Species of mosses in the forests of Canchacancha (1), Kontorkayku (2), Kelloq’ocha (3), and Manthanay (4).
Abstract
The diversity of mosses was evaluated in four Polylepis (Rosaceae) forests in the Cordillera del Urubamba, Cusco region – Peru. Epiphytic mosses were collected from the lower base of the trunk, canopy area, and terminal branches in paper bags, selecting specimens with sporophytes. Twenty-seven species distributed in seven orders, 13 families, and 17 genera were determined. Three species are new records for Peru, that is, Neckera ehrenbergii, Zygodon quitensis, and Didymodon challaense. Fourteen species are reported for the first time in the Cusco region. These chorological novelties demonstrate the importance of Polylepis forests as stores of cryptogamic diversity.
Keywords
- diversity
- mosses
- canopy
- terminal branches
- new records
1. Introduction
In the upper parts of the Cusco region – Peru, the vegetation is mostly in the form of grasslands and small shrubs, with the genus
The
The Musci are the most complex group within the Bryophytes, not only because of the morphology of their gametophytes but particularly because of the structure of their sporophytes. They make up the group of Bryophytes with the most species; it is estimated that worldwide there are between 8000 and 12,800 species in 900 genera. The number of mosses estimated for the Andean region of Colombia, Ecuador, and Peru is likely to reach around 900−950 species. Among these countries, Peru requires more work both in terms of additional inventories and published results; however, around 829 taxa are currently known for the entire territory, of which 797 are reported for the Andean region [2]. There is a gradual increase in the diversity of taxa according to elevation, with a maximum between 2500 and 3000 m, beyond this altitude interval, there is a marked decrease in diversity, notably above 3500 m. Thus, in Bolivia, a greater diversity was recorded between 2000 and 2500 m, followed by the 2500−3000 m interval [3].
In this regard, Menzel [4] cites 181 species for the department of Ayacucho, the vast majority collected by Hegewald & Hegewald, who carried out explorations in various departments of Peru between 1973 and 1977, including Ayacucho, some were carried out in the province of Huamanga. Opisso [5, 6] carried out an inventory of pleurocarpous mosses in the province of San Ignacio – Cajamarca, registering 40 species, 32 genera, and 15 families. He also recorded 55 species of mosses for the Pomahuaca-Cajamarca district with 47 genera and 30 families. Opisso and Churchill [7] carried out a study in the surroundings of the Yanachaga-Chemillén National Park in the department of Pasco, in which they indicate the existence of 134 species of bryophytes distributed among 92 genera and 45 families (32 liverworts, 2 hornworts, and 100 mosses).
In the Cusco region, studies on Bryophytes began in 1921 with Fortunato L. Herrera in his work “Flora del Departamento del Cuzco” reporting 26 species of mosses, among which he mentioned
For its part, Carhuapoma [13] carried out the study of mosses of the Historical Sanctuary of the Pampa de Ayacucho in an altitude range of 3350 to 4100 m, registering 67 species belonging to 45 genera, 21 families, and 11 orders. The Pottiaceae family was the most representative with 10 species, followed by Orthotrichaceae, Brachytheciaceae, and Bartramiaceae with six species each, reporting 23 new records for Ayacucho and the third report for Peru of
2. Methods
3. Study area
The scope of study territorially includes the Huarán basin with the towns of Canchacancha (13° 14′ 34.9” S and 72° 01′ 13.4” W) located at 4548 m.a.s.l. and Contorkayku (13° 16′ 03.7” S and 72° 01′ 02” W) at 4310 m.a.s.l. between the limits of the provinces of Calca and Urubamba; the Wayoqhari-Yanacocha basin, locality K′elloq’ocha (13° 16′ 34.3” S and 72° 03′09.7” W) at 4343 m.a.s.l. belonging to the district of Huayllabamba, and the Manthanay basin, locality Manthanay (13° 12′ 08.4” S and 72° 08′ 42.8” W) at 4778 m.a.s.l. located in the heights of the Yanahuara Valley in the province of Urubamba, Peru.
3.1 Natural life zones
3.1.1 Subtropical sub-Andean very humid páramo (pmh-SaS)
This life zone also known as wet puna in the South of Peru; it presents a total annual precipitation that varies between 640 mm and 800 mm and an average annual biotemperature between 6°C and 3°C. It is a cold climate zone. According to the Holdridge Diagram, this life zone has potential evapotranspiration that varies between a quarter (0.25) and a half (0.5) of the average total precipitation per year. Life includes the dense forests of
3.1.2 Subtropical sub-Andean pluvial tundra (tp-SaS)
Life zone includes the ecological system of the cold Andean desert of the humid puna. It is located on the very humid-subalpine and subtropical paramo and below the Nival floor. It has a cold climate, the soils are mostly rocky due to erosion of the old glacial mountains, in whose cracks there is scattered and discontinuous vegetation conditioned and adapted to the daily alternation of ice-thaw and where
4. Sampling
Using the protocol of Gradstein et al. [15], four forests were selected, in which 16
In each forest, epiphytic mosses were collected from the lower base of the trunk, canopy area, and terminal branches. The collection was made using craft paper bags and/or recycled paper envelopes, preferably selecting specimens with a sporophyte as this is often necessary for identification.
4.1 Identification process
To identify the species in the laboratory, it was necessary to rehydrate the samples, so that they recover their natural form. It was enough to submerge a few plants in water for them to rehydrate in a few seconds. In some species of phyllidia or thicker thallus, this operation took a little longer, but it is almost never more than a minute.
Once the sample was rehydrated, with the help of fine-tipped tweezers and a pair of lancets, the parts that we are interested in observing were separated: phyllidia, caulydia, capsules, etc. Something important to keep in mind, before separating the samples, is that the details of the plant’s shape must be observed, such as the arrangement of the phyllidia on the plant (distichous, more or less together or separated) and the type of branching.
Cross sections of the phyllidia of many species of mosses were also made to observe the structure of the central nerve, the thickness (number of cells) of the phyllidia, the presence of papillae or nipples, etc. (Figures 1–8).
The main characteristics observed were as follows:
Habit (acrocarpic, pleurocarpous moss).
Shape of the phyllidia.
Shape and dimensions of the cells of the phyllidia, presence of papillae, etc.
Characteristics of the central nerve of the phyilidia.
Shape and dimensions of the cells of the phyllidia, presence of papillae, etc.
Characteristics of the margin of the phyllidia.
Presence of gems or other vegetatively reproducing structures.
Characteristics of the sporophyte.
5. Taxonomic identification
To determine the species, we first proceeded to separate the samples collected from other nonvascular epiphytes (lichens and liverworts), as well as from the bark of the trees, then proceeded to identify the samples following dichotomous keys [16], photographs of each of the species were taken using a stereoscope and an optical microscope, exhaustively describing and making drawings of the observations. Some of the samples were sent to the city of Santa Cruz in Bolivia for review by a moss specialist.
6. Results
In the forests of Manthanay K′elloq’ocha Contorkayku and Canchacancha, a total of 27 morphospecies of moss were found, which are distributed in seven orders, 13 families, and 17 genera. Twenty of these mosses are identified to species, six to genus, and one only at the family level (Table 1).
Order/Family | Species | 1 | 2 | 3 | 4 |
---|---|---|---|---|---|
Orthotrichales/Orthotrichaceae | x | x | x | x | |
x | x | x | x | ||
— | x | x | — | ||
— | x | x | x | ||
x | x | — | — | ||
x | — | — | — | ||
Hedwigiaceae | — | x | x | x | |
Leucodontales/Neckeraceae | x | x | — | — | |
— | x | x | — | ||
Dicranaceae | x | — | x | x | |
— | — | x | — | ||
Bryaceae | x | — | — | — | |
— | — | — | x | ||
Bartramiaceae | — | — | x | — | |
— | — | x | |||
Hypnales/Brachytheciaceae | x | x | — | x | |
x | x | x | |||
Thuidiaceae | — | x | x | — | |
Sematophyllaceae | — | x | x | — | |
Dicranales/Pottiaceae | x | — | — | x | |
x | x | x | x | ||
— | — | — | x | ||
— | x | x | |||
x | — | — | — | ||
Hookeriales/Daltoniaceae | x | — | — | x | |
Leucodontales/Leptodontaceae | x | x | — | — | |
Lepyrodontaceae | x | x | x | x |
Three species are new records for Peru, that is,
As seen in Table 1, some morphospecies are almost exclusive to a single forest, such as
Figure 9 shows an important beta diversity for the 4
In the box plot, (Figure 10) the richness of species of the four forests is shown graphically, we can observe that there are atypical data for the Kellococha and Manthanay forests, an important aspect to point out is that the box of this last forest presents a great difference when compared to the Canchacancha and Contorkayku boxes, that is why we say that the Manthanay forest presents a difference in terms of the richness of moss species.
In the box plot, (Figure 11) the richness of species there is no significant difference in species richness between living and dead arboretums. The floristic composition between living and dead arboretums is different; however, it was observed that in dead arboretums the species richness is equal to that of living arboretums.
7. Discussions
Acurio [11] carried out an exhaustive study of the diversity of mosses in the Wiñay Wayna – Machupicchu area – Peru, finding a total of 129 species of mosses in 72 genera and 29 families. Making a comparison with the present study we have that: Acurio collected mosses in an altitude range of 2500 to 3100 m, on the other hand, this study was carried out in an altitude range of 4300 to 4800 m, and in investigations carried out by Churchill et al. [3], it was found that the elevation zone with the highest number of moss species is between 2000 and 3000 m. Lithophytic, epiphytic, and terrestrial mosses were evaluated; however, in the present study, only epiphytic mosses were taken, and according to Churchill et al. [3], the most common substrate among mosses is soil, followed by rocks and finally epiphytes. Therefore, the ecological conditions for Wiñay Wayna seem to be much more favorable, which allows the existence of a greater diversity of species. Despite the differences regarding the study area, it is necessary to highlight that, of the 129 species cited by Acurio, three of these species were also reported for this study:
Fuentes & Churchill [17] found, in the Madidi region (Bolivia), a total of 369 species in 168 genera and 54 families, of which seven species are shared with the present study:
For its part, Carhuapoma [13] carried out the study of mosses of the Historical Sanctuary of the Pampa de Ayacucho in an altitude range of 3350 to 4100 m.a.s.l., registering the Pottiaceae as the most representative family with 10 species, followed by Orthotrichaceae, Brachytheciaceae, and Bartramiaceae with six species, which shows that the results are similar to those reported in the present investigation.
It was believed that the adaptation of
8. Conclusions
Studies of mosses in the high mountains of Peru, particularly in
Acknowledgments
We express our gratitude to Dr. Steven P. Churchill for the certification of the determined species, as well as to the Center for Research and Production of Food and Medicinal Mushrooms of the Faculty of Sciences of the National University of San Antonio Abad of Cusco for the ease of laboratory and use of stereo microscopes. We are grateful to ECOAN-Peru for helping with the logistics of the fieldwork.
Funding
Fieldwork was funded by the Leverhulme Trust, UK, the University of Zurich, the SNSF [grant P2ZHP3_161988], and the DFG [grants HE 6726 5/1; JU 2748/6–1].
Conflict of interests
The authors do not incur conflicts of interest.
Ethical/legal aspects
The authors declare that they have not incurred any violation of an ethical or legal nature.
References
- 1.
Fjeldsa J, Kessler M. “Conserving the Biological Diversity of Polylepis Woodlands of the Highland of Perú and Bolivia” a Contribution to Sustainable Natural Resource Management in the Andes. Copenhagen, Denmark: NORDECO; 1996 - 2.
Churchill SP, Aldana MC, Opisso J, Morales T. Familias y géneros de los Musgos de los Andes Tropicales. Editorial La Rosa. Santa Cruz de la Sierra, Bolivia Financiado por National Science Foundation Missouri Botanical Garden. 2020. p. 8 - 3.
Churchill SPN, Sanjines A, Aldana C. Catálogo de las Briofitas de Bolivia: La Diversidad, Distribución y Ecología. La Rosa Editorial, Santa Cruz de la Sierra, Bolivia: Museo de Historio Natural Noel Kempff Mercado y Missouri Botanical Garden; 2009 - 4.
Menzel M. Preliminary checklist of the mosses of Perú. Journal of the Hattori Botanical Laboratory. 1992; 71 :175-254 - 5.
Opisso Mejía JA. Contribución al Conocimiento de los Musgos Pleurocárpicos de la Provincia de San Ignacio. Cajamarca, Lima, Perú: Universidad Nacional Mayor de San Marcos; 2001 - 6.
Opisso Mejía JA. Briófitos de Pomahuaca. Cajamarca-Perú. Arnaldoa. 2003; 10 (2):2,7-14 - 7.
Opisso MJA, Churchill SP. Bryophytes from the environs of Yanachaga Chemillén National Park, Department of Pasco, Peru. The Bryologist. 2008; 111 (2):310-317 - 8.
Galiano WH. The Flora of Yanacocha, a Tropical High Andean Forest in Southern Perú. A Thesis for the Degree of Master of Science. St. Louis, U.S.A: University of Missouri; 1990. p. 270 - 9.
Tupayachi HA. Flora de la Cordillera del Vilcanota. Arnaldoa. 2005; 12 (1–2):126-144 - 10.
Galiano W, Tupayachi A, Nuñez M. “Flora del Valle del Cusco” Historia Natural del Valle del Cusco . 1 era Edición. Peru: Sociedad Protectora de la Naturaleza Cusco; 2005. pp. 197-230 - 11.
Acurio L. ;Musgos de la zona de Wiñay Wayna, Santuario Histórico de Machupicchu Tesis de Investigación Carrera Profesional de Biología. Peru: UNSAAC; 2002 - 12.
Huallparimachi QG, Molina IY, Holgado Rojas ME, Saji SM. Briofitas del Santuario Histórico de Machu Picchu. UNESCO; 2016 - 13.
Carhuapoma Soto JZ. Musgos en el Santuario Histórico de la Pampa de Ayacucho, distrito Quinua, provincia Huamanga. Ayacucho, Peru, 2017; 2019 Tesis para obtener el título profesional de bióloga en la especialidad de ecología y recursos naturales - 14.
Holdridge LR. Life Zone Ecology. San José, Costa Rica: Tropical Science Center; 1967 [Traduccion del ingles por Humberto Jiménez Saa: Ecología Basada en Zonas de Vida, la. Ed. San José, Costa Rica: IICA; 1982] - 15.
Gradstein SR, Nadkarni NM, Kromer T, Holz IN, Noske N. A protocol for rapid and representative sampling of vascular and non-vascular epiphyte diversity of tropical rain forests. Selbyana. 2003; 24 :105-111 - 16.
Gradstein SR, Churchill S, Salazar Allen N. A guide to the bryophytes of tropical America. Memoirs of the New York Bolivia Garden. 2001; 86 :1-577 - 17.
Fuentes A, Churchill S. Catálogo de las briófitas de la región de Madidi, Bolivia. Ecología en Bolivia. 3ra Edición. 2005. pp. 170-198 - 18.
Churchill SP, Griffin D III, Lewis M. Diversidad de Musgos en los Andes Tropicales. New York, USA: The New York Botanical Garden; 1995. pp. 335-346 - 19.
Purcell J, Brelsford A, Kessleer M. “The World's highest Forest” a better understanding of the properties of Andean queuña woodlands has major implications for their conservation. American Scientist. 2004; 92 :454-461