Research Article |
Corresponding author: Alejandro E. Villarroel ( alejandro.villarroel@userena.cl ) Academic editor: Peter de Lange
© 2022 Alejandro E. Villarroel, Kora Menegoz, Carlos Le Quesne, Ricardo Moreno-Gonzalez.
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:
Villarroel AE, Menegoz K, Le Quesne C, Moreno-Gonzalez R (2022) Valeriana praecipitis (Caprifoliaceae), a species new to science and endemic to Central Chile. PhytoKeys 189: 81-98. https://doi.org/10.3897/phytokeys.189.73959
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The species Valeriana praecipitis (Caprifoliaceae), new to science and endemic to the Ñuble Region, Central Chile, is formally described. Morphological data support its placement in a new species, clearly different from V. philippiana. A detailed description, insights about its habitat and ecology, distribution map and illustration are provided. A table of comparison is also given with the morphological characters discriminating V. praecipitis from V. philippiana. The species is assessed as Endangered (EN) under the IUCN categories.
Andes, biodiversity, cliffs flora, Ñuble Region, taxonomy
The Valerianaceae family was included in the Caprifoliaceae family by the Angiosperm Phylogeny Group III (
Valeriana is one of the most diverse genera within the Caprifoliaceae, with a worldwide distribution, and centres of diversity in tropical areas of Central-America and to the south along the Andes mountains (
The Central Chilean Andes mountains are recognised as a centre for endemism in South-America (e.g.,
In this manuscript we describe a species of Valeriana located in the Central Chilean Andes at around 36° S latitude and clearly distinguishable from other species by its remarkable silvery-green basal leaves. In the following sections, a detailed account on how the species was discovered is given as a formal description. In addition, a distribution map, insights about its habitat and ecology, conservation status, and illustrations are provided.
During the austral spring-summer between 2015–2021 several botanical explorations were carried out in the Andean ranges within the locality of San Fabián de Alico, Punilla Province, Ñuble Region, Chile (Fig.
Herbarium specimens were collected from Laguna Añil, and distributed to the herbaria of CONC, EIF, JBN and SGO (acronyms after
The morphological study was based on observations and measurement of fresh and dried specimens. Detailed photographs of fresh material were taken in the field to document the overall plant morphology and especially the floral structure. In order to accurately describe the vegetative and reproductive morphology of the collected plants, dry and rehydrated specimens were dissected. Ovary, fruits, flowers and leaf details were photographed with a zoom lens and subsequently observed under a binocular microscope. Terminology for describing Valeriana plant parts followed
The conservation status assessment of the species used the International Union for Conservation of Nature criteria (
Chile. Ñuble Region, Punilla Province, San Fabián de Alico, Laguna Añil, crevices and small terraces of granite cliffs, 1724 m elevation, 36°32'00.8"S, 71°23'36.1"W, 7 January 2020, A.E. Villarroel & E. Ponce s.n., (holotype SGO!); 1724 m elevation, 36°32'00.8"S, 71°23'36.1"W, 3 February 2021, A.E. Villarroel & R. Neira (paratypes EIF!, JBN!); 1650 m elevation, 36°32'2.28"S, 71°23'26.62"W, 7 December 2020, K. Menegoz & G. Ossa (paratypes CONC!) (Fig.
The habit and macro-morphology of Valeriana praecipitis is similar to Valeriana philippiana, but differs by its height (including flower stem) that can reach 65.5 cm (vs. 20 cm), rhizome woody, reaching more than 30 cm long and up to 20 mm diameter (vs. semi-woody, to 14 cm long, to 8 mm diam.), basal leaves pinnatisect to pinnatipartite, up to 26 cm long (vs. pinnatilobed to pinnatisect, to 8 cm long), petiole glabrous (vs. pubescent), lobes 1–35 mm long, 1–24 mm wide (vs. 4–8 × 3–7 mm), upper leaves oblanceolate, 14–40 mm long, 5–19 mm wide, margin entire to irregularly undulate or sinuate, (vs. oblong, 6–10 × 3–5 mm, entire), bracts oblanceolate to oblong, up to 20 mm long (vs. oblong, to 7 mm long), bracteoles spathulate to oblong, 3–7 mm long, entire (vs. oblong, 2.5–4.5 mm, erose), inflorescence a relatively diffuse thyrse or compound dichasial cyme (vs. dense compound dichasial cyme, contracted), corolla up to 4.5 mm long (vs. up to 4 mm), stamens 3 mm long (vs. 2 mm), stigma 0.2 mm long (vs. 0.5 mm), growing on cliffs that remain humid all-year (vs. well-drained rocky soils), and endemic to the Ñuble Region (vs. in Chile, V. philippiana can be found in Los Lagos, Aysén and Magallanes Regions) (Table
Morphological differences between Valeriana praecipitis and V. philippiana. Based on
Species | V. praecipitis | V. philippiana |
Habit | Perennial herb, hemicryptophyte, simple or branched | Perennial herb, hemicryptophyte, simple or branched |
Height with inflorescence | 26–65.5 cm | To 20 cm |
Taproot rhizome | Circular, reaching more than 30 cm long, 8–20 mm diameter, sometimes stoloniferous, woody | Circular, reaching 14 cm long, 5–8 mm diameter, sometimes stoloniferous, semi-woody |
Stem | 3.5–6 mm diameter, with very short internodes, forming a basal rosette | 3–6 mm diameter, with very short internodes, forming a basal rosette |
Basal leaves | Pinnatisect to pinnatipartite, oblong, up to 26 cm long | Pinnatilobed to pinnatisect, oblong, up to 8 cm long |
Lobes | Orbicular to obovate, base attenuate, 1–35 × 1–24 mm, overlapped, glabrous, fleshy. In the field, leaf lobes are expanded | Orbicular to obovate, base attenuate, 4–8 × 3–7 mm, overlapped, glabrous, fleshy. In the field, leaf lobes are quite folded |
Petioles | Canaliculated, glabrous | Canaliculated, pubescent |
Upper leaves | Oblanceolate, 14–40 × 5–19 mm, margin entire to irregularly undulate or sinuate | Oblong, 6–10 × 3–5 mm, margin entire |
Bracts | Oblanceolate to oblong, up to 20 mm long | Oblong, up to 7 mm long |
Bracteoles | Spatulate to oblong, 3–7 mm long, margin entire | Oblong, 2.5–4.5 mm long, margin erose |
Inflorescence | A relatively diffuse thyrse or compound dichasial cyme | Dense compound dichasial cyme |
Flowers | Hermaphrodite; corolla infundibuliform; corolla tube 3.5–4.5 mm long, base slightly gibbous; corolla lobes oblong to obovate, 1–1.5 × 1–1.7 mm; stamens 3 mm long, exerted; ovary incipient sterile locules; style 2.2 mm long; stigma lobed laminate to lamellate, less than 0.2 mm | Hermaphrodite; corolla infundibuliform-campanulate; corolla tube 4 mm long, base gibbous; corolla lobes oblong to obovate, 1.5 × 1.5–2 mm; stamens 2 mm long, exerted; ovary incipient sterile locules; style 2.5 mm long; stigma lobed lamellate, 0.5 mm |
Fruits | Ellipsoid, 3 × 1 mm, pubescent; pappus plumose, bristles 11, 3.5 mm long | Ellipsoid, 3–4 × 2 mm, pubescent; pappus plumose, bristles 11–13, 5–7 mm long |
Perennial herb, hemicryptophyte, erect or lax when cliff-hanging, simple or branched from the upper part of the taproot, 4–25 cm tall (26–65.5 cm with inflorescence), 4–28.5 cm wide. Rhizome is dark brown, thick, circular, simple, sometimes branched, reaching more than 30 cm long, 8–20 mm diameter, vertical to lateral, sometimes stoloniferous, woody, tortuous, rough, fetid. Secondary-tertiary roots, numerous, located in the first 3 cm of the upper part of the taproot. Stem merging into the taproot, 3.5–6 mm diameter, with short internodes, forming a basal rosette with 9–25 whorled leaves. Basal leaves deciduous, silvery-green turning yellow-brown at the end of summer, simple, petiolate, pinnatisect, sometimes becoming gradually pinnatipartite at the apex (mainly young leaves), oblong, generally symmetric; blade 3–16 cm long (4–26 cm with petiole), 1.5–6.3 cm wide, glabrous, fleshy, with reticulated veins; petiole green turning purple towards the base, canaliculated, up to 14.5 cm long, 3–13 mm wide at the base, 2–7 mm wide at the blade base, entire, glabrous, midrib visible; lateral lobes opposite to subopposite, superimposed, orbicular to obovate, base attenuate, apex rounded to retuse, margin entire to slightly undulate and involute, 6–26 per blade; larger lobes located in the centre of the blade, 9–35 × 7–22 mm; smaller lobes located at the base of the blade, 1–15 × 1–8 mm; terminal lobe orbicular to obovate, 6.5–23.5 × 6–24 mm, base attenuate, apex rounded to obtuse, occasionally retuse, margin entire to irregularly undulate or lobed. Inflorescence a relatively diffuse thyrse or terminal compound dichasium, sometimes corymboid. Floral stem purple at the base, light green towards the flowers, erect, circular, 28.6–60 cm long, 3.5–6 mm diameter at the base, gradually thinner towards the flowers (1.5–2.9 mm), striated, 5–9 internodes (their length decreasing from base toward the apex), branched in the upper half (2–27 cm long) with 1–6 lateral ascending branch pairs (forming partial inflorescences). Upper leaves green, simple, sessile, oblanceolate, 14–40 × 5–19 mm, opposite, decussate, arranged every 2.7–13.5 cm on the flower stem, leaves’ size decreasing from base toward the inflorescence, margin entire to irregularly undulate or sinuate, base decurrent, apex acute to rounded, occasionally retuse, glabrous, less fleshy than basal leaves, reticulated veins. Bracts green, simple, sessile, oblanceolate to oblong, 7–21.2 × 1–7 mm, decreasing in size towards the inflorescence, opposite, decussate, margin entire, base decurrent, apex variable (acute, rounded or retuse), glabrous, less fleshy than upper leaves, reticulated veins. Bracteoles green, sometimes turning purple towards the apex, simple, sessile, spathulate to oblong, 3–7.5 × 0.5–2 mm, decreasing in size towards the inflorescence, opposite, decussate, margin entire, base decurrent, apex rounded to retuse, glabrous, less fleshy than upper leaves. Flowers hermaphrodite, pentamerous, sessile; calyx green and purple at the top, inconspicuous, fused segments forming a wavy ring, 0.3 mm, pubescent, adnate to the infer ovary, accrescent and persistent on fruit modified into feathery structures forming the pappus; corolla 5, fused petals, white, although buds sometimes with purple-pink tinges, infundibuliform, glabrous, 4–4.5 mm wide; corolla tube 3.5–4.5 mm long, base slightly gibbous; corolla lobes oblong to obovate, 1–1.5 × 1–1.7 mm, perpendicular or slightly inclined in relation to the corolla tube; stamens 3, white, filiform, 3 mm long, exerted, attached in the lower third of the tube; anthers light yellow, ellipsoid, bithecal, dorsifixed, deciduous; ovary inferior, green, tricarpellate, trilocular with 1 fertile locule and 2 incipient sterile locules; style 1, white, filiform, 2.2 mm long; stigma trifid, lobed, laminate to lamellate, less than 0.2 mm. Fruit an achene, yellow-green at the base, turning purple towards the apex, ellipsoid, triquetrous, 3 × 1 mm, pubescent, longitudinally striated on one face, calyx persistent, pappus plumose, 0.5 mm diameter at base, bristles 11, purple-reddish colour, 3.5 mm long, with hairs 0.5 mm long.
Valeriana praecipitis A, B plants growing in natural habitat C rhizome, secondary-tertiary roots D basal leaves with lobes detail E petioles F floral stem, corymboid inflorescence G floral stem, thyrse inflorescence H upper leaves I, J detail of flowers (stamens, style) K bracteoles, ovary, calyx L dry inflorescence, bracts, bracteoles M fruit, pappus. Photographed by Alejandro E. Villarroel and Kora Menegoz.
The specific epithet refers to cliff faces inhabited by these plants. The name means “Valeriana of cliffs” (latin praeceps = steep place, precipice, dangerous; genitive praecipitis).
Flowering from November to December; fruiting from January to February.
Endemic to the Andean ranges of the Ñuble Region, Chile (Fig.
From a phytogeographical point of view, Valeriana praecipitis is part of two vegetational formations and three vegetation belts (
Our field observations in Laguna Añil (1724 and 1650 m elevation) indicate a total of 45 species associated with Valeriana praecipitis. Some of these species include: Chiliotrichum diffusum (G. Forst.) Kuntze, Senecio spp. (Asteraceae), Berberis empetrifolia, Maytenus disticha (Hook.f.) Urb. (Celestraceae), Desfontainia fulgens D. Don (Columelliaceae), Empetrum rubrum Vahl ex Willd., Gaultheria sp., G. poeppigii DC., G. pumila (L.f.) D.J. Middleton, G. tenuifolia (Phil.) Sleumer (Ericaceae), Escallonia alpina Poepp. ex DC., E. rubra (Ruiz & Pav.) Pers., Rayenia malalcurensis Menegoz & A.E. Villarroel (Escalloniaceae), Luzula sp. (Juncaceae), Myrceugenia chrysocarpa (O. Berg) Kausel, Myrteola nummularia (Poir.) O. Berg (Myrtaceae), Nothofagus obliqua (Mirb.) Oerst., N. pumilio (Nothofagaceae), Codonorchis lessonii (Brongn.) Lindl. (Orchidaceae), Ourisia sp., O. coccinea (Cav.) (Plantaginaceae), Chusquea montana Phil. (Poaceae), Saxifraga magellanica Poir. (Saxifragaceae), Quinchamalium chilense Molina (Schoepfiaceae).
Valeriana praecipitis is assessed here as Endangered (EN) under the IUCN categories following criteria B2ab(iii). Criterion B2 was selected because its area of occupancy is < 500 km2 (estimated at 20 km2). Criterion “a” was selected because it is known to exist in two locations (Fig.
Key to Valeriana species present in the Andes range of the Ñuble Region, according to the floristic inventories of
1 | Basal leaves strictly entire | 2 |
– | Basal leaves strictly divided | 3 |
– | Basal leaves entire and divided | 4 |
2 | Basal leaves membranaceous | V. leucocarpa |
– | Basal leaves sub-fleshy or fleshy | 5 |
5 | Basal leaves fleshy | 6 |
– | Basal leaves sub-fleshy | 7 |
6 | Basal leaves opaque green colour | V. carnosa |
– | Basal leaves bright green colour | 8 |
7 | Basal leaves spatulate, ovate or suborbicular, with entire, sinuate or dentate margin; inflorescences paniculiform, densely contracted; fruits ellipsoid and glabrous | V. chilensis |
– | Basal leaves ovate or elliptic, with a markedly sinuate or lobed margin; inflorescences paniculiform, lax; fruits tightly ellipsoid and densely hirsute | V. hebecarpa |
8 | Basal leaves spatulate-obovate, with entire margin; inflorescences spike-like and contracted | V. macrorhiza |
– | Basal leaves spatulate, with entire to pausidentate margin; inflorescences glomeruliform and contracted | V. fonckii |
3 | Basal leaves fleshy | V. praecipitis |
– | Basal leaves membranaceous | 9 |
9 | Stems hirsute, particularly at the nodes; basal leaf segments ovate to oblong; fruits ellipsoid | V. valdiviana |
– | Stems with scarce pubescence; basal leaf segments ovate, oblong, lanceolate-elliptic; fruits ovoid and flat | V. polemoniifolia |
4 | Basal leaves membranaceous | 10 |
– | Basal leaves sub-fleshy | 11 |
10 | Stems cylindrical and striated; basal leaf segments ovate-lanceolate; fruits ovoid or ellipsoid | V. crispa |
– | Stems quadrangular with hairy winged edges; basal leaf segments oblong, ovate, lanceolate or suborbicular; fruits ellipsoid | V. grandifolia |
11 | Maximum plant height 80 cm; entire basal leaves oblong, ovate or elliptic; divided basal leaves pinnatilobate; fruits lageniform | V. laxiflora |
– | Maximum plant height 40 cm; entire basal leaves ovate or elliptic; divided basal leaves pinnatisect; fruits ellipsoid | V. obtusifolia |
Pollen evidence indicates that Valeriana is a Holarctic genus that might have arrived from the northern hemisphere, perhaps before the uplifting of the Andes during the Pleistocene (
The discovery of V. praecipitis in southern South-America follows the recent discovery of Rayenia malalcurensis, a newly described endemic genus and species found in the same area. As mentioned before, since the revision of
In addition to consistent morphological differences, V. praecipitis stands out from other species of the genus due to its specific ecological habitat. Few other Valeriana species are able to grow abundantly in damp rock-cliffs at this high altitude, directly rooted in fissures or small soil pockets (e.g., V. chilensis Borsini in Chile or V. ruizlealii Borsini in Argentina). Rock-cliffs are challenging habitats with high erosion rates, limited soil depth and nutrients availability (
Little is known about rare and endangered species growing in the southern Andean cliff ecosystems, and many open questions remain. Future research needs to be done to identify morphological and physiological adaptations to grow under harsh soil and climatic conditions that could shed light on the future of Andean plant communities in the face of climate change. For instance, how new climatic scenarios could affect their area of occupancy? Are they able to migrate? Or more precisely, how extreme and persistent climatic conditions, like the ongoing mega-droughts, might affect the physiological performance of these species, perhaps pressing viable populations’ threshold to the edge. The recent finding of V. praecipitis and Rayenia malalcurensis, in addition to other rare and endemic species, adds to the importance of monitoring and promoting the conservation of these species (
V. praecipitis is not present in any public protected area in Chile (Fig.
Another threat specific to cliff vegetation is the possible impact that rock-climbing could cause. In Chile (and worldwide) rock-climbing is of increasing interest, attracting more people to the mountains (
Valeriana philippiana. Argentina. Patagonia: Neuquén Province, Cerro Colouhincul, Mar 1927, Comber E00143957 (E); Río Negro Province, Cerro López, Dic 1928, Cordini 1594532 (US). Chile. Los Lagos: Llanquihue Province, Parque Nacional Vicente Pérez Rosales – Volcán Osorno, 200 m elevation, 41°10'S, 72°30'W, Jan 1986, Gardner 107129 (CONC); Osorno Province, Parque Nacional Puyehue - Volcán Casablanca, 1500 m elevation, 40°47'S, 72°10'W, Jan 1988, Gardner & Knees E00023408 (E); Osorno Province, Mirador Puyehue, Feb 1971, Landrum 107764 (SGO).
Fieldwork was financed by FONDECYT N°1181956, and the Instituto de Conservación, Biodiversidad y Territorio, UACh. Alejandro E. Villarroel was supported by CONICYT magister fellowship (22201444). Ricardo Moreno-Gonzalez was supported by DAAD/BECAS Chile, 2014 (57144001) during early stages of the study design. We express our gratitude to the owners and administrators of the private sites where the plant was found, especially to Elías Acuña and Esau Marin. The authors would like to thank the curators of CONC, EIF, JBN, and SGO, especially Jimena Arriagada. We are indebted to Martin Gardner and Nicolás Lavandero for the support and revision of the manuscript. Thanks to Marco Duretto and an anonymous reviewer for their helpful suggestions during the reviewing process of this manuscript. We are grateful to Arón Cádiz-Véliz for the illustration of the species. Thank you so much to our field partners: Gonzalo Ossa, Eduardo Ponce, Rayen Neira and all members of the Bullileo Team.