Avances de la sucesión vegetal en el volcán Paricutín: 25 años después del cese de

actividad

Introducción
La tasa de sucesión ecológica depende de las circunstancias. A pesar de la idoneidad de las
nuevas áreas volcánicas para el estudio de la sucesión primaria, las observaciones a largo
plazo de los mismos sitios son fragmentarias. Los datos sobre colonización, de 1960-1977, nos
permiten comparar la tasa de colonización del cono principal de Paricutín con diferentes series
sucesionales en otras localidades.

Descripción del sitio e historia del vulcanismo

El Volcán Paricutín está situado en el centro oeste de Michoacán, México. Comenzó a entrar en
erupción el 20 de febrero de 1943 y estuvo activo hasta el 4 de marzo de 1952, resultando un
gran cono de ceniza, flujos de lava que cubrieron casi 25 km 2, depósitos de ceniza y el cono
basáltico es el rasgo más destacado.

El pico del borde del cráter (2808 m) se eleva 410m de centiárea sobre el nivel original de los
campos de maíz, la base del cono es ovalada, con un diámetro de centiárea de 650 metros; el
diámetro es de 900 m; el borde del cráter es casi circular con un diámetro de centiárea de 280
metros. Los fondos de las dos ventilaciones del cráter son 30 o 40 m de centiárea por debajo de
los niveles de sillares en el borde. Gran parte del cono está formado por capas sucesivas de
materiales piroclásticos como cenizas, lapilli y bombas. Solo unos pocos respiraderos
fumarólicos situados a lo largo del borde del cráter estaban activos en 1977.

Cuentas previas de la vegetación

El primer informe sobre plantas que invaden el cono de ceniza de Paricutin fue hecho por
Segerstrom, quien encontró dos especies de plantas vasculares creciendo cerca de la cumbre
en febrero de 1957, Gnaphalium sp. y Eryngium sp.

En el verano de 1958, Beaman (1960) recolectó 13 especies de plantas vasculares en o cerca

del borde del cráter. Eggler realizó varios viajes al cono en los veranos de 1959 y 1960 y las
plantas más abundantes fueron Eupatorium adenochaenium, Pityrogramma tartarea,
Gnaphalium canescens, Muhlenbergia minutiss M. minutiflora (esto es un error de Eggler) y
Cheilanthes farinosa. Otras especies eran más escasas y en algunos casos estaban
representadas por una sola planta. Eggler también determinó el contenido de nitrógeno en las
muestras de cenizas recolectadas del cono.

Observaciones
EN 1977 Las 39 especies vasculares encontradas en el cono de Paricutín el 20 y 21 de
septiembre de 1977 se enumeran en la Tabla 1. Se distinguieron tres tipos de comunidades de
plantas sobre la base de 16 niveles fitosociológicos (25 m2): (1) etapa sucesional inicial en
ceniza fina con Muhlenbergia minutissima y Aster exilis como especies dominantes; (2) la
comunidad más extendida de Aegopogon cenchroides y Gaultheria parviflora en sustratos de
ceniza gruesa stabi, y (3) la comunidad de Crusea sp. y Eragrostis ciliaris situado en la
proximidad de respiraderos fumarólicos activos. Las otras especies principales son Eragrostis
pectinacea, Pityrograma tartarea, Eupatorium spp. y Gnaphalium spp. La cobertura total de
plantas vasculares solía ser de hasta el 20% y nunca superaba el 40%. Los árboles más viejos
de Pinus spp. y Baccharis salicifolia se encontraron no lejos de la localidad de recolección de
Pinus montezumae mencionada por Beaman (1960) (ver Fig. 1).

Discusión
Beaman (1960) concluyó: "Paricutín ahora proporciona un diseño modelo para el estudio de la
dispersión, establecimiento, sucesión y desarrollo comunitario de plantas en productos
volcánicos inorgánicos".

La tasa de colonización del cono de ceniza de Paricutín hasta este momento se compara con
otras localidades en la Figura 2. Para hacer comparables las curvas de colonización de las
sucesiones primarias, se eligió como año de inicio para el año en que se observó la primera
especie en la localidad. cada curva. La curva de las dunas de arena se ha construido sobre la
base de la ecuación publicada por McNaughton y Wolf (1973). Debido a que el número de
especies depende del tamaño del área investigada, las únicas curvas directamente
comparables son las de Hawai y Paricutín. Las parcelas para el estudio de las sucesiones
secundarias fueron de menor tamaño.

Sólo fue necesario un corto tiempo para el inicio de la sucesión primaria en el cono de ceniza de
Paricutín. Desde entonces, la tasa de inmigración (inicialmente hasta 10 especies por año) ha
ido disminuyendo. Esto está de acuerdo con la teoría de la biogeografía insular (MacArthur,
1972). El material volcánico desnudo exhibe condiciones térmicas, de humedad y nutricionales
muy severas. Por lo tanto, muchas especies se han extinguido. Sin embargo, sobre la base de
los datos disponibles, no está claro si la tasa de extinción había aumentado o disminuido.

A pesar de la discontinuidad en la investigación de campo, Paricutín todavía brinda una

oportunidad única para el estudio de la colonización y la sucesión.
Establecimiento de coníferas tras la erupción del volcán Paricutín en el centro de México

1. Antecedentes
Las erupciones volcánicas son grandes perturbaciones que modifican sustancialmente las
condiciones ambientales, la disponibilidad de recursos, los sustratos y la estructura de los
ecosistemas y paisajes (Del Moral y Grishin, 1999). La deposición de ceniza y arena volcánica
(tefra), incluidos los flujos de lava posteriores, pueden cubrir grandes áreas, lo que representa
una barrera para el establecimiento de plantas (Titus y Tsuyuzaki, 2003), ya que estos nuevos
sustratos experimentan altos niveles de insolación, fluctuaciones extremas de temperatura, baja
contenido de materia orgánica, baja retención de agua y alta porosidad (Chang et al., 2019).

El tiempo desde la erupción hasta la llegada de la vegetación se conoce como lapso de

establecimiento o ecesis (Pierson, 2007). El patrón de sucesión más común en áreas volcánicas
es la colonización por especies pioneras como líquenes y musgos, seguidas por pastos y
arbustos, y finalmente por especies arbóreas (Cutler et al., 2008). La sucesión de plantas
depende en gran medida de la capacidad de las semillas para germinar, establecerse y crecer
en los nuevos sustratos (Vilmundardottir et al., 2018). Además, la distancia de los bosques
maduros a los claros cubiertos por sustratos volcánicos influye en la llegada de vegetación
colonizadora (Holle y Tsuyuzaki, 2018). La variabilidad climática también juega un papel
fundamental en el establecimiento de las plantas, ya que la disponibilidad de humedad y las
temperaturas adecuadas influyen en el reclutamiento de árboles (Del Moral y Grishin, 1999).

Los sustratos formados por volcanes representan una oportunidad para caracterizar las
trayectorias de sucesión primaria, la duración del establecimiento, el orden de llegada de las
especies, la capacidad de supervivencia de las plantas en ambientes extremos, el efecto de la
distancia al bosque maduro al momento del establecimiento, y patrones espaciales de
colonización (Del Moral y Bliss, 1993). Esta información es crucial para proporcionar pautas
para gestionar áreas perturbadas e informar los esfuerzos de restauración ecológica, incluso
por otras perturbaciones como grandes incendios (Robinson y Handel, 2000). En ecosistemas
templados, la formación de anillos de crecimiento anual en especies de coníferas ofrece la
posibilidad de reconstruir la historia de establecimiento de poblaciones y evaluar la capacidad
de estas especies para regenerarse después de este tipo de perturbaciones (Stoffel y
Bollschweiler, 2008).

El volcán Paricutín surgió en 1943 en el centro-occidente de México y provocó la devastación

de pueblos, tierras agrícolas y bosques de coníferas, incluyendo la caída de ceniza volcánica en
más de 300 km2 (Pioli et al., 2008). La erupción también afectó el crecimiento de los pinos en
los bosques circundantes (Sheppard et al., 2008). En áreas cubiertas por magma petrificado y
tefra (24.5 km2), la sucesión primaria comenzó rápidamente en 1950 en los sustratos más
antiguos, principalmente por musgos y líquenes; las plantas vasculares llegaron en 1957,
alcanzando hasta 28 especies entre hierbas y arbustos en 1960 (Eggler, 1963). En 1977, 25
años después del final de la erupción, se registraron 39 plantas vasculares (R ejmanek et al.,
1982). Posteriormente, 50 años después de la erupción, 49 especies de plantas habían
colonizado los nuevos sustratos (Lindig-Cisneros et al., 2006).

Observaciones de campo han reportado la presencia de coníferas establecidas sobre los

depósitos de tefra y flujos de lava del volcán Paricutín, que son las especies dominantes en el
bosque que rodea los sustratos volcánicos (Medina, 2003). Sin embargo, la historia del
establecimiento de estas coníferas no ha sido documentada. Dado que los árboles representan
la última etapa serial en perturbaciones de gran magnitud (Cook y Halpern, 2018), esta
información podría ser útil para caracterizar el patrón de sucesión de los árboles dominantes en
estos sustratos volcánicos.
Los objetivos de este estudio fueron: (1) reconstruir la historia del establecimiento de coníferas
en tefra y flujos de lava del volcán Paricutín, (2) caracterizar el orden de colonización de las
especies y los patrones de establecimiento, abundancia y dominancia de los árboles, (3) para
evaluar el efecto de la distancia del bosque preservado en los patrones de establecimiento
temporal y espacial, y (4) para evaluar el efecto del clima en el establecimiento de árboles.

2. Methods
2.1. Study area
The Paricutín volcano is located at coordinates 19290 3900 N, 102150 0500 W in the state of
Michoacan, in central Mexico, belonging to the Tran-Mexican Volcanic Belt (Fig. 1). Its volcanic
activity began on February 20, 1943, and ended on March 4, 1952, emerging on a montane
plain covered originally by cornfields at 2,400 m of elevation (Wilcox, 1954). It was the second
volcano whose emersion was historically recorded in that portion of Mexico, after the El Jorullo
volcano, which emerged in 1759, 100 km southeast of Paricutín (Bullard, 1947).

The Paricutín volcano is a monogenetic cineritic cone, formed by a Strombolian eruption which
reached a height of 424 m, at a maximum altitude of 2,782 m (Fig. 2a), expulsing pyroclastic
bombs, ash, sand (tephra), incandescent material, and magma during three stages: the
formation of the cone, the opening of the adventitious cone Sapichu, and the release of lava
flows (Foshag and Gonzalez, 1956; Pioli et al., 2008). It spewed a total of 1.4 million m3 of lava
over 24.5 km2 (2,450 ha) (Wilcox, 1954).

Due to the geomorphological characteristics of the terrain where the Paricutín volcano emerged,
the first lava flows were concentrated on the southern and southwestern cone faces, while the
later lava flows advanced down the slope towards the northern and eastern cone faces,
destroying the towns of Paricutín and San Juan Parangaricutiro (Krauskopf, 1948). The area
covered by tephra was 233 km2 (23,300 ha), with an average depth of 25 cm, while on the
slopes and in the valleys, it reached up to 3 m (Inbar et al., 1994).

The climate is humid subtropical with summer rains, an average annual temperature of 15 C,
and average annual rainfall of 1,500 mm (Medina-García et al., 2000). The soils of the
surrounding forests are Andosols, which were covered by a layer of tephra (Siebe et al., 2003).
The dominant vegetation is pine-oak forest, with a high degree of human disturbance (Fregoso
et al., 2003). A floristic richness of 108 families, 306 genera, and 716 species was recorded in
the mature forests, of which 12 are pine species and two fir species, including the endangered
Pinus martinezii Larsen (ocote pine) and Abies flinckii Rushforth (Jalisco fir) (Medina, 2003).
Currently, the volcanic cone and the land south of the lava flows belong to the Caltzontzin
indigenous community, of Purepecha origin, which has owned these lands since 1599 by a royal
decree during the colonial period, currently managing a surface of approximately 6, 700 ha
(Corona-Chavez, 2018).

2.2. Dendrochronological sampling

By analyzing a remote-sensing image from Google Earth Pro version 7.3.4.8248 (Google, 2019),
16 sampling plots were established in April 2019 along three transects (western, southwestern,
and southern) with sampling plots every 250 m from the forest fragments to the volcanic cone.
The western transect had a length of 1,250 m (6 plots), 1,000 m in the southwestern (5 plots),
and 1,000 m in the southern transect (5 plots) (Fig. 1). In each sampling plot, the presence of
conifers was verified and the elevation, slope-aspect, and slope of the terrain, substrate
condition (sandy or rocky), and the density of coniferous trees in a circular area of 100 m2 were
recorded.

The elevation of the plots ranged from 2,554 to 2,672 m, starting from the edge of mature forest
fragments (Fig. 2a), passing through petrified magma and sandy lands (Fig. 2b) until the highest
point with the presence of conifers at the base of the volcanic cone (Fig. 2c). Seven plots had a
zenith position, while four of them were North-facing, three were Eastfacing, and two were
South-facing. The slope of the plots ranged from 0 (flat surface) to 45 (steep slopes). Nine plots
had a rocky substrate, four had a sandy substrate; the rest had a mixed substrate. The density
of coniferous trees varied between 6 and 28 trees⋅(100 m2 ), with an average density of 16.7
trees⋅(100 m2 ) (Table 1). No evidence of pastfires, or trees affected by pests was detected. No
broadleaf tree species were recorded.

In each sampling plot, the closest 25 conifer trees with a basal diameter of 5 cm were selected.
The species were identified and their basal diameter was measured (cm). Then, with a Pressler
increment borer, 1 to 2 radial cores were extracted per tree, depending on whether the pith was
reached in the first or second attempt. Following Speer (2010), radial cores were collected from
the stem base, as close as possible to the ground, to obtain the tree age (Fig. 2d). In total, radial
cores from 400 individuals were sampled.

2.3. Growth ring cross-dating

In the laboratory, the increment cores were dried, glued to wooden mounts, and gradually
polished with coarse to fine-grit sandpaper. The age of establishment was obtained by cross-
dating the growth rings under a microscope (Stokes and Smiley, 1996). Skeleton plots were
used to compare the sequence of narrow rings with a regional master chronology obtained from
an area 10 km south of the Paricutín volcano of a population of trees from Abies religiosa
(Kunth) Schltdl. et Cham (sacred fir) (Cerano-Paredes et al., 2014). This procedure allowed the
identification of micro-rings, false rings, and missing rings (Speer, 2010).

2.4. Composition of forest community

The importance value index (IVI) for all the sampled species was estimated, which indicates the
importance of a species within a stand, obtained from the sum of relative frequency (1), relative
density (2), and relative dominance (3) of the species according to the basal area (4) (Curtis and
McIntosh, 1951). The IVI (5) of each species was calculated as follows: (formulas)

Once the importance value of each species was obtained, it was standardized to know its
relative importance value (RIV), according to the following formula: (formula)

2.5. Age structure

The years of tree establishment were grouped and charted according to the sampling plot and
the distance to the volcano. Pulses of regeneration were identified when four or more trees
established in the same year, followed and preceded by 5-year periods without recruitment. A
continuous establishment pattern was defined when there were no periods without recruitment.

The influence of distance to the volcano, elevation, transect, slope, substrate, plot tree density,
tree basal diameter, and the species of the aged trees was evaluated, as well as the relationship
between the distance to the volcano, the number of species and the tree density. First, the
normality of distribution data was assessed through Shapiro-Wilk tests, in which tree age (W ¼
0.94274, p > 0.05), elevation (W ¼ 0.89602, p > 0.05), and tree density (W ¼ 0.93614, p > 0.05)
followed a normal distribution, hence these variables were analyzed through linear regressions
(lm function; stats package), analysis of variance (aov function; stats package) and analysis of
covariance (aov function), with a confidence level of 95%. Since slope (W ¼ 0.79246, p < 0.05)
and basal diameter (W ¼ 0.90984, p < 0.05) were not distributed normally, we used the non-
parametric mag function (mcvg package) to analyze their relationship with tree age. These tests
were run in the R program version 3.4.3 (R-project Core Team, 2017).

An arrival map of the trees was generated according to the tree with the oldest establishment
year for each plot, and the colonization year was interpolated for the sampled area using the
inverse distance weighting technique. Also, an accumulation curve of the individuals established
by species was performed, which allowed the identification of the oldest year of establishment of
each species. The order of arrival of the species was evaluated using a sequence randomness
test with the runs.test function (snpar package) (R-project Core R Core Team, 2017).

2.6. Relationship between establishment and climatic factors

The relationship between tree establishment and inter-annual climatic conditions was assessed
through a superposed epoch analysis (SEA), in which the climatic conditions that occurred five
years before, two years after, and during the year of establishment of 5 or more trees were
compared (Brewer et al., 2016). A ring-width index (RWI) was used as a proxy of regional
precipitation, obtained from a climatic chronology elaborated from trees of Abies religiosa near
the Paricutín for the period 1970–2014 (Cerano-Paredes et al., 2014). Also, the years of the
establishment were compared with the El Nino Southern Oscillation Index ~ (ENSO) during the
period 1970–2012 (Cook, 2000; NOOA, 2017), and with a local reconstruction of the Palmer
Drought Severity Index (PDSI) that covers the period 1970–2008 (Stahle et al., 2012).

The SEA was divided into two periods (1970–1994 and 1995–2015) because after 1995 the
number of established individuals increased compared with the first period. A bootstrapping
technique was used with 1000 iterations and 95% confidence intervals, in the software FHAES
version 2.0.2 (Brewer et al., 2016). The years of the establishment were compared also
graphically, to identify whether an increase in tree recruitment was associated with wet years.

3. Results
3.1. Forest community composition
Eight pine species were collected: Pinus devoniana Lindley (Michoacan pine), Pinus herrerae
(Herrera pine), Pinus lawsonii Schltdl. et Cham. (Lawson's pine), Pinus martinezii, Pinus
montezumae Lambert (Montezuma pine), Pinus oocarpa Schiede ex. Schtldl. (Mexican yellow
pine), Pinus pseudostrobus Lindley (false white pine) and Pinus teocote Schiede ex. Schtldl.
(Aztec pine). Only one fir species was found: Abies flinckii. Data on frequency, density and mean
basal area of each species can be found in Table S1. The species with the highest relative
importance were, in descending order, Pinus pseudostrobus, Pinus montezumae, Pinus
devoniana, and Pinus teocote (Table 2). Moreover, Pinus pseudostrobus and Pinus montezumae
had the highest relative dominance according to the basal area, while Pinus devoniana and
Pinus teocote had a high relative density. Pinus martinezii, Pinus herrerae, and Abies flinckii
were the species with the lowest relative importance value.

3.2. Age structure

The establishment of the conifer trees began in 1970, whereas the youngest sampled tree
established approximately in 2015, which was found at the volcano base. The general pattern of
the tree establishment has been continuous since there were only 4 plots with peaks of tree
establishment in the same year, in which abundant tree establishment occurred in 2001 (Fig.
3a), 2003 (Figs. 3e), and 2012 (Fig. 3j and Fig. 3n). In the rest of the plots, we observed that
once the first individual was established, the following trees were established continuously, but
not always in consecutive years. For example, in the plots located 750 and 500 m from the
volcano in the western transect (Fig. 3g and Fig. 3j), we observed that there were 4 and 6 years
between one established tree and the next one.

A clear spatial pattern was not found, suggesting that the oldest trees were found in the plots
farthest from the volcano, but there was a large variation in the year of the arrival among the
sampling plots (Fig. 4). The oldest tree was established in 1970 at the 250-m plot in the western
transect. In this transect, the most recent arrival year occurred at the 1000-m plot, whereas, at
the volcano base, the establishment occurred in 1985. In the southwestern transect, this pattern
was not very clear either, and, even in the plot farthest from the volcano (1,000 m), the
colonization did not occur until 1990. In the southern transect, the plots closest to the volcano
had more recent years of establishment. The relationship between the age of establishment and
the distance to the volcano, however, was not significant (p > 0.05), nor was it affected by the
identity of the transect (p > 0.05) (Fig. 5a).

Elevation did not influence the age of tree establishment (p > 0.05) and was not influenced by
the transect (p > 0.05) (Fig. 5b). However, as the slope of the terrain increased, the tree age
decreased (df ¼ 399, t ¼ 4.07, p < 0.05), although with a low determination coefficient (r 2 ¼
0.04) (Fig. 5c). The age of establishment also differed significantly with respect to slope aspect
(df ¼ 3, F ¼ 94.6, p < 0.05) (Fig. 5d), soil condition (df ¼ 2; F ¼ 22.73, p < 0.05) (Fig. 5e) and
the conifer species (df ¼ 7, F ¼ 2.08, p < 0.05) (Fig. 5f). Age was also positively correlated with
basal diameter (r 2 ¼ 0.34, df ¼ 399, t ¼ 14.43, p < 0.05), then trees with a large diameter
represent older individuals (Fig. 5g). The number of species increased significantly as the
sampling plots approached the volcano base (df ¼ 4, F ¼ 9.42, p < 0.05) with a correlation
coefficient of r 2 ¼ 0.62 (Fig. 5h). Tree density decreased with the greater distance from the
volcano (df ¼ 399, t ¼ 1.97, p < 0.05) but with a very low correlation (r 2 ¼ 0.007) (Fig. 5i). More
details on these analyses can be found in Table S2.

The most dominant species were the first to establish after the Paricutín eruption: Pinus
pseudostrobus in 1970, Pinus montezumae in 1974, Pinus devoniana in 1977, and Pinus
teocote in 1980. This pattern was very similar within the first decade in the western (Fig. 6a) and
southern (Fig. 6c) transects, whereas tree establishment in the southwestern transect began
until 1980 (Fig. 6b). Pine species with less relative abundance (Pinus herrerae, Pinus lawsonii,
and Pinus martinezii) were also the last to become established between 1989 and 1995 (Fig.
6d). The order of arrival of tree species, however, was completely random (p > 0.05) in the three
transects (Fig. 6a‒6c) and after grouping the data from all transects (Fig. 6d).

The individual accumulation curves indicated that between 1970 and 1995, there was a period of
slow recruitment for the populations of all tree species, except for Abies flinckii, which arrived in
the community by 1980 in the southwestern transect and its population has not grown since
then. Starting in 1995, the populations of the dominant species increased steadily at an annual
rate of 12.9%, until reaching an apparent period of relative stabilization in 2010 (Fig. 6).

3.3. Establishment and climate variation

The conifer recruitment was significantly associated with dry conditions during the establishment
year in the period 1970–1994 (Fig. 7a), but not in the period 1995–2015 (Fig. 7b). There was no
significant association with the occurrence of the warm or cold phase of the ENSO before or
during the year of establishment of the conifers in the period 1970–1994 (Fig. 7c), but in the
period 1995–2015, there were wet conditions, although not significant, related to the cold phase
of the ENSO (Fig. 7d). The climatic conditions related to the PDSI did not influence the tree
establishment in the period 1970–1994 (Fig. 7e), however, during 1995–2015, tree
establishment was related to wet conditions (Fig. 7f).

The graphical comparison of the three evaluated indices and tree establishment confirmed that,
as indicated by the SEA, during the first period the establishment of new trees occurred during
dry years, while in the second period, the new individuals were established both in wet and dry
years (Fig. 8a). During 1995–2015, tree establishment was relatively influenced by the cold
phase of the ENSO, when conditions were more humid (Fig. 8b). Finally, it was observed that
during a prolonged phase of humid conditions between 2000 and 2008, according to the PDSI,
there was continuous and abundant recruitment in the community (Fig. 8c).

4. Discussion
The conifer species have shown a great ability to establish in volcanic substrates, even a few
years after the Paricutín eruption. Although Pinus montezumae seedlings had been detected 8
years after the eruption (Eggler, 1963), the oldest living tree in our sample (Pinus
pseudostrobus) was established 10 years later (1970), which means that the suitable conditions
for seed germination and conifer establishment was reached until that time, which allowed
seedlings to survive and grow (Dale et al., 2005). Currently, the conifer populations are growing,
probably becoming a young forest or reaching a mid-seral phase, where recruited saplings are
derived from colonizer trees (Powell, 2012).

The beginning of colonization was slightly longer than in other volcanic substrates. For example,
Pinus hartwegii Lindley (Mexican mountain pine) took 10 years to establish in lahars of the
Popocatepetl volcano in Mexico (Franco-Ramos et al., 2019), Pseudotsuga menziesii Franco
(Douglas-fir) arrived after 14 years in avalanche deposits near Mount St. Helens, in the United
States (Pierson, 2007), whereas Larix cajanderi Mayr (Dahurian larch) appeared after 6 years in
lahars at the Shiveluch volcano, Russia (Salaorni et al., 2017). It is important to mention that
conifer arrival would be expected to take longer in lava flows than in lahars because igneous
rocks are very porous, retain little water, and are difficult for roots to penetrate (Vilmundardottir et
al., 2018).

The rapid colonization of conifers in deposits of tephra and volcanic lavas of Paricutín can be
explained by the presence of “safe sites”, where microclimatic and edaphic conditions gradually
allow tree establishment (Cutler et al., 2008). The number of safe sites increases due to the
incorporation of ash and tephra in the lava flows during the eruptive phase (Del Moral and
Grishin, 1999), as well as the entry of soil particles from sites with preserved vegetation through
hydric or wind processes, which together form a suitable substrate for the growth and
development of tree roots (Deligne et al., 2013). Once the first plants are established, biological
weathering and the mechanical breaking of the rocks by the roots improve the edaphic
conditions (Moldaway and Sparrow, 2006).

In the case of Paricutín, the lava flows located towards the south and southwest of the cone
were covered by ash and tephra during the eruptive period, while subsequent lava flows were
covered by other layers of lava (Inbar et al., 1994). It may explain why primary succession has
been concentrated in the oldest lava flows, while the rest of the area covered by younger lava
remains devoid of arboreal vegetation (Lindig-Cisneros et al., 2006), which is similar to that
reported for the El Jorullo volcano, where the oldest flows covered by tephra also show more
vegetation (Deligne et al., 2013). Moreover, in the areas covered by tephra, erosion has played
an important role in reducing the depth of this material (Inbar et al., 1994) since a depth greater
than 30 cm of tephra significantly limits the establishment of conifers (Gomez-Romero et al.,
2006 ).

In volcanic substrates, there is a strong relationship between tree age and distance from the
preserved forests (Del Moral et al., 2005). However, the nearest trees to the Paricutín cone are
not necessarily the youngest, which may be associated with the wind-dispersion of conifer
seeds, increasing their probability to reach distances up to 120 m from the source of propagules
(McCaughey et al., 1986). In the case of the species found at the Paricutín volcano, all of them
release winged seeds, dispersed through the wind (Farjon et al., 1997). Besides, the arrival of
conifers was not directional, but random, which is common in sites with primary succession
(Garibotti et al., 2011). Nonetheless, the four species that colonized first currently have greater
abundance and basal area, which is consistent with the trend in which the first arriving species
tend to dominate the succession (Powell, 2000). Also, these species have fast growth rates
(Klepac, 2001), of which Pinus pseudostrobus and Pinus montezumae are the dominant species
in the nearby forest fragments (Fregoso et al., 2003).
The conifer community of the Paricutín volcano experienced an expansion process between
1995 and 2010. In this mid-seral phase, the seed bank might have begun to be fundamentally
local, whereas the external sources of propagules decreased in importance (Del Moral et al.,
2005). Additionally, completely stochastic processes, such as the dispersal of seeds from
external sources, cease to operate, while deterministic processes, such as competition and
facilitation, are more relevant (Cutler et al., 2008). Currently, only 70 years after the Paricutín
eruptive period, there are already well-developed conifer stands, which is faster than in other
volcanic sites recorded outside of the tropics, for example, 600 years in Mount Hekla, Iceland
(Vilmundardottir et al., 2018 ), and 200 years in the Cascade Range, United States (Deligne et
al., 2013).

The climate variability had a low influence on the establishment of conifers in the Paricutín
substrates since trees were recruited in wet and dry years. This pattern has also been observed
in other Mexican conifer forests (Saenz-Ceja and Perez-Salicrup, 2020), which contrasts with
the strong correlation between wet years and conifer recruitment in the semi-arid northern
Mexico (Fule and Covington, 1997). On the one hand, rainfall regimes in tropical montane
forests, where the Paricutín is located, may not be as contrasting as extra-tropical regions, so
differences in tree recruitment during wet and dry years might not so clear (Astudillo-Sanchez et
al., 2017). On the other hand, amelioration of substrates properties by a litter layer and biological
weathering could have made suitable seed germination, even in dry years (Crisafulli et al.,
2005).

In terms of ecological restoration, the fast establishment of conifers around the Paricutín
suggests that the regeneration of tropical Mexican conifers can occur rapidly after high-
magnitude disturbances, even in primary succession scenarios. Indeed, pine establishment
began only two years after large fires in mixed pine-oak forests (Juarez-Martínez and
Rodríguez-Trejo, 2003) and seven years after pyroclastic flows at the Popocatepetl volcano
(Franco-Ramos et al., 2019). At later seral stages, the basal area can be similar to that of mature
pines only 28 years after high-severity fires (Quintero-Gradilla et al., 2019). It suggests that
forest succession could be left in place without human intervention, as long as the sources of
propagules are close to the affected areas and the soil conditions allow the establishment of
seedlings (Angeles-Cervantes and Lopez-Mata, 2009 ).

The Paricutín volcano represents a unique site for characterizing the primary succession after a
high-magnitude disturbance. It is important to mention that disturbances such as fires or insect
infestations have remained absent, but when they occur they could interrupt, reverse, and alter
the successional trajectories (Riff et al., 2003). The establishment of conifers also skipped
different stages of succession (the previous colonization by shrubs and later by broadleaf tree
species), as has been recorded in some glaciers (Garibotti et al., 2011), which underscores the
great adaptive capacity of conifers that inhabit the Paricutín region, within a region that host a
high species diversity, including those of the genus Pinus (Medina, 2003). More research is
needed on the successional dynamics after these disturbances, on the most recent established
saplings after 2015, and the assessment of the influence of topographic and climatic variables,
as well as the implications for ecological restoration and forest management.

5. Conclusions
The conifers around the Paricutin volcano have shown a great ability for colonizing substrates
after 70 years of ending its eruptive period. The arrival of conifers began only 18 years after the
last eruption in 1952, a process that has been random, unrelated to the distance from preserved
forests, and not influenced by climate variability. The analysis of age structure was useful to
identify a continuous establishment pattern, with a progressive increase of the population since
1995, followed by a population stabilization process from 2010. It is important to document the
colonization of tree species in the more recent lava flows, where tree regeneration is low, and to
include the most recent established saplings. It is also necessary to evaluate the ability of
tropical coniferous species to regenerate in sites subjected to high-magnitude disturbances,
information that could be useful to guide ecological restoration strategies. This study is a pioneer
in applying dendrochronology to lava flows from recent volcanic activity, so it is recommended to
replicate this type of study in both recent and ancient volcanic substrates, where successional
patterns with annual resolution could be characterized.