The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering

D.A. Coomes, R.B. Allen, W.A. Bentley, L.E. Burrows, C.D. Canham, L. Fagan, D.M. Forsyth, A. Gaxiola-Alcantar, R.L. Parfitt, W.A. Ruscoe, D.A. Wardle, D.J. Wilson, E.F. Wright

    Research output: Contribution to journalArticle

    133 Citations (Scopus)

    Abstract

    1 Angiosperm trees often dominate forests growing in resource-rich habitats, whereas conifers are generally restricted to less productive habitats. It has been suggested that conifers may be displaced by angiosperms except where competition is less intense, because conifer seedlings are inherently slow growing, and are outpaced by faster-growing angiosperm species. Here we investigate whether competition with ferns and deeply shading trees also contributes to a failure of conifers to regenerate in resource-rich habitats. 2 We examined how changes in soil nutrient availability and drainage affected vegetation along the retrogressive stages of a soil chronosequence in southern New Zealand. Vegetation composition shifted from angiosperm-tree dominance on 'recent' alluvial terraces (< 24 ky), via coniferous-tree dominance on older marine terraces (79-121 ky), to coniferous-shrub dominance on the oldest marine terraces (291 ky). Soil drainage deteriorated along the sequence, and N : P leaves and N : Psoil indicate increasing P-limitation. Conifer species appear to be adapted to persistence on infertile and poorly drained soils. 3 The floor of the relatively fertile alluvial forests was deeply shaded (∼1% light transmission) by dense groves of tree-ferns and ground-ferns, and by large-leaved subcanopy trees. Few seedlings of any type were found on the forest floor, even in tree-fall gaps, and establishment was restricted to rotting logs and tree-fern trunks. Angiosperms were particularly successful at colonizing these raised surfaces. 4 Less shade was cast by the conifer-dominated forests on infertile marine terraces (∼5% light transmission), which lacked tall ferns. There were many opportunities for conifer establishment, with high seedling densities recorded on the forest floor and on logs. By contrast, angiosperm seedlings were mainly restricted to logs. 5 Our results suggest that several mechanisms act in concert to reduce regeneration opportunities for conifers in productive habitats. In particular, we suggest that tall ferns and deep shade are responsible for a restriction of regeneration opportunities in relatively productive forests in New Zealand, diminishing the opportunities for conifers to escape the competitive effects of fast-growing angiosperms. Thus 'crocodiles' may alter the outcome of the race between 'hares' and 'tortoises'.
    Original languageEnglish
    Pages (from-to)918-935
    Number of pages18
    JournalJournal of Ecology
    Volume93
    Issue number5
    DOIs
    Publication statusPublished - 2005

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    crocodiles
    tortoises
    hares
    fern
    ferns and fern allies
    angiosperm
    conifers
    coniferous tree
    Angiospermae
    persistence
    ecology
    terraces
    terrace
    tree ferns
    shade
    seedlings
    seedling
    habitats
    forest litter
    habitat

    Cite this

    Coomes, D. A., Allen, R. B., Bentley, W. A., Burrows, L. E., Canham, C. D., Fagan, L., ... Wright, E. F. (2005). The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering. Journal of Ecology, 93(5), 918-935. https://doi.org/10.1111/j.1365-2745.2005.01012.x
    Coomes, D.A. ; Allen, R.B. ; Bentley, W.A. ; Burrows, L.E. ; Canham, C.D. ; Fagan, L. ; Forsyth, D.M. ; Gaxiola-Alcantar, A. ; Parfitt, R.L. ; Ruscoe, W.A. ; Wardle, D.A. ; Wilson, D.J. ; Wright, E.F. / The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering. In: Journal of Ecology. 2005 ; Vol. 93, No. 5. pp. 918-935.
    @article{967231ea4bc74e649d5e6891b9f69d05,
    title = "The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering",
    abstract = "1 Angiosperm trees often dominate forests growing in resource-rich habitats, whereas conifers are generally restricted to less productive habitats. It has been suggested that conifers may be displaced by angiosperms except where competition is less intense, because conifer seedlings are inherently slow growing, and are outpaced by faster-growing angiosperm species. Here we investigate whether competition with ferns and deeply shading trees also contributes to a failure of conifers to regenerate in resource-rich habitats. 2 We examined how changes in soil nutrient availability and drainage affected vegetation along the retrogressive stages of a soil chronosequence in southern New Zealand. Vegetation composition shifted from angiosperm-tree dominance on 'recent' alluvial terraces (< 24 ky), via coniferous-tree dominance on older marine terraces (79-121 ky), to coniferous-shrub dominance on the oldest marine terraces (291 ky). Soil drainage deteriorated along the sequence, and N : P leaves and N : Psoil indicate increasing P-limitation. Conifer species appear to be adapted to persistence on infertile and poorly drained soils. 3 The floor of the relatively fertile alluvial forests was deeply shaded (∼1{\%} light transmission) by dense groves of tree-ferns and ground-ferns, and by large-leaved subcanopy trees. Few seedlings of any type were found on the forest floor, even in tree-fall gaps, and establishment was restricted to rotting logs and tree-fern trunks. Angiosperms were particularly successful at colonizing these raised surfaces. 4 Less shade was cast by the conifer-dominated forests on infertile marine terraces (∼5{\%} light transmission), which lacked tall ferns. There were many opportunities for conifer establishment, with high seedling densities recorded on the forest floor and on logs. By contrast, angiosperm seedlings were mainly restricted to logs. 5 Our results suggest that several mechanisms act in concert to reduce regeneration opportunities for conifers in productive habitats. In particular, we suggest that tall ferns and deep shade are responsible for a restriction of regeneration opportunities in relatively productive forests in New Zealand, diminishing the opportunities for conifers to escape the competitive effects of fast-growing angiosperms. Thus 'crocodiles' may alter the outcome of the race between 'hares' and 'tortoises'.",
    author = "D.A. Coomes and R.B. Allen and W.A. Bentley and L.E. Burrows and C.D. Canham and L. Fagan and D.M. Forsyth and A. Gaxiola-Alcantar and R.L. Parfitt and W.A. Ruscoe and D.A. Wardle and D.J. Wilson and E.F. Wright",
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    Coomes, DA, Allen, RB, Bentley, WA, Burrows, LE, Canham, CD, Fagan, L, Forsyth, DM, Gaxiola-Alcantar, A, Parfitt, RL, Ruscoe, WA, Wardle, DA, Wilson, DJ & Wright, EF 2005, 'The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering', Journal of Ecology, vol. 93, no. 5, pp. 918-935. https://doi.org/10.1111/j.1365-2745.2005.01012.x

    The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering. / Coomes, D.A.; Allen, R.B.; Bentley, W.A.; Burrows, L.E.; Canham, C.D.; Fagan, L.; Forsyth, D.M.; Gaxiola-Alcantar, A.; Parfitt, R.L.; Ruscoe, W.A.; Wardle, D.A.; Wilson, D.J.; Wright, E.F.

    In: Journal of Ecology, Vol. 93, No. 5, 2005, p. 918-935.

    Research output: Contribution to journalArticle

    TY - JOUR

    T1 - The hare, the tortoise and the crocodile: The ecology of angiosperm dominance, conifer persistence and fern filtering

    AU - Coomes, D.A.

    AU - Allen, R.B.

    AU - Bentley, W.A.

    AU - Burrows, L.E.

    AU - Canham, C.D.

    AU - Fagan, L.

    AU - Forsyth, D.M.

    AU - Gaxiola-Alcantar, A.

    AU - Parfitt, R.L.

    AU - Ruscoe, W.A.

    AU - Wardle, D.A.

    AU - Wilson, D.J.

    AU - Wright, E.F.

    N1 - cited By 111

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    N2 - 1 Angiosperm trees often dominate forests growing in resource-rich habitats, whereas conifers are generally restricted to less productive habitats. It has been suggested that conifers may be displaced by angiosperms except where competition is less intense, because conifer seedlings are inherently slow growing, and are outpaced by faster-growing angiosperm species. Here we investigate whether competition with ferns and deeply shading trees also contributes to a failure of conifers to regenerate in resource-rich habitats. 2 We examined how changes in soil nutrient availability and drainage affected vegetation along the retrogressive stages of a soil chronosequence in southern New Zealand. Vegetation composition shifted from angiosperm-tree dominance on 'recent' alluvial terraces (< 24 ky), via coniferous-tree dominance on older marine terraces (79-121 ky), to coniferous-shrub dominance on the oldest marine terraces (291 ky). Soil drainage deteriorated along the sequence, and N : P leaves and N : Psoil indicate increasing P-limitation. Conifer species appear to be adapted to persistence on infertile and poorly drained soils. 3 The floor of the relatively fertile alluvial forests was deeply shaded (∼1% light transmission) by dense groves of tree-ferns and ground-ferns, and by large-leaved subcanopy trees. Few seedlings of any type were found on the forest floor, even in tree-fall gaps, and establishment was restricted to rotting logs and tree-fern trunks. Angiosperms were particularly successful at colonizing these raised surfaces. 4 Less shade was cast by the conifer-dominated forests on infertile marine terraces (∼5% light transmission), which lacked tall ferns. There were many opportunities for conifer establishment, with high seedling densities recorded on the forest floor and on logs. By contrast, angiosperm seedlings were mainly restricted to logs. 5 Our results suggest that several mechanisms act in concert to reduce regeneration opportunities for conifers in productive habitats. In particular, we suggest that tall ferns and deep shade are responsible for a restriction of regeneration opportunities in relatively productive forests in New Zealand, diminishing the opportunities for conifers to escape the competitive effects of fast-growing angiosperms. Thus 'crocodiles' may alter the outcome of the race between 'hares' and 'tortoises'.

    AB - 1 Angiosperm trees often dominate forests growing in resource-rich habitats, whereas conifers are generally restricted to less productive habitats. It has been suggested that conifers may be displaced by angiosperms except where competition is less intense, because conifer seedlings are inherently slow growing, and are outpaced by faster-growing angiosperm species. Here we investigate whether competition with ferns and deeply shading trees also contributes to a failure of conifers to regenerate in resource-rich habitats. 2 We examined how changes in soil nutrient availability and drainage affected vegetation along the retrogressive stages of a soil chronosequence in southern New Zealand. Vegetation composition shifted from angiosperm-tree dominance on 'recent' alluvial terraces (< 24 ky), via coniferous-tree dominance on older marine terraces (79-121 ky), to coniferous-shrub dominance on the oldest marine terraces (291 ky). Soil drainage deteriorated along the sequence, and N : P leaves and N : Psoil indicate increasing P-limitation. Conifer species appear to be adapted to persistence on infertile and poorly drained soils. 3 The floor of the relatively fertile alluvial forests was deeply shaded (∼1% light transmission) by dense groves of tree-ferns and ground-ferns, and by large-leaved subcanopy trees. Few seedlings of any type were found on the forest floor, even in tree-fall gaps, and establishment was restricted to rotting logs and tree-fern trunks. Angiosperms were particularly successful at colonizing these raised surfaces. 4 Less shade was cast by the conifer-dominated forests on infertile marine terraces (∼5% light transmission), which lacked tall ferns. There were many opportunities for conifer establishment, with high seedling densities recorded on the forest floor and on logs. By contrast, angiosperm seedlings were mainly restricted to logs. 5 Our results suggest that several mechanisms act in concert to reduce regeneration opportunities for conifers in productive habitats. In particular, we suggest that tall ferns and deep shade are responsible for a restriction of regeneration opportunities in relatively productive forests in New Zealand, diminishing the opportunities for conifers to escape the competitive effects of fast-growing angiosperms. Thus 'crocodiles' may alter the outcome of the race between 'hares' and 'tortoises'.

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    DO - 10.1111/j.1365-2745.2005.01012.x

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    JO - Journal of Ecology

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