Bridging the gap between movement data and connectivity analysis using the Time-Explicit Habitat Selection (TEHS) model.

BACKGROUND: Understanding how to connect habitat remnants to facilitate the movement of species is a critical task in an increasingly fragmented world impacted by human activities. The identification of dispersal routes and corridors through connectivity analysis requires measures of landscape resistance but there has been no consensus on how to calculate resistance from habitat characteristics, potentially leading to very different connectivity outcomes. METHODS: We propose a new model, called the Time-Explicit Habitat Selection (TEHS) model, that can be directly used for connectivity analysis. The TEHS model decomposes the movement process in a principled approach into a time and a selection component, providing complementary information regarding space use by separately assessing the drivers of time to traverse the landscape and the drivers of habitat selection. These models are illustrated using GPS-tracking data from giant anteaters (Myrmecophaga tridactyla) in the Pantanal wetlands of Brazil. RESULTS: The time model revealed that the fastest movements tended to occur between 8 p.m. and 5 a.m., suggesting a crepuscular/nocturnal behavior. Giant anteaters moved faster over wetlands while moving much slower over forests and savannas, in comparison to grasslands. We also found that wetlands were consistently avoided whereas forest and savannas tended to be selected. Importantly, this model revealed that selection for forest increased with temperature, suggesting that forests may act as important thermal shelters when temperatures are high. Finally, using the spatial absorbing Markov chain framework, we show that the TEHS model results can be used to simulate movement and connectivity within a fragmented landscape, revealing that giant anteaters will often not use the shortest-distance path to the destination patch due to avoidance of certain habitats. CONCLUSIONS: The proposed approach can be used to characterize how landscape features are perceived by individuals through the decomposition of movement patterns into a time and a habitat selection component. Additionally, this framework can help bridge the gap between movement-based models and connectivity analysis, enabling the generation of time-explicit connectivity results.

Behavioral responses of terrestrial mammals to COVID-19 lockdowns.

COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals' 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.

Biologging as an important tool to uncover behaviors of cryptic species: an analysis of giant armadillos (Priodontes maximus).

Advances in biologging have increased the understanding of how animals interact with their environment, especially for cryptic species. For example, giant armadillos (Priodontes maximus) are the largest extant species of armadillo but are rarely encountered due to their fossorial and nocturnal behavior. Through the analysis of speed, turning angles, and accelerometer activity counts, we estimated behavioral states, characterized activity budgets, and investigated the state-habitat associations exhibited by individuals monitored with GPS telemetry in the Brazilian Pantanal from 2019 to 2020. This methodology is proposed as a useful framework for the identification of priority habitat. Using the non-parametric Bayesian mixture model for movement (M3), we estimated four latent behavioral states that were named 'vigilance-excavation', 'local search', 'exploratory', and 'transit'. These states appeared to correspond with behavior near burrows or termite mounds, foraging, ranging, and rapid movements, respectively. The first and last hours of activity presented relatively high proportions of the vigilance-excavation state, while most of the activity period was dominated by local search and exploratory states. The vigilance-excavation state occurred more frequently in regions between forest and closed savannas, whereas local search was more likely in high proportions of closed savanna. Exploratory behavior probability increased in areas with high proportions of both forest and closed savanna. Our results establish a baseline for behavioral complexity, activity budgets, and habitat associations in a relatively pristine environment that can be used for future work to investigate anthropogenic impacts on giant armadillo behavior and fitness. The integration of accelerometer and GPS-derived movement data through our mixture model has the potential to become a powerful methodological approach for the conservation of other cryptic species.

What a few hairs can tell us about the resource use of giant armadillos.

Understanding of resource requirements of vulnerable species is key for conservation planning. Here, we used stable carbon (δ13 C) and nitrogen isotopes (δ15 N) of hair from giant armadillos (Priodontes maximus) to understand individual resource use. We collected hair from 19 giant armadillos analyzing 34 samples from the Brazilian Pantanal and compared the resource use between sexes and age classes. We also assessed the relationship of isotopic values with individuals' body mass and the habitat type used. We observed a wide variation in resource use by giant armadillos, showing that individuals consume prey that feeds on distinct resources and occupies different habitat types, indicating that there is individual variation in foraging behavior. Most giant armadillos presented a mixed diet (C3 /C4 resources, forests/open areas), but a quarter of individuals had strictly C4 diets (open areas). Males are more prone to forage on C4 resources, presenting an isotopic niche 2.6-times larger than that of females. Subadults presented diets more associated with habitats with high arboreal cover (C3 resources, forests), while adults foraged more in open areas (C4 resources). This result is mirrored by the positive relationship between δ13 C values and body mass, suggesting that larger giant armadillos tend to feed more in open areas. We observed that δ13 C values decreased as individuals increased the use of habitats with high arboreal cover. We stress the importance of conserving the natural mosaic in the Pantanal landscape for the species persistence, given that giant armadillos require all habitat types to fulfill their dietary, spatial, and developmental needs.

Evaluating expert-based habitat suitability information of terrestrial mammals with GPS-tracking data.

Aim: Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert-based information is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert-based information with detailed empirical evidence. Here, we compared expert-based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individuals from 49 mammal species. Location: Worldwide. Time period: 1998-2021. Major taxa studied: Forty-nine terrestrial mammal species. Methods: Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habitat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS-based habitat suitability measures were in agreement with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types. Results: IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively. Main conclusions: We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species included in this study, it is appropriate to use IUCN habitat suitability data in macroecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data.

The role of environmental temperature on movement patterns of giant anteaters.

Mammals can show conspicuous behavioral responses to thermal variation, including changes in movement patterns. We used an integrative approach to understand how environmental temperature can drive the movement behavior of a mammal with low capacity for physiological thermoregulation, the giant anteater (Myrmecophaga tridactyla). We tracked 52 giant anteaters in 7 areas throughout the Brazilian savannah. We estimated the distance moved, area used, use of forest areas, and mean environmental temperature for each monitoring day of each individual. We modeled these data with Mixed Structural Equations - considering the possible interactions between our variables and controlling for sex and body mass. Giant anteaters reduced displacement and increased forest use with decreasing environmental temperature, probably because of their low body heat production. It is possible that they reduce distance moved and area used by reducing the duration of activity. With decreasing temperature, forest habitats become warmer than open ones, besides buffer rain and chilly winds. Reducing displacement and using forests are important strategies to reduce body heat loss and the energetic costs of thermoregulation. However, decreasing movement can limit food access and, consequently, fitness. Therefore, we highlight the importance of forests as thermal shelters. With increasing frequency and intensity of extreme weather events, we showed the need to preserve forest patches to offer suitable conditions for tropical mammals' behavioral thermoregulation. In this context, policies favoring deforestation on Brazilian territory are especially worrisome. Finally, we emphasize the need of integrative approaches to understand the complex interactions between organisms and the environment.

Automatic selection of the number of clusters using Bayesian clustering and sparsity-inducing priors.

Clustering is a ubiquitous task in ecological and environmental sciences and multiple methods have been developed for this purpose. Because these clustering methods typically require users to a priori specify the number of groups, the standard approach is to run the algorithm for different numbers of groups and then choose the optimal number using a criterion (e.g., AIC or BIC). The problem with this approach is that it can be computationally expensive to run these clustering algorithms multiple times (i.e., for different numbers of groups) and some of these information criteria can lead to an overestimation of the number of groups. To address these concerns, we advocate for the use of sparsity-inducing priors within a Bayesian clustering framework. In particular, we highlight how the truncated stick-breaking (TSB) prior, a prior commonly adopted in Bayesian nonparametrics, can be used to simultaneously determine the number of groups and estimate model parameters for a wide range of Bayesian clustering models without requiring the fitting of multiple models. We illustrate the ability of this prior to successfully recover the true number of groups for three clustering models (two types of mixture models, applied to GPS movement data and species occurrence data, as well as the species archetype model) using simulated data in the context of movement ecology and community ecology. We then apply these models to armadillo movement data in Brazil, plant occurrence data from Alberta (Canada), and bird occurrence data from North America. We believe that many ecological and environmental sciences applications will benefit from Bayesian clustering methods with sparsity-inducing priors given the ubiquity of clustering and the associated challenge of determining the number of groups. Two R packages, EcoCluster and bayesmove, are provided that enable the straightforward fitting of these models with the TSB prior.

Sexual, allometric and forest cover effects on giant anteaters' movement ecology.

Knowing the influence of intrinsic and environmental traits on animals' movement is a central interest of ecology and can aid to enhance management decisions. The giant anteater (Myrmecophaga tridactyla) is a vulnerable mammal that presents low capacity for physiological thermoregulation and uses forests as thermal shelters. Here, we aim to provide reliable estimates of giant anteaters' movement patterns and home range size, as well as untangle the role of intrinsic and environmental drivers on their movement. We GPS-tracked 19 giant anteaters in Brazilian savannah. We used a continuous-time movement model to estimate their movement patterns (described by home range crossing time, daily distance moved and directionality), and provide an autocorrelated kernel density estimate of home range size. Then, we used mixed structural equations to integratively model the effects of sex, body mass and proportion of forest cover on movement patterns and home range size, considering the complex net of interactions between these variables. Male giant anteaters presented more intensive space use and larger home range than females with similar body mass, as it is expected in polygynous social mating systems. Males and females increased home range size with increasing body mass, but the allometric scaling of intensity of space use was negative for males and positive for females, indicating different strategies in search for resources. With decreasing proportion of forest cover inside their home ranges, and, consequently, decreasing thermal quality of their habitat, giant anteaters increased home range size, possibly to maximize the chances of accessing thermal shelters. As frequency and intensity of extreme weather events and deforestation are increasing, effective management efforts need to consider the role of forests as an important thermal resource driving spatial requirements of this species. We highlight that both intrinsic and environmental drivers of animal movement should be integrated to better guide management strategies.

Zoonotic parasites infecting free-living armadillos from Brazil.

Armadillos are specialist diggers and their burrows are used to find food, seek shelter and protect their pups. These burrows can also be shared with dozens of vertebrate and invertebrate species and; consequently, their parasites including the zoonotics. The aim of this study was to diagnose the presence of zoonotic parasites in four wild-caught armadillo species from two different Brazilian ecosystems, the Cerrado (Brazilian savanna) and the Pantanal (wetland). The investigated parasites and their correspondent diseases were: Toxoplasma gondii (toxoplasmosis), Trypanosoma cruzi (Chagas disease), Leishmania spp., (leishmaniasis), Paracoccidioides brasiliensis (Paracoccidioidomicosis) and Mycobacterium leprae (Hansen's disease). Forty-three free-living armadillos from Pantanal and seven road-killed armadillos from the Cerrado were sampled. Trypanosoma cruzi DTU TcIII were isolated from 2 out of 43 (4.65%) armadillos, including one of them also infected with Trypanosoma rangeli. Antibodies anti-T. gondii were detected in 13 out of 43 (30.2%) armadillos. All seven armadillos from Cerrado tested positive for P. brasiliensis DNA, in the lungs, spleen, liver fragments. Also, by molecular analysis, all 43 individuals were negative for M. leprae and Leishmania spp. Armadillos were infected by T. cruzi, T. rangeli, P. brasiliensis and presented seric antibodies to T. gondii, highlighting the importance of those armadillos could have in the epidemiology of zoonotic parasites.

Effects of body size on estimation of mammalian area requirements.

Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home-range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied block cross-validation to quantify bias in empirical home-range estimates. Area requirements of mammals <10 kg were underestimated by a mean approximately15%, and species weighing approximately100 kg were underestimated by approximately50% on average. Thus, we found area estimation was subject to autocorrelation-induced bias that was worse for large species. Combined with the fact that extinction risk increases as body mass increases, the allometric scaling of bias we observed suggests the most threatened species are also likely to be those with the least accurate home-range estimates. As a correction, we tested whether data thinning or autocorrelation-informed home-range estimation minimized the scaling effect of autocorrelation on area estimates. Data thinning required an approximately93% data loss to achieve statistical independence with 95% confidence and was, therefore, not a viable solution. In contrast, autocorrelation-informed home-range estimation resulted in consistently accurate estimates irrespective of mass. When relating body mass to home range size, we detected that correcting for autocorrelation resulted in a scaling exponent significantly >1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum.

Efectos del Tamaño Corporal sobre la Estimación de los Requerimientos de Área de Mamíferos Resumen La cuantificación precisa de los requerimientos de área de una especie es un prerrequisito para que la conservación basada en áreas sea efectiva. Esto comúnmente implica la recolección de datos de rastreo de la especie de interés para después realizar análisis de la distribución local. De manera problemática, la autocorrelación en los datos de rastreo puede resultar en una subestimación grave de las necesidades de espacio. Con base en trabajos previos, formulamos una hipótesis en la que supusimos que la magnitud de la subestimación varía con la masa corporal, una relación que podría tener implicaciones serias para la conservación. Para probar esta hipótesis en mamíferos terrestres, estimamos las áreas de distribución local con las ubicaciones en GPS de 757 individuos de 61 especies de mamíferos distribuidas mundialmente con una masa corporal entre 0.4 y 4,000 kg. Después aplicamos una validación cruzada en bloque para cuantificar el sesgo en estimaciones empíricas de la distribución local. Los requerimientos de área de los mamíferos <10 kg fueron subestimados por una media ∼15% y las especies con una masa ∼100 kg fueron subestimadas en ∼50% en promedio. Por lo tanto, encontramos que la estimación del área estaba sujeta al sesgo inducido por la autocorrelación, el cual era peor para las especies de talla grande. En combinación con el hecho de que el riesgo de extinción incrementa conforme aumenta la masa corporal, el escalamiento alométrico del sesgo que observamos sugiere que la mayoría de las especies amenazadas también tienen la probabilidad de ser aquellas especies con las estimaciones de distribución local menos acertadas. Como corrección, probamos si la reducción de datos o la estimación de la distribución local informada por la autocorrelación minimizan el efecto de escalamiento que tiene la autocorrelación sobre las estimaciones de área. La reducción de datos requirió una pérdida de datos del ∼93% para lograr la independencia estadística con un 95% de confianza y por lo tanto no fue una solución viable. Al contrario, la estimación de la distribución local informada por la autocorrelación resultó en estimaciones constantemente precisas sin importar la masa corporal. Cuando relacionamos la masa corporal con el tamaño de la distribución local, detectamos que la corrección de la autocorrelación resultó en un exponente de escalamiento significativamente >1, lo que significa que el escalamiento de la relación cambió sustancialmente en el extremo superior del espectro de la masa corporal.

ANESTHESIA AND SURGERY PROTOCOLS FOR INTRA-ABDOMINAL TRANSMITTER PLACEMENT IN FOUR SPECIES OF WILD ARMADILLO.

The anatomy and physiology of armadillos make anesthetic procedures and the placement of telemetry devices challenging. From June 2011 to November 2019, a total of 73 free-living armadillos were captured and anesthetized in the Pantanal, Brazil. Giant (Priodontes maximus), six-banded (Euphractus sexcinctus), southern naked-tailed (Cabassous unicinctus), and nine-banded (Dasypus novemcinctus) armadillos were divided into two groups. In group 1, 30 armadillos were anesthetized for collection of biological samples, body measurements, and placement of a microchip tag. Anesthetic combination BDM was applied: butorphanol tartrate, detomidine hydrochloride, and midazolam hydrochloride, each at 0.1 mg/kg. In group 2, 43 armadillos received ketamine hydrochloride at 10 mg/kg 20 min after BDM injection, and intra-abdominal radio transmitters were surgically implanted. The transmitter was inserted freely into the abdominal cavity. Vital signs were monitored during anesthesia every 10 min and varied within species and between groups. Rectal temperature varied from 33.1 ± 1.36 to 35.34 ± 1.21°C, heart rate (beats/min) from 19 ± 2.14 to 84.71 ± 9.25, respiratory rate (breaths/min) from 11 ± 4.16 to 31 ± 2.82, and oxygen saturation values (SPO2%) from 84.17% ± 2.39 to 98% ± 1.20. Both groups received the antagonist combination NYF: naloxone hydrochloride (0.02 mg/kg), yohimbine hydrochloride (0.125 mg/kg), and flumazenil (0.01 mg/kg). Recovery varied according to intravenous or intramuscular injection from 2 ± 4 to 8.08 ± 2.93 min respectively. BDM protocol was considered satisfactory and provided enough time to complete the procedures (60 ± 85 to 133.20 ± 9.12 min) according to the species and group. Ketamine added to the BDM provided enough time and a surgical plane of anesthesia (97 ± 22 to 137 ± 39.5 min). The surgical procedure technique chosen did not appear to have a negative impact on armadillos studied. Implantable transmitters provide a cost-effective method for long-term monitoring of wild individuals.

Moving in the Anthropocene: Global reductions in terrestrial mammalian movements.

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission.

Chemical Immobilization of Free-ranging Yellow Armadillos (Euphractus sexcinctus) for Implantation of Intra-abdominal Transmitters.

Free-ranging yellow armadillos (Euphractus sexcinctus) from the Brazilian Pantanal were captured for surgical implantation of intra-abdominal radio transmitters. During March 2014, 11 animals were chemically immobilized with a combination of intramuscular ketamine hydrochloride, xylazine hydrochloride, midazolam maleate, and atropine sulfate. The mean induction time and initial heart rate in males was higher than in females. The sex of armadillos did not affect the respiratory rate, which increased from the beginning to end of the monitoring time. The rectal temperatures of females remained almost unchanged, whereas the rectal temperatures of males suffered a slight drop through the procedure. The mean (±SD) total surgical time was 50±6.5 min, and the total anesthetic recovery time was 2.5±0.5 h. The armadillos were monitored for 24 h after surgery and released at their capture site. The anesthetic association was effective, promoting muscle relaxation and analgesia that allowed the performance of surgical procedures.

Mamíferos do Parque Estadual do Desengano, Rio de Janeiro, Brasil / Mammals of Desengano State Park, Rio de Janeiro, Brazil

Mesmo após a devastação de grande parte das suas matas, o Estado do Rio de Janeiro ainda abriga uma grande riqueza de mamíferos terrestres. Contudo, informações sobre abundância e distribuição da mastofauna do Estado ainda são escassas. Este estudo tem como objetivo inventariar os mamíferos do Parque Estadual do Desengano, localizado na região norte do Estado do Rio de Janeiro, uma das regiões menos amostradas e mais alteradas do Estado. O inventário foi realizado utilizando armadilhas do tipo Sherman e Tomahawk, e armadilhas de queda para os pequenos mamíferos não-voadores, redes de neblina para os morcegos, armadilhas fotográficas, avistamentos e relatos de moradores e funcionários do Parque para os mamíferos de maior porte. Nós registramos 56 espécies de mamíferos, das quais 15 constam na lista da fauna ameaçada de extinção do Estado do Rio de Janeiro e 10 na lista da fauna brasileira ameaçada de extinção, incluindo Brachyteles arachnoides. As espécies mais abundantes foram o morcego Sturnira lilium e o roedor Akodon serrensis. Além disso, Thaptomys nigrita, roedor normalmente pouco capturado no Estado, foi relativamente abundante no Parque Estadual do Desengano. Nossos resultados indicam que o Parque abriga 33,7 por cento das espécies de mamíferos com ocorrência conhecida para o Estado, destacando a importância da área para a conservação da fauna do Rio de Janeiro.

Even after the devastation of large part of its forests, Rio de Janeiro State still harbors a rich terrestrial mammal fauna. However, information on the distribution and abundance of mammal species in the State is still scarce. This study aims to survey the mammals of the Desengano State Park, located on the northern part of Rio de Janeiro, one of the least studied and most altered regions of the State. The inventory was conducted using Sherman, Tomahawk, and pitfall traps for non-volant small mammals, mist nets for bats, camera traps, sightings and reports of residents and officials of the Park for large mammals. We recorded 56 species of mammals, of which 15 are on the list of threatened fauna of Rio de Janeiro State and 10 on the list of Brazilian threatened fauna, including Brachyteles arachnoides. The most abundant species were the bat Sturnira lilium and the rodent Akodon serrensis. Moreover, Thaptomys nigrita, a rodent not commonly captured in the State, was relatively abundant in the Desengano State Park. Results indicate that the park harbors 33.7 percent of the species of mammals known to occur in the State, highlighting the importance of the area for the conservation of Rio de Janeiro fauna.