Corresponding author: Chelsea D. Specht (
Academic editor: T. Haevermans
In preparation for a full taxonomic revision of the Neotropical genera of
Maas PJM, Maas-van de Kamer H, André T, Skinner D, Valderrama E, Specht CD (2023) Eighteen new species of Neotropical Costaceae (Zingiberales). PhytoKeys 222: 75–127.
The Neotropical members of the family
Since Maas’ monographic work based largely on observations of herbarium collections, novel observations from extensive field work and herbarium research have resulted in the publication of 18 additional species by Paul Maas (
Recent years have also seen a rise in ecological and evolutionary studies on
In the present publication, 17 new species of
Material from ca. 150 herbaria was consulted for this study. A complete list of specimens examined will be provided in the forthcoming revision (Part II).
In addition, extensive field observations and collections have contributed to this study, supporting our understanding of species distributions and providing opportunities to photograph and observe key traits for distinguishing among taxa, including the colour of floral organs, orientation of the flower relative to the inflorescence (abaxial, erect, adaxial), colour of bracts, and colouration patterns of vegetative material. Field work was led by Paul and Hiltje Maas, with all co-authors participating. Dave Skinner visited most Neotropical countries and made essential observations on this group, particularly visiting type localities and taking detailed photographs, contributing to our understanding of the distribution of characters among taxa. Skinner’s observations have been recorded in iNaturalist and are available at
Brazil. Amazonas: Mauá Road, 22 Mar 1971,
Brazil (Amazonas, Pará) (Fig.
In terra-firme rainforests or campinarana vegetation, on sandy or clayey soils, at elevations of about sea level. Flowering and fruiting: October to March, July.
This species is named “manausensis” after the city of Manaus, State of Amazonas, Brazil, where this species is most commonly distributed.
For additional field data, see
Bolivia, La Paz: Prov. Bautista Saavedra, Area Natural de Manejo Integrado Apolobamba, between Paujeyuyo and Calzada, 1140 m, 16 Nov 2003,
Bolivia (La Paz) (Fig.
In wet, sub-Andine forests with
In Bolivia and Brazil, D. Skinner has observed plants with leaves and ligules that match this description. However, they were not in flower. The distribution of this species may be found to extend beyond the department of La Paz, Bolivia.
Panama, Veraguas: Río Segundo Brazo, 700–750 m, 8 Sep 1974,
Panama (Coclé, Colón, Veraguas) (Fig.
In forests, at elevations of 200–1300 m. Flowering in the rainy season.
This species strongly resembles
Mexico. Oaxaca: Sierra Mazateca, Mun. Santa María Chilchotla, Cuauhtemoc, ca. 4 km NE of Santa María Chilchotla, near Clemencia and Santa Rosa, 1200 m, 16 Jan 1984,
Mexico (Oaxaca) (Fig.
In primary, wet montane to premontane cloud forests, with some mosses on trunks, drier on ridge crests, with
We are greatly indebted to Manuel Gutiérrez, who assisted Dave Skinner in obtaining permission from the indigenous people of Santa Cruz Tepetotutla, Oaxaca, Mexico, to study this new species common in the Chinantla region of Oaxaca.
Colombia, Antioquia: Mun. San Rafael, along road from Guatapé to San Rafael, Río El Bizcocho, Finca El Castillo, 1059 m, 12 Oct 2013,
Colombia (Amazonas, Antioquia, Arauca, Boyacá, Caldas, Cauca, Chocó, Córdoba, Cundinamarca, Guajira, Guaviare, Huila, Meta, Norte de Santander, Quindío, Risaralda, Santander, Tolima, Valle del Cauca, Vichada), Venezuela (Mérida), Ecuador (Azuay, Carchi, Pastaza, Zamora-Chinchipe), Peru (Ucayali). (Fig.
In forests (a.o., “selva baja/alta perennifolia” or wet montane forests) or forested roadsides at elevations of 50–1700 m. Flowering year-round.
Colombia: Caña agria, Cañagria, Cañaguate. Peru: Sacha huiro.
This species was collected by Paul and Hiltje Maas for the first time in the Colombian department of Antioquia and is therefore named in recognition of that locality.
Colombia, Chocó: Mun. San José del Palmar, mouth of Río Torito (affluent of Río Hábita), Finca Los Guaduales, 730–830 m, 4 Mar 1980,
Costa Rica, Panama, Colombia (Antioquia, Chocó) (Fig.
In forests, at elevations of 0–1300 m. Flowering year-round.
Panama: Caña agria (Kuna).
Bolivia, Cochabamba: Prov. Chapare, Villa Tunari, 40 km hacia el Beni, por Chipiriri, 400 m, 24 Nov 1981,
Bolivia (Cochabamba) (Fig.
In high forests (“pie de la ladera con bosque alto”), at an elevation of ca. 400 m. Flowering in November.
This species is named after the Bolivian department of Cochabamba, where it was collected.
Colombia, Putumayo: 15 km S of Villagarzón on road to Puerto Umbria, 360 m, 17 Jul 2018,
Colombia (Putumayo), Ecuador (Morona-Santiago, Zamora-Chinchipe) (Fig.
In forests, in open shade or sunny areas, often found along roadsides and other disturbed areas, at elevations of 360–1500 m. Flowering year-round.
Plants of
This new species is observed to be very common along the eastern flanks of the mountains from Putumayo, Colombia, south through Ecuador to Zamora-Chinchipe. Many more recent observations of this new species are recorded on
Brazil Acre: Mun. Tarauacá, Basin of Rio Juruá, Rio Tarauacá, Seringal Sumaré, 19 Nov 1995,
Peru (Ucayali), Brazil (Acre, Rondônia) (Fig.
In non-inundated (terra firme) forests, campinarana, river margins, or roadsides at elevations of 0–350 m. Flowering year-round.
This species is named for our dear colleague and friend Douglas Daly, whom PM and HM met many times in his home institute at the New York Botanical Garden Herbarium (NY) and who enabled us to undertake field work in the Brazilian regions of Acre and Amazonas. He also inspired CDS during her graduate studies at the New York Botanical Garden, providing insights into excellence in field research and tropical botany.
Ecuador, Pichincha: along the road from Aloag to Santo Domingo, 1150 m (“3500 ft”), 17 Nov 1974,
Ecuador (El Oro, Esmeraldas, Guayas, Loja, Manabi, Pichincha) (Fig.
In forested roadsides, dry forests, or gallery forests at elevations of 0–1500 m. Flowering year-round.
Mexico, Chiapas: Mun. Chilon, 22 km from Temo in the direction of Palenque, 2 km from Patathel, near El Chich, 850 m, 4 Nov 1998,
Mexico (Chiapas, Oaxaca, Tabasco, Veracruz) (Fig.
In forests (a.o. “selva baja/alta perennifolia”) with
Mexico: Apagafuego.
This species is named after its very soft indument on the abaxial surface of the lamina, the word “mollis” meaning soft in Latin.
In Maas’ treatment of
Peru, Madre de Dios: Prov. Manu, near Shintuya, on road to Salvación, 600 m, 9 Feb 1974,
Peru (Cuzco, Huánuco, Madre de Dios, San Martín) (Fig.
In forests, in wet, shady areas, often in disturbed places, at elevations of 450–1300 m. Flowering in the rainy season.
Plants of
This new species has been widely cultivated from the type collection
Ecuador, Tunguruhua: Río Verde Grande, 1500 m, 30 Mar 1956,
Ecuador (Pastaza, Tungurahua, Zamora-Chinchipe) (Fig.
In forests, at elevations of 1150–1600 m. Flowering in the rainy season.
The species name “
Most collections of
The authors are indebted to orchid specialist Marco Jiménez Villata of Zamora, Ecuador, who has explored the forests of Zamora-Chinchipe for over 30 years. He showed Dave Skinner several localities in the province, where he was able to make photographs of this species. Skinner later found the living plants of this new species in Tungurahua, which were consistent with the plants in Zamora-Chinchipe but usually with less dense hairs on the undersides of the leaves. Moreover, in living material, the corolla lobes were found to be sharply recurved, which is very unusual in the genus
Colombia, Cauca: forested roadside from San Juan de Villalobos to Pitalito, 1590 m, 19 Jul 2018,
Colombia (Cauca, Putumayo) (Fig.
Along forested roadsides, at an elevation of ca. 1600 m. Flowering and fruiting: May and July.
This species is named “pitalito” after its type locality near Pitalito, in the Colombian department of Cauca.
Brazil, Acre: vicinity of Serra da Moa, 22 Apr 1971,
Brazil (Acre, Amazonas, Rondônia), Peru (Loreto, San Martín) (Fig.
In non-inundated (terra firme) or periodically inundated (várzea) forests at elevations of 0–260 m. Flowering year-round.
Brazil: Canafiche.
This species is named after our friend and dearest colleague Sir Ghillean Prance, who invited us (PM and HM) in 1971 to join various of his expeditions into the Brazilian Amazon region and who inspired me (PM) very much to continue working as a Neotropical taxonomist.
We have named this species after Sir Ghillean Prance, who enabled the first author (PM) to undertake his first steps in the Amazonian world and assisted and taught him in a very kind and generous way.
The two Peruvian collections (
Bolivia, Beni: km 4 of road from Riberalta to Santa María, 150 m, 22 Jan 1999,
Bolivia (Beni), Brazil (Rondônia) (Fig.
In secondary roadside vegetation, at elevations of 150–700 m. Flowering year-round.
This species is named “pseudospiralis”, referring to its resemblance to
Peru, Pasco: Prov. Oxapampa, Gramazú, 1961 m, 17 Mar 2005,
Peru (Huánuco, Pasco) (Fig.
In primary or secondary, premontane or montane forests, at elevations of (300–)1400–2200 m. Flowering year-round.
Colombia, Putumayo: Km 42 of Mocoa-Puerto Asis Road, Quebrada El Whiskey, Finca Santa Marta, Hilltop forest, c. 400 m (“1260 ft”), 5 Nov 1974,
Distribution maps for new species of
Colombia (Putumayo). Ecuador (Napo), Peru (Loreto) (Fig.
Distribution maps for new species of
In tropical wet forests at an elevation of 250–400 m. Flowering period is uncertain.
This species has been named after its type locality El Whiskey (Putumayo, Colombia), with “cola” the Latin for “living in”.
This new species has long been widely cultivated worldwide, originally grown from seeds collected by Tim Plowman when preparing
In April 2022, Dave Skinner visited the Santa Cruz private reserve of Project Amazonas near Iquitos, Loreto, Peru and found large populations of plants in primary forest that are identical vegetatively and have flowers that match those of
We are very grateful to the many curators of the herbaria from which we borrowed or studied material, especially Anna Stalter and Peter Fraissinet of the Herbarium of the L.H. Bailey Hortorium (BH). We thank Esmée Winkel (Naturalis Herbarium in Leiden) for preparing some drawings of the new species, and are greatly indebted to the many people who provided us with photographs of
Collections are identified by first collector and number only. The abbreviations behind the collector number refer to the following taxa:
Ch man =
Co alf =
Co all =
Co alt =
Co ant =
Co cal =
Co coc =
Co con =
Co dou =
Co gib =
Co mol =
Co obs =
Co ore =
Co pit =
Co pra =
Co pse =
Co rub =
Co whi =
Asplund 7833: Co ore; 15412: Co gib; 15895: Co gib; 16421: Co gib; 16494: Co gib; 16763: Co gib; 18160: Co gib; 20068: Co ore – Assunção 72: Co pra;
Balcázar 49: Co ant – Beck 7320: Co coc – Beltrán Cuartas 78: Co ant – Betancur B. 5970: Co ant; 7500: Co ant; 16507: Co ant; 16651: Co ant – Bilby 33: Co pra – Boeke 1009: Co ant – Bohlin 1226: Co gib – Boyle 2556: Co alt – Breedlove 26463: Co mol;
Calderón 2986: Co pra – Camp E 2395: Co ant – Campbell 8951: Co pra – Carreira 315: Co pra – Castro 81: Co pra – Castillo 642: Co rub – Cavalcante 540: Co pra; Cid 5437: Co pra – Cordero-P. 91: Co ant – Cornejo 425: Co gib; 5792: Co gib – Croat 62674: Co dou; 83948A: Co gib; 85008: Co dou; 92034: Co ant – Cuatrecasas 8396: Co ant; 13714: Co ant;
Daly 8606: Co dou – Davidson 8770: Co cal; 9395: Co rub – Dodson 913: Co ore; 1441: Co ore; 5347: Co gib; 13039: Co gib – Dressler 4190: Co cal – Duque-Jaramillo 2251: Co ant; 3912-A: Co ant;
Ehrich 5c: Co pra;
Fernández-Alonso 11440: Co pit – Fiaschi 3287: Co pra – Forero 5765: Co cal; 6161: Co ant; 6668: Co cal; 7413: Co ant – Foster 8950A: Co rub; 10937: Co obs – Frame 171: Co pra – Fuentes Claros 6216: Co alf – Funk 10490: Co cal;
Galeano 2036: Co ant – García-Barriga 18240: Co ant – Gentry 52203: Co pra; 72332: Co gib; 10165: Co gib; 27304: Co obs – Gilli 97: Co mol; 119: Co gib – Gómez Pignataro 21201: Co cal – Guevara-Ibarra 39: Co ant;
Hammel 15871: Co gib; 16513: Co cal; 17668: Co cal; 20494: Co cal; 20694: Co cal – Harling 14354: Co gib; 18358: Co gib – Haught 4292: Co ant – Hayden 1020: Co cal – Hernández G. 387: Co mol; 1435: Co mol – Hitchcock 20543: Co gib – Holm-Nielsen 27916: Co gib;
Idrobo 2027: Co ant – Ishiki 2400: Co mol;
Jardim 56: Co pse;
Kirkbride 1180: Co cal – Knapp 4519: Co cal; 5524: Co cal – Kress 86-1978: Co cal;
Labiak 6250: Co pra – La Frankie 85-38: Co cal – Lozano-Contreras 7397: Co ant – Lücking s.n.: Co cal;
Maas 596: Co ant; 1548: Co cal; 1652: Co all; 1783: Co ant; 2915: Co gib; 4795: Co gib; 8751: Co pse; 8999: Co dou; 10488: Co ant; 10631: Co ant; 10635: Co gib; 10688: Co cal; 10707; Co con; 10733: Co pit; P 12972: Co dou; P 13213: Co dou – MacDougal 4020: Co cal – Madison 4623: Co ant – Madsen 86796; Co con – Martínez Salas 3160: Co mol –Monteagudo Mendoza 7658: Co rub – Monteiro 601: Co dou;
Nagata 2811: Co gib – Nee 54842: Co pra – Nevling 1526: Co mol – Nuñez 306: Co gib – Nuñez V. 23785: Co obs;
Obando 284: Co ant – Øllgaard 90385; Co con – Ortiz V. 521: Co rub;
Peña-Chocarro 65: Co pse – Penland 179: Co ore – Pennell 8486: Co ant – Pennington 56 SD: Co gib – Plowman 1775: Co obs; 4454: Co gib; 4455: Co gib; 5054: Co obs; 5054 A: Co obs; 5791: Co obs; 7595: Co obs – Prance 3151: Co pra; 5218: Co pra; 6672: Co pra; 8227: Co pra; 8459: Co pse; 8866: Co pra; 11529: Ch man; 12230: Co dou – 12265: Co pra; 15300: Co pra – Proctor Cooper s.n.: Co pra – Pulido 107: Co ant;
Ramamoorthy 1790: Co mol – Ramos-Pérez 3052: Co ant – Rentería Arriaga 2123: Co ant – Rincón Guttiérrez 592: Co alt – Rivera Reyes 1610: Co alt – Rodrigues 2996: Co pra – Roe 928: Co mol – Rojas Gonzáles 3574: Co rub – Rojas-Zamora 136: Co ant – Rubio 2007: Co gib;
Schunke V. 10195: Co ant; 15567: Co dou; 16185: Co dou – Silveira 559: Co pra; 596: Co pra – Silverstone-Sopkin 3085: Co ant; 5811: Co ant; 8963: Co ant – Skinner 3322: Co obs; R 3131: Co rub; R 3332: Co ore; R 3333: Co ore; R 3833: Co obs – Smith 2946: Co rub; 5360: Co rub – Solheim 1367: Co alt – Solomon 7978: Co pse – Sparre 13862: Co gib –
Steward 476: Co pra;
Uliana 1342: Co pra – Ulloa U. 111: Co gib;
Vareschi 1674: Co ant – Vargas-Figueroa 855: Co ant – Vásquez Martínez 27782: Co rub – Vélez J. 7073: Co ant – Vieira 369: Co dou – Villanueva 1844: Co ant – Von Martius s.n.: Co pra;
Wessels Boer 2440: Co ant.