Corresponding author: Daniel Santamaría-Aguilar (
Academic editor: Thomas L.P. Couvreur
With about 70 species
Santamaría-Aguilar D, Lagomarsino LP (2022) New Species of
List of species of
Species | Country of original material | Region | Distributions |
---|---|---|---|
Ecuador | S | Ecuador. | |
Peru | S | Peru. | |
Brazil | S | Colombia ( |
|
Costa Rica | C | Costa Rica. | |
Ecuador | S | Ecuador. | |
Panama | C | Costa Rica, Panama. | |
Ecuador | S | Ecuador. | |
Brazil | S | Colombia ( |
|
Colombia | S | Colombia, Ecuador | |
Venezuela | S | Colombia, Ecuador, Peru, Bolivia, Brazil, Venezuela, Guyana, Suriname. | |
Brazil | S | Colombia (Aymard C. et al. 2020), Brazil. | |
Brazil | S | Colombia ( |
|
Costa Rica | C | Costa Rica, Panama. | |
Brazil | S | Colombia, Peru, Brazil | |
Colombia | S | Colombia. | |
Brazil | S | Brazil. | |
Peru | S | Peru. | |
Brazil | S | Colombia ( |
|
Brazil | S | Colombia ( |
|
Ecuador | S | Colombia, Ecuador. | |
Brazil | S | Colombia, Ecuador, Peru, Bolivia (Achá & Liesner 2014), Brazil, Venezuela (Aymard C. et al. 2007). | |
C, S | Panama, Colombia, Ecuador, Peru, Bolivia, Brazil, Venezuela, Guyana, French Guiana, Suriname. | ||
Ecuador | S | Colombia, Ecuador, Peru, Brazil. | |
Brazil | S | Colombia, Ecuador, Peru, Bolivia, Brazil. | |
Panama | C | Costa Rica, Panama. | |
Brazil | S | Brazil. | |
Guatemala | C, M | Mexico, Guatemala, Honduras. | |
Brazil | S | Brazil. | |
Costa Rica | C, M | Mexico, Guatemala, Honduras, Belize, Nicaragua, Costa Rica, Panama. | |
French Guiana | S | Brazil, French Guiana. | |
Panama | C | Costa Rica, Panama. | |
Peru | C | Colombia, Peru, Bolivia, Brazil, (Achá & Liesner 2014) | |
Colombia | S | Colombia. | |
Brazil | S | Brazil. | |
Brazil | S | Colombia, Peru, Brazil. | |
Panama | C | Panama. | |
French Guiana | S | Colombia ( |
|
Colombia | S | Colombia, Venezuela. | |
Brazil | S | Peru (Vásquez M. et al. 2018), Brazil, Venezuela ( |
|
Peru | S | Colombia, Ecuador ( |
|
Costa Rica | C | Costa Rica, Panama. | |
Brazil | S | Colombia ( |
|
Belize | C | Honduras, Belize, Nicaragua, Costa Rica, Panama. | |
Brazil | S | Colombia ( |
|
Panama | C | Costa Rica (cf.), Panama. | |
Brazil | S | Colombia, Peru, Brazil. | |
Brazil | S | Brazil. | |
Panama | C | Panama. | |
Peru | S | Peru. | |
Brazil | S | Colombia ( |
|
Peru | S | Peru. | |
Peru | S | Colombia, Ecuador, Peru, Bolivia (Achá & Liesner 2014), Brazil, Venezuela. | |
Peru | S | Colombia, Ecuador, Peru, Bolivia, Brazil. | |
Brazil | S | Colombia ( |
|
Peru | S | Peru. | |
Ecuador | S | Colombia, Ecuador. | |
Brazil | S | Brazil, Venezuela. | |
Honduras | C | Honduras. | |
Colombia | S | Colombia, Venezuela. | |
French Guiana | C, S | Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, Brazil, Venezuela, Guyana, French Guiana, Suriname. | |
Brazil | S | Brazil. | |
Venezuela | S | Venezuela. | |
Brazil | S | Brazil. | |
Suriname | S, A | Colombia, Ecuador, Peru, Bolivia, Brazil, Venezuela, French Guiana, Guyana, Suriname, Antilles. | |
Colombia | S | Colombia. | |
Brazil | S | Brazil. | |
Brazil | S | Brazil, Venezuela, Suriname. | |
Peru | S | Peru. | |
Ecuador | S | Ecuador, Brazil. |
Geographic distribution of
Morphological characters of
Fruit morphology of
The genus has also been well-studied by botanists and ecologists as it is a tractable system for studying seed dispersal (e.g.,
The last complete taxonomic revision of
During a review of herbarium material from South America as part of the
Comparation of pattern of the lateral veins and trichomes on abaxial leaf surface (below) of the newly described species of
Below, we describe 10 new species of
Approximately 3000 physical herbarium specimens from Meso and South American
Species descriptions are based primarily on herbarium specimens. If necessary and material permitted, flowers from herbarium specimens were rehydrated before measurement. A ruler was used to measure leaves and inflorescences; a digital Neiko caliper was used to measure fruits and seeds, as well as the thickness of the twigs, petioles and peduncles; and, finally, flowers, trichomes and thickness of the pericarp were measured with a micrometer calibration tool (1div = 1 mm) under a dissecting stereoscope (Bausch & Lomb).
Specimens cited are listed first by country. Within a country, specimens are listed alphabetically by major division and then alphabetically by department, province or state and, finally, in alphabetical order by the collector’s surname. When the coordinates and/or elevation were not included on the herbarium label, but were present in the TROPICOS database, the values from TROPICOS are included. Dot-distribution maps were compiled from studied specimens and generated with SimpleMappr (
Distribution, habitat, phenological data, common name and uses, flower and fruit colors, habit, bark, and exudate data were obtained from herbarium labels.
In the nomenclatural section for each new species, we cite both accession numbers and barcodes when present.
The preliminary conservation status of each new species was assessed using quantitative criteria recommended by the IUCN Red List (
Comparison of fruits of newly described
The specific epithet honors the Aguaruna people, who live in the area where this species was collected.
Staminate buds and flowers of
No common names or uses are mentioned among the herbarium specimens observed.
Most of the specimens of
The only seed that could be measured is from one of the two fruits of the specimen
The specimen
It is a great pleasure to dedicate this new species to the Ecuadoran botanist Álvaro Javier Pérez Castañeda. He is an excellent botanist, collector and expert of the flora of Ecuador, especially the flora of Yasuní. Among other contributions, he has described at least 26 from different angiosperm families (e.g.,
Only a single studied specimen of
Guangare macho (Ecuador;
Herbarium specimens of
Comparison of
Morphological character |
|
|
|
|
---|---|---|---|---|
Leaf blade size, and pubescence abaxially | 26–28.5 × 12.1–14 cm; densely pubescent (Fig. |
25–29.7 × (9.2–) 10.8–15 cm; densely pubescent (Fig. |
20–40 × 7–11 cm†; sparsely pubescent (Fig. |
(14–) 18.2–42.5 × (4.1–) 7.3–14.2 cm; sparsely pubescent |
Spaced lateral veins | (2–) 2.4–2.7 cm apart | 1.9–2.5 cm apart | 0.8–1.5 cm apart | 1.7–3 cm apart |
Fruit size, and pubescence | 4.3–4.5 × 3–3.6 cm; with persistent trichomes (Fig. |
2.7–3.2 × 1.7–2.2 cm; the trichomes do not fall easily like dust (Fig. |
2.7–3.3 × 2–2.3 cm†; with caducous trichomes (Fig. |
(2.7–) 3.5–4.5 × (1.9–) 2.3–2.9 cm; densely tomentose, with persistent trichomes (Fig. |
Pericarp thickness | 2.8 mm | 1.2–1.6 mm | 1.8–3 mm | (2.7–) 3–4.7 mm |
Seed | 2.7 × 1.6 cm | 2.2–2.4 × 1.3–1.4 cm | 2.2–2.5 × 1.5–1.7 cm† | 2.5–2.8 × 1.5–1.7 cm |
The holotype deposited at Missouri Botanical Garden (MO) represents a single collection mounted on two sheets that are clearly labeled (i.e., “Sheet 1 of 2” and “Sheet 2 of 2”) as being parts of the same specimen (ICN Art. 8.3;
Leaf size and the number of lateral veins may be higher than presented here, as most of the material examined had broken leaves.
The collection
The fruit specimen
The two collections of
The specific epithet makes reference to the Bombuscaro River in the Podocarpus National Park of southern Ecuador, where most of the collections of this new species were made.
Staminate flowers of
None recorded.
Comparison of
Morphological character |
|
|
|
|
|
|
---|---|---|---|---|---|---|
Leaf base | Cordate | Usually deeply cordate to truncate (obtuse) | Truncate to subcordate, rarely deeply cordate | Deeply to shallowly cordate | Cordate to rounded | Deeply cordate |
# of lateral veins | (13–) 18–23 | 11–28 | (13–) 18–24 | 17–30 | (10–) 12–18 | 14–19 |
Perianth staminate long and trichomes inside | 2.8–3.4 mm; densely pubescent | 1–2.1 mm; glabrescent | 1.5–2.8 mm; almost glabrous | 2.3–3.2 mm; puberulent | 3–3.5 mm; tomentose (in pistillate flowers) | 2–2.3 mm long; pubescent |
Long filament column, and trichomes | ca. 1.4 mm; slightly pubescent at the base | 0.2–0.6 mm; glabrous | (0.4–) 0.5–0.8 mm; glabrous | 0.4–0.6 mm; glabrous | 0.4–0.7 mm; trichomes not described | 0.3–0.4 mm; glabrous |
Fruit size, and trichomes | 2.4 × 1.8–1.9 cm; densely tomentose, falling easily to the touch as dust (Fig. |
2.5–3 × 1.2–2.5 cm; tomentelous to glabrescent, not falling easily to the touch as dust (Fig. |
2–2.6 × 1.4–1.7 cm; densely tomentose, falling easily to the touch as dust (Fig. |
2–2.8 × 1.5–2.2 cm; glabrescent (Fig. |
2.8–3.6 ×1.8–2.4 cm, densely tomentose | 0.8–1.1 × 0.7–1 cm; minutely pubescent, not falling easily to the touch as dust |
Carina on the line of dehiscence | Slightly carinate | Carinate | Slightly carinate | Conspicuously carinate | Slightly carinate | Not carinate |
Pericarp thick | 1.6–2.2 mm | 0.5–5 mm | 1.5–2.3 mm | 0.5–1.5 mm | 2–3 mm | 0.5 mm |
Other species with similar leaf morphology (i.e., a usually deeply cordate base and well-separated lateral veins) are:
Additionally, compared to the species with with it can be confused,
The specimen
Two collections from Cajamarca Department, Peru collected at 1500–1700 m elevation,
The specific epithet of the new species refers to the Bajo Calima region (Valle del Cauca department, Colombia), the region where most specimens of this new species come from.
Staminate flowers of
Cuangare (Colombia;
Herbarium specimens of
Comparison of
Morphological character |
|
|
---|---|---|
Leaf blades size, and pubescence abaxially | 19–25 × 7.3–11 cm; densely pubescent (Fig. |
20–40 × 7–11 cm†; sparsely pubescent (Fig. |
Spaced lateral veins | 1.3–2.3 cm apart | 1.7–3 cm apart |
Fruit shape, and long | Globose, 2.8–3.1 cm (Fig. |
Ellipsoid, (2.7–) 3.5–4.5 cm (Fig. |
Pericarp thickness | ca. 3.8–4 mm | (2.7–) 3–4.7 mm |
Seed size | ca. 2.5 × 2.1 cm | 2.2–2.5 × 1.5–1.7 cm† |
Habitat | Lowland rain forest, Colombia (Valle del Cauca), and Ecuador (Esmeraldas), at 20 to 260 m elevation | Montane forests, Andes of Colombia (Boyacá), at 1100 m elevation† |
The first collection of
Comparison of
Morphological character |
|
|
---|---|---|
Leaf blade size, and base | 19–25 × 7.3–11 cm, obtuse | (15–) 20–60 × 10–16 cm; (usually) deeply cordate to truncate (obtuse) |
# of lateral veins | ca. 12–13 | 11–28 |
Length of staminate inflorescences | 6–8 cm | 6–30 cm |
Staminate perianth long, and internal surface | 2.5–3.5 mm; densely pubescent | 1–2.1 mm; glabrescent |
Long filament column | 0.4–0.6 (–0.8) mm | 0.2–0.6 mm |
Long anthers | 0.6–1 (–1.2) mm | 0.4–0.5 mm |
Fruit size | ca. 3.3–4.2 × 2.8–3.1 cm (Fig. |
2.5–3 × 1.6–1.5 cm (Fig. |
The specimens
Costa del Pacífico, Río Cajambre, Silva, 5–80 m, 5–15 May 1944 (♂ fl),
The specific epithet honors Álvaro Cogollo Pacheco, the Colombian botanist who collected most known specimens of this new species as well for his valuable contribution to our knowledge of the Colombian flora. We celebrate his important contributions to botany, epitomized by his numerous collections, ~48 of which now represent type specimens (
Specimens with staminate flowers of
Sebo (Colombia;
Prior to our study, most of the specimens with fruits of
Comparison of
Morphological character |
|
|
---|---|---|
Petiole | 1.7–2.4 × (0.28–) 0.32–0.6 cm | 1.5–2.3† × 0.18–0.23 cm |
Leaf blade size, and pubescence abaxially | 25–29.7 × (9.2–) 10.8–15 cm; densely pubescent (Fig. |
20–40 × 7–11 cm†; sparsely pubescent (Fig. |
Spaced lateral veins | 1.9–2.5 cm apart | 1.7–3 cm apart |
Fruit size, and pubescence | 2.7–3.2 × 1.7–2.2 cm; with persistent trichomes (Fig. |
(2.7–) 3.5–4.5 × (1.9–) 2.3–2.9 cm; with caducous trichomes (Fig. |
Pericarp thickness | 1.9–2.1 mm (on the thickest side) | (2.7–) 3–4.7 mm |
A second
Comparison of
Morphological character |
|
|
---|---|---|
Leaf blade wide | (9.2–) 10.8–15 cm | 5–7.2 (–11.7) cm |
Lateral veins | ca. 12–14 per side, 3 veins per 5 cm, spaced 1.9–2.5 cm | 16–19 per side, 3–4 (–6) veins per 5 cm, spaced 1.1–1.8 (–2.1) cm |
Staminate peduncle | 2.3–2.5 cm long | 0.6–1.8 (–2.8) cm long |
Filament column | 1.2–1.4 mm long | 0.6–0.7 (–0.9) mm long |
Anthers | 1.1–1.2 mm long | 1.2–1.6 mm long |
Fruit pubescence | With an inconspicuous layer of trichomes, persistent (Fig. |
With a conspicuous layer of trichomes, caducous, that fall like dust (Fig. |
Pericarp thickness | 1.9–2.1 mm (on the thickest side) | ca. 2.4 mm |
The specimens
The most distinctive characteristic of
Diversity of fruits from
The specific epithet is taken from the common name on the label of
Staminate flowers of
Cumala negra (Peru;
Comparison of
Morphological character |
|
|
|
|
---|---|---|---|---|
Leaf blade upper surface | Glabrous | Pubescent | Glabrous | Glabrous |
No. of lateral veins | 49–65 | 45–60‡‡ | (32–) 40–50 | 40–60‡‡ |
Staminate perianth texture, and pubescence on external surface | Subcarnose, without lines or dots, densely pubescent on all the surface | Submembranous, with lines, pubescent | Subcarnose, without lines or dots, densely pubescent on all the surface | Membranous, with lines, glabrous or pubescent only at the base and apex |
Filament column length | 0.5–0.7 mm | 0.3–0.4 mm‡‡ | 0.9–1.3 mm | 0.3–0.5 mm |
Anther length | 0.7–0.9 mm | 0.5–0.6 mm‡‡ | 0.8–0.9 mm | 0.3–0.4 mm |
Fruit size and apex | 3–3.5 × 2.3–2.9 cm, obtuse (Fig. |
2.7–3.5 × 1.7–2.2 cm‡‡, acute to apiculate (Fig. |
4–5.7 × 2–2.9 cm, acuminate to rostrate (Fig. |
2–3 × 1.5–2.5 cm, apiculate‡‡ (Fig. |
Pericarp thickness | 3–6 mm | 4–5 mm | 3–6 mm | 1.5–4 mm |
The fruits of
The type specimen of
Comparison of abaxial surface, veins, and trichomes of
The specific epithet of the new species comes from the word
Staminate flowers of
Common names include in Colombia: sangre de toro (
Of morphology apparent in herbarium specimens, the leaf blades and some flower and fruit features of
Comparison of
Morphological character |
|
|
---|---|---|
Leaf blades size, base and shape | (10.5–) 23.5–46.5 × (6.7–) 9.5–13.5 (–19.7) cm; base truncate to subcordate, rarely deeply cordate; lanceolate to obovate | 11.5–29.5 (–34) × 6.8–12.5 cm; base attenuate to acute; obovate-elliptic. |
Staminate perianth | Without resinous punctuations | With resinous punctuations |
Infructescence peduncle | ca. 1.4–3.5 (–5) cm long | 0.7–1.3 cm long |
Fruits | 2–2.6 × 1.4–1.7 cm, without stipe, the base truncate to obtuse (Fig. |
1.3–2.3 ×x 0.8–1 cm, shortly stipitate, the base rounded (Fig. |
Pericarp thickness | 1.5–2.3 mm | 1–1.2 mm |
Seed | 2–2.3 × 1.1–1.3 cm | 1.7–1.9 × 0.6–0.7 cm |
The majority of specimens cited here as
The most striking character of
The specific epithet honors Theodore A. Parker III (1 Apr. 1953–3 Aug. 1993), renowned and talented ornithologist and research associate of the LSU Museum of Natural Science. Parker died in a plane crash on August 3, 1993 while surveying a remote forest in Ecuador, along with three other people: Raul Mortensen (the pilot), Eduardo Aspiazu (ecologist), and Alwyn H. Gentry (botanist).
Staminate flowers and fruits of
Common names include banderín and rrohuatquech (Peru;
Comparison of
Morphological character |
|
|
---|---|---|
Leaf blade size | 19.7–24 × 5.2–6.4 cm | 15.7–27.5 × 4.4–6.6 cm |
Spaced lateral veins | 1.5–1.7 (–2) cm apart | 0.9–1.5 (–1.9) cm apart |
Staminate perianth lobes | ca. 0.9 × 0.5 mm | 0.7–1 × 0.5–0.8 mm |
Filament column | 0.2–0.4 mm long, and ca. 0.1 mm wide at the base | 0.3–0.5 mm long, and ca. 0.2–0.3 mm wide at the base |
Anthers long | 0.5–0.7 mm | 0.5–0.6 mm |
Apiculus long | ca. 0.8 mm | ca. 0.1 mm |
Fruit size | 3–3.6 × 2.8–3.1 cm | 3.4–4.2 × 3.1–4 cm |
Fruit carina | ca. 0.4 cm long (Fig. |
0.4–0.7 cm long (Fig. |
Seed | ca. 1.9 × 1.6 cm | ca. 2.5 × 1.9 cm |
The type specimen of
It is a pleasure to name a species of
Sebo cordillero (Colombia;
All the studied specimens with flowers of
Finally,
As mentioned above, collections with flower have been previously identified as
The very distinctive fruits of
The specific epithet refers to Parque Nacional Yasuní, where most of the collections of this new species come from. At present, 2700 vascular plant species are documented from Yasuní, including a high diversity of lianas, epiphytes, and ferns; it is estimated its flora comprises approximately 3213 species; Yasuní is also home to
Geographic distribution of the new species of
Staminate flowers of
Dobompapoca (Ecuador: Huaorani;
It is possible to confuse
Comparison of
Morphological character |
|
|
|
|
---|---|---|---|---|
Leaf blade size, and base | 15.7–27.5 × 4.4–6.6 cm; acute | (15–) 20–60 × 10–16 cm; (usually) deeply cordate to truncate (obtuse) | 11.5–29.5 (–34) × 6.8–12.5 cm; attenuate to acute | 16–35 × 6.5–10.5 cm; shallowly cordate or rounded |
# of lateral veins | 14–18 | 11–28 | 14–19 | 17–30 |
Long filament column | 0.2–0.4 mm | 0.2–0.6 mm | 0.4–0.8 mm | 0.4–0.6 mm |
Long anthers | 0.5–0.6 mm | 0.4–0.5 mm | 0.4–0.6 mm | 1.1–1.6 mm |
Fruit size, and pubescent | 3.4–4.2 × 3.1–4 cm, inconspicuously pubescent | 2.5–3 × 1.2–2.5 cm; tomentulose, |
1.3–2.3 × 0.8–1 cm, densely pubescent | 2–2.8 × 1.5–2.2 cm; glabrescent |
Carina on the line of dehiscence | Markedly carinate (Fig. |
Carinate (Fig. |
Not carinate (Fig. |
Conspicuously carinate (Fig. |
Pericarp thick | 1.3 and 2.3 mm (just two specimens measured) | 0.5–5 mm | 1–1.2 mm | 0.5–1.5 mm |
Herbarium specimens of
Several collections of
French Guiana and Brazil (Amapá).
Gaan busi Mulumba (Paramaka;
The specific epithet alludes to the Spider monkeys, or ateles (
Endemic to Colombia (Boyacá).
The name
It is important to mention that, following our species concept, this name has been widely used to identify specimens, and is thus frequent in the literature and specimen databases. As circumscribed here,
(
1a | Mature leaf blades densely pubescent on abaxial surface (Fig. |
|
2a | Staminate flowers with the filament column shorter than the anthers |
|
3a | Leaf blades widely oblong to elliptical (7.3–11 cm wide), base obtuse; staminate inflorescence broadly paniculate; staminate flowers in lax terminal fascicles, the filament column 0.4–0.6 (–0.8) mm long, anthers 0.6–1 (–1.2) mm long; fruits with persistent trichomes; pericarp ca. 3.8–4 mm thick; growing at 20–260 m elevation |
|
3b | Leaf blades narrowly oblong or rarely elliptical (5–7.2 [–11.7] cm wide), base cuneate; staminate inflorescence narrowly paniculate; staminate flowers in dense terminal fascicles, the filament column 0.6–0.7 (–0.9) mm long, anthers 1.2–1.6 mm long; fruits with trichomes that fall like dust; pericarp ca. 2.4 mm thick; growing at 1300–1330 m elevation |
|
2b | Staminate flowers with the filament column longer than the anthers |
|
4a | Abaxial leaf blade surface drying silver to golden (Fig. |
|
5a | Staminate inflorescences 12–30 cm long; $pollen subtype |
|
5b | Staminate inflorescences 1–4 cm long; $pollen subtype |
|
4b | Abaxial leaf blade surface drying brown or whitish-grayish (Fig. |
|
6a | Leaf blades with cordate base; staminate inflorescences ca. 5.4 cm long, wide paniculate; staminate flowers in dense terminal fascicles, filament column with scattered trichomes (Fig. |
|
6b | Leaf blades with obtuse base; staminate inflorescences 7.5–9 cm long, narrow paniculate; staminate flowers in lax terminal fascicles, filament column glabrous; fruits covered with an inconspicuous layer of persistent trichomes |
|
1b | Mature leaf blades sparsely pubescent to glabrescent on abaxial surface (Fig. |
|
7a | Staminate flowers with the lobes of the perianth glabrous or almost glabrous internally |
|
8a | Twigs and inflorescence axes covered with appressed trichomes; fruits 2.7–3.5 long, covered by an inconspicuous layer of trichomes (Fig. |
|
8b | Twigs and inflorescence axes covered with erect trichomes; fruits 2–2.6 cm long, covered by a conspicuous layer of trichomes (Fig. |
|
9a | Leaf blades truncate to subcordate at the base; perianth of staminate flowers without resinous punctuations when rehydrated; infructescence with peduncle ca. 1.4–3.5 (–5) cm long; fruits 1.4–1.7 cm wide, without stipe, the base truncate to obtuse |
|
9b | Leaf blades attenuate to acute at the base; perianth of staminate flowers with resinous punctuations when rehydrated; infructescence with peduncle 0.7–1.3 cm long; fruits 0.8–1 cm wide, shortly stipitate, the base rounded |
|
7b | Staminate flowers with the lobes of the perianth pubescent internally |
|
10a | Staminate inflorescence 2.2–2.5 cm long; fruits covered with a conspicuous layer of trichomes (Fig. |
|
10b | Staminate inflorescence 3–9.5 cm long; fruits inconspicuously pubescent |
|
11a | Leaf blades (6.7–) 9.5–13.5 (–19.7) cm wide; staminate flowers with the filament column 0.9–1 mm long |
|
11b | Leaf blades 4.4–9.5 (–12.5) cm wide; staminate flowers with the filament column 0.2–0.5 mm long |
|
12a | Leaf blades elliptical to widely elliptical; staminate flowers in lax terminal fascicles; fruits 1.7–2 cm wide, the line of dehiscence slightly carinate |
|
12b | Leaf blades lanceolate to oblong; staminate flowers in dense terminal fascicles; fruits 2.8–4 cm wide, the line of dehiscence markedly carinate, like a wing |
|
13a | Staminate flowers with the filament column ca. 0.1 mm thick at the base, not widening at the base (i.e., straight throughout); fruits 3–3.6 × 2.8–3.1 cm, oblate, the line of dehiscence with a wing ca. 0.4 cm long |
|
13b | Staminate flowers with the filament column ca. 0.2–0.3 mm thick at the base, widening at the base; fruits 3.4–4.2 × 3.1–4 cm, ellipsoid to somewhat flattened, the line of dehiscence with a wing 0.4–0.7 cm long |
|
(Adapted from
1a | Adaxial surface of mature leaf blades pubescent |
|
2a | Leaf blades 25–60 cm long; staminate perianth submembranous; mature fruits covered by a conspicuous layer of trichomes (Fig. |
|
2b | Leaf blades 5–22 (–25.5) cm long; staminate perianth carnose to subcarnose; mature fruits without conspicuous layer of trichomes (glabrescent or densely tomentose to glabrate) (Fig. |
|
3a | Leaf blades with 28–32 lateral veins; staminate inflorescences to 8.5 cm long; staminate flowers with perianth 2–3 mm long; filament column 1.3–1.5 mm long; deciduous during blossom |
|
3b | Leaf blades with 24–58 lateral veins; staminate inflorescences to 14 cm long; staminate flowers with perianth ca. 1–1.5 mm long; filament column 0.3–0.4 mm long; evergreen |
|
1b | Adaxial surface of mature leaf blades glabrous |
|
4a | Leaf blades with 16–38 lateral veins |
|
5a | Leaf blades on abaxial side densely tomentose, with pediculate, dendritic trichomes; staminate flowers with perianth 2–2.5 mm long, carnose; androecium 1.2–1.6 mm long; fruits on the line of dehiscence smooth or slightly carinate, apex obtuse (Fig. |
|
5b | Leaf blades on abaxial side tomentulose to puberulent, with sessile or shortly pediculate, stellate trichomes; staminate flowers with perianth 1.2–1.8 mm long, membranous to submembranous; androecium 0.8–0.9 mm long; fruits on the line of dehiscence carinate, apex apiculate to acute |
|
6a | Leaf blades with 40–50 lateral veins; staminate inflorescence with 10–15 flowers per fascicle |
|
6b | Leaf blades with 30–38 lateral veins; staminate inflorescence with 50–100 flowers per fascicle |
|
4b | Leaf blades with (30–) 40–60 lateral veins |
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7a | External surface of staminate perianth almost glabrous (pubescent only at base and apex) |
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7b | External surface of staminate perianth uniformly pubescent (not described for |
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8a | Staminate flowers with perianth 2–3 mm long; fruits 3–5.7 cm long, conspicuously pubescent, always present; pericarp 3–6 mm thick |
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9a | Leaf blades with 49–65 lateral veins; staminate flowers with filament column 0.5–0.7 mm long; fruits rounded to ellipsoid, covered by a conspicuous layer of trichomes (1–2.3 mm thick), obtuse at apex (Fig. |
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9b | Leaf blades with (32–) 40–50 lateral veins; staminate flowers with filament column 0.9–1.3 mm long; fruits ovoid-ellipsoid, covered by an inconspicuous layer of trichomes, acuminate to rostrate at apex (Fig. |
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8b | Staminate flowers with perianth 1–1.5 mm long (not described in |
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10a | Abaxial surface of leaf blades glabrous to glabrescent |
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11a | Leaf blades 10–18 (–29) × 2.2–3 (–6) cm, narrowly oblong-lanceolate; fruits 2–3.5 × 1.8–2.5 cm, tomentose (Fig. |
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11b | Leaf blades 20–28 × 4–10 cm, elliptic or oblong-elliptic; fruits 2.1–2.3 × 1.5–1.8 cm‡, glabrous to glabrescent (Fig. |
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10b | Abaxial surface of leaf pubescent |
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12a | Leaf blades 10–42 × 3.5–12.5 cm, abaxial surface with sessile trichomes; deciduous in flower |
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12b | Leaf blades 5.5–11 × 4–8.5 cm, abaxial surface with pediculate trichomes; evergreen |
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We would like to express our gratitude to the curators of the following institutions for allowing access to, and use of, their physical collections, loan material, and/or digital images: CR, COL, F, HUH,