Physiognomy, structure and floristic composition of one hectare of lowland tropical rain forest was studied in detail at Los Tuxtlas, Mexico. Physiognomically, the Los Tuxtlas forest should be classified as 'lowland tropical high evergreen rain forest'. The forest showed a closed canopy at 30-35 m. Of all woody, non-climbing species with aDBH >_ 1.0 cm 89.407o (94.5070 of all individuals) were evergreen, 25.4070 (59.5 070 of the individuals) had compound leaves, and over 80070 of species (and individuals) had leaves in the notophyll and mesophyll size classes. The forest structure was characterized by a low density (2976 individuals with a DBH >_ 1.0 cm, 346 individuals with a DBH >_ 10.0 cm, per ha, excluding vines) with an average basal area (38.1 m 2, DBH>_ 1.0 cm, 34.9 m 2, DBH>_ 10.0 cm, per ha, excluding vines). This was attributed to the relative maturity of the forest on the study plot. The study plot contained 234 species (11 208 individuals with a height >0.5 m), of which 55.1070 (34.8o70 of individuals)were trees, 9.4070 (6.8070)shrubs, 3.4070 (44.3070)palms, 20.1°/0 (5.207o) vines, 6.8070 (8.707o) herbs and 5.10/0 (0.307o) of unknown lifeform. Furthermore, 58 species of epiphytes and hemi-epiphytes were found. Diversity of trees, shrubs and palms with a DBH _> 1.0 cm was calculated as Shannon-Wiener index (4.65), Equitability index (0.65), and Simpson index (0.10). The dominance-diversity curve showed a lognormal form, characteristic for tropical rain forest. The community structure was characterized by a relative dominance of Astrocaryum mexicanum in the understorey, Pseudolmedia oxyphyllaria in the middle storeys, and Nectandra ambigens in the canopy. Species population structures of 31 species showed three characteristic patterns, differentiated by recruitment: continuously high, discontinuously high, and continuously low recruitment. Height/diameter and crown cover/diameter diagrams suggested a very gradual shift from height growth to crown growth during tree development. Forest turnover was calculated as 138 years. Compared to other tropical rain forests the Los Tuxtlas forest had 1. similar leaf physiognomical characteristics, 2. a lower diversity, 3. a lower density, 4. an average basal area, and 5. a slow canopy turnover.Nomenclature is given in Appendix I 56
Summary. Growth and morphology of seedlings of ten tropical rain forest species were studied at Los Tuxtlas, Mexico. Seedlings were grown in three environmental conditions: the shaded forest understorey (FU, receiving 0.%2.3% of the daily photosynthetic photon flux, PF, above the canopy), a small canopy gap of approx. 50 m 2 (SG, receiving 2.1-6.1% of daily PF), and a large canopy gap of approx. 500 m 2 (LG, receiving 38.6-53.4% of daily PF). The growth of all species was enhanced in gaps, and in LG the effect was stronger than in SG. Plants grown in LG had a sunplant morphology, with a high root-shoot ratio (R/S), a high specific leaf weight (SLW) and a low leaf area ratio (LAR). Plants grown in SG or FU showed a shade-plant morphology, with a low R/S, a low SLW and a high LAR. Growth responses varied from species unable to grow in the shade but with strong growth in the sun, to species with relatively high growth rates in both shade and sun conditions. Shade tolerant species were able to grow in the shade because of a relatively high unit leaf rate. The pioneer Cecropia had a high growth rate in LG because of a high LAR. Most species showed a complex growth response in which they resembled the shade intolerant extreme in some aspects of the response, and the shade tolerant extreme in other aspects.
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.
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