A new species of Deamia (Cactaceae) from the Mesoamerican region

Abstract

Deamia montalvoae sp. nov. (Cactaceae) from the Mesoamerican region (El Salvador, Guatemala, and Mexico) is described and illustrated based on morphological and molecular evidences. The new species is morphologically characterized by stems up to 2 cm in diameter, 7−8-ribbed, flowers (23−)27−30 cm long, with pericarpel covered by bristles and trichomes, but without spines; fruit 5−6 cm long and pale-red, covered by bristles and trichomes, with white flesh; seeds 3.1−3.5 mm long, dark brown, with a smooth microrelief. The phylogenetic analysis using two introns, rpl16 and trnL-trnF, and a spacer, psbA-trnH, shows that D. testudo is the sister species of D. montalvoae and D. chontalensis.

Full text

Phytotaxa 369 (4): 251–259
http://www.mapress.com/j/pt/
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Duilio Iamonico: 16 Jun. 2018; published: 21 Sept. 2018
https://doi.org/10.11646/phytotaxa.369.4.2
251
A new species of Deamia (Cactaceae) from the Mesoamerican region
GABRIEL CERÉN1, MIGUEL ÁNGEL CRUZ2, JENNY MENJÍVAR1 & SALVADOR ARIAS2*
1 Museo de Historia Natural de El Salvador, Ministerio de Cultura, El Salvador
2 Jardín Botánico, Instituto de Biología, Universidad Nacional Autónoma de México, 04510 México D.F., México
*Corresponding author: sarias@ib.unam.mx
Abstract
Deamia montalvoae sp. nov. (Cactaceae) from the Mesoamerican region (El Salvador, Guatemala, and Mexico) is described
and illustrated based on morphological and molecular evidences. The new species is morphologically characterized by
stems up to 2 cm in diameter, 7−8-ribbed, flowers (23−)27−30 cm long, with pericarpel covered by bristles and trichomes,
but without spines; fruit 5−6 cm long and pale-red, covered by bristles and trichomes, with white flesh; seeds 3.1−3.5 mm
long, dark brown, with a smooth microrelief. The phylogenetic analysis using two introns, rpl16 and trnL-trnF, and a spacer,
psbA-trnH, shows that D. testudo is the sister species of D. montalvoae and D. chontalensis.
Keywords: Deamia, Echinocereeae, molecular phylogeny, Selenicereus, taxonomy
Introduction
Flora of Mesoamerica and adjacent regions include approximately 100 species of Cactaceae Juss., of which
approximately 60% are climbers and epiphytic taxa (see e.g., Bravo-Hollis & Arias 1999, Bauer 2003). Deamia Britton
& Rose (1920: 212) (Pachycereinae Buxb., Echinocereeae Buxb.) is a genus distributed in the Mesoamerican region,
which was recently resurrected by Korotkova et al. (2017) who showed that it is not related to Strophocactus Britton
& Rose (1913: 262) deserving to be treated as separate taxon. On the basis of the current knowledge, two species
are recognized, i.e. Deamia chontalensis (Alexander 1950: 131) Doweld (2002: 41), and D. testudo (Karwinsky ex
Zuccarini) Britton & Rose (1920: 213). Both are clambering or hemi-epiphytic pendent shrubs, with determinate
growth, wax deposits on their stems, hairs and bristles on their flowers, red fruit and clear pulp (Korotkova et al.
2017).
Deamia chontalensis, originally described as Nyctocereus chontalensis, was transferred to Selenicereus by
Kimnach (1991) due to the presence of adventitious roots and spines on its flowers. Doweld (2002) later transferred N.
chontalensis to Deamia because it shares seed micromorphology with D. testudo, and D. chontalensis this is the current
accepted name. D. chontalensis is characterized by its cylindrical and pendent stems, with 5−6(−7) acute ribs, flowers
6−9 cm long, with stigma located below the anthers (brevistyle flower), and fruits 2.0−2.5 cm in diameter, pale red, and
covered with spines, with white pulp (Alexander 1950, Bravo-Hollis 1978). Recently, Véliz (2008) reported Deamia
chontalensis in Guatemala (Huehuetenango), recognizing the same growth form as that described above, but without
observing the flower. Ishiki-Ishihara et al. (2013) reported Selenicereus grandiflorus (Linnaeus 1753: 567) Britton &
Rose (1909: 430) in Mexico (Chiapas), based on several specimens, including one collected at La Trinitaria that has a
climbing stem with eight ribs and flowers that are 28 cm long, and covered with bristles and hairs (Ochoa-Gaona 4128,
MEXU). Meanwhile, Cerén et al. (2015) recorded Strophocactus aff. chontalensis in El Salvador (Santa Ana), based
on samples collected by a team of foresters from the Montecristo National Park and the Natural History Museum of
El Salvador. The plants of this well-preserved population share the aforementioned characteristics of specimens from
Guatemala and Mexico. However, they have flowers up to 30 cm in length and stigma located above or at the same
level of the anthers, which differ from the typical D. chontalensis. All these specimens from El Salvador, Guatemala,
and Mexico were morphologically compared with Deamia chontalensis, D. testudo, and Selenicereus grandiflorus,
and molecular data were also produced (trnL-F, psbA-trnH, and rpl16).

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252 • Phytotaxa 369 (4) © 2018 Magnolia Press
Material and methods
Morphology
Field surveys were carried out in El Salvador and Mexico and from cultivated specimens in Jardín Botánico, Instituto
de Biología UNAM, and Museo de Historia Natural de El Salvador to collect morphological and phenological data.
Herbarium collections preserved at BIGU, FT, LAGU, MHES, MEXU, and MO (acronyms according to Thiers
2018-onward) were also examined. With the aim to evaluate possible differences in reproductive structures, some
flowers were preserved in formaldehyde (FAA). Seeds were studied with a scanning electron microscopy (SEM). The
geographical distribution map was made using ArcGis V.10.0 software (Esri, New York, USA).
DNA extraction, PCR amplification, and sequencing
The phylogenetic analysis was carried out with three molecular chloroplast markers: two introns, rpl16 and trnL-trnF,
and a spacer, psbA-trnH (Table 1). The sequences were derived from Cruz et al. (2016) for Hylocereeae and Tapia et al.
(2017) for Pachycereinae. Further markers were sequenced for samples of Deamia chontalensis and Deamia sp. nov.
(Table 1). Stem tissue samples were dried with silica gel and stored at -20 °C. Total DNA was extracted using the EZ-
10 Spin Column Plant Genomic DNA Miniprep kit (Bio Basic Inc., Ontario, Canada). The intron rpl16 was amplified
with the primers rpl161F and rpl163R (Hernández-Hernández et al. 2011), the initial denaturation of the DNA was 5
min at 94 °C, followed by 26 cycles of 1 min at 94 °C, 50 s at 55 °C and 2 min at 72 ºC, and a final extension of 4 min
at 72 ºC. The trnL-trnF region was amplified using the “C” and “F” primers designed by Taberlet et al. (1991). The
PCR programme used for this region consisted of 2 min at 94 ºC, 29 cycles of 30 s at 94 ºC, 30 s at 52 ºC and 1 min
at 72 ºC, and a final extension of 7 min at 72 ºC. The psbA-trnH spacer was amplified with the primers psbA (Sang
et al. 1997) and trnH (Tate and Simpson 2003), with an initial DNA denaturation for 2 min at 94 °C, followed by 29
cycles of 30 s at 94 ºC, 30 s at 52 ºC and 1 min at 72 ºC, with a final extension of 7 min at 72 ºC. The unpurified PCR
products were sequenced at the University of Washington (UW-High Throughput Genomics Center, http://www.htseq.
org/). The regions were sequenced with the amplification primers. The sequences were assembled and edited with
Sequencher 4.8 (GeneCode).
The sequences were aligned using MAFFT (Katoh et al. 2002), and then a manual inspection was performed.
The highly variable regions, defined here as hotspots, were eliminated. The insertion and deletion events (indels) were
coded following the simple Indel coding method proposed by Simmons and Ochoterena (2000), with the Indel Coder
option in SeqState v.1.40 (Müller 2005). The aligned markers and the encoded indels were concatenated using the
Mesquite program (Maddison & Maddison 2017) to generate a single molecular data matrix.
TABLE 1. GenBank accession numbers of the sequences used in this study.
Name Voucher rpl16 trnL-trnF psbA-trnH
Acanthocereus chiapensis MX: Chiapas, Guzmán 999, MEXU KU598005 KU598057 KU597952
A. tetragonus MX: Chiapas, Guzmán 1002, MEXU KU598021 KU598074 KU597969
Aporocactus flagelliformis MX: Veracruz, Cruz 01, MEXU KU597980 KU598032 KU597927
Bergerocactus emoryi MX: Baja Cal. Arias 1307, CHAPA DQ099994 DQ099925 KF783478
Carnegiea gigantea USA: Arizona, NYGB N.A. AY181591.1 AY181619.2 KY624692
Cephalocereus euphorbioides MX: Veracruz, Tapia 01, MEXU KY624637 KY624651 KY624666
C. nudus MX: Oaxaca, Tapia 11, MEXU KY624643 KY624657 KY624678
C. sanchezmejoradae MX: Oaxaca, Tapia 39, MEXU KY624640 KY624654 KY624674
C. totolapensis MX: Oaxaca, Terrazas 628, CHAPA KY624647 KY624661 KY624688
Deamia chontalensis MX: Oaxaca, Yañez 03, MEXU MH107788 MH107803 MH107793
D. chontalensis MX: Oaxaca, Yañez 04, MEXU MH107789 MH107804 -
D. chontalensis MX: Oaxaca, Yañez 05, MEXU MH107790 MH107805 -
D. montalvoae MX:Chiapas, Arias 1640, MEXU MH107791 MH107806 MH107794
D. montalvoae GT: Huehuetenango, Veliz 20416, BIGU MH107792 MH107807 MH107795
D. testudo MX: Oaxaca, Yañez 01, MEXU KY624648 KY624662 KY624731
D. testudo GT: Izabal, Veliz 19628, BIGU KY624649 KY624663 KY624696
Disocactus biformis GT: Sacatepéquez, Veliz, 19901, BIGU KU598016 KU598069 KU597964
Echinocereus pentalophus MX: Querétaro, Arias 1737, MEXU KF783558 KF783628 KF783509
Epiphyllum phyllanthus SR: Hammel 22377, INB KU598015 KU598068 KU597963
Lemaireocereus hollianus MX: Puebla, Arias 1373, MEXU KU598012 KU598064 KU597959
Lophocereus schottii MX: Baja Cal. Sur, Terrazas 474, CHAPA AY181613 AY181620 MH107796
.....continued on the next page

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TABLE 1. (Continued)
Name Voucher rpl16 trnL-trnF psbA-trnH
Marshallocereus aragonii
eichlamii
MX: Yucatán, Arias 1363, MEXU AY181610 AY181629 KY624690
Morangaya pensilis MX: Baja Cal., Arias 1295, MEXU KF783609 KF783679 KF783538
Myrtillocactus geometrizans MX: Querétaro, Terrazas 557, CHAPA DQ100012 DQ099943 KF783546
Nyctocereus serpentinus MX: Michoacán, Sánchez-Mejorada 4249, MEXU DQ100028 DQ099959 MH107797
Pachycereus grandis MX: Puebla, DFE 31. MEXU AY181605 AY181646 MH107798
P. tepamo MX: Michoacán, Gama 107,MEXU AY181593 AY181647 MH107799
Peniocereus marianus MX: Sinaloa, Guzmán 1038 MEXU DQ100024 DQ099955 MH107800
P. striatus MX: Sonora, McGill S/N DQ100031 DQ099962 MH107801
P. viperinus MX: Puebla, Gama 37, MEXU DQ100033 DQ099964 MH107802
Pereskia lychnidiflora MX: Oaxaca, Arias 1153, MEXU KU598024 KU598077 KU597972
Pseudorhipsalis himantoclada CR: San Jose, Hammel 22076, INB KU597998 KU598050 KU597945
Selenicereus glaber MX: Chiapas,Bravo 5614, MEXU KU598031 KU598084 KU597979
S. grandiflorus MX: Veracruz, Guzmán 1365, MEXU KU598023 KU598076 KU597971
S. ocamponis MX: Guerrero, Gama 104, MEXU KU598027 KU598080 KU597975
Strophocactus wittii Hannover, Herrenhäuser Gärten B05X138, (cult.) KY624650 KY624664 KY624697
Phylogenetic analysis
PAUP* v. 4.0b10 (Swofford 2002) was used for the maximum parsimony (MP) analysis. All the characters were
disordered, with equal weights. A heuristic search with the addition of simple sequences and 15,000 replicas was
used; the branch exchange algorithm used was TBR (tree-bisection-reconnection). For the support of branches, 7000
bootstrap (BS) replicas were used. The Bayesian inference (BI) analysis was done with MrBayes 3.2 (Huelsenbeck
& Ronquist 2001) with the partitioned data. For each partition, we used the model designated by jModeltest (Darriba
et al. 2012); the partitions of trnL-F and psbA-trnH used the substitution model TPM1 µf + G, the partition of rpl16
used the substitution model TVM + I + G, and the partition of indels used the model of restriction sites (binary). Four
independent runs were carried out with 4 chains and six million generations each, with a sampling of the chains every
1000 generations. The convergence of the chains was observed using Tracer (Rambaut & Drummond 2009). A total
of 20% of the first trees were discarded as burn-in. The remaining trees were used to build the majority consensus tree
at 50%.
Results and discussion
Deamia montalvoae Cerén, J. Menjívar et S. Arias sp. nov. (Figures 2, 3, and 4)
Type:—EL SALVADOR. Santa Ana: Mpio. Metapán, Parque Nacional Montecristo, 14º23′17′′N, 89º22′04′′W, 1590 m, 26 March 2014,
Menjívar, Cerén & Magaña 3219 (Holotype MHES15-6619!, isotype MEXU!).
Diagnosis:—Deamia montalvoae is similar to D. chontalensis and D. testudo. It differ from D. chontalensis by the
higher number of ribs (7−8 vs. 5−7 in D. chontalensis) and spines (7−13 vs. 1−7) stem and longer flowers (23−30
cm vs. 6−9 cm), stigmas located above or at the same level of the anthers, and larger fruit (5−6 × 4−4.5 cm vs. 2−3 ×
2−2.5 cm) completely covered by bristles and trichomes. D. montalvoae can be easily distinguished from D. testudo by
the smaller diameter of its stem (1−2 cm vs. 3−10 cm) and less variable stem diameter; in addition, it does not adhere
strongly to its support, has a greater number of ribs (7−8 vs. 6−9), and has a dense layer of bristles and trichomes
covering its fruit.
Description:—Climbing or hemiepiphytic shrubs on rocks or trees. Stems numerous, cylindrical, and prolifically
branching; new stems reddish or green-reddish, older stems lustrous green, pendent or sprawling, up to 1.5 m × 1−2
cm, with adventitious roots not strongly adhered to support, ribs 7−8, 1−1.5 cm high, acute margin; areolae 0.2−0.3
cm in diameter, 0.5−2.0 cm apart, spines 7−13, 0.4−2.6 cm long, some spines renewed, yellowish with reddish apex
becoming grey to black when old; bristles 0.5−1.5 cm long, abundant, yellowish to whitish at maturity, and persistent.
Flowers solitary, emerging from subapical areoles, (23−)27−30 cm long, infundibuliform; pericarpel 2.0−2.2 ×
1.5−1.8 cm, with 8−10 series of podaria, bracts 1.0−1.5 × 0.5 mm, deltoids, green, fully covered by bristles and hairs
(0.5−3.0 cm) long, golden brown, translucent; receptacular tube 9−15 cm, ca. 0.8 cm wide at the base of the tube,
1.0−1.2 cm wide in the throat, green with brown tone, reddish after anthesis, decurrent podaria, slightly conspicuous,

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FIGURE 1. Phylogenetic position of Deamia montalvoae based on three chloroplast markers. A. Maximum parsimony. B. Bayesian
inference. Bootstrap and posterior probabilities are indicated at the nodes.
bracts 2.5−13 × 1−2 mm, green with reddish apex, bristles and hairs 0.3−3.2 cm, brown to white-pale; perianth 11−12
cm long, outer tepals 5.0−10.5 × 0.4−1. cm, linear lanceolate, yellow-brown with apices acuminate, inner tepals
8.9−10.0 × 0.6−1.1 cm, linear oblong, white with apices acuminate; nectarial chamber in the lower part of tube,
4.0−5.5 cm; stamens 5.5−8.5 cm, white, emerging in the upper third of the tube, attached in one series, in an area of
2.5−3 cm long, anthers 3 × 1 mm, light yellow; style 21.7−21.9 × 0.2−0.3 cm, yellow-green, the stigma lobes ca. 15,
0.7−1.0 cm, light yellow, the stigmas located above (up to 1 cm) or at the same level of the anthers. Fruits 5−6 × 4−4.5
cm, globose to ovoid, pale red, oxidize when cut, without evident bracts; areolae ca. 1.5 mm in diameter, fruit fully
covered by bristles and hairs, up to 50 per areolae, 4.5−30 mm, brownish with whitish apex, persistent areolae with
ripening fruit, pulp white, fragrant and sweet; seeds 3.1−3.5 × 2.1−2.5 mm, ovate, dark brown, glossy, with flat cell
and smooth microrelief.
Etymology:—The specific epithet is in honour of Edy Albertina Montalvo for her legacy in the study of plants as
the first Salvadoran woman dedicated to botany.
Phenology:—Flowering time March−April; fruiting time April−August.
Habitat:—Deamia montalvoae occurs in tropical deciduous forests and in transitional areas with cloud forests.
It grows in cliffs, by climbing on different species of trees, including Sideroxylon tepicense (Standley 1924: 1115)
Pennington (1952: 159) (tempisque) and Ficus spp. (amate), and around rivers and creeks, from 1200−600 m a.s.l.
Distribution:—Deamia montalvoae occurs in El Salvador (Santa Ana), Guatemala (Huehuetenango), and
Mexico (Chiapas), in some foothills of the Central American Range (Sierra Madre del Sur, Mexico, and Guatemala to
Alotepeque-Metapán between Honduras and El Salvador).
Conservation status:—According to the IUCN Red List categories and criteria (2017), and the criterion B1, a,
Deamia montalvoae is assessed as endangered (EN). The species occurs in three know and isolated populations (Santa
Ana, El Salvador; Huehuetenango, Guatemala; Chiapas, Mexico), in an area smaller than 5,000 km2. In Santa Ana it is
located in a protected natural area.

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FIGURE 2. Main characteristics of Deamia montalvoae Cerén, J. Menjívar & S. Arias sp. nov. A. Hemiepiphytic plant in its habitat. B.
Climbing plant on a tree. C–D. Juvenile terminal stems showing spines and bristles.
Taxonomic notes:—Deamia montalvoae shares the following characters with D. chontalensis, and D. testudo:
climbing habitus and stems with acute margin ribs. The number of ribs (Table 2), wax deposits on stems (Figure 2B,
D), flower form, bristles and hairs, and the fruit form (Table 2), differs between them. Several characters of the stem
are shared exclusively between D. montalvoae and D. chontalensis, such as the pendent terminal stems, stem diameter,
height of the ribs and size of the spines. Therefore, we consider it possible for sterile specimens to cause confusion and
mistaken identification with D. chontalensis (see e.g., Véliz 2008, Cerén et al. 2015) and Selenicereus grandiflorus
(e.g., Ishiki-Ishihara et al. 2013; Table 2). Other evaluated characters can help to identify the species, such as number
of ribs, flower size and position of the stigmas according to the anthers, the size of the fruit and the presence of bristles
and hairs, as well as the size, color and microrelief of the seed (Table 2).

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FIGURE 3. Reproductive structures of Deamia montalvoae. A. Flower in longitudinal section, showing inner and outer sections. B.
Flowers in post-anthesis. C. Stem and fruits. D. Fruit in longitudinal section, showing seeds with pulp.

A NEW SPECIES OF DEAMIA Phytotaxa 369 (4) © 2018 Magnolia Press • 257
FIGURE 4. SEM micrographs of Deamia montalvoae seeds. A. Lateral view. B. Hilum-micropylar region. C. Close-up of lateral view.
FIGURE 5. Distribution of Deamia montalvoae in Mexico, Guatemala, and El Salvador.
The strict consensus tree obtained from the MP and BI tree supports the relationship of the two terminals of
Deamia montalvoae with D. chontalensis (78% BS / 1.0 PP). These two species are resolved as sisters of D. testudo,
with strong support (100% BS / 1.0 PP). This analysis also corroborates the monophyly of Deamia, after including the
new species, and the relationship within the Pachycereinae subtribe, albeit with low support (66% BS / 0.99 PP; Figure
1). Recent phylogenetic analyses offer new interpretations on the diversification of the climbing, hemi-epiphytic and
epiphytic Cactaceae. There is evidence of at least five climbing or epiphyte lineages, possibly shrubs, that diversified
independently (Korotkova et al., 2010, 2017, Tapia et al., 2017). In Mesoamerica, most of the Hylocereeae genera and
one Echinocereeae genus (Deamia) have diversified into climbing, hemi-epiphytic or epiphytic growth forms (Cruz
et al., 2016, Korotkova et al., 2017, Tapia et al., 2017); therefore, it is of great interest to know the processes and
structures involved in their adaptation.
Other specimens examined:—EL SALVADOR. Santa Ana: Metapán, Cantón El Limo, La Cuevona, 15 October
2013, Cerén et al. 3199 (LAGU, MHES); Parque Nacional Montecristo, Las Huertas, 29 April 2014, Cerén & Menjívar
3312 (MHES); Las Huertas, 08 July 2014, Cerén 3329 (LAGU, MHES, MEXU, MO); Las Huertas, 26 March 2014,
Menjívar et al. 3095 (MHES, MEXU); Las Huertas, 13 April 2013, Menjívar et al. 2866 (LAGU, MHES, MEXU,
FT). GUATEMALA. Huehuetenango: Nentón, carretera a Gracias de Dios, 16 April 2008, Véliz & Velásquez 20144
(BIGU); Nentón, 17 July 2008, Véliz & Velásquez 20285, 20286, 20292 (BIGU); Nentón, carretera a Gracias, 16
September 2008, Véliz & Velásquez 20416, 20417 (BIGU, MEXU); Nentón, 07 February 2011, Véliz & Velásquez
22569 (BIGU). MÉXICO. Chiapas: La Trinitaria, 11 May 1993, Ochoa-Gaona 4128 (MEXU); La Trinitaria, 20
January 2007, Arias & Cruz 1640 (MEXU).

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TABLE 2. Morphological comparison of Deamia montalvoae, and other species of Deamia and Selenicereus grandiflorus.
Deamia montalvoae Deamia chontalensis Deamia testudo Selenicereus grandiflorus
Growth
form
Climbing or hemiepiphytic
plants
Climbing plants Climbing or hemiepiphytic
plants
Climbing or hemiepiphytic
plants
Stem 1–2 cm diameter, 7–8 ribs,
margin acute, pendent or
sprawling with adventitious
roots to support
2.5–3.5 cm diameter,
5–6(–7) ribs, margin acute,
pendent or sprawling with
adventitious roots to support
3–10 cm diameter, 3–5(–6)
ribs, margin acute, usually
adhering to support
(trunks or rocks) by tough
adventitious roots
1.2–3 cm diameter, 6–8(–9)
ribs, margin rounded,
pendent or sprawling with
adventitious roots to support
Stem spines
and bristles
Spines 7–13 per areole, up
to 2.6 cm long; bristles up
to 1.5 cm long, abundant,
usually persistent
Spines 1–7 per areole, up
to 2 cm long; bristles up to
1.5 cm long, scarce, usually
persistent
Spines 6–12 per areole, up
to 2 cm long; bristles up to 1
cm long, scarce, deciduous
5–7(–11) spines per areole,
up to 1.2 cm long; bristles
up to 0.8 cm long, scarce,
deciduous
Flower (23–) 27–30 cm long,
pericarpel without spines,
covered by golden brown
bristles and hairs (up to 3 cm
long); stigmas located above
or at the same level of the
anthers
6–9 cm long, pericarpel
almost covered by whitish
or golden brown spines and
bristles (up to 1.2 cm long);
stigma located below the
anthers
25–28 cm long, pericarpel
covered or not by whitish
or golden brown spines and
bristles (up to 1.5 cm long);
stigmas located above or at
the same level of the anthers
18–24(–30) cm long,
pericarpel not covered
by whitish or brownish-
yellowish spines and bristles
(up to 2.2 cm long); stigmas
located above or at the same
level of the anthers
Fruit 5–6 × 4–4.5 cm, globose to
ovoid, pale red, covered by
bristles and hairs; pulp white
2–3 × 2–2.5 cm, globose,
pale red, almost covered
by spines and bristles; pulp
white
5–9 × 3–3.5 cm, globose,
pale red, not covered by
spines and bristles; pulp
white
5–9 × 5–7 cm, globose to
ovoid, red, not covered by
spines and bristles; pulp
white
Seed 3.1–3.5 × 2.1—2.5 mm,
dark brown, glossy, smooth
microrelief
3.6–3.7 × ca. 2.1 mm, light
brown, semi-matt, smooth
microrelief
2.6–2.8 × 1.5–1.6, dark
brown, glossy, verrucose
microrelief
Ca. 2.4 ×1.2 mm, brown,
semi-matt, smooth
microrelief
Acknowledgements
Thanks to M. Guido, director of Montecristo National Park, for facilitating the necessary field support. We thank F.
Magaña for monitoring the species in situ. We also thank L. Cabrera (Instituto de Biología, UNAM) for her technical
support in the molecular laboratory, B. Mendoza, and Y. Morales (Instituto de Biología, UNAM) for her technical
support in SEM. Partial funding was provided by DGAPA, PAPIIT (IN208315 to S. A.).
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