Corresponding author: José María Cardiel Sanz (
Academic editor: Dmitry Geltman
A critical review of the Ecuadorian species of
Se presenta una revisión crítica de las especies ecuatorianas de
Despite its great diversity,
This work presents a reviewed critical synopsis of the species of
Species of
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accepted | ||
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The voucher cited (P. Jørgensen 1551) corresponds to |
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The voucher cited (P. Jørgensen 1449) corresponds to |
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accepted | accepted | |
The voucher cited (Berg & Akkermans 1016) corresponds to |
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Doubtful taxon not found in Ecuador (fide |
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accepted | ||
accepted | ||
accepted | accepted | |
accepted | accepted | |
accepted | = |
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accepted | = |
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accepted | ||
not cited | accepted | |
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accepted | accepted | |
The voucher cited (C. Ceron & C. Iguago 5461) correspond to |
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The voucher cited (Dodson & al. 7109) corresponds to |
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accepted | ||
accepted | accepted | |
accepted | The voucher cited (Sodiro 151) corresponds to |
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accepted | accepted | |
accepted | ||
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accepted | The voucher cited (Madsen 63051) corresponds to |
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accepted | ||
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The voucher cited (Van der Werff & W. Palacios 10355) corresponds to |
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accepted | accepted | |
accepted | = |
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accepted | = |
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accepted | accepted | |
We studied 987 Ecuadorian collections of
The structure of the checklist follows, in general terms, those of
All the information gathered as part of this work is available online in the regularly updated “Acalypha Taxonomic Information System” Website (Cardiel J.M., P. Muñoz, E. Dorda & M. Pardo de Santallana
Our work records 20 accepted species of
1 | Female flowers pedicellate; calyx with 4 or 5 sepals, the subtending bracts inconspicuous, not becoming foliaceous in fruit | Subgen. |
2 | Female flowers sessile; calyx with 3 sepals, the subtending bracts becoming foliaceous and accrescent in fruit (except in |
Subgen. |
Subgenus
1a | Leaf blade palmately nerved, brightly colored minute resinous droplets present, mainly on lower surface | 18. |
1b | Leaf blade pinnately nerved, brightly colored minute resinous droplets absent | 2 |
2a | Female inflorescences paniculate. Leaf blade with 10–17 veins per side; stipules generally more than 5 mm long. Petioles more than 1 cm long | 11. |
2b | Female inflorescences racemose. Leaf blade with 9–13 veins per side; stipules inconspicuous, ca. 1 mm long. Petioles less than 1 cm long | 12. |
Subgenus
1a | Herb or suffrutex | 2 |
1b | Trees or shrubs | 6 |
2a | Female inflorescences ellipsoid or cylindrical, densely flowered, with the axis completely covered by the flowers | 3 |
2b | Female inflorescences loosely flowered, with the axis conspicuously visible | 5 |
3a | Female bract with long awned lobes | 4 |
3b | Female bract with triangular awnless lobes | 5. |
4a | Young branches and leaves without glandular hairs; leaf blade acute; styles branched | 2. |
4b | Young branches and leaves with glandular hairs; leaf blade acuminate; styles unbranched | 1. |
5a | Female inflorescences terminal, bracts with filiform lobes cut more than ½ length to the base | 17. |
5b | Female inflorescences axillary, bracts with triangular lobes cut ca. ¼ length to the base | 4. |
6a | Leaves with indumentum of stellate hairs | 7 |
6b | Leaves without indumentum of stellate hairs | 8 |
7a | Female inflorescences terminal, subtending bracts and styles with stellate hairs | 16. |
7b | Female inflorescences generally axillary, rarely terminal, subtending bracts and styles without stellate hairs | 5. |
8a | Female or bisexual inflorescences terminal | 9 |
8b | Female or bisexual inflorescences axillary | 12 |
9a | Female bracts without glandular hairs | 18. |
9b | Female bracts with glandular hairs | 10 |
10a | Female bracts subtriangular at maturity, with the central tooth prominent, lanceolate, acuminate | 19. |
10b | Female bracts suborbicular at maturity, with the central tooth not or slightly prominent | 11 |
11a | Leaf blade generally broadly ovate-lanceolate; accrescent bracts with glandular hairs ca. 0.3–0.5 mm long; styles 4–5 mm long. | 14. |
11b | Leaf blade generally narrowly ovate-lanceolate; accrescent bracts with glandular hairs ca. 1 mm long; styles 7–8 mm long | 10. |
12a | Plants with both unisexual and bisexual inflorescences | 13 |
12b | Plants with all the inflorescences unisexual | 14 |
13a | Leaf blade generally triangular-lanceolate, palmately nerved | 4. |
13b | Leaf blade elliptic-lanceolate or oblong-lanceolate, pinnately nerved | 6. |
14a | Leaf blade pinnately nerved | 15 |
14b | Leaf blade palmately nerved | 16 |
15a | Leaf blade generally obovate, the base subcuneate; female inflorescences 7–15 cm long | 3. |
15b | Leaf blade ovate to oblong-lanceolate, the base rounded to subcordate; female inflorescences 25–40 cm long | 12. |
16a | Female inflorescences extremely densely flowered, with the axis hidden; bracts non-accrescent | 9. |
16b | Female inflorescences more or less densely flowered, with the axis visible; bracts conspicuously accrescent | 17 |
17a | Leaf blade generally variegated; female inflorescences up to 10 cm long | 20. |
17b | Leaf blade not variegate; female inflorescences more than 15 cm long | 9. |
01.
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Herb. Coastal, 0–250 m. ‒ Provincial distribution: Guayas (2 collections examined). ‒ Voucher: F.M.Valverde 334 (COL, US).
02.
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Annual herb or small suffrutex. Coastal, 0–1000 m. Roadsides and disturbed vegetation. ‒ Provincial distribution: Cotopaxi, Guayas, Los Ríos, Manabí, Napo, Pichincha, Sucumbíos (14 collections examined). ‒ Voucher: O.Haught 2994 (GH, US).
03.
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Shrub or small tree. Mainly Amazonian and Coastal, excepcionally Andean, (0-) 250–1000(-1500) m. Generally associated with primary rainforests. ‒ Provincial distribution: Esmeraldas, Guayas, Los Ríos, Manabí, Morona-Santiago, Napo, Ore-llana, Pastaza, Santa Elena, Santo Domingo de los Tsáchilas , Sucumbíos, Tungurahua, Zamora-Chinchipe (222 collections examined). ‒ Voucher: E. Gudiño 61 (GB, DAV, MO, QCNE, U).
04.
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Shrub or suffrutex. Coastal, 0–250 m. Associated with dry deciduous forest, savanna and thickets. ‒ Provincial distribution: El Oro, Guayas, Manabí, Santa Elena (8 collections examined). ‒ Voucher: J.E. Madsen 63938 (AAU, MA, QCA, QCNE).
05.
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Shrub or small tree. Andean, (1300)1750–2500 m. Montane rainforests. ‒ Provincial distribution: Bolívar, Chimborazo, Imbabura, Napo, Orellana, Pichincha, Santo Domingo de los Tsáchilas, Tungurahua (52 collections examined). ‒ Voucher: M.Balslev & E.Madsen 10362 (AAU, COL, C, F, MO, NY, QCA, SEL, US).
06.
Shrub or small tree. Amazonian, Andean and Coastal, 0–2000 m. Lowland rainforests, deciduous and semi-deciduous forests (often along river banks) and disturbed areas. ‒ Provincial distribution: Bolívar, Carchi, Chimborazo, Cotopaxi, El Oro, Esmeraldas, Guayas, Imbabura, Loja, Los Ríos, Manabí, Morona-Santiago, Napo, Orellana, Pastaza, Pichincha, Santo Domingo de los Tsáchilas, Sucumbíos, Tungurahua, Zamora-Chinchipe (310 collections examined). ‒ Voucher: V.Zak & J.Jaramillo 2822 (DAV, F, GB, GH, K, MO, NY, QCNE, SEL).
07.
Shrub. Coastal, 0–250 m. Cultivated and naturalized. ‒ Provincial distribution: Guayas, Santo Domingo de los Tsáchilas (4 collections examined). ‒ Voucher: L.P.Kvist & E.Asanza 40725 (AAU, QCA, QCNE).
08.
Annual herb or suffrutex. Andean, 1000–2500 m. Disturbed areas. ‒ Provincial distribution: Bolívar, Cañar, Chimborazo, Pichincha (6 collections examined). ‒ Voucher: F.R.Fosberg & M.A.Giler 22627 (NY, US).
09.
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Shrub or small tree. Amazonian, (250-)500–2000 m. In lowland and lower montane rainforests, frequent in disturbed areas. ‒ Provincial distribution: Carchi, Chimborazo, El Oro, Esmeraldas, Morona-Santiago, Napo, Orellana, Pastaza, Pichincha, Santo Domingo de los Tsáchilas, Sucumbíos, Tungurahua, Zamora-Chinchipe (103 collections examined). ‒ Voucher: H.Balslev & E.Madsen 10462 (AAU, C, COL, F, GB, MO, NY, QCA, SEL, US).
10.
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Shrub or small tree. Andean, 2000–3250(3800) m. Upper and lower montane rainforests, mainly disturbed. ‒ Provincial distribution: Azuay, Bolívar, Cañar, Carchi, Chimborazo, Imbabura, Loja, Pastaza, Pichincha, Tungurahua (83 collections examined). ‒ Voucher: C.E.Cerón & G.Benavides 2551 (F, MO, QCA, QCNE).
11.
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Shrub or small tree. Andean, 1250–2500 m. In lower and upper montane rain- forests. ‒ Provincial distribution: Azuay, Cotopaxi, Morona-Santiago, Napo, Orellana, Pichincha, Santo Domingo de los Tsáchilas (87 collections examined). ‒ Voucher: J.Jaravillo & V.Zak 7481 (F, MO, NY, QCA, QCNE).
12.
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Shrub or small tree. Amazonian, 500–1250(-1750) m. In lower montane rainforests and lowland rainforests. ‒ Provincial distribution: Morona-Santiago, Napo, Sucumbíos, Zamora-Chinchipe (28collections examined). ‒ Voucher: H.Van der Werff & E.Gudiño 11350 (G, GB, MO, NY, QCNE).
13.
Shrub or small tree. Amazonian, 0–250 m. Lowland rainforests, often along river banks. ‒ Provincial distribution: Orellana, Pastaza, Sucumbíos (6 collections examined). ‒ Voucher: C.E.Cerón et N.Gallo 5186 (AAU, MO, QCNE).
14.
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Shrub or small tree. Coastal, 0–500(-750) m. Generally associated with dry forests and thickets. ‒ Provincial distribution: Guayas, Loja, Los Ríos, Manabí (19 collections examined). ‒ Voucher: E.Asplund 15240 (S)
15.
Shrub or small tree. Amazonian region, 0–1000(-1250) m. In lowland rainforests. ‒ Provincial distribution: Morona-Santiago, Napo, Orellana, Pastaza, Sucumbíos, Zamora-Chinchipe (136 collections examined). ‒ Voucher: J.Zaruma & al. 126 (F, GB, MO, NY, QCNE).
16.
17.
Annual herb. Coastal, 0–500(-1200) m. Generally associated with dry forest and open sites. ‒ Provincial distribution: Guayas, Loja, Los Ríos, Manabí, Santa Elena (24 collections examined). ‒ Voucher: C.M.Dodson & P.M.Dodson 13680 (MO, QCNE, SEL).
18.
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Shrub or small tree. Coastal, 0–250 m. Generally associated to secondary woods and thickets, also with savanna and deciduous forest, mainly on sandy and humid soils. ‒ Provincial distribution: El Oro, Esmeraldas, Guayas, Los Ríos, Manabí, Pichincha, Santo Domingo de los Tsáchilas (54 collections examined). ‒ Voucher: L.P.Kvist 40466 (AAU, MO, NY, QCA).
19.
20.
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Shrub. Cultivated and naturalized. Coastal, Andean and Amazonian, 0–1500 m. ‒ Provincial distribution: Guayas, Imbabura, Manabí, Napo, Santo Domingo de los Tsáchilas (6 collections examined). ‒ Voucher: M.Acosta-Solís 13076 (F).
This work was partly financed by the Spanish Government, through the research project EUI 2008-0388. Support was also received from the Universidad Autónoma de Madrid (Spain) and the Regional Government (Comunidad de Madrid), through the research project CCG07-UAM/AMB-1453. We kindly thank the curators and staff of the aforementioned herbaria for facilitating the study of their specimens. Francisco Pando, manager of GBIF-Spain, kindly helped us in everything relating to biodiversity databases. The authors are also grateful to Grady Webster for his advice in the early studies on Ecuadorean