Selaginella PeCtoPuellenSiS (SelaginellaCeae,  
lyCoPodioPhyta): a neW SPeCieS from Serra do  
Peito-de-moça, bahia, brazil, With CommentS on the  
taxonomy of the SPeCieS rePorted in the Country  
Selaginella PeCtoPuellenSiS (SelaginellaCeae, lyCoPodioPhyta):  
una nueva eSPeCie de la Serra do Peito-de-moça, bahía, braSil Con  
ComentarioS Sobre la taxonomía de laS eSPeCieS rePortadaS en el PaíS  
1,2,3  
Iván A. Valdespino * , Christian A. López , Jorge I. Ceballos  
&
José M. Palacios  
Summary  
Background and aims: Our research on neotropical Selaginella led us to examine a  
collection from the Atlantic Forest of Serra do Peito-de-Moça, Bahia, Brazil, initially  
determined as S. muscosa but with unusual features, indicating it represented an  
undescribed species. We also reexamined the taxonomic and nomenclatural status  
of three Selaginella taxa previously described from Brazil and gathered information on  
introduced and naturalized species in the country.  
M&M: Morphological analysis used stereomicroscopes and scanning electron microscopy  
to distinguish the suspected new species from its morphologically nearest ally.  
Results: We confirmed that the plant material from the Atlantic Forest of Serra do Peito-  
de-Moça, Bahia, Brazil, is a new species that we named Selaginella pectopuellensis  
1
. Departamento de Botánica,  
Facultad de Ciencias Naturales,  
Exactas y Tecnología, Universidad  
de Panamá, Panamá  
2
(
3
. Sistema Nacional de Investigación  
SNI), SENACYT, Panamá  
. Smithsonian Tropical Research  
Institute, Panamá, Panamá  
Department of Integrative  
4
.
Biology, University of Texas at  
Austin, Austin, USA  
(Selaginellaceae). Additionally, we clarified the taxonomic and nomenclatural status  
of three Selaginella species previously described from Brazil and provided more  
information on naturalized taxa. Overall, we expanded the confirmed number of native  
Brazilian Selaginella species to 91 and confirmed six introduced and naturalized taxa  
in the country.  
*
iavaldespino@gmail.com  
Citar este artículo  
Conclusions: The description of S. pectopuellensis adds to our knowledge of Brazil's  
pteridophyte flora. We synonymized S. kriegeriana under S. magnafornensis and  
clarified that S. bahiensis subsp. bahiensis and S. bahiensis subsp. manausensis should  
not be recognized as separate taxa. We also confirmed six naturalized Selaginella taxa  
in Brazil.  
VALDESPINO, I. A., C. A. LÓPEZ,  
J. I. CEBALLOS & J. M. PALACIOS.  
2
024. Selaginella pectopuellensis  
Selaginellaceae, Lycopodiophyta):  
new species from Serra do  
(
a
Peito-de-Moça, Bahia, Brazil, with  
comments on the taxonomy of the  
species reported in the country. Bol.  
Soc. Argent. Bot. 59: 393-403.  
Key WordS  
Atlantic Forest, endemism, idioblasts, lycophytes, papillae.  
reSumen  
Introducción y objetivos: Nuestra investigación sobre Selaginella neotropical nos llevó  
a examinar una colección del Bosque Atlántico de Serra do Peito-de-Moça, Bahía,  
Brasil, inicialmente identificada como S. muscosa, pero con características inusuales  
que indicaban que representaba una especie no descrita. Además, reexaminamos la  
taxonomía y nomenclatura de tres taxones de Selaginella previamente descritos en  
Brasil e información sobre especies introducidas y naturalizadas en el país.  
M&M: Se realizó un análisis morfológico utilizando estereomicroscopios y microscopía  
electrónica de barrido para distinguir la supuesta nueva especie de su aliada  
morfológicamente más cercana.  
Resultados: Confirmamos que el material vegetal del Bosque Atlántico de Serra do  
Peito-de-Moça, Bahía, Brasil, es una nueva especie y la nombramos Selaginella  
pectopuellensis (Selaginellaceae). Además, clarificamos el estado taxonómico y  
nomenclatural de tres especies de Selaginella previamente descritas para Brasil y  
adicionamos información sobre taxones naturalizados. En general, ampliamos el  
número confirmado de especies nativas de Selaginella en Brasil a 91 y confirmamos la  
ocurrencia de seis taxones introducidos y naturalizados en el país.  
Conclusiones: La descripción de S. pectopuellensis amplía nuestro conocimiento de la  
flora de pteridofitas de Brasil. Sinonimizamos S. kriegeriana bajo S. magnafornensis y  
aclaramos que S. bahiensis subsp. bahiensis y S. bahiensis subsp. manausensis no  
deben reconocerse como taxones separados. Igualmente, confirmamos seis taxones  
de Selaginella naturalizados en Brasil.  
Recibido: 15 Abr 2024  
Aceptado: 4 Jul 2024  
Publicado en línea: 30 Sep 2024  
Publicado impreso: 30 Sep 2024  
Editora: Olga G. Martínez  
PalabraS Clave  
Bosque Atlántico, endemismo, idioblastos, licófitas, papilas.  
ISSN versión impresa 0373-580X  
ISSN versión on-line 1851-2372  
393  
Bol. Soc. Argent. Bot. 59 (3) 2024  
introduCtion  
our analysis of the protologue, including SEM  
images, of S. kriegeriana L.A. Goés, listed in Flora  
The Selaginellaceae of Brazil are highly diverse e Funga do Brasil (2020) as a native species, leads  
and species-rich (Valdespino, 2015a; Valdespino us to consider it conspecific with S. magnafornensis  
et al., 2018a, b; Flora e Funga do Brasil, 2020). Valdespino & C. López, which has nomenclatural  
Alston et al. (1981) listed 45 native Selaginella precedence. In consequence, only 90 Selaginella  
species, including two subspecies for Brazil, while species are well-documented as native to Brazil,  
Valdespino (2018a, b) estimated 89–96 native while six are introduced and somewhat naturalized  
species to occur in that country. Flora e Funga do (i.e., S. kraussiana, S. pallescens, S. plana, S.  
Brasil (2020) indicates a total of 97 Selaginella viticulosa, S. vogelii, and S. willdenowii), and  
species and one subspecies as occurring within another (i.e., S. kriegeriana) is a recent synonym of  
Brazil, as stated in the document’s summary. a previously described taxon. These data serve to  
This number, however, is somewhat unclear illustrate that our overall knowledge of Selaginella  
since some of the listed taxa are introduced; one diversity in Brazil and, for that matter, in other  
is a synonym of a previously described taxon, South American countries such as Colombia,  
and the listed subspecies do not have taxonomic Venezuela, Ecuador, Peru, and Bolivia, is still far  
standing. Specifically, S. kraussiana (Kunze) A. from complete (Smith & Kessler, 2018; Valdespino  
Braun, S. pallescens (C. Presl) Spring, S. plana etal., 2018a, b). Indeed, asanticipatedbyValdespino  
(Desv.) Hieron., S. viticulosa Klotszch, S. vogelii et al. (2018a), new and undescribed species are still  
Spring, and S. willdenowii (Desv.) Baker are to be uncovered in these countries. In the case of  
non-native, but introduced and naturalized taxa. Brazil, without a doubt, the country is proving to  
Likewise, S. viticulosa is known to naturally be a Selaginella treasure trove, and within it, the  
occur only in Costa Rica and Panama, while it has Bahia State has yielded recent significant findings  
perhaps been introduced to Jamaica and Bermuda (Valdespino 2015a, b; Valdespino et al., 2015).  
in the West Indies, and in Colombia, Venezuela,  
The state of Bahia includes an extensive territory  
as well as in the island of Trinidad in Trinidad with diverse ecosystems, among which the Atlantic  
and Tobago (Baksh-Comeau, 2000) in South Forest (Mata Atlântica) along its eastern part is  
America. Specimens documenting S. viticulosa a biodiversity hotspot due to the high number of  
in Brazil, e.g., material collected in and around endemic and threatened plant species found there  
neighborhoods of Rio de Janeiro (i.e., Brade 16105, (Myers et al., 2000; Giulietti et al., 2006; Matos  
RB barcode RB00702089-image!; Brade s.n., et al., 2010; Ostroski et al., 2018; Flora e Funga  
RB barcode RB00691682-image!), the Botanical do Brasil, 2020). Furthermore, Bahia is the sixth  
Garden of Rio de Janeiro (i.e., Nascimento 7, RB state with the highest number of lycophytes and  
barcode RB01417493; Oliveira 02, RB barcode ferns in Brazil. It harbors 18 known Selaginella  
RB00316275;Simões76,RBbarcodeRB00316244, species, which makes it the seventh in that country  
8
7, RB barcode RB00316245; Winter 68, RB in terms of the number of taxa in that genus (Flora  
barcode RB00533100-image!, 69, RB barcode e Funga do Brasil, 2020), including four endemic  
RB00316191, 73, RB barcode RB00316187, 102, Selaginella: S. bahiensis Spring (Spring, 1840), and  
RB barcode RB00316116 – all RB images!), and the recently described S. blepharodella Valdespino,  
the Parque Nacional da Tijuca (i.e., Mynseen 1365, S. crinita Valdespino, and S. mucugensis Valdespino  
RB barcode RB00726501-image!), originate from (Valdespino et al., 2015). In this study, we further  
well-known areas with introduced, cultivated or describe Selaginella pectopuellensis Valdespino &  
naturalized flora. Additionally, S. bahiensis Spring C. López as an additional new and endemic species  
subsp. manausensis (Bautista) Jermy & J.M. Rankin from the Bahia State, Brazil.  
(≡ S. manausensis Bautista) is conspecific with and  
a later heterotypic synonym of S. palmiformis  
Alston ex Crabbe & Jermy, therefore S. bahiensis materialS and methodS  
subsp. bahiensis is a homotypic synonym of S.  
bahiensis (Valdespino et al., 2015) and should not  
One gathering of Selaginella pectopuellensis was  
be listed as an independent taxon. Furthermore, initially examined at the Herbarium of the New York  
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I. A. Valdespino et al. - Selaginella pectopuellensis from Bahia, Brazil  
Botanical Garden (NY) using a stereomicroscope  
Description. Plants terrestrial. Stems long-  
(
i.e., Olympus SZ 60-STS). Further examination creeping, stramineous, 8–12 cm long, 0.8–1.0  
of loaned material was undertaken at the University mm diam., non-articulate, not flagelliform,  
of Panama herbarium (PMA) using an Olympus non-stoloniferous, sparsely 2- or 3-branched.  
SZX16 to better ascertain leaf and spore characters. Rhizophores ventral, stout, borne throughout  
Air-dried samples of stem sections, leaves, and proximal ⅓−½ of stems, 0.2–0.5 mm diam.  
spores were sputter coated with gold-palladium and Leaves heteromorphic throughout, coriaceous,  
examined with a Zeiss Model Evo 40vp Scanning upper surfaces green or greenish-brown when  
Electron Microscope (SEM) at 10–15 kV at the old or due to fixing and drying processes, lower  
Smithsonian Tropical Research Institute (STRI). surfaces silvery green or light-brown when old or  
Digitized SEM images of plant sections, leaves, due to sample processing. Lateral leaves distant,  
and spores were taken at different magnifications, ascending to slightly perpendicular to stems or  
post-processed, and assembled in multipart figures slightly imbricate along distal portion of stem and  
using Adobe Photoshop, as explained in Valdespino branches, ovate, ovate-cordate to cordate, 1.8–2.5 ×  
(
2016). Terminology and measurements used in 1.0–1.5 mm; bases subcordate, glabrous, acroscopic  
species descriptions follow Valdespino (2019) and bases overlapping stems, basiscopic bases free  
Valdespino & López (2019), while conservation from stems; acroscopic margins on both surfaces  
status was determined according to the IUCN continuously bordered by a hyaline band comprised  
Standards and Petitions Committee (2022).  
of idioblasts, the band 3–7 cell wide, each idioblast  
elongate, straight-walled, and papillate, the papillae  
in one or two rows over each cell lumen and  
submarginally on upper surfaces bordered by a  
broad band of 5–10 round, sinuate-walled, and  
reSultS  
Selaginella pectopuellensis Valdespino & C. papillate cells along distal ½, the papillae 5–22  
López, sp. nov. TYPE: BRAZIL. Bahia, Arataca, on each cell lumen, basiscopic margins on upper  
Serra do Peito-de-Moça, road joining Arataca to surfaces continuously bordered by a hyaline band  
Una, ca. 22.4 km from Arataca, entrance to Sto. comprised of idioblasts, the band 1–3 cell wide, each  
Antônio settlement, RPPN Caminho das Pedras, idioblasts similar to those on acroscopic margins on  
1
5°10’25”S, 39°20’30”W, 1000 m, 13-IV-2007, F. upper surfaces and submarginally, continuously  
B. Matos, S. Vieira, M. Lopes, S. Sant’Ana & L. C. bordered by a band of cells similar to that found  
Gomes 1360 (Holotype, NY barcode NY1686456!; submarginally, along distal ½ of acroscopic margins  
isotypes, CEPEC-n.v., PMA!, UPCB barcode on upper surfaces, the basiscopic margins on lower  
UPCB0010917-image!). Figs. 1-5.  
surfaces, bordered by a band of papillate idioblasts,  
the band 2–6 cell wide, each idioblast as those in  
Diagnosis. Selaginella pectopuellensis differs acroscopic margins and both the acroscopic and the  
from S. muscosa Spring by its coriaceous (vs. basiscopic submarginal portion of laminae in lower  
chartaceous) leaves that lack elongate idioblasts surfaces without a band of rounded, sinuate-walled,  
on upper and lower surfaces or few scattered along and papillate cells, acroscopic margins long-ciliate  
acroscopic halves of lateral leaves (vs. idioblasts along proximal ⅔, otherwise short-ciliate to dentate  
present on both leaf surfaces or only visible distally, each cilium, stiff and 0.05–0.15 mm long,  
on leaf lower surfaces), broadly-ovate to ovate- basiscopic margins sparsely short-ciliate to dentate  
orbicular (vs. ovate-lanceolate) median leaves throughout; apices short-acuminate, each acumen  
long-ciliate throughout (vs. short-ciliate along 0.05–0.2 mm long, each tipped by 1–3, short, tooth-  
proximal ¾, otherwise serrate to serrulate along like hairs; upper surfaces glabrous, comprising  
distal ¼) margins, each cilium 0.15 or 0.2 (0.1 or rounded, irregularly, sinuate-walled, laevigate  
less) mm long, and long-aristate apices with each cells, without idioblasts or stomata; lower surfaces  
arista covered by stiff, short, permanent tooth-like comprising elongate, sinuate-walled cells, with few  
hair (vs. arista lacking stiff tooth-like hairs or this elongate and papillate idioblasts sparsely distributed  
inconspicuous, delicate, and easily abraded) on along acroscopic halves of the laminae and  
upper surfaces.  
abundantly distributed on proximal portion near the  
395  
Bol. Soc. Argent. Bot. 59 (3) 2024  
Fig. 1. Selaginella pectopuellensis Valdespino & C. López. Scanned image of holotype: Matos et al. 1360  
(NY). Digitized image courtesy of the herbarium of the University of Panama, Panama.  
396  
I. A. Valdespino et al. - Selaginella pectopuellensis from Bahia, Brazil  
Fig. 2. Selaginella pectopuellensis Valdespino & C. López. A: Stem, upper surface. B: Lateral leaf, upper  
surface. C: Close-up of lateral leaf proximal section, upper surface. D: Close-up of lateral leaf distal section,  
upper surface. E: Median leaf, upper surface. F: Close-up of median leaf proximal section, inner half, upper  
surface. G: Close-up of median leaf proximal section, outer half, upper surface. H: Close-up of median leaf,  
distal section, upper surface. Abbreviations= a: round, papillate cells; b: stomata; c: elongate and papillate  
idioblasts. Scale bars= A: 1 mm; B–H: 100 µm. (Matos et al. 1360, holotype, NY).  
397  
Bol. Soc. Argent. Bot. 59 (3) 2024  
Fig. 3. Selaginella pectopuellensis Valdespino & C. López. A: Stem, lower surface. B: Lateral leaf, lower  
surface. C: Close-up of lateral leaf proximal section, lower surface. D: Close-up of lateral leaf distal section,  
lower surface. Abbreviations= a: stomata; b: elongate and papillate idioblasts on leaf lamina; c: submarginal  
and marginal, elongate and papillate idioblasts. Scale bars= A: 1 mm; B–D: 100 µm. (Matos et al. 1360,  
holotype, NY).  
bases, each idioblast lumen covered by 1 or 2 rows arista denticulate along its margins, ½ the length  
of papillae, with stomata in 2–5 rows along midribs of the laminae, each 0.5–0.8 mm long, usually  
and on one row along basiscopic margins. Median tipped by 1–3 teeth; upper surfaces composed of  
leaves slightly imbricate, ascending, broadly ovate medial to submedial laevigate, rounded, sinuate-  
to broadly ovate-elliptic or orbicular with the inner walled cells and a submedial to submarginal band  
and outer halves equal in width, 1.0–1.6 × 0.7–1.1 of rounded, sinuate-walled, and papillate cells  
mm; bases glabrous, rounded to oblique on leaf similar to those present in submarginal portion of  
near distal portions of main stems and branches, the lateral leaf laminae upper surfaces, without  
without auricles; margins continuously bordered idioblasts, with stomata in 1–3 rows along distal  
by a broadly hyaline band comprised of idioblasts, ¾ of midribs and one row submarginally on  
the band 2–4 cells wide, the idioblasts, elongate, proximal ¼ of inner margins and 6 or 7 marginally  
straight-walled, and papillate, the papillae on one or and submarginally on proximal ¼–⅓ of outer  
two rows on each cell lumen, long-ciliate throughout margins; lower surfaces similar to lower surfaces  
or entire along proximal ¼–⅓ and long-ciliate of lateral leaves, without idioblasts or stomata.  
along distal ¾–⅔; apices keeled, long-aristate, the Axillary leaf shape, size, bases, margins (except  
398  
I. A. Valdespino et al. - Selaginella pectopuellensis from Bahia, Brazil  
Fig. 4. Selaginella pectopuellensis Valdespino & C. López. A: Megaspore, proximal face. B: Close-up of  
megaspore, proximal face. C: Close-up of megaspore, proximal face. D: Microspore, proximal face. E:  
Megaspore, distal face. F: Close-up of megaspore, distal face. G: Close-up of megaspore, distal face. H:  
Microspore, distal face. Scale bars= A, E: 100 µm; B, D: 10 µm; C, F–H: 2 µm. (Matos et al. 1360, holotype, NY).  
399  
Bol. Soc. Argent. Bot. 59 (3) 2024  
those on proximal faces, closed, the microstructure  
echinate and perforate; microsporangia on two  
dorsal rows; microspores orange, 25–40 µm  
diam., proximal faces rugulate, the microstructure  
echinulate, distal faces rugulate with baculate to  
capitate projections, the microstructure echinulate.  
Distribution and habitat. Selaginella  
pectopuellensis grows as a terrestrial plant among  
shrubby and herbaceous vegetation in montane  
forests with many epiphytes at 1000 m in Serra do  
Peito-de-Moça in southeastern Bahia, Brazil.  
Etymology. The specific epithet is a free,  
compound word derived from the Latin, pectus  
“breast” and puella “maiden,” which together refer  
to the Portuguese name of the mountain range “Serra  
do Peito-de-Moça,” where this novel species occurs.  
Fig.5.DistributionmapofSelaginellapectopuellensis  
Valdespino & C. López.  
Conservation status. Selaginella pectopuellensis  
is only known from a single collection made in  
some with both margins as acroscopic margins of the Private Natural Heritage Reserve (i.e., RPPN  
lateral leaves), apices, and leaf surfaces similar to Caminho das Pedras, Bahia). The Reserve is  
lateral leaves. Strobili terminal, single at stem and adjacent to and forms part of the inclusive Parque  
branch tips, quadrangular to slightly dorsiventrally Nacional da Serra das Lontras in the Atlantic  
flattened, 3–10 mm long. Sporophylls slightly Forest (Mata Atlântica) of Brazil, which comprises  
heteromorphic and spreading, ovate to ovate- significant remnants of mountain forest in the South  
lanceolate, dorsal sporophylls slightly shorter than of Bahia harboring a rich plant diversity that is  
ventral sporophylls, each with a strongly developed highly endangered (Amorim & Matos, 2009). Based  
keel along the midrib, the keel puberulent with short on IUCN criteria (IUCN Standards and Petitions  
teeth or hairs along distal ½, margins hyaline as in Committee, 2022), the extent of occurrence (EOO=  
2
)
median leaves, short-ciliate to dentate throughout, 0.000 km and the area of occupancy (AOO= 4.000  
2
apices long-aristate, aristae 0.3–0.5 mm long, km ) suggest this species is critically endangered  
each tipped by 1–4, short tooth-like projections; (CR). According to Araujo et al. (2009) this region  
dorsal sporophylls 1.0–1.2 × 0.6–0.8 mm, upper severely suffers from deforestation and habitat  
surfaces green with cells as in median leaves upper fragmentation. This is confirmed by more recent  
surfaces, except for the halves that overlap each vegetation loss analysis (Guedes et al., 2021) and  
ventral sporophyll where the surfaces are hyaline, vegetation images (Google, 2024).  
composed of elongate and sinuate-walled cells,  
similar to those of median leaves lower surfaces,  
lower surfaces hyaline, similar to those of the diSCuSSion  
median leaves; ventral sporophylls 1.3–1.5 × 0.6–  
0
.8 mm, upper and lower surfaces greenish-hyaline  
Selaginella pectopuellensis is notable by its long-  
to hyaline, comprised of elongate, sinuate-walled creeping stems, which are sparsely 2- or 3-branched,  
cells. Megasporangia along two ventral rows; ventral, stout rhizophores, each borne throughout  
megaspores yellow, 330–410 µm diam., proximal proximal ⅓−½ of stems, coriaceous leaves (Fig. 1),  
faces reticulate, each reticulum small in diameter imbricate and ascending median leaves, each broadly  
and closed, with a slightly developed equatorial ovate to broadly ovate-elliptic or orbicular with the  
flange, the microstructure echinate and perforate, inner and outer halves equal in width and margins  
distal faces reticulate, each reticulum wider than continuously bordered by a broadly hyaline band of  
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I. A. Valdespino et al. - Selaginella pectopuellensis from Bahia, Brazil  
idioblasts, the band 2–4 cells wide, the idioblasts, the vulnerability of these species and the need  
elongate, straight-walled, and papillate, the papillae for conservation efforts. The documentation of  
on one row on each cell lumen (Figs. 2A, E). In a new species like Selaginella pectopuellensis  
addition, the median leaf of S. pectopuellensis is is significant for understanding the ecological  
long-ciliate throughout margins or entire along complexity of these regions and the evolutionary  
proximal ¼–⅓ and long-ciliate along distal ¾–⅔ of relationships within the genus Selaginella. In this  
margins and has keeled, long-aristate apices, with work, we recognize 91 native Selaginella species  
aristae denticulate along margins, each arista ½ the and 6 introduced and naturalized taxa in Brazil,  
length of the laminae, 0.5–0.8 mm long, and usually while proposing S. kriegeriana as a conspecific with  
tipped by one to three teeth (Figs. 2E−H). Selaginella and a synonym of S. magnafornensis.  
pectopuellensis is unique by its lateral leaf upper  
surfaces having a distinct submedial to submarginal  
band of round, sinuate-walled, and papillate cells author ContributionS  
along distal ½ on acroscopic halves of the laminae,  
which is also present along the laminae length on  
basiscopic halves of the laminae (Figs. 2B−D).  
IAV: conceptualization (lead); data curation  
(lead); formal analysis (lead); funding acquisition  
The type collection of S. pectopuellensis was (lead); investigation (lead); methodology (lead);  
initially determinedasS. muscosa, whichshowssome project administration (lead); resources (lead);  
degree of variability throughout its distribution range supervision (lead); validation (lead); visualization  
(
Valdespino, 1995). Certainly, S. pectopuellensis (lead); writing – original draft (lead); writing –  
and the latter species show certain morphological review and editing (lead). CAL: data curation  
similarities, including broadly-ovate, long-aristate (equal); formal analysis (equal); investigation  
median leaves with hyaline margins. In addition, (supporting); methodology (supporting); writing  
both species have yellow megaspores on two ventral – review and editing (equal). JIC: data curation  
rows of sporophylls and orange microspores on two (supporting); formal analysis (supporting);  
dorsal rows of sporophylls, although S. muscosa methodology (supporting); resources (supporting);  
might also have white megaspores. Nevertheless, visualization (supporting); writing – review  
S. pectopuellensis is easily distinguished from and editing (supporting). JMP: data curation  
S. muscosa by the characters contrasted in the (supporting); formal analysis (supporting);  
diagnosis. Selaginella pectopuellensis is also easily investigation (supporting); resources (supporting);  
separated from the latter by its lateral leaf upper software (supporting); visualization (equal); writing  
surfaces having (vs. lacking) a distinct submedial – review and editing (supporting).  
to submarginal band of round, sinuate-walled, and  
papillate cells along distal ½ on acroscopic halves  
of the laminae, which is also present along the aCKnoWledgmentS  
laminae length on basiscopic halves of the laminae.  
It is further distinct from S. muscosa by its lateral  
We are thankful to Dr. Robbin Moran, Curator  
leaf acroscopic margins long-ciliate throughout (vs. Emeritus of ferns and lycophytes at the New York  
serrate or shortly ciliate along basal ¼ and serrulate Botanical Garden Herbarium (NY), for allowing  
distally), each cilium stiff and 0.1 or 0.2 mm (vs. the study of the holotype, arranging its loan, and  
each cilium, when present, 0.5 mm) in length, use of fragment material for further study at the  
with long-acuminate (vs. acute to short-acuminate) herbarium of the University of Panama (PMA).  
apices, each 0.1 (vs. less than 0.1) mm long.  
We are grateful to Dra. María de Stapf and Vielka  
Murillo, both at PMA, for respectively, procuring  
specimen loans and digitizing the holotype. The  
Smithsonian Tropical Research Institute (STRI)  
has also supported this research by allowing the  
ConCluSionS  
This study emphasizes the rich biodiversity use of scanning electron microscopy facilities.  
found in Brazil’s Atlantic Forest and the importance IAV is further indebted to the Dean of the Faculty  
of continued research in this area. It also highlights of Natural and Exact Sciences and Technology  
401  
Bol. Soc. Argent. Bot. 59 (3) 2024  
and the office of the Vice Chancellor for Research GUEDES, P., L. F. M. HIROTA & M. ROSA. 2021. Atlas  
and Graduate Programs of the University of  
Panama, and the National Research System  
da Mata Atlântica e MapBiomas se complementam  
no monitoramento do bioma. Fundação SOS Mata  
Atlântica-Instituto Nacional de Pesquisas Espaciais  
and MapBiomas. Available in: https://www.sosma.  
org.br/artigos/atlas-da-mata-atlantica-e-mapbiomas-  
se-complementam-no-monitoramento-do-bioma/.  
[Accessed August 17, 2021].  
(
SNI, for its initials in Spanish) of the National  
Secretariat for Science, Technology, and Innovation  
SENACYT) of Panama for supporting his research  
(
on Selaginella. CAL thanks Dr. Brian E. Sedio for  
his constant support and advice. We also appreciate  
the support provided by Pedro Argudo at NovoArt IUCN STANDARDS AND PETITIONS COMMITTEE.  
in preparing composed images and Christopher  
Gioia and Dr. Robbin Moran for proofreading an  
early draft of the manuscript. Finally, we are also  
thankful to the reviewers and the editorial team of  
the Boletín de la Sociedad Argentina de Botánica  
2022. Guidelines for Using the IUCN Red List  
Categories and Criteria. Version 15. Prepared by  
the Standards and Petitions Committee. Available  
in: https://www.iucnredlist.org/documents/  
RedListGuidelines.pdf. [Accessed March 11, 2024].  
for their suggestions to improve the manuscript and MATOS, F. B., A. M. AMORIM & P. H. LABIAK. 2010.  
successfully steering the publication process.  
The ferns and lycophytes of a montane tropical  
forest in southern Bahia, Brazil. J. Bot. Res. Inst.  
Texas 4: 333-346.  
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