vegetation of a hill gRassland of the Paititi natuRal

ReseRve (PamPa biome) and eaRly detection of non-

native sPecies acting as invasive

vegetación de un Pastizal seRRano de la ReseRva natuRal Paititi (bioma

PamPa)y detección temPRana de esPecies no nativas actuando como invasoRas

1

2

& Viviana M. Comparatore

María L. Echeverría * , Sara I. Alonso

Resumen

Introducciónyobjetivos:Lasespeciesinvasorasrepresentanlamayoramenazapara

la conservación. Los objetivos de este estudio fueron identiﬁcar la ﬂora del pastizal

serrano de la Reserva Natural Paititi, realizar la detección temprana de especies

vegetales exóticas que pudieran estar actuando como invasoras y establecer la

situación actual de aquellas especies nativas consideradas amenazadas.

1

. Universidad Nacional de Mar del

Plata, Facultad de Ciencias Agrarias,

Ruta 226, km 73.5, Balcarce,

Buenos Aires, Argentina

2

. Universidad Nacional de Mar del

M&M: Siguiendo el gradiente altitudinal se seleccionaron 14 sitios donde se registraron

Plata, Facultad de Ciencias Exactas

y Naturales, Funes 3250, Mar del

Plata, Buenos Aires, Argentina.

Instituto de Investigaciones

las características ambientales

y la cobertura-abundancia de las especies

vasculares. Con los datos recabados se realizó un Análisis de Coordenadas

Principales (ACooP) que originó unidades de vegetación que fueron descritas

según características ambientales, especies identiﬁcadas y tipo de comunidad

vegetal. Adicionalmente, se destacaron las especies amenazadas y las exóticas

más abundantes, en consecuencia consideradas invasoras.

Marinas

CONICET

y Costeras (IIMyC),

Resultados: La riqueza total fue de 370 especies, correspondiendo el 26,5% a

exóticas. Se identiﬁcaron 30 especies amenazadas. El ACooP reconoció ocho

unidades de vegetación. Las especies consideradas invasoras fueron Dactylis

glomerata, Senecio madagascariensis, Holcus lanatus y Racosperma melanoxylon,

estas dos últimas con los mayores valores de cobertura.

Conclusiones: Para conservar la biodiversidad y minimizar el proceso de invasión, los

esfuerzos de manejo deberían enfocarse en monitorear las especies amenazadas y

controlar el avance de las invasoras. La detección temprana de dichas especies en

ambientes similares sería fundamental para reducir invasiones vegetales.

*

mlecheverria@mdp.edu.ar

Citar este artículo

ECHEVERRÍA, M. L., S. I. ALONSO

V. M. COMPARATORE. 2023.

&

Vegetation of a hill grassland of

the Paititi Natural Reserve (Pampa

biome) and early detection of non-

native species acting as invasive.

Bol. Soc. Argent. Bot. 58: 71-90.

PalabRas clave

DOI: https://doi.

org/10.31055/1851.2372.v58.

n1.38512

Especies exóticas, pastizales serranos, áreas protegidas, especies amenazadas.

summaRy

Background and aims: Invasive species are the greatest threat to conservation. The

objectives of this study were to identify the ﬂora that thrive in a hill grassland of the

Paititi Natural Reserve, to perform an early detection of non-native plant species

that might be acting as invasive, and to establish the current situation of the native

species considered threatened.

M&M: Following the altitudinal gradient, 14 sites were selected; the environmental

characteristics and cover-abundance of the vascular plant species were recorded

in each site. With the collected data, a Principal Coordinate Analysis (PCooA)

was performed to group the sites into vegetation units that were later described

considering environmental characteristics, identiﬁed species and plant community

type. Additionally, the threatened species were speciﬁed, as well as the most

abundant exotic ones, therefore considered invasive.

Results: Total richness reached 370 species, 26.5% corresponding to non-native

ones. Thirty threatened species were identiﬁed. The PCooA grouped the sites

into eight vegetation units. The non-native species considered invasive were

Dactylis glomerata, Senecio madagascariensis, Holcus lanatus and Racosperma

melanoxylon, these last two reached the highest coverage-abundance values.

Conclusions: To conserve biodiversity and minimize the invasion process,

management eﬀorts should be focused on monitoring the threatened species and

controlling the advance of the non-native species acting as invasive. Early detection

of those species in similar environments would be fundamental to facilitate rapid

responses towards reducing invasions.

Recibido: 13 Ago 2022

Aceptado: 22 Nov 2022

Publicado en línea: 17 Feb 2023

Publicado impreso: 31 Mar 2023

Editor: Juan Carlos Moreno Saiz

Key woRds

ISSN versión impresa 0373-580X

ISSN versión on-line 1851-2372

Exotic species, mountain grasslands, protected areas, threatened species.

71

Bol. Soc. Argent. Bot. 58 (1) 2023

intRoduction

& Cock, 2001; Waterhouse, 2003; Brancatelli et al.,

020).

The Pampa biome, an extensive South American

2

Biological invasions have been recognized

in many parts of the world as the main agents grassland region located in Uruguay, southern

of change in natural environments, generating Brazil, and central Argentina (Scottá & da Fonseca,

harmful eﬀects on ecosystems and biodiversity and 2015) (Fig. 1A), has a high degree of landscape

imposing an enormous cost on human productive fragmentation. It was formerly characterized by

activities (Wittenberg & Cock, 2001; Pyšek & grasslands and, to a lesser extent, shrubs (Cabrera,

Richardson, 2010; van Kleunen et al., 2018). The 1968; Cabrera & Zardini, 1978). However, the

invasive plants are the non-native plant species advance of the agricultural frontier has generated

–

also called introduced, alien, or exotic species– a homogeneous matrix of mainly agricultural lands

that after their introduction into non-native natural interspersed with areas of natural or semi-natural

environments, may spread, invading the new vegetation, generally not cultivable (Herrera et

territory and generating a high impact on the al., 2020). As a result, this region faces a serious

new ecological niches (Bentivegna & Zalba, and growing challenge associated with an increase

2

014). Nevertheless, not all non-native species in invasive non-native species (Frangi, 1975;

become invasive, and the ones that do, are invasive Bilenca & Miñarro 2004; Echeverría et al., 2017;

only in some places, since the term invasive is Brancatelli & Zalba, 2018).

associated with speciﬁc environmental conditions

Toward the south of the Pampa biome, the

(Weber & Gut, 2004; van Kleunen et al., 2018). Tandilia Hill System in Buenos Aires Province

However, species with a history of invasiveness (Argentina) (Fig. 1A-B) is a discontinuous chain

tend to repeat this behavior in the places where of low hills (called sierras in Spanish) and rocky

they are introduced (Baskin, 2002). According outcrops. The Tandilia Hill System is approximately

to van Kleunen et al. (2018), the deﬁnition of an 350 km in length × 60 km in maximum width and

invasive species implicitly assumes that the non- less than 600 m elevation, extending NW to SE

native species should be locally abundant (i.e. (Dalla Salda et al., 2006). In these low mountainous

produces oﬀspring in large numbers). Therefore, by areas, the development of agricultural activities

increasing their abundance in the near future, non- is diﬃcult, which facilitates the conservation of

native species can become invasive.

pristine vegetation remnants known as mountain

Protected areas are a key component in grasslands (Frangi, 1975; Alonso et al., 2009a;

responding to degradation and environmental Herrera et al., 2017; Kristensen et al., 2014;

changes. However, invasive plant species are a Echeverría et al., 2017; Brancatelli et al., 2020).

serious problem for protected area managers around These areas are considered biodiversity “hotspots”

the world (Foxcroft et al., 2017; Brancatelli et al., because they harbor numerous native and endemic

2

020). The management of these areas regarding species, some of which are threatened (Delucchi,

invasive species involves diﬀerent strategies: A) 2006; Herrera & Laterra, 2011; Kristensen et al.,

prevention and exclusion; B) early detection and 2014; Echeverría et al., 2017). Threatened species

rapid assessment; C) control, containment, and are considered priority species to preserve, and

eradication (Rejmánek, 2000). Although prevention therefore a fundamental pillar to guide biodiversity

is the most eﬃcient control strategy, on certain conservation and sustainable development policies

occasions, invasive species are already established (IUCN, 2022). At the same time, native species

in the territory (Anderson et al., 2014; Brancatelli are a very important source of ecosystem services

&

Zalba, 2018). Since the plants are usually (Barral & Maceira, 2012; Herrera et al., 2017). The

distributed in patches or aggregates according to the Strict Nature Reserve of the Paititi Private Natural

environmental characteristics, carrying out ﬂoristic Reserve (PNR) is the largest nature reserve of the

surveys in diﬀerent sites of the area of interest Tandilia Hill System. This reserve is considered

is essential for the early recognition of current a Valuable Grassland Area (Bilenca & Miñarro,

or potential invasions and the establishment of 2004) and an Area of Interest for Conservation

management methodologies aimed at minimizing and Ecotourism (Chebez, 2005). The wide

the process of invasion in early stages (Wittenberg environmental heterogeneity present in the PNR has

72

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

Fig. 1. A: Map of South America (light gray), with delimitation of the Pampa biome (dark gray), modiﬁed from

Scottá and Da Fonseca (2015), and of Buenos Aires Province (dotted line); B: Buenos Aires Province with

delimitation of the Tandilia Hill System and Paititi Natural Reserve (PNR); C: Delimitation of the Strict Nature

Reserve area of the PNR (dotted line). Each marker shows one of the sites (S) selected for the study; D:

Topographic proﬁle reﬂecting the altitudinal gradient of the study area and the location of the analyzed sites.

led to the development of a great diversity of native

Considering the value of the PNR as a refuge for

fauna (Cicchino & Farina, 2007; Isacch et al., 2016; the ﬂora and fauna of the grasslands and hill areas

Arcusa, 2016; Ferreti et al., 2019; O’Connor et al., of the southeastern extreme of the Tandilia Hill

2

020). Regarding the plant species, Echeverría et System and the possible threat posed by invasive

al. (2017) carried out a ﬂoristic survey in the PNR plants to biological diversity, it is important to

and identiﬁed 360 species, more than 25% being know the distribution and representativeness of

non-native species. These authors alerted that some the plant species that can be found in the reserve.

of those species may become invasive and thus Therefore, the objectives of this work were: 1)

contribute to the deterioration of the natural areas to identify all the vascular plants (native and

of the reserve.

non-native ones) that thrive in sites with diﬀerent

73

Bol. Soc. Argent. Bot. 58 (1) 2023

environmental characteristics in the PNR; 2) to between 32 °C and below 0 °C, with a mean annual

quantify their abundance in order to carry out the temperature of 14 °C. Rainfall is usually distributed

early detection of non-native species that might throughout the year with an average of 850 mm per

be acting as invasive ones; 3) to establish the year (Falasca et al., 2000; INTA, 2022).

current situation of those native species that were

considered threatened in Buenos Aires Province.

Sampling and data collection

Following the altitudinal gradient, 14 sites were

selected. These sites varied in the physiognomy of

the vegetation (grasses, shrubs, or trees), dominant

species, and predominant substrate (soil, rock, or

water). The sites were georeferenced using a global

mateRials and methods

Study area

The study was conducted in the hill grassland positioning system (GPS) and digitized on an

ecosystem of the PNR located in the southeastern aerial image of the study area (Fig. 1C-D). A plot

Tandilia Hill System, in the southeast ﬂank of was delimited at each site based on the concept of

2

Sierra de Difuntos (37° 54′ S - 57° 49′ W; geodetic minimum area, with plot size ranging from 4 m to

2

datum WGS84), belonging to the orographic group 200 m (Matteucci & Colma, 2002; Escaray, 2007).

of Sierras de Mar del Plata (Guazzelli, 1999), The soil type was identiﬁed. Additionally, electrical

in General Pueyrredon district, Buenos Aires conductivity, pH, organic matter content, and

Province, Argentina (Fig. 1B). The main vegetation available phosphorus were determined in ten soil

landscape is a large open area covered with grasses subsamples of the ﬁrst 10 cm of the proﬁle by the

(

grasslands), although there are also oreophilic Soil Analysis Laboratory of Balcarce Agricultural

steppes, scrublands, and hydrophilic communities Experimental Station belonging to the Instituto

Cabrera, 1968; Cabrera & Zardini, 1978). In Nacional de Tecnología Agropecuaria.

particular, in the mountainous areas, different The ﬂoristic composition and the speciﬁc cover-

(

types of grassland and other plant communities can abundance in each plot were determined from

thrive, such as the shrubs and scrubs that are typical september 2018 to december 2019. To estimate

of mountain and rocky soils (Frangi, 1975).

species coverage, the Braun-Blanquet cover-

The PNR is divided into two areas: one dedicated abundance scale modiﬁed by Westhoﬀ and Maarel

to educational, recreational, and livestock activities, (1978) was used.

and the other one categorized as a Strict Nature

The taxa surveyed were determined at the

Reserve so it remains as an area with minimal speciﬁc level based on the following ﬂora book

anthropic interference. The study was carried out collections: Flora of the Province of Buenos Aires

in the Strict Nature Reserve, where there is a hill (Cabrera 1963, 1965a, 1965b, 1967, 1968 and

area called Sierra Chica and a stream surrounding 1970) and Flora Argentina (Zuloaga et al., 2012a,

the base of the west slope of that hill, forming a b; Zuloaga et al., 2014a, b). For each species,

small pond before continuing its N-E path (Fig. botanical family, origin (non-native, native, or

1

C). Sierra Chica is oriented N-S, with a maximum cosmopolitan), and threatened situation in Buenos

elevation of 156 m.a.s.l. in the rocky area at the Aires Province (Delucchi, 2006) were determined.

top, and a minimum of 86 m.a.s.l. in the pond. The

hill blocks are mainly composed of a crystalline Analysis of data

basement on which Eopaleozoic sediments were

Total richness (number of species), as well as

deposited. This sediment accumulation gave rise to richness per site, representativeness of botanical

the development of Mollisol soils of variable depth, families, and distribution of species by category

loamy texture, high content of organic matter, and of origin were determined. Correlations between

slightly acidic pH (Osterrieth & Cabria, 1995; Dalla site altitude and percentage of species according to

Salda et al., 2006; Álvarez et al., 2012). The climate origin (native versus non-native + cosmopolitan)

of the region is humid-subhumid, mesothermal, were calculated using the Pearson correlation (r)

with low water deﬁciency, noticeable seasonal test (α = 0.1%) through the “cor.test” function of

variation in temperature, and a short cold period. It the “corrplot” package of RStudio software Version

is also characterized by maritime temperate ranging 1.2.5033 (R Core Team, 2014).

74

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

To associate the sites and generate vegetation

The vascular species richness in the plots varied

units with similar characteristics considering their from 6 to 161 species. In the highest sites such

environmental attributes and the species at each as the hillside and summit areas, the percentage

site, the qualitative and quantitative variables were of native plants was over 70%, except for site ten

jointly analyzed by a Principal Coordinate Analysis (S10) (Fig. 2). This trend was conﬁrmed by the

(PCooA) using Gower similarity coeﬃcient (Gower, correlation between altitude and percentage of

1

971; Cuadras, 2014). The analysis was carried out native species, which was signiﬁcant and positive

in the RStudio software Version 1.2.5033 (R Core (r = 0.67; p-value = 0.00829); consequently, it was

Team, 2014) using the “cluster” package (Maechler also signiﬁcant, but negative, between altitude and

et al., 2016).

percentage of non-native + cosmopolitan species.

The vegetation units were described considering

the environmental characteristics, the identiﬁed Relationship between sites

ﬂora and the type of plant community according

The first three coordinates of the PCooA

to Frangi (1975). Additionally, the threatened contribute explaining 45.82, 15.57, and 10.02%

species were speciﬁed, as well as the most abundant of the total variability, respectively. The projection

non-native ones (≥ 5% cover-abundance values), of the study sites in the three-dimensional space

therefore considered invasive.

showed the formation of eight clusters, which

represent the vegetation units (Fig. 3).

Results

Characteristics of the vegetation units

The soils of the units were dark to very dark

In the study area, the total richness reached brown and slightly acid (pH= 5.3–5.8); the

370 vascular species, 72.5% being native, 26.5% superﬁcial horizons had high root density, low

non-native and 1% cosmopolitan (Appendix S1); electrical conductivity (0.3–0.8 mmhos), low to

ten taxa, Acmella decumbens (Sm.) R.K. Jansen, moderate phosphorus content (3.4–7.6 ppm), and a

Aira caryophyllea L., Anthoxanthum odoratum L., high percentage of organic matter (6.3–11.4%). The

Eryngium ebracteatum Lam., Galium richardianum greatest abiotic diﬀerences were established in the

(

GilliesexHook.&Arn.)Hicken,Nassellatrichotoma type and depth of the soil, the slope (Table 1), the

Nees) Hack, Parodia submammulosa (Lem) R. degree of solar exposure, and the relief position. For

Kriesli, Plantago lanceolata L., Tripogonella spicata information on the species of each vegetation unit,

Nees) P.M. Peterson & Romasch., and Wigginsia consult the Appendix S1.

tephracantha (Link & Otto) D.M. Porter, had not Base Grassland: This unit corresponds to an

(

been previously detected by Echeverría et al. (2017) area located at the base of the hill (Fig. 4A; Table

at the study area. The surveyed species belong to 1). The species richness reached 114 vascular

6

9 families, the most representative ones being plants, 42% being non-native ones. The area was

Poaceae (78), Asteraceae (72), Fabaceae (21), physiognomically dominated by taxa of the Poaceae

Cyperaceae (13), Apiaceae (12), Solanaceae (11), family, of the genera Aristida L., Nassella E.

and Brassicaceae (10) (Appendix S1). Some taxa Desv., Jarava Ruiz. & Pav., and Piptochaetium J.

were found only in one site and with very low cover Presl. Floristically, the Base Grassland resembles

values, such as Pterocaulon cordobense Kuntze, the typical “Flechillar” grassland community of

Hypericum connatum Lam., and Lemna gibba L., the Tandil hills dominated by grasses of stabbing

while others were found in most of the sites and in caryopses, which look like small arrows (Flechillar

some of them with more than 50% coverage, such as derives from the Spanish word ‘ﬂecha’, which means

Racosperma melanoxylon (R. Br.) Mart. The greatest arrow in English). Six of the species mentioned as

habitat variation was presented by four species, threatened in Buenos Aires Province were found,

which were found in nine of the fourteen sites, two although only two (Acanthostyles buniifolius (Hook.

non-native species, Cirsium vulgare (Savi) Ten, and & Arn.) R. M. King & H. Rob. and Senecio selloi

Senecio madagascariensis Poir., and two native (Spreng.) DC.) had cover-abundance levels greater

ones, Cypella herbertii (Lindl.) Herb. and Vicia nana than 5% (Table 2). Among the non-native species,

Vogel.

Holcus lanatus L. (Fig. 5A) had the highest level of

75

Bol. Soc. Argent. Bot. 58 (1) 2023

Fig. 2. Vascular species richness in 14 sites of the Strict Nature Reserve area of the Paititi Natural Reserve

(Buenos Aires, Argentina) according to their origin category (native, non-native, cosmopolitan).

Fig. 3. Principal Coordinates Analysis using combined data of presence/absence of vascular species and

environmental characteristics of 14 sites (S) in the Strict Nature Reserve area of the Paititi Natural Reserve

(Buenos Aires, Argentina). The clusters (vegetation units) are indicated by dashed lines on the three ﬁrst

principal coordinates.

76

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

Table 1. Slope, soil type and soil depth of the vegetation units of the Strict Nature Reserve area of the

Paititi Natural Reserve (Buenos Aires, Argentina).

Slope

%)

Soil Depth

(cm)

Vegetation Unit

Base Grassland (S1)

Soil Type

(

7 – 9

< 2

Typic Argiudoll

Typic Argiacuoll

> 100

Pond (S2)

Mixed Grove (S3)

< 2

Typic Argiudoll

Aquic Argiudolls

Typic Argiudoll

Hapludoll lytic

Typic Argiudoll

> 100

50 – 100

10 – 80

1 – 40

1 – 20

> 100

Humid Mountain Grassland (S4, S13)

Dry Mountain Grassland (S5, S8, S14)

Rocky Place (S6, S7, S11)

Spring (S9, S12)

2 – 45

< 2

Acacia Groove (S10)

3

cover-abundance (15-25%), followed by Dactylis rock blocks at the top of the hill (Fig. 4D; Table

glomerata L. (Fig. 5B) (5-15%). 1). In this unit, 81 species were identiﬁed being

Pond: This unit is located in the lowest part of 34.5% non-native and 1.2% cosmopolitan. The

the study area, surrounding a shallow pond (Fig. flora corresponds to the grassland community

4

3

B; Table 1). Thirty-ﬁve species were found being named “Pajonal (tall grasses) of tussock paspalum”,

1.4% non-native and 5.7% cosmopolitan. This unit as it is mainly composed of herbaceous plants

has some ﬂoating and marsh plants that are typical but physiognomically dominated by Paspalum

of the wetlands and watercourses of Buenos Aires quadrifarium Lam (“tussock paspalum”). In fact,

Province (Appendix S1). In this vegetation unit, this species showed high cover-abundance levels

one native species was categorized as threatened (50-75%). Three threatened species were found

(

Cypella herbertii ssp. wolﬀhuegeli (Lindl.) Herb. in this vegetation unit, all with very low cover-

Hauman) Ravenna) and with very low cover- abundance levels (Table 2). Additionally, Holcus

abundance level (Table 2). The non-native species lanatus was the non-native species with the highest

with the highest cover-abundance level was Senecio level of cover-abundance (5-15%).

madagascariensis Poir (5-15%) (Fig. 5C).

Dry Mountain Grassland: This unit is found on

Mixed Grove: This unit corresponds to an very steep slopes and at the top of the hill (Fig. 4E;

implanted grove following the eastern course of the Table 1). The total richness reached 156 species:

stream. It is located at the base of the hill (Fig. 4C; 25% non-native and 0.6% cosmopolitan. The

Table 1). The total richness resulted in 50 vascular vegetation was mainly represented by “ﬂechillas”

species, and 36% of the species were non-native and dicot species, with less than 20% of tree and

ones. The vegetation consisted mainly of perennial shrub species. The ﬂoristic composition is similar

species dominated by phanerophytes (34%), Celtis to that found on shrubland, with physiognomic

tala Gillies ex Planch being the most abundant dominance of Acanthostyles buniifolium, Baccharis

species. The trees were accompanied by shorter coridifolia DC., B. dracunculifolia ssp. tandilensis

herbaceous plants adapted to the canopy shade. (Speg.) Giuliano and Colletia paradoxa, and

Four threatened species were found in this unit, in a lower stratum, the ﬂora varied according

whose coverage was very low, except for Colletia to the position on the ground. The non-native

paradoxa (Spreng.) Escal. with 5% (Table 2). No species with the highest level of cover-abundance

exotic species exceeded 5% cover.

were Racosperma melanoxylon (5-15%), Dactylis

Humid Mountain Grassland: This type of glomerata (5-15%), and Holcus lanatus (15-25%)

grassland occurs in two sectors of the study area: (Fig. 5). Besides, 19 threatened species were found,

one at the base, near temporary water courses, and three of which had coverage greater than 15%

another in depressions or meadows between the (Table 2).

77

Bol. Soc. Argent. Bot. 58 (1) 2023

78

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

79

Bol. Soc. Argent. Bot. 58 (1) 2023

Fig. 4. Images of the sites that are part of the vegetation units obtained from the PCooA. A: Base Grassland;

B: Pond; C: Mixed Grove; D: Humid Mountain Grassland; E: Dry Mountain Grassland; F: Rocky Place; G:

Spring; H: Acacia Grove.

80

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

Fig. 5. Most abundant non-native species found in diﬀerent areas of the Strict Nature Reserve area of the

Paititi Natural Reserve (Buenos Aires, Argentina). A: Holcus lanatus; B: Dactylis glomerata; C: Senecio

madagascariensis; D: seedling of Racosperma melanoxylon; E: adult of R. melanoxylon.

Rocky Place: This unit is located in elevated areas mainly of herbaceous species, and to a lesser extent,

corresponding to the slopes and top of the hill, with sparse trees and shrubs (38 species). Twenty-ﬁve

abundant rocky outcrops, and stony surfaces (Fig. threatened species were found, most of which had

4F; Table 1). It is the richest vegetation unit since a low coverage (Table 2). The non-native species

total of 239 species were identiﬁed: 17% non-native with the highest level of cover-abundance was

and 0,8% cosmopolitan. The vegetation consisted Racosperma melanoxylon (5-15%).

81

Bol. Soc. Argent. Bot. 58 (1) 2023

Spring: This unit is situated in areas of high to species not previously found in the reserve, possibly

intermediate position in the hill, rocky outcrops of due to their low frequency. Therefore, the richness

low blocks, which have ﬂat to concave shape and are amounted to 370 vascular species. This increase

almost continuous. After the rains, this unit presents did not aﬀect the predominance of the Poaceae

temporary springs and small holes with little or no and Asteraceae families, which continued to be

drainage (Fig. 4G; Table 1). The vegetation reached the best represented, and neither did it change the

20 herbaceous species: 30% non-native species. Most proportion of geographic origin of the species since

of the ﬂoristic components are typically found on the native plants far exceeded the set of non-native and

water courses of the region, while some are common cosmopolitan species by bringing together 72.5%

species of the rocky areas. Two threatened native of the species.

species were found (Juncus pallescens Lam. and

The vegetation units varied mainly due to their

Zephyranthes biﬁda (Herb.) Nic. García & Meerow), position in the relief, the slope, the type and depth

both with very low coverage (Table 2). No exotic of the soil, and the established plant community.

species exceeded 5% cover.

This coincides with several authors who noted

Acacia Grove: This unit occupies areas on the that the ﬂoristic composition and distribution of

slope of the mountain (Fig. 4H; Table 1). Fifty taxa vascular species in hills and mountain areas are

were identiﬁed in this unit, 42% non-native, and 4% associated with the environmental heterogeneity

cosmopolitan species. The dominant physiognomic determined mainly by topographic variants, even in

species was the non-native species Racosperma short distances (Frangi & Bottino, 1995; Guerrero

melanoxylon, which was accompanied in the lower Campo et al., 1999; Cantero et al., 2014). Among

stratum by sparse herbaceous species. In fact, the the vegetation units considered in the study area,

cover-abundance level of R. melanoxylon ranged the ﬂora varied in richness, proportion of species

between 50 and 70%. Four threatened species with according to their status, and cover-abundance of

very low coverage were found in this unit (Table 2).

each species. Some vegetation units were similar

to the plant communities described by Frangi

(1975) for the Tandil hills of the Albion Group.

Threatened species

In the study area, 30 of the native species In some cases, the resemblance was high, as the

considered threatened in Buenos Aires Province dominant and main companion species coincided.

(

Delucchi, 2006) were identified (Table 2). In others, due to the absence of some species or

Regarding their category of threat, 14 are listed as the replacement of some physiognomic dominants,

Vulnerable” and four as “Critically Endangered”. the ﬂora seemed to be a variant of the community

“

Some of the threatened species were found in two described as “typical”. For example, Nassella

or more vegetation units, but ten of them were only poeppigiana (Trin. & Rupr.) Barkworth and

found in a single vegetation unit, in some cases in the Koeleria permollis Nees, dominant species in the

Dry Mountain Grassland and in others in the Rocky Tandil Flechillar according to Frangi (1975), were

Place, but always with very low cover-abundance not found in the in Base grassland vegetation unit. A

levels. In contrast, Acanthostyles buniifolius, similar case was detected in the Rocky place where

Baccharis dracunculifolia ssp. tandilensis, Colletia the ﬂoristic composition of the vegetation unit

paradoxa, and Senecio selloi exhibited cover- coincides mostly with the community described

abundance levels ranging from 15 to 25%, and were by Frangi (1975) for Tandil hills named “Roquedal

found in at least three vegetation units.

típico” (Typical rocky community), except for

several species of physiognomic relevance that

were not found in study area, such as Andropogon

ternatus (Spreng.) Nees, Arjona tuberosa Cav.

and Hatschbachiella tweediana (Hook. & Arn.)

R.M. King & H. Rob., among others. Although

discussion

Flora

In a previous survey of the vascular plants of the the detected absent species should be the object

Strict Nature Reserve area of the PNR, Echeverría of detailed studies, this situation could be due to

el at. (2017) identiﬁed 360 taxa at the species level. the location of the PNR, exceeding the distribution

The present study allowed the detection of ten limit of these species. This hypothesis would apply

82

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

to Hatschbachiella tweediana, a species for which Threatened species

there are no records in the southern portion of the

Thirty vascular species listed as threatened

Tandilia Hill System (Alonso et al., 2009a; Alonso in the Buenos Aires Province (Delucchi, 2006)

et al., 2009b; Flora Argentina, 2022; Flora del Cono were found. This result coincides with that found

Sur, 2022; GBIF, 2022; iNaturalist, 2022). In turn, by Echeverría et al. (2017) for the same area of

some of the absent species, such as Andropogon the PNR, which indicates that no species have

ternatus and Arjona tuberosa, have been collected disappeared over the last years when the reserve has

in other hills in General Pueyrredon district, and been closed to the public. These species are mostly

in Balcarce, a neighboring district (Alonso et al., endemic to the mountain grasslands (Echeverría

2

009a; Alonso et al., 2009). In these cases, the et al., 2017), so they only thrive in mountain

action of anthropic alterations (e.g. indiscriminate environments with little intervention, such as

grazing, urbanization, ﬁre) could have aﬀected the the Strict Nature Reserve area of the PNR. This

presence of these plants in the study area.

situation reveals the important role of this reserve

The vegetation units located in high ground in the conservation of mountain grassland ﬂora in

positions had the highest total richness and native general, and of threatened species in particular,

ﬂora and the dominance of perennial species, in as well as warns about the vulnerability of poorly

particular geophytes, in agreement with Cantero et represented taxa and the need to consider them in

al. (2017) for rocky mountain areas of the Sierra future management plans.

de Los Cóndores in Córdoba Province (Argentina).

According to the data collected in the present

The sites located in high and intermediate positions study, the species Acanthostyles buniifolius,

of the relief tend to present shallow soils, high Baccharis dracunculifolia ssp. tandilensis, Colletia

exposure to solar radiation, and a large proportion paradoxa, and Senecio selloii, which have been

of rocky outcrops and stony surfaces. Therefore, cited as threatened in Buenos Aires Province

these sites are subjected to greater arid conditions (Delucchi, 2006), were recorded with high levels

than those located on lower parts of the slopes or of cover-abundance and in several sites. This

in sectors totally or partially shaded by tree canopy information is encouraging as it indicates that these

(

Mazzola et al., 2008; Aguirre Mendoza, 2013; entities are currently out of danger in the Strict

Kristensen & Frangi, 2015). Exotic species are not Nature Reserve area of the PNR. In particular,

adapted to these environmental conditions, hence for Baccharis dracunculifolia ssp. tandilensis

their lower number or representability in those sites. and Senecio selloi, changes in habitat preference

In the rocky areas and elevated slopes of the have been reported in the area: B. dracunculifolia

study area, the development of phanerophytes is extending from the hills to the plains near the

restricted, mainly because of the shallow soils, Atlantic Ocean coast (Scaramuzzino et al., 2015;

and hence greater richness of this kind of species Manfreda et al., 2020) and S. selloi to grasslands

is found at the base and on the slopes of the hill. dedicated to livestock grazing (Fernández, 2011).

In addition, in the sites located at lower ground Therefore, new surveys in the province could

positions, a greater number of non-native species yield information that accounts for the current

were registered, mostly therophytes. The greater distribution of native species in ecosystems close

abundance of annual cycle species is related to to the studied one and thus clarifying the threatened

the greater anthropization that characterizes the status of the species and eventually incorporating

surrounding rural landscape (Vervoorst, 1967; new taxa into the plant red list.

Zalba & Villamil, 2002; Cantero et al., 2017).

The rural ﬁelds close to the PNR would act Invasive species

as plant reproduction reservoirs, and the wind,

Currently, the most important problems in

waterways, birds, mammals, invertebrates, and the PNR associated with invasions involve the

humans as vectors for their dispersal, introduction, non-native species Dactylis glomerata, Senecio

and spread of plant species in a certain area. This madagascariensis, Holcus lanatus and Racosperma

situation would promote the colonization and melanoxylon (Fig. 5). These exotic taxa had the

proliferation of non-native species toward the hill greater cover-abundance values and the ability to

grasslands of the PNR, thus aﬀecting the ecosystem. thrive in diﬀerent environmental conditions, since

83

Bol. Soc. Argent. Bot. 58 (1) 2023

they were registered in various vegetation units. All (González et al., 1995; Hussain et al., 2011). The

of them have already been cited as invasive species seeds have the ability to remain viable in the seed

in diﬀerent locations of Argentina and vary in bank for many years and their germination is

life forms, morphological attributes, reproduction stimulated by occasional ﬁres that scarify the seeds

strategies and invasion capacity (Ahumada et al., (Arán et al., 2017). It also has gemiferous roots

2

016). Management eﬀorts on the PNR should be and high seed production (Ahumada et al., 2016;

focus on monitoring the threatened species and Arán et al., 2017). R. melanoxylon has shown to be

controlling the non-native with the highest value highly invasive in other mountain grasslands of the

of cover-abundance in each vegetation unit. The Tandilia Hill System where it occurs in the wild and

characteristics of each species should be considered it seems to be advancing on adjacent sectors of the

in planning the management of the invaded hills (Ahumada et al., 2016; Gandini et al., 2019;

vegetation units, since the integrated control of an De Rito et al., 2020).

invader implies, among other requirements, having

complete knowledge of the bioecology of the invaders in the study area, mechanical controls in

species (Bentivegna & Zalba, 2014). H. lanatus, D. glomerata and S. madagascariensis

In relation to the management of the identiﬁed

Senecio madagascariensis is an herbaceous have not been successful while the use of herbicides

perennial species known as “ﬁreweed”. It belongs has allowed eﬀective chemical controls (Villalba

to the Asteraceae family and is native to South & Fernández, 2005; Ahumada et al., 2016). In

Africa and Madagascar. It is a toxic species, the particular case of R. melanoxylon, Arán et al.

recognized as invasive in diﬀerent parts of the (2017) proposed controlling this species by cutting

world (Dematteis et al., 2020). S. madagascariensis down the plants, both adults and juveniles, as well

has been reported in the Argentinean Pampas as a as uprooting all the roots to prevent new shoots

weed in extensive farmlands in the past (Verona from emerging. They also recommend minimal

et al., 1982), while it is currently a problem in soil disturbance to prevent mechanical scariﬁcation

cattle ﬁelds and a growing concern in farms under of the seeds in the seed bank. They also suggest

no-tillage systems and in ruderal areas (Ahumada favoring the colonization of the land with one

et al., 2016; Dematteis et al., 2020; Diez de or several fast-growing native species to prevent

Ulzurrun personal communication, 2022). It is a R. melanoxylon seedlings from prospering. On

species with a high production of seeds per plant the other hand, in a study performed by Campos

and easy to disperse thanks to a hairy pappus, et al. (2002), the combined felling of trees and

characteristics that determine their invasive nature. herbicide application was eﬀective at controlling R.

Dactylis glomerata (“orchard grass”) and H. melanoxylon. Therefore, a similar strategy could be

lanatus (“Yorkshire fog” or “velvet grass”) are suitable to control the proliferation of this species in

perennial microthermal species that belong to the the PNR and in other mountain grasslands. Beyond

Poaceae family. These species are native to Europe this, it is important to consider that the control

and were brought into Argentina as forage. They strategy must be adjusted to each situation, and may

escaped from cultivation and have become weed vary depending on the location of the plant invasion

species that are considered invasive in diﬀerent on the ground (top, slope, base of the hill).

parts of the American continent due to their proliﬁc

It is important to mention that in the study area

seed production and their ability to reproduce from some non-native species were found that have a

tillers (Bastow et al., 2008; Ahumada et al., 2016). history of invasion in mountain grasslands of the

Racosperma melanoxylon (Acacia melanoxylon Pampa biome but with low representativeness

R. Br.; “Australian blackwood”) is an arboreal in the PNR. Such is the case of Eucalyptus

species native to Australia that belongs to the camaldulensis Dehnh., Gleditsia triacanthos L.,

Fabaceae family. In the southeast of Buenos Aires, Ligustrum lucidum W. T. Aiton, Rubus ulmifolius

it was introduced for its cultivation as ornamental Schott, Pinus radiata D. Don and Prunus mahaleb

and forestry (Martínez Crovetto, 1947; Carranza, L., among others (Ghersa et al., 2002; Zalba &

2

007). This evergreen tree causes shade on the Villamil, 2002; Hoyos et al., 2010; Mazzolari

lower stratum and has allelopathic compounds et al., 2011; Ferreras et al., 2014; Mazzolari

that aﬀect the growth and survival of other plants & Comparatore, 2014; Ahumada et al., 2016).

84

M. L. Echeverría et al. - Vegetación y detección de especies invasoras en un pastizal serrano

Therefore, periodic monitoring of the evolution

Finally, we suggest promoting environmental

of these species’ expansion and rapid actions for education and citizen participation activities

their control could prevent future problems of plant aimed at valuing and caring for the hill natural

invasions and facilitate the protection of hill and environments of the Pampa biome and raising

mountain grassland biodiversity.

awareness about their importance within the

Prevention and early detection are the most framework of biodiversity conservation policies.

eﬃcient methods for dealing with invasive species.

When the invasion process advances, the possibility

of carrying out successful control measures acKnowledgements

decreases, management costs and impacts generated

by an invasion process increase, and sometimes the

problem becomes irreversible (Anderson et al., Zugasti, owner of the Paititi Natural Reserve,

014; Brancatelli & Zalba, 2018). Therefore, to who gave permission to carry out this research

We would like to thank Esteban González

2

avoid increasing invasions and the appearance of and provided logistical support. This work was

a new invasion focus, vegetation units should be ﬁnancially supported by Universidad Nacional de

monitored by sporadic surveys. Thus, when an Mar del Plata (Argentina) [research projects AGR

invasive species is detected, rapid action can be 557/18 and AGR 616/20] and The Neotropical

taken to restore the aﬀected site or to minimize the Grassland Conservancy.

invasion process in the early stages, thus preserving

the biodiversity.

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