Since the identification and imprisonment of “Typhoid Mary,” a woman who infected at least 47 people with typhoid in the early 1900s, epidemiologists have recognized that ‘superspreading’ hosts play a key role in disease epidemics. Such variability in transmission also exists among species within a community (amplification hosts) and among habitat patches across a landscape (disease ‘hotspots’), underscoring the need for an integrative framework for studying transmission heterogeneity. Here, we synthesize literature on human, plant, and animal diseases to evaluate the relative contributions of host, pathogen, and environmental factors in driving transmission heterogeneity across hosts and space. We show that host and spatial heterogeneity are closely linked and that quantitatively assessing the contribution of infectious individuals, species, or environmental patches to overall transmission can aid management strategies. We conclude by posing hypotheses regarding how pathogen natural history influences transmission heterogeneity and highlight emerging frontiers in the study of transmission heterogeneity.

Resumen.-Documentamos los patrones de disponibilidad de néctar y la abundancia de aves nectarívoras por cerca de tres años en nueve sitios de estudio a lo largo de un gradiente altitudinal de  m en la isla de Hawai para investigar la relación entre la variación en los recursos y la abundancia de aves. La densidad de flores (flores ha-) y el contenido energético del néctar de la planta monodominante llamada Metrosideros polymorpha fueron medidos a lo largo del gradiente. Cuatro especies nectarívoras fueron capturadas mensualmente con redes de niebla y censadas cada tres meses mediante muestreos de distancia con puntos en transectos en cada sitio para examinar los patrones de densidad y abundancia relativa. Los picos de floración se asociaron con la temporada, pero no con la precipitación ni con la elevación. Las densidades de aves presentaron un pico en el invierno y la primavera de cada año en las elevaciones altas, pero los patrones fueron -113 -The Auk 128(1):113 126,Abstract.-We documented patterns of nectar availability and nectarivorous bird abundance over ~ years at nine study sites across an ,-m elevational gradient on Hawaii Island to investigate the relationship between resource variation and bird abundance. Flower density (flowers ha − ) and nectar energy content were measured across the gradient for the monodominant `Ōhi`a (Metrosideros polymorpha). Four nectarivorous bird species were captured monthly in mist nets and surveyed quarterly with point-transect distance sampling at each site to examine patterns of density and relative abundance. Flowering peaks were associated with season but not rainfall or elevation. Bird densities peaked in the winter and spring of each year at high elevations, but patterns were less clear at middle and low elevations. Variability in bird abundance was generally best modeled as a function of elevation, season, and flower density, but the strength of the latter effect varied with species. The low elevations had the greatest density of flowers but contained far fewer individuals of the two most strongly nectarivorous species. There is little evidence of large-scale altitudinal movement of birds in response to `Ōhi`a flowering peaks. The loose relationship between nectar and bird abundance may be explained by a number of potential mechanisms, including () demographic constraints to movement; () nonlimiting nectar resources; and () the presence of an "ecological trap," whereby birds are attracted by the high resource abundance of, but suffer increased mortality at, middle and low elevations as a result of disease. Received  February , accepted  October .

1. Animal movement influences the spatial spread of directly transmitted wildlife disease through host-host contact structure. Wildlife disease hosts vary in home rangeassociated foraging and social behaviours, which may increase the spread and intensity of disease outbreaks. The consequences of variation in host home range movement and space use on wildlife disease dynamics are poorly understood, but could help to predict disease spread and determine more effective disease management strategies.2. We developed a spatially explicit individual-based model to examine the effect of spatiotemporal variation in host home range size on the spatial spread rate, persistence and incidence of rabies virus (RABV) in raccoons (Procyon lotor). We tested the hypothesis that variation in home range size increases RABV spread and decreases vaccination effectiveness in host populations following pathogen invasion into a vaccination zone.3. We simulated raccoon demography and RABV dynamics across a range of magnitudes and variances in weekly home range size for raccoons. We examined how variable home range size influenced the relative effectiveness of three components of oral rabies vaccination (ORV) programmes targeting raccoons-timing and frequency of bait delivery, width of the ORV zone and proportion of hosts immunized. 4. Variability in weekly home range size increased RABV spread rates by 1.2-fold to 5.2-fold compared to simulations that assumed a fixed home range size. More variable host home range sizes decreased relative vaccination effectiveness by 71% compared to less variable host home range sizes under conventional vaccination conditions. We found that vaccination timing was more influential for vaccination effectiveness than vaccination frequency or vaccination zone width. 5.Our results suggest that variation in wildlife home range movement behaviour increases the spatial spread and incidence of RABV. Our vaccination results underscore the importance of prioritizing individual-level space use and movement data collection to understand wildlife disease dynamics and plan their effective control and elimination. This is an open access article under the terms of the Creat ive Commo ns Attri bution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. . conceived the ideas and designed methodology; K.M.P. wrote the model code and conducted the simulations; K.M.P. and K.M.M. analysed the model results; and K.M.M. wrote the first draft of the manuscript. All authors provided critical feedback on the draft and gave final approval for publication.