Research Article |
Corresponding author: Marie-Stéphanie Samain ( mariestephanie.samain@gmail.com ) Academic editor: Sandy Knapp
© 2021 Marie-Stéphanie Samain, Carolina Granados Mendoza, Esteban Manuel Martínez Salas.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Samain M-S, Granados Mendoza C, Martínez Salas EM (2021) On Hydrangea peruviana, an endangered species from Ecuador, and Hydrangea oerstedii, very common in Costa Rica and Panama, and seven threatened Central and South American Hydrangeas, which have been confounded with these. PhytoKeys 171: 91-153. https://doi.org/10.3897/phytokeys.171.56351
|
Hydrangea section Cornidia, currently consisting of 19 accepted taxa, occurs from Mexico to Chile and Argentina, with one species in southeast Asia. Its representatives are root-climbing lianas which may grow up to 60 m high in the tree canopy of temperate to (sub)tropical forests. Our extensive field work throughout its distribution area, study of herbarium specimens and ongoing molecular studies have resulted in the discovery of species new to science, as well as new insights into the circumscription of many taxa. We here present amended descriptions for seven Hydrangea species of Central and South America and discuss the taxonomical situation of two Colombian Hydrangeas, including an identification key, illustrations, and distribution maps. Field work was carried out in Costa Rica, Panama, Ecuador and Peru, including exploration in areas where the genus had not been collected before. These specimens and observations were complemented with the study of specimens of 41 herbaria of North, Central and South America, as well as Europe. Detailed morphological studies of all species were carried out, based on living plants in their natural habitat, as well as on dried specimens from our own collections and all available herbarium material. Type material was studied in detail for all species concerned. Based on an extensive number of morphological characters, combined with distribution patterns, phenological differences and ecological preferences, including molecular data in most cases, Hydrangea peruviana and H. oerstedii are clearly distinct taxa, as well as the other seven species mentioned here, which had been synonymized with either of these two species. The present study results in the recognition of 26 species in section Cornidia and exemplifies the urgent need for profound taxonomic studies in plants, as in many families we do not dispose of well-circumscribed units for conservation to mitigate the already occurring unprecedented loss of biodiversity.
Hydrangea sección Cornidia, que actualmente consiste en 19 taxones aceptados, se distribuye desde México hasta Chile y Argentina, con una especie en el sureste asiático. Sus representantes son lianas trepadoras que pueden crecer hasta 60 m de altura en la copa de los árboles de los bosques templados a (sub)tropicales. Nuestro extenso trabajo de campo en toda su área de distribución, la revisión de especímenes de herbario y los estudios moleculares en curso han dado como resultado el descubrimiento de especies nuevas para la ciencia, así como conocimiento nuevo sobre la circunscripción de muchos taxones. Aquí presentamos descripciones enmendadas para siete especies de Hydrangea de Centro y Suramérica y discutimos la situación taxonómica de dos Hydrangeas colombianas, incluyendo una clave de identificación, ilustraciones y mapas de distribución. El trabajo de campo se llevó a cabo en Costa Rica, Panamá, Ecuador y Perú, incluída la exploración en áreas donde el género no había sido recolectado antes. Estos especímenes y observaciones se complementaron con el estudio de especímenes de 41 herbarios de Norte, Centro y Suramérica, así como de Europa. Se llevaron a cabo estudios morfológicos detallados de todas las especies, basados en plantas vivas en su hábitat natural, así como en muestras secas de nuestras propias colecciones y todo el material de herbario disponible. El material tipo se estudió en detalle para todas las especies en cuestión. En base a una gran cantidad de caracteres morfológicos, combinados con patrones de distribución, diferencias fenológicas y preferencias ecológicas, incluyendo datos moleculares en la mayoría de los casos, Hydrangea peruviana y H. oerstedii son taxones claramente distintos, así como las otras siete especies mencionadas aquí, que habían sido sinonimizadas con cualquiera de estas dos especies. El presente estudio resulta en el reconocimiento de 26 especies en la sección Cornidia y ejemplifica la necesidad urgente de estudios taxonómicos profundos en plantas, ya que en muchas familias no disponemos de unidades de conservación bien circunscritas para mitigar la pérdida de biodiversidad sin precedentes.
Conservation, Ecuador, functional dioecism, hortensia, lianas, Neotropics, Peru, taxonomy
The relatively poorly known Hydrangea section Cornidia Ruiz & Pav. consists of 19 currently accepted taxa (18 species and one variety) and a yet undefined number of species new to science and taxa that have been erroneously synonymized. The representatives of this section occur from northern Mexico to southern Chile and Argentina with one species, Hydrangea integrifolia Hayata, in southeastern Asia (
Hydrangea section Cornidia (hereafter shortened as Cornidia) is monophyletic, including the single Asian species (
The monography of the genus Hydrangea s.s. by
We have carried out extensive field work throughout the distribution area of Cornidia since 2009, and as a consequence, we realized that its representatives are much more common than previously known (albeit most of them are severely threatened, mainly because of habitat destruction) and that its incredible morphological variation definitely did not fit into the then eleven accepted species. However, contrary to Mexico where we have recently described seven new Hydrangea species and registered one new record of a species until then only known from Guatemala (
Apart from the considerable confusion over names, an additional challenge in this group is the functional dioecism, which we have observed in most individuals during our extensive field work throughout the Neotropics, and which is reflected by a notoriously different size and shape of flower receptacle, anthers and pistils between functionally female and male flowers (
The above-mentioned issues emphasize the need for a complete and urgent revision of Cornidia, especially in the light of conservation of these species. Their pristine habitat with very specific conditions (near water, often flat topography near the plants, efficient drainage) makes them not only promising bio-indicators, but also poses an additional threat as these habitats are being destroyed because they are highly appreciated by local people for agriculture (
The present work aims at resolving the complex of nine species from Central and South American species that had been formally synonymized under Hydrangea peruviana (H. schlimii Briq., H. caucana Engl., H. durifolia Briq., H. goudotii Briq., H. oerstedii Briq., H. panamensis Standl., H. peruviana Moric. ex. Ser., H. trianae Briq. and H. weberbaueri Engl.). The main objective of the present study is to show that we are dealing with nine different species, primarily based on morphological characters, and strengthened by preliminary molecular data. The identification key which is presented here thus has no other aim than showing that these species can be distinguished with relative ease based on morphological characters that are straightforward to observe. Based on meticulous observations in the field and of herbarium specimens, we here present amended descriptions for seven Hydrangea species of Central and South America and discuss the taxonomic situation of two more species of this group, including an identification key of species as recognized here, illustrations, and distribution maps, as well as information about their diversity, their global conservation status, and their affinities with other Cornidia species.
Field work focused on Hydrangea has been carried out by the authors during dry and rainy seasons in Costa Rica (2012, 2013), Ecuador (2012), Panama (2019) and Peru (2011, 2012, 2013), coinciding with the flowering and fruiting seasons, based on herbarium material of most of the species included. Exploring field work was not only carried out in the areas where previous collections had been made, but also in zones where Hydrangeas had not yet been recorded and where we suspected they would be present, based on our knowledge of their habitat preferences. Branches with inflorescences, flowers and fruits of all stages were collected and preserved. Moreover, in several individuals where architectural traits seemed to be important, whole branches were collected, of course without affecting the viability of the individual plants, cut and subsequently numbered in order to maintain the architecture available for further study. All specimens were deposited in local herbaria in the respective countries where we collected (CR (including INB), HOXA, PMA, QCNE and USM) with duplicates in the herbaria of the Instituto de Ecología, A.C. (
Our field observations were complemented with a detailed study of relevant herbarium specimens of 41 herbaria in Europe, North, Central and South America (A, AAU, AMAZ, B, BM, BR, C, CAS, COL, CR, DUKE, E, F, G,
Red List categories were obtained according to the IUCN Red List criteria (
The present treatment includes the Cornidia species Hydrangea oerstedii and H. peruviana, plus seven other species which had been erroneously synonymized with the former two. Amended morphological descriptions for seven of these and an identification key to the taxa treated here are provided. We do not repeat here the morphological description of the section as this has been published in an Open Access paper by
It should be mentioned that our ongoing molecular studies in the Cornidia clade show that most of the species studied here are even not closely related, with the exception of Hydrangea panamensis and H. peruviana on the one hand, and H. goudotii and H. trianae on the other hand (
Of the nine taxa treated below, we have observed six in the field throughout their distribution area. Hydrangea caucana Engl., H. durifolia Briq. and H. schlimii Briq. have not been collected recently and are known with certainty from nine herbarium specimens, the type collection and a putative additional herbarium specimen, or the type collection only, respectively. The three of them are endemic to Colombia, where we have not yet been able to collect Hydrangeas due to collection permit and export regulations.
(Note: as mentioned above, this is a partial key that shows that these species are easy to distinguish. It cannot be used to identify all red-flowered Hydrangeas of Central and South America)
1 | Inflorescences mainly consisting of flowers with enlarged sepals; currently only known from the type locality in Colombia | 7. H. schlimii Briq. |
– | Inflorescences with only the marginal flowers with enlarged sepals, these flowers placed terminally on cymes or racemes, one per partial inflorescence, or rarely absent; occurring in Costa Rica, Panama, Colombia, Ecuador or Peru | 2 |
2 | Leaves up to 13 cm long and 6 cm wide | 3 |
– | Leaves longer than 13 cm and wider than 6 cm | 6 |
3 | Leaf apex rounded with a very small acumen; acarodomatia on the abaxial side 0–2 per leaf; Costa Rica and Panama | 5. H. panamensis Standl. |
– | Leaf apex acute to acuminate, rarely mucronate; acarodomatia more numerous; occurring in Central and South America | 4 |
4 | Lamina very slightly spoon-shaped, elliptic to slightly obovate; leaf margin serrate to slightly dentate; endemic to Ecuador | 6. H. peruviana Moric. ex Ser. |
– | Lamina flat, ovate to lanceolate-elliptic; leaf margin glandular dentate; Costa Rica, Panama and Colombia | 5 |
5 | Free-growing branches and inflorescence axes pubescent with persistent reddish brown stellate hairs; endemic to Colombia | 1. H. caucana Engl. |
– | Free-growing branches glabrous, and inflorescence axes pubescent with caducous, appressed, white stellate hairs; Costa Rica and Panama | 4. H. oerstedii Briq. |
6 | Leaves with primary veins and secondary veins parallel to each other and forming a distinct regular pattern with all veins arching towards the apex; Colombia, Ecuador and Peru | 9. H. weberbaueri Engl. |
– | Leaves with a pinnate leaf vein pattern | 7 |
7 | Free-growing branches and inflorescence axes glabrous | 8 |
– | Free-growing branches with appressed white caducous stellate pubescence | 9 |
8 | Abaxial leaf side with a granulate texture due to remaining basal stalks of stellate pubescence; inflorescence axis terete; inflorescence 5–9.5 cm wide; cymes compact; currently known from Colombia only | 2. H. durifolia Briq. |
– | Abaxial leaf side pubescent with caducous, appressed, white, stellate hairs; inflorescence axis angled; inflorescence 6–28 cm wide; cymes lax; widely distributed in Costa Rica and Panama | 4. H. oerstedii Briq. |
9 | Leaves markedly coriaceous; secondary and tertiary veins on the abaxial leaf side forming a reticulate network, connecting the primary veins; apical portion of flowering branch including leaves and buds densely pubescent with erect hairs; Colombia, Ecuador and Peru | 8. H. trianae Briq. |
– | Leaves papyraceous; secondary and tertiary veins on the abaxial leaf side not forming a reticulate network; apical portion of flowering branch including leaves and buds scarcely to densely pubescent with appressed hairs; Colombia and Ecuador | 9. H. goudotii Briq. |
Colombia. Cauca: Caldas, Las Pavas Cimarronas, 1200–1600 m, ♀, flowers, F.C. Lehmann 5106 (lectotype, designated by
Root-climbing liana of probably not more than 15–20 m high; functionally dioecious; free-growing branches slightly quadrangular, densely pubescent with reddish brown stellate hairs; leaves decussate, petiole sulcate adaxially, terete abaxially, color reddish brown, densely pubescent with caducous, reddish-brown, stellate hairs, 0.6–1.5 cm long, leaving a semicircular scar on the branch when leaves shed; lamina flat, ovate to lanceolate-elliptic, 6–12 cm long, 2.6–5.5 cm broad, base rounded to decurrent, sometimes asymmetric, apex acuminate, leaf margin glandular dentate, teeth generally small, larger in only a few leaves, venation brochidodromous, veins 5–7 pairs, adaxial leaf side with marked midvein, primary and secondary veins lightly marked, primary veins join to form submarginal vein, sparsely pubescent with appressed, white stellate hairs, in young leaves more dense and reddish hairs, abaxially with protruding veins, primary veins sometimes alternating protruding and marked in the same leaf, dark brown green, densely pubescent with appressed stellate reddish hairs near the midvein, rest of the lamina more sparsely pubescent, acarodomatia present, numerous, consisting of a simple cavity, but often not very conspicuous as they lay hidden under the midvein pubescence, in axils of midvein and primary veins; inflorescence axis densely pubescent with persistent reddish brown, stellate hairs, more dense towards the apex, 8–21 cm long, many-ribbed, with up to 3 opposite or decussate leaf pairs and up to 3 scars of possibly kataphyll pairs below the inflorescence, deciduous, petiole 5–15 mm long, lamina 6–12.5 cm long, 2.5–5.5 cm broad, kataphylls not seen, apex of the floral axis woody, basally quadrangular, apically triangular, elongated bract scars visible, 3–4 mm broad, 1.5–2 mm high in functionally female plants, 3.5–5 mm broad, 2–2.5 mm high in functionally male plants, inflorescence bracts cucullate densely pubescent, hairs reddish-brown, stellate, increasing in size, lowermost bract 1.5 cm large, 1.2 cm broad, other bracts not visible, inflorescences lateral (Fig.
Hydrangea caucana is known from mountain cloud forest at elevations between 750–1365 m.
Hydrangea caucana has been collected with flowers and fruits between November and March.
Hydrangea caucana should not be considered a synonym of H. peruviana and can be distinguished from the latter species by the flat, ovate to lanceolate-elliptic leaves with a glandular dentate margin. In contrast, H. peruviana is characterized by very slightly spoon-shaped, elliptic to slightly obovate leaves with a serrate to slightly dentate margin. Moreover, H. caucana is currently only known from Colombia, whereas H. peruviana is restricted to Ecuador.
We have not observed this species in the field, and herbarium labels of the known specimens of H. caucana do not record the size of the plants.
The phylogenetic relationships of Hydrangea caucana are yet unknown as there was no fresh material available for our molecular study (Granados Mendoza et al. unpublished results).
Based on the available herbarium collections, this species is Endangered according to the IUCN categories and criteria (
Colombia. Antioquia: Urrao, Aguadas, 1300 m, 8 Dec 1992, sterile, Pipoly et al. 16776 (MO); Frontino, Vereda Venados, Parque Nacional Natural Las Orquídeas, margen izq. Río Venados (Garrucha y Alto Bonito), 6°32'N, 76°19'W, 800–850 m, 30 Jan 1995, ♂, flowers, Pipoly et al. 18117 (MO, NY); same data as preceding, Parque Nacional Natural las Orquídeas, sector Calles, margen derecha del Río Calles, 1310–1365 m, ♀, fruits, 27 Mar 1988, Cogollo et al. 2669 (COL, MO); same data as preceding, 1360 m, ♂, flowers, 19 Feb 1989, Cogollo et al. 4091 (MO); Nariño: Mpio. de Ricaurte, Resguardo Indígena Nulpe Medio, camino a la quebrada La Conga, 1°6'N, 78°13'W, 750 m, 8 Jan 1996, ♂, flowers, González & Ramírez 1636 (QCA); municipio Barbacoas, corregimiento Altaquer, Vías Las Vegas, al borde del río Veza, 870 m, flower buds, Mar 1995, Fernández et al. 12459 (COL); Valle de Cauca: Cordillera Occidental, vertiente occidental, Hoya del Río Digua, lado derecho, La Elsa, 1000–1200 m, 9 Nov 1943, ♀, flowers, fruits, Cuatrecasas 15326 (F, P, US); Cordillera Occidental, vertiente occidental, Hoya del Río Digua, lado derecho, entre Queremal y La Elsa, 1200–1160 m, ♀, 27, 29 Mar 1947, ♀, flowers, Cuatrecasas 23994 (F, US).
Colombia. Norte de Santander: Pamplona, ♂, flowers, N. Funck & L.J. Schlim 1393 (lectotype, designated by
The most complete description to date can be found in the treatment by
Hydrangea durifolia is known from the type and an additional collection in northern Colombia (Fig.
The herbaria COL and F each house a black and white photo of a specimen of the Linden 1393 collection, presumably taken in the BR and G herbaria, respectively.
Because of the lack of material and therefore, the uncertainty about this species´ circumscription, it is currently not possible to present an amended description. However, we also have no elements to consider this species as a synonym of one of the other species treated here, although it is morphologically close to H. oerstedii Briq.
The phylogenetic relationships of Hydrangea durifolia are yet unknown as there was no fresh material available for our molecular study (Granados Mendoza et al. unpublished results).
It is possible this species is Critically Endangered according to the IUCN categories and criteria (
Colombia. Tolima: massif du Quindío á Portiohuelo, ♀, flower buds, flowers, J. Goudot s.n. (lectotype, designated by
Root-climbing liana of up to 10 m high; functionally dioecious; free-growing branches many-ribbed, slightly angular to quadrangular, apically with dense appressed white caducous stellate hairs; leaves papyraceous, decussate, petiole terete, rarely slightly quadrangular or sulcate adaxially, basally with broadly sulcate adaxially, color dark green, scarcely pubescent with small, white stellate hairs, 1.5–2.7 cm long, leaving a triangular scar on the branch when leaves shed; lamina elliptic to obovate, 14–25 cm long, 7–14 cm broad, base cuneate, slightly decurrent, sometimes asymmetric, apex mucronate to acuminate, leaf margin (widely) serrate, venation brochidodromous, veins 8–9 pairs, adaxial leaf side with midvein protruding along its whole length, primary and secondary veins also protruding, angle primary veins up to 50 degrees, secondary veins reticulate and connecting the primary veins resulting in a network with parallel secondary veins nearly perpendicular with respect to the primary veins, white stellate pubescence scarce, only near the leaf base, abaxially with protruding veins, opaque olive green, scarcely to densely pubescent with appressed stellate white hairs, the latter depending on the specimen, nearly sessile to shortly stalked, acarodomatia numerous, present in axils of midvein and primary veins in the lower 2/3 of the lamina, consisting of a simple cavity, glabrous or sometimes with stellate hairs in the entrance; inflorescence axis densely pubescent with appressed, white, stellate hairs (Fig.
Hydrangea goudotii A branch with leaves seen abaxially, old inflorescence axes and the vegetative portion of the flowering branch B inflorescence bud with cucullate inflorescence bracts and densely pubescent inflorescence axis C inflorescence of functionally male plant, with enlarged marginal flowers and a few flowers that still show stamens A field image of collection Granados Mendoza et al. 2012-105 B, C field images of collection Granados Mendoza et al. 2012-43.
Hydrangea goudotii is endemic to Colombia and Ecuador. Further exploration for this species throughout both countries is required, as its currently known distribution pattern is fragmentary.
This species occurs in mountain cloud forest, sometimes on heavy slopes at elevations between 1000–2500 m.
This species has been collected in flower and fruit in January, March, June, July, August, and December. Although further studies on its biology are needed, it is likely that H. goudotii is characterized by two distinct flowering and fruiting periods: December–March and June–August. However, it remains to be investigated whether these two periods take place each year.
Hydrangea goudotii should not be considered a synonym of H. oerstedii or H. peruviana var. oerstedii from which it can be distinguished by the appressed white caducous stellate pubescence on free-growing branches, these being glabrous in H. oerstedii. Moreover, H. goudotii is currently only known from Colombia and Ecuador, whereas H. oerstedii occurs in Costa Rica and Panama.
According to our molecular study, Hydrangea goudotii is closely related to H. trianae (Granados Mendoza et al. unpublished results).
Although this species has an EOO of about 154,700 km2, it is Endangered according to the IUCN categories and criteria (
Colombia. Antioquia: Urrao, Vereda Calles, Parque Nacional Natural “Las Orquídeas’, margen derecha del Río Calles, en el filo NW de la Cabaña de Calles, 1450 m, 6 Dec 1993, ♀, inflorescence buds, flower buds, Cogollo et al. 7857 (MO); Zona limítrofe del Parque Nacional Natural Las Orquídeas, vereda Calles, Alto de Palmitas, ca 1 km de la Cabaña de Calles del INDERENA, 6°32'N, 76°19'W, 1300–1400 m, 1 Dec 1993, inflorescence bud, Pipoly et al. 17468 (MO); municipio Urrao, carretera Urrao–Caicedo, 1 km antes del Alto de Caicedo, bosques a lo largo de Quebrada Villa Riaga, 6°28'N, 76°10'W, 2180 m, 5 Dec 1986, ♀, fruits, Callejas et al. 3165 (F, HUA); Alto de Cuevas, 10 km W of Blanquita, 12 km W of Nutibara, transect B, 1720 m, 3 Mar 1992, ♂, flowers, Gentry et al. 76116 (MO); municipio de Venecia, Vereda El Rincón, camino a Cerro Bravo, 5°56'19"N, 75°42'17"W, 1600–2300 m, 5 Jul 2007, flower buds, David et al. 2192 (HUA); La Ceja, 2300 m, Sep 1964, ♀, fruits, Espinal 1777 (COL). Caldas: Pueblo Rico, La Selva, 1400 m, 4 Jan 1946, ♀, flower buds, fruits, von Sneidern 5265 (AAU, C, F, GH, MICH, US); Mcpio. Pensilvania, Vereda Líbano, al lado del camino del puente de Líbano, ca. 2400 m, 11 Jul 1982, ♀, flowers, Albert de Escobar & Brand 2095 (HUA); Cauca: municipio El Tambo, Correg. La Romelia, km 75 vía a La Gallera, 1700–2000 m, 29 Jan 1995, ♂, flower buds, flowers, Ruiz et al. 361 (COL); Nariño: en la quebradita de El Osa, 1900 m, 8 Jan 1949, ♂, flowers, Uribe 1913 (COL); Risaralda: municipio de Santuario, estribación Oriental de la Cordillera Occidental, transecto de las Colonias, hacia Alto del Tigre, 2500 m, 31 Jan 1983, ♀, fruits, Torres et al. 1359 (COL).
Ecuador. Carchi: near Maldonado, 1400 m, 30 Jul 1989, ♂, flowers, van der Werff & Gudiño 10778 (QCNE); Napo: San Francisco de Borja, small forest high above banks of river beyond meadow at S. E. of town, hotel and square, 6 Jul 1980, ♂, flowers, Sobel 2392 (NY); vicinity of Baeza, 1900 m, 27 Mar 1972, ♀, flowers, fruits, Dwyer & McBryde 9601 (QCA); carretera El Chaco–Oyacachi, cerca del Río San Juan Grande, 0°16'S, 77°51'W, 1730 m, 15 Dec 1993, ♀, fruits, Freire-Fierro & Yánez 2694 (QCA); El Chaco, fincas en la parte Este del pueblo, 0°13'S, 77°48'W, 1700–2000 m, 15 Mar 1991, ♂, flowers, Gavilanes & Quezada 470 (AAU, QCA); Cerro Antisana, montane forest 2 mi. S. E. Borja, 5700 ft, 3 Aug 1960, ♀, flowers, Grubb 1216 (K, NY); Mera, surroundings of Sevilla de Oro, 6.9 airline km S [of] Shell, 1°33'40.5"S, 78°4'29.3"W, 1080 m, 15 Jun 2012, inflorescence bud, Granados Mendoza et al. 2012-43 (
Cornidia radiata
Oerst., Vidensk. Meddel. Naturhist. Foren. Kjobenhaven 1856: 42. 1856. Type. Costa Rica. Cartago: in monte Candelaria, 6000–7000 ft, Feb 1847, ♂, flowers, A.S. Oersted 1782 (lectotype, designated by
Hydrangea peruviana var. oerstedii (Briq.) Freire Fierro, Fl. Ecuador 73: 34. 2004. Type. Based on Hydrangea oerstedii Briq.
Based on (replacement name for) Cornidia radiata Oerst.
Root-climbing liana of up to 60 m high, often bending downwards, functionally dioecious; free-growing branches quadrangular, glabrous (Fig.
Hydrangea oerstedii A branch with leaves seen abaxially B flowering branch of functionally male plant with three inflorescences with enlarged marginal flowers and many flowers with large stamens C inflorescence of functionally male plant with enlarged marginal flowers, most flowers in bud with petals visible, and a few open flowers with long stamens D close-up of functionally male inflorescence E close-up of functionally male flowers with reduced pistils F close-up of functionally female flowers with reduced stamens and large pistils. A field image of collection Samain & Martínez 2012-044 B field image of collection Samain & Martínez 2013-025 C field image of collection Samain & Martínez 2012-044 D field image of collection
This species occurs in mountain cloud forest at elevations between 1100 and 2500 m. In contrast to most of the other Hydrangea species, it is common to find this species in disturbed habitats, even when there is no primary forest left and only the very humid microhabitat and protection against direct sunlight remain.
This species has been collected with flowers and fruits throughout the year.
Not to be considered as a variety of H. peruviana, from which it can be distinguished by the flat, ovate to lanceolate-elliptic leaves with white pubescence abaxially and a glandular dentate margin. In contrast, H. peruviana is characterized by very slightly spoon-shaped, elliptic to slightly obovate leaves with reddish pubescence adaxially and a serrate to slightly dentate margin. Moreover, H. oerstedii is a relatively common species in Costa Rica and Panama, whereas H. peruviana is a very rare species from Ecuador.
According to our molecular study, Hydrangea oerstedii is not related to any of the species included in this study (Granados Mendoza et al. unpublished results).
With an EOO of slightly more than 20,760 km2 and an AAO of 428 km2, this species is Near Threatened according to the IUCN categories and criteria (
Costa Rica. Alajuela: Alajuela, Distrito 3° Carrizal, 13.1 km N of Heredia via Rte. 114 (10.9 km. N of Barva), 20 ft. from rd., 10°5'26.4"N, 84°9'56.71"W, 1500–2000 m, 15 May 1990, ♀, fruits, Grantham 90.0433 & Parsons (CAS); San Ramón, Distrito 1° San Ramón, Vicinity of San Ramón, La Palma de San Ramón, 14 Dec 1926, ♀, fruits, Brenes 5221 (CR, NY); R.B. Monteverde, Cordillera de Tilarán, Sendero Rancho Alegre, 10°17'40"N, 84°45'W, 1000 m, 4 Apr 1995, ♀, fruits, Martínez 379 (CR, MO); La Palma, Picada San Antonio, 1 Dec 1922, ♀, fruits, Brenes 3807 (CR, F, GH); Reserva Biológica Bosque Nuboso de Monteverde, al lado del Sendero Chomogo, 1.15 km al NE de la caseta de entrada, 10°18'22.23"N, 84°47'11.12"W, 1710 m, 2 Aug 2012, ♀, fruits, Samain & Martínez 2012-045bis (CR,
Panama. Bocas del Toro: Cordillera de Talamanca, headwaters of the Río Culubre, 6 airline km NW of the peak of Cerro Echandi on the Costa Rican–Panamian international border, 09°05'00"N, 82°50'30"W, 2450–2600 m, 2–3 Mar 1984, ♀, fruits, Davidse et al. 25254 (CR, MEXU, MO, PMA); E branch, headwater of Colubre River, 2300–2500 m, 2 Mar 1984, inflorescence bud, Gómez et al. 22303 (MO); La Pata del Cedro, ridgetop, 09°04.270'N, 82°44.174'W, 1750 m, 11 Mar 2004, ♀, fruits, Monro & Alfaro 4315 (CR, MO, PMA); Chiriquí: 1.4 mi S of Cerro Punta, 1850 m, 28 Dec 1963, ♀, fruits, Graham 297 (MICH); along road and in pasturelands and stream banks above Las Nubes, NW of Cerro Punta, 1900–2100 m, 26 May 1973, ♀, fruits, Luteyn 3797 (DUKE, MICH); Parque Internacional La Amistad, 100 m al SSW de la oficina, 8°53'36.3"N, 82°36'58.1"W, 2207 m, 10 May 2019, ♂, inflorescence buds, flowers,
Panama. Colón, along Río Fato, 10–100 m, floral buds, H.F. Pittier 3919 (holotype: US! [00097000], isotypes: A!, BM! [BM000028808], C!, F! [V0066620F], GH! [00042780], K! [K000486137], NY! [00007170]) floral buds.
Root-climbing liana of up to 20 m high, never reaching above the lower branches of its host tree canopy (Fig.
Hydrangea panamensis A plant growing along the stem of a Cecropia B infructescence with young and mature fruits, and enlarged marginal flowers. Hydrangea peruviana C stolons with adventitious roots and decussate leaves. A, B field images of collection Samain & Martínez 2012-063 C field image of collection Granados Mendoza et al. 2012-112.
This species is known from Costa Rica and Panama (Fig.
Hydrangea panamensis grows in tropical rainforest between 200 and 1000 m elevation. It usually occurs near water streams at lower elevations. Of all the species of the present study, this is the one that grows at the lowest elevation.
Hydrangea panamensis has been collected with flowers and fruits between June and September.
Not to be considered a synonym of H. peruviana from which it can be distinguished by the rounded leaf apex with a very small acumen and the very few acarodomatia (0–2/leaf) on the abaxial leaf side vs. the acute to acuminate, rarely mucronate leaf apex and the many acarodomatia (present in both axils of midvein and primary veins and those of primary and secondary veins) of H. peruviana. Moreover, their known distribution areas are far away from each other, with H. panamensis only growing in Central America and H. peruviana being endemic to Ecuador.
The label of the specimen Gentry 5569B mentions that the flowers are red, whereas the label of the specimen Mori & Bolten 7674 says “summit of ovary of fertile flowers reddish-pink”. It seems there is some color variation in this species in both the marginal and reduced flowers. Both flower types are generally white, and this is also the color we have observed in the field ourselves.
According to our molecular study, Hydrangea panamensis is closely related to H. peruviana (Granados Mendoza et al. unpublished results).
Although this species has an EOO of about 50,123 km2, it is Endangered according to the IUCN categories and criteria (
Costa Rica. Puntarenas: Golfito, Playa Cacao, Cuenca Superior de Quebrada Nazareno, 8°37'50"N, 83°11'30"W, 200 m, 11 Jun 1994, ♂, flowers, Herrera & Rivera 7152 (CR, F, K, MO); Golfito–La Gamba (km 37), gap by the waterfall – Sendero Ozelot near the Rainforest Lodge, 8°41'N, 83°13'W, 70–300 m, 20 Jun 1997, inflorescence buds, Weissenhofer W105 (CR, WU); Golfito, Bosque de los Austriacos, Fila Gamba, Avilán woodcutter way, 8°41'N, 83°13'W, 250 m, 3 Jun 1997, ♂, old flowers, Huber & Weissenhofer 747-3.6.97 (CR); Osa, al lado del camino Rincon–Bahía Drake, 2.23 km al ONO de Rancho Quemado, 8°41'35.39"N, 83°35'7.42"W, 237 m, 8 Aug 2012, ♀, fruits, Samain & Martínez 2012-063 (CR,
Panama. Darién: Cana and vicinity, 2000–6500 ft, 7 Apr 1908–8 Jun 1908, old inflorescence branches, Williams 775 (NY); Coclé: between Cerro Pilón and el Valle de Antón, 700–900 m, 15 May 1967, ♀, flowers, Duke & Dwyer 13952 (MO); region north of El Valle de Anton, 1000 m, 27 Sep 1946, ♀, fruits, Allen 3712 (E, GH, F, MO, P, UPS); alrededores de El Valle de Antón, Altos de La Mesa, 1.3 km al N de la bifurcación hacia Río Indio, 8°38'38” N, 80°06'51"W, 735 m, ♂, flowers, 13 Aug 2014, Galdames et al. 7676 (PMA); Distrito La Pintada, cerca de las cabañas de ANAM, 8°40'05"N, 80°35'34"W, 800–900 m, 14 Aug 2007, ♀, fruits, Hernández et al. 528 (PMA); Panama: Canal Zone, Barro Colorado Island, E. Of Wheeler 13, 18 Aug 1970, ♀, flowers, Foster 1797 (DUKE, F, PMA, UC); Barro Colorado Island, forest 200 m E of Wheeler Trail 1300, 18 Aug 1970, ♀, fruits, Croat 11850 (AAU, DUKE, E, F, MO); pipeline road, 14 Sep 1971, ♀, fruits, Gentry 1792 (DUKE, F, GH, MO); Panamá, North of El Llano, 500–800 m, 25 Jul 1972, ♀, flowers, Gentry 5569B (MO); Cerro Campana, 45 km SW of Panama City on Inter-American Hwy, 8 Aug 1975, ♀, fruits, Mori & Bolten 7674 (AAU, MO); Distrito de Capira, Parque Nacional Altos de Campana, Sendero de Interpretación, 8°40'54"N, 79°55'40"O, ca. 750 m, 23 Aug 1990, ♀, fruits, Galdames 880 (PMA); Parque Nacional Altos de Campana, Sendero de interpretación, 1 km al este del campamento de los guardaparques de INRENARE, parcela 10-8, 8°40'N, 79°55'W, 800–900 m, 19 Aug 1993, ♂, flowers, Correa & Montenegro 9780 (PMA, UCH).
Cornidia peruviana (Moric. ex DC.) Small, North American Flora 22(2): 161. 1905.
Ecuador. “In Peruvia prope Huyaquaquil”, ♀, fruits, J.A. Pavón s.n. [J.J. Tafalla s.n.] (holotype: G! [G00301424], isotypes: F!, MA! [MA-811940])
Root-climbing liana of up to 30 m high, up to 20 cm diameter, functionally dioecious; leaves decussate, coriaceous, petiole sulcate adaxially, clasping its branch, color reddish brown, densely pubescent with partially caducous, reddish simple and stellate hairs, 0.5–2 cm long, leaving a semicircular scar on the branch when leaves shed; lamina very slightly spoon-shaped (Figs
Hydrangea peruviana A branch with inflorescence B branch with leaves seen abaxially and inflorescence C close up of infructescence with maturing fruits and enlarged marginal flowers. A, B field images of collection Granados Mendoza et al. 2012-111 C field image of collection Granados Mendoza et al. 2012-112.
Hydrangea peruviana is a rare species which is known from Ecuador only (Fig.
This species has been reported in rainforest and cloud forest at elevations between 682 and 1300 m.
This species has been collected with flowers and fruits in March and July. Only female plants have been observed. There are no collections known of male individuals.
Since the revision by
In contrast to what might be expected because of its name, H. peruviana is not known from Peru, the type locality area of Guayaquil now being the second largest city of neighboring Ecuador, and this country´s main harbor. However, at the time of its collection in the late 18th century, modern-day Peru and most of Spanish-ruled South America belonged to the Viceroyalty of Peru.
As mentioned by
According to our molecular study, Hydrangea peruviana is closely related to H. panamensis, the two of them unrelated to the other species of this study (Granados Mendoza et al. unpublished results).
Although this species has an EOO of about 13,515 km2, it is Endangered according to the IUCN categories and criteria (
Ecuador. Carchi: Camino Chical–Peñas Blancas–Tobar Donoso, colec. a 5 horas de camino, 1°0'N, 78°12'W, 6 Dec 1993, 1200 m, sterile, Freire-Fierro 2616 (AAU, QCA); Esmeraldas: environs of Lita, on the Ibarra–San Lorenzo R.R., 550–650 m, 11 Jun 1978, ♀, fruits, Madison et al. 5251 (F, QCA); Pichincha: km 87–84 old road Quito–Santo Domingo, 1200–1300 m, 21 Mar 1980, ♀, fruits, Dodson 9733 (MO); Los Ríos, Road Patricia Pilar–Montañas de Ila, km 18, N side of Torre de Bijagual, below antenas, 00°38'S, 79°17'W, 620–680 m, 28 Feb 1993, ♀, fruits, Øllgaard & Borchsenius 100686 (QCA, QCNE); Santo Domingo de los Tsáchilas: 5.3 airline Km SW of Corina Parral, 0°39'20.8"S, 79°17'27.7"W, 693 m, 11 Jul 2012, ♀, fruits, Granados Mendoza et al. 2012-111 (
Colombia. Norte de Santander, Ocaña, without date, ♂, old flowers, J.J. Linden 1139 (lectotype, designated by
The most complete description to date can be found in the treatment by
This species is currently known from the type locality in Colombia only (Fig.
This species concerns a very distinct taxon with more enlarged “marginal” flowers than any other member of Cornidia. In fact, from the type specimen it seems that most flowers possess enlarged sepals. It is only known from its type collection made around 1850 and to our knowledge has not been collected since. It cannot be excluded that it concerns a local “mophead” mutation, but in the absence of recently collected material and as the leaf morphology of the type specimens cannot be matched with better-known species, we decide here to recognize this taxon as a distinct species, although we cannot present an amended description at this time, given that we only dispose of the type specimen, which already was given an excellent description by
As in the case of H. durifolia, exploring field work in forests around Ocaña in the Colombian department of Norte de Santander might lead to the rediscovery of this species, which will be helpful to define its taxonomic status.
The phylogenetic relationships of Hydrangea schlimii are yet unknown as there was no fresh material available for our molecular study (Granados Mendoza et al. unpublished results).
H. lehmannii
Engl., Nat. Pflanzenfam. (ed. 2) 18a: 207. 1930. Type. Colombia. Valle del Cauca: Cali, 4 Oct. 1900, ♀, fruits, F.C. Lehmann 9074 (lectotype, designated by
Hydrangea platyphylla
Briq., Annuaire Conserv. Jard. Bot. Genève 20: 401. 1919. Type. Colombia. Tolima: Mariquita, ♂, old flowers, J.J. Linden 894 (lectotype, designated by
Colombia. Tolima: Quindío, ♂, flowers, J.J. Triana s.n. (holotype: G-DC n.v. (photo in
Root-climbing liana of up to 20 m high, functionally dioecious; free-growing branches many-ribbed, slightly angular, old branches quadrangular, with erect, white stellate hairs (Fig.
Hydrangea trianae A branch with leaves seen abaxially and inflorescence buds B functionally female inflorescence with enlarged marginal flowers C basal portion of functionally male inflorescence with densely pubescent apex of the inflorescence axis, and a few open male flowers with large stamens D close-up of functionally female flowers with petals, reduced stamens and large pistils. A, C field images of collection Samain et al. 2011-064 B, D field images of collection Samain et al. 2011-067.
Hydrangea trianae is known from cloud forest and remnants of this vegetation type at elevation between 400–3680 m. Moreover, it has been noted to occur in disturbed or late secondary forests.
This species has been collected with flowers and fruits throughout the year.
Hydrangea trianae should not be considered a synonym of H. peruviana, from which it can be distinguished by the notoriously coriaceous leaves, the abaxial reticulate network of secondary and tertiary veins connecting the primary ones, and the abaxial white pubescence.
Hydrangea lehmannii was synonymized with H. peruviana by
With respect to the type material of Hydrangea lehmannii,
With respect to the type material of H. platyphylla,
One of the F specimens (V0066626F), which supposedly is an isotype of H. platyphylla, does not belong to this species based on leaf venation, probably due to a labeling error; however, it is not possible to propose any other identification as the material is very limited.
The specimen of Triana 4668 in NY does not correspond with H. trianae or any of the other species treated here, in contrast with the specimen with the same number in COL. Triana’s numbers do not correspond to field collection numbers as currently used, but instead are generic numbers based on the Endlicher system (
According to our molecular study, Hydrangea trianae is closely related to H. goudotii (Granados Mendoza et al. unpublished results).
Although this species has an EOO of nearly 726,300 km2, it is Endangered (B2abiii) according to the IUCN categories and criteria (
Colombia. Antioquia: Argelia, km 17 of road Sonsón–Argelia (2 km past fork in road to Nariño), 2160 m, 3 Oct 1987, ♀, fruits, Zarucchi et al. 6164 (HUA, MO); Frontino, Correg. Nutibara, Nutibara–La Blanquita road, region of Murrí, Alto de Cuevas, 6°45'N, 76°20'W, 1700–1800 m, 19 Apr 1988, ♂, old flowers, Luteyn et al. 12047 (HUA, K, MO); Urrao, Parque Nacional Natural las Orquideas, vereda Calles, 06°32'N, 76°19'W, 1350–1450 m, 5 Dec 1993, ♀, inflorescence buds, flowers, Pipoly et al. 17737 (MO); Jardín, km 20 of road Jardín–Riosucio (dpto. Caldas), ca 15 km SSE of Jardín, Alto de Ventanas, 29 Oct 1988, 2700–2790 m, ♂, flower buds, Zarucchi et al. 6962 (HUA, MO); road between Jardín and Río Sucio, ca 9 km from Jardín, 2300–2400 m, 29 Jan 1989, ♂, flowers, MacDougal & Roldán 3581 (HUA, MO); Frontino, region of Murrí, road between Nutibara and La Blanquita, 19.2 km from centro de Nutibara, 1560 m, 11 Feb 1989, ♂, old flowers, MacDougal et al. 3929 (CR, HUA, MO); Frontino, Corregimiento Nutibara, Región Murí, márgen de río, 1500 m, 11 Jul 1986, ♀, old inflorescences, Acevedo et al. 1260 (HUA, NY, US); along road between Caramanta and Supia (Risaralda), ca 4 mi S of Caramanta, 2030 m, 27 Jan 1990, ♀, flowers, Croat 70038 (MO); Medellín y Guarne, Parque Ecológico Piedras Blancas, laguna de Guarne, 2350 m, 26 Aug 1995, ♂, old inflorescences, Roldán & Medina 2385 (HUA, MO); Parque Ecológico Piedras Blancas, sector Lajas, 2350 m, 23 Sep 1995, ♂, flowers, Fonnegra et al. 5643 (HUA, MO); Parque Nacional Natural “Las Orquideas”, sector Calles, margen derecha del río Calles, 1250 m, 30 May 1988, ♀, fruits, Cogollo et al. 3087 (COL, MO); Urrao, Corregimiento Encarnación, sitio El Río, 1 hora de camino del Páramo de Frontino, 3120 m, 7 Apr 1989, inflorescence buds, Callejas et al. 7774 (HUA, NY); same data as preceding, 3000–3150 m, 8 Apr 1989, ♀, flower buds, flowers, Callejas et al. 7835 (HUA, MO); Envigado, Vereda “El Escobero”, Costa Occidental Cerro “San Luis”, 2100–2250 m, 15 May 1996, ♂, inflorescence bud, flower buds, flowers, Correa & Carona 1073 (COL, HUA, MO); cerca de San José de San Andrés, 1 May 1948, inflorescence buds, floral buds, Correa & Velasquez 45 (US); bosque bajo de la cumbre cerca de Santa Elena, camino entre Medellín y Río Negro, 2300–2500 m, 3 Aug 1945, ♀, fruits, Medina 239 (US); municipio de Caldas, Alto de San Miguel, 2200 m, 8 Jul 1996, ♂, flowers, Monsalve 38 (F); municipio de Peque, 7°2'6.7"N, 75°58'27"W, 2500 m, 19 Nov 1995, ♀, fruits, Benítez et al. 584 (COL); municipio de Peque, vereda Romeral, 6°59'18.4"N, 75°58'02.5"W, 2650 m, 16 Nov 1995, floral buds, Benítez et al. 452 (COL); municipio Caldas, Reserva “Alto de San Miguel”, cuenca alta del río Medellín, 6°5'N, 75°38'W, 2100 m, ♂, old flowers, Roldán et al. 2516 (HUA); municipio de Caldas, vereda La Corrala, Finca “La Zarza”, Alto del Gallinazo, ca. 2600 m, 4 Feb 1986, inflorescence buds, Albert de Escobar y Stein 6238 (HUA); municipio de Urrao, Finca La Clara, cerca del río Mina y La Clara, 2340 m, 6 Dec 1984, ♀, fruits, Orozco et al. 1782 (COL); camino de Urrao al páramo de Frontino, 2400 m, 19 May 1985, ♂, flower buds, flowers, Cárdenas 128 (HUA); Argelia, vereda La Vibora, antiguo camino de Sonsón a Argelia, 2200–2700 m, 28 Aug 1989, ♂, flowers, Hoyos y Avendaño 1300 (HUA); municipio Ríonegro, vereda Yarumales, 20–30 km SE de Medellín en la vía a Rionegro, 6°15'N, 75°28'W, 2140–2410 m, 12 Nov 1990, ♀, fruits, Callejas & Roldán 9628 (HUA); municipio de Urrao, Páramo de Frontino, El Río, 3020 m, 6 Jan 1985, ♀, fruits, Londoño et al. 669 (HUA); Páramo de Frontino, Urrao – zona situada entre el 15 y la Esperanza, 2980–3680 m, 18 May 1985, ♀, fruits, Rentería 4082 (HUA): Medellín, corregimiento de Santa Elena, Sector de Piedras Blancas, 6°17'N, 75°32'W, 2400 m, May 1998, ♂, flowers, Cardona y Alzate 551 (HUA); Medellín, corregimiento de Santa Elena, Reserva Natural Montevivo, sector Casapalo, 6°12'48"N, 75°29'32.2"W, 2500 m, 30 Sep 2002, ♀, fruits, David & Idarraga 291 (HUA); municipio de Caldas, vereda La Corrala, Finca “La Zarza”, Camino La Zarza – Alto del Gallinazo, ca. 2440 m, 10 Nov 1987, ♂, old flowers, Albert de Escobar et al. 8022 (HUA); municipio de Urrao, Páramo de Frontino, 6°29'N, 76°7'W, 2890–2250 m, 16 May 1987, ♂, old flowers, Estrada et al. 108 (HUA); municipio de Medellín, corregimiento de Santa Elena, vereda El Llano y Perico, cabeceras y recorrido Q. San Pedro, 2550 m, 8 May 1996, ♀, fruits, Giraldo et al. 785 (HUA); Caldas: Vereda la Corrala, Finca La Zarza, al lado del camino entre la cascada y el Alto del Gallinazo, 2550 m, 27 Apr 1987, floral buds, Albert de Escobar 7576 (HUA, UC); Chocó: municipio San José del Palmar, hacia el galápago, 1380 m, 11 Nov 1985, ♀, fruits, Lozano et al. 4920 (COL); Cundinamarca: San Francisco, hacienda “La Laja”, 2880 m, 30 May 2004, ♂, flowers, Parra et al. 465 (COL); Huila: municipio de la Argentina, Quebrada del pueblo, 1850 m, 25 Sep 1984, ♀, fruits, Lozano et al. 3981 (COL); Nariño: Mpio. de Ricaurte, Reserva Indígena Gualcalá, Santa Fé, camino al Río Gualcalá, 1°18'N, 77°54'W, 1100–1200 m, 18 Dec 1995, inflorescence buds, floral buds, Ramírez & González 9135 (QCA). Quindío: Parque Nacional de Cocora, Quebrada Santa Lucía, 3000 m, 08 May 1990, ♀, flowers, Franco 3095 (NY); Risaralda: municipio de Pereira, corregimiento La Florida, SFF Otún Quimbaya, 4°44'17"N, 75°34'01"W, 1900 m, 22–28 Feb 2004, floral buds, Alzate et al. 1962 (F); municipio de Pereira, orilla del camino entre Ceylan y el Cedral, 2150 m, 4 Dec 1989, ♀, fruits, Franco et al. 2929 (COL); Mpio. Pereira, Parque Nat. Reg. UCUMARI, 2200–2450 m, ♀, fruits, González 1635 (COL); municipio de Pereira, Parque Regional Ucumari, vereda la Pastora, camino de el Ceilán a El Cedral, borde de la quebrada Miraflores, 2340 m, 30 Jul 1989, floral buds, Galeano 222 (COL); municipio de Apía, vereda “La Cumbre”, 2285 m, 24 Feb 1983, ♀, fruits, Torres et al. 2192 (COL); Tolima: Quindio, El Roble, old inflorescence axis, Triana 358 (US); same data as preceding, ♂, flowers, Triana 359 (US); same data as preceding, ♂, flowers, Triana 2805 (US); Valle del Cauca: San Antonio, about 25 km W of Cali, 1800 m, 25 Jan 1988, ♂, flowers, van der Werff & Giraldo 9746 (MEXU, MO); same data as preceding, van der Werff & J. Giraldo 9747 (CR, MO); Cordillera Occidental, vertiente occidental, Hoya del Río Digua, lado izquierdo, Piedra de Moler, 900–1180 m, 19–28 Aug 1943, ♀, fruits, Cuatrecasas 14974 (F); Cali, forests of West Cordillera above Cali, 2000–2200 m, June 1900, ♀, young fruits, B.T. 623 (NY); Finca Zingara, km 18 de la carretera Calia–Buenaventura, km 4 vía a Dapa, corregimiento de la Elvira, cordillera Occidental, 1900 m, 23 Jan 1994, ♀, flowers, Giraldo-Gensini & Agredo 130 (MO); same data as preceding, 26 Feb 1995, ♀, fruits, Giraldo-Gensini & Agredo 611 (MO); same data as preceding, 25 Aug 1996, ♀, fruits, Giraldo-Gensini & Corrales 765 (MO); Cuenca del Río Cali, cercanías de Peñas Blancas, 10 Jan 1963, sterile, López Figueiras 8139 (US); Cordillera Occidental, vertiente oriental, Hoya del río Cali, río Pichindé, entre los Cárpatos y El Olivo, 2920–2025 m, 5 Aug 1946, inflorescence buds, Cuatrecasas 21935 (F, P, US); Cordillera Occidental, Los Farallones, extremo N, vertiente oriental, Almorzadero, bosques, 2980 m, 13 Oct 1944, ♂, flowers, Cuatrecasas 18113 (F, US); Yumbo, Finca La Samaria, NE of Darien, near Lago Calima (reservoir), 1700–1750 m, 14 Feb 1984, ♀, flowers, Juncosa 2181 (MO); Peñas Blancas, los Cárpatos, 1600–1700 m, 9 Sep 1988, inflorescence buds, Franco et al. 2509 (COL, MO); Bosque de San Antonio, W of Cali, near television tower, 2250 m, 15 Jul 1984, ♀, fruits, Gentry 48079 et al. (MO); Buga, Carretera a El Retiro, a 1 km de carretera Buga–El Placer (desvío a Retiro está a 12 km antes de El Placer), Cordillera Central, vertiente occidental, al lado de carretera, 2300 m, 15 Sep 1991, ♂, flowers, Silverstone-Sopkin & Giraldo-Gensini 6400 (MO); Restrepo, Río Calima, Cusumbo, 680 m, 21 Feb 1989, ♀, fruits, Devia & Prado 2581 (US); El Silencio, Hacienda Himalaya, Cordillera Occidental W of Yumbo, transect 1, 1860 m, 2 Jan 1989, ♂, flowers, Gentry et al. 65414 (MO); Argelia, Vereda Las Brisas, 2050–2200 m, 21 Jan 1983, ♀, fruits, Díaz P. 3854 (COL, MO); Manizales, vereda La Esperanza, Reserva Torre Cuatro, cerca de la quebrada La Siberia, 5°1'41"N, 75°23'10"W, 2650–2750 m, 28 Mar 1999, ♂, old flowers, Alvear et al. 314 (COL); Manizales, 1851–1857, ♂, old flowers, Triana s.n. (K); Neira, Cementos Caldas, 5°9'45"N, 75°29'43.1"W, 2203 m, 29 Jul 2004, ♀, fruits, Mancera 553 (COL); Manizales, Reserva Río Blanco, 2600 m, 28 Feb 2003, sterile, Sanin 39 (HUA); Manizales, Río Blanco, El Paso nivel, 2500 m, 10 Jul 2003, sterile, Sanin 958 (HUA). Cauca, Parque Nacional Munchique, El Tambo, vereda La Romelia, la Gallera, 1950 m, 25 Jul 1993, ♀, fruits, Velayos et al. 6891 (COL); El Tambo, Parque Nacional Munchique, vereda La Romelia, la Gallera, 2385 m, 26 Jul 1993, ♂, old flowers, Velayos et al. 6917 (COL). With three localities: Quindio: Mariquita, Roble, 2080 m; Antioquia, Manizales, 2140 m: Buenaventura, Cordillera Occidental, 2190 m, Jul 1833, ♂, old flowers, Triana 4668 (COL); same data as preceding, floral buds, Triana 1866 (photo in F of a specimen in G).
Ecuador. Carchi: prominent hillcrest directly N of Lita, on N side of Río Mira and just to E of Río Baboso, on steep W-facing slope, 760 m, 7 Aug 1994, sterile, Boyle 3476 (MO); along Chical–Pailon–San Marcos trail, 01°06'N, 78°18'W, 600–1200 m, 18–19 Nov 1983, ♂, flowers, Kvist et al. 48670 (AAU, QCA, QCNE); El Pailon, ca. 45 km above Maldonado along a footpath to Tobar Donoso, 0°49'S, 78°9'W, 800 m, 1 Dec 1979, ♀, fruits, Madison & Besse 7226 (QCA); Esmeraldas: Quininde, Bilsa Biological Station, Montañas de Mache, 35 km W of Quininde, 5 km W of Santa Isabel, slope between banana plantation and stream SE of Station, 400–600 m, 9 Dec 1994, ♀, fruits, Bass & Pitman 320 (MEXU, MO, QCNE); same data as preceding, 31 Dec. 1994, ♀, flowers, Pitman & Marsh 1133 (MO, QCNE); San Lorenzo, Alto Tambo, Frente Finca del Sr. Lalama, a 1.5 km del sector de El Cristal, 650 m, 13 May 1992, ♂, flowers, Quelal & Luteyn 489 (MO, QCNE); Imbabura: Cordillera Occidental, La Union, lower Intag Valley, 4600 ft, 20 Sep 1944, ♀, flowers, Drew E-684 (NY, US); Los Ríos: Quevedo, Centinela–La Pirámide, vía Santa Domingo de los Colorados–Quevedo entrando por Patricia Pilar km 41, 650 m, 25 Feb 1992, ♂, flowers, Quelal & Tipaz 347 (MO, QCNE); Collections from path following ridge line at El Centinela at crest of Montañas de Ila on road from Patricia Pilar to 24 de Mayo at km 12, Patricia Pilar is at Km 45 on road from Sto. Domingo to Quevedo, 600 m, 4 Feb 1983, ♀, fruits, Dodson 13654 (MO, QCNE); same data as preceding, 6 Apr 1980, inflorescence bud, Dodson s.n. (MO); Napo: Quijos, along road 1 km SE of Cosanga, 00°35'S, 77°52'W, 2000 m, 31 Jul 1990, floral buds, Webster & Richerson 28512 (QCNE); El Chaco, San Juan Chico, camino hacia el Río Oyacachi, 00°17'S, 77°52'W, 2000 m, ♀, fruits, Alvarez 97 et al. (QCNE); Baeza–Sardinas Alto, col. en borde de carretero, al NO de Sardinas Alto, a 1.5 km, 00°22'S, 77°52'W, 1780 m, 15 Dec 1993, inflorescence buds, Freire-Fierro & Yánez 2693 (QCA, QCNE); hills above R. San Juan, at confluence with R. Oyacachi, ca 10 km W of El Chaco, 00°17'S, 77°51'W, 1800–2000 m, ♀, fruits, Ståhl et al. 2354 (QCA); Napo–Pastaza, Mera, 26 Mar 1940, ♀, fruits, Lugo 126 (C, MO); Pichincha: along road and trail from Maquipucuna Lodge to Ecolodge Santa Lucia, 2 km N of Maquipucuna entrance, 1400 m, 15 Mar 2006, ♀, fruits, Croat 95938 (MO); Maquipucuna, 5 km E of Nanegal, Transect # 5, 1550 m, 10 Feb 1991, ♀, flowers, Gentry & Valencia 73174 (AAU, MO, QCNE); Quito, Nanegal, Bosque protector Maquipucuna, 5 km airline SE of Nanegal, disturbed rainforest above Río Tulambi, 00°07.5'N, 78°38.5'W, 1500 m, 31 Aug 1993, ♂, old flowers, Webster et al. 30003 (QCNE); Reserva Biológica Maquipucuna, Nanegal, colectada en primer lindero, camino del Inca, 1600 m, 9 Jan 1992, ♂, flowers, Tipaz et al. 593 (AAU, MO, QCNE); Maquipucuna Tropical Reserve, western slopes of Andes, northern boundary of reserve, 10 km north of Nanegalito, 1200 m, 2 Dec 1988, inflorescences, floral buds, Neill et al. 8643 (MO, QCNE); Reserva Biológica Maquipucuna, 1300 m, 31 Jan 1991, ♀, flowers, Alvarado 406 (COL, MO, QCNE); along road and trail from Maquipucuna Lodge to Ecolodge Santa Lucia, 2 km N of Maquipucuna entrance, 00°07'19"N, 78°37'06"W, 1400 m, 15 Mar 2006, ♀, fruits, Croat et al. 95939 (MO, QCNE); Mindo, on western slopes of Andes, 1200 m, 16 Apr 1994, ♂, flowers, Neill & Asanza 10349 (MO, QCNE); 11 km W of Tandapi, trail along Chictoa River, tributary of Río Pilatón, 1350–1550 m, 26 Oct 1974, ♀, fruits, Gentry et al. 12073 (MO, QCA); Tungurahua: Baños, 2.8 airline km SSW of Río Negro, near El Coral, 1481 m, 1°25'59.5"S, 78°12'1.5"W, 13 Jun 2012, sterile, Granados Mendoza et al. 2012-38 (
Peru. Amazonas: Bagua, Aramango, 8.86 km SE of Aramango, 1.23 km SW of Nueva Esperanza, 1806 m, 5°28'28.9"S, 78°23'08.4"W, 3 Jul 2011, sterile, Samain et al. 2011-110 (
Peru. Amazonas: Cheto, ♀, inflorescence bud, flower buds, flowers, A. Weberbauer 4372 (holotype: B (destroyed), F neg. 4146); lectotype: F! (fragment)). Peru. Amazonas: Chachapoyas, Soloco, Paraje Olia, 1.44 km S of Quitachi, W of road to Pampa del Tío, 6°20'2.85"S, 77°41'2.91"W, 2304 m, 22 Jun. 2011, ♀, flowers, M.S. Samain et al. 2011-062 epitype, designated here: USM!; isoepitypes:
Root-climbing liana of up to 20 m high, functionally dioecious; free-growing branches quadrangular, slightly fissured; leaves decussate, petiole flattened adaxially, sulcate in young leaves (Fig.
Hydrangea weberbaueri A apical portion of a branch with young leaves where the characteristic leaf venation and margin can be observed B adaxial leaf side showing the typical leaf venation of this species C glabrous apex of the inflorescence axis of a functionally female inflorescence. A field image of collection Granados Mendoza et al. 2012-23 B field image of collection Samain et al. 2011-056 C field image of collection Samain et al. 2011-068.
Hydrangea weberbaueri is known from Colombia, Ecuador and Peru (Fig.
This species grows in tropical rainforest and mountain cloud forest at elevations between 974–3500 m.
This species has been collected with flowers and fruits throughout the year.
The following vernacular names are indicated on the specimen Ellemann 75381 (AAU, LOJA, MO, QCA): bejuco matapalo (Spanish), yura huanutic caspic (Quichua). Its use as fuel wood is also indicated on the label of these specimens. This is the only species of this study of which vernacular name and use have been recorded.
Not to be considered a synonym of H. peruviana from which it can be distinguished by the notorious regular venation pattern with primary veins and secondary veins parallel to each other, all of them arching towards the apex.
No duplicate material of the original type collection has been located in the herbarium MOL of the Universidad Nacional Agraria La Molina in Lima, Peru, nor any other herbarium where we have searched. The fragment held in F that is the obligate lectotype is fragmentary and does not have the diagnostic features of this taxon, therefore we designate an epitype here for diagnostic purposes. The epitype specimen was collected approximately 8.5 km south of the original type location, which at the time of our field work was the nearest location to Cheto with primary forest. Unfortunately, satellite images (
A Weberbauer collection with the same number as the original type is present in the herbarium USM, but this is an entirely different species (with branched inflorescences) which does not belong to this group. The label on the USM specimen is not the original Weberbauer label, suggesting this represents a labeling error.
The leaf venation in the specimens from Colombia does slightly differ from that in the material from Ecuador and Peru, although it is still very recognizable. Therefore, we decided to include these in H. weberbaueri, although future studies on Colombian hydrangeas may well show it concerns a distinct and new species.
According to our molecular study, Hydrangea weberbaueri is not closely related to any of the species mentioned here, but it occurs in a clade which is sister to the one with H. goudotii and H. trianae (Granados Mendoza et al. unpublished results).
Hydrangea weberbaueri A branch with inflorescences with enlarged marginal flowers B functionally male inflorescence with enlarged marginal flowers, and reduced flowers with large stamens C functionally female inflorescence with enlarged marginal flowers D close-up of functionally male flowers with large stamens and reduced pistils E close-up of functionally female flowers with reduced stamens and large pistils. A, B field images of collection Granados Mendoza et al. 2012-16 C field image of collection Granados Mendoza et al. 2012-21 D field image of collection Samain et al. 2011-068.
Although this species has an EOO of nearly 660,000 km2, it is Endangered according to the IUCN categories and criteria (
Colombia. Cundinamarca: Quebrada la Aguadita, 2 km arriba de la carretera a Fusagasugá, 1800 m, 28 Nov 1960, ♀, fruits, Idrobo 4186 (COL); Cordillera Oriental, municipio San Bernardo, Vereda de Venecia, margen izquierda del río Chorrera, afluente del Sumapáz, 2450 m, 21 Jul 1981, floral buds, Jaramillo et al. 6844 (COL); Sasaima, vereda San Bernardo, quebrada La María y Río Dulce, 1800 m, sterile, Jan 1958, sterile, García-Barriga 15757 (COL); Santander: municipio Suaita, corregimiento San José de Suaita, carretera a La Veterana, bosque al lado de la carretera, vereda San Imidio, 6°9'44.5"N, 73°25'09.4"W, 1800–1910 m, 6 Apr 2003, ♀, fruits, Betancur et al. 10122 (COL, HUA).
Ecuador. Azuay: the eastern Cordillera, 1–8 km, north of the village of Sevilla de Oro, 8000–9000 ft, 27 Jul–12 Aug 1945, inflorescence buds, Camp E-4464 (AAU, F, MO); Sevilla de Oro, ca 1.7 airline km E of Osorancho (El Cisne), N Sevilla de Oro, 2°45'42"S, 78°37'38.1"W, 2804 m, 8 Jun 2012, sterile, Granados Mendoza et al. 2012-22 (
Peru. Amazonas: Bongará, Jumbilla, Bosque de Protección Alto Mayo, surroundings of Chichilac, 4.79 km NNE of Jumbilla, 5°51'51.33"S, 77°47'16.7"W, 2462 m, 10 Dec 2013, ♀, old fruits, Samain et al. 2013-130 (
The review of this morphological species complex surrounding Hydrangea peruviana and H. oerstedii of mostly red to purple flowered Central and South American climbing Hydrangeas attempts to unravel the taxonomic chaos in the group, based on the most complete sampling to date of herbarium specimens, as well as our own field collections and observations throughout their distribution area. Very importantly, type material was studied in all cases, generally the specimens themselves, rarely high-resolution scans on JSTOR Global Plants or shared by the curators of the respective herbaria. Many taxa had been synonymized, as important characters were, following their first descriptions in the 19th and early 20th centuries (especially by
This study, as are those of other “species complexes” we are currently studying, e.g., Hydrangea asterolasia Diels, Hydrangea preslii Briq. and Hydrangea species with branched inflorescences, is extremely complicated, and the result of a decade of field and herbarium work, matching and re-matching specimens with type material and descriptions, comparisons with studies by
As a result of our study, the number of accepted species in Cornidia currently totals 26. A complete key for morphological identification of all Neotropical species can currently not be provided as the species we have not yet treated (with the exception of this study, and those by
In his otherwise excellent revision of the then known species of Cornidia, including the description of 11 new species in the section, all unusually elaborate for that time,
A surprisingly high number of herbarium labels of the specimens studied here, as well as some references (e.g.,
Of the nine species mentioned above, five are relatively widespread, whereas four have a relatively to very restricted distribution. On the one hand, the taxonomic and conservation statuses of two endemic Colombian species, H. durifolia and H. schlimii, are not completely clear in the absence of more and/or recent collections. A third species, H. caucana, endemic to Colombia as well, also does not have recent collections, but the existing ones do allow to understand its distribution pattern in the western cordillera. The fourth species with a restricted distribution is H. peruviana from Ecuador. On the other hand, H. oerstedii is the most widespread species studied here, known from more than 150 collections from Costa Rica and Panama. Despite the fact that this species grows in montane cloud forest, which is seriously being reduced throughout is whole distribution area (see
The overall distribution area of the species studied here is from northern Costa Rica to central Peru, which is the area with the highest diversity of Neotropical Hydrangeas, apart from Mexico (
Although a few populations of most of the species studied here do occur in protected areas, their long-term conservation is seriously compromised because of the significant threats from which their habitats suffer throughout their distribution area, related to fragmentation, land use change and climate change. Moreover, it is likely their dioecism makes these species even more prone to local extinction because of habitat destruction and fragmentation, as female and male individuals generally do not occur near each other, and their functional biology is not well-understood to date. During our field work, especially in Peru, where Hydrangeas often occur on private properties, in contrast with e.g. Costa Rica and Panama, where many collections were made in areas protected at national level, we have tried to sensitize the land-owners about the uniqueness of this group as a whole. Derived of the current contribution, we will also be able to do so at species level, which is especially important for the threatened taxa.
Finally, it is difficult to believe – though not surprising taking into account our recent discoveries of seven new species in Mexico (
We are grateful to the authorities of Costa Rica, Ecuador, Panama and Peru for issuing the collection permits 020-2012-SINAC, 041-2012-SINAC, 001-12-IC-FLO-DNB/MA, SE/P-11-19, 0124-2011-AG-DGFFS-DGEFFS and 0395-2013-MINAGRI-DGFFS/DGEFFS, respectively. We thank the curators of all abovementioned herbaria for loaning the material required for this study and Laurence Loze of the herbarium G of the Conservatoire et Jardin botaniques de la Ville de Genève, for kindly sharing photos of the original syntypes of Hydrangea goudotii. Thanks to Alina Freire-Fierro, Maarten Christenhusz for their valuable comments on this manuscript and to Sandy Knapp for the detailed review and suggestions that have greatly improved this study. In Colombia, we thank Ricardo Callejas and Favio González for permission to take pictures in the herbaria HUA and COL, respectively. In Costa Rica, we thank Frank González, Barry Hammel, Esteban Jiménez, Julio Morales, and Nelson Zamora for help with logistics and sharing their knowledge. In Ecuador, Diana Fernández, Efraín Freire, Edison Jiménez, and Juan C. Cerón Factos for their invaluable help obtaining the research and collection permits and/or performing field work. In Panama, María de Stapf, Vielka Murillo, and Jorge Valdés Sánchez for their support in obtaining the collection and export permits and field work logistics. In Peru, Joaquina Albán, Carlos Bambarén, Asunción Cano, Mercedes Flores, Víctor Morales, Guillermo Pino, Rigoberto Rivera (who also helped in Ecuador), Rocío Rojas, Merly Saavedra, Rodolfo Vázquez for their help with all herbarium, permit and field logistics. In Peru, we are especially grateful to the local guides, los grandes conocedores, the great connoisseurs, as we used to refer to them in the field, many of whom have Hydrangeas on their private properties. We dedicate this manuscript to these people from the Peruvian countryside, especially to Lino Rojas from Chichilac, Jumbilla, and the children of Nueva Esperanza, Aramango, both in the department of Amazonas.
Funding
Field work was funded by the Bijzonder Onderzoeksfonds of the Ghent University, Belgium, the Research Foundation Flanders, the Systematics Association and private funds of the first author and of Rita Carron, Gent, Belgium.