The richness and seasonal variation of ant‐plant interactions were compared in four habitats in México: lowland tropical dry forest (La Mancha, Veracruz), coastal sand dune matorral (San Benito, Yucatán), semiarid highland vegetation (Zapotitlán, Puebla), and lower montane humid forest (Xalapa, Veracruz). The effects of temperature and precipitation on the seasonal distribution of the number of ant‐plant interactions differed among habitats. The general linear models fitted to the ant‐plant interaction curves explained 78.8 percent of the variation for La Mancha, 80.1 percent for Zapotitlán, 18 percent for San Benito, and 29.5 percent for Xalapa. Even though rainfall is low in Zapotitlán and San Benito, minimum temperature was the most important factor accounting for the seasonal distribution and low number of interactions. At La Mancha, with milder minimum temperatures and higher water availability, temperature alone did not account for the seasonal distribution and number of interactions, whereas the effect of the precipitation × temperature interaction was highly significant. Xalapa exhibits the lowest temperatures and the highest precipitation, but the role of these factors was only marginal. We suggest that the vegetation at Xalapa, a mixture of tropical and temperate floristic elements, constrains ant‐plant interactions due to a limited presence of nectaries. Also, ants are less abundant in cool and relatively aseasonal habitats. The other habitats have tropical floristic elements that are abundant and frequently have nectar‐producing structures. We report considerable variation among habitats in the number and seasonal distribution of ant‐plant interactions, and suggest that it is due to the effect of variation in environmental parameters, the richness of plants with nectaries in the vegetation, and habitat heterogeneity.
Nectaries on fronds of Polypodium spp. have been studied previously only in cultivated specimens. We conducted field observations in middle-elevation forests in Mexico and found five ant species associated with nectaries of Polypodium plebeium and P. lepidotrichum. To investigate whether nectaries promote protection against herbivores, we performed ant-exclusion experiments with nectary-bearing ferns (P. plebeium) and other ferns without nectaries (Polypodium plesiosorum, P. furfuraceum, and Phlebodium pseudoaureum). When ants were excluded from the developing fronds of Polypodium plebeium, damage from foliage-feeding sawfly and lepidopteran caterpillars was significantly greater than in control fronds. Ferns without nectaries did not show a difference in damage between ant-excluded and control fronds. Our results demonstrate that fern nectaries can support ant defense of the plant body as do the extrafloral nectaries of many angiosperms.
A population of the giant leather fern Acrostichum danaeifolium was observed during an 18-mo period at La Mancha (19°36′00″N, 96°22′40″W), Veracruz, Mexico. The study site was 230 m from a brackish-water lagoon, in the understorey of the mangroves, which are dominated by Avicennia germinans (Avicenniaceae). Acrostichum danaeifolium showed a clumped distribution pattern, and one third of the population became fertile during this time. Plants had a mean number of 9.4 ± 0.45 sterile leaves, which developed continuously at a rate of 14.6 ± 0.44 leaves y −1 . The mean life span of sterile leaves was 7.7 mo. Although soil water was always available, phenological patterns of fertility, leaf production and leaf growth were strongly correlated with the seasonal climate. Individual leaf growth and leaf size increased significantly during the rainy season. Fertile leaves emerged exclusively from April to August and had a mean life span of 4.1 mo. Spore release was restricted to the wet season, when conditions for germination were favourable. Environmental triggers of fertility are discussed and phenological patterns are compared with ferns and angiosperms from other habitats.Resumen: Se observó una población de Acrostichum danaeifolium durante 18 meses en La Mancha (19°36′00″N, 96°22′40″W), Veracruz, México. El sitio del estudio distaba unos 230 m de una laguna de agua salobre, en el sotobosque de un manglar, que es dominado por Avicennia germinans (Avicenniaceae). Se observó una distribución agrupada de A. danaeifolium, y de la población aproximadamente un tercio de las plantas estuvieron en estado fértil. Las plantas tenían un número promedio de 9.4 ± 0.45 hojas estériles. Estas se desarrollaron continuamente durante todo el año con una formación promedia de unas 14.6 ± 0.44 hojas por año. La longevidad de hojas estériles fue de 7.7 meses. Aunque agua siempre estaba disponible en el suelo, los patrones fenológicos de fertilidad y la producción y el crecimiento de las hojas fueron fuertemente correlacionadas con el clima estacional. El crecimiento individual y el tamaño de las hojas aumentaron significativamente durante la estación lluviosa, y las hojas fértiles brotaron exclusivamente de abril a agosto y sobrevivieron 4.1 meses. Las esporas fueron liberadas únicamente durante la estación húmeda, cuando las condiciones para la germinación eran favorables. Se discuten factores medioambientales que pueden inducir la fertilidad y se comparan los patrones fenológicos con helechos y angiospermas de otros hábitats.
Based on known data sets and maximum entropy distribution data of fern and lycopod species registered in the Yucatán Peninsula, track and parsimony analyses were undertaken to evaluate the contribution of these groups to the establishment of biogeographical relationships of the peninsula with other areas. The resulting generalized tracks clearly agree with the geological origin of the peninsula and the previously recognized relationship with the Greater Antilles is not supported for ferns and lycopods. Instead, a Central American generalized track connects the Yucatán Peninsula with south‐eastern México and Central America. Floristically, the peninsula harbours 66 species of ferns and lycopods. Seven are registered for the first time in the Yucatán Peninsula and one is a new species for México. These species do not follow the latitudinal pattern expected if ecological factors, such as humidity and rainfall, were the most important in determining their distributions. Groups of areas recognized with parsimony analysis of endemicity could not be defined as provinces as a result of the lack of endemic species. Nevertheless, a regionalization scheme based on maximum entropy distribution data and supported by track analyses is proposed. Two separate districts are recognized within the Yucatán Peninsula: arid/dry Yucatán in the north and El Petén (humid) in the south. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98, 775–786.
The long evolutionary history of ferns may explain why some previous studies of fern nectaries have shown little or no benefit to ferns from nectary visitors, as any coevolved herbivores are those resistant to ant defence. The results suggest that ants protect Pl. crassinervata fronds against herbivory. The presence of nectaries, and the relationship with ants, may contribute to this fern's widespread occurrence and persistence in the face of disturbance, though many other factors also play a role. Ant defence may be more likely to benefit a widespread species of disturbed habitats that encounters a wide range of non-adapted herbivores.
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