by limited knowledge of the biology, propagation and management requirements of individual species. Consequently,
restoration initiatives rarely incorporate less well-known
species or those that are difficult to source and grow. We describe challenges associated with the restoration of threatened trees in the Araucaria Forest of southern Brazil, and
analyse the effectiveness of methods used to define target
species, identify seed sources and generate information on
the phenology of rare or threatened tree species. A review
of secondary data identified rare or threatened taxa as targets for seed collection. We then surveyed . km of trails
in forest remnants, identifying and mapping , seedproducing trees of species. Surveys confirmed the scarcity
of several tree species (including seven species with an abundance of ,. individuals per km), and nine species
showed no signs of fruiting during years of phenological
monitoring. These findings, together with limited knowledge and application of optimal seed collection methods,
are significant factors impeding the recovery of these species
within their natural habitat. Wider application of the results
of this case study could support restoration of the Araucaria
Forest with seedlings from a wider diversity of species.
Keywords Araucaria Forest, Brazil, conservation, phenology, propagation, seed collection, subtropical forest restoration, threatened species
Introduction
he restoration of damaged but previously complex ecosystems may be one of the greatest challenges facing
tropical forest conservation (Rojas-Briales & Weise, ).
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Methods
A list of target species for seed collection was identified, including all threatened or rare tree species. Threatened species were defined as those listed in the regional, national or
global Red Lists (SEMA, ; MMA, ; IUCN, ), including species categorized as Near Threatened. The threat
categories of the various Red Lists were standardized according to IUCN (). Rare species were defined as
those not included in official Red Lists but with few herbarium records (INCT, ) and a low frequency of occurrence, identified through surveys of the structure of the
Araucaria Forest (Durigan, ; Pasdiora, ; Barddal
et al., ; Geraldi et al., ; Seger et al., ;
Watzlawick et al., ; Kozera et al., ; Cordeiro &
Rodrigues, ; Cordeiro, ).
A list of potential forest remnants was developed based
on the occurrence of at least one target species. Data on
the conservation status of these remnants were compiled
based on our knowledge and on a technical report of the status of the Araucaria Forest (Castella & Britez, ).
The final selection was based on geographical variability
(including various floristic associations within the
Araucaria Forest, to maximize genetic variability for each
species), forest condition (fragments of medium to advanced successional stage, with a minimum size of
ha), and guarantee of protection (remnants located in protected areas or under other forms of conservation management, thus maximizing the likelihood of selected mother
trees surviving and being available for long-term monitoring and seed collection).
Each selected forest remnant was surveyed intensively by
walking along all existing trails to search for mother trees of
target species. Species were initially identified in the field
and later confirmed by comparison with herbarium specimens. The criteria for selection of mother trees were
good health (free of damage, disease and infestation by insects) and full reproductive maturity (tree size, presence of
old fruits, seeds or seedlings under the tree).
We attempted to map the highest number of mother
trees in the maximum possible number of remnants. To obtain seeds with broad genetic variation, they were collected,
when possible, from at least three populations of each species (Nogueira & Medeiros, ), and from at least trees
per population (Vencovsky, ). When this was not possible, surveys aimed to identify at least trees per species in
the whole study area. Closely related individuals of the same
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were found for of the target species (all with ,. individuals per km). Particularly rare species include A. excelsa,
O. sellowii, Scutia buxifolia, Bunchosia pallescens,
Maytenus aquifolia, Inga lentiscifolia and Quillaja brasiliensis, which were observed in ,% of the forest remnants,
with ,. individuals per km.
Preliminary phenological assessment Phenological monitoring revealed that some species exhibited fruiting or flowering seasons that differed from those reported in the
literature and herbarium data. The observed fruiting periods
for O. odorifera and O. porosa differed by a few months to
those reported, and larger variations between literature and
field observations were observed for Oreopanax fulvus,
O. sellowii, Cassia leptophylla, Casearia lasiophylla and
Lafoensia pacari (Supplementary Table S). For some species, including Tetrorchidium rubrivenium, Ocotea nutans
and Zanthoxylum kleinii, we recorded a complete absence
of reproductive structures throughout the phenological
monitoring period.
Results
Target species The preliminary list of threatened and rare
tree species included taxa (Supplementary Table S), of
which were threatened and rare. Of the threatened species, were listed only in the Paran state Red List (SEMA,
), one was listed only in the national Red List (MMA,
) and were categorized as globally threatened
(IUCN, ). Five species (A. angustifolia, Ocotea catharinensis, Ocotea odorifera, Ocotea porosa and Trithrinax brasiliensis) were included in all three Red Lists. Of the rare
species, analysis of secondary data revealed that some taxa
had , records in herbarium collections and were generally not recorded in phytosociological studies. These include
Colletia paradoxa, Maytenus boaria, Maytenus dasyclada,
Agonandra excelsa, Castella tweedii, Picrasma crenata,
Drimys angustifolia, Ouratea sellowii and Agarista pulchella.
Identification of seed sources in the field Twenty-six
Araucaria Forest remnants (Fig. ) in municipalities
were identified as priorities for seed collection. Ten
of these are within officially protected areas. A total of
. km of trails were surveyed (c. . km per remnant).
Mother trees of of the target species were mapped
(Supplementary Table S). No individuals of the other
target species were found in the remnants studied. In total
, individuals were marked as potential seed sources. The
aim of selecting a minimum of trees of each species in at
least three remnants was achieved only for A. angustifolia,
Cedrela fissilis, Ilex paraguariensis, O. porosa and
Podocarpus lambertii, all of which except P. lambertii were
observed in .% of the surveyed remnants, with an abundance of . individual per km. In contrast, , individuals
Discussion
Definition of target species and identification of seed
sources
The results indicate that the official list of threatened species
for Paran (SEMA, ) needs to be updated, as it does not
include several rare species that are threatened by habitat
loss and forest fragmentation. Possible candidates include
Butia eriospatha, Cedrela lilloi, Chionanthus filiformis,
Albizia edwallii, Albizia burkartiana, Agarista pulchella,
Acca sellowiana, Ouratea sellowii, Agonandra excelsa,
Castella tweedii, Picrasma crenata and Colletia paradoxa.
Several tree species were scarce in the study area
(Supplementary Table S), which is a common problem in
seed collection and restoration of rare and threatened species (Jalonen et al., ). Low levels of forest cover, edge effects, fragmentation, and the presence of invasive species are
all likely to drive changes in species composition, with rare
and threatened species particularly affected (Phartyal et al.,
). As demonstrated elsewhere in the Atlantic
Rainforest, the general absence of fauna from degraded
areas limits seed dispersal and pollination, further biasing
species composition in favour of naturally dominant species
(Silva & Tabarelli, ).
The results also demonstrate the difficulty of obtaining a
broad range of genotypes among seeds collected in the field.
Despite considerable effort, we achieved our aim of locating a
minimum of trees from at least three populations for only
five of the species studied. Way () recommended sampling individuals in a population as a benchmark figure
to support the production of seedlings for restoration, but
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our results demonstrate the technical and logistical constraints of achieving this for extremely rare species. The number of target species of which no individuals were found in the
Araucaria Forest indicates the need for research on the distribution, ecology and conservation of these threatened species.
Ex situ conservation, through seed collection and storage,
is an important conservation strategy, but the existence of
species with recalcitrant seeds (e.g. P. lambertii and A. angustifolia; Royal Botanic Gardens Kew, ) demonstrates the
importance of ensuring the protection of mother trees in forest remnants. Given the difficulties in storing seeds of recalcitrant species in ex situ seed banks (Phartyal et al., ), the
trees mapped are natural seed banks for restoration. Using
and improving information on the distribution of mother
trees is esssential for future restoration projects in Paran.
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Acknowledgements
We thank Jeniffer Grabias, Marcelo Bosco Pinto, Natalia
Alejandra Bedrij, Tatiane Andressa Silva and Elivelton
Gurski for assistance during the study. We also thank the
Instituto Ambiental do Paran and Instituto Chico
Mendes de Conservao de Biodiversidade for their support, and the Fundao Grupo Boticrio de Proteo
Natureza and the Global Trees Campaign for financial
support.
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Biographical sketches
PA B L O HO F F M A N N is a conservationist working on the propagation of
rare and threatened species of the Araucaria Forest, and monitoring
their development. CH R I S T O P H E R BL U M S research interests include
increasing the knowledge and understanding of the autoecology and
conservation status of rare and threatened species in southern Brazil.
SA N T I A G O VE L A Z C O S research interests lie in plant ecology, with a
focus on the biogeography of northern Argentina and central and
southern Brazil. DA V I D GI L L supports the development of conservation projects for threatened tree species as part of the Global Trees
Campaign. MA R I L I A BO R G O is a conservationist who specializes in
quantifying biodiversity and developing strategies for payment for
environmental services.
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