Revising the convergence hypothesis of avian use of honeydew: evidence from Dominican subtropical dry forest. -Oikos 93: 250-259.A honeydew-producing scale insect (family Margarodidae, tribe Xylococcini, genus Stigmacoccus) was found associated with the tree Bursera simaruba in subtropical dry forests at elevations of 100 -400 m on the south side of the Sierra de Bahoruco, Pedernales Province, Dominican Republic. At two study sites, 91% of Bursera trees supported locally dense populations of Margarodidae. Fifteen species of birds were observed foraging on the honeydew, but most observations were of the winter resident Cape May warbler (Dendroica tigrina) and black-throated blue warbler (D. caerulescens), and the permanent resident bananaquit (Coereba fla6eola) and blackcrowned palm tanager (Phaenicophilus palmarum). The Cape May warbler actively defended the honeydew resource but frequency of use of honeydew was influenced by the close presence of flowering agave and scale insect density. Data suggest that honeydew may be a critical component of the diet of this species especially during the late winter dry-season. Hymenopteran insects also were observed feeding on honeydew, but rates of consumption did not approach that of avian species. The occurrence of this phenomenon in Dominican dry forest is discussed in light of the convergence hypothesis of bird use and defense of homopteran honeydew in which it is proposed that birds are able to maintain relationships with scale insects in moist, warm temperate forests because it is in these climates where ant abundance is low. We suggest that our observation of a well-developed bird -homopteran system in classic subtropical dry forest supports the proposed mechanism of reduced competition with ants allowing bird use of honeydew, but we suggest that a broader array of especially insular habitats which may be relatively depauperate in terms of ants also can be expected to support bird -homopteran systems.

Abstract:A honeydew-producing scale insect, Stigmacoccus garmilleri (Margarodidae), is associated with oak trees (Quercus spp.) in highland forests of Mexico. Although feeding by ants on scale-insect honeydew is more frequently documented in the literature, the honeydew produced by feeding instars of S. garmilleri is sufficient to provide nourishment for birds. This study elucidates bird use of honeydew in the tropical montane forests near Chiconquiaco, Veracruz, Mexico, and uncovers patterns in honeydew foraging. Over a 2-mo period, 40 trees harbouring scale insects, located in both forest and pasture areas, were intensely studied (160 h of bird-foraging observations along with quantitative measurements of honeydew production). Fifteen resident bird species and 18 migrant species were observed visiting observation trees. Approximately 72% of the resident bird species and 83% of the migrant bird species observed were recorded to forage on scale-insect honeydew. Audubon's warbler (Dendroica coronata auduboni) was the most active consumer and defender of the resource. Of 118 aggressive chases observed, only 9.65% occurred in forest observation trees, and 90.3% in pasture trees. Audubon's warbler demonstrated preferential defence and territorial patrolling of scale-insect honeydew in scattered pasture trees.

Stigmacoccus is an unusual scale insect genus from Central and South America that has been little studied. It belongs to the family Stigmacoccidae within the archaeococcoid group of genera which used to comprise the family Margarodidae (Morrison, 1927) but which are now considered to represent at least 9 families. The present paper describes or redescribes the adult females, adult males, cyst stages and crawlers of the three known species (S. asper, S. garmilleri, and S. paranaensis), plus the prepupa of S. garmilleri and S. paranaensis, and (briefly), the pupa of S. paranaensis. It is considered that the female has two cyst stage instars; the number in the male is uncertain. Adult female S. asper and S. paranaensis appear to have groups of loculate pores on the walls of the vagina. A lectotype for S. asper is designated. In addition, cyst stages of three further undescribed species are described (but not formally named) and illustrated. Some observations on the biology and life cycle are also included. The honeydew of Stigmacoccus species has been shown to be an important energy source for overwintering passerine birds which defend this resource. A summary of our present knowledge is presented, including how the honedew is eliminated (through a long anal tube) and details are given with regard to rates of honeydew flow, sugar concentration, cyst densities and annual timing of peak flows. The annual life cycle, as far as it is known, is discussed. It is concluded that this honeydew could be economically important as a source of sugar for honey production but this would need to be carefully managed to maintain an ecological balance.

Stigmacoccus garmilleri Foldi (Hemiptera: Margarodidae) is an ecologically important honeydew-producing scale insect associated with oak trees (Quercus spp.) in highland forests of Veracruz, Mexico. The honeydew exudates of S. garmilleri serve as a significant nutrient source to many species of birds, insects, and sooty molds. Oak trees found in the forest interior, forest edge, and those scattered in pasture areas support scale insect colonies, though the pattern of insect infestations on trees within these varying landscape types has not been elucidated. This study aims to describe the distribution of scale insect infestation and any distinctions in honeydew production based on tree location. Scale insect density, honeydew volume, and sugar concentration were surveyed throughout a continuous landscape that included both patches of forest and scattered pasture trees. In addition, the anal filament through which the honeydew drop is secreted was also measured and was experimentally removed to test and measure regrowth. Scale insect densities on tree trunks were greatest on pasture trees, while intermediate densities were found on trees at the forest edge, and low densities on interior forest trees, suggesting that trees in disturbed areas are more susceptible to scale insect infestation. Trees with small diameters at breast height had significantly higher insect densities than trees with medium to large diameters. Trunk aspect (North, South, East, and West) was not a significant determinant of scale insect density. In forested areas higher densities of scale insects were found at three meters height in comparison to lower heights. Sugar concentrations and drop volumes of honeydew in forest and pasture areas were not significantly different. However, scale-insect anal tubes/filaments were significantly longer in pasture than they were in forests. Sugar concentrations of honeydew appeared to be positively correlated with temperature and negatively correlated with relative humidity. Experiments indicated that anal filaments could grow approximately 4 mm every 24 hours, and average tube growth was significantly faster in pasture than in forest, suggesting that there may be a physiological effect on the insect due to landscape disturbance. The results obtained in this study describe the increases in scale insect infestation of trees with forest disturbance. The effect of these increased scale insect densities on the host tree physiology is still to be resolved.

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