Research Article |
Corresponding author: Vítězslav Plášek ( vitezslav.plasek@osu.cz ) Academic editor: Matt von Konrat
© 2024 Vítězslav Plášek, Jakub Sawicki, Felipe Osorio, Monika Szczecińska, Hana Režnarová.
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:
Plášek V, Sawicki J, Osorio F, Szczecińska M, Režnarová H (2024) Orthotrichum camanchacanum, a remarkable new moss species from Chile (Bryopsida, Orthotrichaceae). PhytoKeys 242: 51-67. https://doi.org/10.3897/phytokeys.242.120717
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Orthotrichum camanchacanum is presented as a newly described species from Chile. The species is primarily distinguished by its emergent capsule with cryptoporous stomata, a double peristome, linear-lanceolate stem leaves with a long hyaline aristae in apex, conspicuously differentiated perichaetial leaves, and a densely hairy vaginula. The species was discovered in the mountain massif of the Andes in the Coquimbo region, notable for its unique climatic conditions. Molecular data and a brief discussion comparing the newly described species with the most closely related taxa are also provided.
Bryophytes, new taxa, Orthotrichaceae, South America, taxonomy
Orthotrichum Hedw. is a cosmopolitan moss genus, mainly distributed in temperate regions of both northern and southern hemispheres. Similarly, as is the case with most of such moss genera, Orthotrichum has been recognized as a heterogeneous taxon (
Orthotrichum in Chile was comprehensively treated by
Occurrence of Orthotrichum taxa in Chile based on literature data summarizing taxonomic and floristic research.
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Orthotrichum anomalum | ♢ | ♢ | ♢ | |||||||||||
O. assimile | ♢ | ♢ | ♢ | ♢ | ♢ | |||||||||
O. cupulatum var. austroamericanum | ♢ | ♢ | ♢ | ♢ | ||||||||||
O. diaphanum var. podocarpi | ♢ | ♢ | ||||||||||||
O. freyanum | ♢ | ♢ | ♢ | |||||||||||
O. gigantosporum | ♢ | ♢ | ♢ | |||||||||||
O. inclinatum | ♢ | ♢ | ♢ | |||||||||||
O. perexiguum | ♢ | ♢ | ♢ | |||||||||||
O. tristriatum | ♢ | ♢ | ♢ | ♢ | ||||||||||
O. truncatum | ♢ | ♢ | ♢ |
Previously, Orthotrichum aequatoreum was also reported from the territory of Chile (
During a bryofloristic survey in the Andes Mountains in 2021, a remarkable epiphytic moss from the genus Orthotrichum was collected. Specimens were carefully gathered, air-dried, and sent to the University of Ostrava for identification and inclusion in the herbarium collections (herbarium OSTR). The material proved to be the first record of this taxon for Chile and, upon closer examination, also to represent a species new to science. Plants were subsequently documented using an Olympus SZ61 trinocular microscope for macrophotographs and Olympus BX53 and IPOS-810 microscopes for microphotographs. Detailed SEM photo-documentation of peristome structures and spores was carried out using a Jeol SEM microscope. All photographs were captured from the holotype (OSTR #8123).
Total genomic DNA from a single individual was extracted using the Qiagen Mini Spin Plant Kit (Qiagen, Germany). Details concerning DNA quantification and nanopore sequencing are identical to those in the previous studies (
Obtained raw reads were trimmed using porechop 0.2.4 and assembled using Flye 2.91 assembler (
Chloroplast sequences of 22 specimens of Orthotrichaceae, including seven from Orthotrichum were used for phylogenetic analysis. Phylogenetic analysis was carried out using the Bayesian inference (BI) according to model and parameters used in previous study (
Plantae olivacea, obscure viridis, usque ad 1-cm altae. Folia erecta, lineari-lanceolata, carinata, apicibus longis acuminatis. Capsulae emergentes, cylindricae. Stomata cryptopora. Vaginula dense pilosa cum capilli longi. Peristomium duplex, exostoma 8 paribus dentium siccitate erectum, endostoma 16 segmentis. Calyptra dense pilosa. Sporae 19–24 µm, leniter papillosae.
Chile, Región de Coquimbo (Region IV), Provincia del Elqui, Comuna de Coquimbo, 2 km southeast of Totoralillo town, GPS: 30°04'26"S, 71°21'13"W (-30.073972, -71.353583), on hills profoundly influenced by the humidity generated by camanchaca, vegetation formed mainly by shrubs (Adesmia argyrophylla Phil. and Echinopsis chiloensis subsp. chiloensis (Colla) H.Friedrich & G.D.Rowley), moss was found epiphytically on bark of shrubs, leg. F. Osorio 4378, 10 Aug 2021, holotype (OSTR #8123); isotype (VALD s.n.).
Plants in dense tufts to 1 cm tall, olive green above, dark green to brown below (Fig.
Moss Orthotrichum camanchacanum was discovered growing epiphytically on the bark of shrubs within vegetation predominantly composed of shrubs such as Adesmia argyrophylla and Echinopsis chiloensis subsp. chiloensis. This epiphytic growth pattern suggests a specific ecological niche for this moss within the ecosystem. See map (Fig.
Macro photographs of Orthotrichum camanchacanum. View on fertile plants: A plants with capsules at various stages of development: young immature capsule (left), capsule closed by lid (middle), and mature capsule with open peristome (right) B dried furrowed capsules emergent from long leaves. Scale bars: 1 mm (A, B). Photographs were taken from the holotype (OSTR #8123).
Micro photographs of Orthotrichum camanchacanum leaves A–G leaves from upper part of a stem (linear-lanceolate with remarkable narrow hyaline apex) H–N leaves from middle and lower part of a stem (many of them are damaged and without apical part) O–S detail views of laminar cells, abaxial views (O apical part P upper part Q middle part R lower part and S base of leaf) T–V perichaetial leaves W leaf sections (from apical to basal part). Scale bars: 1 mm (A–N); 100 µm (O–S); 0.5 mm (T–V); 50 µm (W). Photographs were taken from the holotype (OSTR #8123).
Macro and micro photographs of Orthotrichum camanchacanum sporophyte characters A, B view on mature capsules C, D mature capsule with peristome E detail of peristome F stoma (immersed) on capsule urn G lid H, I spores J, K calyptra covered by long hairs L, M hairy vaginula. Scale bars: 1 mm (A, B); 0.5 mm (C, D); 100 µm (E); 50 µm (F); 0.1 mm (G); 20 µm (H, I); 0.5 mm (J–M). Photographs were taken from the holotype (OSTR #8123).
The authors are delighted to name the species (camanchacanum) after “camanchaca”, a term derived from Aymara native language, signifying marine cloud (fog). Forming along the Chilean coast as a cloud, the camanchaca transforms into a dense fog as it moves inland towards the mountains. This fog provides the humidity essential for plant survival.
Chloroplast genome of Orthotrichum camanchacanum sp. nov. was 123,409 bp long and had a typical quadripartite structure with one small single-copy (SSC), one large single-copy (LSC), and two inverted repeats (IR). The use of third-generation sequencing did not identify any structural heteroplasmy associated with inversions in the SSC region. The newly sequenced plastome comprises 82 gene encoding proteins (including hypothetical chloroplast reading frames like ycf1, 2, 3, 4, 12, and 66), 32 transfer RNA (tRNA), and four ribosomal RNA (rRNA) genes. Notably, the rps12 gene is split into two separate transcription units, namely 5’-rps12 and 3’-rps12, and their transcripts undergo trans-splicing (Fig.
Analysis of molecular diagnostics characters (MDCs) revealed 641 SNPs characteristic for Orthotrichum camanchacanum followed by 646 for O. rogeri and 890 in the case of O. cupulatum. Higher numbers of MDCs were detected for O. crenulatum and O. stellatum, 932 and 1304 respectively.
The phylogenetic relationships based on complete plastomes sequences results in a tree (Fig.
The combination of characters of the newly described species, Orthotrichum camanchacanum, is distinctive, making it easily recognizable. Upon initial observation, the most notable feature of the gametophyte is the shape of the stem leaves, particularly those in the upper third of the stem. They exhibit a linear-lanceolate form with a narrow apex that terminates in a long, hyaline arista (Fig.
Comparison of diagnostic characters in Chilean species of the genus Orthotrichum. Data for the newly described species are presented in bold.
Species / Diagnostic characters | leaf shape | leaf margin | lamina | perichaetial leaves | vaginula | capsule position | exostome teeth | endostome segments | preperistome | spore size (µm) | calyptra | asexual repr. |
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Orthotrichum anomalum | lanceolate | revolute | unistratose | not differentiated | occasionally hairy | long exserted | 8 pairs | absent | present | 14–18 | hairy | not observed |
O. assimile | ovate-lanceolate | revolute | unistratose | not differentiated | naked | emergent | 8 pairs | 8 (or 16) | absent | 15–20 | naked or sparsely hairy | gemmae (uncommon) |
O. camanchacanum | linear-lanceolate | recurved | unistratose | differentiated | densely hairy | emergent | 8 pairs | 16 segments | absent | 19–24 | densely hairy | not observed |
O. cupulatum var. austroamericanum | ovate-lanceolate | recurved | unistratose | not differentiated | naked | shortly emergent | 16 teeth | mostly absent | present | 15.5–20 | hairy | not observed |
O. diaphanum var. podocarpi | ovate-lanceolate | revolute | unistratose | not differentiated | naked | immersed to emergent | 8 pairs (splitting) | 16 segments | absent | 18–22 | hairy | gemmae |
O. freyanum | ovate-lanceolate | revolute | unistratose | not differentiated | naked | emergent to shortly exserted | 8 pairs | 8 segments | absent | 12–15 | hairy | gemmae |
O. gigantosporum | ovate-lanceolate | plane or slightly recurved | unistratose | differentiated | hairly | long exserted | 16 teeth | 16 segments | absent | 31–34 (–40) | naked | not observed |
O. inclinatum | linear-lanceolate | plane or revolute | unistratose | not differentiated | naked | long exserted | 8 pairs | 8 segments | absent | 13–19 | with few scattered hairs | gemmae (occasionally) |
O. perexiguum | narrow lanceolate | plane | bistratose | differentiated | naked | emergent to just exserted | 8 pairs | 8 segments | absent | 14–16 | naked | not observed |
O. tristriatum | ovate-lanceolate | broadly revolute | unistratose | differentiated | naked | emergent to shortly exserted | 8 pairs | 8 segments | absent | 14–18 | with few scattered hairs | not observed |
O. truncatum | ovate-lanceolate | reflexed | unistratose | not differentiated | hairly | shortly exserted | 8 pairs | 8 segments | mostly absent | 20–23 | with few scattered hairs | not observed |
Although Orthotrichum perexiguum also exhibits a narrow and lanceolate leaf shape (
The perichaetial leaves produced by Orthotrichum camanchacanum are distinctive and markedly different when compared with all South American species of this genus. Notably, they possess a unique apex formed by a group of conspicuous hyaline cells (Fig.
The hairiness of the vaginula is considered a distinctive taxonomic character for identifying Orthotrichum species (
When comparing the sporophyte characters of the Chilean species within the genus Orthotrichum, most of them, including the newly described species, have emergent or shortly exserted capsules. However, three species (O. anomalum, O. gigantosporum, and O. inclinatum) produce significantly exserted capsules on a long seta (
It is widely acknowledged that species within the genus Orthotrichum have immersed stomata, distinguishing them from representatives of the related genus Lewinskya, whose urns possess superficial stomata (
The territory where the new species was collected possesses distinctive geographical and climatic conditions. The entire Coquimbo region features several transverse valleys that cut across perpendicular to the main Andes mountain chain, carving through the country horizontally (
The authors have declared that no competing interests exist.
No ethical statement was reported.
The project is co-financed by the European Union under the European Regional Development Fund RPO WO 2014–2020 Operation 1.2 R&D infrastructure. Agreement no. RPOP.01.02.00-16-0001/17-00 from 31.01.2018.; EU structural funding CZ.1.05/2.1.00/19.0388; Ministry of Education, Youth and Sports of the Czech Republic: LO1208; ANID PhD grant 21231650.
FO collected the original specimen and provided additional habitat information; VP identified the material, prepared macro, micro and SEM photographs, and completed the description of the new taxon; JS performed the molecular analyzes and prepared the molecular background for the verification of the new taxon, HR helped with preparation of the material; VP, JS, FO and HR prepared the draft version of the manuscript, which was reviewed, edited, and approved by all co-authors.
Vítězslav Plášek https://orcid.org/0000-0002-4664-2135
Jakub Sawicki https://orcid.org/0000-0002-4759-8113
Felipe Osorio https://orcid.org/0009-0001-0759-920X
Monika Szczecińska https://orcid.org/0000-0002-5377-4304
Hana Režnarová https://orcid.org/0009-0004-2168-4015
All of the data that support the findings of this study are available in the main text.