Hydrangea goudotii Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 404. 1919.

Samain, Marie-Stephanie, Granados Mendoza, Carolina & Martinez Salas, Esteban Manuel, 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, pp. 91-153 : 91

3. Hydrangea goudotii Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 404. 1919. Figures 5 , 6

Type.

Colombia. Tolima: massif du Quindío á Portiohuelo, ♀, flower buds, flowers, J. Goudot s.n. (lectotype, designated by McClintock 1957, pg. 238 [as “holotype”]): G! [G00418961]; isolectotype: F! [F0066624F]).

Description.

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. 5B ), 6-15 cm long, gradually broadening towards the apex, with 2 opposite leaf pairs along the axis and one opposite kataphyll pair immediately below the inflorescence, deciduous, petiole 1 cm long, lamina 3.5-9.3 cm long, 2.5-6 cm broad, abaxially densely pubescent with white stellate hairs, apex of the floral axis woody, cone-shaped, elongated bract scars visible, thickening at the top, 8-9 mm broad, 5-6 mm high in functionally female plants, 6-7 mm broad, 2-2.5 mm high in functionally male plants, inflorescence bracts cucullate, dark green to light pink (Fig. 5B ), coriaceous, abaxially densely pubescent with whitish stellate hairs, margin membranous and glabrous, veins not visible as a consequence of the pubescence, bracts increasing in size distally, consecutively and rapidly deciduous during inflorescence development, inflorescences lateral (Fig. 5C ), opposite, 1-4 pairs of inflorescences per flowering branch, up to 3 decussate small leaf pairs (or their scars) between the two inflorescence pairs, flowering branch continues growing vegetatively very rapidly during inflorescence development, with already up to 4 decussate leaf pairs above the inflorescences when the upper inflorescences are still in bud, linear, with dense white stellate hairs, inflorescence umbellate, buds up to 3.4 cm broad and 3 cm high before opening, in flowering stage 5-12 cm diameter, 2.5-8 cm high, with 5-10 main axes in functionally male plants, 5-6 main axes in functionally female plants, partial inflorescences umbels, secondary and tertiary inflorescence axes with reddish-white, appressed, stellate hairs; enlarged marginal flowers always present (Fig. 5C ), terminally placed in a cyme, up to 3.6 cm diameter in female plants, up to 1 cm in male plants, sepals with marked veins, pistils 2 or sometimes not at all developed, reduced stigma not developed, receptacle rudimentary, globose, nearly the same size as that of reduced flowers, further characters not observed in detail, pedicel in male plants up to 1.5 cm; flower pedicel of reduced flowers 0.1-2.2 mm long in functionally male flowers, 1 mm long in functionally female flowers, enlarging during fruit development, 1.5 mm long in mature fruits, receptacle broadly campanulate in functionally male flowers, semiglobose in functionally female flowers, ovary inferior, calyx lobes 4, triangular, 0.3-0.5 mm long, petals 4, pinkish to wine red, imbricate, cucullate, membranous, 2 mm long, 1 mm broad; functionally male flowers (Fig. 5C ): hypanthium 1 mm diameter, 1.5 mm high, stamens 8, well-developed, filaments 1-1.5 mm long, anthers 1 mm long, 0.2 mm broad, pistils 2, reduced, 0.1 mm long, stigmas not penicellate; functionally female flowers: hypanthium 1.5 mm diameter, 1-1.5 mm high, stamens extremely reduced, as such their number could not be determined and their filaments and anthers have not been measured, pistils 2, 1 mm long, enlarging up to 1.5 mm during fruit maturation, stigmas apically clavate and shortly penicellate, 0.2 mm long; fruit a semiglobose capsule, apically with a conspicuous border, dark reddish brown, 2 mm high, 2.5 mm broad above, 3 mm long, opening between the two pistils to release seeds, seeds not seen.

Distribution.

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.

Habitat.

This species occurs in mountain cloud forest, sometimes on heavy slopes at elevations between 1000-2500 m.

Phenology.

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.

Notes.

Briquet (1919) based his description of H. goudotii on three collections, all by Goudot (without number): "Columbia: Ibaque ( Ibagué), Cahi (Cali) [G00223928, G00418963]; rives du Combayma (Combeima) [G00418962]; massif du Quindiu (Quindio), a Portoihuelo (?) [G00418961]" (correct current locality names and G barcode numbers are placed between brackets). It seems all three collections are from the Central Cordillera between Ibagué and Armenia in the department of Tolima (R. Callejas, Universidad de Antioquia, Colombia, pers. comm.). McClintock (1957) selected the latter of the three syntypes mentioned in the original description by Briquet (1919) as lectotype, although she referred to it as holotype which is not correct according to the ICN (Art. 9, Turland et al. 2018). However, as in several other species for which McClintock (1957) realized lectotypifications and synonimizations, we doubt that she actually has had access to the involved specimens. In the case of H. goudotii , she considered the most representative specimen to be the collection which has been fragmented to take material to the herbarium F. Indeed, the specimen in F [F0066624F] consists, apart from the fragments of leaves and inflorescences, of a photo of the original Goudot collection. Careful examination of the latter shows that it consists of the branch which was designated as lectotype by McClintock (1957) [G00418961], which has been turned around 180 degrees, and an additional branch which was fragmented and is the F specimen. The latter is considered as isolectotype in that herbarium. However, because of the fragmentation of the original collection, the material that remained in G does lack fully developed enlarged marginal flowers.

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).

Preliminary conservation status.

Although this species has an EOO of about 154,700 km2, it is Endangered according to the IUCN categories and criteria ( IUCN 2012), with an AAO of 100 km2, as well as an extensive reduction in both EOO and AOO due to habitat destruction and fragmentation.

Additional specimens examined.

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 (GENT, HOXA, IEB, MEXU, QCNE); El Chaco, ca. 2.1 airline km W of Sardinas, 0°23'2.7"S, 77°51'23.9"W, 1996 m, 10 Jul 2012, inflorescence bud, Granados Mendoza et al. 2012-105 (GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, ca. 2 airline km W of Sardinas, 0°23'1.3"S, 77°51'22.4"W, 1987 m, 10 Jul 2012, sterile, old inflorescences axes of previous year visible, Granados Mendoza et al. 2012-107 (GENT, HOXA, IEB, MEXU, QCNE); Tena, Cordillera de los Huacamayos, localidad Pigui Yacu (Verde Yacu) Río Cushillo Yacu ( Río Grande se origina en la cabecera de Sisahua), 1620 m, 30 Dec 1995, ♀, flowers, Jaramillo & Tapia 18583 (QCA); Quijos, Cosanga. Cerca de la población de Cosanga, a 500 m del Río Cosanga, 2000 m, 11 May 1990, ♂, flowers, Palacios 4895 (MO, QCNE); Pichincha: Nanegal, Sendero de la Unión de los Ríos, secondary lower montane rain forest along Río Umachaca, 00°07.5'N, 78°37.5'W, 1450 m, 24 Jun 1996, ♂, flowers, Kelch et al. 31803 (QCNE); Quito, Parroquia Nanegal, montane rain forest along trail from Río Umachaca to Loma Sta. Lucía, c. 6 km airline SE of Nanegal, near top ridge, 00°07.5'N, 78°37'W, 1635 m, 6 Sep 1993, inflorescence buds, Webster et al. 30392 (QCNE); Quito, Nanegal, Reserva Biológica Maquipucuna, 1200-1700 m, 20 May 1991, ♀, flowers, Tipaz & Quelal 207 (AAU, MO, NY, QCNE); Quito, Maquipucuna Reserve, trail to Hacienda Esparragos, 00°07'N, 78°38'W, c. 1400 m, 27 Aug 1989, ♂, flowers, Webster et al. 27062 (QCA); Tungurahua: Baños, ca. 2 km E of Río Verde on touristic path before tunnel on road to Baños-Mera, 1°23'25.1"S, 78°16'51.2"W, 1721 m, 16 Jun 2012, sterile, Granados Mendoza et al. 2012-45 (GENT, HOXA, IEB, MEXU, QCNE); Zamore-Chinchipe: in the vicinity of the mining camp, at the Río Tundaime, along road to military base El Condor, 3°37'31"S, 78°26'26"W, 1000 m, 5 Nov 2004, ♀, fruits, van der Werff et al. 19315 (LOJA, QCNE).

Figures

Figure 5. 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.

Figure 6. Distribution of Hydrangea goudotii and Hydrangea peruviana.