Research Article |
Corresponding author: Alejandro Zuluaga ( zuluaga.alejandro@correounivalle.edu.co ) Academic editor: Peter Boyce
© 2019 Allison Muñoz-Castillo, Leonardo Guevara-Ibarra, Laura Clavijo, Alejandro Zuluaga.
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:
Muñoz-Castillo A, Guevara-Ibarra L, Clavijo L, Zuluaga A (2019) Two new species of Chlorospatha section Orientales (Araceae) from western Andes in Colombia. PhytoKeys 135: 21-34. https://doi.org/10.3897/phytokeys.135.38050
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Two new species of Chlorospatha (section Orientales) from the western slope of the Cordillera Occidental in the departments of Valle del Cauca and Choco (Colombia) are described here. The new species represent the first records of section Orientales for Colombia, which was previously known only from the eastern Andes in Ecuador. The two new species are similar to C. longipoda, C. hannoniae and C. boosii. Chlorospatha minima sp. nov. is differentiated by its small overall size (less than 30 cm tall), blade strongly inequilateral with smooth adaxial surface, and spadix less than 2.2 cm long. Chlorospatha silverstonei sp. nov. is differentiated by its large overall size (30–60 cm tall), 1–3 leaves per plant, and quilted adaxial blade surface.
Araceae, Chlorospatha, new species, taxonomy, tropical Andes, section Orientales
The tribe Caladieae (Araceae) comprises 11 genera and 326 species restricted to Tropical America (
Until 1981, and for almost 100 years, the genus Chlorospatha was monotypic with C. kolbii Engl. as the only species described.
Most species of Chlorospatha have narrow distribution ranges; furthermore, the level of endemism in Chlorospatha is the highest among the genera of Araceae in Colombia, with 43 (63%) species endemic to the country (vs. 23 [33%] endemic to Ecuador). Despite the large number of species in Colombia, fewer collections were available compared with Ecuador, highlighting the need for more botanical exploration in the country and the potential for the discovery of several new species (
We assembled a database with all collections from
Our database comprised, in total, 572 collections of Chlorospatha, representing 70 species (including the two described here), with 214 collections from Colombia, seven of them belonging to the two new species. The number of collections per species was very low, with 22 species known only from the type collection, 13 only from two, 27 from less than 10, and merely eight species known from more than 10 collections.
COLOMBIA. Valle del Cauca: municipio Dagua, corregimiento El Queremal, old road Cali-Buenaventura, 6 km from El Queremal, 3°33'45.6"N, 76°45'27.1"W, 1050–1100 m, 20 May 2017, A. Zuluaga, L. Guevara, M. Llano & A. Muñoz 1645 (holotype: CUVC!; isotypes: COL!, MO!)
Chlorospatha minima can be distinguished from other species in section Orientales by its overall small size (less than 30 cm tall), smooth adaxial leaf surface, 1–2 inflorescences per axil, and spadix 20.4–22.8 mm long. Additionally, it differs from C. silverstonei sp. nov., the other species of this section in the western slopes of the Andes, by having three collective veins (vs. two in C. silverstonei sp. nov.), and the primary lateral (secondary) and minor veins glabrous on the abaxial surface (vs. scale-like indument).
Terrestrial herb, 10–25(–30) cm tall; stem subterranean, decumbent, with cataphylls quickly deciduous; internodes 6.6–7.4 × 5.4–7.8 mm, drying matte, dark brown; cataphylls brownish green, 5.2–6.2 cm long, acuminate at apex, drying matte, reddish brown. Leaves 2 to 5, erect-spreading; petioles 8.3–28.2 cm long, fleshy, glabrous, semiglossy, green with faint darker transverse markings, drying matte, dark brown, sheathed 5.2–9.4(–12.0) cm or (1/5–)1/3–1/2 of its total length, rarely more than 1/2; sheath decurrent onto the petiole apex; free portion of the petiole 0.6–3.7 mm diam. midway; blades broadly triangular-ovate, inequilateral, 5.8–14.2(–16.3) × 2.6–10.1 cm, 1.5 to 2.2 times longer than wide, weakly hastate at base, acuminate at apex, usually slightly broader across anterior lobe than at base, not constricted at petiole insertion, glabrous, conspicuously discolor, distance tip to tip across posterior lobes 2.2–9.2 cm wide; both surfaces smooth, glossy, drying semiglossy; abaxial surface with several layers of cells forming a reticulum, 0.3 to 0.4 mm diam.; anterior lobe 5.1–11.9 × 2.6–10.1 cm, 1 to 2 times longer than wide, 2.2 to 5.7 times longer than posterior lobe, wider near petiole insertion, rarely asymmetrical; posterior lobes directed toward base, 0.9–4.6 (–5.2) × 0.8–4.4 cm, 0.7 to 1.4 times longer than wide, narrowly rounded to obtuse at apex, slightly broader at petiole insertion, ± symmetrical, sinus parabolic to spathulate; midrib and major venation usually darker than the surface, round-raised and drying ± flattened abaxially; primary lateral veins 3, rarely 4, per side, arising at 30°–60°, rarely 70°, straight to weakly curving towards the margin; secondary veins abaxially sunken, drying visible and darker than the surface, the primary lateral and minor veins glabrous on the abaxial surface; 3 collective veins that originate from first, second and third basiscopic veins, respectively, ± parallel to margin; basal veins coalescent into a prominent posterior rib, 1–2(–3) acroscopic, 2–3(–4) basiscopic veins; minor veins slightly visible abaxially. Inflorescences erect, 1 to 2 per axil; cataphylls of inflorescence not visible outside the sheath; peduncle held within the sheath, 34.0–51.5(–76.0) × 0.4–1.2 mm, drying dark brown to black; spathe erect (all measurements for the spathe and spadix made from spirit material), 27.0–29.8 mm long, apiculate at apex, 6.6–7.0 mm (1.1 to 1.3 times) longer than spadix; spathe tube green or pale green on outer surface, rarely maroon-tinged, 10.0–12.9 × 4.2–4.3 mm, drying dark brown to black on outer surface; spathe blade maroon-tinged, with green veins on outer surface, green on inner surface, drying dark brown or black, ca. 16.9 mm long, erect after anthesis, then marcescent; spadix erect, 20.4–22.8 mm long, sessile, adnate basally to the spathe for 2.3–3.1 mm (1/5 to 1/3 of the length of pistillate portion); pistillate portion light green, 8.3–8.6 × 2.0 mm; pistils ca. 1.3 mm diam.; stigma light green, sessile, ca. 0.4 mm diam.; fertile staminate portion white, 11.2–12.9 × 2.8–3.0 mm, cylindrical, rounded at apex, drying whitish brown; synandria ca. 1.3 mm diam., coherent; sterile portion white, 2.2–2.5 × 2.7 mm, wider at apex, drying whitish brown; sterile flowers with straight borders, 1.2 mm diam. (viewed from above). Infructescence (measurements made from spirit material) erect or pendent, brown, ca. 25.5 × 8.0 mm, drying mate, dark brown on outer surface; berries green, 2.3–3.9 mm diam.; seeds white, (6–)20 to 24 per berry, 1.2–1.5 × 0.7–0.9 mm, ovoid to ellipsoid, longitudinally striate, minutely white-strophiolate.
The epithet minima refers to the small size of this species, less than 30 cm tall, the smallest in the genus.
Chlorospatha minima is endemic to the western slopes of the Colombian Andes in the department of Valle del Cauca. It has been found only in one locality on the old road Cali-Buenaventura at 1000 m, inhabiting humid forest and growing close to a waterfall (Fig.
Despite more than four years of extensive field work, Chlorospatha minima is known only from one population located outside the Farallones de Cali National Natural Park, in an area frequently visited by tourists. Because the estimated extent of occurrence is less than 100 km2, the only population known has less than 100 individuals, and the quality of habitat is declining, C. minima could be assessed as Critically Endangered, according to the IUCN criteria (
Chlorospatha minima belongs to section Orientales, characterized by having a stylar region lacking a mantle (Fig.
Inflorescence and leaf characters of Chlorospatha minima and C. silverstonei. A–I C. minima J–O C. silverstonei A inflorescence B female flowers C male flowers D sterile flowers E infructescence F adaxial blade surface, note the reticulum of cells in the peeling of abaxial surface G abaxial blade surface H primary lateral and minor veins glabrous on abaxial surface I seed J inflorescence K female flowers L male flowers M adaxial blade surface, note the layers of cells N abaxial blade surface O primary lateral and minor veins with scale-like indumentum on abaxial surface. (Photographs by Juan Felipe Ortega and the imaging laboratory at the Biology graduate program at Universidad del Valle).
Main morphological characters of Chlorospatha silverstonei and C. minima. A, B, E C. silverstonei C, D, F, G C. minima A adult plant B abaxial blade surface C abaxial blade surface D adult plant E cataphyll and inflorescence in post anthesis, note spathe acuminate and longer than the spadix F Inflorescence in anthesis (right) and post-anthesis (left) G shoot, showing young infructescence with deciduous spathe blade and young inflorescence.
Chlorospatha minima differs from C. longipoda in having leaves broadly triangular-ovate vs. narrowly ovate or ovate-elliptic, base of blade slightly hastate vs. subcordate to subsagittate, and (6)20–24 seeds per berry (vs. 7–8). It differs from C. hannoniae in having leaves weakly hastate at base vs. sagittate or subsagittate, apex of spathe apiculate vs. cuspidate, and erect spadix vs. slightly curving forward. Finally, C. minima differs from C. boosii in having 2–5 leaves that are held erect (vs. 8 to 12 leaves) (Table
Morphological comparison of Chlorospatha minima, C. silverstonei, C. longipoda, C. hannoniae and C. boosii.
C. minima | C. silverstonei | C. longipoda | C. hannoniae | C. boosii | |
---|---|---|---|---|---|
Plant size | 10–25 (–30 cm) | 30–60 cm | 40 cm | 50 cm | 30–50 cm |
Bulbils | Absent | absent | absent | Present | present |
Cataphylls | quickly deciduous | persisting ± intact | Remnants of old cataphylls persisting ± intact to semi-intact. | ultimately deciduous | quickly deciduous |
Number of leaves | 2–5 | 1–3 | 3–5 | 8–14 | 8–12 |
Leaf shape | broadly triangular-ovate | broadly ovate to rounded | narrowly ovate or ovate-elliptic | ovate-cordate, occasionally broadly subtriangular | ovate (occasionally subsagittate in juvenile plants) |
Base of blade | inaequilateral, slightly hastate | cordate, rarely slightly hastate | subcordate to subsagittate | sagittate or subsagittate | cordate-subcordate |
Apex of blade | acuminate | acuminate to cuspidate, almost always mucronate at apex | weakly to moderately acuminate to bluntly acute or apiculate at apex | weakly acuminate to apiculate at apex | weakly acuminate or apiculate at apex |
Blade size | 5.8–14.2(–16.3) × 2.6–10.1 cm | 9.3–27.7 × 4.9–17.7(–21.2) cm | (10.0–)15.5–21.5 × (3.5–)5.0–13.0 cm | 16.0–20.5 × 10.0–15.5 cm | (16.0–)19.0–27.5 × (7.0–)13.0–19.5 cm |
Adaxial leaf surface | Smooth | quilted | quilted | broadly quilted and sub-bullate | broadly quilted |
Abaxial leaf surface | broadly reticulate, with dead cells layers | reticulate, with dead cells layers | reticulate, narrowly colliculate along all venation | reticulate, narrowly minutely colliculate along all venation | reticulate, narrowly minutely colliculate along all venation |
Diameter of reticulum on abaxial surface of the leaf | 0.3–0.4 mm diam. | 0.2–0.3 mm diam. | present, not seen | present, not seen | present, not seen |
Secondary and minor venation on lower surface | sunken | prominulous | convex or moderately to narrowly round-raised | prominulous | slightly raised |
Indument of veins in lower surface | Absent | present | unknown | unknown | unknown |
Number of acroscopic veins | 1–2(–3) | 1–3(–4) | 2–3 | unknown | 2–3 |
Number of basiscopic veins | 2–3(–4) | 3–4(–5) | 2–3 | unknown | 3 to 4 |
Number of primary lateral veins | 3(–4) pairs | 2–4(–6) pairs | 4–6 pairs | (3–)4 pairs | 3–4 pairs |
Angle of primary lateral veins | 30°–60°(70°) | 30°–70°(90°) | 17°–45° | 45°–65° | 25°–55° |
Number of collective veins | 3 | 2 | 2–3(–4) | 2(–3) | 3 |
Inflorescences per axil | 1 to 2 | 1 to 4 | 1 to 6 | 3 to 5 | 4 to 7 |
Cataphylls of inflorescence | not visible outside petiole sheath | visible outside petiole sheath | visible outside petiole sheath | visible outside petiole sheath | visible outside petiole sheath |
Portion of the spathe exceeding the spadix | 6.6–7 mm | 4.3–5.0(–25.7) mm | 10.0–35.0 mm | 7.0–10.0 mm | 20.0–40.0(60.0) mm |
Spadix position | Erect | erect | erect, occasionally curving forward at anthesis | slightly curving forward | erect |
Spadix length | 20.4–22.8 mm | 25.0–37.3 mm | (33.0–)43.0–53.0 mm | 38.0–50.0 mm | 56.0–69.0 mm |
Length of spadix adnate to spathe | 2.3–3.1 mm | 3.3–5.1 mm | 6.0–8.0 mm | 2.0–3.0 mm | 3.0–7.0 mm |
Proportion of adnate portion | 1/3 | 1/3 | 1/2 | 1/4 or less | 1/4 to ca. 1/2 |
Pistillate portion length | 8.3–8.6 × 2.0 mm | 8.7–16.1 × 2.5–2.9 mm | (7.0–)10.0–18.0 × 2.0–3.5 mm | 7.0–12.0 × 2.0–4.0 mm | 10.0–15.0 × 3.0–3.5 mm |
Staminate portion length | 11.2–12.9 × 2.8–3.0 mm | 13.6–19.6 × 4.1–4.4 mm | 20.0–30.0 × 2.0–3.0 mm | 25.0–35.0 × 3.0–3.5 mm | 34.0–45.0 × 3.5–4.0 mm |
Sterile portion length | 2.2–2.5 × 2.7 mm | 3.1–3.9 × 3.7–4.3 mm | 4.0–8.0 × ca. 2.0 mm | 7.0–12.0 × 2.0–3.0 mm | 5.0–9.0 × 2.5–3.0 mm |
Border of sterile flowers | Straight | irregular | irregular | irregular | irregular |
Color of spathe in fruit | Brown | black | entirely green or occasionally purple-tinged | unknown | unknown |
Seeds per berry | (6–)20 to 24 | 6 to 20 | 7 to 8 | unknown | unknown |
COLOMBIA. Valle del Cauca: municipio Dagua, corregimiento El Queremal, 3°33'45.6"N, 76°45'27.1"W, 1159 m, 17 Mar 2018, Zuluaga et al. 2328 (CUVC!).
COLOMBIA. Valle del Cauca: municipio El Cairo, Reserva Natural de la Sociedad Civil “Cerro El Inglés”, camino al límite departamental entre Valle y Chocó. 4°44'13.3"N, 76°18'7.7"W, 2120–2230 m, 8 Oct 2017, A. Zuluaga & M.E. Cardona 1946 (holotype: CUVC!; isotypes: COL!, MO!)
Chlorospatha silverstonei can be distinguished from the other species in section Orientales by having 1–3 leaves per plant, an overall larger size (30–60 cm tall) and a small spadix (25–37.3 mm long). Additionally, it differs from C. minima sp. nov., the other species in this section from the western slopes of the Colombian Andes, in having two collective veins (vs. three in C. minima sp. nov.), the primary lateral and minor veins with scale like indument on the abaxial surface (vs. glabrous).
Terrestrial herb, 30–60 cm tall; stem decumbent, with remnants of cataphylls persisting ± intact; internodes 5.2–20.2 × 6.8–12.5 mm, drying matte, dark brown; cataphylls brownish green, (3.1–)4.4–8.5 cm long, acuminate at apex, drying faintly glossy, reddish brown. Leaves 1 to 3, erect-spreading; petioles 15.2–46.8(–52.4) cm long, free portion of the petiole 1.72–7.18 mm diam. midway, fleshy, glabrous, semiglossy, irregularly dark purple-mottled with longitudinal dark purple lines, drying dark brown to black, sheathed 2.7–11.5(–15.0) cm, less than 2/5 of its total length; sheath decurrent on to the petiole apex; blades broadly ovate to rounded, glabrous, conspicuously bicolor, 9.3–27.7 × 4.9–17.7(–21.2) cm, 1.2 to 2.3 times longer than wide, cordate at base, rarely slightly hastate, acuminate to cuspidate at apex, almost always mucronate at apex, usually slightly broader across anterior lobe than at base, distance tip to tip across posterior lobes 3.1–13.9(–20.5) cm, not constricted at petiole insertion; adaxial surface quilted, glossy, drying brownish green; abaxial surface reticulate, glossy, drying green to yellow-green, with several layers of cells forming a reticulum, 0.2 to 0.3 mm diam.; anterior lobe 7.2–17.4(–18.5) × 4.9–17.7(–21.2) cm, 0.8 to 1.7(–2.0) times longer than wide, 1.5 to 3.6(–4.0) times longer than posterior lobes, broader near petiole insertion, ± symmetrical; posterior lobes directed toward base, 2.1–9.6 × 1.7–8.3 cm, 0.68 to 1.45 times longer than wide, rounded to obtuse at apex, weakly broader at base, slightly inequilateral, the inner side narrower, sinus spathulate to ovate; midrib and major venation narrowly sunken adaxially, round-raised, drying ± flattened and usually darker than surface abaxially; primary lateral veins 2 to 4 per side, rarely 5–6, arising at 30°–70°, rarely 90°, straight to weakly curved towards the margin; secondary and minor veins darker than the surface, prominulous, forming a conspicuous reticulum abaxially, more visible when dried, the primary lateral and minor veins with scale-like indumentum abaxially, only visible in dried material under the microscope; collective veins 2, the outermost arising from the first and second basiscopic veins, ± parallel to margin, the innermost arising from the third basiscopic vein; basal veins coalescent into a prominent posterior rib, 1–3(–4) acroscopic veins, 3–4(–5) basiscopic veins. Inflorescences erect (all measurements made from spirit material), 1 to 4 per axil; cataphylls of the inflorescence visible outside the petiole sheath, irregularly dark purple-mottled, drying dark brown to black; peduncle 42.8–53.1 (–112.7) × 0.8–1.3 mm, held within the sheath, drying dark brown to black; spathe erect, 32.2–62.9 mm long, acuminate at apex, 4.3–5.0(–25.7) mm longer than spadix (1.2 to 1.7 times longer than spadix); spathe tube pale green on outer surface, rarely maroon-tinged, 8.7–23.3 × 5.0–6.3 mm, drying dark brown to black on outer surface; spathe blade maroon-tinged, with green veins on outer surface, green on inner surface, 18.7–39.7 mm long, drying dark brown or black, erect after anthesis, marcescent in fruit; spadix erect, 25.0–37.3 mm long, sessile, adnate basally to the spathe for 3.3–5.1 mm (ca. 1/3 of the length of the pistillate portion); pistillate portion light green, 8.7–16.1 × 2.5–2.9 mm; pistils coherent, ca. 1.0 mm diam.; stigma light green, sessile, ca. 0.3 mm diam.; fertile staminate portion white, 13.6–19.6 × 4.1–4.4 mm, slightly conical, rounded at apex, drying whitish brown; synandria ca. 1.1 mm diam., coherent; sterile portion white, 3.1–3.9 × 3.7–4.3 mm, wider at apex, drying whitish brown; sterile flowers with irregular borders, ca. 1.4 mm (viewed from above). Infructescence erect or pendent, brown, 18.5–32.6 × 4.8–10.3 mm, drying dark brown on outer surface; berries drying pale tan; seeds white, 6 to 20 per berry, 1.5–2.0 × 0.9–1.2 mm, ovoid to ellipsoid, longitudinally striate, minutely strophiolate, drying brown.
Illustration of Chlorospatha minima A adult plant with inflorescence B infructescence C inflorescence at anthesis D inflorescence on post-anthesis E abaxial surface of leaf blade; note reticulate venation and collective veins F upper view of male flowers G upper view of sterile flowers H upper view of female flowers I seed. (Drawn by Eileen Muñoz from the holotype A. Zuluaga et al. 1645).
Illustration of Chlorospatha silverstonei A adult plant with inflorescence B abaxial surface of leaf blade; note reticulate venation and collective veins C Inflorescence in female anthesis D Inflorescence in post-anthesis E upper view of male flowers F upper view of sterile flowers G upper view of female flowers H seed. (Drawn by Eileen Muñoz from the holotype A. Zuluaga et al. 1946).
Chlorospatha silverstonei is named in honor of Dr. Phillip Silverstone-Sopkin (1939–2018), an American botanist who lived and worked in Colombia for 39 years. He was a faculty member at Universidad del Valle until 2014 and an ad Honorem professor since 2015. Additionally, he was the director of the herbarium Luis Sigifredo Espinal Tascon at the same University for 17 years. Dr. Silverstone-Sopkin collected more than 13000 botanical specimens, especially from the department of Valle del Cauca, and carried out several explorations in the region where this species was found.
Chlorospatha silverstonei is endemic to the western slopes of the Colombian Andes, along the border between the departments of Valle del Cauca and Chocó. It grows in cloud forests between 1900 and 2300 m. It has been collected in two natural reserves, “Cerro El Inglés” and “Alto Galapagos” (Fig.
Chlorospatha silverstonei has been found in two localities along the Serranía de los Paraguas mountain range, with an estimated extent of occurrence larger than 38000 km2. In these two localities there are several populations of this species with abundant individuals; therefore, C. silverstonei is preliminary categorized as Least Concern (LC), according to the IUCN criteria (
Chlorospatha silverstonei is similar to C. minima, C. longipoda, C. hannoniae, and C. boosii, but it differs from these four species in having fewer leaves (1–3 vs. 2–14) (Table
COLOMBIA. Chocó: municipio Sipí, Reserva Natural Cerro El Inglés, debajo del sitio Santicos, 4°45'22.0"N, 76°18'12.9"W, 2000 m, 17 Oct 2016, A. Zuluaga et al. 1321 (CUVC!). Valle del Cauca: municipio El Cairo, Reserva Natural Cerro El Inglés, camino desde la divisoria de aguas hasta la cabaña de investigadores, 4°44'23.9"N, 76°18'15.0"W, 2100–2200 m, 22 Jan 2016, A. Zuluaga et al. 946 (CUVC!); camino a Los Santicos, 4°45'15.5"N, 76°18'02.3"W, 2250 m, 17 Oct 2016, A. Zuluaga et al. 1305 (CUVC!); reserva natural Alto Galapagos, near to the border Chocó-Valle del Cauca, 2018, A. Zuluaga et al. (CUVC!).
We thank the non-governmental organization Serraniagua for allowing us to conduct fieldwork in the Reserves Cerro El Inglés and Alto Galapagos in Serranía de los Paraguas mountain range. We also thank Juan Felipe Ortega and the imaging laboratory at the Biology graduate program at Universidad del Valle for providing some of the images. Finally, we thank Monica Carlsen and Peter Boyce for their meticulous reviews and for their suggestions to improve this manuscript.