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Additions to the ora of Panama, with comments on plant
collections and information gaps
Orlando O. Ortiz1, Rodolfo Flores2, Gordon McPherson3, Juan F. Carrión4,
Ernesto Campos-Pineda5, Riccardo M. Baldini6
1 Herbario PMA, Universidad de Panamá, Vía Simón Bolívar, Panama City, Panama Province, Estafeta Universitaria, Panama. 2 Programa de
Maestría en Biología Vegetal, Universidad Autónoma de Chiriquí, El Cabrero, David City, Chiriquí Province, Panama. 3 Herbarium, Missouri
Botanical Garden, 4500 Shaw Boulevard, St. Louis, Missouri, MO 63166-0299, USA. 4 Programa de Pós-Graduação em Botânica, Universidade
Estadual de Feira de Santana, Avenida Transnordestina s/n, Novo Horizonte, 44036-900, Feira de Santana, Bahia, Brazil. 5 Smithsonian Tropical
Research Institute, Luis Clement Avenue (Ancón, Tupper 401), Panama City, Panama Province, Panama. 6 Centro Studi Erbario Tropicale (FT
herbarium) and Dipartimento di Biologia, Università di Firenze, Via La Pira 4, 50121, Firenze, Italy.
Corresponding author: Orlando O. Ortiz, ortizopma@gmail.com.
Abstract
In the present study, we report 46 new records of vascular plants species from Panama. The species belong to the fol-
lowing families: Anacardiaceae, Apocynaceae, Aquifoliaceae, Araceae, Bignoniaceae, Burseraceae, Caryocaraceae,
Celastraceae, Chrysobalanaceae, Cucurbitaceae, Erythroxylaceae, Euphorbiaceae, Fabaceae, Gentianaceae, Laciste-
mataceae, Lauraceae, Malpighiaceae, Malvaceae, Marattiaceae, Melastomataceae, Moraceae, Myrtaceae, Ochnaceae,
Orchidaceae, Passioraceae, Peraceae, Poaceae, Portulacaceae, Ranunculaceae, Salicaceae, Sapindaceae, Sapotaceae,
Solanaceae, and Violaceae. Additionally, the status of plant collections in Panama is discussed; we focused on the
areas where we identied signicant information gaps regarding real assessments of plant biodiversity in the country.
Keywords
Central America, oristics, new records, taxonomy, vascular plants.
Academic editor: Rosa del C. Ortiz | Received 30 December 2018 | Accepted 9 July 2019 | Published 2 August 2019
Citation: Ortiz OO, Flores R, McPherson G, Carrión JF, Campos-Pineda E, Baldini RM (2019) Additions to the ora of Panama, with comments
on plant collections and information gaps. Check List 15 (4): 601–627. https://doi.org/10.15560/15.4.601
Introduction
The Republic of Panama is located in the intertropical
zone, between 07°12′08″ and 09°38′46″ north latitude
and 077°09′24″ and 083°03′07″ west longitude. Panama
is sandwiched between the Caribbean Sea to the north,
the Pacic Ocean to the south, the Republic of Costa
Rica on the west and the Republic of Colombia on the
east. Panama has a land area of 75,845 km2, the fourth
largest among the seven countries of Central Amer-
ica, representing 14.7% of the entire Central American
region (ANAM 2010). The territorial organization of the
country includes provinces, which are divided into dis-
tricts and these in turn, are subdivided in corregimien-
tos. Currently, the territorial organization comprises
10 provinces, 79 districts, 664 corregimientos and ve
indigenous comarcas. In 2014, the province of Pan-
amá Oeste was created, which was segregated from the
province of Panamá and comprises the districts that are
located to the west of the Panama Canal.
The Isthmus of Panama is a land connection between
North America and South America, as an oceanic barrier
Check List 15 (4): 601–627
https://doi.org/10.15560/15.4.601
4
15
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NOTES ON GEOGRAPHIC DISTRIBUTION
602 Check List 15 (4)
for the Caribbean Sea and the Pacic Ocean. Its forma-
tion stands as one of the greatest natural events, as it trig-
gered an interchange of previously isolated terrestrial
organisms and opened each continent to invasions by
northbound and southbound migrant species, allowing
for the Great American Biotic Interchange (Webb 2006;
Cione et al. 2015). Although the exact age of formation
of the Isthmus is still hotly debated by several scholars
(e.g. Coates and Stallard 2013; Bacon et al. 2015; Mon-
tes et al. 2015; O’Dea et al. 2016; Jaramillo et al. 2017;
Molnar 2017), it appears that the nal closure occurred
no later than 2.8 Ma (Coates and Stallard 2013; O’Dea et
al. 2016).
Panama has a warm, wet, tropical climate with high
temperatures throughout the year, with an average of
27 °C and two seasons: wet and dry (in certain regions
microclimates generated by nearby mountains or bod-
ies of water are exceptions). The wet period is longer
in duration, usually beginning in mid- to late-April
and ending the rst half of December. The shorter dry
season occurs on average from mid-December to mid-
April. On the Caribbean slope, annual rainfall amounts
to 3500 mm, while on the Pacic coast, to approximately
2300 mm; however, some regions of Darién Province
can reach 7000 mm of annual rainfall (ANAM 2010).
Approximately 70% of the Panamanian territory is
formed by lowlands and hills with less than 700 m eleva-
tion. The main highlands are formed by the Cordillera
Central (Cordillera de Talamanca in Panamanian ter-
ritory), a mountain range that extends from the border
with Costa Rica to the center of Panama, with elevations
gradually decreasing from west to east and includes the
highest peak of the Isthmus (Volcán Barú), which attains
3474 m (Correa et al. 2004).
The rst documents that included information on the
plants of Panama were produced during the period of
the Spanish conquest, which mainly reported the uses of
local species (Escobar 1971). These publications include
the volumes of Fernández de Oviedo in the “Historia
General y Natural de las Indias” (published between
1535 and 1547) dealing with the plants, animals, and
people of the Isthmus (Velásquez-Runk 2015). Through-
out the 17th century, although Spain tried to exclude for-
eigners from its colonies, European knowledge about the
Isth mus advanced mostly through the work of non-Span-
ish explorers (Seemann 1852–1857); i.e. English bucca-
neers William Dampier and Lionel Wafer documented
some aspects of the geography and riches of the Isthmus
(Velásquez-Runk 2015). Wafer and Dampier, in their
works originally published in 1697 and 1699 respectively,
discussed several aspects about the climate, important
localities of the Isthmus, the wildlife, and the use of local
plants (Wafer 1699; Shipman 1962; Neill 2000).
The formal study of the ora of Panama began in the
early 18th century through collections made mainly by
European naturalists (Dwyer 1985). The rst botanical
collections made in Panamanian territory were done by
Scottish botanist James Wallace around the year 1700
(Croat 1978; Dwyer 1968a, 1985). Wallace (1701), in his
work titled “A Part of a Journal kept from Scotland to
New Caledonia in Darien, with a short Account of that
Country” mentioned some aspects of the fauna and ora
of the Darién Province, emphasizing the lack of knowl-
edge of the local ora and the morphological complexity
of the plants. During the expedition led by Alessandro
Malaspina between 1789 and 1794, Luis Née and Thad-
deus Haenke collected numerous specimens from Pan-
ama (Cerro Ancón, the Bay of Panama, and Taboga
Island) (Seemann 1852–1857; Dwyer 1964; Escobar
1971; Garmendia-Muñoz 1992). Subsequently, the Pana-
manian material collected by Haenke was used for the
description of numerous new species in the work “Rel-
iquiae Haenkeanae” published by Presl (1825–1830).
In the 19th century, during the Swedish scientic
expedition of Michaelson carried out in the years 1825
and 1826, the botanist J. Billberg made numerous col-
lections in the Province of Colon, in the area of Porto-
belo (Croat 1978). Years later, Beurling (1854) published
the work “Primitiae orae Portobellensis” possibly the
rst oristic treatment for Panama. As a result of the
rst authorized expedition to study the possible routes
of the interoceanic canal, Lloyd (1831) published the
work “Notes respecting the Isthmus of Panama”, where
he mentioned some uses of Panama’s plants, mainly tim-
ber tree species. Another important study of this cen-
tury is that of Bentham (1844), in which he described a
signicant number of new species from Panama, using
the material collected during the expeditions carried out
by HMS Sulphur in the years 1836–1842. A few years
later, as a result of the expedition of the HMS Herald
(1852–1857), the German naturalist Berthold Seemann
published his magnicent work “Botany of the Voyage
of HMS Herald”. In his work, Seemann (1852–1857),
devoted an entire chapter to the ora of the Isthmus of
Panama, including an annotated list of 1204 species (as
well as nomenclatural aspects) and described numer-
ous new species from Panama. In addition, the same
chapter added maps, historical notes, ethnobotanical
aspects, and information on the vegetation of the Isth-
mus (Blake and Atwood 1942). Other 19th-century col-
lectors included H. Cuming, R. Hinds, G. Barclay, A.
Sinclair, J. von Warscewicz, A. Fendler, M. Halsted, E.
Duchassaing, and S. Hayes (Dwyer 1964; Croat 1978).
Although, the formal study of the ora of Panama
began in the early 18th century, most of the contributions
were made during the 20th century, mainly through the
Missouri Botanical Garden (MBG) (Escobar 1971). In
the early 1920 s, MBG entere d the eld in Panama, beg i n-
ning a collection program in association with the Canal
Zone Experiment Gardens at Summit (Frodin 2001).
The Garden founded a tropical station in the Canal Zone
in 1926 (Dressler 1972) and began a series of botanical
expeditions in 1934 (Dwyer 1964; Lewis 1968; Croat
1978). Botanical activity leading to a Flora of Panama
began with the eorts of R.E. Woodson, Jr with R.W.
Schery, R.J. Seibert, J.A. Steyermark, P.C. Allen, A.A.
Ortiz et al. | Additions to the ora of Panama 603
Hunter, and Carol Dodge (Croat 1978). Later, Robert
E. Woodson, Jr published seven “Contributions toward
a Flora of Panama” (from 1937 to 1943) using material
collected between 1934 and 1941 (Robyns 1965; Lewis
1968). After these preliminary contributions, the Flora
of Panama Project itself was commenced in 1943 (Lewis
1968), culminating with the publication of the Check-
list by William G. D’Arcy in 1987. The Flora of Panama
project was the rst major overseas oristic project of
the MBG and remains nearly the most complete oristic
work from Panama (Escobar 1971). In the Checklist pre-
pared by D’Arcy (1987), 7345 species of vascular plants
in 195 families are listed. While the recent “Catálogo de
las plantas vasculares de Panamá” (Correa et al. 2004)
reports 9520 species of vascular plants, including 1144
endemic species, and showcases the richness of the Pan-
amanian ora, it also highlighted the incompleteness of
our knowledge. Currently, the ora of Panama consists
of about 10,400 species of vascular plants, including
nearly 1500 endemic species (PMA Herbarium unpub-
lished data).
After completion of the checklist by Correa et al.
(2004) and through the joint eorts of PMA and other
institutions such as the herbaria of the University of Flor-
ence (Centro Studi Erbario Tropicale, herbarium FT),
MBG, the Natural History Museum of London (BM), the
Biodiversity Institute of Costa Rica (INBio), the Smith-
sonian Tropical Research Institute (STRI), the Ministry
of Environment of Panama (MiAmbiente), International
Cooperative Biodiversity Group (ICGB), and individu-
als, new records and species were added to the Flora of
Panama. These eorts centered around dierent regions
of the country. For example: a) the Panamanian Carib-
bean slopes (e.g., Baldini and Ortiz 2014, 2015; Ortiz and
Croat 2015a, 2015b, 2016, 2017a; Ortiz et al. 2016, 2018a);
b) the district of Donoso in the Province of Colón in con-
nection with the Minera Panamá Project (e.g., Clark and
Mora 2014; Daly 2014; Daniel and McPherson 2014;
Holst 2014; Kennedy 2014; Idárraga et al. 2015; Schatz et
al. 2015; Barrie et al. 2016; Croat and Ortiz 2016; Grayum
and De Gracia 2016; Kawasaki et al. 2016; Croat et al.
2017; Batista and Mori 2017; De Gracia et al. 2017; Alm-
eda and Penneys 2018; Prance 2018); and c) La Amistad
International Park located in the Cordillera of Talamanca
between Costa Rica and Panama’s border (e.g., Soto and
Monro 2008; Monro 2009, 2012; Rodríguez et al. 2011;
Santamaría et al. 2014, 2015; Monro et al. 2017; Ortiz
and Croat 2017a, 2017b; Ortiz et al. 2018b; Rodríguez
and Solano-Peralta 2018). In addition, the Autonomous
University of Chiriquí herbarium (UCH) and the Jardín
Botánico Lankester of Costa Rica collected and thor-
oughly studied the orchid ora of Panama (Bogarín et
al. 2013a, 2013b, 2014a, 2014b, 2017; Serracín et al. 2013,
2016). For its part, the ICGB project has car ried out eld-
work all over Panama and has found several new species
and new records published and unpublished (in process)
for the country (Cáceres-González and Ibáñez 2014;
Daniel and Carrión 2015; Kennedy and Flores 2015; Flores
et al. 2016, 2017, 2018; Maas et al. 2019).
Although many new records have been found and
many new species have been described from Panama in
recent years, there are still many gaps in assessing the
diversity of the local ora. In this work and as a contri-
bution to the eorts still needed to better document Pan-
ama’s ora, we reported 46 species of vascular plants,
which represents new reports for the country, almost all
of which were found in areas with substantial informa-
tion gaps. Additionally, we discuss the main areas where
these gaps of information on plant collections in Panama
are most prevalent.
Methods
Newly documented species records were conrmed by
comparing collected voucher specimens with those iden-
tied and housed at Missouri Botanical Garden Her-
barium (MO) and University of Panama Herbarium
(PMA) (herbaria acronyms according to Thiers 2018).
In addition, images of type specimens of each species
here included were examined by consulting the JSTOR
Global Plants database (JSTOR 2018). Data on species
distribution were obtained from Almeda (2007), Balick
et al. (2000), Cornejo et al. (2012), Daly (1993), Danin et
al. (1978), Davidse (1978), Davidse et al. (2015), Funk
et al. (2007), García-Mendoza and Meave (2011), Gold-
enberg et al. (2013), González-Ramírez (2007), Idárraga
et al. (2011), Jerey (1978), Jørgensen and León-Yánez
(1999), Jørgensen et al. (2014), Lombardi (2014), Morales
(2007, 2014, 2015), Penneys and Judd (2013), Plowman
and Barrie (2010), Sánchez-Ken (2017), Secco (2004),
TROPICOS (2019a), Wang (2004), Zamora (2010) and
Zuloaga et al. (2003). Distribution maps for all new spe-
cies records are presented in Figures 3–5.
To establish areas with plant information gaps, we
used the collection databases of PMA and MO for vas-
cular plant specimens collected in Panama (considering
only specimens that have coordinates available); these
totaled 123,521 records. These records were related with
a layer grid, where each cell measured 10 km2, and only
cells with 50 records or more were mapped. All this
information was processed using ArcGIS 10.4 for Desk-
top to generate Figure 6, to nd where regions without
information remain.
Results
New Records to the Panamanian Flora
ANACARDIACEAE
Ochoterenaea colombiana F.A. Barkley, Bull. Torrey
Bot. Club 69: 442 (Barkley 1942).
Figure 2A
New records. Coclé. La Pintada, vertiente Atlántica;
08°36′ N, 080°27′ W; 600 m; 12 Nov. 2007; C. Guerra
et al. 1147 (PMA). Colón. Área del proyecto minero de
604 Check List 15 (4)
Figure 1. Selected new records for Panama. A. Amphilophium chocoense (McPherson 20358). B. Rhodospatha monsalveae (Espinosa 6014). C.
Anthodiscus choco ensis (Hammel 26289). D. Ery throxylum mbriatum (Vergara-Pérez 576). E. Hasseltia lateriora (Flores 574). F. Tachia parviora
(Araúz 1428). G. Portulaca papillato-stellulata (seed) (Ortiz & Baldini 1129). H. Cuatresia trianae (McPherson 21565). Photo credits: Cynthia Lane
(A), Alex Espinosa (B), Barr y Hammel (C), Irving Vergara- Pérez (D), Rodolfo Flores (E), Jan Meerman (F), Riccardo Baldini (G), Christel Ramos (H).
Ortiz et al. | Additions to the ora of Panama 605
Figure 2. Selected new records for Panama. A. Ochoterenaea colombiana (Araúz & De Gracia 2024). B. Miconia paleacea (Ortiz 1214). C.
Mabea tenorioi (McPherson 21574). D. Calypt ranthes bracteata (Espinosa 6006). E. Byrsonima garcibarrigae (Ortiz 1420). F. Sobralia purpurella
(Ortiz 1243). G. Paypayrola confertiora (Hammel 6313). H. Passiora macrophylla (Ortiz 2907). Photo credits: Barry Hammel (A, G), Orlando
Ortiz (B, E, F, H), Betzabeth Henríquez (C), Alex Espinosa (D).
606 Check List 15 (4)
Figure 3. Distribution map with new record specimens (circles). A. Ochoterenaea colombiana, Macropharynx renteriae, Ilex skutchii,
Rhodospatha monsalveae and Syngonium armigerum. B. Amphilophium chocoense, Bursera graveolens, Anthodiscus chocoensis, Pristimera
tenuiora and Parinari parvifolia. C. Gurania bignoniacea, Erythroxylum mbriatum, Aparisthmium cordatum and Mabea tenorioi.
Ortiz et al. | Additions to the ora of Panama 607
Figure 4. Map showing the distribution of selected new reports (circles). A. Abarema adenophora, Diplotropis purpurea, Tachigali cos-
taricensis, Taralea oppositifolia and Zygia conzat tii. B. Tachia parviora, Lozania g labrata, Ocotea aciphylla, Pleurothyrium glabritepalum and
Byrsonima garcibarrigae. C. Pentaplaris doroteae, Danaea grandifolia, Blakea dimorphophylla, Henriettea odorata and Miconia paleacea.
608 Check List 15 (4)
Figure 5. Map showing the distribution of selec ted new reports (circles). A. Ficus donnell-smithii, Calyptranth es bracteata, Eugen ia albicans,
Ouratea rinconensis and Sobralia purpurella. B. Passiora macrophylla, Pera oppositifolia, Digitaria costaricensis, Lasiacis ligulata, Lasiacis
sloanei and Portulaca papillato-stellulata. C. Clematis populifolia, Hasseltia lateriora, Vouarana anomala, Pouteria bulliformis, Cuatresia
trianae and Paypayrola confertiora.
609
Ortiz et al. | Additions to the ora of Panama
Petaquilla, helipad C08, Río Belencillo; 08°48′41″ N,
080°43′59″ W; 26 Jul. 2009; B. Araúz & J. De Gracia
2024 (MO, PMA); ibid., Sierra 1 (a un costado de la par-
cela de restauracion 2.5), bosque secundario intervenido;
08°49′54″ N, 080°41′05″ W; 291 m; 27 Jul. 2015; J. Batista
1399 (MO, PMA). Panamá Oeste. Carretera que con-
duce al pueblo de Ciri Grande; 08°49′52″ N, 080°02′39″
W, 13 Jun. 2011; R. Pérez et al. 2468 (MO, SCZ); Car-
retera Santa Rita-Ciri Grande, km 30; 08°48′17″ N,
080°04′42″ W; 218 m; 17 Apr. 2012; C. Galdames et al.
7032 (MO, SCZ); ibid.; 14 Jun. 2012; J. Aranda 4203
(MO). Figure 3A.
Global distribution. Bolivia, Colombia, Panama, and
Venezuela.
Identi cation. O choterenaea c olombiana i s d istin-
guished by having imparipinnate leaves, owers many in
thyrsi at the ends of the short lateral branches, ve per-
sistent sepals, ve petals and stamens, prominent disk,
tricarpellate pistil with one fertile carpel and very at-
tened fruits, long-pilose along the margin. So far, this is
the only species described for the genus Ochoterenaea
(Barkley 1942).
APOCYNACEAE
Macropharynx renteriae A.H. Gentry, Phytologia 47:
99 (Gentry 1980).
New records. Coclé. Distrito de El Valle, colectado
después de las granjas de pollos, a orillas de vía que
va hacia la entrada del Sendero El Gaital; 08°37′49″
N, 080°65′69″ W; 800 m; 28 Jun. 2017; E. Campos 960
(PMA). Colón. Teck Cominco Petaquilla mining con-
cession, forested slopes along ridge road; 08°49′56″ N,
080°39′32″ W; 130 m; 9 Mar. 2008; G. McPherson 20490
(MO, PMA); Distrito de Colón, colectado a orillas de la
vía de Santa Rita; 09°19′47″ N, 079°46′59″ W; 247 m; 15
Jul. 2015; E. Campos 485 (PMA, SCZ). Figure 3A.
Global distribution. Colombia, Costa Rica, Ecuador,
Honduras, and Panama.
Identi cation. This species is distinguished by having
puberulent terete branches, leaves with ferruginous-
puberulous petioles, broadly elliptic to ovate-elliptic
blades, rounded at base and acute or caudate-acumi-
nate at apex, conspicuously bracteate inorescences
(one to many owers) with long pedicels, owers with
foliaceous sepals, narrowly elliptic, acute apically and
a tubular-infundibuliform white corolla (Gentry 1980).
Macropharynx renteriae is the only representative of the
genus in Central America (previously reported in Costa
Rica) (Morales 1997).
AQUIFOLIACEAE
Ilex skutchii Edwin ex T.R. Dudley & W.J. Hahn in
Hahn, Novon 6: 181 (Hahn 1996).
New records. Colón. Westernmost part of province,
site of proposed copper mine (INMET), tailings area,
lowland forest; 08°52′10″ N, 080°40′00″ W; 75 m; 13
Apr. 2009; G. McPherson & Jean-Yves Serein 20936
(MO, PMA). Figure 3A.
Global distribution. Costa Rica, Nicaragua, and
Panama.
Identi cation. This species is characterized by hav-
ing elliptic to ovate or obovate blades, acute to apicu-
late at the apex, margins at, venation the same color as
the blades, prominent stigmatic residue, green or light
brown young stems (when dry), peduncles and pedicels
usually recurved (ca 0.5 mm in diameter) and spheri-
cal to globose fruits (2.5–3.5 mm long) (Hahn 1996). In
Panama, I. skutchii is most similar to I. fortunensis W.J.
Hahn, which diers in having thicker peduncles (2–3
mm diam.) and smaller fruits (0.9–1.1 mm long) (Hahn
1993).
ARACEAE
Rhodospatha monsalveae Croat & D.C. Bay in Croat
and Mora, Aroideana 27: 123 (Croat and Mora 2004).
Figure 1B
New records. Colón. Teck Cominco Petaquilla min-
ing concession, forest near helipad H44; 08°53′17″ N,
080°45′09″ W; 160 m; 30 Jun. 2008; G. McPherson 20754
(MO); Coclé del Norte, área del helipad T02A, cami-
nando hacia la ruta oeste; 08°53′34″ N, 080°40′55″ W;
138 m; 19 Jul. 2012; A. Espinosa 6014 (MO). Veraguas.
Santa Fe, Parque Nacional Santa Fe, bosque secundario
abierto, bastante plano; 08°41′44″ N, 080°53′00″ W; 290
m; 8 Feb. 2014; A. Morris et al. 2086 (MO); Santa Fe,
área del Rio Piedra, trocha bordeando la ribera del río,
bosque muy húmedo con una alta presencia de epítas y
briótos en los árboles; 08°44′06″ N, 080°46′19″ W; 16
Dec. 2013; L. Martínez et al. 1558 (MO). Figure 3A.
Global distribution. Colombia, Ecuador, and Panama.
Identi cation. The species is characterized by its light-
brown, conspicuously ssured stems, petioles sheathed
to the geniculum, usually drying reddish brown with
the sheath usually deciduous with dark-brown persistent
fragments and blades which are more or less equilateral
and decurrent at the base, drying usually grayish above
and reddish brown and minutely dark-granular (pellu-
cid-punctate lines) on the lower surface (Croat and Mora
2004). Rhodospatha monsalveae is similar to R. pellu-
cida Croat & Grayum, which diers in having blades
with numerous pellucid-punctate lines on lower surfaces.
Syngonium armigerum (Standl. & L.O. Williams) Croat,
Ann. Missouri Bot. Gard. 68: 585 (Croat 1981).
(≡) Philodendron armigerum Standl. & L.O. Williams, Ceiba 3, 1952.
New records. Colón. Teck Cominco Petaquilla mining
concession, forest near current end of Petaquilla road;
08°49′52″ N, 080°41′05″ W; 290 m, 28 Jun. 2008; G.
McPherson 20717 (MO, PMA). Figure 3A.
Global distribution. Costa Rica and Panama.
Identi cation. The species is characterized by its scan-
610 Check List 15 (4)
dent hemiepiphytic habit, brown-drying stems with lon-
gitudinal wrinkles, short petioles (5–7 cm long), oblong
to oblong-elliptic simple bl ades with obscure black punc-
tations on the lower surfaces, weakly lobed at the base
(the lobes broadly triangular or rounded), few primary
lateral veins (ve to six pairs) and usually one inores-
cence per node (Croat 1981). In Panama, Syngonium
armigerum is similar to S. brewsterense Croat & Delan-
nay, which diers in having longer petioles (21–45 cm
long) and two inorescences per node (Croat et al. 2019).
BIGNONIACEAE
Amphilophium chocoense (A.H. Gentry) L. G. Lohm-
ann, Ann. Missouri Bot. Gard. 99: 402 (Lohmann 2014).
(≡) Distictella chocoensis A.H. Gentry, Phytologia 47, 1980.
Figure 1A
New records. Colón. Teck Cominco Petaquilla min-
ing concession, forest along Río Caimito, near mouth;
09°01′06″ N, 080°40′56″ W; 5 m; 1 Mar. 2008; G.
McPherson 20358 (MO, PMA); ibid., forest near heli-
pad, 8°50’05” N, 80°38’48” W; 140 m; 20 Jun. 2008; G.
McPherson 20547 (MO, PMA); Site of proposed copper
mine (MPSA), forested slopes; 08°48′27″ N, 080°36′20″
W; 100 m; 13 Dec. 2009; G. McPherson & M. Merello
21292 (MO, PMA); Coclé del Norte, colectada a orillas de
la carretera frente al punto de liberaciones de serpientes
(Proyecto Minera Panamá, S.A.); 08°50′ N, 080°45′ W;
2009–2010; Y. Yaleman et al. 64 (MO, PMA); Donoso,
área de concesión de Minera Panamá, frente a Botadero
500, a orilla de la calle; 08°49′15″ N, 080°39′22″ W; 55
m; 2 Jul. 2012; J.F. Carrión 815 (PMA). Figure 3B.
Global distribution. Colombia and Panama.
Identi cation. This species is distinguished by having
short (0.03–0.7 mm long) trichomes along the branchlets
and petioles, broad pseudostipules (width 1/2 to equal
length), b ifoliolate l eaves with petiolules shorter than pet-
ioles, ovate, lanceolate or elliptic leaets, pubescent on
the lower surface, tertiary veins raised on lower surfaces
(more raised than higher order venation) and immersed
on upper surfaces, and ovaries without a stipe. Amphi-
lophium chocoense diers from the other Panamanian
congener, A. magnoliifolium (Kunth) L.G. Lohmann, by
having pubescent leaets on lower surfaces (Pool 2009).
BURSERACEAE
Bursera graveolens (Kunth) Triana & Planch., Ann. Sci.
Nat., Bot., sér. 5, 14: 303 (Triana and Planchon 1872).
(≡) Elaphrium graveolens Kunth, Nov. Gen. Sp., IV ed. 7, 1824.
New records. Darién. Chepigana, Parque Nacional
Darién, Punta Garachiné, bosque seco tropical; 08°05′18″
N, 078°25′23″ W; 200 m; 22 Jun. 2013; J.F. Carrión 1175
(PMA). Figure 3B.
Global distribution. Colombia, Costa Rica, Cuba,
Ecuador, El Salvador, Guatemala, Honduras, Mexico,
Nicaragua, Panama, Peru, and Venezuela.
Identi cation. This species is distinguished by having
once-pinnate leaves with winged rachis, bright green
leaets with crenate margins, pubescent to glabrous (not
densely tomentose), attenuate to long acuminate at the
apex, non-exfoliating bark, tetramerous owers, deltoid
sepals, dehiscent fruits in two valves and a white pseu-
doaril (Daly 1993). In Panama, B. graveolens is most
similar to B. tomentosa (Jacq.) Triana & Planch., which
also has leaves with winged rachis. Bursera tomentosa
diers in having densely tomentose leaets and lanceo-
late sepals.
CARYOCARACEAE
Anthodiscus chocoensis Prance, Brittonia 32: 530
(Prance 1980).
Figure 1C
New records. Colón. Donoso, westernmost part of pro-
vince, site of proposed copper mine (INMET), along
route of proposed road, lowland forest; 08°53′04″ N,
080°42′27″ W; 150 m; 12 Apr. 2009, G. McPherson
20902 (MO); Área del proyecto minero Petaquilla, 08°
51′25″ N, 080°41′40″ W; 21 Jan. 2008; N. Guerrero 1213
(MO); ibid.; 08°51′ N, 080°41′ W; 28 Jun. 2008; N. Guer-
rero et al. 1413 (MO); MPSA Concession, Valle Grande,
Sierra 19; 08°49′54″ N, 080°41′05″ W; 291 m; 17 May
2012; B. Hammel et al. 26219 (MO); ibid., trail to Pet-
aquilla Camp; 08°49′59″ N, 080°41′11″ W; 270 m; 20
May 2012; B. Hammel et al. 26289 (MO); Teck Cominco
Petaquilla mining concession, forested slopes along
ridge road; 08°49′56″ N, 080°39′32″ W; 130 m; 9 Mar.
2008; G. McPherson 20479 (MO). Figure 3B.
Global distribution. Colombia, Costa Rica, and Panama.
Identi cation. Anthodiscus chocoensis diers from
other species within the genus by the larger (12–15 × 6–7
cm) acuminate leaves, coriaceous blades, the shorter leaf
apex, the longer petiolules and shorter petioles and the
thicker inorescence and pedicels (Prance 1980). The
specimens cited here represent the rst records of the
genus Anthodiscus in Panama.
CELASTRACEAE
Pristimera tenuiora (Mart. ex Peyr.) A.C. Sm., Britto-
nia 3: 382 (Smith 1940).
(≡) Hippocratea tenuiora Mart. ex Peyr., Fl. Bras. 11, 1878.
New records. Colón. Donoso, Teck Cominco Petaquilla
mining concession, forest along river’s banks, Caimito
River; 08°58′53″ N, 080°40′40″ W; 5 m; 27 Feb. 2008; G.
McPherson 20253 (MO, PMA). Figure 3B.
Global distribution. Colombia, Costa Rica, Brazil,
French Guiana, Nicaragua, Panama, Peru, Suriname, and
Venezuela.
Identi cation. According to Lombardi (2015), Pris-
timera can be recognized in sterile specimens by its
greenish-drying blades with cream-colored veins, but
the identication of the species in this condition is very
Ortiz et al. | Additions to the ora of Panama 611
dicult or impossible. Pristimera tenuiora diers from
other Neotropical species of the genus in having tubu-
lar owers and ovate segments on fruits and seeds with
the wing partially surrounding the seminiferous nucleus
(Lombardi 2015).
CHRYSOBALANACEAE
Parinari parvifolia Sandwith, Bull. Misc. Inform. Kew
1931: 374 (Sandwith 1931).
New records. Colón. Donoso, MPSA Concession, intact
forest, bosque primario; 08°50′59″ N, 080°38′29″ W; 29
Aug. 2008; N. Guerrero NG1642 (PMA). Figure 3B.
Global distribution. Brazil, Colombia, Costa Rica,
French Guiana, Guyana and Panama.
Identi cation. Parinari parvifolia is distinguished by
having petioles with two inconspicuous glands near the
blade, terete petioles, elliptic to oblong-lanceolate blades,
numerous primary veins (27–30 pairs) and a slightly
impressed midrib for its entire length on upper surface
(Prance 1972). Previously, the only species reported for
this genus in Panama was P. chocoensis (Correa et al.
2004), which diers from P. parvifolia in having a mid-
rib impressed only on the proximal portion of the blade
and canaliculate petioles.
CUCURBITACEAE
Gurania bignoniacea (Poepp. & Endl.) C. Jerey, Kew
Bulletin 33: 354 (Jerey 1978).
(≡) Anguria bignoniacea Poepp. & Endl., Nov. Gen. Sp. Pl. 2, 1838.
New records. Colón. Donoso, Coclé del Norte, área
del helipad Escorpio 02, tomando la ruta sur; 08°55′29″
N, 080°40′08″ W; 60 m; 21 Jul. 2012; A. Espinosa 6026
(MO, PMA). Panamá. East slope of Cerro Jefe, cloud
forest; 09°14′02″ N, 079°22′20″ W; 823 m; 8 Feb. 1966;
E. Tyson 3401 (MO). Figure 3C.
Global distribution. Bolivia, Brazil, Colombia, Ecua-
dor, French Guiana, Guyana, Panama, Peru, Suriname,
and Venezuela.
Identi cation. Wunderlin (1978a) made taxonomic com-
binations in Anguria and proposed Psiguria bignoniacea
(Poepp. & Endl.) Wunderlin, which was transferred to
Gurania as G. bignoniaceae (Poepp. & Endl.) C. Jerey
(Jerey 1978). An earlier report of this species [referred as
P. bignoniacea (Poepp. & Endl.) Wunderlin] in the Flora
of Panama treatment (Wunderlin 1978b), was erroneously
based on P. triphylla (Miq.) C. Jerey (Jerey 2001),
which diers in having glabrous leaves and owers with
a green hypanthium (vs reddish in Gurania). As currently
circumscribed, G. bignoniacea is a complex, which is
characterized by membranous trifoliolate or simple leaves
and an indumentum (when developed) of rather coarse,
scattered, fairly long hairs on the petioles, leaves, and hy-
panthia (Jerey 1978). In Panama, G. bignoniacea could
be confused with G. coccinea Cogn., but the latter diers
in having shorter glabrescent pedicels and glabrous fruits.
ERYTHROXYLACEAE
Erythroxylum mbriatum Peyr. in Martius, Fl. Bras. 12:
162 (Peyritsch 1878).
Figure 1D
New records. Coclé. Above El Copé, along road to
Caribbean side, Atlantic slope; 08°38′ N, 080°35′ W;
720–800 m; 31 Aug. 1988; G. McPherson 12862 (MO).
Colón. Área del proyecto minero Petaquilla, Valle
Grande, antiguo campamento cuatro crestas; 21 Jun.
2009, B. Araúz 1939 (PMA); ibid., Quebrada Brazo; 08°
49′01″ N, 080°38′30″ W; 147 m; 17 Jun. 2014; I. Vergara-
Pérez 576 (MO, PMA). Panamá. Chepo, Isla Majé-
Bayano; 09°09′30″ N, 078°49′52″ W; 50–100 m, 23 Aug.
1976; L. Aguilar 166 (MO). Figure 3C.
Global distribution. Brazil, Colombia, Costa Rica, Ecua -
dor, French Guiana, Nicaragua, Panama, Peru, and Venezuela.
Identi cation. Erythroxylum mbriatum is easily recog-
nized by its small and striated persistent stipules (0.15–
0.2 cm long), with the two recurved lateral setae, often
mbriated at apex and fruits with endocarp pointed at
apex (Jara-Muñoz 2015). This species is most similar to
E. citrifolium A. St.-Hil., which diers in having longer
(0.4–1.2 cm long) and deciduous stipules (Plowman and
Barrie 2010).
EUPHORBIACEAE
Aparisthmium cordatum (A. Juss.) Baill., Adansonia 5:
307 (Baillon 1865).
(≡) Conceveibum cordatum A. Juss., Euphorb. gen. 43, 1824.
New records. Panamá Oeste. Río Indio, Los Chorros,
orilla de la carretera; 08°46′26″ N, 080°07′49″ W; 401
m; 5 Jun. 2019; R. Flores 4154 (PMA, UCH); Carretera
El Espino-río Indio, Oeste de La Chorrera, km 30 desde
la Carretera Interamericana; 08°48′21″ N, 080°05′39″ W;
300 m; 8 Jul. 2014; C. Galdames 7615 (PMA, SCZ);
ibid.; C. Galdames 7616 (PMA, SCZ). Figure 3C.
Global distribution. Bolivia, Brazil, Costa Rica, Colom-
bia, Ecuador, French Guiana, Guyana, Panama, Peru,
Suriname, and Venezuela.
Identi cation. Aparisthmium is a monotypic genus that
includes the following characters: leaves with two stipel-
like appendages at the apex of the petiole, four sepals,
three to ve stamens, capsules with three mericarps with
ovate to elliptic seeds (Secco 2004). This genus could be
confused with species of Alchornea and Conceveiba, but
both diers in having capsules with two mericarps and
stellate trichomes, respectively.
Mabea tenorioi Mart. Gord., J. Jiménez Ram. & Cruz
Durán, Anales Inst. Biol. Univ. Nac. Autón. México,
Biol. 71: 93 (Martínez-Gordillo et al. 2000).
Figure 2C
New records. Colón. Distrito de Donoso, área de conc-
esión de Minera Panamá, Botija Pit, área 1; 08°50′33″ N,
080°38′56″ W; 23 Sept. 2013; J.F Carrión 863 (PMA);
612 Check List 15 (4)
Donoso; 08°49′25″ N, 080°41′09″ W; 22 Jul. 2008; N.
Guerrero 1359 (PMA); Teck Cominco Petaquilla mining
concession, forested slopes; 08°50′17″ N, 080°38′52″ W;
171 m; 3 Dec. 2007; G. McPherson & H. van der Wer
19940 (MO, PMA); ibid., slopes along exploration road;
08°50′22″ N, 080°38′51″ W; 184 m; 15 Sept. 2007; G.
McPherson 19559 (MO, PMA); MPSA mining site, for-
ested slopes along trail called Quatro Crestos; 08°49′24″
N, 080°42′32″ W; 201 m; 3 Sept. 2014; G. McPher-
son & M. Grayum 21574 (MO, PMA); Área de estudio
poblaciones EdI, zona de proyecto, punto 3 (ZP-P3),
bosque húmedo tropical, sobre una pendiente, área de
Botija; 08°50′35″ N, 080°39′27″ W; 190 m; 9 Feb. 2011;
L. Martínez et al. 751 (MO, PMA); Botija Rd., carretera
pionero; 08°50′06″ N, 080°39′17″ W; 12 Sept. 2012; H.
van der Wer et al. 24448 (MO, PMA); Área de conc-
esión del Proyecto Minera Cobre Panamá S. A., Botija,
MSA, área 43; 08°50′53″ N, 080°39′34″ W; 134 m; 30
Nov. 2015; I. Vergara-Pérez 1038 (MO, PMA); ibid.,
Valle Grande, camino hacia Petaquilla; 08°49′59″ N,
080°41′09″ W; 1 Oct. 2014; I. Vergara-Pérez 903 (MO,
PMA). Figure 3C.
Global distribution. Mexico and Panama.
Identi cation. This species is characterized by hav-
ing linear-lanceolate blades with distinct glands, whit-
ish on lower surface, staminate owers with 10 anthers,
unequal (central distinctly larger than laterals) and
apparently non-articulate pedicels, erect bracts with lat-
eral glands, free part of styles diuse and reexed style-
branches (Martínez-Gordillo et al. 2000). In Panama, M.
tenorioi could be confused with M. montana Müll. Arg.,
which diers in having staminate owers with more than
10 anthers and equal-sized pedicels.
FABACE A E
Abarema adenophora (Ducke) Barneby & J.W. Grimes,
Mem. New York Bot. Gard. 74: 74 (Barneby and Grimes
1996).
(≡) Pithecellobium adenophorum Ducke, Arq. In st . Biol. Veg. 4, 1938.
New records. Colón. Teck Cominco Petaquilla mining
concession, forest along creek; 08°49′43″ N, 080°39′37″
W; 237 m; 30 Nov. 2007; G. McPherson & H. van der
Wer 19890 (MO, PMA); San Juan del General, Conce-
sión del Proyecto Mina de Cobre Panamá, Valle Grande,
Sierra 18, bosque secundario maduro; 08°49′18″ N,
080°40′58″ W; 280 m; 25 Jul. 2014; J. De Gracia et al.
793 (MO, PMA). Figure 4A.
Global distribution. Brazil, Costa Rica, Honduras, Nica-
ragua, Panama, Peru, and Venezuela.
Identi cation. Abarema adenophora is distinguished
from other Central American species of the genus by
having usually two to four pairs of leaets per pinna, a
large (5–15 × 4–12 mm) funnelform-campanulate rst
petiolar nectary and dimorphic owers (Barneby and
Grimes 1996). This species could be confused with A.
macradenia (Pittier) Barneby & J.W. Grimes, which
diers in having more than seven pairs of leaets per
pinna (Zamora 2010).
Diplotropis purpurea (Rich.) Amsho, Meded. Bot.
Mus. Herb. Rijks Univ. Utrecht 52: 43 (Amsho 1939).
(≡) Tachigali purpurea Rich., Actes Soc. Hist. Nat. Paris 1, 1792.
New records. Colón. Site of proposed copper mine
(MPSA), forested slopes; 08°56′54″ N, 080°41′42″ W;
140 m; 5 Dec. 2009; G. McPherson & M. Merello 21135
(MO, PMA); Site of proposed copper mine (MPSA), for-
ested slopes; 08°48′27″ N, 080°36′20″ W; 100 m; 13 Dec.
2009; G. McPherson & M. Merello 21287 (MO, PMA);
Santa Rita-Aguas Claras, parcela #31, 09°21′36″ N, 079°
45′2″ W; 7 Aug. 2003; R. Pérez 1065 (PMA). Figure 4A.
Global distribution. Bolivia, Brazil, Colombia, Ecua-
dor, French Guiana, Guyana, Panama, Suriname, and
Venezuela.
Identi cation. This species is distinguished by its arbo-
real habit (up to 40 m), leaets with prominently unequal
bases, panicles with several branches and fruits membra-
nous, samaroids (Vásquez 1997). The specimens cited
here represent the rst records of the genus Diplotropis
in Panama.
Tachigali costaricensis (N. Zamora & Po ved a) N. Zamora
& van der Wer, Harvard Pap. Bot. 15: 149 (van der Wer
and Zamora 2010).
(≡) Sclerolobium costaricense N. Zamora & Poveda, Novon 1, 1991.
New records. Coclé. El Harino, Parque Nacional G.D.
Omar Torrijos Herrera, colectado camino al río Tife,
área de Cano Sucio, a orillas del río Cano Sucio, terrenos
del señor Ife; 08°43′21″ N, 080°37′24″ W; 231 m; 17
Oct. 2012; L. Martínez 1078 (MO, PMA). Colón. MPSA
Con-cession, Botadero 600, plants collected after area
was cleared with chainsaws; 08°49′19″ N, 080°39′22″
W; 22 May 2012; B. Hammel et al. 26323 (MO, PMA);
Site of proposed copper mine (MPSA), forested slopes;
08°55′43″ N, 080°40′48″ W; 75 m; 4 Dec. 2009; G.
McPherson & M. Merello 21114 (MO, PMA); Coclé del
Norte, helipad C16, rumbo norte, curso superior del río
Boca Chica, área boscosa cerca al río; 08°46′21″ N, 080°
44′14″ W; 29 Aug. 2012; A. Zapata 3060 (MO, PMA).
Veraguas. Parque Nacional Santa Fe, área del río Santa
María (ent-rando por Piragual), bosque intervenido; 08°
33′17″ N, 081°02′48″ W; 404 m; 20 Nov. 2012, L.
Martínez et al. 1166 (MO, PMA). Figure 4A.
Global distribution. Costa Rica, Nicaragua, and Panama.
Identi cation. This species is distinguished by having
pectinate (like little pinnate leaves) and usually persis-
tent stipules, leaets with dense gold or gray sericeous
pubescence on lower surfaces and inorescences and
ower buds with brown pubescence (van der Wer and
Zamora 2010; Zamora 2010). Tachigali costaricensis
could be confused with T. panamensis van der Wer &
N. Zamor a, but the lat ter diers i n having scattered i ndu-
mentum on lower leaet surfaces and inorescences and
ower buds with reddish-brown pubescence.
613
Ortiz et al. | Additions to the ora of Panama
Taralea oppositifolia Aubl., Hist. Pl. Guiane 2: 745, t.
298 (Aublet 1775).
New records. Coclé. El Harino, Parque Nacional G.D.
Omar Torrijos H, La Rica, camino al río Tife, área del río
Blanco; 08°42′43″ N, 080°37′01″ W; 375 m; 18 Oct.
2012; A. Zapata et al. 3086 (MO, PMA). Colón. Donoso;
10 September 2008; B. Araúz 1807 (MO, PMA); Coclé
del Norte, Minera Panamá, helipat H55; 08°51′35″ N,
080°42′09″ W; 182 m; 16 Dec. 2010; A. Espinosa 5914
(MO, PMA); Teck Cominco Petaquilla mining conces-
sion, forested slopes; 08°50′17″ N, 080°38′52″ W; 171 m;
3 Dec. 2007; G. McPherson & H. van der Wer 19950
(MO, PMA); ibid., forest along ridge road; 08°49′22″ N,
080°39’32” W; 300 m; 25 Feb. 2008; G. McPherson &
M. Merello 20248 (MO, PMA); ibid., forest along río
Caimito; 09°00′25″ N, 080°41′03″ W; 5 m; 2 Mar. 2008;
G. McPherson 20426 (MO, PMA); Site of proposed
copper mine (MPSA), forested slopes; 08°48′27″ N, 080°
36′20″ W; 100 m; 13 Dec. 2009; G. McPherson & M.
Merello 21289 (MO, PMA). Figure 4A.
Global distribution. Brazil, Colombia, French Guiana,
Guyana, Panama, Peru, Suriname, and Venezuela.
Identi ication. Taralea oppositifolia is characterized by
having glabrous leaflets with inconspicuous venation on
upper surfaces and flowers with upper lip of calyx cori-
aceous and a densely pilose ovary (Lima and Aymard
1999). Taralea oppositifolia represents the first record of
the genus in Central America.
Zygia conzattii (Standl.) Britton & Rose, N. Amer. Fl.
23: 40 (Britton and Rose 1928).
(≡) Calliandra conzattii Standl., Contr. U.S. Natl. Herb. 20, 1919.
New records. Colón. Teck Cominco Petaquilla mining
concession, forest; 08°55′53″ N, 080°40′08″ W; 80 m; 24
Jun. 2008; G. McPherson 20625 (MO, PMA). Figure 4A.
Global distribution. Belize, Costa Rica, Mexico, and
Panama.
Identi ication. This species is distinguished by its leaves
with three or five subcoriaceous leaflets per pinna, ellip-
tic to broadly elliptic or asymmetric-oblong leaflets,
sessile whitish flowers and relatively narrow (1–1.5 cm)
fruits. Zygia conzattii could be confused with Z. latifolia
(L.) Fawc. & Rendle, which differs in having straight
and broader fruits (1.5–3.3 cm wide) and leaflets
typically with all secondary veins more or less patent
to slightly ascending (vs leaflets usually with two or
three second-ary veins strongly ascending in Z.
conzattii) (Zamora 2010).
GENTIANACEAE
Tachia parvi lora Maguire & Weaver, J. Arnold
Arbor. 56: 123 (Maguire and Weaver 1975).
Figure 1F
New records. Coclé. Camino al Alto Tífe; 08°42′56″ N,
080°37′45″ W; 383 m; 24 Jan. 2013; N. Jaén et al. 112
(PMA); Parque Nacional Omar Torrijos, bosque cercano
al Río Tífe, entrando por los terrenos del Sr. Ife Quiróz;
08°42′22″ N, 080°38′03″ W; 483 m; 17 Jul. 2013; A.
Zapata et al. 3184 (PMA). Colón. Distrito de Donoso,
campamento Petaquilla hacia Valle Grande, en dirección
13° SE; 3 Jul. 1996; A. Zapata et al. 983 (PMA); Área del
proyecto minero Petaquilla; 6 Dec. 2007; B. Araúz & J.
Meerman 1428B (PMA); ibid.; 6 Jan. 2007; B. Araúz &
P. Moreno 1505 (PMA); Proposed copper mining site;
Ar a úz 142 8 (MO, PMA); Teck Cominco Petaquilla min-
ing concession, forest; 08°49′06″ N, 080°41′02″ W; 180
m; 22 Jun. 2008; G. McPherson 20594 (MO, PMA);
Área de concesión del Proyecto Minera Cobre Panamá
S.A., Valle Grande; 08°50′01″ N, 080°41′10″ W; 171 m;
16 Sept. 2014; I. Vergara-Pérez 1012 (PMA). Figure 4B.
Global distribution. Bolivia, Colombia, Costa Rica,
Panama, and Peru.
Identi cation. This species is easily distinguished from
all other Tachia species by the combination of the follow-
ing characters: small corollas, frequently branched and
terete stems, and arcuate leaves (Struwe and Kinkade
2013). Tachia parviora is the only representative of the
genus in Central America (previously reported in Costa
Rica) (Davidse 2009).
LACISTEMATACEAE
Lozania glabrata A.H. Gentry, An n. Missouri Bot. Gard.
75: 1431 (Gentry 1988).
New records. Colón. Teck Cominco Petaquilla min-
ing concession, ridgetop forest along road; 08°49′28″
N, 080°39′29″ W; 190 m; 21 Sept. 2007; G. McPherson
19768 (MO); ibid.; 08°49′18″ N, 080°39′35″ W; 255 m;
28 Nov. 2007; G. McPherson & H. van der Wer 19857
(MO); ibid., forested slopes; 08°50′17″ N, 080°38′52″ W;
171 m; 3 Dec. 2007; G. McPherson & H. van der Wer
19941 (MO); ibid., Colina camp, forested slopes along
road; 08°49′23″ N, 080°39′32″ W; 101 m; 26 Feb. 2008;
M. Merello et al. 3063 (MO); ibid., forest in proposed
tailings area, 08°51′09″ N, 080°39′05″ W; 150 m; 19 Jun.
2008; G. McPherson 20530 (MO); Botija, carretera Pio-
nera, lowland wet rain forest; 08°50′06″ N, 080°39′17″
W; 10 Sept. 2012, H. van der Wer et al. 24429 (MO).
Figure 4B.
Global distribution. Colombia and Panama.
Identi cation. According to Gentry (1988), L. glabrata
is easily the most distinctive species of the genus by its
glabrous leaves and sub-entire blade margins and diers
from all its congeners by having stamens with the la-
ment deeply split at the apex (which gives the illusion of
two stamens).
LAURACEAE
Ocotea aciphylla (Nees & Mart.) Mez, Jahrb. Königl.
Bot. Gart. Berlin 5: 243 (Mez 1889).
(≡) Oreodaphne aciphylla Nees & Mart., Linnaea 8, 1833.
New records. Colón. Minera Panamá, Brazo; 08°48′
614 Check List 15 (4)
48″ W, 080°38′08″ W; 271 m; 26 Sept. 2014, C. Ramos
331 (PMA, MO). Figure 4B.
Global distribution. Bolivia, Brazil, Colombia, Ecua-
dor, French Guiana, Guyana, Panama, Peru, Suriname,
and Venezuela.
Identi cation. This species is characterized by hav-
ing terete, pubescent and solid branches, a canaliculate
petiole, an elliptic or narrowly elliptic-ovate blade (1–2
times longer that wide) that is strongly recurved at base,
pedicels 3– 7 mm long, short inorescences (2–10 cm
long) with sericeous-pubescent owers and relative long
fruits (to 3 cm long) (Allen 1964; Vásquez 1997). Based
on decurrent base blades and pubescent bisexual ow-
ers, O. aciphylla could be confused with O. insularis
(Meissner) Mez, but the latter species diers in having
broadly elliptic to obovat e or oblong blades, longer in o-
rescences (7–22 cm long) and smaller fruits (1–1.8 cm
long) (González-Ramírez and Poveda 2007).
Pleurothyrium glabritepalum van der Wer, Ann. Mis-
souri Bot. Gard. 80: 75 (van der Werff 1993).
New records. Veraguas. Área del río Guayabalito,
bosque secundario maduro, trocha que conduce a la
comunidad de Calle Larga; 08°44′10″ N, 080°46′59″ W;
387 m; 22 Mar. 2014; L. Martínez et al. 1776 (MO,
PMA). Figure 4B.
Global distribution. Colombia, Ecuador, and Panama.
Identi cation. According to van der Wer (1993), the
useful characters to identify P. glabritepalum are the fol-
lowing: the more or less erect pubescence, which is also
present on the upper leaf surface, the gland dots on the
upper leaf surface, the alternate, acuminate thin-textured
leaves, the few-owered inorescences, and the glabrous
inner surface of the unequal tepals. This species is most
similar to P. triorum van der Wer, which diers in
having equal tepals with the inner surface densely gray
papillose and blades glabrous on the abaxial surface (van
der Wer 2009).
MALPIGHIACEAE
Byrsonima garcibarrigae Cuatrec., Webbia 13: 621 (Cua-
trecasas 1958).
Figure 2E
New records. Coclé. Parque Nacional G.D. Omar Tor-
rijos Herrera, área boscosa en las proximidades del río
Cano Sucio; 08°43′00″ N, 080°36′41″ W; 245 m; 25 Jul.
2013; A. Zapata 3319b (MO, PMA); ibid., Distrito de
La Pintada, correg. El Harino, bosque cercano al río
Tife; 08°43′18″ N, 080°37′56″ W; 247 m; 20 Jul. 2013;
A. Zapata 3243 (MO, PMA); ibid., Caño Sucio, área
del Tífe, bosque húmedo; 08°43′18″ N, 080°37′56″ W;
247 m; 20 Jul. 2013; O. Ortiz et al. 1420 (MO, PMA);
ibid., Río San Juan, colectada cercano a la desemboca-
dura del río Escobal; 08°43′24″ N, 080°40′05″ W; 143
m; 20 Jul. 2013; L. Martínez 1325 (MO, PMA); ibid.,
área de Calle Larguita (Palmarazo), ribera del Río San
Juan (cerca del campamento), bosque primario ribereño;
08°43′20″ N, 080°40′23″ W; 140 m; 20 Jul. 2013; A.
Espinosa 6168 (MO, PMA). Colón. Teck Cominco Pet-
aquilla mining concession, forest near helipad; 08°50′16″
N, 080°39′17″ W; 160 m; 21 Jun. 2008; G. McPher-son
20576 (MO); MPSA Concession, Valle Grande; 08°49′
31″ N, 080°40′11″ W; 291 m; 18 May 2012; B. Hammel
26241 (MO); Donoso, San Juan del General, concesión
del Proyecto Mina de Cobre Panamá, Brazo, bosque
secundario maduro; 08°48′32″ N, 080°38′36″ W; 245 m;
9 Jul. 2014; J. De Gracia 787 (MO, PMA); Área de
concesión del Proyecto Minera Cobre Panamá S.A.,
antiguo campamento de Botija; 08°50′19″ N, 080°38′55″
W; 170 m; 4 Jun. 2014; I. Vergara-Pérez 541 (MO,
PMA); MPSA, sendero Botija, detrás vivero FCD, cerca
de quebrada Botija y río Botija; 08°50′12″ N, 080°38′41″
W; 117 m; 24 May 2015; J. Batista 1370 (MO, PMA);
Proyecto Minera Cobre Panamá, Botija, parcela
interpretativa de restauración ecológica 2.5, sierra 1; 08°
48′59″ N, 080°39′11″ W; 237 m; 31 Mar. 2016; I. Verg-
ara-Pérez 1083 (MO, PMA); ibid., camino hacia cantera
Botija; 08°50′32″ N, 080°39′18″ W; 178 m; 11 May 2014;
I. Vergara-Pérez 447 (MO, PMA); ibid., Botija, send-ero
detrás del vivero de plantas de MPSA; 08°49′43″ N, 080°
39′12″ W, 167 m; 29 Apr. 2014, I. Vergara-Pérez 380
(MO, PMA). Comarca Ngäbe-Buglé. Río Caña, finca de
Patricia Robinson; 09°01′06″ N, 081°43′26″ W; 30 m, 14
May 2014; A. Ibáñez et al. 8925AI (PMA). Figure 4B.
Global distribution. Brazil, Colombia, Panama, and Peru.
Identi ication. Byrsonima garcibarrigae is character-
ized by having papery, narrowly elliptic leaves with long
petioles to 2 cm long, white or pink flowers with puberu-
lent pedicels only 5–6 mm long, and outer anther locules
flattened and alate with the connective shorter than the
locules and blunt. This species is related to B. dressleri
W.H. Lewis, which differs in having obovate coriaceous
leaves, shorter petioles (1 cm long), longer (7–10 mm
long) pinkish pedicels, and outer anther locules non-
alate and cylindrical with the connective equaling the
locules (Lewis 1973).
MALVACEAE
Pentaplaris doroteae L.O. Williams & Standl., Ceiba 3:
140 (Williams and Standley 1952).
New records. Colón. Donoso, Teck Cominco Petaquilla
mining concession, slope, collected near plot C003; 08°
50′22″ N, 080°38′51″ W; 184 m; 18 Sept. 2007; G.
McPher-son 19661 (MO). Figure 4C.
Global distribution. Colombia, Costa Rica, and Panama.
Identi ication. This species is easily distinguished from
two other Pentaplaris species by the combination of the
following characters: leaf blades with stellate-pubes-
cence below, the hairs multi-rayed stellate, cordate,
asymmetrical to strongly asymmetrical at base, obovoid
to pyriform flower buds and fruit with calyx ca 2–3.5 cm
long (Bayer and Dorr 1999).
Ortiz et al. | Additions to the ora of Panama 615
MARATTIACEAE
Danaea grandifolia Underw., N. Amer. Fl. 16: 18
(Underwood 1909).
New records. Coclé. Parque Nacional G.D. Omar Tor-
rijos Herrera, área boscosa en las proximidades del río
Guabal, aguas arriba, ribera izquierda, terrenos colin-
dantes con la propiedad del Sr. Santos Navas; 08°42′19″
N, 080°35′16″ W; 378 m; 24 Jul. 2013; A. Zapata 3309
(MO, PMA); ibid., La Rica, bosque cercano al río Juan
Julio, bosque húmedo con presencia de muchas quebra-
das; 08°42′43″ N, 080°35′30″ W; 200 m; 22 Jul. 2013;
O. Ortiz et al. 1453 (MO, PMA); Colón. Donoso, Coclé
del Norte, Minera Panamá, helipat C01; 08°58′24″ N,
080°42′22″ W; 130 m; 9 Mar. 2010; A. Espinosa et al.
5527 (MO, PMA); Teck Cominco Petaquilla mining
concession, forest in proposed tailings area; 08°51′09″
N, 080°39′05″ W; 155 m; 19 Jun. 2008; G. McPherson
20527 (MO); Coclé del Norte, área del helipad Escor-
pio 02, tomando la ruta sur; 08°55′31″ N, 080°40′09″ W;
39 m; 21 Jul. 2012; A. Espinosa et al. 6021 (MO, PMA);
Coclé del Norte, Minera Panamá, helipad C02; 08°56′37″
N, 080°44′00″ W; 134 m; 16 Mar. 2010; A. Espinosa et
al. 5719 (MO, PMA); ibid., área del helipad C15, bosque
húmedo tropical; 18 Mar. 2010; A. Zapata et al. 2554 (MO,
PMA); ibid., Minera Panamá, helipad C02; 08°56′58″ N,
080°43′52″ W; 131 m; 14 Mar. 2010; A. Espinosa 5670
(MO, PMA); ibid., área del helipad C11, bosque húmedo
tropical; 08°57′54″ N, 080°37′01″ W; 12 Mar. 2010; A.
Zapata & J.I. González 2545 (MO, PMA). Figure 4C.
Global distribution. Colombia, Dominican Republic,
Haiti, Panama, Puerto Rico, and Venezuela.
Identi cation. Underwood (1909) described Danaea
grandifolia, as a taxon related to D. nodosa (L.) Sm., and
reported it from Colombia (type from Valparaiso, Santa
Marta) and Panama (without citing a Panamanian speci-
men). Rolleri (2004) synonymized D. grandifolia under
D. nodosa, arguing that the type material of D. gran-
difolia corresponded to well-developed specimens of D.
nodosa. However, Christenhusz (2010) recognized that
D. grandifolia is restricted to South America and dis-
tinct from D. nodosa. According to Christenhusz (2010),
D. grandifolia is distinguished by the combination of
the following characters: terrestrial plants, creeping rhi-
zomes with several rows of leaves, these placed more or
less radially and the roots all on the lower side; leathery
or transparent blades, always with normally developed
terminal pinnae (never replaced by bulbils), iridescent
(when juvenile); entire pinna margins (sometimes slightly
sinuate at apex). Danaea grandifolia could be confused
with D. nodosa, which diers in having dentate, denticu-
late, serrate, serrulate or crenulate pinna margins.
MELASTOMATACEAE
Blakea dimorphophylla (Almeda) Penneys & Almeda
in Penneys and Judd, PhytoKeys 20: 25 (Penneys and
Judd 2013).
(≡) Topobea dimorphophylla Almeda, Proc. Calif. Acad. Sci., 4th Ser.
55, 2001.
New records. Colón. Minera Panamá, north waste dump,
plant site; 08°51′20″ N, 080°38′24″ W; 143 m; 24 Oct.
2014; C. Ramos 366 (MO, PMA). Figure 4C.
Global distribution. Costa Rica and Panama. Monro et
al. (2017) reported this species in Panama; however, they
did not cite any representative specimens from Panama.
Identi cation. This species is characterized by hav-
ing leaf blades markedly unequal in size at each node,
densely pubescent on both surfaces and basally fused
oral bracts (Almeda 2001). Blakea dimorphophylla is
similar to B. intricata (Almeda) Penneys & Almeda, but
the latter diers in having slightly unequal leaf blades
and free oral bracts (Almeda 2009).
Henriettea odorata (Markgr.) Penneys, Michelang.,
Judd & Almeda, Syst. Bot. 35: 798 (Penneys et al. 2010).
(≡) Henriettella odorata Markgr., Notizbl. Bot. Gart. Berlin-Dahlem
15, 1941.
New records. Colón. Teck Cominco Mining Conces-
sion, Camp Colina, forested slopes along road; 08°49′30″
N, 080°40′11″ W; 298 m; 1 Mar. 2008; M. Merello & J.I.
González 3160 (MO); ibid., forest near helipad; 08°50′16″
N, 080°39′17″ W; 160 m; 21 Jun. 2008; G. McPherson
20 574 (MO). Figure 4C.
Global distribution. Colombia, Costa Rica, Ecuador,
Nicaragua, and Panama.
Identi cation. Henriettea odorata is recognized by its
essentially glabrous leaf blades on both surfaces, gla-
brous pedicels (in fruits) and consistently tetramerous
owers with unguiculate petals. This species is similar
to H. fascicularis (Sw.) C. Wright, which diers in hav-
ing sparsely setulose blades on upper surfaces, oblong
anthers and minutely glandular-puber ulent fruiting pedi-
cels (Almeda 2007, 2009).
Miconia paleacea Cogn., Monogr. Phan. 7: 757 (Cog-
niaux 1891).
Figure 2B
New records. Bocas del Toro. Changuinola, Bosque
Protector Palo Seco, Río Changuinola arriba, Charco
La Pava, bosque trasero de la casa del señor Reynaldo
Abrego; 09°09′31″ N, 082°30′35″ W; 424 m; 3 Feb. 2013,
O. Ortiz et al. 1214 (PMA); ibid., comunidad de Guay-
acán, 09°08′22″ N, 082°29′47″ W; 232 m; 4 Feb. 2014;
O. Ortiz & C. Galdames 2089 (PMA, SCZ). Figure 4C.
Global distribution. Belize, Brazil, Colombia, Ecuador,
Honduras, Nicaragua, Panama, Peru, and Venezuela.
Identi cation. This species is distinguished by the
combination of the following characters: distal twigs
with trichomes basally attened and reexed, petioles
and basal portions of the elevated nerves on lower sur-
faces of the blades usually stained red-purple, and by its
large leaves abruptly acuminate at the apex and ino-
rescences moderately to densely covered by reexed
rigid, basally attened hairs, with these underlaid by soft
616 Check List 15 (4)
woolly interwoven hairs (Almeda 2007). Both Miconia
paleacea and M. calvescens DC. have purple-red blades
beneath, but the latter diers in having inorescences
and hypanthium moderately covered by late-deciduous
stellate hairs (Almeda 2009).
MORACEAE
Ficus donnell-smithii Standl., Contr. U.S. Natl. Herb.
20: 21 (Standley 1917).
New records. Colón. Site of proposed copper mine
(MPSA), forested slopes; 08°50′13″ N, 080°37′04″ W;
70 m; 11 Dec. 2009; G. McPherson & M. Merello 21269
(MO, PMA). Figure 5A.
Global distribution. Belize, Bolivia, Brazil, Colombia,
Costa Rica, Ecuador, French Guiana, Guatemala, Guy-
ana, Honduras, Panama, Peru, Suriname, and Venezuela.
Identi cation. This species is recognized by its densely
pubescent stipules, lanceolate to narrowly oblong leaf
blades with glandular dots in the axils of the basal lateral
veins and pedunculate gs (arranged in pairs) (Standley
1917; González-Ramírez 2007). Ficus donnell-smithii is
vegetatively similar to F. yoponensis Desv., which dif-
fers in having oblong-elliptic to very narrowly oblong or
lanceolate-oblong blades with glandular dots at the base
of mid-veins and a solitary g.
MYRTACEAE
Calyptranthes bracteata M.L. Kawas. & B. Holst, Brit-
tonia 46: 137 (Kawasaki and Holst 1994).
Figure 2D
New records. Colón. Coclé del Norte, área del helipad
CR10, tomando hacia el oeste; 08°56′40″ N, 080°42′28″
W; 19 Jul. 2012; J. Aranda 4251 (MO, PMA); ibid,
Minera Panamá, helipad H55; 08°51′38″ N, 080°41′34″
W; 85 m; 12 Dec. 2010; A. Espinosa 5842 (MO, PMA);
ibid, helipad H55; 08°51′37″ N, 080°41′34″ W; 95 m; 12
Dec. 2010; A. Espinosa 5838 (MO, PMA); ibid., área del
helipad TO2A, caminando hacia la ruta oeste; 08°53′38″
N, 080°40′44″ W; 130 m; 19 Jul. 2012; A. Espinosa 6006
(MO, PMA); ibid., área del helipad TO2A, caminando
hacia la ruta norte; 08°53′38″ N, 080°40′44″ W; 130 m;
17 Jul. 2012; A. Espinosa 5987 (MO, PMA); ibid., área
del helipad BE03 (río Escribano), tomando la ruta norte;
08°52′17″ N, 080°50′56″ W; 52 m; 25 Jul. 2012; A. Espi-
nosa 6065 (MO, PMA); ibid., A. Espinosa 6066 (MO,
PMA); Teck Cominco Petaquilla mining concession,
forest near helipad; 08°49′12″ N, 080°40′52″ W; 190 m;
22 Jun. 2008; G. McPherson 20591 (MO); ibid., área del
Helipad T02A, tomando la ruta Sur, a orilla de una tro-
cha; 08°53′20″ N, 080°40′26″ W; 143 m; 18 Jul. 2012;
L. Martínez 870 (MO, PMA); ibid., área del río Escrib-
ano, cercano al río Belén límite de la Provincia de Colón
y Veraguas; 08°52′17″ N, 080°51′04″ W; 37 m; 25 Jul.
2012; L. Martínez 928 (MO, PMA). Figure 5A.
Global distribution. French Guiana and Panama.
Identi cation. Calyptranthes bracteata diers from all
other species in the genus by the short, few owered,
brown-tomentose inorescences with large, persistent
bracts and elliptic to narrowly elliptic (12–20 × 3.5–8
cm), coriaceous leaf blades. Based on tomentose ino-
rescences, Calyptranthes bracteata is similar to C. ellip-
tica B. Holst & M.L. Kawas., which diers in having
broadly elliptical to elliptical (4–9.4 × 3.1–7 cm) charta-
ceous leaf blades and deciduous bracts (Kawasaki and
Holst 1994).
Eugenia albicans (O. Berg) Urb., Bot. Jahrb. Syst. 19:
617 (Urban 1895).
(≡) Stenocalyx albicans O. Berg, Linnaea 30, 1860.
New records. Colón. Teck Cominco Petaquilla min-
ing concession, ridgetop forest along road; 08°49′18″ N,
080°39′35″ W; 255 m; 28 Nov. 2007; McPherson & van
der Wer 19848 (MO); ibid., forest on slope; 08°49′08″
N, 080°37′14″ W; 100 m; G. McPherson & H. van der
Wer 20063 (MO); ibid., forested slopes along ridge
road; 08°49′51″ N, 080°40′55″ W; 250 m; G. McPher-
son 20469 (MO); Minera Panamá, Valle Grande, Sierra
18, sendero de CABO hacia plataformas; 08°50′06″ N,
080°40′43″ W; 303 m; 9 Oct. 2014, C. Ramos 348 (MO,
PMA). Figure 5A.
Global distribution. Brazil, Caribbean (Leeward
Islands, Trinidad and Tobago, Windward Islands),
French Guiana, Guyana, Panama, and Suriname.
Identi cation. This species is recognized by having
markedly bicolorous oblong-lanceolate blades which are
smooth and lustrous above and very pale or glaucous on
the lower surfaces; long appressed bright coppery silky
hairs that cover the inorescences and owers and long
(3–4 mm long) thin-textured, glabrous (internally) and
broadly hooded calyx-lobes that cover the corolla in
bud (McVaugh 1969). Eugenia albicans is vegetatively
similar to E. coibensis Barrie (both have greyish blades
when dry), but the latter diers in having ovate, elliptic
or obovate leaf blades and elliptic petals and sepals.
OCHNACEAE
Ouratea rinconensis Whitef., Novon 2: 277 (Wh itefoor d
1992).
New records. Colón. Coclé del Norte, Minera Panamá,
helipad C22; 08°50′37″ N, 080°45′35″ W; 120 m; 20 Oct.
2010; A. Espinosa 5784 (MO, PMA); Coclé del Norte,
Minera Panamá, helipad H02; 08°56′13″ N, 080°40′21″
W; 54 m; 15 Dec. 2010; A. Espinosa 5893 (MO, PMA);
Coclé del Norte, helipad C10, a orillas del río Belen-
cillo, bosque secundario intervenido; 08°48′25″ N,
080°43′18″ W; 117 m; 21 Oct. 2010; L. Martínez 654
(MO, PMA); Coclé del Norte, helipad CR15W; 08°55′26″
N, 080°40′37″ W; L. Martínez 708 (PMA); Westernmost
part of province, site of proposed copper mine (INMET),
lowland forest in Tailings Area; 08°53′16″ N, 080°39′15″
W; 65 m; 10 Apr. 2009; G. McPherson & Jean-Yves
Serein 20833 (MO); Coclé del Norte, área de concesión
Ortiz et al. | Additions to the ora of Panama 617
Minera Panamá; 08°54′37″ N, 080°35′30″ W; O. Ortiz
76 (MO, PMA); Coclé del Norte, bosque húmedo tropi-
cal, área del helipad C14; 08°48′49″ N, 080°46′29″ W;
63 m; 21 Oct. 2010; A. Zapata 2588 (MO, PMA); Área
de concesión del Proyecto Minera Cobre Panamá S.A.,
Valle Grande, antiguo sendero de reubicación de ora;
08°49′21″ N, 080°39′58″ W; 260 m; 16 Aug. 2014; I. Ver-
gara-Pérez 770 (MO, PMA). Figure 5A.
Global distribution. Costa Rica and Panama.
Identi cation. This species is recognized by its leaf
blades with the secondary nerves regularly spaced, con-
spicuously elevated on lower surface, racemose ino-
rescences and owers with two sepals (Morales 2007).
Ouratea rinconensis Whitef. is most similar to O. exi-
pedicellata Dwyer, which diers mainly in having ve
sepals (Whitefoord 1992).
ORCHIDACEAE
Sobralia purpurella Dressler & Bogarín, Orchids (West
Palm Beach) 80: 309 (Dressler and Bogarín 2011).
Figure 2F
New records. Bocas del Toro. Bosque Protector Palo
Seco, sendero El Verrugoso; 08°46′46″ N, 082°10′14″ W;
802 m; 6 Feb. 2013; O. Ortiz et al. 1243 (PMA, UCH).
Figure 5A.
Global distribution. Costa Rica and Panama.
Identi cation. Sobralia purpurella is distinguished by
the following characters: lanceolate-elliptic leaf blades of
15–30 × 1.9–3 cm; inorescence terminal on leafy stems
and also on short, leaess stems; owers with white
sepals, pale pink to purplish petals with an intensely pur-
ple lip (blood red within), undulate and emarginated. In
Panama, S. purpurella is most similar to S. leucoxantha,
but the latter has leaess inorescences usually about as
long as the leafy stems (Dressler and Bogarín 2011).
PASSIFLORACEAE
Passiora macrophylla Spruce ex Mast., J. Linn. Soc.
Bot. 20: 31 (Masters 1883).
Figure 2H
New records. Darién. Pueblo de Bahía Piñas; 7°34′57″
N, 078°10′40″ W; 46 m; 29 Jun. 2015; A. Jiménez 268
(MO, PMA); Área de Manejo Especial de Bahía Piñas,
bosque cercano a Aceite; 7°37′41″ N, 078°11′10″ W; 87
m; 30 Jun. 2018; O. Ortiz et al. 2907 (PMA). Figure 5B.
Global distribution. Colombia, Ecuador, and Panama.
Identi cation. Passiora macrophylla is recognized by
the following characters: weak tree, to 5 m high, stipules
setaceous to triangular, absence of tendrils, leaf blade
unlobed, subpeltate, usually 40–81 cm long. Peduncles
dichotomously or trichotomously branched 1–3 times,
2–12 (–17) owered, 8–25 cm long. Flowers white inside,
tinged pale green outside (Vanderplank et al. 2017). Cur-
rently in Panama there are two other species that may
present arboreal habit, P. tica Gómez-Laur. & L.D.
Gómez and P. pittieri Mast. These two species dier
from P. macrophylla in having smaller blades (up to 50
cm long) and a densely tomentose ovary (vs glabrescent
in P. macrophylla).
PERACEAE
Pera oppositifolia Griseb., Nachr. Königl. Ges. Wiss.
Georg-Augusts-Univ. 1: 181 (Grisebach 1865).
New records. Chiriquí. Cerro Batipa; 08°21′27″ N,
082°14′31″ W; 26 m; 5 Dec. 2003; R. Pérez 1159 (PMA).
Figure 5B.
Global distribution. Costa Rica, Cuba, and Panama.
Identi cation. It is recognized by its opposite leaves,
blades with anastomosing secondary veins which form
several submarginal veins, densely covered with lep-
idote indumentum on the lower surface (González-
Ramírez 2010). Pera oppositifolia represents the second
species of this genus in Panama; previously there were
only records of P. arborea Mutis (Correa et al. 2004).
The latter species diers from P. oppositifolia mainly in
having alternate leaves.
POACEAE
Digitaria costaricensis R. Pohl, Fieldiana, Bot. 38: 5
(Pohl 1976).
New records. Chiriquí. Playa La Barqueta, cerca al
arrozal localizado a 200 m de la playa; J. Polanco 2836
(PMA). Figure 5B.
Global distribution. Costa Rica, Guatemala, and Pan-
ama. Recent data from Mexico (Sánchez-Ken 2017) and
El Salvador (Baldini and Menjívar, in progress) suggest
a more widespread presence of this species in Central
America.
Identi cation. This species is distinguished by having
velutinous leaf blades, non-winged racemes and long
spikelets (3.5–4.2 mm), unbranched lower racemes,
grayish superior lemma and 5–7-ribbed superior glume
(Pohl and Davidse 1994). Digitaria costaricensis is mor-
phological similar to D. aequiglumis (Hack. & Areche.)
Parodi, which diers from D . c ostaricensis in having
densely pubescent foliage, wider leaf blades, a cu-like
rst glume on the spikelet, and longer racemes up to
14 cm. The record from Panama seems to be similar to
those from Costa Rica, because it presents long racemes,
reduced upper owers, and inferior lemma with large
nerves.
Lasiacis ligulata Hitchc. & Chase, Contr. U.S. Natl. Herb.
18: 337 (Hitchcock and Chase 1917).
New records. Bocas del Toro. Bosque Protector Palo
Seco, río Changuinola arriba, Charco La Pava, bosque
trasero de la casa del señor Reynaldo Abrego; 09°09′31″
N, 082°30′35″ W; 424 m; 3 Feb. 2013; O. Ortiz et al. 1216
(FT, PMA); Changuinola, Bosque Protector Palo Seco,
Charco La Pava, Comunidad de Guayacán; 09°09′31″ N,
618 Check List 15 (4)
082°30′35″ W; 424 m; 4 Feb. 2014; O. Ortiz & C. Gal-
da mes 2105 (FT, PMA). Coclé. Región de El Copé, Parque
Nacional Omar Torrijos Herrera, centro de visitantes, sen-
dero la rana dorada; 08°40′05″ N, 080°35′34″ W; 750–790
m; C. Galdames 5901 (PMA, SCZ). Figure 5B.
Global distribution. Bolivia, Brazil, Colombia, Domin-
ican Republic, Ecuador, French Guiana, Guyana, Pan-
ama, Peru, Puerto Rico, Suriname, West Indies and
Venezuela. The specimens cited here represent the rst
records of this species for Central America.
Identi cation. According to Davidse (1978), this spe-
cies is distinguished by having erect stems, hollow or
partially hollow culms, conspicuous ligules of the upper
leaves (usually 2–3 mm long), small blades (usually 7–14
cm long and 1.0–2.2 cm wide), not conspicuously cor-
date; small inorescences (2–21 cm long), long ovoid
panicles (usually 2–17 cm long), completely exserted at
the base. Specimens from Panama match the type mate-
rial from Trinidad (Hitchcock and Chase 1917). This spe-
cies is most closely related to L. maculata (Aubl.) Urb. [=
L. sorghoidea (Desv.) Hitchc. & Chase var. sorghoidea]
from which diers in its less dense and reexed branches,
scanty blade pubescence, and longer membranous ligules.
Lasiacis sloanei (Griseb.) A. Hitchc., Bot. Gaz. 51: 302
(Hitchcock 1911).
(≡) Panicum sloanei Griseb. Flora of the British West Indian Islands,
1864.
New records. Panamá Oeste. Parque Nacional Altos
de Campana, carretera a Chicá; 08°41′13″ N, 079°55′00″
W; 2 Feb. 2013; M. de Stapf & R. Baldini 939 (PM A);
ibid., orillas de la carretera a Chicá, después del mirador;
08°41′22″ N, 079°54′54″ W; 624 m; 16 Jun. 2013; M. de
Stapf & D. Stapf 1073 (PMA). Figure 5B.
Global distribution. Cuba, Haiti, Dominican Repub-
lic, Belize, Caribbean, Mexico, Guatemala, Honduras,
El Salvador, Nicaragua, Costa Rica, Panama, Colombia,
Venezuela, Ecuador. An old specimen was found at the
herbarium MA, collected by L. Neé (“Panama, Neé s.n.”,
MA) during the A. Malaspina’s expedition around the
World (Baldini unpubl. data).
Identi cation. Lasiacis sloanei has a caespitose, clam-
bering, or climbing culm. It is characterized by glabrous
sheaths and blades that are broad (usually 1.7–4.0 cm
wide), shiny, and glabrous except for a line of puberu-
lence or scabridity along the midrib on the upper surface.
This species also has a collar that is densely puberulent
or hispid, and the inorescence is sparsely owered with
large spikelets (usually 4.3–4.8 cm long) born on short
pedicels appressed to the panicle branches (Davidse
1978). This species belong to the Caribbean and Meso-
american Lasiacis group and is closely related to L.
divaricata (L.) Hitchc. The latter species diers from L.
sloanei in having a zigzag pattern of branching, shorter
and narrower leaves, smaller spikelets, culm without
adventitious roots, and reexed panicle branches (gener-
ally at maturity).
PORTULACACEAE
Portulaca papillato-stellulata (Danin & H.G. Baker)
Danin in Danin and Reyes-Betancort, Lagascalia 26: 76
(Danin and Reyes-Betancort 2006).
(≡) Portulaca oleracea L. subsp. papillato-stellulata Danin & H.G.
Baker, Israel J. Bot. 27, 1978.
Figure 1G
New records. Comarca Guna Yala. Isla de Ailigandi;
09°13′40″ N, 078°01′40″ W; 4 m; 20 Jan. 2013; O. Ortiz
& R. Baldini 1129 (FT, PMA). Figure 5B.
Global distribution. Guatemala, Mexico, Liberia, Pan-
ama, and United States.
Identi cation. Danin et al. (1978) stated that the seed
coat characters, seed size and chromosome numbers
are constant diagnostic characters within the Portulaca
oleracea L. polyploid complex. Danin and Reyes-Betan-
cort (2006) reviewed the complex and proposed raising
some of its subspecies to specic rank, which were sub-
sequently included in the treatment developed by Danin
and Raus (2012). Portulaca papillato-stellulata (2n = 54)
is characterized by having seeds of 0.9 × 0.8 mm, star-
shaped testa cells and long rays with terminal papillae
(Rodríguez-Navarro et al. 2009; Danin and Raus 2012).
Portulaca papillato-stellulata and P. oleracea are mor-
phologically very similar (when sterile, almost undistin-
guishable from each other). However, when in fruit, P.
oleracea diers in having smooth testa cells.
RANUNCULACEAE
Clematis populifolia Turcz., Bull. Soc. Imp. Naturali-
stes Moscou 27: 272 (Turczaninow 1854).
New records. Chiriquí. Vicinity of Fortuna Dam. along
trail across valley south of lake, forest; 08°45′04″ N,
082°15′04″ W; 1400 m; 28 Dec. 1986; G. McPherson &
J. Aranda 10204 (MO). Figure 5C.
Global distribution. Colombia, Costa Rica, Ecuador, El
Salvador, Honduras, Nicaragua, Panama, and Venezuela.
Identi cation. It is recognized by its 3-foliolate leaves
with membranous entire leaets, inorescences usu-
ally with umbellate ultimate nodes and 8–25 carpels
(Moreno 1993). According to Moreno (1993), Clematis
populifolia has long been included within C. haenkeana
C. Presl, but the latter diers in having 20–50 carpels
and ve-foliolate leaves.
SALICACEAE
Hasseltia lateriora Rusby, Descr. S. Amer. Pl. 62
(Rusby 1920).
Figure 1E
New records. Darién. Serranía del Majé, Cerro Chucantí,
sendero al helicóptero; 08°47′50″ N, 078°27′42″ W; 900 m;
3 Mar. 2011; R. Flores et al. 574 (PMA). Figure 5C.
Global distribution. Colombia, Ecuador, and Panama.
Identi cation. Hasseltia lateriora is distinguished
619
Ortiz et al. | Additions to the ora of Panama
from the other species by having leaves with short peti-
oles (usually 0.5–3.5 cm long), elliptic, obovate, or rarely
subovate blades, inorescences terminal and also axil-
lary from the upper 3–5 leaf axils, usually 5–9 branches
or owers per umbel, owers with white-villous stami-
nal laments and numerous disk lobes. This species is
similar to H. guatemalensis Warb., which diers in hav-
ing terminal inorescences with usually 3–5 branches
or owers per umbel and few disk lobes (fewer than 16
lobes) (Alford 2006).
SAPINDACEAE
Vouarana anomala (Steyerm.) Acev.-Rodr., BioLlania,
Ed. Espec. 6: 146 (Acevedo-Rodríguez 1997).
(≡) Toulicia anomala Steyerm., Ann. Missouri Bot. Gard. 75, 1988.
New records. Panamá. Cerro Jefe, antes del desvío
hacia Altos de Pacora; 09°12′46″N, 079°22′41″W; 500 m;
7 Sept. 2007; FLORPAN et al. 7192 (PMA). Figure 5C.
Global distribution. Costa Rica, Colombia, Ecuador,
Panama, Suriname, and Venezuela.
Identi cation. This species is distinguished by its arbo-
real habit, pinnately compound leaves, leaets with
domatia on the lower surface (along mid-veins) and ow-
ers with four sepals and petals. Currently, Vouarana
comprises only two Neotropical species, and V. anomala
is the only representative of the genus in Central Amer-
ica (previously reported in Costa Rica) (Morales 2015).
SAPOTACEAE
Pouteria bulliformis Q. Jiménez & T.D. Penn., Novon 7:
169 (Jiménez-Madrigal and Pennington 1997).
New records. Colón. Quebrada López; 09°19′27″ N,
079°47′58″ W; 10 Jun. 2003; R. Pérez & I. Tejada 1049
(PMA); Río Palenque, Nueva Providencia; 09°17′9.60″
N, 079°47′38.78″ W; 23 Aug. 2003; A. Somoza et al.
114 (PMA); Nueva Providencia, cuenca del río
Palenque; 09°16′19″ N, 079°47′48″ W; 57 m; 2 Oct.
2003; R. Pérez et al. 1138 (PMA). Panamá. Parque
Nacional Chagres, Cerro Jefe, sendero de las
Pentagonias; 09°12′40″ N, 079°22′41″ W; 750 m; 3
Jan. 2001; R. Aizprúa et al. B2528 (MO); San Juan de
Pequení; 09°23′49″ N, 079°31′35″ W; 200 m; 25 Mar.
1998; S. Aguilar et al. 979 (PMA). Fig-ure 5C.
Global distribution. Colombia, Costa Rica, Nicaragua,
and Panama.
Identi cation. This species is characterized by having
densely ferruginous-tomentose young shoots and ino-
rescences, bullate blades, rounded apically (sometimes
short cuspidate) and tomentose (at least on the veins)
on lower surfaces. Pouteria bulliformis is similar to P.
simulans Monach., but the latter diers in having short-
pubescent young shoots and inorescences, non-bullate
blades, narrowly short-attenuate apically that are some-
what glabrous on lower surfaces (Jiménez-Madrigal and
Pennington 1997).
SOLANACEAE
Cuatresia trianae Hunz., Caldasia 15: 143 (Hunziker
1986).
Figure 1H
New records. Coclé. Vicinity El Copé, 5–6 mi. N of
El Copé, along trail which leads into the lowlands from
old Riviera saw works area; 08°40′14″ N, 080°35′34″
W–08°41′18″ N, 080°35′58″ W; 600–800 m; 8 Jul. 1994;
T.B. Croat & G.H. Zhu 77214 (MO). Colón. MPSA min-
ing site, forested slopes along Quebrada Petaquilla;
08°49′16″ N, 080°40′26″ W; 150 m; 2 Sept. 2014; G.
McPherson et al. 21565 (MO). Figure 5C.
Global distribution. Colombia, Ecuador, Panama, and
Peru.
Identi cation. It is recognized by having strongly roug h-
ened blades, dense inorescences with few owers (5–10
owers), densely pubescent owers with lobed calyx,
bell-shaped corolla (with conspicuous inter-petal mem-
brane) and non-exerted stamens (Hunziker 1986; Canal
and Orozco 2010). In Panama, Cuatresia trianae could be
confused with C. exiguiora (D’Arcy) Hunz., but the lat-
ter diers in having lax inorescences, glabrous owers
and ellipsoid berries (Canal and Orozco 2012).
VIOLACEAE
Paypayrola confertiora Tul., Ann. Sci. Nat., sér. 3 7:
373 (Tulasne 1847).
Figure 2G
New records. Coclé. Parque Nacional General de Divi-sion
Omar Torrijos Herrera, La Rica, bosque cercano al río Juan
Julio; 08°42′59″ N, 080°35′30″ W; 240 m; 23 Jul. 2013; O.
Ortiz et al. 1475 (MO, PMA). Colón. Distrito de Donoso,
área de concesión de Minera Panamá, Campa-mento 600,
área 4; 08°49′26″ N, 080°39′32″ W; 230 m; 7 Jun. 2012;
J.F. Carrión 785 (PMA); ibid., Coastal Road 17k-700; 08°
56′08″ N, 080°41′32″ W; 61 m; 13 Nov. 2013; R. Flores
3424 (PMA); Teck Cominco Petaquilla min-ing concession,
slopes along exploration road; 08°50′22″N, 080°38′51″ W;
184 m; 15 Sept. 2007; G. McPherson 19558 (MO); ibid.,
ridgetop forest, near transect C002; 08°50′22″ N, 080°38′
51″ W; 205 m; 18 Sept. 2007; G. McPherson 19679
(MO); ibid., ridgetop forest along road; 08°49′29″ N, 080°
40′12″ W; 323 m; 1 Dec. 2007; G. McPherson & H. van der
Wer 19910 (MO); ibid., for-est along ridge road; 08°49′22″
N, 080°39′32″ W; 300 m; 25 Feb. 2008; G. McPherson
& M. Merello 20241 (MO); Site of proposed copper
mine (MPSA), forested slopes; 08°48′27″ N, 080°36′
20″ W; 100 m; 13 Dec. 2009; G. McPherson & M.
Merello 21283 (MO); Botija, Carretera Pionera,
similar to prairie clay; 08°50′06″ N, 080°39′17″W; 137
m; 11 Sept. 2012; H. van der Wer et al. 24441 (MO,
PMA). Comarca Guna Yala. Río Cangandí, hills W
of river S of confluence with río Titamibe; 09°27′30″
N, 079°07′00″ W; 50–150 m; 27 Jan. 1985; G. de Nev-
ers et al. 4669 (MO); Between río Diablo & río Acuatí
620 Check List 15 (4)
near Narganá; 09°26′34″ N, 078°35′26″ W; 3 Nov. 1967;
J. Duke 14895 (MO); Cordillera frente a Isla Narganá;
09°22′ N, 078°34′ W; 65 m; 12 Aug. 1994; C. Galdames
1587 (MO. PMA); Veci ndad de R ío Dia blo, desde el cam-
pamento Duque Sui hasta Isper Yala; 09°22′ N, 078°35′
W; 70–100 m; 3 Jul. 1992; Herrera et al. 1210 (MO);
ca ½ way between the continental divide and the Atlan-
tic coast opposite Cartí; 09°22′ N, 078°58′ W; 200 m;
23 Feb. 1973; H. Kennedy 2616 (MO). Panamá . 12 mi.
on Cartí Road from the Interamerican Highway, then 3
hours walk along road to Atlantic side; 09°23′ N, 08°57′
W – 09°27′ N, 078°57′ W; 30 m; 15 Feb. 1980; T. Anto-
nio 3782 (MO); El Llano-Cartí Road, km 26.5. Trail NE
from road, premontane wet forest, assoc.: Ormosia coc-
cinea, Gnetum leyboldii; 09°22′ N, 078°58′ W; 175 m; 9
Mar. 1985; G. de Nevers et al. 5078 (MO); El Llano-Cartí
Road, km 26.5, trail to río Cartí Chico and up ridge on E
side; 09°22′ N, 078°58′ W; 200 m; 7 Apr. 1985; G. de
Nevers et al. 5236 (MO); El Llano-Cartí Road, km 26.5,
tropical moist forest, assoc.: Bonafousia undulata, Atta-
lea allenii, Theobroma bernoulii; 09°22′ N, 078°58′ W;
200 m; 14 Apr. 1985; G. de Nevers et al. 5363 (MO);
El Llano-Cartí highway, 14–17 km north of El Llano;
09°17′20″ N, 078°55′22″ W; 335 m; 13 Feb. 1973; R.
Dressler 4270 (MO); Road from El Llano to Cartí, 14.8
km north of the Panamerican Highway; 09°17′18″ N,
078°56′08″ W; 300–500 m; 3 Sept. 1977; J. Folsom &
P. Maas 5221 (MO); 16 km above Pan-Am Highway
on road from El Llano to Carti-Tupile, near campsite;
09°17′50″ N, 078°56′15″ W; 350–400 m; 21 Feb. 1973,
H. Kennedy 2542 (MO); El Llano-Cartí road, km 15,
rain forest; 09°19′40″ N, 078°57′05″ W; 370 m; 3 Sept.
1977; P. Maas 2794 (MO); Along El Llano-Cartí road,
Forested slopes; 09°15′ N, 079°00′ W; 400 m; 24 Nov.
1985; G. McPherson 7586 (MO); Along El Llano-Cartí
road, forested slopes; 09°15′ N, 079°00′ W; 400 m; 24
Nov. 1985; G. McPherson 7588 (MO); On El Llano-Carti
road, near Nusagandi, along trail to waterfall; 09°15′ N,
079°00′ W; 250 m; 1 Nov. 1992; G. McPherson & Rich-
ardson 16007 (MO). Figure 5C.
Global distribution. Brazil, Colombia, French Guiana,
Guyana, Panama, Suriname, and Venezuela.
Identi cation. It is recognized by having elliptic to
obovate-elliptic blades, long-attenuate at base, petioles
5–9 mm, congested inorescences and owers with yel-
low petals (Aymard et al. 2014). In Panama, P. conferti-
ora could be confused with P . l ongifolia Tul., but the
latter diers in having obovate to obovate-lanceolate
blades, lon ger petioles ( >12 mm) and l oose inorescences.
Information gaps on plant collections in Panama
Geographically, the west of the country (i.e. the provinces
of Chiriqui, Bocas del Toro, and Comarca Ngäbe-Buglé),
has been relatively well collected (Fig. 6), especially in
the mid- and high-elevation areas, such as the Boquete
district, the corregimiento of Volcán, and the Fortuna
Forest Reserve (Reserva Forestal Fortuna). However,
there are still areas with few collections, primarily in
lowland forest relics in the southern part of Chiriquí.
Bocas del Toro Province has several relatively well-
collected locations, both in the lowlands (Changuinola
and Chiriquí Grande districts) and at mid and high ele-
vations (La Amistad International Park), but there are
still important sites with few collections, mainly in the
protected area of the Bosque Protector Palo Seco, which
includes some areas of the Comarca Ngäbe-Buglé. The
Comarca Ngäbe-Buglé presents many unexplored locali-
ties, and the existing collections of this region are con-
centrated in Cerro Colorado, Ratón, Cerro Santiago,
Escudo de Veraguas island (Degó in the indigenous lan-
guage), and the Damani Guariviara wetlands.
The central region of the country, which includes
the provinces of Veraguas, Herrera, Los Santos, Coclé,
Panamá Oeste, and certain parts of western Colón, has
at least six areas with more than 1700 botanical records
(Fig. 6). Collections in the Province of Verag uas are con-
centrated to the north, in the protected area of Santa Fe
and to the south, in Coiba National Park, while the rest
of the province is still poorly explored. Similarly, despite
some collecting eorts in Santa Fe National Park, this
protected area is currently underexplored. The Azu-
ero Peninsula, which includes the provinces of Herrera,
Los Santos and the south-eastern part of the Province of
Veraguas, has some scattered collections, mainly in pro-
tected areas, but other important sites such as the moun-
tains of Cerro Hoya National Park remain very poorly
known. In Coclé province, collections are concentrated
in the Antón district (Valle de Antón) and the Omar Tor-
rijos Herrera National Park, with still-unexplored areas
in the north-east of the province (Penonomé district),
such as Chiguirí arriba and the Río Indio area. A large
number of recent collections are from the Province of
Colón, most of them concentrated in the western part of
Donoso district. In the Province of Panamá Oeste, botan-
ical records are located mainly in the Altos de Cam-
pana National Park, but there are collection gaps in the
northwest (Río Indio area and the corregimientos of Ciri
Grande, Cirí de Los Sotos, and Ollas Arriba).
The Canal area (the old Canal Zone), which includes
the central part of Colón and the western part of the
province of Panama, undoubtedly represents the portion
of the country with the most botanical records (Fig. 6).
The protected areas bordering the Panama Canal, such as
Barro Colorado Island, Bosque Protector San Lorenzo,
and the National Parks of Camino de Cruces, Altos de
Campana and Soberanía, have been widely collected.
In the eastern part of the country, in the province of
Panamá there are many botanical records, but these are
concentrated in the areas near Panama City, Chagres
National Park (including Cerro Azul and Cerro Jefe),
and the central part of the province, mainly in the district
of Chepo (along the road from El Llano to Cartí); the rest
of the eastern part is still virtually unexplored. On the
other hand, the eastern part of the province of Colón has
been widely explored in the Portobelo area, but many
Ortiz et al. | Additions to the ora of Panama 621
parts of the province are still undercollected, primar-
ily in the eastern part near the Comarca Guna Yala. The
remaining sites of the eastern part of Panama (the least
explored region of the country) have very few botani-
cal collections. Only nine localities have more than 300
collections, seven of them in the province of Darién (for
example, the Pirre and Cana Mountains) and two in the
Comarca Guna Yala.
Discussion
In summary, most collections have been made in the
Canal area and the western part of Panama (specically
in Chiriquí Province). Furthermore, there are only ve
locations with more than 2500 botanical records (Fig. 6).
Many of the sites identied here as having few collec-
tions correspond to areas with high endemism and diver-
sity and are important biodiversity conservation sites
(Myers 1969; Lewis 1971; Powell et al. 2018a, 2018b).
In recent years, many new records and new species
have been added to the Panamanian ora. This suggests
that there are still information gaps and are probably due
to the following reasons:
• Panama is a small country (75,845.072 km2) and most of
its territory is anthropic (towns, highways, roads, energy
plants, cultivated areas, etc.), so the remaining fully nat-
ural areas are currently very reduced and highly frag-
mented. According to historical records, it is estimated
that in 1850 forest covered 91% of Panama (Arias 2004);
however, there has been rapid deforestation throughout
the 20th and into the 21st centuries (Killip 1919; Por-
ter 1970; Croat 1972; Kapos et al. 2015). In 1947, 1970,
and 1992, the forests covered 70% (53,091 km2), 53%
(40,816 km2), and 49.3% (36,951.60 km2), respectively, of
the total territory (Garver 1947; Lamb 1953; Hartshorn
1981; ANAM 2010; Kapos et al. 2015). Subsequently, it
was reported that the deforestation rate was about 413
km2 per year, and 134 km2 per year between 2000 and
2008, which caused the total forest cover to decrease to
33,507 km2 in 2000, and to 32,433 km2 in 2008 (Verg-
ara-Asenjo and Potvin 2014). It is likely that due to the
continuous reduction of the forest cover in the country
during the last 50 years, botanical eorts in Panama
have concentrated mainly in the protected natural areas,
which occupy approximately 31.8% (ca 24,901 km2) of
the national territory (ANAM 2010).
• The collection sites have probably been the same through-
out the history of botanical collecting in Panama. In fact,
many sites with considerable botanical records identied
in this work coincide with the historically more-visited
sites (cf. Dwyer 1964, 1968a, 1968b, 1985; Lewis 1968,
1971; Bogarín et al. 2014a). All these sites have in com-
mon easy access, being surrounded by highways, roads,
cities and towns. In the past, several botanists had already
noticed that one of the most important reasons related to
the lack of knowledge of the ora of Panama is the inac-
cessibility of many mountainous areas far from roads
(Dressler 1972; Lewis 1971; Dwyer 1985). This suggests
that there are still information gaps, particularly in those
regions currently undercollected. For example, 31 of the
Figure 6. Main areas with gaps of information on plant collections in Panama.
622 Check List 15 (4)
46 new records included in this work were recently col-
lected (between 2007 and 2016) in a poorly explored area
of Colón Province. These collections are part of a MBG
Project that consists in documenting the ora of Donoso
district, in which they have collected about 1400 botani-
cal specimens and identied more than 30 new species
(TROPICOS 2019b).
• The are few resident Panamanian collectors. Histori-
cally, foreign botanists made most of the collections in
Panamanian territory over short periods of time (Dwyer
1964, 1985; Dressler 1972). Dressler (1972) emphasized
the importance of training local resident collectors in
Panama, arguing that they are more familiar with the
local ora and would avoid collecting in the same areas
repeatedly, thereby increasing the possibility of nding
novelties. Expert collectors (sometimes referred to as
“mega-collectors”) have proven successful in the search
for new species for science, mainly due to the experience
of having collected in several places (Bebber et al. 2012;
Whiteeld 2012). Currently in Panama, there are very
few botanists who are exclusively dedicated to collecting
plants, and local herbaria do not have sucient budgets
to hire botanists dedicated to research on the local ora.
The collections made recently are mainly due to ecolog-
ical or oristic studies related to projects of economic
or social interest. Experienced general plant collectors
are currently few in Panama, even though, in order to
ascertain the biodiversity importance of the undercol-
lected or poorly known areas discussed and their vul-
nerability, additional oristic and taxonomic studies are
needed. Consequently, the training of local botanists is
necessary as well as the collaboration between local and
international specialists. Furthermore, a detailed map-
ping of Panama’s biodiversity (like the one carried out in
Costa Rica by INBio) is clearly needed. Another factor
limiting an integrated knowledge of the Flora of Panama
is the lack of an updated Flora of Panama, representing
the current state of oristic knowledge, rather than that
of the last decades of the last century.
In the name of the common good, we hope that these
suggestions will be considered by local government o-
cials and institutions, such as the Secretariat for Science,
Technology and Innovation (SENACYT), Ministry of
Environment of Panama, University of Panama (UP),
UNACHI, STRI, Institute of Scientic Research and High
Technology Services (INDICASAT), National Associa-
tion for the Conservation of Nature (ANCON), Techno-
logical University of Panama (UTP).
Acknowledgements
We oer our gratitude to all collectors (Adrián Jiménez,
Alex Espinosa, Alicia Ibáñez, Alvin Zapata, Barry Ham-
mel, Blanca Araúz, Carmen Galdames, Carlos Guerra,
Christel Ramos, Edwin Tyson, Greg de Nevers, Guang-
hua Zhu†, Helen Kennedy, Henk van der Wer, Heraclio
Herrera, Irving Vergara, James Duke†, James Folsom,
Jean-Yves Serein, Jorge Aranda, José González, José De
Gracia, José Polanco, Juvenal Batista, Karina Víquez,
Laurencio Martínez, Mary Merello, Michael Gra-
yum, Nelson Jaén, Nicolás Guerrero, Paul Maas, Rob-
ert Dressler, Rolando Pérez, Thomas Antonio, Thomas
Croat, and Yeleinshka Yaleman). We also acknowledge
the curators that identied the material, as well as the her-
baria that allowed access to their databases and collec-
tions (PMA, MO, SCZ, and UCH). We are also grateful
to the following institutions and companies for their help
in dierent ways with eldwork conducted by us: Uni-
versity of Panama (UP), University of Florence (UNIFI),
Missouri Botanical Garden (MBG), Minera Panamá
S.A. (MPSA), Adopt a Panama Rainforest Association
(ADOPTA), Tropic Star Lodge (Bahía Piñas, Darién), and
Hidroecológica del Teribe S.A. Lastly, we oer thanks
to Dr Oris Rodríguez for providing information on the
emergence of the Isthmus of Panama; Dr Julio Castillo,
Prof. Jerry Harrison, Dr Alicia Ibáñez, Dr Iván Valde-
spino, and the reviewers, Dr Michael Grayum and Dr
Álvaro Idárraga, for their important suggestions and cor-
rections. The UNIFI is acknowledged for supporting the
publication by a funding to RMB (Ex 60% fund 2018).
Authors’ Contributions
OOO collected and identied part of the material, made
part of the identication notes and wrote the manuscript.
RF collected and identied part of the material and made
part of the identication notes. GMP collected and iden-
tied part of the material, contributed to the design and
implementation of the research, and reviewed all manu-
script. JFC collected and identied part of the material
and made part of the identication notes. ECP prepared
the maps and made part of the identication notes. RMB
supported the publication through UNIFI, collected and
identied part of the material, made part the identi-
cation notes, co-wrote the discussion and reviewed all
manuscript.
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