rates of species/km
, within a
Fig. 2. Lapaea clade extracted from the Majority-rule consensus tree generated by Bayesian inference from the Partial Combined
(PC) dataset (2.1) and from ITS1 dataset (2.2). Numbers next to nodes are Posterior probability/Bootstrap values.
Lapaea, a New Genus from Brazil 5
megadiverse country (Alves & Kolbek, 2010; Alves
et al., 2014; Ribeiro et al., 2014).
Lapaea Scatigna & V.C.Souza gen. nov.
Type Species.Lapaea stellata (B.L.Turner) Scatigna
Etymology.Lapaea alludes to lapa, a Portuguese word for
natural shelters provided by fractures on rocks and bould-
ers, which are common in the campo rupestre of the ER.
Diagnosis.Lapaea differs from all other genera in
Gratioleae for the following unique set of traits: terres-
trial to rupicolous life form; crawling to procumbent
habit; clearly petiolate, usually 3-whorled leaves; long-
pedicellate, axillary, ebracteolate and non-resupinate
flowers; calyx with sepals connate at the base to a third
of their length; bilabiate corolla; four fertile stamens
with free anthers, each one composed of two glabrous
thecae separated by a 2-brachiate connective; obdeltoid
and slightly curved stigma; and capsule dehiscence both
loculicidal and septicidal on apical portion, function-
Subshrubs or rarely herbs, terrestrial to rupicolous, per-
ennial, up to 1 m high, with variable indument on vege-
tative parts, usually aromatic. Stems crawling to
ascending, terete to sub-quadrangular, branched. Leaves,
3-whorled, rarely opposite in younger plants, clearly
petiolate. Flowers axillary, non-resupinate, single or
geminate, ebracteolate, long-pedicelate, pentamerous,
bisexual; pedicels clearly ascending after maturation of
fruits; sepals usually equal, clearly connate at the basal
third; corolla slightly to strongly bilabiate, whitish, lilac,
purple, dark blue, deep violet or red, upper lip 2-lobed,
lower lip 3-lobed. Stamens 4, didynamous, included;
anthers with 2 thecae, separated by a 2-brachiate con-
nective, glabrous; staminode 1, much reduced. Ovary
superior, syncarpous, 2-carpelar, 2-locular, ovoid, glab-
rous; placentation axillary, ovules numerous; style ter-
minal, solitary, apex obdeltoid, curved, glabrous.
Capsule ovoid, dehiscence both loculicidal and septici-
dal on apical portion, functionally poricidal, glabrous,
apex zygomorphic; seeds cylindrical to ovoid, reticulate,
longitudinally ribbed, base acuminate.
Distribution and habitat
Lapaea is restricted to the Espinhac¸o Range in Minas
Gerais and Bahia, Brazil (Fig. 3). All known species of
this genus occur in areas of campo rupestre, at eleva-
tions between 900 and 2000 m asl, growing on edges of
relatively humid and shaded fractures of quartzite out-
crops, regionally called lapas (Fig. 4.6).
Lapaea differs from Stemodia sensu stricto for the habit
(crawling to procumbent subshrubs vs. erect herbs); by
the petiolate leaves (vs. sessile); long-pedicellate, axil-
lary flowers (vs. usually sessile to sub-sessile, frequently
in spiciform inflorescences); bracteoles absent (vs. pre-
sent); and sepals connate at base, to a third of their
length (vs. free). It also differs from its sister genus,
Philcoxia, for the presence of four, bithecous stamens
(vs. only two, monothecous stamens); absence of peltate
leaves (vs. presence); aerial stems and petioles (vs.
underground); and non-resupinate flowers
Souza and Giulietti (2009) recognized a morpho-
logical assemblage of species that included the herein
defined Lapaea plus Stemodia microphylla and S. vero-
nicoides. These last two also present crawling habit,
petiolate leaves and long-pedicellate flowers. Lapaea
differs from these two species for the absence of brac-
teoles on pedicels (vs. presence) and in the non-resupin-
ate flowers (vs. resupinate). Molecular evidence
corroborates the segregation of S. microphylla from
Lapaea (Scatigna et al., 2018).
Lapaea species may be confused with Stemodia trifo-
liata (Link) Rchb., differing in the crawling to procum-
bent habit in Lapaea (vs. erect in S. trifoliata), the
curved, obdeltoid stigma (vs. erect, strongly bilobed), the
infundibular and bilabiate corolla (vs. salverform and cru-
ciform) and the simple-serrate to crenate leaf margins
(vs. usually double-serrate) (Scatigna et al., 2017).
Identification key for the species of Lapaea
1. Dendritic trichomes present ........................ L. stellata
10. Dendritic trichomes absent
2. Stems and pedicels densely glandular-puberulent,
covered exclusively with short, capitate trichomes;
corolla throat glabrous ..................................... L. lobata
20. Stems and pedicels hirsute to villous, covered with
long, non-capitate or minutely capitate trichomes, in
addition to short, capitate trichomes; corolla throat
pubescent or villous
3. Leaves strigose, mainly covered with long, stiff,
strongly curved, non-capitate trichomes, in addition
to short, capitate trichomes; corolla throat
pubescent ........................................... L. cipoensis
6 A. V. Scatigna et al.
30. Leaves villous, mainly covered with long, flexible,
minutely capitate trichomes, in addition to short,
capitate trichomes; corolla throat villous
4. Corolla blue to purplish, with white and yellow
nectar guide, tube 5.5–8 mm long; leaves ovate to ovate-
elliptical, margin flat, petioles 0.2–2.3 cm; sepals
5.5–9.3 mm long long; ..................................... L. harleyi
40. Corolla red, without nectar guide, tube 15–18.2 mm
long; leaves elliptic, margin revolute, petioles 0.2–0.5 cm
long; sepals 7.5–10.1 mm long; ..................... L. rubriflora
Lapaea cipoensis (Scatigna) Scatigna, comb. nov.
Stemodia cipoensis Scatigna, in Scatigna et al. Syst.
Bot. 42: 373. 2017. TYPE: BRAZIL. Minas Gerais,
Santana do Riacho, RPPN Brumas do Espinhac¸o, base
do morro Ermo Ac¸u, 434205000W, 190200900S, 1430 m,
30 Apr 2016, A.V. Scatigna & C.A. Ferreira Junior
1120 (holotype: UEC; isotypes: K, NY, SPF). Fig. 4.1.
Lapaea cipoensis is characterized by a hispid texture in
its vegetative parts, which results from the combination
of stiff, strongly curved, non-capitate trichomes with
short, capitate trichomes. It is similar to L. lobata and L.
stellata, though differing in the overall, more robust,
habit. Furthermore, it differs from L. lobata in the hispid
indument of stems, petioles and pedicels (vs. glandular-
puberulent); deep blue to purple corolla with a white
patch in the pubescent throat (vs. entirely violet, with
glabrous throat); and the dorsiventrally compressed cor-
olla tube (vs. not compressed). It differs from L. stellata
in the lack of dendritic trichomes (vs. present), and the
upper corolla lip being patent to ascending (vs. reflexed).
Subshrubs, up to 50 cm tall. Stems ascending, covered
with short capitate trichomes, interspersed with long,
stiff, non-capitate trichomes, denser towards apex.
Petioles 0.2–1.3 cm long, densely covered with short,
capitate trichomes and long, non-capitate trichomes;
blades broadly ovate to ovate-lanceolate,
0.8–6.3 0.3–3 cm, chartaceous, abaxial surface sparsely
covered with short capitate trichomes and some longer,
curved, non-capitate trichomes, these exclusively on
veins, adaxial surface strigose, covered with short
Fig. 3. Distribution map of Lapaea spp. in eastern Brazil. Diamond ¼L. cipoensis;Star¼L. harleyi; Circle ¼L. lobata; Cross ¼
L. rubriflora; Triangle ¼L. stellata.
Lapaea, a New Genus from Brazil 7
capitate trichomes interspersed with longer, non-capitate,
curved trichomes scattered over the surface, apex acute,
base cuneate to obtuse, margins crenate-serrate, plane.
Pedicels 0.8–3 cm long, densely covered with short capi-
tate trichomes, sparsely covered with long, non-capitate
trichomes; sepals lanceolate, 5–61.3–2.2 mm, sparsely
covered with short, capitate trichomes, with few longer,
curved, non-capitate trichomes on veins and margins;
corolla strongly bilabiate, dark blue to purple, with
broad white patch on throat; tube 10–12 mm long, dorsi-
ventrally compressed towards throat, base slightly
inflated, externally glabrous, internally villous at base;
upper lip patent to ascending, obscurely 2-lobed,
5.1–5.3 6–6.3 mm, apex truncate to slightly emargi-
nated, externally bearded; lower lip shallowly 3-lobed,
2-canaliculate, 5.5–67.8–8.5 mm, pubescent, with
short, non-capitate trichomes on throat, lobes with trun-
cate to emarginate apex, 4.5 3.5 mm. Anterior pair of
stamens 6.2–6.3 mm long, posterior 4.2–4.3 mm long;
thecae 0.7 mm long. Ovary 2.1–2.2 1.3–1.4 mm;
style 5.4–5.5 mm long. Apex 0.7 mm long.
Capsule 7–7.2 3.4–3.5 mm; seeds cylindrical,
0.7–0.8 0.4–0.5 mm.
BRAZIL. Minas Gerais, Santana do Riacho, RPPN
Brumas do Espinhac¸o, sop
e do Ermo Ac¸u, 11 Apr 2014,
C.A. Ferreira Junior et al. 1373 (BHZB[digital image],
UEC); ibid., base do morro Ermo Ac¸u, 434205000W,
190200900S, 1430 m, 29 Apr 2016, A.V. Scatigna &
C.A. Ferreira Junior 1116 (UEC); idem A.V. Scatigna
& C.A. Ferreira Junior 1119 (BHCB, UEC).
Distribution, habitat, and ecology
Lapaea cipoensis is restricted to the Brazilian state of
Minas Gerais, only known from the Reserva Particular
onio Natural (RPPN) Brumas do Espinhac¸o, a
private Natural Reserve, at elevations around 1400 m
asl, in Serra do Cip
o (in the central portion of Southern
Espinhac¸o). It occurs in the edges of lapas in an area of
campo rupestre. Scatigna et al. (2017) observed that, as
happens in closely related species, the capsules of L.
cipoensis are upfacing, as a result of the pedicel, with
functionally poricidal dehiscence that may be related to
seed dispersal by raindrops and/or wind. Specimens
were collected with flowers and fruits in April.
Lapaea cipoensis has been assessed as CR (Critically
Endangered) by Scatigna et al. (2017), meeting criteria
B1ab(iii) and B2ab(iii). The species presents AOO and
, and is known from a single location.
The area and quality of the suitable habitat are under
continuous decline due to fire, cattle grazing, invasion
of exotic grasses, and erosion caused by tourist activ-
ities, such as hiking, as observed by Gualtieri-Pinto
et al. (2008) and Ribeiro et al. (2005).
Lapaea harleyi (B.L.Turner) Scatigna, comb. nov.
Stemodia harleyi B.L.Turner in Turner & Cowan,
Phytologia 75: 292. 1993. TYPE: BRAZIL. Bahia,
e, about 5 km along Andara
12580S, ca. 900 m, R.M. Harley et al. 20667 (holotype:
CTES; isotypes: CEPEC, K[digital image], SPF).
Lapaea harleyi is very similar to L. rubriflora in the
habit and villose indument on vegetative parts, but dif-
fers from the latter in the corolla tube being much
shorter (5.5–8 mm vs. 15–18.2 mm long) and lilac to
blue, with white and yellow nectar guide (vs. red, with-
out nectar guide); longer petioles (0.2–2.3 cm vs.
0.2–0.5 cm long); in the flat leaf blade margin (vs. revo-
lute); shorter sepals (5.5–9.3 mm vs. 7.5–10.1 mm long);
and leaf shape (ovate to broad-elliptical vs. elliptical),
texture (membranous vs. sub-coriaceous), and surface
(smooth vs. bullate).
Subshrubs, up to 70 cm tall. Stems crawling to ascend-
ing, densely covered with long, flexuous, minutely capi-
tate trichomes, interspersed with short capitate trichomes
and sessile glands. Petioles 0.2–2.3 cm long, indument
similar to stems; blades ovate to broad-elliptical,
0.9–7.2 0.8–5.1 cm, membranous, abaxial surface cov-
ered with long, flexuous, minutely capitate trichomes,
interspersed with short capitate trichomes and sessile
glands concentrated on veins, adaxial densely covered
with long, flexuous, minutely capitate trichomes, inter-
spersed with short capitate trichomes and sessile glands
throughout, apex acute to obtuse, base obtuse, round to
truncate, attenuate, margins usually crenate, rarely sub-
entire or argute, plane. Pedicels 1.5–3.5 cm long at
anthesis, up to 4.5 cm when fruiting, indument same as
stems; sepals lanceolate to linear-lanceolate,
5.5–9.3 1–2.5 mm, externally covered with long, flexu-
ous, minutely capitate trichomes, interspersed with short
capitate trichomes and sessile glands on veins and mar-
gins, internally sparsely covered with rare sessile glands,
apex with an apparently epidermal thickening; corolla
strongly bilabiate, lilac to blue with white to yellow
8 A. V. Scatigna et al.
patch on throat; tube 5.5–8 mm long, dorsiventrally
compressed towards throat, base slightly inflated, exter-
nally glabrous to subglabrous, internally villous at the
insertion of filaments; upper lip ascending, obscurely 2-
lobed, 5.3–65.8–6.2 mm, apex emarginate, lobes api-
ces mucronulate; lower lip 3-lobed, 2-canaliculate,
5.5–6.1 6–6.2 mm, villous, with long, minutely capi-
tate trichomes on throat, lobes 3.1–3.8 2.8–3.2 mm,
apices emarginate to mucronulate. Anterior pair of sta-
mens 4.6–4.7 mm long, posterior 2.7–3.1 mm long; the-
cae 0.6–0.8 mm long. Ovary 2.3–2.5 1.3 mm;
style 5.0–5.2 mm long., apex 0.7–0.9 mm long.
Capsule 5.5–73.3–3.5 mm; seeds cylindrical,
0.6–0.8 0.3–0.5 mm.
BRAZIL. Bahia: Andara
ı, Igatu, 19 Feb 2011, A.A.C.
Miranda & Chiquinho 14 (HUEFS); ibid., Distrito de
Igatu, Grota do Brej~
ao, 434205000W, 190200900S,
1430 m, 15 Dec 1999, J.G. Jardim et al. 2321 (CEPEC,
NY[digital image]); ibid., Serra do Sincor
15 km North of Mucug^
e on road to Andara
41200W, 12570S, ca. 1100 m, 18 Feb 1977, R.M.
Harley et al. 18865 (CEPEC, K[digital image], SPF,
ois, Fazenda Salobrinho, 41230W, 12320S,
R. Funch 7 (HUEFS); ibid., ca. 1 km do in
estrada lateral que sai da Rodovia Lenc¸
23 km NW de Lenc¸ois, 900–1000 m, 16 Oct 1994, V.C.
Fig. 4. Lapaea spp. in natural habitat. (4.1)L. cipoensis;(4.2)L. harleyi;(4.3)L. lobata;(4.4)L. rubriflora;(4.5)L. stellata;(4.6)
Specimens of L. lobata growing in a lapa in Ouro Preto, Minas Gerais.
Lapaea, a New Genus from Brazil 9
Souza et al. CFCR14118 (ESA, K[digital image], NY,
SPF); ibid., Chapadinha, 412602500W, 122703500S,
910 m, 27 Oct 1994, A.M. Carvalho et al. PCD1079
(ALCB, CEPEC, ESA, HUEFS, HRB, K[digital image],
erio Bizantino, trilha para o cru-
zeiro, 412204000W, 130002500S, 19 Jan 2015, A.V.
Scatigna & J.A.M. Carmo 651, 652, 653 (UEC); ibid.,
Pico do Gobira, 412204000W, 130500500S, 20 Jan 2005,
R.M. Castro & S.M. Lambert 1088 (ESA, HUEFS);
ibid., Serra do Esbarrancado, no topo da serra,
413003300W, 124305100S, 01 Nov 2011, R.P. Oliveira
et al 1969 (HUEFS); ibid., 10 km N na rod. para
1900800W, 125705700S, 27 Jan 2001, J.G.
Jardim et al. 2538 (CEPEC, ESA, NY[digital image]);
ibid., encosta da Serra na subida para o campo do
Gobira, 24 Jan 2000, L.P. Queiroz et al. 5648 (CEPEC,
ESA, HUEFS); ibid., 10–15 km ao NW de Mucug^
estrada para Andara
200W, 12570S, 1000 m 27 Jul
1979, S.A. Mori et al. 12688 (CEPEC, K[digital image],
NY); ibid., campo defronte ao cemit
erio, 20 Jul 1981,
A.M. Giulietti et al. CFCR1397 (ESA, K, SPF);
Palmeiras, Morro da M~
acia, 41230W, 12340S, 18
Nov 2006, J.S. Novais et al. 39 (HUEFS); ibid., Morro
do Pai In
acio, km 224 da rodovia BR 242, ca. 41270W,
12300S, 19 Dec 1981, G.P. Lewis et al. 899 (CEPEC,
K[digital image], RB); ibid., Pai In
Fig. 5. Line drawing of Lapaea rubriflora.(5.1) Reproductive branch; (5.2) Calyx with sepals connate at base and erect style; (5.3)
Dissected corolla tube and androecium; (5.4) Stamen with arm-like connective separating thecae; (5.5) Gynoecium; (5.6) Cross
section of ovary; (5.7) Capsule and persistent calyx; (5.8) Cross section of capsule. Drawn by Rog
erio Lupo after R.M. Harley et al.
10 A. V. Scatigna et al.
12270S, 19 Nov 1983, L.R. Noblick & A. Pinto 2794
(HUEFS, UEC); ibid., Pai In
acio, BR242, W of Lenc¸
at km 232, 12 Jun 1981, S.A. Mori & B.M. Boom 14396
(CEPEC, K[digital image], NY[digital image]); ibid.,
Morro do Pai In
acio, 16 Nov 1983, G.C. Pereira Pinto
et al. 443/83 (HRB); ibid., Pr
oximo ao Rio
Mucugezinho, Rod. Lenc¸
ois–Seabra, ca. 21 km NW de
ois, 412701000W, 122702700S, 900–1100 m, 17 Feb
1994, V.C. Souza et al. CFCR14168 (ESA, K, NY,
SPF); ibid., Morro do Pai In
122702000S, 1080 m, 25 Oct 1994, A.M. Carvalho et al.
PCD1005 (ALCB); ibid., Pai In
122703100S, 1150 m, 21 Nov 1994, E. Melo et al.
PCD1145 (ALCB, CEPEC, ESA, K[digital image],
SPF); idem, E. Melo et al. PCD1166 (ALCB, ESA).
Distribution, habitat, and ecology
Lapaea harleyi is restricted to the Brazilian state of
Bahia, being known from the Chapada Diamantina, in
the municipalities of Andara
Palmeiras. It grows at the shady bases of large rocky
blocks in areas of campo rupestre. Specimens were col-
lected with flowers and fruits especially between
October and March; few records are available of collec-
tions in June and July.
Lapaea harleyi was assessed as VU (vulnerable) by the
CNCFlora Red List Project (Souza et al., 2013) and
included in the same category in the last official list of
threatened species of the Brazilian Flora (Minist
Meio Ambiente (MMA), 2014). The EOO
) was probably overestimated by Souza
et al. (2013), because they included in their analysis
specimens of L. rubriflora misidentified as L. harleyi.
More recently, the species was included in the official
list of threatened species of the Flora of Bahia under the
category VU (vulnerable; Secretaria do Meio Ambiente
[SEMA], 2017). However, according to our evaluation,
L. harleyi should be considered endangered (EN), based
on: meeting criteria B1ab(iii) and B2ab(iii), AOO ¼
and EOO ¼971.241 km
; populations severely
fragmented; and an observed continuous decline of area
and quality of the suitable habitat due to historical min-
ing activities, intensified cattle grazing, and invasion of
exotic grasses facilitated by constant fire (Ganen &
Lapaea lobata (J.A.Schmidt) Scatigna, comb. nov.
Stemodia lobata J.A.Schmidt, in Martius, Fl. Bras. 8(1):
299. 1862. TYPE: BRAZIL. Minas Gerais, Monte
Itacolumi, L. Riedel s.n. (lectotype: LE barcode
LE00004903[digital image], here designated; isolecto-
type: LE barcode LE00004904[digital image]). Fig. 4.3.
¼Stemodia damaziana Beauverd, Bull. Herb.
Boissier 2 (7): 151. 1907. TYPE: BRAZIL. Morro de
ao, L.B. Damazio 294 (holotype: G barcode
G00343865[folder with two sheets sharing a single ori-
ginal label] [digital image]; isotypes: G barcodes
G00343874[digital image], G00343863[digital image]).
Lapaea lobata can be recognized for being very aro-
matic, its glandular-puberulent indument formed mainly
by short, capitate trichomes on stems and pedicels, and
its broadly and deeply crenate, sometimes almost lobed
leaf blades. Lapaea lobata differs from the species that
are most similar to it, L. cipoensis and L. stellata, for
the entirely deep violet corolla, with glabrous throat,
and the tube not dorsiventrally compressed (vs. pale
lilac to deep blue or purple, with white patch on the
throat, and tube dorsiventrally compressed).
Herbs to subshrubs, up to 40 cm tall. Stems ascending
to erect, glandular-puberulent, entirely densely covered
with short, capitate trichomes, with rare stiff, non-capi-
tate trichomes on nodes. Petioles 0.3–1 cm long, indu-
ment same as the stem; blades ovate 1–50.8–3 cm,
chartaceous, abaxial surface covered with short capitate
trichomes and sessile glands throughout with some rare
longer, curved, non-capitate trichomes, these exclusively
on veins, adaxial surface covered with short capitate tri-
chomes and sessile glands throughout with some rare
longer, curved, non-capitate trichomes scattered over the
surface and near margin, apex acute, obtuse to round,
base truncate to round, margins crenate, plane. Pedicels
1.1–2 cm long, densely covered with short capitate tri-
chomes and sessile glands; sepals lanceolate,
4.2–6.5 1.1–2.1 mm, sparsely covered with short, capi-
tate trichomes, with few longer, curved, non-capitate tri-
chomes on veins and margins, apex acute to acuminate;
corolla strongly bilabiate, entirely deep violet; tube
0.9–12 mm long, base slightly inflated, externally glab-
rous, internally pubescent on filaments insertion; upper
lip patent, obscurely 2-lobed, 5.8–75.1–6.1 mm, apex
truncate to slightly emarginated; lower lip 3-lobed,
6.1–6.8 8.1–8.8 mm, glabrous, lobes,
4–4.3 4.4–4.8 mm apices truncate to shallowly emargi-
nate. Anterior pair of stamens 7.3–8.2 mm long, poster-
ior 4.4–5 mm long; thecae 0.5–0.7 mm long. Ovary
2.1–2.5 1.2–1.3 mm; style 6.9–7.5 mm long. apex
Lapaea, a New Genus from Brazil 11
0.2–0.4 mm long. Capsule 5.1–73.1–3.6 mm mm;
seeds ovoid, 0.5–0.7 0.3–0.4 mm.
In the protologue of Stemodia lobata, Schmidt (1862)
cited a specimen as “In monte Itacolumi prov.
Minarum: Riedel. Floret Febr. Pl. Oreas”but did not
specify in which herbarium it was housed. Turner and
Cowan (1993b) designated as lectotype a specimen at
HBG, cited with a question mark, and indicated two
sheets housed at LE as isolectotypes. However, due to
the use of a question mark, their typification does not
take effect (Art. 7.11 of the Shenzen code; Turland
et al., 2018). We have located the two specimens of S.
lobata at LE, both bearing Riedel’s label written “In
monte Itacolumi Febr 35”, but we could not locate the
HBG specimen cited by Turner and Cowan (1993b).
When consulted, the herbarium curator also declined
that this specimen was housed there (M. Schultz, per-
sonal communication, 18 March 2019). Therefore, we
have designated the specimen at LE (barcode
LE00004903) as lectotype, being the most complete ori-
ginal specimen found.
In the protologue of S. damaziana, Beauverd (1907)
cited the type as 'Typus in Herb. Barbey-Boissier' and
'n294 [ …] Leg. Damazio. –Det. Beauverd'. Turner
and Cowan (1993b) cited one specimen from L.B.
Damazio 294 at G-BOIS as holotype, and three others
as isotypes, one of them also at G-BOIS, and the
remainder at G-DC. These specimens are now housed in
the general collection, G. The specimen G (barcode
G00343865) consists of two sheets, the second being
double-sized and folded over the first, that is, the first
sheet (annotated by B.L. Turner in 1992 as holotype) is
physically included in the second one (annotated iso-
type). This is usual in G’s general collection, because of
the small format of the sheets (Gautier et al., 2016).
Furthermore, in this specific case, it is likely that the
specimen was originally mounted on a single sheet in
the Boissier herbarium, which had a much larger format
and, when incorporated in the general collection, it had
to be separated into two sheets (L. Gautier, personal
communication, 3 June 2019). Since both sheets share a
unique original label (by L.B. Damazio), they should be
considered as holotype sheet 1/2 and sheet 2/2 (Art.
8.3., ex. 9, of the Shenzen code; Turland et al., 2018)
and not holotype and isotype, as annotated by Turner.
The specimen G barcode G00343874 bears a label with
'ex Herbier Boissier', along with information on the pub-
lication of the new species (S. damaziana), in
Beauverd’s handwriting, and is possibly a fragment sep-
arated from the holotype after Beauverd’s publication,
which was then gifted to Augustin De Candolle (L.
Gautier, personal communication, 3 June 2019). In this
case, it should be considered an isotype (Art. 8.3., ex.
8). The remaining specimen, G barcode G00343863,
was not part of Boissier’s herbarium, and does not bear
any annotation in Beauverd’s handwriting; hence, we
assume that Beauverd did not see this specimen, and,
therefore, it should be treated as an isotype.
BRAZIL. Minas Gerais, Mariana, Parque Estadual do
ao, 432704900W, 202602500S, 1429m, 23
Apr 2010, E.S. C^
andido et al. 399 (OUPR); ibid.,
Serrinha, 7 Feb 2003, M.C.T.B. Messias 769 (OUPR);
[Ouro Branco], Serra de Itatiaia prope Chapada, 12 May
1895, C.A.W. Schwacke 11492 (P); Ouro Preto,
Andorinhas, 11 Jun 1975, J. Badini s.n. (OUPR); ibid.,
Parque Municipal Cachoeira das Andorinhas, 432903800W,
202103900S, 02 May 2016, A.V. Scatigna et al. 1142,
1143, 1144 (UEC); ibid., Andorinhas, s.col. 09 May 1971
(OUPR); ibid., Camarinhas, 07 Jun 1973, J. Badini s.n.
(OUPR); ibid., Camarinhas, 1940, J. Badini s.n. (OUPR);
ibid., Serra do Itacolomi, 1937, J. Badini, s.n. (OUPR);
ibid., Alto do Itacolomi, 1938, J. Badini s.n. (R); ibid.,
Morro de S~
ao, A.P. Alves s.n. (R); ibid., Morro
ao, L.B. Damazio 763 (OUPR); ibid., Morro
ao, 1904, L.B. Damazio s.n. (BHCB); ibid.,
Serra de Lavras Novas, 10 Apr 1972, J. Badini s.n.
(OUPR); Serra de Lavras Novas, 11 May 1974, J. badini
s.n. (OUPR); Serra das Lavras Novas, localidade de
Venda do Campo, 27 Mar 2001, M. Groppo Jr. & M.
Ulwin 686 (SPF, UEC); ibid., Lavras Novas, Serra do
3103200W, 202704700S, 03 May
2016, A.V. Scatigna & T.V. Bastos 1145, 1146, 1147,
1148 (UEC); Lavras Novas, s.col. (OUPR).
Dubious location, Minas Gerais, Serra do Cip
o, 3 Mar
1958, E.P. Heringer & A. Castellanos 5991 (R, UB).
Distribution, habitat, and ecology
Lapaea lobata is restricted to Minas Gerais, Brazil, with
most records from Mariana and Ouro Preto, in the Iron
Quadrangle, southernmost portion of the ER, at eleva-
tions around 1400 m asl. The only record from Serra do
o(Heringer & Castellanos 5991 [R, UB]) was col-
lected in 1958 and, despite the extensive fieldwork con-
ducted in the region in the following 60 years, the
species has never been recollected there; moreover, the
R herbarium sheet bears the observation 'Et. Horto
Florest. Paraopeba', suggesting that the specimen was
originated in cultivation. This raises doubt on the correct
provenance of this specimen. The species grows on
12 A. V. Scatigna et al.
humid and shaded edges of quartzite boulders, in areas
of campo rupestre. Specimens with flowers and fruits
were collected between March and June, with one
record in October and another in December.
We assessed Lapaea lobata as CR (critically endan-
gered), meeting criterion B1ab(i,ii,iii). The species has
an EOO of 32.575 km
, with severely fragmented popu-
lation and observed continuous decline of AOO, EOO
and area and quality of the suitable habitat. The region
has historically suffered intense degradation from gold
mining activities, and is currently under pressure from
iron mining activities and human occupation (Carmo &
Kamino, 2015). In 2015, the region of Mariana has suf-
fered the largest environmental tragedy in Brazil, due to
the rupture of a dam in a mining company area, with
serious damage to local vegetation (Lopes, 2016).
Lapaea rubriflora Scatigna & V.C. Souza, sp. nov.
Figs 4.4, 5, 6 and 7.
TYPE: BRAZIL. Bahia, Aba
ıra, Campo de Ouro
Fino, 41540W, 13150S, 1600–1700 m, 10 Jan 1992,
R.M. Harley et al. H50738 (holotype: HUEFS; isotypes:
CEPEC, ESA, K[digital image], NY[digital image],
ETYMOLOGY: The epithet [rubriflora] refers to the
red-coloured corolla (Figs 4.4 and 5.1, 5.2), a unique
feature in the whole tribe Gratioleae.
DIAGNOSIS: Lapaea rubriflora is characterized by
the unique red corolla. It is similar to L. harleyi in its
general morphology, but differs in leaf shape (elliptical
vs. ovate to broad-elliptical); petiole length (0.2–0.5 cm
vs. 0.2–2.3 cm long); blade margin (revolute vs. flat)
and surface (bullate vs. smooth); sepal length
(7.5–10.1 mm vs. 5.5–9.3 mm long); length of the cor-
olla tube(15–18.2 mm vs. 5.5–8 mm long) and corolla
colour (red, without nectar guide vs. lilac to blue with
white and yellow nectar guide).
Subshrubs, 30–80 cm tall. Stems terete to sub-quadran-
gular, branched, covered with short capitate trichomes
interspersed with long, flexuous, minutely capitate tri-
chomes, denser towards apex. Petioles 0.2–0.5 cm long,
densely covered with short capitate trichomes and long
minutely capitate trichomes; blades ovate to elliptical,
1.8–4.1 1–2.5 cm, sub-coriaceous, bullate, abaxial sur-
face densely covered with short capitate trichomes and
longer minutely capitate trichomes, these concentrated
on prominent veins and on margin, adaxial surface cov-
ered with short capitate trichomes interspersed with
longer minutely capitate trichomes throughout the sur-
face, apex acute to obtuse, round, base cuneate to
obtuse, margin crenate to serrate, clearly revolute.
Pedicels 1.5–2.5 cm long, covered with short and long
capitate trichomes, densely towards flower, upcurved
after fruiting; sepals lanceolate, 7.5–10.1 1.8–2.2 mm,
covered with short, capitate trichomes, with few longer,
minutely capitate trichomes on veins and margins,
denser on base, apex acute to acuminate; corolla slightly
bilabiate, red; tube 15–18.2 mm long, cylindrical,
slightly gibbous at anthers region, externally covered
with short, capitate trichomes and longer minutely capi-
tate trichomes scattered over the surface, glabrous at
base, internally sparsely pubescent, denser on filaments
insertion; upper lip erect, shallowly 2-lobed,
4.5–57.5–8 mm, apex emarginated, externally glandu-
lar-pubescent; lower lip 3-lobed, throat glabrous, lobes
orbicular, 3.5–4 mm diam., externally glandular pubes-
cent. Anterior pair of stamens 7.9–8 mm long, posterior
pair 5.9–6 mm long; thecae 2–2.1 mm long. Ovary
3.4–3.5 1.5–1.6 mm; style 6.5 mm long, apex 0.5 mm
long. Capsule 7.5 4 mm; seeds ovoid, slightly com-
pressed, 0.8 0.2 mm.
PARATYPES: BRAZIL. Bahia, Aba
ıra, Serra ao Sul
do Riacho da Taquara, 41550W, 13150S, 1890 m, 10
Jan 1992, R.M. Harley et al. H51257 (CEPEC, E, ESA,
HUEFS, K[digital image], NY[digital image], SPF);
ibid., Campo do Cigano, 41550W, 13150S,
1700–1800 m, 25 Feb 1992, P.T. Sano H52323
(CEPEC, ESA, HUEFS, K[digital image]); ibid., Campo
de Ouro Fino, 41540W, 13150S, 1600–1700 m, 24 Jan
1992, J.R. Pirani et al. H50778 (ESA, HUEFS,
K[digital image], SPF); ibid., Catol
es, Serra do Barbado,
between 415400600W, 131702700S and 415402900W,
131705000S, 1750–2035 m, 26 Feb 1994, V.C. Souza
et al. CFCR14590 (CEPEC, E, ESA, HUEFS, K[digital
image], MO, NY, SPF); ibid., Distrito de Catol
Encosta da Serra do Atalho, subida pela boca do Le~
20 Apr 1998, L.P. Queiroz et al. 5075 (HUEFS[2
sheets], K[digital image], NY[digital image]); ibid.,
es, Trilha para o Campo do Ouro Fino,
415400700W, 130300800S, 11 Jan 2007, A.K.A. Santos &
Sr. Raimundo 961 (HUEFS); ibid., Catol
es, descida para
Mata dos Frios, 415300800W, 131805700S, 1450 m, 23
Oct 1999, E.B. Miranda et al. 292 (HUEFS); ibid.,
es, Trilha para Pico do Barbado, Forquilha,
415401500W, 131702700S, 1670 m, 17 Jan 2016, A.V.
Scatigna & J.A.M. Carmo 1035, 1042 (UEC).
Of the six types of trichomes described by Scatigna
et al. (2017), we observed the presence of three types
Lapaea, a New Genus from Brazil 13
(types 1, 3, and 4), all of them glandular, on the surfa-
ces of stems and leaves of Lapaea rubriflora (Fig. 6).
We did not observe any type of non-glandular trichomes
on the examined samples. Type 1 has a broad base, a
pedicel with two or three cells, and a rounded head,
seemingly unicellular (Fig. 6.1
); type 3 has a short uni-
cellular (rarely two-celled) pedicel and a wide, multicel-
lular, morular-shaped head (Fig. 6.1
); and type 4,
which is a minutely capitate, filiform, flexible, and uni-
seriate trichome, with variable length (200–1200 mm)
(Fig. 6.2). The three types of trichomes were more
abundant on the stem (Fig. 5.3) and on the abaxial sur-
face of the leaves (Fig. 6.4), with type 1 concentrated
on the blade and type 4 concentrated on the veins.
Lapaea rubriflora is readily distinguished from any other
species of Lapaea and Stemodia sensulatoduetoits
unique red corolla (Figs 4.4 and 5.1, 5.2). In an account of
the vascular plants of the Catol
(Zappi et al., 2003), specimens of L. rubriflora were iden-
tified as Stemodia harleyi (¼L. harleyi), a similar species
in the overall morphology; for comparison, see notes under
the latter. In what concerns the micromorphology, L. har-
leyi and L. rubriflora sharetype4trichomes,whichare
lacking in all other species of the genus; L. harleyi lacks
type 1 trichomes, which are present in L. rubriflora and
all remaining species of Lapaea.
Distribution, habitat, and ecology
Lapaea rubriflora is restricted to Bahia, being only
known from the Catol
es region, in the Chapada
Diamantina. It grows on fractures of rocky blocks (Fig.
7.2, 7.3, 7.4) in areas of campo rupestre, at elevations
of 1500–2000 m asl. Flowering and fruiting specimens
were collected mainly between January and April, with
one record in October. The unique red-coloured corolla,
with a relatively long tube (Figs 4.4, 5.3 and 7.1, 7.2),
may indicate the first case of ornithophily in the
Gratioleae, but studies in reproductive biology are cru-
cial to test this hypothesis.
We assessed Lapaea rubriflora as CR (critically endan-
gered) under criterion B1ab(iii). The species has an
Fig. 6. Scanning electron microscopy of vegetative parts of Lapaea rubriflora.(6.1
) Type 1 trichome; (6.1
) Type 3 trichome; (6.2)
Type 4 trichome; (6.3) Stem; (6.4) Abaxial surface of leaf. From Scatigna & Carmo 1035 (UEC).
14 A. V. Scatigna et al.
EOO of 16 km
, and is known to occur in only one
locality. Furthermore, a continuous decline in the area
and quality of the suitable habitat has been observed,
due to historical mining activities and currently growing
cattle grazing, in addition to invasion of exotic grasses,
facilitated by constant fire (Ganen & Viana, 2006).
Lapaea stellata (B.L.Turner) Scatigna, comb. nov.
Stemodia stellata B.L.Turner, in Turner & Cowan,
Phytologia 75(4): 312. TYPE: BRAZIL. Minas Gerais,
eastern slopes of Pico do Itamb
e, ca. 1700 m, 11 Feb
1972 W.R. Anderson et al. 35828 (holotype: MO[digital
image]; isotypes: MBM, NY[digital image], K[digital
image], UB). Fig. 4.5.
Lapaea stellata is readily distinguished from any other
known species of Lapaea and Stemodia s.l. by the
presence of dendritic trichomes over the surfaces of
Herbs or subshrubs, up to 35 cm tall. Stems crawling to
ascending, densely covered with dendritic, non-capitate
trichomes interspersed with short, capitate trichomes and
sessile glands, denser towards apex. Petioles 0.1–0.6 cm
long, indument same as stem; blades ovate, elliptical or
obovate, 0.2–20.2–1.5 cm, chartaceous, apex acute to
obtuse, round, base acute to obtuse-attenuate margins
serrate to crenate-serrate, plane, both abaxial and adaxial
surfaces densely covered with dendritic, non-capitate tri-
chomes interspersed with short, capitate trichomes and
sessile glands, denser towards apex of branches.
Pedicels 0.5–1.4 cm long, covered with dendritic, non-
capitate trichomes interspersed with short, capitate tri-
chomes and sessile glands; sepals lanceolate,
Fig. 7. Lapaea rubriflora in natural habitat. (7.1) Flowering branch; (7.2) Habit; (7.3) Specimen of L. rubriflora under a lapa;(7.4)
J.A.M.Carmo in the habitat of L. rubriflora.
Lapaea, a New Genus from Brazil 15
4–6.5 0.8–2 mm, apex acute to acuminate, covered
with dendritic, non-glandular trichomes on veins and
margins and capitate trichomes on blades; corolla
strongly bilabiate, whitish to lilac, with broad white
patch on throat, tube internally dark purple; tube
5.2–9 mm long, dorsiventrally compressed towards
throat, base slightly inflated, externally glabrous, intern-
ally villous at the insertion of filaments; upper lip
reflexed, 2-lobed, 4.5–5.1 6.1–6.4 mm, apex emargi-
nated, externally bearded; lower lip 3-lobed, 2-canalicu-
late, 6.5–6.7 8–8.2 mm, pubescent, with short, non-
capitate trichomes on throat, lobes with truncate to
emarginate apex, 3–3.2 4–4.3 mm. Anterior pair of sta-
mens 4.4–4.8 mm long, posterior 2.9–3.1 mm long; the-
cae 0.7–1 mm long. Ovary 1.6–1.8 1–1.1 mm; style
4.2–5.8 mm long. apex 0.5–0.6 mm long. Capsule
4.1–5.3 3–3.4 mm; seeds cylindrical,
0.6–0.8 0.3–0.4 mm.
Souza and Giulietti (2009) ascertained that L. stellata is
similar to S. microphylla in the overall morphology, but
the two species are readily differentiated by the 3-
whorled leaves and ebracteolate, non-resupinate flowers
in L. stellata, vs. opposite leaves and bi-bracteolate and
resupinate flowers in S. microphylla (Scatigna
et al., 2017).
BRAZIL. Minas Gerais, Santo Ant^
onio do Itamb
Parque Estadual do Pico do Itamb
e, [ …] along the
main trail., 432005300W, 182305500S, 1367–2038 m, 02
Mar 2009, F. Almeda et al. 9675 (UEC); ibid., Parque
Estadual do Pico do Itamb
e, trilha para o Pico do
e, 431901800W, 182400100S 07 Jul 2014, A.V.
Scatigna et al. 477 (UEC); ibid., Parque Estadual do
Pico do Itamb
e, trilha para o Pico do Itamb
431902500W, 182305700S, 07 Jul 2014, A.V. Scatigna
et al. 478 (UEC); ibid., Parque Estadual do Pico do
e, trilha para o Pico do Itamb
182305100S, 07 Jul 2014, A.V. Scatigna et al. 479
(UEC); ibid., Pico do Itamb
1500 m, 07 Apr 1998, V.C. Souza et al. 21126 (ESA);
ibid., Pico do Itamb
e, Subida para o pico, 432005400W,
182304500S, 08 Apr 2010, G.O. Rom~
ao et al. 2619
(ESA); ibid., Pico do Itamb
e, 431901800W, 182400200S,
1551 m, 14 Dec 2007, C.D.N. Rodrigues et al. 118, 119
(ESA); ibid., Caminho para o Pico do Itamb
1500–2000 m, 26 Feb 2002, V.C. Souza et al. 28474
Distribution, habitat, and ecology
Lapaea stellata is restricted to the state of Minas Gerais
and only known from the Pico do Itamb
e region, in the
municipality of Santo Antonio do Itamb
e, which is
encompassed by the Diamantina Plateau, a central-north-
ern portion of the Southern Espinhac¸o. As its most
closely related species, it grows in lapas of large rocky
blocks in areas of campo rupestre in elevations of
1300–2000 m. Specimens were collected with flowers
and fruits between December and April, and in July.
We assessed Lapaea stellata as CR (critically endan-
gered) under criteria B1ab(iii) and B2ab(iii). The species
has an AOO and EOO of 8 km
, and is known from
only one locality. Furthermore, a continuous decline of
the area and quality of the suitable habitat was observed,
due to invasion of exotic grasses after fire
We recognize Lapaea as a new genus segregated from
Stemodia with morphological and phylogenetic support.
It comprises five threatened species, L. cipoensis,L.
harleyi,L. lobata,L. rubriflora, and L. stellata, each
one presenting a narrow geographic distribution that is
consistent with currently recognized biogeographic units.
The geographically structured diversity found in the ER,
previously documented in unrelated lineages, was also
observed in Lapaea and Philcoxia, which identifies
these two genera as potential model-groups for recon-
structing biogeographic scenarios in campo rupestre.
We thank C.A. Ferreira-J
unior, G. Piassa, J.A.M.
Carmo, L. Echternacht and T. Bastos for field support;
A.C.D. Castello, F.A.O. Silveira, M.C.E. Amaral, P.T.
Sano and R.B. Pinto for suggestions on an early version
of the manuscript; A.R. Sim~
oes for English revision and
additional suggestions; R. Lupo for the line drawing; the
curators of all cited herbaria for all support, especially
Drs M. Schultz (HBG), L. Gautier (G) and V.
Dorofeyev (LE); Instituto Estadual de Florestas (IEF/
MG) and Instituto Chico Mendes de Conservac¸~
Biodiversidade (ICMBio) for collecting permits.
The authors declare no conflict of interest.
16 A. V. Scatigna et al.
This work was supported by the Fundac¸~
ao de Amparo
Pesquisa do Estado de S~
ao Paulo [FAPESP grant
number 2012/12927-9]; Conselho Nacional de
ıfico e Tecnol
ogico [CNPq grant
number 159924/2018-9]; Coordenac¸~
Aperfeic¸oamento de Pessoal de N
ıvel Superior [CAPES
grant number 001]; International Association for Plant
Taxonomy [IAPT Research Grants].
e Vito Scatigna http://orcid.org/0000-0002-
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