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Article

Epiphytes as Environmental Bioindicators in Forest Remnants of the Pisaca Reserve: Preserving the Unique Pre-Inca Artificial Wetland of Paltas, Ecuador

by
María Ganazhapa-Plasencia
1,
Erika Yangua-Solano
1,2,
Leslye Ruiz
1,2,
Rolando Andrade-Hidalgo
3 and
Ángel Benítez
1,2,*
1
Carrera de Biología, Departamento de Ciencias Biológicas y Agropecuarias, Universidad Técnica Particular de Loja, San Cayetano Alto s/n, Loja 1101608, Ecuador
2
Biodiversidad de Ecosistemas Tropicales-BIETROP, Herbario HUTPL, Departamento de Ciencias Biológicas y Agropecuarias, Universidad Técnica Particular de Loja, San Cayetano Alto s/n, Loja 1101608, Ecuador
3
Departamento de Ciencias Jurídicas, Universidad Técnica Particular de Loja, San Cayetano Alto s/n, Loja 1101608, Ecuador
*
Author to whom correspondence should be addressed.
Forests 2025, 16(4), 628; https://doi.org/10.3390/f16040628
Submission received: 21 February 2025 / Revised: 28 March 2025 / Accepted: 29 March 2025 / Published: 3 April 2025
(This article belongs to the Special Issue The Role of Bryophytes and Lichens in Forest Ecosystem Dynamics)
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Abstract

:
Epiphytic organisms are characteristic elements of the Andean dry forest, playing a crucial role in ecosystem diversity and functionality, but they are threatened by deforestation-related factors. The diversity of epiphytic lichens and bryophytes was recorded in the Pisaca Reserve, which has an artificial pond locally known as “Laguna Pisaca”, serving as a critical micro-watershed. This pond provides water services to the city of Catacocha, motivating local communities to protect its biodiversity. In each zone (low, middle and high), 10 plots of 5 × 5 m were established, where the presence and coverage of lichens and bryophytes were sampled in 4 trees per plot (120 trees). Richness and diversity (Shannon–Weaver and Simpson indices) were calculated. Generalized linear models (GLM) were used to analyze the effect of the zone on richness and diversity, and multivariate analysis was used to analyze species composition. A total of 90 species were recorded (65 lichens and 25 bryophytes), distributed in three zones: 74 in the high, 67 in the low and 41 in the middle zone. Species richness and composition showed significant variations in relation to the three zones, influenced by forest structure, small altitudinal changes and forests disturbance. The forests of the Pisaca Reserve harbor a great diversity of lichens and epiphytic bryophytes, which serve as refuges for biodiversity in the Andean dry montane forest of South Ecuador.

1. Introduction

The Andean dry forest is one of Ecuador’s distinctive ecosystems, located between 1600 and 2300 m altitude [1,2]. These forests harbor a high diversity; however, there are multiple threats, including selective logging, habitat degradation, forest fragmentation, fires, land conversion for agriculture and cattle ranching [3]. In these forests, unique flora and fauna thrive and they play an essential role in biological processes and conservation [4,5].
Epiphytes are an important component of this forests, in terms of diversity and functionality [2,6]. However, only a few studies have focused on dry forest epiphytes; for example, two studies on lichen and bryophyte epiphytes have been conducted in seasonal dry forests of Ecuador [7,8], and following this pattern, local studies on bryophytes in montane Inter-Andean dry forests have been conducted in northern Ecuador [2,6], focusing on vascular and non-vascular epiphytes. Thus, to our knowledge, this study is the first to assess lichen and bryophyte diversity in montane dry forests of southern Ecuador.
Bryophytes and lichens are strongly influenced by macro- and micro-environmental variables, which affect their richness, diversity and community composition on local and regional scales [8,9]. At the regional level, altitude, latitude, temperature and precipitation influence their diversity [10,11,12,13,14,15]. On the other hand, at the local level, forest structure, microclimate and host traits (tree species, pH, bark type) have been identified as principal drivers of their diversity, and due to which they are strongly adapted to the microclimate conditions [8,16,17,18,19]. Therefore, the lack of mechanisms to regulate water uptake and loss makes them sensitive indicators of climatic conditions and can be used as indicators of environmental change [8,17,20,21], related with forests disturbance [22], air pollution [23] and climatic change [24].
The Pisaca Reserve, situated in southern Ecuador within the canton of Paltas in the province of Loja, conserves the last remnants of Andean dry forest in the region, which are rich in biodiversity; for instance, there are 34 species of vascular plants, which belong to 33 genera and 21 botanical families [3,25]. This protected area has an artificial pond called by the locals “Laguna Pisaca” that serves as a critical micro-watershed, providing essential water services to the city of Catacocha and motivating local communities to protect its biodiversity [25,26]. For example, in 2018, UNESCO’s Intergovernmental Hydrological Programme (IHP) designated the Paltas Catacocha WS&H system as an ecohydrology demonstration site, linked to the presence of an artificial pond or Cocha in the Pisaca reserve, within the Paltas Catacocha demonstration area that was restored between 2005 and 2008 [26].
Previous research in the Pisaca Reserve has primarily focused on raising tourism awareness among local residents [27]. Additionally, studies have examined the effects of soil conditioners on the growth and survival of Caesalpinia spinosa Kuntze [28]. Thus, Caesalpinia spinosa (commonly known as Vainillo) has been proposed as a sustainable productive alternative for the Province of Loja [29]. In addition, the area is recognized as part of the “Archaeological Route of the Canton of Paltas”, underscoring its significance as an invaluable site for the city [30]. However, our research is the first to assess the diversity of bryophytes and lichens in the Pisaca Reserve, as studies have been limited to tourism, hydrology and vascular plant diversity.
In the present study, we analyzed the response of epiphytic communities (lichens and bryophytes) to three different zones of Andean dry forest. We hypothesized that, under similar environmental conditions, differences in species diversity and community structure would result from differences in elevation and disturbance, which are associated with changes in forest structure and microclimate. Specifically, we wanted to answer the following questions: (A) Do the different elevations and disturbances influence the richness, diversity and composition of epiphytic communities? and (B) Is forest structure and canopy cover the main factor controlling the structure of epiphytic communities?

2. Materials and Methods

2.1. Study Area

The study was conducted in the Pisaca Reserve, situated in the canton of Paltas in the province of Loja (Figure 1). This reserve is part of the Ecuadorian Andes biogeographical region and is classified as an Andean dry forest, with an area of 39.87 hectares and elevation ranging from 1560 to 2220 m a.s.l. [3,22]. The Pisaca Reserve has a warm, dry tropical climate, with an average annual temperature of 18.6 °C and it is notable for its rich vegetation [22].
We selected three zones with different elevations and disturbance levels [3] (Figure 2). The low zone (1750–1850 m a.s.l.), historically affected by high disturbance, currently presents a reforestation process where species such as Vachellia macracantha (Humb. & Bonpl. ex Willd.) Seigler & Ebinger, Fulcaldea laurifolia (Bonpl.) Poir. and Duranta repens L. predominate [3]. The middle zone (2040–2140 m a.s.l.), historically affected by medium disturbance, a transition zone between dry forest and Andean forest, is home to species such as Lafoensia acuminata (Ruiz & Pav.) DC., Myrcianthes sp. and Mauria heterophylla Kunth [3]. Finally, the high zone (2160–2320 m a.s.l.) is characterized by fragments of native forest (low disturbance) with a humid microclimate and the presence of floristic elements typical of the Andean region, where Myrcianthes sp., Lafoensia acuminata, Xylosma sp., Myrcia fallax (A.Rich.) DC. and Mauria heterophylla predominate [3].

2.2. Desing and Data Sampling

In each zone, a total of 10 plots of 5 × 5 m were established with distance of 5 m between them. Within each plot, 4 trees were randomly selected to assess lichen and bryophyte cover on the trunk of each tree using 10 × 50 cm quadrats [31,32]. In addition, elevation, light, bark type and DBH (Diameter breast height) were recorded. Sampling was carried out in July and August 2023. The lichen and bryophytes samples were transferred to the Herbarium of the Universidad Técnica Particular de Loja (HUTPL) for identification, using taxonomic general keys for lichens [33,34,35,36] and bryophytes [37,38,39,40]. In addition, we used keys for specific groups [41,42,43,44].

2.3. Data Analysis

Sampling efficiency was assessed in three zones using a Chao 2 estimator. Richness and diversity were calculated using Shannon–Weaver and Simpson indices. Variation in richness, abundance and diversity in each zone was represented with box plots. To analyze the effects of zone on richness, abundance and diversity, generalized linear models (GLM) with a Poisson error distribution and a logarithmic link function were used [45].
NMDS was used to visualize differences in species composition between zones with Bray–Curtis distance and 999 Monte Carlo permutations. We performed a permutation-based multivariate analysis (PERMANOVA) to assess the effect of different zones in bryophyte and epiphytic lichen composition. All these analyses were carried out using the statistical software R version 3.6.3 with the statistical package ‘vegan’ [46]. To determine the indicator species in each zone, we carried out an indicator species analysis [47]. The indicator species analysis (ISA) provides an indicator value (IV) for each species in each zone. We used the IndVal function of the “labdsv” package [48]. The indicator value ranges from 0 (one species was absent from one zone) to 1 or 100 (one species occurred in all trees of one zone and was absent from other trees).

3. Results

A total of 90 species were identified (Table A1): 65 lichens (32 foliaceous, 29 crustaceous and 4 fruticose) and 25 epiphytic bryophytes (11 liverworts and 14 mosses). The Chao 2 richness estimator indicated a higher estimated number of species in the high zone (ZA), with 74 species, in contrast to the low zone (ZB), which presented 67 species, followed by the middle zone (ZM) with 41 species. The box plots show that the high zone (ZA) has a higher richness, abundance and diversity compared to the low (ZB) and medium (ZM) zones (Figure 3).
A greater richness of pleurocarpous mosses and foliose liverworts, as well as gelatinous and foliose lichens were observed in the high zone, whereas the low zone contained more species of crustose, fruticose lichens and acrocarpous mosses (Figure 4).
The GLM model revealed that richness, abundance and Shannon and Simpson indices were positively influenced by the high zone (ZA), whereas ZM and ZB showed negative effects on these diversity metrics (Table 1). In addition, a significant effect of DBH on richness and of light on abundance was observed (Table 1).
The NMDS analysis indicated that in the high zone (ZA) a greater uniformity of species distribution was observed. In contrast, in the low zone (ZB) and middle zone (ZM), species showed a wider dispersion (Figure 5). Thus, results showed that the epiphyte community composition had changed relative to the zone.
The results of the PERMANOVA test showed that the zone variable exerts a significant influence on the composition of the epiphytic bryophyte and lichen communities, explaining 12% of the variability observed in these communities. On the other hand, the amount of light and diameter at breast height (DBH) were found to have a significant effect on the composition of epiphytic communities; however, the variance is very low at 0.1% (Table 2).
The ISA showed that 33 species distributed in 15 bryophytes and 18 lichens had significant IVs for the different zones (Table 3). All the zones had different indicator species that were specific to those environmental conditions (Table 3).

4. Discussion

The results showed that the Andean dry forest zones of the Pisaca Reserve in southern Ecuador have a high diversity and can be considered as refuges of lichen and bryophyte diversity, as our study reported 28 epiphytic bryophyte species and 66 lichen species. This is in contrast with the low richness of bryophytes (13 species) and the lack of presence of lichen species reported by Werner and Gradstein [2] in the montane dry forests of northern Ecuador.
The bryophyte and lichen species richness and diversity were related to elevation, disturbance and forest structure. In the high zone (ZA), richness, cover and diversity were higher due to the native forest, which provides favorable conditions for the development of lichens and bryophytes (pleurocarpous mosses and foliose liverworts), related to a more closed canopy and a microclimate with higher humidity and lower light intensity [2,3,49]. In line with this, several studies have shown that foliose liverworts are very sensitive to forest disturbance, including the disappearance of several species [6,49,50].
In contrast, the lower and middle zones have historically been subject to disturbance [3], which implies a negative impact on richness and diversity, as documented by research in Andean montane dry forests [2,8]. Despite this, the ZB showed greater richness, cover and diversity than the ZM, due to the fact that most species are lichens that prefer drier habitats related with forests disturbance [2].
In terms of species composition, it is different in the three zones, where the ZA with the wet canopy and large diameter trees is mainly dominated by bryophytes (foliose liverworts and pleurocarpous mosses) and gelatinose lichens (Figure 6). In this zone, we recorded bryophyte species with high indicator values (e.g., Bryopteris flicina, Neckera chilensis, Porella leiboldii, Porotrichum expansum and Thysananthus auriculatus) and cyanolichens (e.g., Leptogium chloromelum and Leptogium milligranum), which are conditioned to humid microclimates [20,51]. These species are associated with the high diversity, abundance and mature trees recorded in the upper part of the Pisaca Reserve [3]. In addition, the forest remnants in the upper zone are influenced by the moisture provided by the Pisaca wetland [26], creating a favorable habitat for a greater diversity of bryophytes (e.g., Porella leiboldii) and macrolichens (e.g., Lobariella subexornata) with high moisture requirements [29,49,50,51].
On the other hand, the lower zone showed foliose macrolichens with high indicator values (e.g., Leucodermia leucomelos, Physcia lacinulata), species with a crustose growth form (Chrysothrix candelaris, Coniocarpon cinnabarinum, Ramboldia haematites) and fruticose lichens such as Ramalina celastri, Teloschistes flavicans, Usnea cornuta and Usnea strigosa. Similarly, previous studies have shown that these species are associated with disturbed forests [31,51], and some species have been recorded in dry forests where crustose lichens are dominant [8,52]. In addition, species of the genera Ramalina, Teloschistes and Usnea present secondary metabolites (e.g., usnic acid and parietin) that allow them to protect themselves against excessive light availability [31], which is common in forests of the ZB.
Forests in the ZB have less vegetation cover, density and basal area [3], as much of the forest cover in the ZM and ZB has been lost due to the exploitation of wood for fuel and housing, and land use change from forest to crops and pastures. Although our study has shown that bryophytes and lichens can be indicators of disturbance and environmental change, other factors, such as host tree species, pH and bark moisture, are influential elements in the diversity of lichens and epiphytic bryophytes, suggesting areas for future research that were not evaluated in this study [31,49,53]. For this reason, our results are limited to the Andean dry montane forests of the Pisaca Reserve, as we do not have any other remnants of forest with similar conditions.

5. Conclusions

The Andean dry forest of the Pisaca Reserve can be considered a refuge for a high diversity of epiphytes, with a total of 94 species distributed in different zones, where the richness, diversity and composition of lichen and bryophyte communities are different. The high zone (ZA), with more canopy cover, large trees and high plant diversity, is mainly dominated by bryophytes and cyanolichens, which are restricted to the humid understory due to the influence of the moisture provided by the Pisaca pond. On the other hand, in the low zone (ZB) and middle zone (ZM), lichens with crustose, foliose and fruticose growth forms, which have secondary metabolites, were abundant in these zones with less canopy cover and low plant diversity. Therefore, the high diversity of lichens and epiphytic bryophytes and their relationship with different microclimatic factors in the Pisaca reserve underline the importance of these organisms in maintaining the functioning and conservation of the Pisaca pond.

Author Contributions

Conceptualization, Á.B. and M.G.-P.; methodology, Á.B., M.G.-P. and L.R.; formal analysis, M.G.-P. and Á.B.; investigation, M.G.-P. and Á.B.; writing—original draft preparation, Á.B., M.G.-P., E.Y.-S. and R.A.-H. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Universidad Técnica Particular de Loja, grant POA-VIN-056.

Data Availability Statement

Data are contained within the article.

Acknowledgments

We thank the Private Technical University of Loja (UTPL) for funding this Open Access publication. We thank Naturaleza y Cultura for access to the Pisaca Reserve.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A

Table A1. Species of bryophytes and lichens occurring in the three different zones.
Table A1. Species of bryophytes and lichens occurring in the three different zones.
SpeciesZBZMZA
Bryophytes
Bryopteris flicina (Sw.) Nees.064
Frullania ericoides (Nees) Mont.30216
Lejeunea laetevirens Nees & Mont.9921
Diplasiolejeunea cavifolia Steph.175
Metzgeria lechleri Steph.134
Metzgeria rufula Spruce.137
Plagiochila aff. simplex (Sw.) Lindenb.197
Porella leiboldii (Lehm.) Trevis9730
Radula javanica Gottsche.0110
Radula quadrata Gottsche.010
Thysananthus auriculatus (Wilson & Hook.) Sukkharak & Gradst. 9517
Brachythecium plumosum (Hedw.) Schimp.502
Campylopus richardii Brid.204
Cryphaea patens Müll. Hal.403
Cyrto-hypnum minutulum (Hedw.) W.R.Buck & H.A.Crum.010
Fabronia ciliaris (Brid.) Brid.11812
Fissidens steerei Grout.5156
Leptodotium longicaule Mitt., J. Linn. Soc., Bot. var Longicaule.301
Macromitrium podocarpi Müll. Hal.3116
Neckera chilensis Mont.02522
Porotrichum expansum (Taylor) Mitt.082
Orthostichella pentasticha (Brid.) W.R. Buck.3154
Rhynchostegium scariosum (Taylor) A.Jaeger.031
Squamidium macrocarpum (Mitt.) Broth.5156
Syntrichia amphidiacea (Müll. Hal.) R.H.Zander.101
Lichens
Arthonia ilicina Taylor002
Bacidia sp19215
Bacidia sp2400
Bulbothrix isidiza (Nyl.) Hale100
Caloplaca sp1201
Caloplaca sp2100
Chrysothrix candelaris (L.) J. R. Laundon 600
Coccocarpia palmicola (Sprengel) Arv. y DJ Galloway 001
Coenogonium luteum (Dicks.) Kalb & Lücking002
Coenogonium roumeguerianum (Müll. Arg.) Kalb016
Coniocarpon cinnabarinum DC.100
Dirinaria picta (Sw.) Clem. y esquivar403
Fissurina columbina (Tuck.) Staiger012
Flavoparmelia sp.302
Flavoparmelia ecuadorensis T.H. Nash, Elix & J. Johnst. 102
Flavoplaca citrina (Hoffm.) Arup, Frödén & Søchting305
Glyphis cicatricosa Ach.301
Graphis elegans (Borrer ex Sm.) Ach. 110
Graphis leptoclada Müll. Arg.1255
Herpothallon granulare (Sipman) Aptroot & Lücking 141
Heterodermia sp1101
Heterodermia sp2002
Heterodermia granulifera (Ach.) Culb.11116
Hypotrachyna sp.100
Hypotrachyna cirrhata (Fr.) Divakar, A. Crespo, Sipman, Elix & Lumbsch203
Hypotrachyna lipidifera (Hale & M. Wirth) Divakar, A. Crespo, Sipman, Elix & Lumbsch020
Lecanora chlarotera Nyl.211
Lecanora leprosa Fée702
Lecanora tropica Zahlbr.502
Lepraria sp.203
Leptogium chloromelum (Ach.) Nyl.3216
Leptogium milligranum Sierk 8220
Leptogium phyllocarpum (Pers.) Mont. 502
Leptogium aff. pseudofurfuraceum P.M. Jørg. & Wallace002
Lobariella exornata (Zahlbr.) Yoshim.002
Lobariella subexornata (Yoshim.) Yoshim. 0718
Leucodermia leucomelos (L.) Kalb 1000
Parmotrema chinense (Osbeck) Hale & Ahti202
Parmotrema fasciculatum (Van.) Hale101
Parmotrema mellissii (C.W. Dodge) Hale403
Parmotrema reticulatum (Taylor) M. Choisy 507
Parmotrema robustum (Degel.) Hale506
Parmotrema subisidiosum (Müll. Arg.) Hale & Fletcher102
Parmotrema subsumptum (Nyl.) Hale502
Pertusaria sp.101
Pertusaria texana Müll. Arg. 201213
Phaeographis sp.500
Phaeographis dendritica (Ach.) Müll. Arg.306
Phaeographis scalpturata (Ach.) Staiger023
Phyllopsora buettneri (Müll.Arg.) Zahlbr. 031
Phyllopsora parvifolia (Pers.) Müll. Arg.021
Phyllopsora aff. parvifoliella (Nyl.) Müll. Arg.200
Phyllopsora furfuracea (Pers.) Zahlbr.052
Physcia lacinulata Müll. Arg. 1001
Polyblastidium albicans (Pers.) SY Kondr., Lőkös & Hur002
Porina aff. nucula Ach.533
Pyrenula sp.217
Ramalina celastri (Sprengel) Krog & Swinscow300
Ramboldia aff. haematites (Fée) Kalb, Lumbsch & Elix200
Sticta beauvoisii Delise 105
Sticta aff. damicornis (Sw.) Ach.112
Syncesia farinacea (Fée) Tehler102
Teloschistes flavicans (Sw.) Norman200
Usnea cornuta Körb.700
Usnea strigosa (Ach.) Eaton400

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Figure 1. Study area in the Pisaca reserve. (A) Ecuador country with the province of Loja; (B) Cantón Paltas; and (C) Three zones with different elevations and disturbance levels, ZA; high zone, ZM; middle zone, ZB; low zone.
Figure 1. Study area in the Pisaca reserve. (A) Ecuador country with the province of Loja; (B) Cantón Paltas; and (C) Three zones with different elevations and disturbance levels, ZA; high zone, ZM; middle zone, ZB; low zone.
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Forests 16 00628 g002
Figure 2. Study area in the Pisaca reserve in the canton of Paltas, province of Loja shown three zones, (A) High zone, (B) middle zone, (C) low zone.
Figure 2. Study area in the Pisaca reserve in the canton of Paltas, province of Loja shown three zones, (A) High zone, (B) middle zone, (C) low zone.
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Forests 16 00628 g003
Figure 3. Box plots of richness, abundance and diversity indices (Shannon-Weaver and Simpson).
Figure 3. Box plots of richness, abundance and diversity indices (Shannon-Weaver and Simpson).
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Forests 16 00628 g004
Figure 4. Barplot showing the richness of different growth forms of bryophytes and lichens.
Figure 4. Barplot showing the richness of different growth forms of bryophytes and lichens.
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Forests 16 00628 g005
Figure 5. Non-parametric multidimensional scaling (NMDS) analysis of bryophytes an lichens composition in three zones with different elevations and disturbance levels.
Figure 5. Non-parametric multidimensional scaling (NMDS) analysis of bryophytes an lichens composition in three zones with different elevations and disturbance levels.
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Forests 16 00628 g006aForests 16 00628 g006b
Figure 6. Several species of bryophytes and lichens with high indicator values.
Figure 6. Several species of bryophytes and lichens with high indicator values.
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Table 1. Generalized linear models (GLM) results for richness, abundance, Shannon-Weaver and Simpson as a function of zones. t = t-statistics; St = Standard error.
Table 1. Generalized linear models (GLM) results for richness, abundance, Shannon-Weaver and Simpson as a function of zones. t = t-statistics; St = Standard error.
RichnessStimatorSttp-Value
Zone ZA8.600511.277186.734<0.0001
Zone ZB−2.652620.68202−3.8890.00016
Zone ZM−5.257050.81233−6.472<0.0001
Light 0.062940.078620.8010.42500
DAP0.028760.024931.1540.25104
Coverage
Zone ZA94.651016.53335.725<0.0001
Zone ZB−23.63118.8288−2.6770.0085
Zone ZM−23.133210.5158−2.2000.0298
Light0.45930.32281.4230.1573
DAP−0.12431.0177−0.1220.9030
Shannon-Weaver
Zone ZA1.8919720.1897149.973<0.0001
Zone ZB−0.3643730.101308−3.5970.000476
Zone ZM−0.8058770.120665−6.679<0.0001
Light0.0082080.0116780.7030.483580
DAP0.0015730.0037040.4250.671860
Simpson
Zone ZA0.81482800.063477412.837<0.0001
Zone ZB−0.08385790.0338972−2.4740.0148
Zone ZM−0.20835020.0403738−5.161<0.0001
Light0.00137820.00390740.3530.7249
DAP0.00026170.00123920.2110.8331
Table 2. Non-parametric multidimensional scaling analysis (NMDS). Df = degrees of freedom; SS = sum of squares; R2 = coefficient of variation; F = F-statistics.
Table 2. Non-parametric multidimensional scaling analysis (NMDS). Df = degrees of freedom; SS = sum of squares; R2 = coefficient of variation; F = F-statistics.
FactorDfSSR2Fp-Value
Zone25.7210.120498.14660.001
Light10.7510.015812.13810.005
DAP10.6300.013271.79510.015
Residual11540.3800.85043
Total11948.4821.00000
Table 3. The indicator species of bryophytes and lichens of the three zones with their indicator values.
Table 3. The indicator species of bryophytes and lichens of the three zones with their indicator values.
SpeciesZoneIndicator Value p-Value Growth Forms
Bryophytes
Frullania ericoides (Nees) Mont.ZB56.030.0001 Foliose liverwort
Porella leiboldii (Lehm.) TrevisZA53.280.0001 Foliose liverwort
Neckera chilensis Mont.ZM34.930.0003 Pleurocarpous moss
Macromitrium podocarpi Müll. Hal.ZA31.490.0001 Acrocarpous moss
Radula javanica GottscheZM27.50.0001 Foliose liverwort
Fissidens steerei Grout.ZM27.080.0012 Acrocarpous moss
Squamidium macrocarpum (Mitt.) Broth.ZM27.080.0012 Pleurocarpous moss
Thysananthus auriculatus (Wilson & Hook.) Sukkharak & Gradst. ZA26.890.0007 Foliose liverwort
Orthostichella pentasticha (Brid.) W.R. Buck.ZM26.720.0001 Pleurocarpous moss
Lejeunea laetevirens Nees & MontZA18.440.0407 Foliose liverwort
Porotrichum expansum (Taylor) Mitt.ZM17.070.0012 Pleurocarpous moss
Metzgeria rufula SpruceZA11.250.0228 Thalose liverwort
Diplasiolejeunea cavifolia StephZM10.940.0399 Foliose liverwort
Brachythecium plumosum (Hedw.) SchimpZB10.340.0281 Pleurocarpous moss
Bryopteris flicina (Sw.) Nees.ZM9.0570.0495 Foliose liverwort
Lichens
Lobariella subexornata (Yoshim.) Yoshim. ZA37.50.0001 Foliose
Leptogium chloromelum (Ach.) Nyl.ZA32.860.0001 Gelatinose
Leptogium milligranum Sierk ZA28.530.0004 Gelatinose
Leucodermia leucomelos (L.) Kalb ZB250.0001 Foliose
Pertusaria texana Müll. Arg. ZB22.940.0247 Crustose
Physcia lacinulata Müll. Arg. ZB21.550.0001 Foliose
Graphis leptoclada Müll. Arg.ZB17.560.0079 Crustose
Usnea cornuta KörbZB17.50.0006 Fruticulose
Chrysothrix candelaris (L.) J. R. Laundon ZB150.001 Crustose
Lecanora leprosa FéeZB13.750.0051 Crustose
Coenogonium roumeguerianum (Müll. Arg.) KalbZA13.70.0032 Crustose
Phaeographis dendritica (Ach.) Müll. Arg.ZA11.250.0193 Crustose
Phyllopsora furfuracea (Pers.) Zahlbr.ZM11.180.02 Squmulose
Sticta beauvoisii Delise ZA10.940.0136 Foliose
Usnea strigosa (Ach.) EatonZB100.0118 Fruticulose
Lecanora tropica Zahlbr.ZB9.3750.033 Crustose
Leptogium phyllocarpum (Pers.) Mont. ZB9.3750.0331 Gelatinose
Ramalina celastri (Sprengel) Krog & SwinscowZB7.50.0338 Fruticulose
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Ganazhapa-Plasencia, M.; Yangua-Solano, E.; Ruiz, L.; Andrade-Hidalgo, R.; Benítez, Á. Epiphytes as Environmental Bioindicators in Forest Remnants of the Pisaca Reserve: Preserving the Unique Pre-Inca Artificial Wetland of Paltas, Ecuador. Forests 2025, 16, 628. https://doi.org/10.3390/f16040628

AMA Style

Ganazhapa-Plasencia M, Yangua-Solano E, Ruiz L, Andrade-Hidalgo R, Benítez Á. Epiphytes as Environmental Bioindicators in Forest Remnants of the Pisaca Reserve: Preserving the Unique Pre-Inca Artificial Wetland of Paltas, Ecuador. Forests. 2025; 16(4):628. https://doi.org/10.3390/f16040628

Chicago/Turabian Style

Ganazhapa-Plasencia, María, Erika Yangua-Solano, Leslye Ruiz, Rolando Andrade-Hidalgo, and Ángel Benítez. 2025. "Epiphytes as Environmental Bioindicators in Forest Remnants of the Pisaca Reserve: Preserving the Unique Pre-Inca Artificial Wetland of Paltas, Ecuador" Forests 16, no. 4: 628. https://doi.org/10.3390/f16040628

APA Style

Ganazhapa-Plasencia, M., Yangua-Solano, E., Ruiz, L., Andrade-Hidalgo, R., & Benítez, Á. (2025). Epiphytes as Environmental Bioindicators in Forest Remnants of the Pisaca Reserve: Preserving the Unique Pre-Inca Artificial Wetland of Paltas, Ecuador. Forests, 16(4), 628. https://doi.org/10.3390/f16040628

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