Introduction

The Neotropical orchid genus Sarcoglottis C.Presl comprises about 50 geophytic species with fasciculate, fleshy roots (rarely these produced spaced apart along a rhizome), flat broad leaves forming a basal rosette, and fleshy, comparatively large flowers for subtribe Spiranthinae (Salazar 2003, Salazar et al. 2018). The flowers are resupinate, often fragrant, with an ascending, shortly pedicellate ovary and horizontal or ascending perianth, and the basal portions of the lateral sepals are connate and decurrent on the ventral surface of the ovary to form a nectary, which can be completely fused with the ovary or form a conspicuous sac (see also Salazar et al. 2018). The labellum consists of a proximal, usually spathulate, channeled portion (“hypochile”), the base of which bears two subulate, retrorse nectar glands inserted in the nectary chamber, and an expanded, variously recurved or deflexed distal lobe (“epichile”). The column is straight, obliquely adnate to the ovary, with a triangular to oblong-triangular rostellum, which is truncate or shallowly emarginate after removal of the wishbone-shaped pollinarium, and the anther is semi-ellipsoid, with a distinct apical beak. The scarce information available on natural pollination of species of Sarcoglottis indicates that it is carried out by long-tonged bees of tribe Euglossini (Apidae) that probably feed on the nectar present at the bottom of the floral tube (Pérez-Escobar, Roubik & Gerlach 2017, Salazar 2003, Singer & Sazima 1999).

Sarcoglottis has never been revised, little information has been published on almost any aspect of its biology, and the taxonomy of the species is far from thoroughly studied, especially that of the South American taxa. As for Peru, Schweinfurth (1958) recorded a single species, S. acaulis (Sm.) Schltr. (as Spiranthes acaulis (Sm.) Cogn.), listing Peruvian S. speciosa C.Presl, the type species of the genus, as its synonym. Then Brako & Zarucchi (1993) recorded three species, including S. acaulis, S. homalogastra (Rchb.f. & Warm.) Schltr., and S. villosa (Poepp. & Endl.) Schltr., but they did not cite specimens of the last two species and stated that they were unconfirmed for the country. Bennett & Christenson (1995) reported S. neglecta Christenson, but the specimen from Moyobamba they illustrated represents a different species, which was described later as S. portillae Christenson based on a cultivated plant without precise locality but said to have been found in Ecuador “near the Peruvian border” (Christenson 2003a: 243). Subsequently, Bennett & Christenson (1998) “resurrected” S. speciosa, indicating as its distribution Peru and probably the neighboring countries. A further Peruvian species, S. micrantha Christenson, was distinguished by its very small flowers with a 5-lobed apical labellum (Christenson 2001). A recent compilation of the Peruvian Orchidaceae listed five species, namely S. acaulis, S. grandiflora, S. homalogastra, S. micrantha, and S. villosa (Goicochea et al. 2019). A recent addition was S. neillii Salazar & Tobar, discovered on the Ecuadorian side of the Cordillera del Cóndor but also known from Peruvian herbarium specimens (Salazar et al. 2019).

During fieldwork conducted in the department of Cusco, Peru, in 2017, two populations of a Sarcoglottis that did not correspond to any of the species previously known either for Peru or anywhere else were discovered. In the following, we describe and illustrate the unknown entity as a new species, highlight the features that distinguish it from S. fasciculata (Vell.) Schltr., to which it shows similarities in the appearance of the flowers, provide a table contrasting the features that distinguish the new species from S. fasciculata and its Bolivian and Peruvian congeners, and present a preliminary assessment of its conservation status.

Materials and methods

The morphological description and the measurements were made from live plants and flowers preserved in 70% ethanol. Fresh flowers were dissected and photographed with a digital camera (Sony SLTα 58 with type mount and sensor APS-C, Sony Group Corporation, Tokyo, Japan) provided with a SLT-A58Y 18-55 mm lens). Image processing and plate preparation was done with Photoshop v. 24.1.0. Two specimens collected at Machupicchu were deposited in CUZ and USM; and a specimen (flowers in spirit) from Sawayacu (district of Santa Teresa) was deposited in KUELAP. The new species was compared with the protologues and original material of all previously known South American species of Sarcoglottis, and with specimens or digital images of specimens of this genus housed in Peruvian and other international herbaria examined such as AMES, AMO, ANDES, BHCB, BHZB, BM, COL, CUZ, ECUAMZ, F, GH, HB, HOXA, K, KUELAP, M, MEXU, MO, NY, P, QCA, QCNE, R, RB, SEL, SP, US, USM, VEN, and W.

Taxonomic treatment

Sarcoglottis wernerherzogii Collantes, Edquén et Salazar, sp. nov. (Fig. 1-4).

TYPE: PERU. Department of Cusco: Province of Urubamba, district of Machupicchu, Santuario Histórico de Machupicchu, llaqta Machupicchu, 2446 m., 21 Oct 2017, B. Collantes & F. Incahuamán 344 (holotype: USM!; isotype: CUZ!).

Diagnosis: The new species is similar to eastern South American Sarcoglottis fasciculata (Vell.) Schltr., from which it is distinguished by inhabiting in Andean montane cloud forest at 2050-2500 m a.s.l. (vs. inhabiting in Atlantic rain forest near sea level); flowering after the shedding of the leaves (vs. flowering with the leaves present); homogeneously dull greenishto yellowish perianth segments (vs. whitish or pale green background with contrasting greenish or reddish veins); rigidly conduplicate-concave lateral sepals that, when spread out, are strongly falcate (vs. slightly concave, not rigid, obliquely lanceolate lateral sepals); strongly cymbiform hypochile that when spread out is obovate, as long as wide (vs. broadly channeled hypochile that when spread out is oblanceolate, about twice longer than wide or longer); ovate, strongly deflexed epichile provided with a central longitudinal thickening throughout its length (vs. triangular, erect or gently arcuate epichile with a thickening only at the apex); and apical anther beak truncate (vs. apical anther beak rounded).

Geophytic, acaulescent herb 50−60 cm in height including the inflorescence. Roots fasciculate, fleshy, cylindrical-fusiform, dull brownish white, to 12 cm long, 4−6 mm in diameter. Leaves 4−6, deciduous, withered and lost at flowering time but a new rosette may be developing laterally as a new shoot, upper surface homogenous bright green with 5 pale yellowish veins, the central one broader, lower surface slightly paler green, oblong-elliptic, cuneate, acute, 6−17 × 3−4 cm. Inflorescence scapose, arising from the apex of the shoot, the flowers opening after the shedding of the leaves, 50−55 cm long; scape 4−6 mm in diameter, terete, glabrous below, becoming lanuginose above (trichomes simple, septate, glandular, whitish), partially concealed by bracts, these narrowly lanceolate, semi-tubular, acute to acuminate, 5−7 cm long; raceme relatively dense, 10−13 cm long, with 12−15 simultaneous flowers. Floral bracts subequal to conspicuously longer than the ovary, 27−35 × 6−7 mm, green to pale brown with reddish suffusion towards the apex, concave, slightly incurved, with involute margins, when spread-out lanceolate, acute to long-acuminate, 5−7-veined, sparsely papillose to shortly pubescent on the outer surface near and along the margins, glabrous otherwise. Flowers with dull green or yellow sepals and petals, sometimes suffused with brownish green or rusty red, hypochile of labellum whitish with olive green base, epichile olive green with white longitudinal thickening. Ovary 26−27 × 5−7 mm, green to pale brownish or golden yellow, ascending, subclavate, twisted at base, densely pubescent, the trichomes simple, septate, tapering, ending in a small, round secretory cell. Dorsal sepal abruptly incurved from near the base, forming with the petals a sort of hood tightly covering the column and labellum, concave below the middle, sparsely pubescent near the base, the rest of the outer surface colliculate, lanceolate-oblong, obtuse, 3-veined, the lateral veins branching near the base, 24.0−28.0 × 6.0−6.5 mm. Lateral sepals adnate to the distal 2/3 of the ventral surface of the ovary and connate at their bases, forming a rounded, prominent nectary, free portions strongly downcurved, concave, rigidly conduplicate, 3-veined, the lateral veins branching near the base, 27.0−28.5 × 7.0−8.0 mm, densely pubescent outside, the pubescence gradually reduced to papillae towards the apex, falcate-lanceolate, long-acute. Petals marginally adherent along their upper margin to the dorsal sepal on their distal 2/3, free below, obliquely oblanceolate-falcate, free part of inner margin waved, apex obliquely acute, surface at and near the outer margin below the middle densely pubescent, with short septate, glandular trichomes, 14.5−22.5 × 4.0−5.0 mm, 3-veined at base, the lateral veins branched above into 5 veins. Labellum adnate at its base with the connate portions of the lateral sepals, 25−26 mm total length excluding the nectar glands; hypochile strongly cymbiform, ca. 4 mm wide and deep in natural position, when spread out broadly obovate and expanded at each side into a rounded, membranaceous margin that in natural position is adhered to the respective side of the column, 12.0−13.0 × 12.0−13.5 mm, densely papillose outside, with a concavity at the base of each nectar gland provided with finger-like trichomes directed opposite to the nectar gland, inside somewhat contracted by effect of the folding inwards of the margins; the bottom of the hypochile protrudes slightly between the bases of the free portions of the lateral sepals; epichile with a fleshy, canaliculate central longitudinal thickening that reaches the labellum apex, abruptly deflexed more than 90°, ovate, acute and shortly apiculate, with membranaceous margins, 10.0−11.0 × 8.0−8.5 mm; base of labellum projected into two olive green, retrorse, subulate, smooth nectar glands 5−6 × ca. 1 mm. Column 6.0−9.0 × 1.8−2.5 mm, dorsally free from the ovary for 1-2 mm, ventral surface with a deep, narrow channel limited at each side by a fleshy, rounded keel, ventral surface of the column foot with a single thickened keel without a channel. Anther narrowly ellipsoid, centrally fleshy-thickened, with membranaceous, recurved margins and an oblong, truncate apical beak, 5.0−7.0 × 1.5−2.0 mm, plus a short free filament about 1.0−1.5 mm long; clinandrium shallow, with membranaceous, rounded margins ending in a short, obtuse projection towards the apex. Pollinarium ca. 5.5 mm long, consisting of two creamy white, clavate, deeply cleft pollinia united at apex to a subtrapezoid, gray viscidium reminiscent of a cow’s hoof. Rostellum ca. 3 mm long, narrowly triangular, broadly and shallowly notched after removal of the viscidium; stigma cordiform, emarginate, with two ovate. convergent receptive areas. Fruits not seen.

Eponymy: The species is named in honor of the great poet and filmmaker Werner Herzog (Munich, 1942), whose iconic films Aguirre, the Wrath of God (1972) and Fitzcarraldo (1982) reveal the natural magnificence of Machupicchu and the Peruvian Amazon.

Distribution and ecology: Known only from the Eastern Andean Cordillera in the Department of Cusco, provinces of Urubamba and La Convención, southern Peru. Terrestrial, in deep leaf litter in open areas within the montane cloud forest biome and among coffee and avocado orchards, at 2050−2500 m elevation. Flowering has been observed in the field in September and October. The flowers produce a conspicuous odor during daylight hours, described by one collector as reminiscent of the fruits of Pouteria lucuma (Ruiz & Pav.) Kuntze, and by the other as of bleach or semen. No objective characterization of the floral fragrance has been carried out.

Discussion

Sarcoglottis wernerherzogii is similar in outward floral appearance to eastern South American S. fasciculata (Vell.) Schltr. (the type from Rio de Janeiro, J. Velloso s.n., drawing!) by its horizontal perianth, dorsal sepal lanceolate-oblong, incurved, lateral sepals downcurved, but differs from the latter in eco-geographic features (high-Andean cloud forest at 2050-2500 m a.s.l. vs. Atlantic rain forest near sea level), phenology (flowering after the shedding of the leaves vs. flowering with functional leaves), flower coloration (homogeneously dull green or yellow flowers vs. pale green or rosy-green flowers with contrasting reddish veins, especially on the labellum epichile, and several morphological floral characteristics. These include the strongly cymbiform hypochile that, when spread out, is obovate and as wide as long (Fig. 1J, 2G−I, 3D−E) vs. broadly channeled hypochile that, when spread out, is oblanceolate and about twice as long as wide), the strongly deflexed, centrally thickened epichile (vs. gently arching, thin epichile), and the truncate anther beak (vs. rounded). It is worth nothing that the hypochile characteristics of S. wernerherzogii are unique in the genus.

Bolivian S. herzogii Schltr. (the type from Tigüipa, department of Chuquisaca, 700 m a.s.l., T. Herzog 1154, not located) also flowers after the leaves are shed, but it is distinguished from S. wernerherzogii in its dry, lowland Amazonian habitat at 400−700 m a.s.l. (Vásquez, Ibisch & Gerkmann 2003; https://tropicos.org/name/23511337, accessed 15 Nov 2023), the 1-veined, glabrous petals (vs. 5-veined, prominently ciliate petals; Fig. 4 D-E), and the much smaller, narrowly oblanceolate labellum 18 × 4.5 mm (fideSchlechter 1925: 333).

The main differences of S. wernerherzogii to S. fasciculata and to other Peruvian species of the genus are highlighted in Table 1. We omitted from that table S. acaulis, S. homalogastra, and S. villosa since we have been unable to find any specimens or other cogent evidence of their presence in Peru. As noted by Bennett & Christenson (1998), S. acaulis is known with certainty only from northern South America, including the island of Trinidad, Venezuela, the Guyanas, and probably neighboring Brazil. On the other hand, the first report of S. homalogastra and S. villosa for Peru was based on an unconfirmed personal communication by Dodson (Brako & Zarucchi 1993: 850), and their inclusion in Goicochea et al.’s (2019) compilation merely followed them. Sarcoglottis homalogastra (the type from Brazil, Minas Gerais, Lagoa Santa, E. Warming s.n., W-R, microfiche of flower and floral analysis!) is widespread in open fields in northern Argentina, eastern Brazil, and Paraguay (Schinini 2010). On the other hand, S. villosa, described from a plant collected near the mouth of the Amazon River, on the Atlantic coast of Brazil (Pará, Colares, Poeppig s.n., W, digital image!), is known with certainty only from northeastern Brazil (Ceará and Pará; Hoehne 1945).

Additional specimen examined: PERU. Department of Cusco: Province of La Convención: Distrito Santa Teresa, parte alta del sector Sawayacu, 2064 m, 30 Sept 2017, J. D. Edquén 4165 (photographs and flowers in spirit, KUELAP!); province of Urubamba, district of Machupicchu, Santuario Histórico de Machupicchu, al lado derecho del Camino Inca, cerca y rumbo a Intipunku, 21 Oct 2004, B. Collantes s.n. (photograph!); same locality, without date, J. G. Ochoa s.n. (photograph!)

Conservation status: The population of Machupicchu consists of about ten plants found over about 30 m along the sides of the Camino Inca (Inca Trail) within the historical sanctuary, hence under nominal legal protection. However, there are two potential threats to that population: first, the increasingly frequent forest fires escaped from crop fields of the surrounding areas, where farmers set fire to the vegetation during the dry season in the belief that it will improve soil fertility. Second, the even more frequent trimming of the vegetation along the Inca Trail by personnel of the sanctuary to ease the transit of tourists. The last practice could, and should, be supervised to avoid inadvertent damage to the rosettes or inflorescences of the plants of this species. There is also a possibility of damage by trampling by the tourists, which could be reduced by adequate signaling and the placement of discrete barriers. On the other hand, the population of Sawayacu is located in the adjacent “Área de Conservación Regional (ACR) Choquequirao,” but among coffee and avocado orchards.

It is worth nothing that S. wernerherzogii was not recorded in the several recent inventories and guides to the orchids (and other plants) of Machupicchu (Christenson 2003b; Collantes, Soto & Koechlin 2007, Ochoa Estrada 2021) and the ACR Choquequirao (Vilca et al. 2023), which suggests either that the species is very rare, or that it is easily overlooked, perhaps because of flowering when the leaves are absent and the dull coloration of its flowers. In any event, further field work is required to objectively determine the conservation status of this species. For the time being, we suggest its inclusion in the category of “data deficient” (DD, IUCN 2019).

Acknowledgements

We thank Peru’s Servicio Nacional Forestal y de Fauna Silvestre (SERFOR) of the Ministerio de Agricultura y Riego (MINAGRI) for the permits for scientific collecting (No. 099-2019-MINAGRI-SERFORD-GGSPFFS; No. 009-2017-SERNANP-JEF); Federico Rizo-Patrón and Ricardo Fernández for hospitality and assistance during our study of Peruvian herbaria; the Curators and staff of AMES, AMO, ANDES, BHCB, BHZB, BM, COL, CUZ, ECUAMZ, F, GH, HB, HOXA, K, KUELAP, M, MEXU, MO, NY, P, QCA, QCNE, R, RB, SEL, SP, US, USM, VEN, and W for courtesies extended while studying the collections in their charge; Abel Monteagudo for providing high-quality digital images of the specimens of Sarcoglottis housed at CUZ; Julio Ochoa for providing a copy of his photographic guide to the flora of Machupicchu and an unpublished photograph of the new species; Lucely L. Vilca for sharing a copy of his digital guide to the orchids of Choquequirao; and two anonumous reviewers for constructive criticisms to the manuscript. BC thanks José Koechlin, Chair of Inkaterra, for continued support to his orchid studies, and José Bastante, Chief of the Parque Arqueológico de Machupicchu and Rosa Huillcanina, of the Oficina de Atención al Visitante y Servicio Turístico, for access to the protected area and authorization to visit and study the population of the new species.

Literature cited

Publication Dates

History