Influence of “el nIño Southern oScIllatIon” PerIodS on

Plant denSIty of hoffmannSeggIa aPhylla (fabaceae), an

endangered SPecIeS

InfluencIa de loS PeríodoS de “el nIño oScIlacIón del Sur” en la denSIdad

vegetal de hoffmannSeggIa aPhylla (fabaceae), una eSPecIe en PelIgro

1,3

2,3

Felipe S. Carevic * & Roberto Contreras

Summary

Background and aims: Studies on the natural regeneration of plant species under

hyperaridity conditions have been scarce, mainly because of the low germination

percentage of the species under these conditions. Presumably, sporadic

phenomena such as the El Niño Southern Oscillation (ENSO) could have a positive

eﬀect on the natural germination of these species, although this eﬀect has not yet

been satisfactorily explored.

1

. Laboratorio de Ecología Vegetal,

Facultad de Recursos Naturales

Renovables, Universidad Arturo

Prat, Iquique, Chile

2

y

. Centro Regional de Investigación

Desarrollo Sustentable de

M&M: To test the crucial assumption of our statement, a hyperarid region (average

rainfall below 5 mm/year) was used as a model to determine the eﬀect of ENSO

years on the natural regeneration rate of adult individuals of the endemic legume

Hoﬀmannseggia aphylla (retama) in the Atacama Desert, northern Chile. Thus, the

vegetation density of an endemic leguminous species in a sector of the Pampa del

Tamarugal, Tarapacá Region, northern Chile, was analyzed for six years.

Results: The density of this species increased during ENSO years, mainly due to

water ﬂows from the highest sectors of the Tamarugal pampas, such as the town of

Pica, in addition to the increase in humidity and summer rainfall.

Atacama (CRIDESAT), Universidad

de Atacama, Copiapó, Chile

3

. Millennium Nucleus on Applied

Historical Ecology for Arid Forests

Aforest). Santiago de Chile

(

*fcarevic@unap.cl

Citar este artículo

Conclusion: Our results highlight the transcendence of the ENSO in the regeneration

of leguminous plants.

CAREVIC, F. S. & R. CONTRERAS.

023. Influence of “el niño

2

southern oscillation” periods on

plant density of Hoffmannseggia

aphylla (Fabaceae) an endangered

species. Bol. Soc. Argent. Bot. 58:

Key wordS

Atacama desert, ENSO, Fabaceae, Hoﬀmannseggia, Hyperarid, legume.

reSumen

257-265.

Introducción y objetivos: Los estudios sobre la regeneración natural de especies

vegetales en condiciones de hiperaridez han sido escasamente estudiados en

terreno, principalmente por el bajo porcentaje de germinación de las especies

en estas condiciones. Presumiblemente, fenómenos esporádicos como El Niño

Oscilación del Sur (ENOS) podrían tener un efecto positivo en la germinación natural

de estas especies, aunque este efecto aún no ha sido explorado satisfactoriamente.

M&M: Para testear la suposición crucial de nuestra aﬁrmación, se utilizó una

región hiperárida (precipitación promedio inferior a 5 mm/año) como modelo para

determinar el efecto de los años ENOS en la tasa de regeneración natural de

individuos adultos de la leguminosa endémica Hoﬀmannseggia aphylla (retama)

en el desierto de Atacama, norte de Chile. Así, se analizó durante seis años la

densidad vegetal de esta especie en un sector de la Pampa del Tamarugal, Región

de Tarapacá, norte de Chile.

DOI: https://doi.

org/110.31055/1851.2372.v58.

n2.38609

Resultados: La densidad de esta especie se incrementó durante los años ENOS,

principalmente por los caudales de agua provenientes de los sectores más altos

de la pampa del tamarugal, como el poblado de Pica, además del aumento de la

humedad y las precipitaciones estivales.

Conclusión: Nuestros resultados destacan la trascendencia del ENOS en la

regeneración de plantas leguminosas.

PalabraS claveS

Desierto de Atacama, ENSO, Fabaceae, hiperárido, Hoﬀmannseggia, leguminosa.

Recibido: 22 Ago 2022

Aceptado: 1 Jun 2023

Publicado impreso: 30 Jun 2023

Editora: Natalia Aguirre Acosta

ISSN versión impresa 0373-580X

ISSN versión on-line 1851-2372

257

Bol. Soc. Argent. Bot. 58 (2) 2023

IntroductIon

(Ministerio de Medio Ambiente, 2022). The

main threat to natural H. aphylla populations

The natural regeneration processes of plant in northern Chile is associated with the current

species are currently considered one of the main depletion of the water table, which may be a

problems facing climate change because the real direct cause of the decline in legume populations

eﬀects of this problem on the reproductive rates of (Chavez et al., 2013). Former studies denoted

the species are still unknown (Peters et al., 2012). that the water ﬂux into the aquifer of Pampa del

In this sense, the Atacama Desert is no stranger Tamarugal, is calculated to be between 880 and

to this type of process, mainly because of four 1000 l/s, while the water outﬂow is estimated

determining factors: 1) the eﬀect of the decrease to be as high as 4000 l/s (Dirección General de

in precipitation recorded at the beginning of Aguas, 2011; Calderón et al., 2015; Viguier et al.,

this century in northern Chile (Soto & Ulloa, 2019). In this ecosystem, groundwater aquifers

1

997); 2) the decrease in relative soil humidity are the only source of water, which are fed by the

in this area (Neilson et al., 2017); 3) the increase rainfall, glaciers and snowmelt from the Andes

in salinity in high Andean limnetic bodies Mountain range. This shrub has conditions of

(

Gajardo & Redón, 2019); and 4) the decrease successful natural adaptability in arid soils with

in the groundwater table (Viguier et al., 2019). high salinity (even with electrical conductivities

This type of groundwater has been described as of 182 Ds/m) and nitrogen impoverishment in the

one of the main sources of water supply for the edaphic stratum (León et al., 2017). However,

maintenance of the vegetation present in this the natural regeneration of plant species in this

ecosystem, but the eﬀect of precipitation pulses area is almost null due to the almost nonexistent

or the occurrence of climatic phenomena such precipitation averaging 1 mm/year and the

as the El Niño Southern Oscillation (ENSO) presence of a saline crust layer on the edaphic

could have some impact on vegetation density surface (León et al., 2017). In this sense, it fails

(

Squeo et al., 2006). In the past decade Chile has to form a homogeneous shrub stratum, as the

experienced a mega drought, where the presence distribution of individuals on the ground is not

of a strong ENSO during most of 2015 stood out very dense, with individuals that seem to form

(

Garreaud et al., 2019).

mosaics following the ﬂoods that usually occur

The natural regeneration of desert leguminous in this ecosystem (Lewis & Sotuyo, 2010).

species under ENSO conditions has scarcely Once established, adult individuals of Prosopis

been evaluated under ﬁeld conditions (Kraus et tamarugo Phil., for example, would depend on

al., 2007). The occurrence of this phenomenon the ﬂow of water present at the phreatic level,

is a powerful tool for evaluating this type of which is usually present in the ﬁrst meters of soil

ecological condition in plants, as precipitation depth owing to its ascent by capillarity (Calderón

and humidity tend to increase in arid areas of et al., 2015). Preliminary results have indicated

northern Chile when ENSO is present (Acosta a favorable increase in the regenerative rates

et al., 2015). The genus Hoﬀmannseggia Cav. of the species, although this trend should be

(

Fabaceae) has a wide distribution, including further explored in detail, considering edaphic

the arid and semi-arid regions of Argentina, factors and ﬂooding episodes during the summer

Chile, Peru, and Bolivia in South America, and (Carevic, 2020). Regarding this condition, natural

Mexico and the southwestern United States seed germination could occasionally occur as a

in North America (Kraus et al., 2007). In the result of summer precipitation and water ﬂooding

Atacama Desert, Hoﬀmannseggia aphylla (Phil.) during ENSO periods and to a lesser extent

G.P. Lewis & Sotuyo (Spanish “retama”) is during Interdecadal Paciﬁc Oscillation (IPO)

an endemic phreatophyte shrub legume that periods. However, this eﬀect has not yet been

ﬁxes atmospheric nitrogen, and its distribution explored in this species.

is limited to the northern part of this desert,

Previous studies carried out in arid ecosystems

speciﬁcally the Pampa del Tamarugal in northern have reported a positive eﬀect of precipitation

Chile (Lewis & Sotuyo, 2010). Recently, this during ENSO events on the density of plant

species has been considered as “endangered” species growing at low altitudes above sea

258

F. S. Carevic & R. Contreras, Densidad de Hoﬀmannseggia aphylla en el norte de Chile

level; although the same precipitation was not spring (September). For this objective, 38 linear

signiﬁcant for the increase in primary productivity transects of 95 m were georeferenced within the

of species established at high altitudes (2,500 m grid, with points every 5 m. Flower sprouting

above sea level) (Squeo et al., 1999). However, was assessed semi-annually during the period

the influence of seasonal environmental of 2012-2016 by tagging the flowers which

parameters and their eﬀects on plant density emerged using a thread of a particular color,

and reproduction parameters in legume species with a distinctive color used for period so that at

growing under hyperarid conditions are poorly harvest, ﬂower production as well as pods could

understood, considering that climate may play an be attributed to the respective period of sprouting

important role in monitoring and predicting plant (Leport et al., 2006). For this purpose, we selected

viability and reproduction. We hypothesized that 12 adult individuals of H. aphylla. This tagging

the increased surface water ﬂow resulting from was also used to record the total number of ﬂowers

precipitation during ENSO years would have an produced per plant (total number of tags per

incremental eﬀect on the germination rate and plant), and the number of pods per plant (number

ﬂower parameters (number of ﬂowers, number of of tags with a date of ﬂower sprouting), and the

pods per plant and percentage of ﬂower abortion) proportion of aborted ﬂowers. For environmental

of H. aphylla seeds in northern Chile. Thus, our characterization, two meteorological stations were

objective was to evaluate the long-term impact of established near the study area to record daily

environmental variables during ENSO events in environmental conditions (ambient temperature,

northern Chile on the reproductive and ecological precipitation, wind speed, and solar radiation),

parameters of H. aphylla, an endangered endemic both located on slopes, to determine the eﬀect of

species of the Atacama Desert.

ﬂooding on the study area. The ﬁrst was located

in Canchones (UTM: 7739099, 443867; 1005 m

a.s.l.), located 1.5 km from the study area. Another

meteorological station was in the town of Pica

materIalS and methodS

(

UTM: 7734562, 465733; 1345 m a.s.l.), 15 km

Study area

from the study area. Similarly, ﬁve digital soil

The study was carried out within the natural humidity probes (ECHO) were established at 0.6

habitat of H. aphylla in Pampa del Tamarugal, m depth distributed in the study area to record

northern Chile. The climate of this area is the seasonal evolution of this parameter. Each

hyperarid. The site was dominated by P. tamarugo, humidity sensor was separated by approximately

H. aphylla, and Atriplex atacamensis Phil. 70 m from each other.

The latter two species are small shrubs that are

sparsely distributed on the grid. The study area Statistical analysis

is characterized by extreme hyperarid conditions,

To evaluate interannual differences in H.

with annual precipitation not exceeding 0.6 mm/ aphylla density and ﬂower parameters (number of

year, which occurs mainly during the summer ﬂowers, number of pods per plant and percentage

season (Carevic et al., 2021). However, there of ﬂower abortion), we used a linear general

are years with rainfall pulses that exceed 7 mm/ model by means of repeated measures. The ﬂower

year, especially in sectors with an altitude above parameters data were log-transformed ln (x+1)

sea level higher than that present in Canchones to increase linearity and reduce the correlation

(

Arenas 2019; Lanino & Poblete, 2022).

between the mean and variance. Relationships

between meteorological and edaphic data

(temperature, precipitation, solar radiation,

Experimental design

During the period 2012-2018, an experimental wind speed, and soil humidity) and H. aphylla

grid of 3.1 ha was established in the Canchones density and ﬂower production were evaluated

sector in the Pampa del Tamarugal, northern using multiple matrix correlations (r). Data was

Chile to evaluate H. aphylla density using the analyzed by using SPSS Statistics for Windows,

transect line method (Cox, 1981). Sampling was Version 23.0 (IBM SPSS Statistics for Windows,

carried out during the fall (May) of each year and Version 23.0. Armonk, NY: IBM Corp).

259

Bol. Soc. Argent. Bot. 58 (2) 2023

Fig. 1. Precipitation, temperature and humity in the study area. A: Annual precipitation detected in Pica

(black columns) and Canchones (white columns) (left, above). B: Mean annual summer temperatures (black

line) and mean humidity (white bars) in Canchones.

reSultS

The environmental humidity and temperatures

detected during summer had diﬀerent ﬂuctuations,

During the present study, the presence of two with the highest environmental humidity values

ﬂooding events during the summers of 2015 and recorded in 2015 and the highest summer

2

016 was observed, reﬂected in the increase in soil temperature averages in 2017. Likewise, the density

humidity during those periods and the presence of determined during 2015 and 2016 was strongly

summer rainfall that exceeded 4 mm/year in Pica correlated with the environmental humidity detected

(

Figs. 1; 2).

during summer 2015 in Canchones (December

st st

The most important variation associated with 21 to March 21 ); r=0.56, p<0.01, and with the

H. aphylla density parameters was related to an annual precipitation recorded in Pica the same

increase in the number of individuals per area after year (r=0.51, p=0.02). Furthermore, we observed

the 2015 event (Table 1), when new individuals a signiﬁcant relationship between soil humidity

of low size were recorded, typical of regeneration and plant density during the same period (r=0.39,

phenomena, which subsequently decreased (2017 p=0.03). Regarding ﬂower parameters, we did not

and 2018).

ﬁnd signiﬁcant diﬀerences between numbers of

260

F. S. Carevic & R. Contreras, Densidad de Hoﬀmannseggia aphylla en el norte de Chile

Fig. 2. Seasonal evolution of average soil humidity by season of the year.

ﬂower sprouts at population level. Furthermore,

ﬂower parameters did not show any relationship

with climatic parameters (Table 2).

Table 1. Area density of H. aphylla individuals

during the study. Diﬀerences between years are

denoted by diﬀerent letters.

Plant density H. aphylla

dIScuSSIon

2

Year

p value

Individuals/m

0.001

F value

24.55

29.09

30.87

41.76

41.08

21.22

21.97

Our ﬁndings constitute the ﬁrst large-scale study

of the natural regeneration processes generated

under hyperarid conditions in the Atacama Desert.

Thus, precipitation generated during ENSO

events seems to be an essential factor for the

germination of H. aphylla in northern Chile,

consistent with previous studies indicating that

the increase in sporadic water pulses present

during this phenomenon plays an essential role

in the regeneration of arid zone plant species

ab

2012

2013

2014

2015

2016

2017

2018

0.12

b

a

0.21

0.22

0.04

0.001

(Gutiérrez et al., 2000; Squeo et al., 2006; León

0.001

et al., 2011). In the present study, rainfall recorded

in Pica, that is, in a sector with a higher altitude

above sea level, had the greatest relationship with

the increase in vegetation density of this legume

(Fig. 3). This record is not surprising, as, in “wet”

periods, precipitation tends to carry water to lower

ab

0.08

0.09

0.001

261

Bol. Soc. Argent. Bot. 58 (2) 2023

Table 2. Summarized data on ﬂower sprout at intrapopulation level.

Periods measured

April November March November

Parameter

December August

May

2012

2013

2014

2015

2016

Number of ﬂowers (mean±SE)

12±2

13±4

10±3

11±2

13±2

13±3

10±2

Number of pods per

plant (mean±SE)

0

3±2

4±2

3±3

2±2

Percentage of ﬂower abortion (%)

76.9

69.2

76.9

84.6

areas such as Canchones, where this water forms 50% seem to break the physiological dormancy of

real lagoons that tend to dry up or inﬁltrate as the this species. Previous investigations, carried out at

summer progresses (Houston, 2001). However, the the laboratory level, agree that water availability,

increase in water availability must necessarily be temperature and increased environmental humidity

accompanied by an increase in humidity during the are limiting factors for breaking dormancy in

summer in the same study site for this germination species of this genus (Alves et al., 2018; Moreno

phenomenon to take place, in which values above et al., 2018).

Fig. 3. Schematic illustration of comparative situation ENSO observed in the study area. A: In 2012.

Normal periods of rainfall in Pica (1) and environmental humidity in Canchones (2) do not generate an

incremental variation in the density of H. aphylla population (3). B: In 2015. Increasing rainfall at Pica (4) and

environmental humidity in Canchones (5), the plant density of H. aphylla increasing notably (6).

262

F. S. Carevic & R. Contreras, Densidad de Hoﬀmannseggia aphylla en el norte de Chile

There has been some controversy regarding the humidity. It should be noted the positive eﬀect

eﬀects of environmental variables on germination. that the ﬂooding registered as a result of the

Formerly, some authors have claimed that precipitation in the Pica sector, contributed to a

temperature and sunlight are the limiting factors greater extent to the increase in the density of

for germination processes in plants in the arid H. aphylla plants, which has an explanation due

zones of Asia (El-Keblawy & Al-Rawai, 2005). to the higher altitude above the ground sea level

Such studies on the regeneration of native species that Pica presents in relation to the study area

have been limited to a few years of evaluation and (300 m higher), since when precipitation falls on

are usually restricted to semiarid ecosystems with that area, the eﬀect of gravity probably allows

the highest availability of precipitation. In our the displacement of these masses of water to

study, we found no direct evidence of the eﬀect the ﬂatter areas of the Tamarugal pampa, where

of sunlight and temperature on H. aphylla density, it was carried out the study. Our research was

presumably because these parameters remained developed over a long period of time, with the aim

homogeneous during much of the summer, the of evaluating the eﬀect of climatic phenomena on

time when increased density was detected (Arenas, the population of an endangered plant species. It

2

019). Nevertheless, this natural regeneration is essential that monitoring is embedded in a long

decreased during 2017 and 2018, probably because management program by the accountable agency

of predation by goat cattle, which is usually or research institutions that is working towards

abundant in this area and is associated with the the recovery and monitoring of a given threatened

consumption of native legumes (Carevic et al., species or threatened ecosystem. We highlight

2

015; Contreras et al., 2020). At reproduction that these beneﬁts of monitoring will be far more

level, we did not ﬁnd any relationship between likely to be achieved if the monitoring program is

climatic traits and ﬂower production. The highest well designed with clearly articulated objectives,

values of ﬂower abortion and the reduced number with due attention to long term biological traits

of ﬂowers per plant are not surprising, because and its dynamics based on future climatic eﬀects.

it has a slow growth with few production of

leaves and ﬂowers, consistent with the hyperarid

habitat of Atacama Desert (Lewis & Sotuyo, author contrIbutIonS

2

010). In conclusion, the presence of ENSO has

a transcendental inﬂuence on the regeneration of

FC did all experimental work and prepared ﬁrst

leguminous plants, especially due to the increase draft of the paper; RC participated on ﬁeld sampling

in precipitation in Pica (which only falls within and prepared ﬁnal version of the paper.

a few hours) and, to a lesser extent, Canchones,

which cause water ﬂooding to lower sectors of the

Tamarugal pampas, a fact that was detected by an acKnowledgementS

increase in soil humidity. In addition, this eﬀect

must be combined because it must be accompanied

Special thanks to Vicerrectoria Investigación,

by an increase in the average environmental Innovación y Postgrado of Universidad Arturo

humidity during the summer (January-March).

Prat, ANID NCS2022_024 Millennium Nucleus

on Applied Historical Ecology for Arid Forests

(Aforest) and Junta Vecinal Challajure. To Cynthia

concluSIonS

Chávez, Paula Rojas for their support in scientiﬁc

graphic design and Luis Ulloa for their collaboration

We demonstrate at the ﬁeld study level, the in ﬁeld support.

positive inﬂuence of the ENSO phenomenon on

the natural regeneration rate of H. aphylla in the

Atacama Desert. The presence of this climatic SuPPlementary data

phenomenon generates water ﬂoodings that allow

the germination of seeds, although this aspect must

Supplementary data to this article can be found

be accompanied by an increase in environmental at: https://github.com/carevicunap/haphylla.

263

Bol. Soc. Argent. Bot. 58 (2) 2023

bIblIograPhy

COX, W. G. 1981. Laboratory manual of general

ecology. C publishers, Brown.

ACOSTA, G., J. GUTIÉRREZ, P. L. MESERVE, D.

A. KELT & M. A. PREVITALLI. 2015. El Niño

Southern Oscillation drives conﬂict between wild

carnivores and livestock farmers in a semiarid area

in Chile. J. Arid Environt . 126:76-80.

DGA. 2011. Actualización de la oferta y la demanda

de recursos hídricos subterráneos del sector

hidrogeológico de aprovechamiento común Pampa

del Tamarugal. Dirección General de Aguas,

Ministerio de Obras Públicas, Gobierno de Chile.

Available at: https://snia.mop.gob.cl/repositoriodga/

handle/20.500.13000/5309 [Accessed: 10 June

2023].

EL-KEBLAWY, A. & A. AL-RAWAI. 2005. Eﬀects

of salinity, temperature and light on germination

of invasive Prosopis juliﬂora (Sw.) DC. J. Arid

Environt . 61: 555-565.

https://doi.org/10.1016/j.jaridenv.2004.10.007

GAJARDO, G. & S. REDÓN. 2019. Andean hypersaline

lakes in the Atacama Desert, northern Chile:

Between lithium exploitation and unique biodiversity

conservation. Conserv. Sci. Pract. 1: e94.

https://doi.org/10.1016/j.jaridenv.2015.08.021

ALVES, M. M., E. U. ALVES, L. R. DE ARAÚJO, M.

DE L. DOS S. LIMA & M. M. URSULINO. 2018.

Germination and vigor of Caesalpinia pulcherrima

(L.) Sw. seeds under diﬀerent light and temperature

conditions. Ciência Rural 48: e20170741.

https://doi.org/10.1590/0103-8478cr20170741

ARENAS, J. 2019. Boletín agroclimático de la localidad

de Pica. Universidad Arturo Prat, Iquique.

CALDERÓN, G., M. GARRIDO & E. ACEVEDO.

2

015. Prosopis tamarugo Phil.: a native tree from

the Atacama Desert. Groundwater table depth

thresholds for conservation. Rev. Chil. Hist. Nat. 88:

https://doi.org/10.1111/csp2.94

1

8. https://doi.org/10.1186/s40693-015-0048-0

GARREAUD, R. D., J. P. BOISIER, R. RONDANELLI,

A. MONTECINOS, … & D. VELOSO-ÁGUILA.

2019. The Central Chile Mega Drought (2010-

2018): A climate dynamics perspective. Int. J.

Climatol. 1-19. https://doi.org/10.1002/joc.6219

GUTIÉRREZ, J. R., G. ARANCIO & F. M. JAKSIC.

2000. Variation in vegetation and seed bank in a

Chilean semi-arid community aﬀected by ENSO

1997. J. Veg. Sci. 11: 641-648.

CAREVIC, F., J. DELATORRE & J. ARENAS. 2015.

Physiological strategies during frost periods for two

populations of Prosopis burkartii, an endangered

species endemic to the Atacama Desert. J. Arid

Environt. 114: 79-83.

https://doi.org/10.1016/j.jaridenv.2014.11.008

CAREVIC, F. 2020. Fenómeno ENSO favorece la

cobertura vegetal en la Pampa del Tamarugal,

norte de Chile. En: RED DE UNIVERSIDADES

DE CHILE (ed.), El niño y cambio climático sus

efectos en Chile. El niño-oscilacion del sur (enso) y

teleconexión, pp 55-56. Santiago de Chile.

https://doi.org/10.2307/3236571

HOUSTON, J. 2001. La precipitación torrencial del año

2000 en Quebrada Chacarilla y el Cálculo de recarga

al acuífero Pampa Tamarugal, norte de Chile. Rev.

Geol. Chile 28: 163-177.

CAREVIC, F., E. ALARCÓN & A. VILLACORTA.

2

021. Is the visual survey method eﬀective for

http://dx.doi.org/10.4067/S0716-02082001000200002

KRAUS, T. A., M. A. GROSSO, S. C. BASCONSUELO,

C. A. BIANCO & R. N. MALPASSI. 2007.

Morphology and anatomy of shoot, root, and

propagation systems in Hoﬀmannseggia glauca.

Plant Biol. 9: 705-712.

measuring fruit production in Prosopis tree species?

Rangel. Ecol. Manag . 74: 119-124.

https://doi.org/10.1016/j.rama.2020.09.001

CHÁVEZ, R. O., J. G. P. W. CLEVERS, M. HEROLD,

E. ACEVEDO & M. ORTIZ. 2013. Assessing water

stress of desert tamarugo trees using in situ data and

very high spatial resolution remote sensing. Remote

Sens . 5: 5064-5088.

https://doi.org/10.1055/s-2007-965080

LANINO, M. & I. POBLETE. 2022. Antecedentes

climáticos históricos de la Estación Experimental

Canchones, Pampa del Tamarugal: herramienta para

la toma de decisiones en agricultura del desierto.

Idesia 40: 27-38.

https://doi.org/10.3390/rs5105064

CONTRERAS-DÍAZ, R., F. S. CAREVIC, V. PORCILE

&

M. ARIAS-ABURTO. 2020. Development of SSR

loci in Prosopis tamarugo Phillipi and assessment of

their transferability to species of the Strombocarpa

section. For. Syst . 29: e012.

http://dx.doi.org/10.4067/S0718-34292022000200027

LEÓN, M. F., S. I. SILVA,A. SANDOVAL, I.ARACENA,

… & P. LEÓN-LOBOS. 2017. El manejo del suelo

salino usando arena afecta el crecimiento de raíces y

https://doi.org/10.5424/fs/2020292-16706

264

F. S. Carevic & R. Contreras, Densidad de Hoﬀmannseggia aphylla en el norte de Chile

la sobrevivencia de plántulas de Prosopis tamarugo

Phil. (Fabaceae). Gayana Bot. 74: 86-94.

and potential reversal of desertification in arid

and semiarid ecosystems. Glob. Change Biol. 18:

151-163.

http://dx.doi.org/10.4067/S0717-66432017005000320

LEÓN, M. F., F. A. SQUEO, J. R. GUTIERREZ & M.

HOLMGREN. 2011. Rapid root extension during

water pulses enhances establishment of shrub

seedlings in the Atacama Desert. J. Veg. Sci. 22:120-

https://doi.org/10.1111/j.1365-2486.2011.02498.x

POL, R. G., G. I. PIRK & L. MARONE. 2010. Grass

seed production in the central Monte desert

during successive wet and dry years. Plant Ecol.

208: 65-75.

1

29.

https://doi.org/10.1111/j.1654-1103.2010.01224.x

LEWIS, G. P. & S. SOTUYO. 2010. Hoﬀmannseggia

aphylla (Leguminosae: Caesalpinieae), a new name

for a Chilean endemic. Kew Bull. 65:221-224.

MINISTERIO DE MEDIO AMBIENTE. 2023. 18º

proceso de clasificación de especies. Available

at: https://clasificacionespecies.mma.gob.cl/

wp-content/uploads/2023/01/Hoffmannseggia_

aphylla_18RCE_FINAL.pdf [Accessed: 1 June

https://doi.org/10.1007/s11258-009-9688-y

SOTO, G. & F. ULLOA, F. 1997. Diagnóstico de la

Desertificación en Chile. Corporación Nacional

Forestal (CONAF)-FAO-PNUMA, La Serena.

SQUEO, F.A., N. OLIVARES, S. OLIVARES, A.

POLLASTRI, … & J. R. EHLERINGER. 1999.

Grupos funcionales en arbustos desérticos

definidos en base a las fuentes de agua utilizadas.

Gayana Bot. 56: 1-15.

2

023].

SQUEO, F. A., Y. TRACOL, D. LÓPEZ, J. R.

GUTIÉRREZ, ... & J. R. EHLERINGER. 2006.

ENSO effects on primary productivity in Southern

Atacama Desert. Adv. Geosci. 6: 273-277.

https://doi.org/10.5194/adgeo-6-273-2006

VIGUIER, B., H. JOURDE, V. LEONARDI, L.

DANIELE, … & V. DE MONTETY. 2019. Water

table variations in the hyperarid Atacama Desert:

Role of the increasing groundwater extraction

in the Pampa del Tamarugal (Northern Chile). J.

Arid Environ. 168: 9-16.

MORENO, D., A. DUPLANCIC, A. L. NAVAS, M.

HERRERA-MORATTA & A. DALMASSO. 2018.

Evaluación de la germinación de Caesalpinia

gilliesii Wall. ex Dietr. Multequina 27: 41-48.

NEILSON, J. W., K. CALIFF, C. CARDONA, A.

COPELAND, … & R. MAIER. 2017. Signiﬁcant

impacts of increasing aridity on the arid soil

microbiome. mSystems 2: e00195-16.

https://doi.org/10.1128/mSystems.00195-16

PETERS, D. P. C., J. YAO, O. E. SALA & J. P.

ANDERSON. 2012. Directional climate change

https://doi.org/10.1016/j.jaridenv.2019.05.007

265