Pioneer species distribution in treefall gaps in Neotropical rain forest; a gap definition and its consequences

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Abstract
An attempt was made to evaluate the consequences of applying a strict definition (Brokaw 1982a) to the delimitation of forest gaps in the field. The northernmost Neotropical rain forest, at Los Tuxtlas, Mexico, was searched for young (1–2 years old) single-event gaps that would meet the criteria of the definition. In 60 ha of rain forest, only 12 such gaps containing pioneer species could be found. Thirty-three pioneer species (shrubs and trees) were used as indicator species for gap conditions. Gap size, measured as projected canopy opening (sensu Brokaw 1982a), underestimated from 44 to 515% the size of the area colonized by pioneer species. On average the size of the colonized area was 3.4 times larger than the size of the projected canopy opening. The majority of the pioneer species showed a relative preference for gap borders, an area generally not included in the projected canopy opening. Pioneer plant abundance and density, and species richness and density, did not differ significantly between gap centres and gap borders. Floristical variation was not related to gap size or location in the gap. These findings can be explained if gap environment (to which pioneer plants respond) is seen as the result of many interacting factors, of which size of the canopy opening is only one. It is concluded that the definition for delimiting gaps in the field as proposed by Brokaw (1982a) cannot be regarded a generally applicable definition, and that its value as a comparative standard is doubtful.