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A notable feature of the highly fragmented montane forests of central Veracruz, Mexico, is the interaction between oak trees (Quercus spp.) and phloem-feeding scale insects (Gamper and Koptur 2010), identified as Stigmacoccus garmilleri Foldi (Hemiptera: Margarodidae) (Foldi 1995; Hodgson et al. 2007). Immature S. garmilleri instars colonize trunks and branches by burrowing under tree bark. The scale insects insert their mouthparts, called stylets, into phloem cells and feed on phloem. The phloem of the host plant is rich in carbohydrates but low in compounds containing soluble nitrogen and amino acids, which are necessary to the insects for protein building (Gullan and Kosztarab 1997). Phloem feeding insects therefore ingest and excrete large quantities of carbohydrates in the process of acquiring sufficient amino acids (Wäckers 2000). This waste excretion, termed honeydew, forms droplets at the end of long anal tubes, or anal filaments (Figure 1). Honeydew-producing insects tend to eliminate copious honeydew, live in groups, and are typically sedentary or semi-sedentary (Williams and Williams 1980). For several months of the year S. garmilleri resides within the tree in the form of these feeding instars. Adult female and male insects develop and can be found mating on the surface of the tree (Figure 2). Details of the life cycle of S. garmilleri were described by Hodgson et al (2007).
Hair-like anal filaments from scale insect Stigmacoccus garmilleri and drops of honeydew secreted from their ends (Chiconquiaco, Veracruz, Mexico). Scale insects are capable of reaching high densities on oak tree trunks and branches. High quality figures are available online.
We found no significant difference in anal tube density among the four aspects (north, south, east, and west) in any habitat (p > 0.05 in all cases; Kruskal-Wallis test). The density of insects among the three trunk heights was not significantly different in edge habitat (χ2 = 0.70, p = 0.71) or in the pasture area (χ2 = 0.001, p = 0.99), although there was a significant difference in forest habitat (χ2 = 10.98, p
Means, standard deviations, minimum and maximum values for honeydew-drop volume, sugar concentrations, and anal-tube lengths for scale insects (Stigmacoccus garmilleri) on oaks in forest (n = 756) and pasture habitats (n = 762) of Chiconquiaco, Mexico. Independent-samples t-tests revealed that habitats differed significantly in anal-tube length; however habitats did not differ significantly in sugar concentrations or volume of honeydew.
Spearman's correlation coefficients were calculated for the predictive strength of the recorded variables (temperature, relative humidity, drop volume, sugar concentration, and anal-tube length). Data were combined from 20 forest trees and 20 trees located in pasture habitat in Chiconquiaco, Mexico. In all, 1518 insects producing honeydew were examined, and data on temperature and humidity were recorded when sugar concentration, drop volume, and tube length were measured.
Variables that might affect the distribution of scale insects within single trees include altitude, host species, exposure to sunlight, aspect (north, south, east, west), trunk diameter, and possibly region (Kelly 1990; Dungan and Kelly 2003). Crozier (1981) found higher densities of scale insects on northern aspects of Nothofagus solandri trunks and hypothesized that higher mean temperatures on this side contributed to the effect. In agreement with Kelly (1990), aspect was not correlated with anal-tube density in our study. The varying slopes and corresponding microclimate created on tree trunks, irrespective of aspect, may have confounded the findings in this study.
Tree size and scale insect densities were inversely related with small trees harboring the highest densities, supporting Wardhaugh (2006) who noted a decrease in scale insect density with increasing diameters at breast height or branch diameter. In contrast, Kelly (1990) found that intermediate-sized trees harbored significantly greater tube densities while the small trees and thin upper branches bore almost no scale insects. Even the thinnest branches of oak trees in Chiconquiaco were found to frequently harbor scale insects. Physiological differences between the insect species and/or tree species in the distinct geographic areas of New Zealand and Mexico may cause this difference.
Examination of the distribution of scale insects in this system indicates that mosaics of forest and pasture provide good habitat for S.garmilleri. The great abundance of honeydew on scattered pasture trees may be of particular importance in this region. Scattered pasture trees have been found to be keystone structures because of the disproportionally large ecosystem services they provide relative to the area they occupy, in addition to the maintenance of habitat and connectivity to other habitat types (Manning et al. 2009). Scattered trees are threatened in many places, making appropriate levels of tree regeneration and preclusion of premature mortality of mature trees essential for maintaining these trees within landscapes (Gibbons et al. 2008). Despite the benefits of habitat and food resources to other organisms, increases of scale insects on scattered pasture and forest edge trees may induce physiological stress, transform forest growth dynamics, and decrease reforestation potential (Gamper, in prep).
The spatial distribution of scale insect populations on trunks and branches of trees of increasing diameters at breast height may indicate a strong temporal component to the spatial dynamics of scale insects driven by changing host tree phenology. Future studies on phytophagous insects infesting large host trees should consider more explicitly changes in population dynamics both spatially and temporally (Wardhaugh et al. 2006). In order to detect interactions between tree size and location (interior, edge, pasture), future experimental design should incorporate information on geographic location, with accompanying scale insect density and other descriptive attributes for a thorough, spatially-explicit statistical analysis. 041b061a72