Introduction
Over more than a century a widespread process of land use change has
taken place worldwide, by which native ecosystems, mainly forests, have
been replaced by agricultural fields (Lambin & Meyfroidt, 2011; Runyan
& D’Ordorico, 2016). In tropical regions, where swidden agriculture is
often practiced, agricultural fields are usually abandoned after a few
years of use due to a reduction in their productivity and yield (Aide et
al, 2000). After land abandonment, a process of secondary succession
starts, leading to vegetation cover recovery (Chazdon, 2014; Letcher et
al., 2015). If undisturbed, the community is capable of reaching a
relatively stable state (Poorter et al. 2021; Muñoz et al. 2021).
Globally, around half of all tropical forests are secondary communities
which are recovering from previous disturbances (FAO, 2015), and this
figure continues to increase as many native forests are being cleared,
replaced and abandoned as the agricultural frontier continues to expand
(Meyfroidt & Lambin, 2011).
A common pattern consisting of three stages has been recognized in
tropical forest secondary succession (Finegan, 1996; Lebrija-Trejos et
al., 2011; Mora et al., Trilleras, 2015; Rozendaal et al., 2016;
Martínez-Ramos et al., 2021; Rüger et al., 2023). The first stage is
characterized by the dominance of weeds, shrubs, and climbing plants.
Next, the establishment of fast-growth, short- and long-lived trees
characterizes a second stage, which eventually gives way to a third
stage characterized by the establishment and development of the
seedlings of slow-growing long-lived tree species under the canopy of
the fast-growing trees. Those species that establish in early succession
(pioneers) can either display a fast demographic strategy (grow,
reproduce and die quickly) or have a longer lifespan (long-lived
pioneers), while mature forest species, with a slow demographic
strategy, establish later in succession (Rüger et al., 2023). The
differential presence and abundance of species in each of the three
stages is determined by the prevailing environmental conditions at each
successional stage, which in turn are driven by a feedback dynamics
between the environment and the community (Wallace & Romney, 1980;
Lebrija-Trejos et al. 2011; Letcher et al., 2015; Matsuo et al. 2021).
Therefore, the presence of certain pioneer species may facilitate or
hinder the performance of other species depending on how they modify the
environment. Ultimately, dominant pioneers play a substantial role in a
community’s successional dynamics and ecosystem functioning (Grime
1998).
Several studies have focused on secondary succession in tropical dry
forests (Kennard, 2002; Peña-Claros, 2003; Lebrija-Trejos et al., 2010a;
Lebrija-Trejos et al. 2010b; Dupuy et al., 2012; Pineda-García, Paz &
Meinzer, 2013; Mora et al., 2015; Rozendaal et al., 2016). In these
systems, the most important limiting factor is water availability
(Murphy & Lugo, 1986; Allen et al. 2017), which entails low plant
growth rates (Rozendaal et al., 2016), and also explains why succession
in these forests is often dominated by very few drought-tolerant species
(Ceccon et al., 2006; Markesteijn, Poorter, Paz, Sack & Bongers, 2011;
Rozendaal et al., 2016). In particular, in the tropical dry forest of
the Nizanda region, southern Mexico, Lebrija-Trejos, Bongers,
Pérez-García, and Meave (2008) found a three year-long initial
successional phase characterized by shrub growth. After this stage,
early successional Mimosa acantholoba var. eurycarpa trees
become dominant over a few decades, creating a canopy layer that
provides shade and reduces the air temperature in the forest understory,
apparently allowing mature forest species to establish and eventually
dominate late-successional stages.
Previous studies established M. acantholoba var.eurycarpa ’s dominance, based on its high basal area and density
of individuals compared to other species of the tropical dry forest of
Nizanda (Romero, 2014). However, little is known about the traits that
underlie its dominance; although initially its thin leaves and high leaf
dry mass content were proposed (Romero, 2014), a recent study showed
that wood anatomical traits in this species ensure efficient water
conduction in the rainy season while reducing its hydraulic
vulnerability in the dry season (Romero et al., 2020). Nevertheless, it
is likely that some other aspect of its life cycle, such as its
population dynamics, may be also involved in this pattern.
An important demographic process of pioneer species population dynamics
in early succession is the entry of new individuals to the population.
We name these new individuals recruits when they derive from seed
germination and seedling establishment. Alternatively, there are
individuals entering the population through the resprouting of
vegetative structures (stumps or roots) that remain alive in the sites
after the agricultural use of the field; we name these resprouts. The
absence of reproductive individuals in early succession implies that its
population dynamics necessarily depend on processes other than
reproduction from mature local trees. These processes encompass seed
dispersal from the surrounding forest matrix (Dent & Estrada-Villegas,
2021), the presence of a seed bank or the resprouting of remnant
vegetative structures (Purata, 1986; Guevara & Laborde, 1993; Bartha,
Meiners, Pickett, & Cadenasso, 2003; Vieira & Scariot, 2006). In the
case of M. acantholoba var. eurycarpa , no seed bank has
been found in the Nizanda tropical dry forest (Meave et al. 2012; Mena,
2009). Further, although seed rain was recorded for this species at the
study site, the amount of seed produced was very low relative to the
number of trees in the plots (Cervantes, 2018), suggesting that seed
dispersal may only play a minor role in the establishment of aMimosa -dominated canopy. What is presently known is that M.
acantholoba var. eurycarpa is the species that displays more
resprouts in the early successional community (Lebrija-Trejos, 2004).
These resprouts, being live tissues produced by previously existing
individuals, usually with a well-developed root system, are able to grow
and reproduce rapidly once anthropogenic disturbances stop, and
represent a pathway for the increase in the species’ representation in
the community. Hence, acknowledging the contribution of resprouting may
help better understand M. acantholoba var. eurycarpa ’s
population dynamics as a whole.
To gain insights into the role that population dynamics plays in the
dominance of a pioneer species in a tropical dry forest, in this study
we characterized and modeled the dynamics of M. acantholobavar. eurycarpa along a successional gradient. We did this by
modeling this species’ vital rates and their changes over succession
using data from permanent plots originally established to represent a
chronosequence, but then monitored yearly over 13 years. We then
assembled these models into an integral projection model (IPM) that
accounted for both resprouting and recruitment, and estimated transient
population growth rates. We predicted that, at the onset of succession,
the environmental conditions would promote M. acantholoba var.eurycarpa ’s performance in the successional plots. Therefore, the
population size of this species should increase during the early years
of succession and decrease in later stages due to changes in
environmental conditions in the forest interior.