Lizards and small rodents were live captured in Tuscany, central Italy, from May through August 2005. Prevalence of infestation by larval Ixodes ricinus L. (Acari: Ixodidae) and mean numbers of larvae per host were not significantly different for common wall lizards, Podarcis muralis Laurenti, and Apodemus spp. mice, whereas infestation levels by nymphs were significantly greater on lizards. Borrelia lusitaniae, which was previously shown to be dominant in host-seeking I. ricinus in the same study area, was detected by polymerase chain reaction (PCR) in 19.8% (95% confidence interval: 14.4, 26.0) of larval ticks and in 52.9% (27.8, 77.0) of nymphs that were collected from lizards. Moreover, 18.8% (7.2, 36.4) and 25.0 (3.2, 65.1) of lizards' tail biopsies and blood samples, respectively, were positive for B. lusitaniae. Conversely, attached ticks and ear biopsies from Apodemus spp. mice were PCR negative. Passerine birds belonging to 10 species were live captured in March 2005, and Borrelia valaisiana was detected in 57.1% (18.4, 90.1) of I. ricinus nymphs feeding on Eurasian blackbirds, Turdus merula L. Results of this study suggest that lizards play an important role as reservoirs for B. lusitanae and may affect the dominance of this genospecies in the Mediterranean area.

BackgroundVector-borne diseases for which transmission occurs exclusively between vectors and hosts can be modeled as spreading on a bipartite network.Methodology/Principal FindingsIn such models the spreading of the disease strongly depends on the degree distribution of the two classes of nodes. It is sufficient for one of the classes to have a scale-free degree distribution with a slow enough decay for the network to have asymptotically vanishing epidemic threshold. Data on the distribution of Ixodes ricinus ticks on mice and lizards from two independent studies are well described by a scale-free distribution compatible with an asymptotically vanishing epidemic threshold. The commonly used negative binomial, instead, cannot describe the right tail of the empirical distribution.Conclusions/SignificanceThe extreme aggregation of vectors on hosts, described by the power-law decay of the degree distribution, makes the epidemic threshold decrease with the size of the network and vanish asymptotically.

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Host-seeking ticks were collected during monthly dragging sessions from November 2004 through October 2006 in Tuscany, central Italy. Normalized difference vegetation index (NDVI), which was calculated from Landsat ETM+ 7 remote sensing data recorded in August 2001, was significantly correlated with numbers of host-seeking immature Ixodes ricinus L. (Acari: Ixodidae) during periods of relatively low rainfall such as summer 2005 (Spearman's p = 0.78, P < 0.001 for nymphs in July) and to a lower extent in spring-summer 2006. In spring 2005, when rainfall was relatively high, the correlation was weak and not statistically significant. Generalized estimating equations (GEEs), taking into account repeated sampling of the same dragging sites, were used to model the effects of NDVI and season on counts of host-seeking I. ricinus nymphs. Seasonal variations of the effect of NDVI yielded a significant NDVI-by-season interaction in the first year of the study (November 2004-October 2005), but not in the second year (November 2005-October 2006) when there was a 2.5-fold increase of the number of nymphs per 100-m dragging for every 0.1 unit increase in NDVI (95% confidence interval = 1.6, 3.0). Risk maps that were obtained based on GEE results confirmed that the predicted number of I. ricinus nymphs per 100 m was relatively homogeneous through the study area during the 2005 spring peak of activity. Conversely, in 2006, the predicted abundance of nymphs was greater in moist bottomland habitat (characterized by high NDVI) than in dry, typically Mediterranean, upland habitat.

To estimate the basic reproduction number (R(0)) of Borrelia lusitaniae and Borrelia afzelii, we formulated a mathematical model considering the interactions among the tick vector, vertebrate hosts, and pathogens in a 500-ha enclosed natural reserve on Le Cerbaie hills, Tuscany, central Italy. In the study area, Ixodes ricinus were abundant and were found infected by B. lusitaniae and B. afzelii. Lizards (Podarcis spp.) and mice (Apodemus spp.), respectively, are the reservoir hosts of these two Borrelia burgdorferi sensu lato (s.l.) genospecies and compete for immature ticks. B. lusitaniae R(0) estimation is in agreement with field observations, indicating the maintenance and diffusion of this genospecies in the study area, where lizards are abundant and highly infested by I. ricinus immature stages. In fact, B. lusitaniae shows a focal distribution in areas where the tick vector and the vertebrate reservoir coexist. Mouse population dynamics and their relatively low suitability as hosts for nymphs seem to determine, on the other hand, a less efficient transmission of B. afzelii, whose R(0) differs between scenarios in the study area. Considering host population dynamics, the proposed model suggests that, given a certain combination of the two host population sizes, both spirochete genospecies can coexist in our study area. Additional incompetent hosts for B. burgdorferi s.l. have a negative effect on B. afzelii maintenance, whose R(0) results > 1 only with high mouse population densities and/or low lizards abundance, but they do not seem to influence B. lusitaniae transmission cycle on Le Cerbaie. Secondly, our model confirms the importance of nymphs' infestation, of host population density and diversity, and spirochetes host association for the maintenance of the transmission cycle of B. burgdorferi s.l.

Lizards and small rodents were live captured in Tuscany, central Italy, from May through August 2005. Prevalence of infestation by larval Ixodes ricinus L. (Acari: Ixodidae) and mean numbers of larvae per host were not significantly different for common wall lizards, Podarcis muralis Laurenti, and Apodemus spp. mice, whereas infestation levels by nymphs were significantly greater on lizards. Borrelia lusitaniae, which was previously shown to be dominant in host-seeking I. ricinus in the same study area, was detected by polymerase chain reaction (PCR) in 19.8% (95% confidence interval: 14.4, 26.0) of larval ticks and in 52.9% (27.8, 77.0) of nymphs that were collected from lizards. Moreover, 18.8% (7.2, 36.4) and 25.0 (3.2, 65.1) of lizards' tail biopsies and blood samples, respectively, were positive for B. lusitaniae. Conversely, attached ticks and ear biopsies from Apodemus spp. mice were PCR negative. Passerine birds belonging to 10 species were live captured in March 2005, and Borrelia valaisiana was detected in 57.1% (18.4, 90.1) of I. ricinus nymphs feeding on Eurasian blackbirds, Turdus merula L. Results of this study suggest that lizards play an important role as reservoirs for B. lusitanae and may affect the dominance of this genospecies in the Mediterranean area.

During the investigations on ticks and tick-borne pathogens (TBP) range expansion in the Northern Apennines, we captured 107 Podarcis muralis lizards. Sixtyeight animals were infested by immature Ixodes ricinus, Haemaphysalis sulcata and H. punctata. Borrelia burgdorferi s.l. was detected in 3.7% of I. ricinus larvae and 8.0% of nymphs. Together with the species-specific B. lusitaniae, we identified B. garinii, B. afzelii and B. valaisiana. Rickettsia spp. (18.1% larvae, 12.0% nymphs), namely R. monacensis, R. helvetica and R. hoogstraalii, were also found in I. ricinus. R. hoogstraalii was detected in H. sulcata nymphs as well, while the two H. punctata did not harbour any bacteria. One out of 16 lizard tail tissues was positive to R. helvetica. Our results support the hypothesis that lizards are involved in the epidemiological cycles of TBP. The heterogeneity of B. burgdorferi genospecies mirrors previous findings in questing ticks in the area, and their finding in attached I. ricinus larvae suggests that lizards may contribute to the maintenance of different genospecies. The rickettsiae are new findings in the study area, and R. helvetica infection in a tail tissue indicates a systemic infection. R. hoogstraalii is reported for the first time in I. ricinus ticks. Lizards seem to favour the bacterial exchange among different tick species, with possible public health consequences.