Teira Research Articles

Drivers of continuous colour variation in the Madeiran wall lizard (Teira dugesii)

Abstract Animal coloration is often shaped by a myriad of factors that lead to differences in colour through changes in the chromatophores. Depending on how this variation is partitioned, coloration is often categorized as continuous or polymorphic. However, the boundaries between these two categories are not always clear. Here, we investigated whether the ventral coloration of the Madeiran wall lizard (Teira dugesii) varies continuously or corresponds to discrete colour morphs, via by-eye colour classification and visual modelling. By combining these two approaches, we show that T. dugesii coloration varies continuously and that colour classifications based on anthropomorphic approaches alone are ill suited to describe animal coloration. We also tested the influence of size, body condition, and sex as possible factors that might explain differences in T. dugesii coloration. We found that body condition, and especially size and sex, explain a great proportion of the variability observed in this species. These differences point to an effect of ontogeny, which might play a major role in colour development owing to the longevity of this species. Moreover, the sexual dichromatism that this species shows is indicative of an effect of sexual selection on coloration, perhaps explained by differences in circulating hormones.

Vertebrate damage to Azorean vineyards: the role of the endemic Azores Woodpigeon Columba palumbus azorica.

In the Azores, complaints about grape loss to birds have become recurrent. Although winegrowers frequently blame the endemic Azores Woodpigeon Columba palumbus azorica, data about the magnitude of grape damage and the species responsible for them are lacking. This study provides detailed information about grape damage caused by vertebrates on Pico Island, home to the main wine-growing area of the region. Vineyards were monitored during the ripening period in 2017 and 2018. Damage was assessed by determining the number of plucked, pecked and bitten grapes for a total of 850 bunches. Camera traps were placed in 113 enclosures providing detection and consumption rates of vertebrate species. GLM analysis was performed to test the effect of grape variety, distance to the woods, year and time before harvest on grape damage and consumption events. Damage was estimated at 10.3% (± 0.9) in 2017 and 8.7% (± 0.9) in 2018 and mostly consisted of plucked grapes. Ten vertebrate species were detected consuming grapes. Overall, 524 consumption events were registered. The Madeira lizard Teira dugesii, the Azores Blackbird Turdus merula azorensis and rodents (rats Rattus sp. and the house mouse Mus musculus) were responsible for most of those events. The Azores Woodpigeon accounted for three consumption events. GLM analysis showed that damage was greater in vines adjacent to woods and lower in traditional white varieties. This study constitutes an important baseline for the implementation of efficient measures to mitigate grape predation and sheds light on the minor role of the Azores Woodpigeon in grape damage.

Density and ecological drivers of free‐ranging cat abundance and activity in Madeira Island, Macaronesia

AbstractMammalian predators introduced to oceanic islands pose a significant threat to biodiversity and have led to numerous extinctions. Free‐ranging cats are particularly problematic due to their predatory habits and negative impact on conservation. However, there is limited information on the ecology and population status of free‐ranging cats in insular ecosystems, where they often represent the apex terrestrial predator. Using a peri‐urban protected area in the subtropical island of Madeira as a case study, we employed camera traps to assess the density of free‐ranging cats and investigate the ecological drivers influencing their abundance and activity in nonurban insular habitats. Based on 582 trapping‐nights, we identified 25 individual cats from 156 cat detections. Spatially explicit capture–recapture models revealed a density of 1.4 cats per km2. Cat activity was positively affected by both the proportion of rocky areas in the landscape and the distance to human resource subsidies, whereas no significant driver was found for abundance. Our results indicate that cats are highly abundant throughout the protected area and suggest that their core home ranges are associated with rocky terrain, away from the most humanized sections of the park. Free‐ranging cats do not appear to heavily rely on anthropogenic food sources, signaling that they may rely mostly on wild prey to fulfill their dietary needs. Their preference for rocky areas could be explained by the increased availability of shelter and prey, such as the Madeira wall lizard (Teira dugesii). Notably, cat abundance and activity were particularly high in the vicinity of the only known breeding colony of the locally threatened Manx shearwater (Puffinus puffinus) on Madeira Island. Our findings suggest that cats pose a significant threat to the native vertebrate fauna of the protected area and thus their management, particularly during the breeding season of the Manx shearwater, should be considered.

Habitat-associated genomic variation in a wall lizard from an oceanic island.

The lizard Teira dugesii exhibits morphological divergence between beach and inland habitats in the face of gene flow, within the volcanic island of Madeira, Portugal. Here, we analysed genomic data obtained by genotyping-by-sequencing, which provided 16378 SNPs from 94 individuals sampled from 15 sites across Madeira. Ancient within-island divergence in allopatry appears to have mediated divergence in similar species within other Atlantic islands but this hypothesis was not supported for T. dugesii. Across all samples, a total of 168 SNPs were classified as statistical outliers using pcadapt and OutFLANK. Redundancy analysis revealed that 17 of these outliers were associated with beach/inland habitats. The SNPs were located within 16 sequence tags and 15 of these were homologous with sequences in a 31Mb region on chromosome three of a reference wall lizard genome (the remaining tag could not be associated with any chromosome). We further-investigated outliers through contingency analyses of allele frequencies at each of four pairs of adjacent beach-inland sites. The majority of the outliers detected by the redundancy analysis were confirmed at two pairs of these matched sites. These analyses also suggested some parallel divergence at different localities. Six other outliers were associated with site elevation, four of which were located on chromosome five of the reference genome. Our study lends supports to a previous hypothesis that divergent selection between grey shingle beaches and inland regions overcomes gene flow and leads to the observed morphological divergence between populations in these adjacent habitats.

Factors shaping the gut microbiome of five species of lizards from different habitats.

Host-gut microbiota interactions are complex and can have a profound impact on the ecology and evolution of both counterparts. Several host traits such as systematics, diet and social behavior, and external factors such as prey availability and local environment are known to influence the composition and diversity of the gut microbiota. In this study, we investigate the influence of systematics, sex, host size, and locality/habitat on gut microbiota diversity in five lizard species from two different sites in Portugal: Podarcis bocagei and Podarcis lusitanicus, living in syntopy in a rural area in northern Portugal (Moledo); the invasive Podarcis siculus and the native Podarcis virescens, living in sympatry in an urbanized environment (Lisbon); and the invasive Teira dugesii also living in an urban area (Lisbon). We also infer the potential microbial transmission occurring between species living in sympatry and syntopy. To achieve these goals, we use a metabarcoding approach to characterize the bacterial communities from the cloaca of lizards, sequencing the V4 region of the 16S rRNA. Habitat/locality was an important factor explaining differences in gut bacterial composition and structure, with species from urbanized environments having higher bacterial diversity. Host systematics (i.e., species) influenced gut bacterial community structure only in lizards from the urbanized environment. We also detected a significant positive correlation between lizard size and gut bacterial alpha-diversity in the invasive species P. siculus, which could be due to its higher exploratory behavior. Moreover, estimates of bacterial transmission indicate that P. siculus may have acquired a high proportion of local microbiota after its introduction. These findings confirm that a diverse array of host and environmental factors can influence lizards' gut microbiota.

The Madeira lizard Teira dugesii may have the greatest population density of any terrestrial vertebrate

The Madeira lizard Teira dugesii may have the greatest population density of any terrestrial vertebrate

Life on a beach leads to phenotypic divergence despite gene flow for an island lizard

Limited spatial separation within small islands suggests that observed population divergence may occur due to habitat differences without interruption to gene flow but strong evidence of this is scarce. The wall lizard Teira dugesii lives in starkly contrasting shingle beach and inland habitats on the island of Madeira. We used a matched pairs sampling design to examine morphological and genomic divergence between four beach and adjacent (<1 km) inland areas. Beach populations are significantly darker than corresponding inland populations. Geometric morphometric analyses reveal divergence in head morphology: beach lizards have generally wider snouts. Genotyping-by-sequencing allows the rejection of the hypothesis that beach populations form a distinct lineage. Bayesian analyses provide strong support for models that incorporate gene flow, relative to those that do not, replicated at all pairs of matched sites. Madeiran lizards show morphological divergence between habitats in the face of gene flow, revealing how divergence may originate within small islands.

Dietary Variation Is Driven by Landscape Heterogeneity in an Insular Omnivorous Endemic Lizard, Revealed by DNA Metabarcoding

Living on islands entails numerous challenges for animals, among which population density approaching the carrying capacity of trophic resources stands out. To overcome this limitation, many insular lizards can supplement their insectivorous diet with increasing portions of plant material. The Madeira wall lizard, Teira dugesii, is a medium-sized lacertid, endemic to the Madeira and Selvagens archipelagos. As common in this family, adults are sexually dimorphic with males being bigger than females. Previous dietary studies on morphological scatology identified a higher proportion of plant over animal prey items, changing according to the location and sex. Here, we used DNA metabarcoding to examine the diet of this lizard species quantifying it at a higher taxonomical resolution and enhancing the detection of soft-body prey that often go undetected in morphology-based studies. In a sample of 151 faecal samples from eight populations including different habitats and altitudes in Madeira, we identified 289 prey items belonging to eight animal and three plant Classes, encompassing 58 distinct orders and 140 families. Of these, 63 were identified up to the species level. The results support a strong trend towards herbivory in this species with plants representing almost 74% of the diet occurrences in contrast to the 26% of animal prey. Remarkably, the plant fraction of the diet remained stable across localities but varied with size and mass in males. As males grew bigger and heavier, they significantly increased their plant matter intake. Likely, larger bodies and abdomens allowed allocating longer and more complex digestive tracts harbouring intestinal flora to better decompose plant organic compounds. This allowed heavier animals to have a richer diet regime. However, diet richness and composition were not affected by either sex or size, while the locality had a significant effect on both diet components likely in response to local variation in prey availability. By including an increasing plant fraction into a primarily insectivorous diet, this insular lizard has not only enlarged its trophic niche but is also able to exploit more efficiently the highly variable resources provided by insular environments.

Trophic interactions of an invasive gecko in an endemic-rich oceanic island: Insights using DNA metabarcoding

Understanding the trophic interactions of introduced predators is key for evidence-based management of biological invasions. This is particularly important in oceanic islands, where predator-prey networks often include numerous endemic and range-restricted species. Geckos are successful island colonizers and in recent years numerous species have established populations in a wide array of oceanic islands. One such species is the Moorish gecko (Tarentola mauritanica), which has colonized multiple islands across the Mediterranean basin, Caribbean and Macaronesia. The species was first reported in Madeira Island in 1993 and over the last 30 years has colonized most of the islands' southern coast and expanded to the nearby island of Porto Santo. Here, we used DNA metabarcoding to provide the first insights into the diet of this successful colonizer in its introduced range. The species' diet was mainly composed of ground-dwelling arthropods belonging to the families Porcellionidae (Isopoda), Julidae (Diplopoda) and Formicidae (Hymenoptera). The diet richness and composition were not affected by neither sex nor size of adult geckos, instead they both change across populations. However, trophic niche-width differed among size classes, with smaller geckos feeding on a wider range of prey. We identified over 160 different Operational Taxonomic Units in the diet of T. mauritanica, with 21.6% of them belonging to introduced invertebrates and 13.6% to native species. Native prey taxa included the endemic Madeira wall lizard (Teira dugesii), the sole native reptile to Madeira. We also detected several agricultural pests and disease vectors in the diet of this exotic predator, and 19 taxa identified as prey had not yet been recorded to Madeira. Of these, several are serious agricultural pests, highlighting how this introduced gecko can be used as a natural sampler, in particular for the early detection of invasive arthropod pests. This study emphasizes the importance of trophic studies for monitoring the impacts of introduced predators in fragile insular systems.

Diet of the exotic Madeiran wall lizard: first insights into trophic interactions in an Atlantic seabird sanctuary

The Madeiran wall lizard Teira dugesii is a relatively new species to the Azores Archipelago, where it was accidentally introduced about 150 to 200 years ago. This lacertid quickly became naturalised and now occurs in all the nine main islands of the Azores. At Praia Islet, off Graciosa Island, the Madeiran wall lizard was recently observed preying on chicks of the threatened Monteiro’s storm-petrel Hydrobates monteiroi. To characterise the lizards’ trophic niche, we conducted a study of the diet of the Madeiran wall lizard at Praia using stable isotope analysis and next-generation sequencing. Our results indicate that the Madeiran wall lizard has a varied diet, consisting of at least 23 invertebrate taxa, 17 plant species, and occasionally, birds (two species detected, including storm-petrels). Marine derived food items were present in few samples, and it is vital to monitor the trophic interactions at Praia Islet to ensure the conservation of this threatened storm-petrel.

Foraging Behavior and Pollen Transport by Flower Visitors of the Madeira Island Endemic Echium candicans.

Simple SummaryThe successful conservation of many endangered island plants depends on the pollination services provided by animals. In this study, we identify the flower visitors of Echium candicans, a charismatic plant exclusive to the island of Madeira, and also evaluate their performance as pollinators by analyzing their behavior on the flowers and the pollen they transport on their body. We found that many different animals visit this plant’s flowers, from insects to reptiles, but bees were the most frequent visitors. Large bees visited more flowers and transported more pollen of Echium candicans compared to other pollinators, like butterflies and hoverflies. However, by visiting many flowers in the same plant large bees might contribute to inbreeding, whilst the other animals visited fewer flowers in each plant favoring outcrossing. We conclude that the different flower visitors of Echium candicans provide complementary services as pollinators and highlight the importance of having diverse communities of pollinators to ensure successful pollination in many island plants.The study of flower visitor behavior and pollen transport dynamics within and between plants can be of great importance, especially for threatened or rare plant species. In this work, we aim to assess the flower visitor assemblage of the Madeiran endemic Echium candicans and evaluate the performance of the most common visitors through the analysis of their foraging behavior and pollen loads. The flower visitor assemblage of E. candicans is diverse, including several insect groups and the endemic lizard Teira dugesii, but bees are the most common visitors. In general, large bees (Amegilla quadrifasciata, Apis mellifera, and Bombus spp.) had the highest average visitation rates (>18 flowers/min) and their pollen loads had higher percentages of homospecific pollen (>66%) when compared with butterflies and hoverflies. The honeybee (Apis mellifera) and two bumblebees (Bombus terrestris and B. ruderatus) were the most efficient flower visitors of E. candicans, but their foraging behavior seems to favor geitonogamy. Other visitors, such as butterflies and the small bee Lasioglossum wollastoni, may have a complementary role to the honeybee and bumblebee species, as their high mobility is associated with fewer flower visits on each plant and may promote xenogamy. Two non-native bees (A. mellifera and B. ruderatus) are important flower visitors of E. candicans and may contribute mostly to self-pollination rendering the endemic plant more vulnerable to inbreeding effects.

Exceptional longevity of a male Madeira rock lizard Teira dugesii maintained in captivity for 41 years

Exceptional longevity of a male Madeira rock lizard Teira dugesii maintained in captivity for 41 years

Using camera-trapping to assess grape consumption by vertebrate pests in a World Heritage vineyard region

Vertebrate pests cause significant economic loss in several agricultural crops worldwide; therefore, their populations are often controlled through culling. Correctly identifying the main species responsible for the damage is essential to avoid persecuting the wrong targets, yet it is challenging. During 2016 and 2017, we tested the usefulness of camera-trapping (CT) in Terceira Island, Azores, Portugal to identify vertebrate pests. Vineyard owners in this region cull the Azores woodpigeon (Columba palumbus azorica), the common pigeon (Columba livia), the house sparrow (Passer domesticus), the Azores blackbird (Turdus merula azorensis), and the Madeira wall lizard (Teira dugesii) to reduce damage to grapes. Using CT photos and videos, we identified nine species damaging the grape, but four of those were only observed occasionally (< 10 consumption events over 2 years). The Madeira lizard (371.09 and 232.47 consumption events100-CT-days in 2016 and 2017, respectively), the house sparrow (284.01 and 21.73 consumption events100-CT-days in 2016 and 2017, respectively), and the Azores blackbird (17.35 and 8.23 consumption events100-CT-days in 2016 and 2017, respectively) had the most frequent consumption events. All three species were most active in the morning (8:00–9:00) and in the afternoon (16:00–17:00 for the Madeira lizard and the house sparrow, and 18:00–19:00 for the Azores blackbird). We demonstrated the advantage of using CT in cultivated habitats to provide valuable information about the identity, behaviour, daily-activity patterns, and relative consumption rates of vineyard pests. We also provided evidence that the endemic Azores woodpigeon and the common pigeon should not be targeted by the farmers in Terceira.

Questioning the proverb 'more haste, less speed': classic versus metabarcoding approaches for the diet study of a remote island endemic gecko.

Dietary studies can reveal valuable information on how species exploit their habitats and are of particular importance for insular endemics conservation as these species present higher risk of extinction. Reptiles are often neglected in island systems, principally the ones inhabiting remote areas, therefore little is known on their ecological networks. The Selvagens gecko Tarentola (boettgeri) bischoffi, endemic to the remote and integral reserve of Selvagens Archipelago, is classified as Vulnerable by the Portuguese Red Data Book. Little is known about this gecko’s ecology and dietary habits, but it is assumed to be exclusively insectivorous. The diet of the continental Tarentola species was already studied using classical methods. Only two studies have used next-generation sequencing (NGS) techniques for this genus thus far, and very few NGS studies have been employed for reptiles in general. Considering the lack of information on its diet and the conservation interest of the Selvagens gecko, we used morphological and DNA metabarcoding approaches to characterize its diet. The traditional method of morphological identification of prey remains in faecal pellets collected over a longer period was compared with metabarcoding of samples collected during rapid surveys. Molecular results revealed that this species is a generalist, feeding on invertebrate, plant and vertebrate items, whereas the morphological approaches were unable to detect the latter two. These results opened up new questions on the ecological role of the Selvagens gecko that deserves to be further explored, such as the possible predation on seabirds, plant services or trophic competition with the sympatric Madeira lizard Teira dugesii. Metabarcoding identified a greater diversity of dietary items at higher taxonomic resolution, but morphological identification enabled calculation of relative abundances and biomasses of ingested arthropods, and detected a dietary shift on invertebrate preys between seasons. Results of this study highlight the global applicability of rapid metabarcoding surveys for understudied taxa on remote islands that are difficult to access. We recommend using the metabarcoding approach, even if ‘speedy’ sampling only is possible, but we must highlight that disregarding long-term ecological data may lead to ‘hasty’ conclusion.

Depredation of Monteiro's Storm-Petrel (Hydrobates monteiroi) Chicks by Madeiran Wall Lizards(Lacerta dugesii)

Monteiro's Storm-petrel (Hydrobates monteiroi) is a small seabird endemic to the Azores archipelago, where it is known to breed only on two mammal-free islets. Total breeding numbers are between 250 and 300 pairs. During the 2015 breeding season on Praia Islet, six Madeiran wall lizards (Lacerta dugesii) were observed depredating a Monteiro's Storm-petrel chick about to fledge. In August 2015, a freshly killed chick without its head and four larger dead chicks were found. Eight other chicks had disappeared from their nests since our previous visit in early July. In 2016, there was further evidence of suspected lizard depredation. Three chicks were found predated and dragged out of their nests, and an additional 22 chicks disappeared from their nests before fledging. In 2016, Monteiro's Storm-petrel breeding success was only 26.8%, the lowest ever recorded. We suspect that some of the chick deaths and nest failures may be attributed to these introduced lizards. All chicks were found dragged out of their nests, a behavior that is not consistent with kills from aerial predators. Lizards were suspected because they are the only possible predator on land. Therefore, we recommend evaluating the impact of Madeiran wall lizards on the two Monteiro's Storm-petrel main colonies: Praia and Baixo Islets.

Response to Seán Manning’s Article: “Why Psychotherapy Must Be Secular”

This article proposes that rather than being inherently dangerous, religions originate in times when people experience inter-tribal wars or other existential anxieties, and evolve in tandem with societal developments; as people become more peaceful, so do their religions. Based on Volkan’s (1985, 2006) analysis of psychological aspects of the conflict after 9/11 between America and some Muslims, it is suggested that religion serves as an ethnic and national identifier rather than being the source of the conflict. Jung’s idea that “gods” correspond to unconscious psychic factors rather than existing as entities is cited.&#x0D; Waitara&#x0D; E kī ana tēnei tuhinga ehara i te ira mōrearea, engari ahu kē mai ai ngā hāhi i ngā putanga mai o ngā wheako mai i ngā riri-ā-iwi ki te iwi o ētahi atu anipānga kē atu rānei, ka putaputa haere ake i te whanaketanga hāpori; ka hūmāriē haere ake te tangata, ka pērā anō hoki ō rātau hāhi. Whakapapa atu ki tā Wākena (1985, 2006) tātarihanga i ētahi wāhanga mātai whaiaroaro o te taupatupatu i muri mai i 9/11 i waenga i a Ameika me ētahi Muhirama, ka puta ake te whakaaro he wāhi tuku momo tangata whakaatu iwi tangata kē te hāhi, ehara i te pūtaketanga o te riri. Ka whakahauhia ake te kī a Jung, he rite kē ngā “atua” ki ngā aranga kauwaka mauri moe kaua ki ngā kitenga kanohi.

Te Ira Tangata: a Zelen randomised controlled trial of a culturally informed treatment compared to treatment as usual in Māori who present to hospital after self-harm.

Indigenous people have high rates of suicide and self-harm compared to other population groups. The aim of this trial was to see if a package of interventions delivered in a culturally appropriate way improved outcomes at one year in Maori who presented with intentional self-harm to emergency departments. Participants were Maori who presented with intentional self-harm to emergency departments in New Zealand. The study design was a double consent Zelen randomised controlled trial. The intervention included regular postcards, problem solving therapy, patient support, risk management, improved access to primary care and cultural assessment in addition to usual care. The control group received usual care. The main outcome measure was the self-rated change in scores on the Beck Hopelessness Scale at one year. 182 people were randomized to the intervention group 95 of whom consented to take part in the study. 183 people were randomized to the control group 72 of whom consented to take part in the study. For those who consented comparing the intervention group with the control group there was a statistically significant greater change in hopelessness scores at 3months (difference -1.7 95% CI -3.4 to -0.01, p=0.05) but not at one year (difference -1.6 95% CI -3.4 to 0.3, p=0.11). Maori who consented and received the intervention were also significantly less likely to present to hospital for non-self-harm reasons in the year after the index episode (44.2 vs. 61.1%, p=0.03). Those participants randomised to the intervention were less likely to re-present with self-harm at 3months (10.4 vs. 18%, p=0.04) but not at 12months compared to the control group. In Maori who presented to hospital with intentional self-harm a culturally informed intervention had an effect on hopelessness and re-presentation with self-harm in the short term but not at 12months. There was a significant decrease in hospital presentations for non-self harm over the next year. Australian and New Zealand Clinical Trials Registry ACTRN12609000952246.

To Move or Not to Move: Cranial Joints in European Gekkotans and Lacertids, an Osteological and Histological Perspective

Lepidosaurs are frequently described as having highly kinetic skulls, and different forms of cranial kinesis have been described as being characteristic of their radiation. The model of amphikinesis proposed by Frazzetta, J Morphol 1962; 111:287-319, which was long considered a synapomorphy of the large suborder Sauria, is now much debated given its uncertain distribution among the various lizard taxa and the lack of data about its morphological correlates. In this article, we analyze the anatomical correlates of different forms of cranial kinesis, with particular regard to the putative saurian amphikinesis, describing the possible diverse skull movements of several species of European gekkotans (Hemidactylus turcicus, Mediodactylus kotschyi, and Tarentola mauritanica) and lacertids (Lacerta agilis, L. bilineata, Podarcis muralis, P. siculus, and Teira dugesii). Using serial and whole-mount histology, we found clear differences between gekkotans and lacertids in the structure of several cranial joints underlining the existence of two degrees of intracranial mobility. The lacertid species possess the anatomical features for streptostyly (quadrate joints) and metakinesis (parietal-supraoccipital and parabasisphenoid-pterygoid joints) and lack the anatomical correlates for mesokinesis (mobility of frontal-parietal and palatine-pterygoid joints) and amphikinesis (coupled mesokinesis, metakinesis, and streptostyly). In contrast, geckos present all the anatomical correlates for amphikinesis as described by the traditional quadratic crank model. Finally, we present a comprehensive summary of the different forms of squamate cranial kinesis, advancing two alternative hypotheses about the evolutionary origin of amphikinesis.

Evidence of high longevity in an Island lacertid, Teira dugesii (Milne-Edwards, 1829). First data on wild specimens

Using the technique of capture, mark and recapture, here is reported a case of high longevity in the Madeiran lizard, Teira dugesii. It is one of the highest reported values for lacertid lizards (16 years or more) in a wild population.

Role of the Lizard Teira dugesii as a Potential Host for Ixodes ricinus Tick-Borne Pathogens

PCR screening of ticks and tissue samples collected from 151 Teira dugesii lizards seems to indicate a potential role of this lizard species in the maintenance and transmission cycle of some Ixodes ricinus tick-borne agents, such as Rickettsia monacensis, Rickettsia helvetica, and Borrelia lusitaniae, that are circulating on Madeira Island.