Abstract
Abstract Tree seedling establishment outside forest boundaries is controlled by many interacting factors. Understanding the relative importance of different pressures is essential for improving techniques for forest restoration and for understanding the potential of forest boundaries to shift and adapt to changing climate conditions. We investigated constraints on the spread of a key southern hemisphere forest type into grasslands by undertaking a multifactorial experiment sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture. We manipulated factors to determine the relative effects of sheltered microsites, competition with pasture species, availability of ectomycorrhizal fungi, soil nutrients and rabbit herbivory, on seedling emergence and survival. Overall survival was low (11.7% after 6 months and 1.5% after 10 months), but there were strong treatment effects on both seedling establishment and survival. The availability of shelter and competition with pasture species had strongest effects, with the presence of pasture species initially aiding seedling emergence due to a sheltering effect, but negatively affecting survival at later stages. Most remaining seedlings at the conclusion of the experiment had formed ectomycorrhizae regardless of whether inoculum was supplied, and seedling survival and health was positively related to ectomycorrhizal colonization. Fertilizing had less of an effect than other factors, and results regarding rabbit herbivory were inconclusive. Synthesis and applications . This study provides new insights into how factors interact to limit tree seedling establishment in grasslands and forest expansion into neighbouring ecosystems. This improves understanding of the ability of an ectomycorrhizal keystone forest species to migrate and adapt to climate change. This study also assists restoration practitioners in selecting techniques that will enhance seedling establishment, and highlights that direct seeding approaches in open grasslands are unlikely to result in high rates of Nothofagus establishment.
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Claims 10
Claim · model-written summary
Nothofagus seedling survival was 11.7% at six months and 1.5% at ten months.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Outcome
- survivale1
- Value
- 11.7% after 6 months and 1.5% after 10 monthse1
- Setting
- historically deforested retired pasturee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“Overall survival was low (11.7% after 6 months and 1.5% after 10 months)”
Claim · model-written summary
Treatments strongly affected seedling establishment and survival.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Treatment
- treatmente1
- Outcome
- seedling establishment and survivale1
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“there were strong treatment effects on both seedling establishment and survival”
Claim · model-written summary
Shelter and pasture competition had the strongest effects on Nothofagus seedlings.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Interactor
- pasture speciese1weed
- Outcome
- strongest effectse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“The availability of shelter and competition with pasture species had strongest effects”
Claim · model-written summary
Pasture species initially aided seedling emergence through shelter.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Treatment
- presence of pasture speciese1
- Interactor
- pasture speciese1weed
- Outcome
- seedling emergencee1
- Direction
- aidinge1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“presence of pasture species initially aiding seedling emergence due to a sheltering effect”
Claim · model-written summary
Pasture species negatively affected later seedling survival.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Treatment
- presence of pasture speciese1
- Interactor
- pasture speciese1weed
- Outcome
- survivale1
- Direction
- negatively affectinge1
- Qualifier
- later stagese1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“presence of pasture species initially aiding seedling emergence due to a sheltering effect, but negatively affecting survival at later stages”
Claim · model-written summary
Most surviving seedlings formed ectomycorrhizae regardless of supplied inoculum.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Interactor
- ectomycorrhizaee1symbiont
- Outcome
- formed ectomycorrhizaee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“Most remaining seedlings at the conclusion of the experiment had formed ectomycorrhizae regardless of whether inoculum was supplied”
Claim · model-written summary
Ectomycorrhizal colonization was positively related to survival and health.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Interactor
- ectomycorrhizale1symbiont
- Outcome
- seedling survival and healthe1
- Direction
- positively relatede1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“seedling survival and health was positively related to ectomycorrhizal colonization”
Claim · model-written summary
Fertilizing had less effect than the other factors.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Treatment
- Fertilizinge1
- Outcome
- effecte1
- Direction
- lesse1
- Comparator
- other factorse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“Fertilizing had less of an effect than other factors”
Claim · model-written summary
Rabbit herbivory results were inconclusive.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Interactor
- rabbit
Oryctolagus cuniculuse1herbivore - Outcome
- rabbit herbivory were inconclusivee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

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Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“results regarding rabbit herbivory were inconclusive”
Claim · model-written summary
Direct seeding in open grasslands is unlikely to produce high Nothofagus establishment.
- Taxon
- Nothofagus cliffortioides
Nothofagus cliffortioidese0 - Outcome
- direct seeding approaches in open grasslands are unlikely to result in high rates of Nothofagus establishmente1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture”
e1“direct seeding approaches in open grasslands are unlikely to result in high rates of Nothofagus establishment”