A Review of The Structure and Dynamics of Araucaria Mixed

NEW ZEALAND JOURNAL OF BOTANY
2021, VOL. 59, NO. 1, 2–54

https://doi.org/10.1080/0028825X.2020.1810712
REVIEW ARTICLE
A review of the structure and dynamics of araucaria mixed

forests in southern Brazil and northern Argentina
Alexandre F. Souza
Departamento de Ecologia, CB, Universidade Federal do Rio Grande do Norte, Campus Universitário, Natal,
Brazil
ABSTRACT
The Araucaria Mixed Forests (AMF) are subtropical conifer-hardwood
mixed forests that are part of the Atlantic forest floristic dominion in
Brazil and Argentina. The AMF is dominated by the conifer
Araucaria angustifolia (hereafter araucaria), accompanied by a
diversity of angiosperms. Its distribution largely corresponds to the
13°C isotherm and is associated with overall lower temperature,
frost-prone, and wetter climates than neighboring vegetation types.
Palynological research and bioclimatic modelling have yielded Alexandre F. Souza
somewhat conflicting results but suggest intense forest-grassland
dynamics, with grasslands possibly dominating the landscape ARTICLE HISTORY
during the drier and colder last glacial maximum. Araucaria is a Received 3 June 2020
cultural keystone species whose distribution has been greatly Accepted 11 August 2020
fostered by Amerindian activity in the last few millennia.
Angiosperms form a lower layer beneath araucaria canopies, and HANDLING EDITOR
they seem to be geographically structured into a wetter and colder Lohengrin Cavieres
eastern and a drier and hotter western portion. Araucaria attain
KEYWORDS
large sizes and high longevity but is unable to regenerate under the Araucaria angustifolia;
shade cast by angiosperm canopies and need large-scale Atlantic forest; Brazil; climate
disturbances for successfully complete their life-cycles. This cycle change; forest structure;
has been synthesised in the non-equilibrium Lozenge model of forest dynamics; functional
forest dynamics. Regeneration opportunities arise from the frequent traits; logging; community
multiple treefall gaps produced by the tornado and windstorm- composition; subtropical
prone climate of subtropical South America and by the grazing-, forests
fire-, and nurse-plant successional dynamics of the forest-grassland
mosaic. Dispersal is mediated by a complex web of interactions with
birds and mammals. A review of the dynamics of southern Oceanic
Temperate Forests indicates that the dynamics of the AMF fits a
prevailing pattern in these forests that corresponds to the Lozenge
model. Deforestation and degradation by logging and cattle
grazing have impacted most of the AMF original range. Similarly to
what happened to Agathis australis in New Zealand, the
establishment of commercial plantations still need incentives and
research to thrive and produce large-scale environmental benefits.
Introduction
The Araucaria angustifolia mixed forests (AMF) are subtropical conifer-hardwood mixed
forests that are part of the Atlantic forest floristic dominion in South America. They
CONTACT Alexandre F. Souza alexsouza.cb.ufrn.br@gmail.com

© 2020 The Royal Society of New Zealand
NEW ZEALAND JOURNAL OF BOTANY 3
covered originally ca. 25 million ha in southern Brazil (Ribeiro et al. 2009) in the states of
Paraná, Santa Catarina, and Rio Grande do Sul and the Argentinian province of Misiones
(longitude ca. 54°30′ W), and disjunct patches in tropical latitudes in southeastern Brazil
(São Paulo, Rio de Janeiro, and Minas Gerais states, latitude ca. 19°15′ S, Figure 1). The
AMF is characterised by at least 50% of canopy cover by the conifer Araucaria angustifolia
(Bertol.) Kuntze (Araucariaceae, hereafter araucaria), accompanied by a diversity of
angiosperms, although the conifer Podocarpus lambertii Klotzsch ex Endl. (Podocarpa-
ceae) may also be abundant (Oliveira-Filho 2009). Along the twentieth century,
growing economic activity and land development led to the widespread deforestation of
the AMF, led by timber logging of araucarias and by the replacement of native forests
by agriculture, pasturelands, Pinus and Eucalyptus plantations, and urban expansion
(Cabral and Cesco 2008; Matos et al. 2016; Nodari 2016). Currently the AMF is
reduced to 12.6% of its original range (Ribeiro et al. 2009), of which a mere 0.7% can
be regarded as well-conserved old-growth forest (Nodari 2016). This situation led arau-
caria to be regarded as a critically endangered species by the IUCN (Thomas 2013).
The current threatened status of the species, along with its long evolutionary history led
researchers to classify araucaria in the third place among the most evolutionarily distinct
and globally endangered (EDGE) gymnosperm of the planet, having lost ca. 97% of its
original range (Forest et al. 2018).
In the present contribution I review the existing knowledge on the structure and
dynamics of the AMF. As shown by Grubb et al. (2013) for different warm temperate
forests around the world, the ecology of the same forest type can present many differences
and particularities between different regions of the planet and therefore a review of the
Figure 1. Left: Distribution of the Araucaria mixed forest in South America. Right: Elevation variation in
southeastern and southern Brazil and northern Argentina. Country acronyms are in bold: ARG = Argen-
tina, BRA = Brazil, and PRY = Paraguay. Acronyms of Brazilian states are in light grey: RS = Rio Grande
do Sul, SC = Santa Catarina, PR = Paraná, SP = São Paulo, MG = Minas Gerais, MS = Mato Grosso do Sul.
Mountain acronyms in the image to the right: SG = Serra Geral, SMan = Serra da Mantiqueira, SO =
Serra dos Órgãos, and SMar = Serra do Mar. Note that all named mountain chains (Serra) are parts
of the broadly north-south coastal Serra do Mar mountain chain. Based on the distribution of the Arau-
caria Mixed Forest found in official vegetation map of Brazil (IBGE 2004), fusing the categories Mixed
Ombrophilous Forest, Dense Ombrophilous Forest/Mixed Ombrophilous Forest transition, and Semide-
ciduous Forest/Mixed Ombrophilous Forest transition.
4 A. F. SOUZA
existing knowledge about each of them is necessary. Although I included a comparison of

the AMF dynamics with the dynamics of other southern hemisphere mixed forests in this
review (section The AMF in the context of southern mixed forests), a detailed review of the
regeneration dynamics of southern hemisphere mixed conifer-hardwood forests like those
occurring in New Zealand and Andean South America is out of the scope of the present con-
tribution and information in this respect can be found in several excellent reviews (Hill and
Brodribb 1999; Brodribb et al. 2012; Kitzberger et al. 2016; McGlone et al. 2016; Veblen et al.
2016; Wyse et al. 2018). I begin by reviewing the climatic and soil conditions in which the
AMF is found, and then review the ecological history of this system as retrieved through
palynological studies. I proceed to summarise the estimated effects of Amerindian activity
on the distribution of the araucaria, and then move on to review the floristic distribution of
the angiosperm species that accompany the araucarias along the AMF. The regeneration
dynamics of the AMF is then conceptually presented in the framework of the Lozenge
model and the life-history ecology of long-lived pioneers. I then review the disturbance
regimes that prevail in subtropical eastern South America landscapes, with emphasis on cli-
matic mediated disturbances and on fire- and livestock grazing-mediated disturbances. In
this regard I placed special emphasis on the forest-grassland dynamics, which is one of the
main contemporary scenarios for the expansion of the AMF. Seed production and dispersal
is then reviewed aiming at examining the role of interactions in the dynamics of this forest
system, before moving to open questions and the future of the AMF research.
Abiotic conditions and neighboring vegetation types

Across its range, the AMF is mostly found in mountainous habitats, covering most of the
Serra Geral highlands, the southern portion of the Serra do Mar, and the highest parts of
the Serra da Mantiqueira, Serra dos Órgãos and other surrounding highlands (Oliveira-
Filho et al. 2015) (Figure 1). Its distribution largely corresponds to the 13°C isotherm
(winter July average temperature) (Backes 2009) and is associated with overall lower temp-
erature, frost-prone, and wetter climates than neighboring vegetation types (Ntima 1968;
Oliveira-Filho et al. 2015; Wrege et al. 2016). This leads the AMF to occupy progressively
lower altitudes with increasing latitude. Examples are the Serra da Mantiqueira (22°S)
where the AMF occurs above ca. 600 m (Santiago et al. 2018), and the Serra do Sudeste
highlands (30°S) where it occurs above ca. 400 m (Behling et al. 2016). The regions occu-
pied by the AMF frequently experience near or below 0°C absolute winter minimum
temperatures with frosts whose frequency may vary from one to 56 frost days per year
(Backes 2009). A requisite for the AMF maintenance seems to be the occurrence of at
least three consecutive winter months with average minimum temperatures ≤ 10°C and
a humid Cfb climate (Alvares et al. 2013) with no dry season and at least 1250 mm of
annual rainfall (Ntima 1968; Backes 2009). In the Paraná state, localities with araucaria
present higher average altitude (824 m versus 512 m) and winter rainfall (323 mm
year−1 versus 241 mm year−1) and reduced evaporation (883 mm year−1 compared to
1209 mm year−1) and insolation (2200 h year−1 compared to 2426 h year−1) than localities
without the species (Fritzsons et al. 2018). Within the overall cool and wet range in which
Araucaria angustifolia and the AMF occur there is an east–west rainfall gradient that
peaks along the mountain ranges that accompany the coast and declines westwards
(Figure 1) (Oliveira-Filho et al. 2015; Fritzsons et al. 2018).
The main AMF block in southern Brazil and its neighboring vegetation types occupy
distinct climatic and edaphic environments. The AMF is replaced by seasonal semidecid-
uous forests to the west along a relatively long thermal gradient, in which higher records of
maximum temperature are likely to restrict the occurrence of many cold-adapted species
including araucaria (Jarenkow and Budke 2009; Oliveira-Filho et al. 2015; Sevegnani et al.
2016). Eastwards the abrupt altitudinal change from the sea level to the Serra Geral and
Serra do Mar highlands ca. 1000 m asl produce short and marked temperature changes
that promote the replacement of the AMF by the coastal rainforests, while southwards
the AMF and semideciduous forests are replaced by the Campos grasslands (Gonçalves
and Souza 2014; Oliveira-Filho et al. 2015). The floristic identity of the AMF is also delim-
ited by increased levels of soil nutrients and aluminium content, and a reduced proportion
of topsoil sand (Gonçalves and Souza 2014; Oliveira-Filho et al. 2015; Cantidio and Souza
2019). Besides floristic differences, the classification of Southern Brazilian Atlantic forests
into rainforests, AMF, and semideciduous has a phylogenetic basis, since phylobetadiver-
sity analyses showed that the AMF differed more in relation to rain and semideciduous
forests than these latter from each other (Duarte et al. 2014). The AMF presents high phy-
logenetic diversity/overdispersion between sites (Duarte et al. 2014), and this is attribu-
table to the importance of temperate conifers like Araucaria and Podocarpus, and to
the frequency of magnoliid genera like Drimys and Cinnamodendron (Rambo 1951).
Mainly in its easternmost parts, where the southern highlands approach the Serra do
Mar and Serra Geral mountain ranges and where the AMF is nearer to the outreaches
of the lowland rainforests, tropical Myrtaceae becomes a frequent eudicot group in the
AMF (Duarte et al. 2009; Higuchi et al. 2012).
A brief recent ecological history of the araucaria mixed forests

Unlike many other tropical and subtropical ecosystems, our understanding of the recent
ecological history of the AMF has been greatly fostered by extensive research in the last
two decades. This sets the stage for the interpretation of the current community and popu-
lation dynamics of this vegetation type (McGlone et al. 2016). See Dutra and Stranz (2009)
for a review of the evolution and palaeoecological history of the Araucariaceae family.
Confirming Klein’s (1960) earlier claims, palynological records indicate that extensive
areas of southern and southeastern parts of Brazil were covered by open grasslands
during most of the time in the last 70000 years, including the Last Glacial Maximum
(Behling 1995; Ledru et al. 1996; Behling 1997; Behling and Lichte 1997; Ledru et al.
1998; Behling and Pillar 2007; Bauermann and Behling 2009; Leonhardt and Lorscheitter
2010; Jeske-Pieruschka and Behling 2012; Jeske-Pieruschka et al. 2013; Scherer and
Lorscheitter 2014; Gu et al. 2017).
The prevalence of grasslands has been linked to cooler and seasonally drier (semiarid)
climates, that probably hindered the establishment of araucaria populations and forest for-
mations in most areas since the last glacial maximum except for scattered sites in the path
of polar advection (Ledru et al. 1998; Leonhardt and Lorscheitter 2010) or on coastal
slopes and deep valleys of mountainous regions with sufficient moisture (Behling 1995;
Ledru et al. 1996; Ledru et al. 1998; Behling et al. 2004; Bauermann and Behling 2009;
Scherer and Lorscheitter 2014). This scenario would have extended northwards into
what today is the Cerrado savanna or semideciduous forests (Ledru 1993; Behling and
6 A. F. SOUZA
Lichte 1997; Behling and Safford 2009). Palynological evidence does not support signifi-
cant expansion of the AMF into Atlantic lowlands (Behling and Negrelle 2001) or areas
to the north of ca 19°S during the last glaciation (Behling and Safford 2009), with the
eventual exception of restrict riparian areas in mountainous patches (Behling and
Lichte 1997).
Bioclimatic modelling of the distribution of the major Brazilian forest types during the
last glacial cycle produced a somewhat different scenario from the one described from the
palynological evidence for the spatial dynamics of the AMF. Arruda and colleagues esti-
mated a shift of the AMF c. 400 km northwards from its present distribution into the
semideciduous forest during the last glacial maximum (Arruda et al. 2018). These
authors found that when the AMF climatic suitability shifted north, the climatic suitability
for southern grassland cover also expanded northwards, replacing the AMF in its south-
ernmost range. They also found that while the palynological scenario of scattered AMF
refugia in a sea of grasslands may have held true in the Atlantic eastern border of the
Serra do Mar and Serra Geral highlands, extensive AMF was predicted to have covered
hinterland areas. Approximately the same region estimated to have been occupied by
the AMF in the last glacial maximum has been identified as historically stable in a biocli-
matic modelling that did not discriminate the AMF from overall wet forests in South
America, and was named the Southern Atlantic Forest refugium (Costa et al. 2018). A
similar result was also obtained when the climatically distinct southern part of the Atlantic
Forest was modelled separately (Carnaval et al. 2014). These findings are in agreement
with a bioclimatic model specifically calibrated for AMF species (Bergamin et al. 2019),
according to which the AMF reached its maximum extension during the LGM when it
spread to the north and west, into regions currently covered by Cerrado savanna and sea-
sonal forests. A niche modelling of araucaria predicted a refugium for the species during
the last glacial maximum in the southern parts of the current AMF range in the Serra Geral
highlands (Barros et al. 2015).
Most palynological studies have been conducted along the Atlantic border of the Serra
do Mar and Serra Geral highlands, as observed by some authors (Flantua et al. 2016; Ber-
gamin et al. 2019), and thus may have failed to register the continuous presence of the
AMF in the Southern Atlantic Forest refugium during glacial times. Current knowledge
of the past distribution of the AMF thus provides partial support to the hypothesis that
the current AMF would represent relicts of a former widespread distribution (Klein
1984) in Rio Grande do Sul and most of Santa Catarina, but supports it in Paraná.
Forest-grassland dynamics is poorly captured by bioclimatic modelling (Bergamin et al.
2019), and thus an alternative explanation to the contradiction between bioclimatic
models and palynological data would be that the Paraná Southern Atlantic Forest refu-
gium in the last glacial maximum was not homogeneous and that many small forest
refugia may have existed along with grassland patches, allowing for rapid forest recovery
under wetter conditions (Costa et al. 2018).
At the end of the Pleistocene and beginning of the Holocene, between ca. 13,000 and
10,000 years BP, palynological data indicated that conditions became hotter although
not evenly moister (Bauermann and Behling 2009) and forest expanded modestly in
montane areas of the southeastern (Behling and Safford 2009) and southern parts of
the AMF range (Behling and Lichte 1997). The earlier AMF occurrence in central Brazilian
areas retreated as climate became warmer and drier there, leading to the replacement of
the AMF by semideciduous forests (Ledru 1993; Werneck et al. 2011; Bergamin et al.
2019), with exception of sites in higher altitudes (Ledru et al. 1996). Between ca. 4000
and ca. 1000 years BP bioclimatic modelling predicted an expansion front along the south-
ernmost distribution of the AMF (Arruda et al. 2018). These results agree with the expan-
sion of Podocarpus lambertii, a species from an ancient conifer genus associated with
elevated moisture levels and with araucaria, in the southern Brazil pollen record starting
3000 years BP (Ledru et al. 2007).
In the mid Holocene, temperature and humidity increases allowed for the proliferation
of mycorrhizal activity, as attested by the presence of Glomus during the last 2000 years,
what is believed to have acted synergistically with forest expansion (Leonhardt and
Lorscheitter 2010). The proportion of pollen grains linked to common genera inhabiting
the AMF have been recorded in this period (e.g. Myrsine, Mimosa, Myrtaceae, Podocarpus,
and Ilex, besides araucaria itself) (Behling et al. 2001; Behling and Pillar 2007; Iriarte and
Behling 2007; Ledru et al. 2007; Bauermann and Behling 2009; Leonhardt and Lorscheitter
2010; Jeske-Pieruschka and Behling 2012; Scherer and Lorscheitter 2014). Stable areas that
were forest refugia in the last glacial maximum are floristically differentiated today in the
overall Atlantic Forest dominion (Cantidio and Souza 2019).
According with the palynological record, the culmination of the AMF forest expansion
under a wetter climate was the substitution of most of the grasslands by forests in the last
1000 years, following increased rainfall (Behling 1995; Behling 1997; Behling et al. 2001;
Behling et al. 2004; Behling and Pillar 2007; Leonhardt and Lorscheitter 2010; Jeske-Pier-
uschka and Behling 2012; Scherer and Lorscheitter 2014; Gu et al. 2017; Portes et al. 2018).
The AMF became the main vegetation type on the highlands although still forming forest-
grassland mosaics. Other species that today are typical of the AMF responded to the
increased rainfall at different colonisation speeds. The tree fern Dicksonia sellowiana
Hook., for example, today regularly associated with araucaria, was not apparent in the
palynological record until ca. 400 years BP (Behling and Pillar 2007; Bauermann and
Behling 2009). In the late Holocene the AMF seems to have migrated westwards, since
the palynological record registers only a very recent and low density presence of araucaria
pollen in the easternmost parts of the Misiones province in Argentina, paralleled by a raise
of other genera typical of mixed Araucaria forests like Clethra, Ilex, Myrsine and Podocar-
pus (Gessert et al. 2011).
We can conclude that the current AMF distribution results from the last of a series of
climatic-driven cycles of vegetation change and community reorganisation, being very
recently assembled communities, formed after millennia of directional retraction, expan-
sion, and reassembling of humid- and cool-adapted species (Gu et al. 2017), alternated
with several different stable periods in which vegetation dynamics seemed to be more sto-
chastic (Behling and Pillar 2007). In support to this view, it has been found that phylobe-
tadiversity between southern Brazil Atlantic forest sites is driven by more terminal and not
basal phylogenetic nodes (Duarte et al. 2014).
Genetic evidence corroborates the late Holocene migration of the AMF to its present
range in the highlands in different latitudes. In southern Brazil the genetic signatures of
Pleistocene population size reductions were lost, whereas these signatures persisted in
the naturally fragmented southeastern populations in mountaintops with increased
inbreeding (Stefenon et al. 2008; Sousa et al. 2020), limited gene flow and reduced
genetic diversity (Souza et al. 2009). Genetic results suggest long-term isolation and
8 A. F. SOUZA
rare gene flow between southern and southeastern regions and among southeastern popu-
lations themselves, which also likely suffered genetic drift (Souza et al. 2009). These evol-
utionary and genetic processes are believed to have been responsible for the high genetic
differentiation found between southern and southeastern araucaria populations with high
divergence of the southeastern populations (Sousa et al. 2004; Stefenon et al. 2007; Souza
et al. 2009; Sousa et al. 2020). A current climate niche modelling identified separate ranges
for the southeastern and southern genetic lineages, indicating that differential selection
pressure between the two biogeographical regions may have taken place and the popula-
tional differentiation may be adaptive (Sousa et al. 2020) but see Wrege et al. (2016). Selec-
tive drivers are likely related to climate and soil characteristics of the two regions (Backes
2009; Sousa et al. 2020).
Araucaria biology and the AMF as cultivated systems

In well conserved AMF, Araucaria angustifolia is the most abundant species, also account-
ing for many of the structural forest characteristics. Araucaria is a wind-pollinated dioe-
cious species with four distinctive life-stages: seedlings are unbranched and frequently
attached to seed remains, juveniles have lateral branchlets forming a monopodial crown
and may grow up to 2.5 m, immatures have woody horizontal or bending whorl branches,
while reproductive trees have minimum diameter at breast height = 10.5 cm and horizon-
tal branches, becoming flat-topped with age (Souza et al. 2008). The species forms a dis-
tinctively separated and higher canopy layer with up to 25.3 m and average 17.7 m above a
lower canopy formed by angiosperms (average 11.7 m) (Souza 2007) (Figure 2). Araucaria
comprises up to 92.1% of forest basal area (Souza 2007), and forests with Araucaria
present increased average above ground biomass (250.3 Mg ha−1) than broadleaf forests
in the same region (118.9 Mg ha−1) (Rosenfield and Souza 2014). Drought reduces seed-
ling root and shoot growth, but high levels of soil compaction (1.25 g cm−3) inhibits root
mass build up (Mósena and Dillenburg 2004). Demographic data are scant for entire
populations, but in adult plants diameter growth is up to 0.25 mm month−1 and does
not differ between male and female trees, being concentrated between the months from
Figure 2. Left: View of an adult Araucaria angustifolia canopy overtopping a lower angiosperm canopy.
Right: Trunk base of a large Araucaria angustifolia tree with surrounding forest understory. Both images
were taken in São Francisco de Paula National Forest, Rio Grande do Sul, Brazil.
September to March (spring and summer), when it is positively influenced by temperature

and rainfall increases, and is reduced under low temperatures (Zanon and Finger 2010;
Oliveira et al. 2017). Interannual diameter growth rates vary from 30 mm year−1–
50 mm year−1 (Figueiredo Filho et al. 2010; Mattos et al. 2010; Adenesky Filho 2014;
Beckert et al. 2014). In closed forests annual seedling mortality is ca. 80%, sharply decreas-
ing to 0.3% among reproductive trees (Paludo et al. 2016) (Figure 2). Among the biggest
reproductive trees (diameter at breast height ≥ 20 cm) there can be large interannual vari-
ation in mortality, ranging from 0.58% to 8.13% (Beckert et al. 2014).
Apart from increased rainfall and cold winters, there is evidence that the activities of
Amerindians greatly promoted the expansion of the AMF in the last 2000 years (Noelli
2000; Bitencourt and Krauspenhar 2006; Iriarte and Behling 2007; Reis et al. 2014; Lau-
terjung et al. 2018). Genetic analyses of chloroplast sequencing supported the hypoth-
eses that Araucaria angustifolia populations show (1) signs of recent and fast expansion,
(2) that this expansion promoted gene flow without enough time for populations to
diverge genetically, (3) that the glacial refugia modelled by Barros et al. (2015) and
suggested by palynology (Behling 1995; Behling and Lichte 1997; Behling et al. 2004)
were the source of other populations in southern Brazil, (4) and that Araucaria angu-
stifolia would not have reached its current distribution after the last glacial maximum
without human help (Lauterjung et al. 2018). The first evidence of human activities
in the southern range of the AMF is the appearing of charcoal fragments ca 7400
years BP, indicating the beginning or the increase in the use of field burning by indi-
genous peoples (Bauermann and Behling 2009). However, the denser and more perma-
nent human occupation of southern Brazil was promoted by the advance of the AMF,
which provided indigenous peoples with Araucaria seeds, a newly and abundant
resource (Iriarte and Behling 2007). The seeds of araucaria contain abundant starch
(37%), dietary fibre, cupper, and magnesium. They are also a source of protein
(ca. 3%) and lipids (ca. 1.3%), in comparable amounts to other starchy grains such
as beans and rice (Cordenunsi et al. 2004; Bello-Péreza et al. 2006). The Amerindian
peoples of the Jê group Kaingang/Xokleng (also known as the Taquara/Itararé cultural
traditions, Noelli 2000) seem to have originated in southern Brazil between around 3950
and 2850 years BP in association with the increase in abundance of araucaria following
increased rainfall (Iriarte and Behling 2007). Araucaria has therefore become what has
been regarded as a cultural icon or culturally defined keystone species (Cristancho and
Vining 2004; Garibaldi and Turner 2004) for millennia (Wilson et al. 2019): it was used
as a ritual object by the indigenous Jê people (Fernandes and Piovezana 2015), and
remains as the most commonly used plant species among southern Brazil rural inhabi-
tants (Wilson et al. 2019). Araucaria seeds were a major element in the diet of the indi-
genous peoples living in and around the range of the araucaria trees, and the
ethnobotanical literature registers the transport, storage, and processing of the seeds
(Bitencourt and Krauspenhar 2006; Schmitz 2009; Reis et al. 2014). Transportation
and intentional or unintentional sowing of the seeds by the Amerindians are believed
to have led araucaria to regenerate following shifting agriculture patterns (Ntima
1968). These groups expanded from east to west and are regarded as responsible for
the establishment of vast araucaria managed forests (Noelli 2000; Bitencourt and Kraus-
penhar 2006; Stefenon et al. 2008), which can thus be conceived as cultural landscapes
(Reis et al. 2014).
10 A. F. SOUZA
A correlation between araucaria pollen percent and the number of records of Taquara/
Itararé human groups was found (Lauterjung et al. 2018). There is also a significant
overlay between the distribution of those human groups and their characteristic pit
houses and the increase of araucaria in the palynological record (Bitencourt and Kraus-
penhar 2006; Schmitz 2009; Lauterjung et al. 2018). The large crops and the fact that
the seeds can be stored and consumed during months in the future made the araucaria
seeds a key resource for the indigenous peoples (Schmitz 2009). The coincidence of the
increase in the abundance of araucaria in the late Holocene and the expansion of the
Taquara/Itararé culture was so intense that the AMF could have replaced the grassland
vegetation in mere 100 years (Iriarte and Behling 2007). The very low genetic differen-
tiation and lack of geographical structure of araucaria populations in the southern high-
lands of Paraná, Santa Catarina, and Rio Grande do Sul is also compatible with the
transportation of great quantities of seeds over long distances by the Amerindians (Lau-
terjung et al. 2018), similarly to what has been found for the Andean Araucaria araucana
(Marchelli et al. 2010) and the Amazonian Bertholettia excelsa (Shepard and Ramirez
2011), both species also cultivated and dispersed by Amerindians (Reis et al. 2014;
Thomas et al. 2015).
It is possible that the ability of araucaria to colonise the grasslands and form scattered
populations of isolated trees, which function as nucleation centres (Silva and Anand 2011;
Korndörfer et al. 2014), have fostered the AMF expansion over the grasslands during post-
glacial times (Barros et al. 2015). However, due to disturbances like fire, the edges of AMF
patches have been estimated to have expanded over grasslands by < 100 m in the last 4000
years (Silva and Anand 2011), and that araucaria would take ca. 14000 years to attain its
current geographical distribution (Lauterjung et al. 2018), a much longer period than the
4000–1500 years indicated by palynology, highlighting the need of human promotion of
seed dispersal (Bitencourt and Krauspenhar 2006; Lauterjung et al. 2018). Having said
that, it is worth stressing that the extent that the Taquara/Itararé groups mediated the
expansion of the AMF is an important research issue that requires further investigation.
Community composition and spatial structure

The AMF is a two-component system in which araucaria comprises a large portion of the
forest basal area (Souza 2007; Jarenkow and Budke 2009) and constitutes a distinct and
higher canopy layer above the lower canopies of angiosperms and Podocarpus lambertii
(Souza 2007). A similar architecture has been found in New Zealand’s mixed conifer –
hardwood forests (Ogden and Stewart 1995) and other southern hemisphere regions
(review in Enright and Hill 1995), including the Araucaria araucana forests on the
Andean southern South America (Veblen et al. 1995). The floristic composition of the
angiosperm-dominated layer of the AMF has, however, considerable geographic structure
along climatic, topographic, and edaphic environmental gradients. The abundance of ara-
ucaria, which is an evergreen species, produces an apparent homogeneity to the AMF.
However, 583 tree species have been recorded in the AMF, 54%–58% of which occurring
in one or two localities only (Jarenkow and Budke 2009b) or occurring in less than 20% of
sampled sites (Rezende, de Oliveira-Filho, et al. 2015). The botanical families with most
species are Myrtaceae, Fabaceae, Lauraceae, Melastomataceae, and Solanaceae and the
most common species, apart from araucaria, are Prunus myrtifolia (Rosaceae), Matayba
elaeagnoiodes (Sapindaceae), Clethra scabra (Clethraceae) and the tree fern Dicksonia
sellowiana (Jarenkow and Budke 2009; Rezende, de Oliveira-Filho, et al. 2015; Sevegnani
et al. 2019). The angiosperm-dominated layer of the AMF is formed by a mosaic of
evergreen forests which are seasonal with respect to temperature and whose local
species composition is largely selected by the environmental filters of frost frequency,
temperature, soil attributes, and the vicinity of other forest types besides spatial structure
related to latitude, altitude, and stochastic dynamics like dispersal limitation (Klein 1960;
Jarenkow and Budke 2009; Higuchi et al. 2012; Souza, Cortez, et al. 2012; Gonçalves and
Souza 2014; Rezende, Eisenlohr, et al. 2015; Santos, Saraiva, et al. 2015; Sevegnani et al.
2016). Climatic and elevation variation have been shown to select different species pools
at regional scales during the regeneration process already (Vefago et al. 2019). At the
local scale, topographic variation produces floristic variation in the AMF regeneration
(Souza et al. 2017). Overall, β-diversity is increased by the isolation and environmental
variation produced by the mountainous environment but diminished by the filter pro-
duced by more intense cold in parts of the AMF (Santos, Saraiva, et al. 2015). The iso-
lated patches of the AMF in the mountaintops of southeastern Brazil are floristically
distinct from southern AMF (Jarenkow and Budke 2009; Cantidio and Souza 2019),
with Melastomataceae genera like Tibouchina and Miconia being particularly frequent
(Jarenkow and Budke 2009).
Using both presence–absence and abundance data, Cantidio and Souza (2019) pro-
posed a general bioregionalisation for the entire Atlantic Forest, which mapped two
main floristic groups in the southern Brazil AMF (their occurrence ecoregions 20
and 21/abundance ecoregions 13 and 14). These compositional groups are distributed
from north to south in the southern Brazilian highlands either along the highest and
cooler eastern edge of the Serra do Mar and Serra Geral highlands (occurrence ecore-
gion 20/abundance ecoregion 13) or along the broader and relatively warmer western
hinterland (occurrence ecoregion 21/abundance ecoregion 14). Neighboring floristic
groups occupy smaller and peripherical portions of the AMF as seasonally dry forests
and lowland rainforest penetrate the margins of the AMF to the west and to the east,
respectively (Cantidio and Souza 2019). This confirmed earlier evidence for a split
into Eastern (Cantidio and Souza’s occurrence ecoregion 20/abundance ecoregion 13)
and Western (Cantidio and Souza’s occurrence ecoregion 21/abundance ecoregion
14) Mixed Forests in both Santa Catarina (Rezende, Eisenlohr, et al. 2015; Sevegnani
et al. 2016), Paraná (Jarenkow and Budke 2009) and Rio Grande do Sul (Gonçalves
and Souza 2014). West-to-east decreasing minimum temperatures and increasing
frosts are likely to limit the penetration of ‘warm-seasonal’ species into the AMF and
determine the existence of internal AMF floristic gradients (Higuchi et al. 2012;
Rezende, Eisenlohr, et al. 2015; Sevegnani et al. 2016). Western Mixed Forests cover
most of the southern highlands and are floristically ecotonal between the Eastern
Mixed Forests and Seasonally Dry Forests, as claimed earlier (Rambo 1954; Klein
1960), being responsible for the association between longitude and floristic composition
in the region (Jarenkow and Budke 2009; Higuchi et al. 2012; Souza, Cortez, et al. 2012).
Cantidio and Souza (2019) found a greater floristic affinity between the semideciduous
forests that cover the hinterland along the Uruguai and Paraná Rivers and the ecore-
gions forming the AMF than with the coastal rainforest, reinforcing the ecotonal
nature of the Western Mixed forests.
12 A. F. SOUZA
The transitional Western Mixed Forests have a reduced number of indicator and exclu-
sive species and share > 70% species with either Eastern Mixed or Seasonally Dry Forests
(Gonçalves and Souza 2014). In its westernmost areas, the Western Mixed Forests are basi-
cally Semideciduous Forests with the presence of Araucaria angustifolia (Jarenkow and
Budke 2009). Eastern Mixed Forests, on their turn, receive high yearly rainfall (c.
2000mm) and suffer cooler winter temperatures with occasional frosts or snow (Jarenkow
and Budke 2009; Higuchi et al. 2012; Rezende, Eisenlohr, et al. 2015; Sevegnani et al. 2016).
Except for the colder sites, they present increased forest height and basal area and are more
productive systems than Western Mixed Forests (Rosenfield and Souza 2014; Sevegnani
et al. 2019). The altitude/cold environmental filter reaches its strongest effects above ca.
1200 m, where climate limits the occurrence of various species, leads to decreased diver-
sity, and the emergence of local endemism like Leandra reitzii, Euplassa nebularis, and
Tibouchina reitzii (Sevegnani et al. 2016). Biogeographically, Eastern Mixed Forests,
including the AMF on mountaintops in southeastern Brazil, are formed by the meeting
of species migrating from tropical rainforests in southeastern Brazil (Gonçalves and
Souza 2014; Oliveira-Filho et al. 2015) and Andean or Southern South America (Gon-
çalves and Souza 2014; Santiago et al. 2018). Although the species of Andean origin are
probably what remained from colder paleoclimates that prevailed in Southern South
America (Sanmartín and Ronquist 2004; Ortiz-Jaureguizar and Cladera 2006), most
Eastern Mixed Forests individuals and species present a wide geographical distribution,
conferring to these forest not a relictual origin (Rambo 1951; Prado 2000) but a mixed
one, as species of Austral-Antartic origin coexist with species originating from south-
eastern Brazil (Gonçalves and Souza 2014; Santiago et al. 2018).
There is evidence that the AMF expansion following increased wetness in the late Holo-
cene was marked by selective migration of lineages from tropical southeastern Brazil
glacial refugia into the subtropics mediated by climatic niche conservatism. The tropical
niche conservatism hypothesis states that the tropics have more species than temperate
regions due to larger area and time to speciation events, that because many lineages ori-
ginated in the tropics, they are adapted to tropical climates and have tropical niches, and
that due to niche conservatism, most tropical lineages and species have been unable to
colonise cooler regions in subtropical and temperate latitudes (Peterson et al. 1999;
Wiens and Donoghue 2004). Indeed, it has been found that several Atlantic Forest tree
lineages in southeastern Brazil were restricted to the tropics, and that the number of
lineages and species occurring in both subtropical and tropical climates was lower than
expected by chance (Giehl and Jarenkow 2012). Additionally, few AMF lineages are exclu-
sively subtropical and diversified in the subtropics, species richness is reduced in subtro-
pical sites and is correlated with increased phylogenetic clustering (Giehl and Jarenkow
2012). These results indicate the inability of most tropical Atlantic tree lineages to colonise
the AMF, probably due to climatic niche conservatism based on cold intolerance. The phy-
logenetic structure of the AMF is associated to functional structure. In a comparative
analysis between southern Brazilian forest types, the AMF was associated with a stress-tol-
erant strategy while frost-free lowland rainforests were associated with a competitive strat-
egy within the CSR scheme (Rosenfield et al. 2019) see also Orihuela et al. (2015). This was
attributed by the authors to lower minimum temperatures, increased temperature season-
ality, and extreme climatic events in the AMF, which are believed to have selected for
species pools adapted to cope with frost episodes and seasonal winter resource limitation.
At local scales, disturbance and environmental filters are posed by hydrological gradi-
ents. These are commonly associated with topographic variation and have been shown to
create pattern in the spatial distribution of species and ecological groups, probably through
the selection of species able to tolerate flooding in river floodplains, and through inter-
mediate disturbance levels at medium elevation sites subjected to episodic flooding
(Budke, Jarenkow, et al. 2010). At the landscape scale, mortality and basal area loss
rates did not respond do climate and soil variables, but were smaller in larger fragments,
which had a greater relative interior areas (Gross et al. 2018). These effects translated into
degraded overall forest structure and diversity in smaller AMF fragments relative to larger
areas (Souza, Cortez, et al. 2012).
Regeneration and dynamics

Life-history and demography
In tropical and subtropical regions the secondary forests that develop after large scale dis-
turbances like landslides, windstorms, tornados, earthquakes, and slash-and-burn agricul-
ture, are frequently dominated by long-lived pioneer tree species (Veblen and Ashton
1978; Finegan 1996; Lusk 1999; Collins and Burns 2001; Gutiérrez et al. 2008). Long-
lived pioneers constitute a second life-history axis that is orthogonal to the well-known
fast–slow life-history continuum. ‘Fast’ or pioneer species grow quickly and present elev-
ated fecundity but survive poorly and have short life cycles, dominating early successional
stages. ‘Slow’ species grow slowly but present higher survivorship rates and dominate later
successional stages (Wright et al. 2010; Reich 2014; Forgiarini et al. 2015). In addition,
forest tree communities are also organised along a second, stature–recruitment tradeoff
major tradeoff axis (Souza et al. 2014; Forgiarini et al. 2015; Rüger et al. 2018). The
stature–recruitment tradeoff distinguishes long-lived pioneers from short-lived reprodu-
cers. The traits underlying this tradeoff are such that in order to reach a tall stature,
species must be selected for high longevity and therefore high survival, at the price of
delayed reproduction and reduced recruitment rates (Rüger et al. 2018). The recognition
of this second life-history dimension is necessary to accurately predict forest dynamics
(Rüger et al. 2020). In the AMF, long-lived pioneers dominate successional processes
after short-lived pioneers like Mimosa scabrella, Lithraea brasiliensis and Vernonanthura
discolor (Ferreira et al. 2013) dominate the first 10–30 years of succession (Budowski 1965;
Hartshorn 1978; Swaine and Whitmore 1988; Guariguata and Ostertag 2001; Kennard
2002; Kubota et al. 2005).
Both fast-growing short-lived and long-lived pioneers colonise disturbed sites from the
beginning (Ogden et al. 1987; Pollmann and Veblen 2004; Santos et al. 2012), and both
tend to form dense and continuous woody canopies, but long-lived pioneers dominate
successional sites after the demise of short-lived pioneers (Finegan 1996; Katovai et al.
2016). Long-lived pioneers frequently germinate in the shade, where their larger seedlings
can survive for variable periods of time (Ogden et al. 1987; Lusk 1999; Santos et al. 2012).
Both groups of species form nearly monodominant and nearly even-aged canopies when
cohorts recruited after disturbance reach maturity (Finegan 1996; Santos et al. 2012;
Katovai et al. 2016). The ability of long-lived pioneers to withstand shade is insufficient
for them to reach maturity, and they rely upon large-scale disturbances for recruitment
14 A. F. SOUZA
into reproductive sizes (Veblen and Ashton 1978; Ogden et al. 1987; Collins and Burns
2001; Pollmann and Veblen 2004; Nagel et al. 2014), displaying size distributions domi-
nated by adults and with scarce seedling, sapling, and/or pole abundances (Veblen
1986; Collins and Burns 2001; Pollmann 2002; Gutiérrez et al. 2008). Establishment
and regeneration into adult sizes is thus intermittent, with pulses following multiple tree-
fall gaps (Veblen and Ashton 1978; Ogden et al. 1987; Collins and Burns 2001; Pollmann
2002; Pollmann and Veblen 2004; Gutiérrez et al. 2008; Nagel et al. 2014). Cohorts of
young long-lived pioneers may dominate degraded or chronically disturbed habitats
like forest edges (Santo-Silva et al. 2016; Souza 2017) and intensively logged areas
(Collins and Burns 2001; Katovai et al. 2016). They often reach large adult sizes, frequently
being emergents (Hartshorn 1978; Lusk 1999), with lifespans often extending for many
centuries (Ogden et al. 1987; Enright and Ogden 1995; Lusk 1999; Pollmann and
Veblen 2004; Nagel et al. 2014).
As happened with other species of the genus Araucaria (Veblen 1982), Araucaria
angustifolia has received the relict status (Rambo 1951) with chronically failed regener-
ation in face of competing angiosperms. On the contrary, others have viewed the species
as having pioneer behaviour (Klein 1960; Ntima 1968; Soares 1979), with regeneration
concentrated on disturbed sites like roadsides and forest edges. Current knowledge indi-
cates that araucaria is a long-lived pioneer that relies on large-scale disturbances for suc-
cessful regeneration and then survives for long periods of time in the canopy without
further recruitment opportunities until the return of a new large-scale disturbance
(Souza 2007; Souza et al. 2008; Souza and Longhi 2019). Araucaria is markedly
different from the angiosperm-dominated fraction of the AMF system in several func-
tional ways. The araucarias have reduced mortality rates, leaf sizes, Specific Leaf Area,
crown depth, and wood density, and increased seed size and maximum height relative
to the angiosperm-dominated community underneath (Souza et al. 2014). Araucaria
seedlings maximum survivorship, growth and biomass accumulation is achieved under
full sunlight (Inoue and Torres 1980; Duarte and Dillenburg 2000; Olguin et al. 2019),
but seedlings can withstand moderate shade (45% of full sunlight) for up to 5 months
without noticeable changes in biomass allocation or chlorophyll content (Duarte and
Dillenburg 2000). After 12 months, however, survivorship is reduced by 60% (Olguin
et al. 2019). Deep shade (10% of full sunlight or less) induce increased Specific Leaf
Area (Duarte and Dillenburg 2000) and greatly reduced biomass and height (Inoue
and Torres 1980; Duarte and Dillenburg 2000; Franco and Dillenburg 2007), although
survivorship may remain elevated for up to 9 months (Inoue and Torres 1980). Contrary
to short-lived pioneers, but in accordance with long-lived pioneers, seeds germinate
quickly and seedlings establish in the forest understory (Chami et al. 2011), where
growth is increased under more open canopy conditions (Duarte et al. 2002). Seedling
survivorship is supported by the long period of attachment and dependence on seed
remains, which may last for up to 4 months (Franco and Dillenburg 2007). Seedlings
are found more often under more open canopy microsites than would be expected by
chance (Ntima 1968; Souza et al. 2008 but see Duarte et al. 2002), and growth is
higher in the centre than in the periphery of large treefall gaps (Olguin et al. 2019).
Similarly to what has been shown to take place with the Andean Araucaria araucana
(Burns 1993), seedlings are easily killed by fire, but trees > 40 cm diameter have been
reported to be fire resistant (Ntima 1968).
Despite seedling establishment in the understory, available data suggest chronic regen-
eration failure into the pole and larger size classes under shaded conditions (Ntima 1968).
Demographic analyses of Araucaria angustifolia are still scant, but in closed canopy forests
the existing data suggest heavy mortality in plants < 0.5 m height (Paludo et al. 2011). Ara-
ucaria density is much reduced in the natural regeneration of the AMF relative to adult
density in the same site in old growth or late successional forests (Caldato et al. 1996;
Backes 2001; Narvaes et al. 2005; Cordeiro and Rodrigues 2007; Paludo et al. 2009;
Souza, Souza, et al. 2012; Higuchi et al. 2015; Santos et al. 2018; Mazon et al. 2019). Sap-
lings and poles are usually less abundant than seedlings (Higuchi et al. 2015), confirming
low recruitment into higher regenerant classes early in the regeneration process. A recruit-
ment bottleneck during the sapling and pole life stages is also confirmed by left-skewed
size distributions dominated by large individuals and with a long tail of relatively rare,
small individuals (Duarte et al. 2002; Souza 2007; Souza et al. 2008). In selectively
logged forests, the species size distributions were more symmetric or even right-skewed,
indicating both a reduction in the relative abundance of logged adult trees but also a
recruitment wave of small and medium-sized individuals in response to transitory
canopy opening increase that accompany logging activities (Souza et al. 2008). A
similar increase in the proportion of young trees has been found in open-canopy Pinus
plantation stands (Duarte et al. 2002; Carvalho et al. 2016; Malysz and Overbeck 2018)
and in AMF fragments subjected do multiple impacts like logging, fire, and thinning
(Schaaf et al. 2006; Valente et al. 2010). In contrast, population size distributions of late
successional shade-tolerant late successional AMF species like Ocotea odorifera (Laura-
ceae) are dominated by young individuals that form a seedling bank that withstand
shade for long periods (Bett and May 2017).
Although analyses of population structure and the relative proportions of juveniles and
adults are unable to predict population trends in time (Johnson et al. 1994; Condit et al.
1998; Souza 2007; Virillo et al. 2011; Bin et al. 2012), it has been shown that species life-
histories tend to produce repeated patterns of population size distribution in habitats with
different disturbance regimes and resource availability (Swaine et al. 1990; Poorter et al.
1996; Wright et al. 2003; Souza, Souza, et al. 2010). Figure 3 summarises the expected
population structures of the araucarias in different forest successional states based on
the life-history behaviour of long-lived pioneers and the above-cited work. Population
size distributions in undisturbed old-growth forests are expected to be dominated by
reproductive trees that established centuries earlier during large-scale disturbances,
while young plants are much less numerous due to chronic regeneration failure in the
shade cast by angiosperms. Recently logged plots are expected to show bimodal distri-
butions due to short-lived regeneration pulses triggered by transient canopy opening fol-
lowing logging operations. Subadult trees are expected to grow into the reproductive class
and young plants into pole size, but further growth becomes again hampered by canopy
closure a few years after logging cessation. Repeated logging deprives populations from
adult plants while allowing for continuous regeneration of young plants due to sustained
canopy opening. Treefall gaps stimulate episodic regeneration pulses of young plants but
sustained growth into larger size classes is suppressed due to limited extension of light
increases in time and space. Finally, at chronically disturbed areas like forest-grassland
borders and along roadsides, sustained light availability is expected to promote continuous
recruitment and population size distributions marked by an abundance of adults as well as
16 A. F. SOUZA
Figure 3. Expected size distributions in populations of Araucaria angustifolia in distinct forest succes-
sional phases and disturbance regimes.
younger individuals, provided adults are not removed (see the Forest-grassland dynamics,
fire and grazing section below). Fire, grazing, and seed collection by local peoples may alter
these patterns through elimination of young and intermediary-sized plants. An assessment
of the extent with which these expectations meet observed patterns is currently difficult
because most data on araucaria population structure has been collected in forest disturbed
by selective logging or some form of chronic disturbance in the last 100 years, and data
collected in either truly undisturbed stands or forest margins are lacking. One exception
is the work by Souza in the São Francisco de Paula National Forest in Rio Grande do Sul
(Souza et al. 2008), that compared araucaria population size distribution between undis-
turbed old-growth and stands that were logged at known time intervals. Additionally,
careful measurements of canopy gaps and araucaria gap-phase regeneration are also
lacking in the literature.
The non-equilibrium Lozenge model of forest dynamics

The dynamics of the AMF can be fit into an overall model of forest dynamics proposed by
New Zealand researchers, and it would be useful to summarise it here. Generalisations on
the recruitment requirements, population trajectories, and interactions of the southern
hemisphere conifers have been summarised in the Lozenge model, also known as the Tem-
poral Stand Replacement model (Ogden 1985; Ogden and Stewart 1995; Enright et al.
1999). Developed by New Zealand researchers, this model departs from the observation
that cohorts of large and light-demanding conifers like Agathis australis (D. Don)
Loudon establish after large-scale disturbances. Successive cohorts of smaller angiosperm
species establish along with the conifers. They dominate the understory and suppress
conifer recruitment, which becomes limited to multiple tree-fall gaps. In the conifer cohort
thinning continues as does increasing basal area and height. Due to their high longevity
and large size, established conifers dominate forest biomass and architecture for long
time intervals (often for centuries, perhaps 1000 years, Ogden 1985). They function,
however, as remnant populations that lack significant regeneration. As conifer stands
start to senesce, they create large gaps and a second and less synchronous wave of recruit-
ment, which can be prolonged, can take place (Ogden et al. 1987). In the absence of a
large-scale disturbances, ultimately the conifer canopy trees begin to die and the
biomass of the cohort declines. The Lozenge name comes from the shape of the plot of
conifer biomass along time.
The Lozenge model is embedded in the modern non-equilibrium model of the
dynamics of ecological communities that emerged in the 1980s (Veblen 1982; Sousa
1984). The paradigm shift from equilibrium to non-equilibrium views highlighted the
commonness of environmental change and questioned earlier views that expected the
dynamics of natural systems to converge toward stability or some kind of stasis (Veblen
et al. 2016). The model is also a combination of the long-lived pioneer life-history with
the large size and longevity of conifers. It depicts the vertical occupation of well-lit
areas of forested ecosystems regardless of the angiosperm secondary succession that devel-
ops in the understory, and may represent niche partitioning by vertical stratification and
differential longevity (Ogden 1985). This is achieved through elevated growth rates associ-
ated with high diameters and maximum height and extended longevity (Easdale et al.
2007). These traits are advantageous in habitats prone to intense and large-scale disturb-
ances like those on mountainous landscapes (Armesto et al. 1992; Easdale et al. 2007;
Nagel et al. 2014), where they promote establishment in well-lit patches that later translate
into massive even-aged presence in the upper canopy or as an emergent layer (Armesto
et al. 1992; Ogden and Stewart 1995). A common feature to the conifers which regener-
ation is described by the Lozenge model is the formation of small or open crowns,
which are capable of casting little shade on co-occurring forest canopies (Souza 2007;
Urrutia-Jalabert et al. 2015). They thus contribute large amounts of biomass but have
only moderate influence on light competition and little effects on forest dynamics, what
has been called the ‘additive basal area’ phenomenon (Enright and Ogden 1995). Since
a millennium or so should be needed to complete the regeneration-domination-demise
cycle in the Lozenge model, even in the case of lack of large-scale disturbance restarting
the process in a given locality, fire, windstorms, and tornadoes will have stochastically
destroyed large forest tracks in the landscape, giving opportunity for the conifers to
form a regeneration mosaic (Ogden 1985). In other words, the dynamics of mixed
forests like the AMF is highly non-equilibrium and disturbance-mediated, with conifers’
long residence time in the canopy ensuring that at least one large gap will be produced
within the dispersal distance to allow its recruitment before the complete Lozenge cycle
is finished (Loehle 2000; Gutiérrez et al. 2008). In more general terms, the Lozenge
model fits into the hysteresis model of community dynamics (Vellend 2016), in which dis-
turbances produce phase changes in community composition (in this case the massive
regeneration of araucaria) that takes long time intervals to return to the system’s original
state (in this cases an angiosperm-dominated canopy with scant senescent araucaria).
Long-term population dynamics models of araucaria in the AMF are still scant and do
not compare closed forests with open habitats like forest edges and grasslands. However,
18 A. F. SOUZA
research on the species’ population ecology has increased in recent years and it points to
some correspondence with the Lozenge model. Figure 4 summarises the dynamics of the
AMF in light of the Lozenge model. A population matrix model fit to araucaria demo-
graphic data collected in an old growth AMF found population stability (population
growth rate λ ∼ 1) and estimated that young juveniles take ca. 600 years to attain the
largest tree size in the sample (Paludo et al. 2016). The survivorship of reproductive
trees corresponded to ca. 97% of the contribution to λ (elasticitiy), highlighting small con-
tribution of the natural regeneration for the maintenance of established populations of the
species. A similar conclusion was reached for New Zealand emergent conifer Dacrydium
cupressinum, which is maintained in the canopy through high longevity and scant regen-
eration events (Lusk and Smith 1998). A distance-independent individual tree-growth
model calibrated to a 70-year-old successional AMF simulated forest dynamics for 50
years and found continuous basal area growth of the emergent araucaria and Ocotea
porosa, regardless of initial forest density (Orellana et al. 2016). This confirmed the
dual nature of the AMF, with the long-term growth of the emergent araucaria occurring
almost independently from the angiosperm layer dynamics.
An agent-based model developed to simulate the population dynamics of araucaria in a
spatially explicit context and that included simplified competition with angiosperm
understory predicted establishment and growth of araucaria populations for the first
340 years (Valeriano et al. 2014). This was followed by a gradual decline of the population
due to insufficient recruitment under a developed angiosperm understory, resulting in the
virtual extinction of the araucarias after 750 years. In New Zealand, it has been found that
although podocarp conifers can tolerate considerable shade, the faster response of angios-
perms to small treefall gaps enables them to out-compete conifers in old-growth stands
(Lusk et al. 2015). For araucarias in southern Brazil, population reduction was prevented
Figure 4. Schematic representation of the long-term dynamics of the Araucaria Mixed Forest in
southern and southeastern Brazil and northern Argentina. The time scale of the forest phases is
measured in centuries and the completion of the entire sequence might take > 1000 years.
with simulated large-scale fire disturbances occurring periodically (Valeriano et al. 2014).
Growth models designed to simulate logging and management strategies in the AMF pre-
dicted araucaria population growth when adopting harvest cycles from 5 to 25 years,
minimum diameter cutting limit of 40 cm, and harvest intensities of up to 50% of stand
basal area (Orellana et al. 2017; Orellana and Vanclay 2018a). These results agree with
models developed to account for the dynamics of other long-lived South American
species like Nothofagus spp. that are mediated by competition with shade tolerant
species (Pollmann and Veblen 2004). They confirm the affinity of araucaria for cyclical
disturbances, be it simulated as logging or fire regimes, and confirm the overall predictions
of the Lozenge model. As long-lived pioneers, araucarias establish in disturbed sites like
large forest gaps, forest edges in forest-grassland ecotones and roadsides alongside fast-
growing smaller pioneers but outlive these and reach biomass dominance due to large
sizes and extended longevity. As observed for Araucaria araucana in the Chilean Andes
(Veblen 1982), the regeneration of araucaria is not deterministic but a non-equilibrium
process in which site dominance may depend on the severity of the disturbance as well
on the abundance of the seed rain and the density of secondary shade-casting
angiosperms.
A distance-dependent competition model between araucaria, light-demanding, and
shade-tolerant tree functional groups confirmed the vertical scape strategy of araucaria,
which had the greatest diameter increment, with rates twice those of the shade-tolerant
species (Orellana and Vanclay 2018b). The authors also found severe intraspecific
growth competition between the araucarias, indicating the self-thinning of dense stands
that follows the establishment stage. Self-thinning can also produce irregular or multimo-
dal diameter distribution among Araucaria cohorts (Pelissari et al. 2018) and spatial repul-
son in the survivorship of trees, that is higher away from forest edges where tree density is
high (Souza 2017). The lack of meaningful competition between shade-tolerant angios-
perms and emergent araucaria (Orellana and Vanclay 2018b) reinforces the view that ara-
ucaria suffers competition from the angiosperms during the regeneration phase rather
than as established trees (Ogden 1985). Lightly disturbed forests subjected to angiosperm
selective logging showed increased growth of canopy and sub-canopy araucaria in the first
decade or so (Longhi et al. 2018). However, when araucarias are logged together with other
timber species and no repeated disturbance occurs stands enter long-term arrested succes-
sion as indicated by reduced abundances of functional groups like Pioneers and biomass-
accumulators like araucaria and Large Seeded Pioneers (Souza and Longhi 2019). This
result also confirms the Lozenge model in that young araucaria suffered competition
from angiosperms and was unable to regenerate in the shaded understory. The authors
suggested management actions were needed to provide regeneration opportunities for ara-
ucaria and functional groups like pioneers and other long-lived pioneers. It should be kept
in mind, however, that long-lived pioneer recruitment failure and therefore management
initiatives aimed at improving their recruitment and biomass relate to timescales relevant
to humans in short-term dynamics only. In the long term, long-lived pioneers dominate
most successional stages and respond for most of the standing biomass than any other
functional group, except in very young forests (< 40 years) or forest patches that
remain undisturbed for a long time (perhaps > 400 years) (Rüger et al. 2020).
The application of the Lozenge model to araucaria in southern Brazil matches the idea
that some conifer clades have evolved pioneer strategies that enable them to coexist with
20 A. F. SOUZA
competitive angiosperms through the occupation of open disturbed habitats (Brodribb

et al. 2012). It is also possible that this coexistence is facilitated in elevated regions
subject to seasonal freezing, where competition with angiosperms is alleviated due to limit-
ations to angiosperm growth under low temperatures and to gymnosperm niche special-
isation for low temperatures (Brodribb et al. 2012; Fragnière et al. 2015). The adaptations
to cold presented by araucaria restrict its occurrence to regions where cold is stressful
enough for most tropical clades that form the angiosperm layer of the AMF (Giehl and
Jarenkow 2012). Araucaria cambial activity and radial growth are proportional to temp-
erature and rainfall in the beginning of the growth season (Martinkoski et al. 2015) but
are diminished in hotter summers, possibly due to high sensitivity to water stress (Silva
et al. 2009; Oliveira et al. 2010; Cattaneo et al. 2013; Martinkoski et al. 2017; Oliveira
et al. 2017). The climatic affinity of the species with reduced temperatures and elevated
rainfall has resulted in the estimation of its potential distribution through niche models
in cooler and mountainous regions of southern and southeastern Brazil (Bergamin et al.
2019; Wilson et al. 2019; Sousa et al. 2020).
The AMF disturbance regimes and succession

Even though ecological disturbances produced by volcanic eruptions and earthquakes are
key to the dynamics of temperate Andean forests in South America (Veblen et al. 2016),
they are absent from tectonically stable regions covered by the AMF. Disturbance is,
however, central in shaping the AMF structure and dynamics. It is produced by
different natural and man-made phenomena that periodically destroy parts of the AMF
system and allow the restarting of succession following the Lozenge model. They are
primarily windstorms and tornadoes by one side, and fire and livestock grazing by the
other side.
Climatic disturbances
Southern Brazil suffers strong ENSO effects, with excess rain (that includes extreme
floods) and temperatures in El Niño years, alternated with droughts and cooler tempera-
tures along La Niña years (Grimm et al. 1998; Garreaud et al. 2009). Besides this pattern,
climate change has increased the climatic intensity of the region. Analysis of climate data
found that minimum temperatures and rainfall have increased in the last few decades of
the twentieth century in northeastern Rio Grande do Sul (Souza and Longhi 2019).
Additionally, rainfall and the occurrence of tornadoes and storms have increased in
southern South America along the second half of the twentieth century resulting from
increased El Niño frequency (Haylock et al. 2006; Gobato et al. 2016), and the greatest
increases took place in Southern Brazil (Penalba and Robledo 2010). Rainfall in the
AMF is regionally accompanied by frequent tornados and windstorms, which peak in sub-
tropical South America due to a particularly rainy climate and where cold and hot air
masses collide on a regular basis (Dias 2011; Candido 2012; Gobato et al. 2016). The
spatial distribution of atmospheric conditions leading to severe thunderstorm and
tornado environments show a concentration of these phenomena in subtropical South
America (Brooks et al. 2003). Warm and moist air masses coming from tropical latitudes
including the Amazon basin collide with cold fronts coming from higher latitudes, gener-
ating storm supercells that produce windstorms, thunderstorms, waterspouts, and
tornadoes (Gobato et al. 2016; Oliveira et al. 2016; Gobato et al. 2019). Tornadoes occur in
southern Brazil almost every year, mostly during spring and summer (i.e. between Septem-
ber and March) (Marcelino et al. 2005). Accordingly, maximum lightning activity in South
America occurs in the subtropical region, between Paraguay, northern Argentina, and the
south of Brazil, with clustered occurrences between spring and summer (Beneti et al.
2014).
This pattern of intense climatic events implies highly frequent small scale secondary
succession due to treefall gap dynamics (Ge et al. 2013). In the AMF wind-thrown mor-
tality is concentrated among the tallest trees (Beckert et al. 2014; Ebling et al. 2014), cor-
responding to the Wind Dispersed Large Trees and araucaria functional groups (Souza
et al. 2014; Souza and Longhi 2019). Treefall gap formation in the AMF, however, has
been scantly studied. Côrte et al. (2007) found that gaps represented 19% of the canopy
of a one-ha study AMF, and varied in size from 5 to 121 m2, while Sanquetta et al.
(2011) found that gaps corresponded to 58.52% of the canopy of the stand they
studied. While the range of gap sizes reported by Cortes and colleagues corresponds to
small gaps, the overall proportion of the canopy covered by gaps is much higher than
the values reported for tropical and temperate forests (Muscolo et al. 2014). This is
because neither sources (Côrte et al. 2007; Sanquetta et al. 2011) discriminated between
canopy gaps and overall canopy openness, rendering estimates of gap openings probably
overstated.
Regardless of the actual distribution of canopy gaps in the AMF, the fall of tall trees
increases the recruitment of pioneer species due to increased regeneration opportunities
in treefall gaps (Chami et al. 2011). Whether succession will take the path including the
regeneration of araucaria stands or not seemingly depends on the scale of the tree fall
gaps created. Expressive araucaria regeneration is expected in large multiple tree fall
gaps, and has not been registered in single tree ones (Souza and Longhi 2019). Surviving
angiosperm trees in tornado-struck AMF forest stands present increased basal area than
trees in untouched stands, indicating competitive release after disturbance (Liebsch et al.
2019). The authors also detected increased intensity of demographic rates, with an increase
in mortality and recruitment. Undisturbed AMF annual mortality rates are generally under
3% (Salami et al. 2014; Gross et al. 2018; Liebsch et al. 2019), while annual mortality values
can reach 4.6%–10% in areas damaged by tornadoes (Liebsch et al. 2019). Angiosperm
basal area growth in undisturbed AMF is also greater in forest microsites with reduced
canopy cover, indicating competitive growth limitation (Salami et al. 2014).
As has been found for other vegetation types and at the global scale (Grime and Pierce
2012; Díaz et al. 2016), the regeneration dynamics in the AMF is mediated by the distri-
bution of species traits and functional groups that represent ecological strategies. A func-
tional analysis of the correlation structure of morphological traits of 66 tree species in
the AMF found that species were distributed along two main axes of ecological variation
(Forgiarini et al. 2015). A horizontal trade-off depicted the resource-acquisition versus
resource-conservation strategies with specific leaf area and trunk slenderness covarying
and representing acquisitive strategies. A vertical trade-off depicted the height-mortality
trade-offs, with taller species presenting reduced mortality and deeper crowns, while
growth was negatively related to wood density. A causal path model indicated that expected
trade-offs between survivorship and growth and between stem slenderness and wood
density were not detected. This was interpreted as resulting from the lack of extreme
22 A. F. SOUZA
pioneers and from the abundance of slow-growing hardwood species in the AMF (Forgiar-
ini et al. 2015). The fastest growing species in the AMF would not grow so fast as to com-
promise wood density and survivorship but grow fast enough as to attain increased size.
A classification analysis of the same trait data to which demographic rates were added, led
to the recognition of seven functional groups in the AMF: Araucaria, Palms, Pioneers, Large-
seeded Pioneers, Wind-dispersed Large trees, Large shade-tolerant trees, and Small Shade-
tolerant trees (Souza et al. 2014). These groups share the trait and vital rates multivariate
space and cluster the different ecological strategies depicted in the trait continua mentioned
above. The lack of extreme pioneers in this functional scheme derives from the fact that the
above-cited analyses did not include the woody shrubs and small trees that are found along
forest-grassland edges and in small forest patches developing withing grasslands, where the
fast pioneers of the AMF are found (see below). Along secondary succession, the AMF pre-
sents functional changes related to increases in resource conservation traits like wood density
and leaf dry matter content, while displaying increased overall trait diversity that accompa-
nies increased taxonomic diversity (Vicente-Silva et al. 2016).
Forest-grassland dynamics, fire, and grazing

While windstorms and tornadoes may produce multiple treefall gaps and reinitiate the
Lozenge model, most of the successional AMF stands are associated with forest-grassland
border and nucleation dynamics (Figure 5). As I stated in The ecological history of the Arau-
caria Mixed Forests section, the region currently occupied by the AMF has been a highly non-
equilibrium system formed by a large-scale forest-grassland mosaic since at least the Pleisto-
cene, with grasslands or forests prevailing in the landscape depending on climatic
Figure 5. Landscape view of the Araucaria mixed forest-grassland mosaic in São Francisco de Paula, Rio
Grande do Sul, Brazil.
fluctuations. During the last glacial period, palaeofires appeared in the environmental record
with very low but constant frequency, indicating that fires were rare events during that time
(Jeske-Pieruschka et al. 2013). Along the Holocene, fire events became very common in both
southern and southeastern Brazil highlands starting ca. 10,400 BP (Jeske-Pieruschka et al.
2013; Portes et al. 2018). These fires were likely caused by Amerindians and correspond to
the beginning of the Umbu and Humaitá cultures at the end of the Pleistocene (Behling
1997; Prous and Fogaça 1999; Behling et al. 2004). Prehistoric fires continued to be frequent
during the late Holocene even under humid climates (Jeske-Pieruschka et al. 2013), as Amer-
indians maintained more open highland habitats probably through slash and burn agricul-
ture (Iriarte and Behling 2007; Gessert et al. 2011; Jeske-Pieruschka et al. 2013; Portes
et al. 2018). In the last two centuries, European colonisers were associated with increases
in the frequency of fire and with the abundance of pollen grains of grassland herbs, due to
the increase in the use of grasslands as fire-managed pasturelands (Bauermann and
Behling 2009; Jeske-Pieruschka et al. 2010; Portes et al. 2018). Fluctuations in the Amerindian
population, in the European pace of colonisation, and in rainfall led to rapid forest cover
expansion or retreat relative to grassland (Portes et al. 2018).
Current precipitation and temperature favour forest expansion and reduction of native
grassland vegetation (Oliveira and Pillar 2004; Pillar and Velez 2010; Blanco et al. 2014;
Overbeck et al. 2016). However, woody species establishment in grazed grasslands is con-
strained by continuous grazing and trampling by livestock, with the magnitude of the
effect depending on grazing pressure (Quadros and Pillar 2001; Santos et al. 2007;
Müller et al. 2012). Grazing has also been shown to be a pervasive disturbance factor in
the AMF (Vibrans et al. 2011), and a determinant factor during the forest regeneration
process, selecting for species assemblages dominated by fast-growing pioneer species
(Vefago et al. 2019). Fire, however, is believed to be the most important factor controlling
the dynamics of the forest-grassland mosaics and the creation of sharp borders between
these two vegetation types (Pillar and Quadros 1997; Jeske-Pieruschka et al. 2010;
Jeske-Pieruschka and Behling 2012). This is because the recurrent use of fire by Amerin-
dians and by European colonisers for slash-and-burn agriculture and to manage pastures,
respectively, is believed to have kept the AMF from expanding in a faster pace over the
grassland, as well as to have produced the forest-grassland mosaic found nowadays over
much of the highlands of southern Brazil (Jeske-Pieruschka et al. 2010; Jeske-Pieruschka
and Behling 2012; Müller et al. 2012; Jeske-Pieruschka et al. 2013).
Araucaria leaves are highly flammable (Soares 1979), but the cold and rainy climate
prevailing in the AMF geographical distribution makes forest fires very rare (Müller
et al. 2012). A spatially explicit vegetation dynamics model confirmed that the coexistence
of the AMF forest-grassland mosaic is a non-equilibrium system dependent on periodic
disturbances like fire acting through feedbacks between fire frequency and grass
biomass (Blanco et al. 2014). Blanco and colleagues also found that, in the absence of dis-
turbances like fire, forest succession leads to replacement of grasslands by the AMF in less
than 100 years. Analyses of aerial photographs and satellite images have shown marked
establishment of forest vegetation along the forest borders (Oliveira and Pillar 2004;
Matos et al. 2016). The study of soil organic matter carbon isotopes have also pointed
to gradual forest succession through both border expansion and patch formation in the
AMF-grassland mosaic (Silva and Anand 2011). Under current climate, the transition
from forest to grassland is unlikely to occur without deforestation by humans, and
24 A. F. SOUZA
therefore only forest-grassland coexistence or forest dominance are possible (Müller et al.
2012). A mathematical model indicated that strong human influence disrupts the stability
of the forest-grassland coexistence, implying that too frequent fires can lead to grassland
domination in the landscape (Innes et al. 2013). It is noteworthy that pyrophobic high-
biomass systems as moist forests like the AMF can be converted into pyrophytic low
biomass systems dominated by shrub species adapted to fire by too frequent fires or
high-severity fires (Kitzberger et al. 2016). Indeed, fire-sensitive forests in temperate
South America and New Zealand have been lost as human activities reach vegetation
types that were not exposed to frequent fires through their evolutionary histories (Kitzber-
ger et al. 2016). However, analyses of land use dynamics in the twentieth century found
strong reduction in the grassland area due to marked substitution by agriculture and
exotic tree plantation, as well as by AMF expansion as a network of small patches in
close contact with other land uses (Matos et al. 2016; Nodari 2016). The authors attributed
this forest expansion pattern to socioeconomic changes that have led to rural exodus by
small landowners and to land concentration.
The core of the non-equilibrium dynamics between forests and grasslands lie in the suc-
cessional belts that are formed along forest borders and around nucleation elements and
forest patches embedded in the grasslands. In the grasslands, the seed rain produced by
birds is concentrated around nucleation elements like isolated trees and tall shrubs, con-
tributing to the establishment of forest-dwelling angiosperm shrubs and trees along forest-
grassland edges, and to forest nucleation under nurse araucaria trees (Duarte, Santos, et al.
2006; Santos et al. 2011; Dallabrida et al. 2017). Young araucarias frequently form nearly
monospecific stands along the forest-grassland borders whenever fire and grazing press-
ures are reduced, potentially starting the Lozenge model successional path (Figure 6).
Figure 6. Forest-grassland border in a small rural property in Gramado, Rio Grande do Sul, Brazil. Note
the massive regeneration front dominated by monospecific stands of Araucaria angustifolia.
Most saplings of Dacrycarpus dacrydioides and Dacrydium cupressinum, two long-lived

conifers in New Zealand, were also found in clearing margins (Lusk and Laughlin
2017). For araucaria it is possible that greater abundance on forest edges may not be attrib-
uted to them out-competing angiosperm species in well-lit environment but is likely to
reflect incapacity to reach maturity in the shade coupled with high longevity superior
resistance to drought and/or frost (Lusk et al. 2015; Lusk and Laughlin 2017). Decreasing
or cessation of fire and livestock grazing frequently leads to shrub encroachment on the
grasslands, mainly by Asteraceae species like Baccharis spp. (Müller et al. 2012). These
shrub species have been shown to reduce overall grassland plant richness and forb
species in particular (Guido et al. 2017), but function poorly as nucleation centres
(Duarte et al. 2010). Abundance is a key factor for the facilitation of forest species estab-
lishment under nurse plants, and the most abundant shrub and tree species in the grass-
lands are also those that function more frequently as nurses, harboring the largest number
of forest seedlings underneath their canopies (Marcilio-Silva et al. 2015). Even though
regenerating seedlings of bird-dispersed forest species are found under several angiosperm
species that serve as nurse plants, seedling density and diversity are significantly higher
under the crowns of isolated araucaria (Duarte, Santos, et al. 2006). Nurse plants are
also important to forest succession starting close to forest edges, explaining most of the
variation in sapling abundance in grassland plots along the forest border (Carlucci, Teix-
eira, et al. 2011).
Araucaria establishment in the grasslands occur independently from the distribution of
inorganic soil nitrogen (Garbin et al. 2006), and despite mortality due to ant herbivory and
competition for light with established grasses (Zandavalli and Dillenburg 2015). The seed
rain in forest patches initiated by nucleation in the grassland is dominated by small seeded
species with long-distance dispersal abilities (Duarte et al. 2007). The initial establishment
of araucarias in the grasslands may be guided by existing perches, since the distribution of
isolated trees of the species in the grasslands is independent of forest patch distribution
(Souza 2017). The abundance of forest tree seedlings commonly found under the canopies
of isolated Araucaria in the grasslands is attributable not only to perching, but also to
nursing, due to the improvement of environmental conditions like summer temperatures
and vapour pressure deficits as well as increased concentrations of soil nutrients under
individuals (Korndörfer et al. 2014), likely due to the strong association of araucarias
with mycorrhiza (Zandavalli et al. 2004). Araucaria nurse effects are self-reinforcing,
since araucarias in the grassland show aggregated spatial distribution (Souza 2017).
This occurs despite increased mortality relative to trees in forest patches, possibly due
to lightning and windthrow, and negative effects of conspecific density on survivorship
(Souza 2017).
Although the margins of clearings have increased light availability than treefall gaps,
plants growing on margins are also exposed to a greater extension to abiotic extremes,
especially frost, higher summer temperatures, and larger vapour pressure deficits (Lusk
and Laughlin 2017). This implies specific selection pressures on species ecological strat-
egies and trait syndromes, and in New Zealand it leads to filtering of species with rela-
tively more stress-resistant traits those that regenerate in treefall gaps (Lusk and
Laughlin 2017). The spatial structure and inferred dispersal limitation seem to be
minor forces explaining the variation in sapling species composition beneath isolated
trees in the grasslands, indicating that niche factors linked to nursing (isolated trees
26 A. F. SOUZA
traits) and perching effects had a dominant role in community assembly (Duarte et al.
2010). The regeneration under nurse trees is dominated by pioneer and generalist
species, but as nucleation succession proceeds species richness increases as resident
forest species colonise the forest patches (Duarte, Machado, et al. 2006). The nurse
effect of isolated trees is also enhanced because nutrient uptake is higher under their
crowns, favouring further woody encroachment into the grassland (Silva and Anand
2011). A phylogenetic analysis revealed a high degree of phylogenetic habitat filtering
during the development of AMF patches in the grasslands, with large and shadier
patches being occupied by more basal clades with limited photosynthetic capacity
like magnoliids, while small patches were dominated by sun-loving asterids (Duarte
2011). In the grasslands, rock outcrops are also associated with forest patches
embedded in the grassland matrix (Carlucci, Duarte, et al. 2011; Dallabrida et al.
2017). This is due to their functioning as nucleation units because of their attractive-
ness as perches (Carlucci, Duarte, et al. 2011) as well as their function as safe sites for
tree and shrub regeneration due to protection from grazing and reduced grass biomass
accumulation and thus flammability (Müller et al. 2012). From the functional point of
view, resprouting ability characterised angiosperm shrub and trees species colonising
the grasslands, with shorter and multi-stemmed plants dominating the succession in
frequently burned areas and tall single-stemmed plants dominating areas not burned
for 3–4 years (Müller et al. 2007). In summary, small forest patches, isolated trees,
and rocky outcrops are the main sources of grassland landscape heterogeneity that
influence forest succession and vegetation-fire feedbacks and hence the dynamics of
the whole mosaic (Blanco et al. 2014).
Seed production and dispersal

Biotic interactions are key to all the stages involved in the regeneration process of the ara-
ucarias and the development of the AMF, but the details of such interactions are still
largely unknown. Araucaria seed crops are large with copious amounts of seedfall, and
quite variable in space (from 44.3 to 427 kg ha−1) (Solórzano-Filho 2001; Mantovani
et al. 2004; Silva and Reis 2009; Figueiredo Filho et al. 2011). The drivers of this variation
have been speculated to include variation in adult tree density, mean tree age, and forest
successional stage (Mantovani et al. 2004; Figueiredo Filho et al. 2011), but no attempt has
been made so far to evaluate it analytically. Developing seeds may suffer varying pre-dis-
persal predation rates (from 0.7% to 7.3%, Hertel 1976; Mantovani et al. 2004) by Laspeyr-
esia araucariae (Lepidoptera, Tortricidae). The dynamics of this interaction, however,
remains to be quantified and modelled. Seeds are large (average ca. 6 g) (Mantovani
et al. 2004; Figueiredo Filho et al. 2011; Zechini et al. 2012), starch- and protein-rich
(Rosado et al. 1994). Seedfall occurs between March and June in the autumn and early
winter (Mantovani et al. 2004), a period during which most of the angiosperm species
are not producing animal-dispersed fruits (Solórzano-Filho 2001; Paise and Vieira 2005;
Liebsch and Mikich 2009). Such asynchronous seed production is believed to favour the
occurrence of granivorous animals, enabling resource offer by the AMF during the
entire year (Paise and Vieira 2005). The AMF fruit production, including araucarias, is
aseasonal with fruits available all year round (Marques et al. 2004). Despite earlier
claims that araucaria presents a seed production cycle of 2–3 years (Mattos 1994), time-
series analyses of the species’ seed production in several distinct localities yielded random,
uncorrelated temporal fluctuations (Souza, Matos, et al. 2010). Populations show geo-
graphical synchrony in seed production up to ca. 100 Km and are influenced by rainfall
and insolation in the two previous years, but a strictly mast seeding phenological
pattern was also not identified (Souza, Matos, et al. 2010). The majority of angiosperm
species in the AMF is zoochorous, with strong mutualistic links with fruit-eating and
seed disperser animals (Duarte et al. 2009).
Araucariaceae was the first conifer family to develop large seed cones when they
evolved in the Jurassic, possibly selected by the seed predation by the then-diversifying
large sauropod dinosaurs that could also act as their seed dispersers (Leslie 2011). Cur-
rently araucaria seeds are heavily consumed by an array of at least 23 insect, bird, and
mammal species that include capuchin and howler monkeys, parrots, jays, and rodents,
as well moth and beetle larvae (review in Vieira and Iob 2009). All of these species are
seed predators, and 8 of which also disperse the seeds (Vieira and Iob 2009). The seed pro-
duction phenology of the araucarias, including production of large quantities of seeds
during a short period of time within years but at unpredictable amounts between years
(Souza, Matos, et al. 2010) is believed to have evolved as a seed predator satiation strategy,
whereby in more productive years predators are not able to consume the entire crops and a
fraction of the seeds escape predation and germinate (Vieira and Iob 2009; Vieira et al.
2011; Brocardo et al. 2018). Variable seed production may also represent a escape in
time and space due to the impossibility of predators to ‘predict’ either the next time in
which individuals will produce seeds (escape in time) or which tree will produce seed
again (escape in space) (Albrectsen 2000). The chances of araucaria seeds escaping preda-
tion and recruiting into seedlings are higher during the seed production period and in
forest patches with high density of adult trees, and decrease markedly in logged or
degraded forest fragments, what further suggests that massive seed production in dense
groves of the species satiate seed predators (Vieira et al. 2011; Brocardo et al. 2018).
The main primary seed predators and disperser seem to be parrots (Amazona pretrei,
A. vinacea, and Pionus maximiliani) and parakeet (Pyrrhura frontalis) which gather on
trees to feed directly on the pinecones. Amazona pretrei migrates from the Rio Grande
do Sul to Santa Catarina during winter to feed on AMF with extensive araucaria popu-
lations (Marini et al. 2010). Primary as well as long-distance dispersal also depends on
blue jay species (Corvidae, Cyanocorax caeruleus and C. chrysops), which consume but
also carry seeds for variable distances, potentially over 100 m and including open areas
(Solórzano-Filho 2001). Parrots drop seeds locally at an average distance of 22.5 m
while parakeets may do so up to 10 m (Solórzano-Filho 2001). Parrots have a wasteful
and mobile foraging behaviour, dropping many intact or only partially damaged seeds
under the canopies of reproductive trees or under distant perches. They also promote
long-distance dispersal of up to 500 m (mean = 247 m) during flight when they spit out
or accidentally drop the seeds (Tella et al. 2016). Araucaria’s seeds maintain elevated meta-
bolic rates and thus can germinate quickly after dispersal, reducing the chances of post-
dispersal predation and desiccation (Gasparin et al. 2019; Shibata et al. 2019). The arau-
caria’s seeds can withstand partial predation and thus benefit from attracting consumers
for dispersal (Mack 1998; Tella et al. 2016; Tella et al. 2019). The large araucaria seeds ger-
minate well when partially eaten by parrots, probably due to the partial removal of the seed
coat eliminating the main barrier to moisture, while favouring subsequent water intake
28 A. F. SOUZA
and seedling emergence (Tella et al. 2019). A recent research found that 67.2% of the ger-
minating seeds on the forest floor were damaged by bird or rodent predators (Tella et al.
2016). However, informal field observations suggest seedling uprooting by seed predators
in order to consume attached seed remains, and seedling desiccation are among the main
seedling mortality factors (Sanquetta et al. 2005; Cordeiro and Rodrigues 2007).
Secondary dispersal is achieved by the action of medium-sized rodents like the agouti
(Dasyprocta sp.) and by small rodents, all of which are scatterhoarders that hide individual
seeds in order to consume them later, but fail to recover the entirety of the stocked grains
(Bordignon and Monteiro-Filho 2000; Iob and Vieira 2008; Vieira et al. 2011; Ribeiro and
Vieira 2014). Small rodents remove nearly 100% of fallen araucaria seeds, < 5% of which
are not consumed and germinate (Iob and Vieira 2008; Vieira and Iob 2009). Squirrels
(Sciurus ingrami) hide intact seeds and may remove them later to locations > 25 from
the seed source (Bordignon and Monteiro-Filho 2000). Despite the accumulation of
data based on field observations of seed eaters and removal experiments, the complexity
of the predator and dispersal network of the araucarias has not yet been comprehensively
integrated or modelled.
Genetic analyses estimated that establishment distance from assigned mother-trees
varies considerably (from 0.35–291 m, average = 83 m), but 47% of the seeds were effec-
tively dispersed within a of 60 m radius around the mother tree (Bittencourt and
Sebbenn 2007). Similar analyses in successional forests found that a few seeds reached
long distances (318 m), and that 50% were dispersed up to 133 m (Sant’Anna et al.
2013). As mentioned earlier, araucaria seedlings occur under more open canopy con-
ditions than expected by chance (Souza et al. 2008), and any additional shading seems
to completely prevent seedling survival. Bamboos are known to reduce seedling growth
and survivorship of some species (Itô and Hino 2004), but their main effects may occur
through seed predator behavioural changes (Caccia et al. 2009). Araucaria seedlings
were not found under bamboos growing in the forest understory in Santa Catarina
(Santos, Ferreira, et al. 2015), but their survivorship was not affected by bamboo presence
in a field experiment (Sanquetta et al. 2005). Bamboos have been found to reduce angios-
perm tree density and basal area in the AMF, although they do not prevent increases in
these forest metrics during succession (Liebsch et al. 2019). Bamboo dieback, however, sig-
nificantly increases canopy openness and promotes increased angiosperm regeneration
diversity and composition (Budke, Alberti, et al. 2010; Santos, Ferreira, et al. 2015).
Further details of the interaction between bamboos and araucarias are needed for
the AMF.
The AMF in the context of southern mixed forests

From a global perspective, the AMF keeps correspondence with few other equivalent forest
types. These have been variously named as Mixed Conifer/Hardwood Temperate Forests
(Gilliam 2016), Warm Temperate Rain Forest (WTRF, Grubb et al. 2013), and Oceanic
Temperate Forests (McGlone et al. 2016; McGlone et al. 2017). The Oceanic Temperate
Forests is a biome that has been recently proposed for lowland New Zealand and other
areas (McGlone et al. 2016), consisting of a mixture of conifer and mostly microphyll
broadleaved angiosperms with frequently abundant lianas, commonly including tree
ferns and vascular epiphytes. Its climatic envelope includes wet climates with mild
warm summers and cold winters with occasional but not severe frost (McGlone et al. 2016;
McGlone et al. 2017). Its global distribution includes the AMF in mountainous southern
and southeastern Brazil as well as forests in southeastern Australia, and the southern Chile
Valdivian and North Patagonian forests (McGlone et al. 2017). Few forests resemble the
Oceanic Temperate Forests in having abundant tall conifer emergents over relatively
small-leaved evergreen angiosperms (McGlone et al. 2017), and the AMF configures
one such good match of climatic, physiognomic, and structural features. This match
extends to some degree to their floristic composition for, as like in New Zealand, the
AMF has a conifer dominant, a podocarp common, and Cunnoniaceae, Myrtaceae, Win-
teraceae trees and tree ferns are present (Rambo 1951; Klein 1984; Jarenkow and Budke
2009).
Historical processes have filtered the taxonomic and functional traits of the New
Zealand Oceanic Temperate Forests through the selective elimination of N-fixing and
ectomycorrhizal groups and the colonisation of the flora with species bearing typically tro-
pical traits such as trunk buttresses and cauliflory, which are not common in temperate
latitudes (McGlone et al. 2016; McGlone et al. 2017). Similarly, the AMF has been
enriched and colonised by lowland Atlantic tree species migrating southwards from tro-
pical latitudes (Rambo 1950; Giehl and Jarenkow 2012; Gonçalves and Souza 2014;
Rezende, Eisenlohr, et al. 2015). Southwards, temperate forests in Chile and Argentina
form a floristically different region with high phylogenetic lineage diversity that seem to
have evolved independently from the tropical domain over millions of years (Rezende
et al. 2017; Rezende et al. 2018). This results from the high family richness of the southern
South American flora, which is accompanied by low genera and species levels richness and
high frequency of monotypic genera and monogeneric families like Desfontainiaceae,
Gomortegaceae, Eucryphiaceae, and Aextoxicaceae (Rezende et al. 2017). From the func-
tional point of view an evaluation of trait convergence between South American and New
Zealand temperate forests found close convergence of trait values, mainly wood density
and the proportion of evergreen species, and partial convergence in trait combinations
(Lusk et al. 2016). The authors attributed the differences in trait combinations, particularly
the greater abundance of small trees with large soft leaves in New Zealand to differences in
the biogeographic histories of New Zealand and southern South America.
As described above for the AMF, the several southern hemisphere Oceanic Temperate
Forests form non-equilibrium systems as the regeneration of their dominant conifers
depends largely on catastrophic disturbances (Veblen et al. 2016). Examples are the
forests dominated by Fizroya cupressoides (Molina) I. M. Johnst. or Araucaria araucana
(Molina) K. Koch in Chile and Argentina, by Agathis australis or Libocedrus bidwillii
Hook. f. in New Zealand, and by Athrotaxis spp. in Tasmania. Fitzroya cuppressoides
may be one of the largest biomass accumulator trees in the world (Urrutia-Jalabert
et al. 2015), growing very slowly and being regarded as a long-lived pioneer that
depends on large-scale disturbances like landslides and large fires for regeneration
(Donoso et al. 1993; Silla et al. 2002; Urrutia-Jalabert et al. 2015). In the Andes landslides
are frequent and F. cuppressoides establish abundantly on them along with Podocarpus
nubigea Lindl., Drymis winteri J. R. Forst.&G. Forst, and other species (Donoso et al.
1993). A very long lifespan is key to the successful regeneration of F. cuppressoides in
the region, since landslide return times may exceed one thousand years, a period
during which adults survive as remnant populations and regeneration remains suspended
30 A. F. SOUZA
(Urrutia-Jalabert et al. 2015). Young plants concentrate under full sunlight while regener-
ation is suppressed under shading by angiosperms (Donoso et al. 1993; Parker and
Donoso 1993; Silla et al. 2002), but regeneration in disturbed sites is hampered grazing
(Donoso et al. 1993) and large fires occurring at several centuries intervals (Veblen and
Ashton 1982; Donoso et al. 1993; Lara et al. 1999). Large and rare fires, however, may
also act as regeneration windows for F. cuppressoides through the elimination of compet-
ing broadleaf vegetation (Silla et al. 2002).
Other nonequilibrium Oceanic Temperate Forest type is found in the Chilean and
Argentinian Andes between 37°27′ S and 40°03′ S where the endemic conifer Araucaria
araucana forms mixed stands with southern beeches, mainly Nothofagus pumilio
(Poepp. & Endl.) Krasser, Nothofagus antarctica (G.Forst.) Oerst., and Nothofagus
dombeyi (Mirb.) Oerst. (Navarro-Cerrillo et al. 2014). The natural regeneration of Arau-
caria araucana depends on large- and small-scale disturbances that produce both massive
tree falls and smaller tree fall gaps, such as wind throw, fire, earthquakes, landslides, single-
tree falls, stand-level diebacks, and human disturbances like logging (Veblen et al. 1992;
Burns 1993; Rebertus and Veblen 1993; Navarro-Cerrillo et al. 2014). Early successional
fast-growing broadleaved species of Nothofagus and Drymis occupy and dominate the
recently opened spaces before Araucaria araucana (Rebertus and Veblen 1993), which
colonises the relatively open understory formed by deciduous Nothofagus (Burns 1993;
Veblen et al. 1995; Drake et al. 2012) and gradually overtop them, probably after ca.
150 years (Burns 1993). After a stand-replacement level disturbance, the high seed
input from Araucaria araucana surviving trees promotes high density of seedlings and
saplings and give the species the ability of to recruit quickly (Navarro-Cerrillo et al.
2014). Heavy browsing and grazing by livestock, however, can seriously prevent natural
regeneration and successional processes of A. araucana forests (Veblen et al. 1992; Roig
et al. 2013).
Studies carried out in the rain forests of the Asia-Pacific region supported the Lozenge
model for several species of Araucaria and Agathis (see review in Enright et al. 1999). New
Zealand emergent conifers such as Dacrydium cupressinum and Dacrycarpus dacrydioides
coexist with smaller and faster-growing angiosperms that regenerate more readily in small
openings due to the conifers’ life spans of > 700 yr, whose low recruitment rates are
sufficient to maintain established abundance (Ogden and Stewart 1995; Lusk and Smith
1998; Lusk et al. 2015). In New Zealand, mixed conifer-hardwood forests occupy
lowland areas where Nothofagaceae forests are absent as well as montane and subalpine
areas (Wyse et al. 2018). In these mixed forests angiosperms and conifers occupy distinct
forest strata and their regeneration is dependent on disturbances that take place at
different spatial scales and frequencies (Ogden et al. 1987; Ogden and Stewart 1995).
New Zealand is highly geologically active, and large-scale disturbances, mainly earth-
quakes, vulcanism, lahars, landslides, and decaying tropical cyclones are key elements of
the dynamics of the country’s forests (Wyse et al. 2018). Myrtaceae or tree fern-dominated
communities develop first in recently disturbed sites with species composition varying
between ecosystem types and latitude (Bray et al. 1999). Landscape-level disturbances
initiate conifer successional stands along with the light-demanding angiosperms, with
Agathis australis as the dominant species, but other conifer species like Dacrydium cupres-
sinum may occur depending on latitude, altitude, and site conditions (Ecroyd 1982;
Steward and Beveridge 2010).
Libocedrus bidwillii is the conifer that dominates montane and sub-alpine forests,
where it forms even-aged stands after wind-throws or landslides (Stewart and Rose
1989; Ogden and Stewart 1995). A landscape-scale simulation of forest succession fol-
lowing the Topo eruption in New Zealand estimated that angiosperms took nearly
1000 years to replace the stands of the initial colonising conifer Libocedros bidwillii
(Thrippleton et al. 2014). Distance from disturbance in both time and space forms a
successional sequence with increasing angiosperm basal area replacing conifer basal
area (Smale et al. 2016). In the absence of disturbances, both Agathis australis and
Libocedrus bidwillii show chronic regeneration failure under established angiosperm
canopies (Ecroyd 1982; Smale et al. 2016; Wyse et al. 2018). In the long run,
Agathis australis eventually predominates on ridges and drought-prone or low-fertility
sites (Steward and Beveridge 2010). In New Zealand, some of the taller podocarp
species (e.g. Podocarpus totara G. Benn. ex D. Don) as well as Dacrycarpus dacrydioides
(A. Rich.) de Laub., and Dacrydium cupressinum Sol. ex G. Forst. also appear to follow
the pattern of regeneration dependent on disturbances like aggradation following
intense flooding or earthquakes on benign sites where there is vigorous competition
from faster-growing broadleaved species (Duncan 1993; Ebbett and Ogden 1998;
Wells et al. 2001). In colder or more infertile sites these species show the ability to
recruit in small gaps likely due to more open forest canopies where there is less com-
petition (Ebbett and Ogden 1998).
Several Tasmanian Oceanic Temperate Forests are dominated by the endemic and
emergent conifer Athrotaxis selaginoides D.Don, which forms mixed stands with a
variety of evergreen and deciduous Nothofagus gunnii (Hook. f.) Oerst. Broadleaf
species like Nothofagus cunninghamii (Hook.) Oerst, Eucryphia lucida (Labill.) Baill.,
and Atherosperma moschatum Labill. show continuous regeneration and self-replace-
ment, although A. moschatum may slowly replace N. cunninghamii and E. lucida due
to differences in shade tolerance (Read and Hill 1988). Athrotaxis selaginoides is shade
intolerant (Read 1985), presents limited seed dispersal, no vegetative reproduction,
slow growth, and irregular cone production (Gibson et al. 1995). In forests where it coex-
ists with evergreen broadleaf species, the regeneration of Athrotaxis selaginoides is sup-
pressed by shading and the species depends on catastrophic disturbances like heavy
snowfalls, landslides, and severe windstorms for successful regeneration (Cullen 1987;
Read and Hill 1988; Cullen 1991). The same occurs with the conifer Phyllocladus asple-
niifolius (Labill.) Hook. f., that as Athrotaxis selaginoides regenerates continuously in
open or multiple treefall gaps (Read and Hill 1988). To a lesser extent, Nothofagus cun-
ninghamii, Leptospermum scoparium, and Eucalyptus vernicosa also find regeneration
opportunities in landslides (Cullen 1991). However, in stands with an elevated pro-
portion of the deciduous Nothofagus gunnii, Athrotaxis selaginoides is able to regenerate
continuously (Cullen 1987). Conifer coexistence with angiosperms also seems to benefit
from poorer soils (less fertile and/or poorly drained), where the competitive advantage of
Nothofagus cunninghamii is absent (Read 1995). As happened to other southern conifers,
Athrotaxis species have been severely impacted by recent changes in disturbance regimes
induced by human activities. One example is Athrotaxis cupressoides D.Don, which is
paleoendemic to Tasmania and suffered widespread population reductions after the
introduction of fire management by Europeans (Holz et al. 2015). Similarly, Athrotaxis
selaginoides has been shown to have become restricted to topographic refugia after
32 A. F. SOUZA
the transition from mild Aboriginal to much more intense European fire regime (Holz
et al. 2020).
This brief description of the forest dynamics of southern mixed or Oceanic Temperate
Forests illustrates that disturbance has a central role in structuring these forests and main-
taining their biodiversity at landscape scales (Kitzberger et al. 2016; Veblen et al. 2016; Wyse
et al. 2018). As suggested earlier (Ogden 1985; Cullen 1987), the dependence on disturb-
ances for successful regeneration is by now evident to be a common feature of the long-
lived conifers of the mixed temperate forests in the southern hemisphere and a major deter-
minant of their successional pathways (Wyse et al. 2018). This confirms the validity of the
Lozenge model (Ogden 1985) as a useful framework for the broad class of systems encom-
passed by the Oceanic Temperate Forests. Population matrix models applied to contrasting
habitat types confirmed the Lozenge model for Araucaria laubenfelsii by finding population
growth in open shrub but population stability in forest environments (Rigg et al. 2010). The
model also received support from a landslide hazard index, which successfully explained the
landscape distribution of Agathis australis in New Zealand (Claessens et al. 2006). Others
used the model to explain mixed conifer-hardwood stand development in the Pacific north-
west of the USA (Ishii and Ford 2002) and in the AMF (Souza 2007; Souza et al. 2008). We
can conclude that in non-equilibrium disturbance-driven mixed conifer-hardwood forests
disturbance-dependent conifers in different continents are ecologically similar in many
ways irrespective of taxonomy, meaning that these conifers can be considered as a function-
ally coherent group (Aiba 2016).
Some results suggest that in the absence of periodic large scale disturbances, shade-
intolerant conifers are able to coexist with angiosperm trees in cold and nutrient-poor
environments, either as emergents above the closed canopy of angiosperm trees or as
codominants in the open canopy (Aiba et al. 2013). It has been suggested that in these
forests cool summers or year-round low temperature on (subtropical mountains limit
the growth of evergreen angiosperms, while mild winters in subtropical and tropical
mountains) could give the competitive advantage to conifers over deciduous broadleaf
trees, leading to conifer dominance in the temperate and subtropical mixed forests
(Regal 1977; Aiba 2016). Thus, the coexistence between conifers and angiosperms
without periodic large-scale disturbances might depend on environmental factors such
as climate and soils strongly limiting angiosperm growth, while competitive exclusion
by angiosperms could explain the absence of conifers in warm and nutrient-rich environ-
ments (Aiba et al. 2013; Aiba 2016). There are also some southern conifers that appear to
develop beneath the open canopies of Nothofagaceae forests. One example is the above-
cited Araucaria araucana that is able to overtop the relatively open understory formed
by early successional broadleaved species of Nothofagus and Drymis in Chile and Argen-
tina (Burns 1993). Other examples of conifers that regenerate under Nothofagaceae cano-
pies are Podocarpus laetus and the related Prumnopitys ferruginea in New Zealand (Wyse
et al. 2018), and Podocarpus nubigenus and Saxegothaea conspicua in Chile (Amigo et al.
2004). This might at least partly reflect the fact that Nothofagaceae canopies are less dense
than those of warm-temperate or subtropical rainforests.
Lastly, there is evidence that several gymnosperms growth more slowly than co-occur-
ring angiosperms even under limiting abiotic conditions (Piper et al. 2019), and it is poss-
ible that a more positive carbon balance in gymnosperms is driven by longer leaf lifespan
than in angiosperms (Piper et al. 2019). The model however does not apply to all conifer
species, with some of them being life-long shade tolerants that reach the canopy through
continuous survivorship and growth in the understory (Adie and Lawes 2009). Podocar-
paceae species in particular increase their growth rates after juvenile stages, and that this
happens due to their shade-tolerance, longer leaf lifespan, and efficient nutrient use (Bro-
dribb et al. 2012; Lusk et al. 2015; Piper et al. 2019). The coexistence of evergreen angios-
perm and gymnosperm tree species may then result from high growth rates of
angiosperms combined with gymnosperm high stress tolerance (Regal 1977; Steward
and Beveridge 2010; Lusk et al. 2015; Piper et al. 2019).
Conservation and management

The AMF has suffered from conservation issues that fall into the deforestation, habitat
degradation, climate change, and sustainable use categories. Since the beginning of the
European colonisation, the AMF suffered extensive deforestation, being reduced from
its original estimated ca. 25.4 million ha to ca 3.2 million ha (12.6% of the original area)(-
Ribeiro et al. 2009). The deforestation process increased after 1870 when, promoted by the
Brazilian government, German and Italian colonisers begun to occupy southern Brazilian
highlands and replace forests by agriculture and livestock raising, and proceeded in colo-
nisation waves towards the countryside along the twentieth century (Cabral and Cesco
2008; Nodari 2016). Araucaria was the main timber species used for both settlement devel-
opment and export (Cabral and Cesco 2008), but several angiosperm species were also
logged mainly between the 1940s through the 1960s (Nodari 2016), with species of
Cedrela, Ilex, Lauraceae genera like Ocotea and Nectandra, Myrtaceae like Eugenia and
Myrcia (Albano Backes personal communication) as well as Lithraea brasiliensis Marchand
(Anacardiaceae), and Acca sellowiana (O.Berg) Burret. (Myrtaceae) (Ansolin et al. 2017)
among the most exploited ones. Between 1915 and 1960, Brazil exported ca. 18.5 million
m3 of timber wood obtained almost entirely from the AMF (Danner et al. 2012). During
the 1970s, araucaria corresponded to 90% of about 1.0 million m3 of timber wood
exported by the country annually (Stefenon et al. 2003). Although Araucaria timber
logging peaked between the decades of 1950 and 1970 (Guerra et al. 2002), from ca.
1970 on a new deforestation impetus has occurred due to the exponential increase in
the export-oriented soy monoculture (Nodari 2016). This process, however, seems to
have suffered a spatial shift beginning in the 1980s when habitat destruction has shifted
from the AMF to natural grasslands and forest remnants started a modest expansion in
the highlands due to rural exodus of small farmers (Scariot et al. 2015; Matos et al.
2016). Timber extraction of Araucaria was forbidden by law in 2001 due to the threatened
status of the species (Silva and Reis 2009), although government environmental agencies
continue to issue logging authorisations through sustainable forest management plans
(Guerra et al. 2002).
Extensive logging of the AMF along the twentieth century exhausted most of southern
Brazil natural timber stocks, what stimulated the establishment of exotic tree plantations,
mainly Pinus and Eucalyptus species (Guerra et al. 2002; Matos et al. 2016). This was
despite the higher quality of the Araucaria wood compared with that of Pinus species,
whose boards were sold in 2002 at half the price practiced for Araucaria boards
(Guerra et al. 2002). The declining exports of AMF timber species also triggered federal
government and some private initiatives that established Araucaria plantations both
34 A. F. SOUZA
directly and through tax incentives (Brepohl 1980). From 1966 to 1979 ca. 48,703 ha were
planted with araucaria in the Paraná state alone, totalling ca. 90,000 ha in southern and
southeastern Brazil 1979 (Guerra et al. 2002). These programmes, however, were not
pursued further and the planted area declined sharply from the 1980s on due to legal
restrictions to the felling of araucaria and the introduction of more productive Pinus
and Eucalyptus plantations, being estimated in 11,179 ha in 2011 (Danner et al. 2012).
Araucaria plantations develop a biodiversity-rich understory and constitute habitats
and corridors for a considerable number of native plant and animal species (Fonseca
et al. 2009).
Most of the remaining forest has been disturbed by selective logging and other impacts
(Dean 1996; Nodari 2016) and is concentrated in the coastal mountain ranges or scattered
into small fragments (80% of the fragments are < 50 ha)(Ribeiro et al. 2009). Araucaria
Mixed Forest fragments frequently lack Araucaria or present it in reduced densities due
to logging, and show signs of secondary succession after disturbance like high tree
density, reduced basal area, and reduced or no advanced regeneration, besides an impo-
verished flora mainly composed by a nested subgroup of the species found in larger, pro-
tected areas, as well as pioneer species as indicator species (Souza, Cortez, et al. 2012). Due
to its dependency on increased light levels for successful regeneration, Araucaria fails to
repopulate most AMF fragments from which it has been logged (Souza and Longhi
2019). AMF fragments have been shown to suffer from pervasive degrading edge effects
like cattle grazing and trampling (Souza, Souza, et al. 2010), fire, hunting, firewood extrac-
tion, and seed harvesting (Guerra et al. 2002; Silveira et al. 2007). These have varying and
complex effects on fragment structure and composition (Ferreira et al. 2016) including
increasing mortality and basal area loss (Gross et al. 2018). AMF forest patches exposed
to stronger human occupation have fewer mammal species and a predominance of
medium-sized omnivores and insectivores species with reduced community complexity
and a paucity of carnivores (Felidae) (Bogoni et al. 2016). Reductions in the coverage of
the AMF are threatening the reproduction and survival of key seed dispersers like the
parrot Amazona pretrei (Marini et al. 2010). Models of the spatial distribution of the
AMF since the last glacial maximum indicated that these forests and many of its tree
species like Araucaria, Annona rugulosa (Schltdl.) H.Rainer, and Ocotea porosa (Nees &
Mart.) Barroso are likely to continue to experience range contraction under projected cli-
matic changes in the future (Wrege et al. 2016; Bergamin et al. 2019).
The seeds of Araucaria represent an important complimentary food and income source
for farming families in southern Brazil (Silva and Reis 2009; Zechini et al. 2012), although
low income people from a diverse array of occupations also resort use it as a seasonal
resource (Silveira et al. 2007). In rural properties seed collection usually takes place
along with the production of yerba mate (Ilex paraguariensis) under araucaria trees, creat-
ing domesticated landscapes with economic, cultural, and social relevance for thousands of
farmers (Reis et al. 2018). It has been asserted that isolated araucarias in the grassland have
higher seed production than araucarias in forest patches, but this information remains to
be tested (Guerra et al. 2002). In protected reserves, seeds are collected by family groups in
forest patches in rural properties as well as in sustainable use reserves, directly from the
ground or by climbing the trees and mechanically removing the pine cones (Santos
et al. 2002; Silveira et al. 2007; Silva and Reis 2009). Seed collectors identify four main var-
ieties of Araucaria angustifolia that are not scientifically established, and prefer the caiová
variety for its bigger size, firmer texture, sweeter taste, easy peeling, and longer storage
times (Adan et al. 2016).
The average daily weight of seeds and pine cones collected by each family group varies
from 12.3 Kg to 33.2 Kg and may reach up to 80 Kg (Silva and Reis 2009; Zechini et al.
2012). For example, only in 2014, more than 8,700 tons of araucaria seeds were collected
in Brazil, corresponding to a total product value of R$19,325,000.00 (at that time over US$
8.2 million) (Reis et al. 2018). The income obtained from seed collection, however, is
highly uncertainty because araucaria seed production presents is temporally random
with marked interannual variation (Souza, Matos, et al. 2010). Besides their use as a
direct food sources to seed collector families, the collected araucaria seeds are also pur-
chased by intermediaries and resold at local and regional retail chains (Santos et al.
2002). The decline in the timber logging of araucaria has stimulated the establishment
of araucaria plantations for seed harvesting (Danner et al. 2012). The seed collection of
araucaria is likely to impact negatively on seed eating animals as well as on the regener-
ation of araucaria itself, but despite these potential effects research on the subject is
scant and quantitative models and estimations are badly needed (Silveira et al. 2007;
Silva and Reis 2009). At the same time, the scientific management of araucaria seed collec-
tion may help improve rural income and promote social and economic sustainability
(Guerra et al. 2002). An assessment of the consequences of araucaria seed collection
carried out in São Paulo found that the number of seedlings and germinated seeds
increased in harvested areas due to suppression of understory vegetation and reduction
in the density of seedeater animals (Fichino et al. 2017). This study identified a critical
threshold of 85% of the total seed production above which the araucaria populations
would be jeopardised, but recognised that lower harvesting intensities may be needed to
keep ecosystem services like carbon storage and fauna maintenance.
The plantation efforts involving araucaria in Brazil have clear parallels with similar
initiatives involving Agathis australis in New Zealand. As happens with araucaria, the
heartwood of mature kauri is reputed to be of excellent quality (Steward et al. 2014),
what led to unsustainable massive timber logging in the nineteenth and twentieth centu-
ries and the consequent drastic reduction of natural stands (Steward and Beveridge 2010).
Agathis australis timber and gum were the main export items of New Zealand for many
years (Ecroyd 1982). Logging of the species in natural forests ceased in 1981 but today,
old-growth forests dominated by kauri covers less than 1% of their original distribution,
while second-growth and scrub with regenerating kauri populations correspond to ca. 6%
of former 1,000,000 ha (Steward and Beveridge 2010). Kauri plantations have been estab-
lished mainly in the Te Ika-a-Māui northern island but there is evidence that the species
productivity is not diminished in the southern island, which may correspond to its cli-
matic niche and have included its geographical range in glacial times (Steward et al.
2014). This pattern corresponds to what has been referred to as the ‘Pleistocene ratchet’
– when forest species lose range during part of a glacial cycle but fail to recover it entirely
in a subsequent part of the same cycle (McGlone et al. 2016).
Despite the history of exploitation of kauri and its economic potential, only recently
appropriate sylvicultural technical information as rotation length and return on invest-
ment have begun to be developed (Chikumbo and Steward 2007; Steward and Beveridge
2010; Steward et al. 2014). Basal area and height models have shown that kauri growth and
productivity in planted stands are higher than those estimated at the beginning of the
36 A. F. SOUZA
twentieth century (Steward et al. 2014). This is because slow root development aggravated
by root limitation in nurseries limit height growth and volume production in the first 15
years after planting (Steward 2011), but the species presents increased growth rates after
the establishment stage (Steward et al. 2014). Analyses of age effects on wood quality
suggest that commercial harvest or thinning could occur as early as 50 years (Steward
and McKinley 2005; Steward et al. 2014), a value comparable to the 45 years rotation
length applied to araucaria plantations in Brazil (Guerra et al. 2002). Both natural and
planted populations are affected by the kauri dieback disease Phytophthora taxon
Agathis (PTA) (Beever et al. 2007; Scott and Williams 2014), what poses a risk to both
the survival of the species and its potential productivity (Steward et al. 2014).
Open questions and future research

The preceding sections highlighted the considerable amount of research that has been
dedicated to the ecology of the AMF, mostly of it in the last two decades. While such accu-
mulated knowledge has allowed a deeper understanding of this system, many research
questions remain unanswered or even not posed at all. Ecology is a historical science
(Belovsky et al. 2004), and our understanding of the AMF has benefited from a productive
investigation effort aimed at uncovering its dynamics since the last glaciation (see the The
ecological history of the Araucaria Mixed Forest section). However, almost all palynological
sampling sites lied along the eastern borders of the Serra Geral and Serra do Mar high-
lands, which may have particular conditions and may have had differential historical tra-
jectories due to the proximity with the Atlantic Ocean and to the particularly high
elevations. The necessity of reconstructing the history of more interior areas of the
AMF becomes even greater in face of the somewhat discordant palaeoenvironmental scen-
arios yielded by palynological studies and niche models. Basically, niche models tended to
project that the AMF covered large land expanses in southern Brazil during the last glacial
while palynological reconstructions predict grassland-dominated landscapes. The
relationship between the distribution of the araucaria and Amerindian activities has
been clearly established but there are several aspects of this relationship that deserve
further investigations, especially what was the spatial extent of the current distribution
of the AMF that can be attributed to indigenous peoples relative to the natural expansion
of the forest in the last millennia.
Although the AMF is a biodiversity-rich forest region, by far most of the information
existing on its ecology, structure, and dynamics concern the autoecology or the population
ecology of Araucaria angustifolia. Although this emphasis is understandable given the
major role this species plays in the ecology of the AMF, the need to improve our efforts
to understand the roles played by other species in the dynamics of the AMF is also striking.
The knowledge about the variation in the composition of the AMF has been significantly
improved in the context of the broad floristic variation of the Atlantic Forest, but the func-
tional aspects of this variation are still poorly understood, mainly the extent to which the
floristic turnover is accompanied by turnover in functional traits and trait syndromes or
ecological strategies. The pioneer study of Rosenfield and colleagues (Rosenfield et al.
2019) has identified increased frequencies of stress-tolerant traits in the AMF relative to
lowland rainforest and semideciduous forests in Rio Grande do Sul, but the structure of
the variation in these strategies along the entire AMF is still speculative. Despite increasing
awareness that the araucarias are long-lived pioneers that need large-scale canopy open-
ings to complete their regeneration cycle and escape competition with angiosperms, the
disturbance regime of the AMF remains poorly studied. Even basic information like
gap size distribution and return times, and their influence on the growth and survivorship
of the araucaria and the angiosperms is almost absent from the literature. If there is a
growing number of papers on the structure and dynamics of the AMF, few of them
have used the framework of the Lozenge model explicitly. There is a paucity of studies
assessing the initial stages of the model, when regenerating fronts dominated by young
araucaria advance over the grasslands along forest borders, despite this situation been
increasingly common in the southern Brazilian highland landscapes that face increasing
rural exodus.
Although the importance of facilitation through nurse plants has been repeatedly
found in the research of the AMF expansion dynamics, it is still uncertain how nurse
species establish themselves in the grasslands in the first place. The most invoked mech-
anism is a regeneration boom following pastureland burning, but establishment mechan-
isms aside from herbaceous removal through fire have been hardly described. The
hypothesis advanced by Soares (Soares 1979) that fire would be a key driver of araucaria
dynamics and that the species would be fire-resistant as adult also awaits testing. Finally,
while much-needed simulation models of the AMF have been successfully advanced
regarding both landscape-scale forest-grassland dynamics and temporal dynamics of
forest stands, several realistic aspects of this system await implementation, such as the
simulation of araucaria dynamics as a separate component of the AMF, the spatially
explicit relationships between trees (which have not been properly studied in the first
place), and long-distance dispersal. Furthermore, it is likely that forests at different
points of the distinct floristic compartments of the AMF display different dynamics,
which would in itself form a valuable venue of investigation. In this regard, given the
ability of some southern hemisphere conifers to regenerate through the relatively open
canopies of deciduous Nothofagaceae species, it is possible that Araucaria angustifolia
regeneration might not depend on large-scale disturbances in its entire range. Instead,
it may present continuous regeneration in the western and drier sections of the AMF,
where deciduous and semideciduous species prevail, as well as in in steep and laterally
lit AMF stands.
Acknowledgements
I am thankful to Maíra G. Pivato and Vinícius C. Silva for their kind support in the preparation of
Figure 1.
Disclosure statement
No potential conflict of interest was reported by the author(s).
ORCID
Alexandre F. Souza http://orcid.org/0000-0001-7468-3631
38 A. F. SOUZA
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A Review of The Structure and Dynamics of Araucaria Mixed

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A Review of The Structure and Dynamics of Araucaria Mixed

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NEW ZEALAND JOURNAL OF BOTANY

2021, VOL. 59, NO. 1, 2–54

A review of the structure and dynamics of araucaria mixed

CONTACT Alexandre F. Souza alexsouza.cb.ufrn.br@gmail.com

existing knowledge about each of them is necessary. Although I included a comparison of

Abiotic conditions and neighboring vegetation types

A brief recent ecological history of the araucaria mixed forests

Araucaria biology and the AMF as cultivated systems

September to March (spring and summer), when it is positively inﬂuenced by temperature

Community composition and spatial structure

Regeneration and dynamics

The non-equilibrium Lozenge model of forest dynamics

competitive angiosperms through the occupation of open disturbed habitats (Brodribb

The AMF disturbance regimes and succession

Forest-grassland dynamics, ﬁre, and grazing

Most saplings of Dacrycarpus dacrydioides and Dacrydium cupressinum, two long-lived

Seed production and dispersal

The AMF in the context of southern mixed forests

Conservation and management

Open questions and future research

Higuchi P, Da Silva AC, Sousa-Ferreira T, Trierveiler De Souza S, Pereira-Gomes J, Montibeller Da

Korndörfer C, Dillenburg L, Duarte L. 2014. Assessing the potential of Araucaria angustifolia

Rambo B. 1950. A porta de Torres. An Botânicos do Herbário Barbosa Rodrigues. 2:9–20.

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