Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Human disturbance promotes an increasing abundance

of shrubby plants in the páramo landscape of Southern Ecuador

Pedro X. Astudillo1*; https://orcid.org/0000-0002-9945-9414

Paul Porras1; https://orcid.org/0000-0002-0428-9070

David C. Siddons1; https://orcid.org/0000-0003-3305-2969

Eduardo Barnuevo1; https://orcid.org/0000-0001-8946-2081

Santiago Barros1; https://orcid.org/0000-0003-2647-6579

1. Laboratorio de Ecología, Escuela de Biología, Universidad del Azuay, Cuenca, Ecuador; pastudillow@uazuay.edu.ec

(Correspondence*); paulporraspolo@gmail.com; dsiddons@uazuay.edu.ec; barflo93@gmail.com;

jsanty.b1@gmail.com

Received 17-IX-2024. Corrected 13-II-2025. Accepted 30-V-2025.

ABSTRACT

Introduction: The páramo grassland ecosystem is an important center of plant diversity and endemism.

However, human activities, such as burning and livestock grazing, are altering the plant composition of the pára-

mos. These changes may be associated with an increase in the abundance of shrubby species and a corresponding

decrease in native grass cover.

Objective: To evaluate the effects of human disturbance on the composition of woody plant species in páramo

habitats.

Methods: We conducted 36 transects across the páramo landscape of the Macizo del Cajas Biosphere Reserve

in Southern Ecuador between April 2017 and November 2019, recording woody plant species (e.g., bushes and

shrubs) along each transect. To explore variations in woody plant composition, we employed non-metric multidi-

mensional scaling, using the proportion of disturbed area, páramo grassland, and elevation as predictor variables.

Results: We recorded a total of 13 377 woody plants. The proportion of disturbed areas has an influence on the

composition of the woody plant species. Shrubby species such as Diplostephium ericoides, Hypericum quitense,

Valeriana microphylla, and Valeriana hirtella are more prevalent in transects with a greater proportion of dis-

turbed areas.

Conclusion: There was a greater presence of fast-growing woody plant species in response to human-induced

disturbance. This suggests that native herbaceous species are gradually being replaced by woody encroachment,

particularly in human-accessible páramos. Conservation and restoration efforts should take this phenomenon

into account to prevent the accelerated spread of woody encroachment and enhance the availability of páramo

grassland habitats.

Key words: high Andes; woody plant encroachment; plant community composition; grasslands; biosphere

reserve.

RESUMEN

Las perturbaciones humanas promueven el incremento en la abundancia

de plantas arbustivas en el paisaje de páramo del sur de Ecuador

Introducción: El ecosistema de páramo herbáceo es un importante centro de diversidad y endemismo de plantas.

Sin embargo, las actividades humanas como el pastoreo y las quemas están alterando la composición de plantas

https://doi.org/10.15517/rev.biol.trop..v73i1.61916

CONSERVATION

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

INTRODUCTION

The Tropical Andes is an important center

of biodiversity and endemism (Jiménez-Rivillas

et al., 2018; Myers et al., 2000). The region is

one of the most biological diverse hotspots

globally (Baquero et al., 2004; Jiménez-Rivillas

et al., 2018; Neill, 1999; Sarmiento, 2000), har-

boring one sixth of the terrestrial plant diversity

on less than 1 % of the global surface (Myers et

al., 2000). Páramo is the major vegetation type

in the high Andes (> 3 000 m.a.s.l.) of North-

ern South America (Llambí et al., 2012). In

Ecuador, the páramos are a mosaic of habitats,

mainly characterized by native tussock grasses

(páramo grassland), but also associated with

small woody plants within marshes, bog plants

in more humid areas (cushion páramo), taller

vegetation in semi-open shrubland (shrubby

páramo), and numerous patches of Polylepis

woodlands (García et al., 2020; Neill, 1999;

Sklenár & Ramsay, 2001).

This páramo ecosystem, like other biodi-

versity hotspots, is impacted by human activi-

ties such as burning to facilitate pastures for

grazing, agricultural expansion and habitat

modification at roadsides (Bagchi et al., 2018;

Barros et al., 2020; Hofstede & Llambí, 2020;

Matson & Bart, 2013; Sylvester et al., 2017).

In Southern Ecuador, livestock grazing is the

main activity affecting the páramo landscape

(Astudillo et al., 2017; Suárez & Medina, 2001)

resulting in the homogenization (i.e., botani-

cally and structurally) of páramo vegetation

(Astudillo et al., 2017; García et al., 2020; Hof-

stede & Llambí, 2020; Jørgensen et al., 2011;

Smart et al., 2006; Sylvester et al., 2017). This

region is also expected to experience changes

in habitat structure and composition due to

climate change (Carrillo-Rojas et al., 2019;

Hofstede & Llambí, 2020). It is predicted that

vegetation adaptable to more extreme climate

ranges (e.g., longer and more intense droughts

with shorter and more intense rainfalls) will

become more dominant (Foster, 2001; Hudson

et al., 2014). These actual and future pressures

have led the páramo ecosystem to become a

priority for biological conservation (Astudillo

et al., 2024; Hofstede & Llambí, 2020; Madriñán

et al., 2013).

The intensity and frequency of human-

induced disturbances have been shown to result

in changes to plant composition (González et al.,

2023; He et al., 2019; Hofstede & Llambí, 2020;

de los páramos. Estos cambios pueden estar asociados a un incremento en la abundancia de especies arbustivas y

una menor cobertura de pastizales nativos.

Objetivo: Evaluar los efectos de la alteración humana sobre la composición de plantas leñosas en hábitats de

páramo.

Métodos: Entre abril de 2017 y noviembre de 2019, recorrimos 36 transectos a través del paisaje de páramo de

la Reserva de la Biósfera del Macizo del Cajas en el sur de Ecuador, registrando las especies de plantas leñosas

(e.g., arbustos y matorrales) a lo largo de cada transecto. Para explorar las diferencias en la composición de las

plantas leñosas, usamos un escalamiento multidimensional no métrico, con la proporción en la cobertura de área

alterada, páramo herbáceo y elevación como variables predictoras.

Resultados: En total, registramos 13 337 plantas leñosas. La proporción de área alterada influye en la composi-

ción de las especies de plantas leñosas. Las especies arbustivas como Diplostephium ericoides, Hypericum quitense,

Valeriana microphylla y Valeriana hirtella tiene mayor presencia en transectos con un incremento en la propor-

ción de las áreas alteradas.

Conclusiones: Hay una mayor presencia de especies de plantas leñosas de crecimiento rápido en respuesta al

efecto de la alteración inducida por los humanos. Esta observación sugiere que las especies herbáceas nativas están

siendo gradualmente reemplazadas por un engrosamiento con plantas leñosas, particularmente en páramos con

acceso humano. Los esfuerzos de conservación y restauración deberían tener en cuenta este fenómeno para evitar

una aceleración de la invasión de plantas leñosas y, asegurar la disponibilidad de hábitats de páramo herbáceo.

Palabras clave: altos Andes; engrosamiento de plantas leñosas; composición de la comunidad de plantas; pasti-

zales; reserva de la biósfera.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Suárez & Medina, 2001; Sylvester et al., 2017).

For example, research shows that excluding fire

leads to an increase in woody plants, such as

shrubs and bushes (Brandt et al., 2013; Matson

& Bart, 2014) with a consequent reduction of

available grazing areas (Matson & Bart, 2014;

Montané et al., 2007). Here, following a fire,

the regeneration of vegetation typically begins

with the establishment of species that have a

patchy distribution, such as those belonging

to the Poaceae family (Jantz & Behling, 2012;

Sarango-Cobos et al., 2019). However, in the

long term, woody plants colonize páramo soils

when they are left to regenerate, mainly occu-

pying areas lacking native grass cover (Matson

& Bart, 2014). This final phenomenon occurs

in conjunction with irregular topography (e.g.,

along elevation gradients and on steep slopes

with minimal grass cover) and facilitates the

encroachment of woody shrubs and bushes

across the páramo landscape (Matson & Bart,

2013). Furthermore, in more extreme climate

regime scenarios, plants with a wider geo-

graphic range, rapid dispersion, and presence

in the montane forest tree line close to the

lower limit of the páramo will also contribute

to woody encroachment (Caballero-Villalobos

et al., 2021; Foster, 2001; Jantz & Behling, 2012;

Loughlin et al., 2018; McKinney & Lockwood,

1999; Montaño-Centellas et al., 2024). Nev-

ertheless, the natural presence of shrubs may

also create favorable conditions (e.g., shade

and protection from the wind), which in turn

may lead to an increase in the species richness

of páramo plant communities (Sylvester et al.,

2017; Vargas-Ríos & Ávila-Rodríguez, 2021).

Therefore, there is an urgent need to moni-

tor páramo habitats to fully understand the

effects of human disturbance and their associ-

ated changes in plant composition in order to

improve conservation and restoration strategies

for natural resource managers. Specifically, it is

important to address how the increasing cover

of woody plants is affecting páramo vegetation

communities. Consequently, in the páramos

of Southern Ecuador, an important region for

plant diversity (Jiménez-Rivillas et al., 2018),

we evaluated the effects of human disturbance

on the vegetation composition of páramo habi-

tats. We expected an increasing abundance of

woody bushes and shrubs in areas with greater

disturbance (mainly influenced by burning

and livestock grazing). Principally, we expected

disturbed areas to have a higher prevalence

of widely distributed, generalist woody plant

species (i.e., faster growing plants and those

associated with the lower limits of páramo and

the upper limits of montane forest).

MATERIALS AND METHODS

Study area: This study was conducted in

the highlands (> 3 500 m.a.s.l.) of the Macizo

del Cajas Biosphere Reserve in SouthWestern

Ecuador (2°55’25’’ S & 79°21’57’’ W) (Fig. 1),

a priority region for biological conservation

(e.g., Astudillo et al., 2024). The study area

is dominated by páramo grassland ecosystem

and covers ~166 000 ha. The Macizo del Cajas

has two protected areas forming the core area

of this reserve, Cajas National Park (2°50’45’’’

S & 79°14’33’’ W) and Quimsacocha National

Recreation Area (3°00’45’’ S & 79°14’12’’ W)

(Rodríguez et al., 2014). These two protected

areas encompass ~19 % of the total area of

páramos within Macizo del Cajas (Fig. 1) (Bar-

ros et al., 2020). The average monthly tem-

perature ranges between 5 and 12 °C. Average

annual precipitation ranges between 1 200 and

1 500 mm, with two rainy seasons; intense

precipitation between March and May and a

second, less intense peak, between September

and February. The lowest precipitation occurs

between June and August (Ballari et al., 2018;

Celleri et al., 2007).

The study area has an elevation range

between 3 600 and 4 050 m. The vegetation

is dominated by tussock grasses of species

of Calamagrostis and Festuca with cushion

plants and rosette species, such as Plantago and

Oreobolus, in smaller humid areas (Minga et

al., 2016; Sklenár et al., 2005). In more hetero-

geneous páramo, the vegetation is associated

with native shrub species mainly of the fam-

ily Asteraceae such as Chuquiraga jussieui J.F.

Gmel, Monticalia arbutifolia (Kunth) C. Jeffrey,

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

Gynoxys miniphylla Cuatrec. (Ansaloni et al.,

2022; Baquero et al., 2004; Minga et al., 2016;

Neill, 1999). In addition, the páramo is inter-

spersed with patches of Polylepis woodlands

(Astudillo et al., 2020; Pinos, 2020). The main

human activity is livestock grazing associated

with burning to enhance pasture (Astudillo et

al., 2017) with evident negative effects on pára-

mo habitats (e.g., less complex habitat structure

with lower plant diversity).

Vegetation sampling: We randomly estab-

lished, in a suitable area (i.e., avoiding extreme

slopes or mountain tops), 36 transects across

protected areas and surrounding zones within

the páramo ecosystem (Fig. 1). Each transect

was 1 km in length and spaced at least 400 m

apart and located in páramo habitats (e.g.,

páramo grassland, shrubby páramo, cushion

páramo). All sampling was carried out between

April 2017 and November 2019.

We used the vegetation sampling protocol

provided by Astudillo et al. (2017), Astudillo

et al. (2019) and Barros et al. (2020). These

protocols have been widely used in monitoring

páramo habitats and woody plants across the

regional páramo. Thus, along each transect,

ten circular plots with a radius of 12 m were

installed, covering a total sampling area of

452.39 m². These plots were regularly spaced

100 m apart. In each circular plot, the propor-

tion of two habitat types was visually estimated

(Fig. 2): (i) páramo grassland (Fig. 2A), a natu-

ral open habitat with herbaceous vegetation

(Ansaloni et al., 2022; Barros et al., 2020; Hof-

stede et al., 2002) dominated by tussock grasses,

including species of Calamagrostis, Festuca, and

Stipa, as well as cushion and bog plants, such

as Azorella, Oreobolus, and Plantago (Beltrán et

al., 2009; Minga et al., 2016) and (ii) disturbed

area (Fig. 2B), defined as human-modified

habitat with signs of paths, trails, exotic plants

Fig. 1. Study area and location of 36 transects for vegetation sampling in the páramo landscape of the Macizo del Cajas

Biosphere Reserve, Southern Andes of Ecuador. The yellow squares represent the central coordinate of each 1 km transect.

The red polygons are the protected areas within the reserve (PNC = Cajas National Park. ANRQ = Quimsacocha National

Recreation Area). / Fig. 1. Área de estudio y ubicación de los 36 transectos para el muestreo de vegetación en el paisaje de

páramo de la Reserva de la Biosfera Macizo del Cajas, sur de los Andes de Ecuador. Los cuadrados amarillos representan

la coordenada central de cada transecto de 1 km. Los polígonos rojos son las áreas protegidas dentro de la reserva (PNC =

Parque Nacional Cajas. ANRQ = Área Nacional de Recreación Quimsacocha).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

(e.g., Pinus sp., Polylepis racemosa Ruiz & Pav.),

burns, evidence of livestock grazing (e.g., feces),

and eroded soils. Additionally, within each

circular plot, we established four 12 m sub-

transects, oriented along the cardinal direc-

tions. Along each sub-transect, we sampled the

vegetation by counting and identifying shrubs

(< 3 cm diameter at breast height [DBH]) and

trees (> 3 cm DBH) touched by an observer

walking with arms extended along the transect.

Data analysis: In order to evalu-

ate the adequacy of the sampling effort, a

species-accumulation curve was used based

on the total abundance of shrubs, bushes and

trees recorded across all transects (n = 36). We

used rarefaction to standardize the observed

richness (Colwell et al., 2012). The Chao 1 esti-

mator was calculated based on 1 000 random

permutations (Chao, 1984) to evaluate whether

the observed richness was representative of the

regional diversity (Colwell et al., 2012).

Community ordination: We used a non-

metric multidimensional analysis (NMDS)

to explore differences in the composition of

Fig. 2. Examples of photos of vegetation monitored in the páramo landscape of Macizo del Cajas Biosphere Reserve. We

present two examples per habitat (upper and lower panels): A. A less human disturbed páramo grassland (e.g., without

evidence of burns and grazing) located inside of Cajas National Park. B. A more human disturbed páramo grassland (e.g.,

with evidence of burns and eroded soils resulting in more presence of certain faster growing woody plants) located outside

of the national system of protected areas. All habitat examples are located at the relatively same elevation (~3 750 m.a.s.l.). /

Fig. 2. Fotos de ejemplo de la vegetación monitoreada en el paisaje de páramo de la Reserva de la Biósfera Macizo del Cajas.

Presentamos dos ejemplos por hábitat (paneles superior e inferior): A. Un páramo herbáceo menos alterado por el humano

(e.g., sin evidencia de quemas y pastoreo) localizado dentro del Parque Nacional Cajas. B. Un páramo herbáceo más alterado

por el humano (e.g., con evidencia de quemas y de suelo erosionado que resulta en una mayor presencia de ciertas plantas

leñosas de crecimiento rápido) localizado fuera del sistema nacional de áreas protegidas. Todos los hábitats de ejemplo están

localizados relativamente a la misma elevación (~3 750 m.s.n.m.).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

woody plant species (2D solution). The NMDS

used Bray-Curtis dissimilarity with the total

abundance of shrub, bushes and tree species

recorded across the 36 transects (i.e., samples).

Samples closer in the ordination represent less

dissimilarity in vegetation composition. In

order to investigate differences in the com-

munity, the average the proportion of páramo

grassland, the proportion of disturbed area,

as well as the mean elevation (meters above

the sea level) per transect were post-hoc lin-

early fitted and their significance was tested

via random permutations (1 000 iterations).

Correlations among the predictors (proportion

of páramo grassland, proportion of disturbed

area and elevation) were evaluated. We found

a negative correlation between the proportion

of disturbed area and the proportion of páramo

grassland (R = -0.77). Consequently, to obtain

an approximation of the ‘proportion of the

páramo grassland’ independent of the propor-

tion of the disturbed area, we performed a lin-

ear model on páramo grassland with disturbed

area as a predictor and used the residuals as

the predictor variable. Therefore, all predic-

tors showed no correlation (R-range = -0.28 to

0.17). All analyses were carried out in R 4.4.1

(R Core Team, 2024). For estimators and spe-

cies accumulation curves we used the ‘iNext’

package (Chao et al., 2024), while for com-

munity ordination we used the ‘vegan’ package

(Oksanen et al., 2024).

RESULTS

In total, we recorded 13 377 woody plants.

The records are grouped in 29 species, 17

genera and 10 families (Table 1). The most

Table 1

Total abundance and codes of woody plant species recorded across 36 transects located in the páramo landscape of the

Macizo del Cajas Biosphere Reserve, Southern Andes of Ecuador / Tabla 1. Abundancia total y códigos de las especies leñosas

registradas a través de 36 transectos ubicados en el paisaje de páramo de la Reserva de la Biosfera Macizo del Cajas, sur de

los Andes de Ecuador.

Family Scientific name Species code Abundance

Asteraceae Baccharis tricuneata (L. f.) Pers. BATR 476

Chuquiraga jussieui J.F. Gmel. CHJU 660

Diplostephium oblanceolatum S.F. Blake DIOB 402

Diplostephium rupestre (Kunth) Wedd. DIRU 900

Diplostephium ericoides (Lam.) Cabrera DIER 178

Diplostephium glandulosum Hieron. DIGL 45

Gynoxys miniphylla Cuatrec. GYMI 789

Gynoxys cuicochensis Cuatrec. GYCU 188

Gynoxys baccharoides (Kunth) Cass. GYBA 19

Loricaria thuyoides (Lam.) Sch. Bip. LOTH 1 420

Monticalia arbutifolia (Kunth) C. Jeffrey MOAR 1 637

Monticalia vaccinioides (Kunth) C. Jeffrey MO VA 864

Monticalia andicola (Turcz.) C. Jeffrey MOAN 76

Monticalia empetroides (Cuatrec.) C. Jeffrey MOEM 137

Berberidaceae Berberis lutea Ruiz & Pav. BELU 101

Berberis rigida Hieron. BERI 43

Caprifoliaceae Valeriana microphylla Kunth VAMI 753

Valeriana hirtella Kunth VAHI 64

Ericaceae Pernettya prostrata (Cav.) Sleumer PEPR 41

Grossulariaceae Ribes lehmannii Jancz. RILE 73

Hypericaceae Hypericum aciculare Kunth HYAC 2 964

Hypericum quitense R. Keller HYQU 221

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

abundant species was Hypericum aciculare

Kunth (Hypericaceae) (22 %) followed by M.

arbutifolia (Asteraceae) (12 %), Loricaria thuy-

oides (Lam.) Sch. Bip. (Asteraceae) (11 %),

Diplostephium rupestre (Kunth) Wedd. (Astera-

ceae) (7 %) and Monticalia vaccinioides (Kunth)

C. Jeffrey (Asteraceae) (6 %) (Table 1). The

proportion of páramo grassland ranged from

30 % to 88 % (mean = 63 %) and the propor-

tion of disturbed areas ranged from 0 % to 45 %

(mean = 20 %) (Table 2).

The species accumulation curve reached

its asymptote (Fig. 3), indicating that the

sampling effort was sufficient and therefore,

Family Scientific name Species code Abundance

Melastomataceae Brachyotum jamesonii Triana BRJA 767

Miconia salicifolia Bonpl. ex Naudin MISA 267

Polygalaceae Monnina crassifolia (Bonpl.) Kunth MOCR 46

Rosaceae Hesperomeles obtusifolia (Pers.) Lindl. HEOB 21

Polylepis reticulata Hieron. PORE 111

Polylepis incana Kunth POIN 56

Rubiaceae Arcytophyllum vernicosum Standl. ARVE 58

Table 2

Locality name, transect code, average proportions (see methods) of páramo grassland as well as disturbed area and mean

elevation of the 36 transects located in the páramo landscape of the Macizo del Cajas Biosphere Reserve, Southern Andes of

Ecuador / Tabla 2. Nombre de la localidad, código de los transectos, el promedio de las proporciones (véase los métodos) de

páramo herbáceo así también de área alterada y la elevación media en los 36 transectos ubicados en el paisaje de páramo de

la Reserva de la Biósfera Macizo del Cajas, sur de los Andes de Ecuador.

Locality Transect code Páramo grassland (%) Disturbed area (%) Elevation (m)

Bermejos BER001 80.5 12.3 3 794

Bermejos BER002 74.4 14.05 3 842

Bermejos BER003 75.31 14.9 3 795

Burgay BUR001 63.3 17.9 3 627

Burgay BUR002 67.6 25.1 3 818

Burgay BUR003 62.2 20.4 3 754

Dublaicocha DUB001 30.05 27.85 3 840

Dublaicocha DUB002 38.6 30.25 3 876

Dublaicocha DUB003 46.35 30.75 3 842

Galgal GAL001 53.8 36.7 3 767

Galgal GAL002 44.6 44.95 3 730

Galgal GAL003 42.25 46 3 684

Miguir MIG001 30.43 30.67 3 803

Miguir MIG002 66.1 9.3 4 049

Miguir MIG003 64.8 13.02 3 929

Patococha PAT001 58.35 28.85 3 910

Patococha PAT002 66.3 18.35 3 803

Patococha PAT003 61.5 25.3 3 812

Rircay RIR001 67.7 11.4 3 759

Rircay RIR002 75.35 9.73 3 782

Rircay RIR003 74.4 11.3 3 685

Santa Ana SAN001 65.25 24.85 3 787

Santa Ana SAN002 62.6 13.9 3 704

Santa Ana SAN003 62.2 31.2 3 797

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representative of the richness within the study

area. Furthermore, the observed richness was

the same as the estimated richness (observed

richness = 29; Chao 1 estimator = 29).

Community composition of woody plant

species: The NMDS analysis showed a ten-

dency to separate the woody plant community

in relation to the proportion of disturbed area

(2D solution, stress = 0.2, R2 = 0.33, p < 0.001)

(Fig. 4), while the proportion of páramo grass-

land (R2 = 0.07, p =0.32) and elevation (R2 =

0.10, p < 0.16) were not significant. Transects

with a higher proportion of disturbed area were

situated towards the central-bottom of the ordi-

nation, and were associated with a greater pres-

ence of D. ericoides, H. quitense, V. microphylla

and Valeriana hirtella Kunth., and transects

with a lower proportion of disturbed area were

located towards the central-top of the plot, and

were associated with a greater presence of M.

arbutifolia, G. miniphylla, M. vaccinioides and,

Hesperomeles obtusifolia (Pers.) Lindl. (Fig. 4).

However, we included elevation in the

ordination plot (via contour lines) as it fol-

lowed the same tendency as disturbed area

Locality Transect code Paramo grassland (%) Disturbed area (%) Elevation (m)

Taitachugo TAI001 62.5 13.1 4 032

Taitachugo TAI002 57.9 16.2 3 941

Taitachugo TAI003 81.8 9.3 3 919

Tarqui TAR001 84.6 11.75 3 998

Tarqui TAR002 60.6 17.1 3 646

Tarqui TAR003 82.9 0 3 789

Tomebamba TOM001 71.78 10.5 3 809

Tomebamba TOM002 87.83 2.8 3 909

Tomebamba TOM003 50.25 23.7 3 733

Ventanas VEN001 53.3 25.35 3 841

Ventanas VEN002 62.85 7.74 3 865

Ventanas VEN003 63.75 23.5 3 898

Fig. 3. Species-accumulation curve of woody plant species recorded in 36 transects located in the páramo landscape of the

Macizo de Cajas Biosphere Reserve, Southern Andes of Ecuador. The solid line represents the accumulation of observed

species via rarefaction. The dashed line shows the Chao 1 estimation. / Fig. 3. Curva de acumulación de especies de plantas

leñosas registradas en 36 transectos ubicados en el paisaje de páramo de la Reserva de la Biósfera Macizo de Cajas, sur de

los Andes de Ecuador. La línea continua representa la acumulación de especies observadas a través de rarefacción. La línea

discontinua muestra la estimación Chao 1.

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(i.e., from the central-top to the central-bottom

of the ordination) (Fig. 4). This facilitates the

interpretation of the spatial distribution of

the transects and the description of woody

plant species associations in relation to eleva-

tion. Thus, from ~3 800 m to ~4 000 m (in the

central-top of the ordination) there is greater

prevalence of Diplostephium oblanceolatum M.

arbutifolia, M. vaccinioides, H. obtusifolia and

Monticalia andicola (Turcz.) C. Jeffrey., while

at lower elevations (i.e., from ~3 600 to 3 800

m) (in the central-bottom of the ordination)

the woody plant community is associated with

more presence of D. ericoides, V. microphylla, H.

quitense and V. hirtella (Fig. 4).

DISCUSSION

Our findings showed significant changes in

the composition of the woody plant community

along the increasing proportion of disturbed

areas in the monitored transects. The propor-

tion of páramo grassland and elevation did not

affect the plant composition. Dense shrub spe-

cies such as V. microphylla, H. quitense and V.

hirtella were more dominant in transects with

Fig. 4. Non-metric multidimensional scaling (NMDS) of the community of woody plant species recorded across 36 transects

(green circles) located in the páramo landscape of the Macizo del Cajas Biosphere Reserve, Southern Andes of Ecuador.

The red arrow represents the proportion of disturbed area (DA) as the environmental variable significantly influencing the

ordination (p < 0.001). The elevation (contour lines) is also shown, although it is not significant (p > 0.05). The four-letter

codes are the scientific names of the woody plant species (see Table 1). / Fig. 4. Escalamiento multidimensonal no métrico

(NMDS) para la comunidad de plantas leñosas registradas a través de 36 transectos (círculos verdes) ubicados en el paisaje

de páramo de la Reserva de la Biósfera del Macizo del Cajas, sur de los Andes del Ecuador. La flecha roja representa la

proporción del área alterada (DA) como una variable ambiental que influye significativamente en la ordenación (p < 0.001).

La elevación (curvas de nivel) también se muestra, aunque no influye significativamente (p > 0.05). Los códigos de cuatro

letras son los nombres científicos de las especies de plantas leñosas (véase Tabla 1).

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

higher proportions of disturbed areas. Smaller

shrub species such as M. arbutifolia, M. vaccini-

oides and H. obtusifolia are associated with tran-

sects with less disturbance. These changes in

the plant composition of the páramo ecosystem

suggests that the greater prevalence of some

dense and fast-growing woody plant species

is a response to human disturbance and could

indicate that the native herbaceous species are

gradually replaced by woody encroachment,

particularly in human-accessible páramos.

Our approximation of the distribution of

the woody plant community reveals that dense

shrubs (e.g., H. quitense) are frequently observed

in groups characterized as dwarf scrub and are

adapted to rocky outcrops and hills where the

evidence of human disturbance is higher. In

fact, some Hypericum species are reported as

opportunistic species that recover quickly after

human disturbance such as fires (Matson &

Bart, 2014). Their presence here may be related

to an increasing cover of shrubs and a reduc-

tion in the cover of native grasses (e.g., Renison

et al., 2006) due to human disturbance reduc-

ing the dominance of the tussock grasses and

providing more opportunity for fast-growing

woody plants (Sylvester et al., 2017).

In general, across the small valleys in the

study region, páramo grassland is often asso-

ciated with shrubby plants (Hofstede et al.,

2002; Minga et al., 2016) creating a botanically

and structurally diverse páramo (Gareca et al.,

2010; Hofstede et al., 2002). However, woody

encroachment is a common effect in disturbed

grassland systems, including high elevation

grasslands such as páramo (Matson & Bart,

2014). Our results, controlling for the propor-

tion of the disturbed area, highlight that some

transects are still characterized by a greater

presence of woody plants such as H. quitense,

V. hirtella and V. microphylla. Besides, more

human accessible páramos (e.g., close to roads,

lower elevations, outside of protected areas) are

more prone to habitat disturbance such as fires

and livestock grazing leading to less availability

of native tussock grasses as well as cushion and

bog plants. Consequently, in unprotected pára-

mos of the study region, the native grasses are

more sensitive to human disturbance (Astudillo

et al., 2017; Hofstede & Llambí, 2020). Here,

our findings suggest that fast-growing woody

plant species are fostering changes in the plant

community leading to more complex páramos

(e.g., height profile vegetation) but with less

availability of open habitats such as tussock

grasses and cushion bogs.

A potential explanation for the pattern of

woody plant species dispersal into the páramo

grassland is the proximity to the tree line of

Andean forests (Matson & Bart, 2013) across

the lower páramo. In the study region, large

and dense shrubs such as V. hirtella and V.

microphylla are commonly found in the tree

line of Andean forests and Polylepis woodlands

(Minga et al., 2016). The frequent association

of forest plants with herbaceous flora of pára-

mo ecosystems has been reported previously

(Domic & Capriles, 2021; Montalvo et al., 2018;

Quispe-Melgar et al., 2020) and indeed, some

native shrubs have been found to be positively

associated with greater habitat complexity of

páramos surrounding Polylepis forest within

the Cajas National Park (Astudillo et al., 2019;

Astudillo et al., 2020). However, as with other

disturbance scenarios found outside the limit

of protected areas (Caballero-Villalobos et al.,

2021; Toivonen et al., 2011), our results high-

light that some woody plant species are becom-

ing more dominant. Within this framework,

conservation and restoration efforts across the

páramo should consider the dynamics between

shrubs and other fast-growing plants with

native grasses.

Conservation implications: In the high

Andes, the restoration of disturbed habitats

has been based on the importance of woody

plant species for important pollinators (i.e.,

hummingbirds) (Cárdenas-Calle et al., 2020;

Crespo et al., 2022; Hazlehurst et al., 2016) with

a few shrubby plant species being identified as

central for the restoration of high Andean eco-

systems (Crespo et al., 2022). However, some of

these identified woody plant species are char-

acterized by faster growing strategies (i.e., year-

round flowering) (e.g., Cárdenas-Calle et al.,

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61916, enero-diciembre 2025 (Publicado Jun. 10, 2025)

2020; Hazlehurst et al., 2016) and are already

widely distributed between the limit of Andean

forests and the páramo ecosystem (Minga et al.,

2016; Suárez et al., 2022). Furthermore, mutual-

istic interactions in disturbed páramos, such as

seed dispersal by birds, are seen as a mechanism

that facilitates increasing cover of shrubby plant

species and subsequent woody encroachment

(Matson & Bart, 2013; Matson & Bart, 2014).

In consequence, habitat restoration focused on

fast-growing plants may promote an accelera-

tion of woody encroachment in human-acces-

sible páramos with evident negative effects on

native grasses and organisms that depend on

more open páramo grassland habitats. Further

studies are needed in order to generate conser-

vation strategies more compatible to the pára-

mo region. For instance, in open páramos of

the study region, small, endemic rodents (e.g.,

Phyllotis haggardi) have been observed foraging

on flowers of herbaceous species (i.e., Taraxa-

cum officinale (Weber), Xenophyllum humile

(Kunth) V.A. Funk and Eryngium humile Cav.)

and are considered as potential pollinators

(Nivelo-Villavicencio et al., 2021). Our findings

indicate that the availability of páramo grass-

land habitats is declining in disturbed areas,

while the presence of fast-growing shrubs is

increasing. This ecological information could

be crucial for improving restoration and con-

servation efforts aimed at enhancing the avail-

ability of páramo grasslands.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

We are grateful for the constant sup-

port of our research by Raffaella Ansaloni,

Andrés López and Francisco Salgado of the

Universidad del Azuay. We thank Boris Landá-

zuri for his field support. We also acknowl-

edge Guillermo Salgado, Vicente Jaramillo, and

Ramiro Jiménez from Dundee Precious Met-

als Ecuador (DPMECUADOR S.A.) for their

continued support of our research. Logistical

support came from Hari González, park man-

agement, and staff of Cajas National Park and

Quimsacocha National Recreation Area, as well

as Empresa Pública de Telecomunicaciones,

Agua Potable, and Alcantarillado y Saneamien-

to de Cuenca (ETAPA-EP). PXA was funded

by DPMECUADOR S.A., ETAPA-EP (Grant

No. 78010200001) and the Vicerrectorado de

Investigaciones of the Universidad del Azuay

(2018-0003; 2019-0090; 2021-0167). All activi-

ties were conducted under permits 196-2019-

DPAA/MA and 183-2018-DPAA/MA issued

by the Ecuadorian Ministry of Environment,

Water and Ecological Transition.

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