Jaguar Research Articles

Habitat destruction threatens jaguars in a mixed land-use region of eastern Bolivia

AbstractLarge carnivores such as the jaguar Panthera onca are particularly susceptible to population decline and local extinction as a result of habitat loss. Here we report on the long-term monitoring of a local jaguar population in a mixed land-use area in the eastern lowlands of Bolivia from March 2017 to December 2019. We recorded 15 jaguar individuals and four reproduction events (five offspring from three females), suggesting that our study area harbours a resident breeding population. Seven iterations of spatially explicit capture–recapture models provided density estimates of 1.32–3.57 jaguars per 100 km2. Jaguar capture rates were highest in forested areas, with few to no jaguar captures in pastures used for livestock. Massive deforestation after the survey period reduced the proportion of dense forest cover by 33%, shrinking the availability of suitable jaguar habitat and placing the resident jaguar population at risk. We use the jaguar as an indicator species to highlight the threat of habitat destruction in the Chiquitano region and we emphasize the importance of intact forest patches for jaguar conservation.

IDENTIFICATION OF ANTHROPOGENIC IMPACT ON NATURAL HABITATS BY ANTIMICROBIAL RESISTANCE QUANTIFICATION IN TWO NEOTROPICAL WILD CATS AND THEIR GEOSPATIAL ANALYSIS.

Human activities such as habitat degradation and fragmentation threaten biodiversity in Neotropical areas. This work proposes an analytical methodology to identify natural areas in Central America with anthropogenic impact, analyzing the presence of antimicrobial resistance genes (ARGs) in accordance with their theoretical relationship with human-related activities. Sixteen ARGs were quantified in feces of different individuals of 13 jaguars (Panthera onca) and 13 pumas (Puma concolor) in three conservation areas in Costa Rica by real-time PCR. At least one ARG was detected in all samples. Of the ARGs encoding tetracycline resistance, the most frequent were tetQ and tetY (85% and 69%, respectively). The sulfonamides (sulI and sulII; 69% each), phenicols (catI and catII; 19% and 54%, respectively), and quinolones (qnrS; 12%) were also detected. The presence of human settlements, livestock farms (pigs, cattle, and poultry), roads, human health centers, flood zones, and rivers were identified within each area to generate an index of human activity. We found no difference between the presence of ARG by roads, agricultural activities, and human settlements (P>0.05). However, tetW showed higher percentages with porcine and bovine farms; both tetY and tetW were more frequent in jaguars than in pumas. Of concern is that many of the most contaminated samples were taken from national parks, such as Braulio Carrillo and Tortuguero, where animals should not have direct contact with humans.

A multispecies corridor in a fragmented landscape: Evaluating effectiveness and identifying high-priority target areas.

While Misiones, Argentina contains one of the largest remnants of Upper Paraná Atlantic Forest ecoregion, one of the world's biodiversity hotspots, only ~50% of this native forest is protected. Each protected area is at risk of becoming an island of native forest surrounded by a matrix of altered habitats due to ongoing land conversion. In an effort to maximize long-term connectivity between existing protected areas, DeMatteo [1] used a multifaceted cost analysis to determine the optimal location for the region's first multispecies corridor using noninvasive data on jaguars (Panthera onca), pumas (Puma concolor), ocelots (Leopardus pardalis), southern tiger cats (Leopardus guttulus), and bush dogs (Speothos venaticus). This work builds on this framework by integrating new field data that broadens the scope of species-specific data across the region's heterogeneous landscape, which varies in vegetation, disturbance, human proximity, and protective status. In addition, two different land use layers are compared across the distributions of the five carnivores, the overlap in their independent distributions, and their relationship to the multispecies corridor. Interpretation of these land use data to species-specific habitat suitability goes beyond DeMatteo [1], with a subdivision of suitability into marginal and optimal areas. This refined scale allows a reanalysis of key areas in the multispecies corridor, where connectivity was previously defined as at highly-at-risk, allowing for a more directed development of management strategies. These analyses and their interpretation extend beyond northern-central Misiones, as the threats are not unique to this region. The need to develop management strategies that balance human-wildlife needs will continue to grow as humans expand their footprint. The techniques applied in this analysis provide a way to identify key areas that require specific management strategies, either through restoration, protection, or a combination of both.

Threatened Habitats of Carnivores: Identifying Conservation Areas in Michoacán, México

The present study contributes to bridging the gap in research related to the presence and distribution patterns of carnivore mammals in western México and identifies priority areas for biodiversity conservation in western Michoacán, México. The distribution of 11 carnivore species (Canis latrans; Urocyon cinereoargenteus; Herpailurus yagouaroundi; Leopardus pardalis; Leopardus wiedii; Puma concolor; Panthera onca; Conepatus leuconotus; Bassariscus astutus; Nasua narica; Procyon lotor) in western México was modeled through the application of a two-scale approach, including a large modeled region that corresponded to the western part of the country, for which consensus models were obtained that represent the species’ bioclimatic envelopes (historic occurrence records); and the second modeled study area that includes only the western portion of the state of Michoacán in which compounded models of the species’ habitat suitability (field occurrence records) for this region were proposed. Using species’ habitat suitability models as biodiversity units, prioritization exercises were carried out on important areas for the conservation of these species, as well as the comparison and analysis of the existing natural protected areas (NPA) and existing proposed conservation areas in the study area. The different exercises for prioritizing areas for conservation yielded similar results and show the potential percentages of the landscape that can be subjected to conservation programs. The highest conservation priority values were mainly located in the Costas del Sur and Cordillera del Sur provinces. This study signifies a flexible basis from which future studies on planning and designing a network of natural protected areas can be carried out in this region.

Big cats like water: occupancy patterns of jaguar in a unique and insular Brazilian Amazon ecosystem

Patterns of detectability and occupation of the jaguar (Panthera onca) vary throughout its distribution, being determined primarily by vegetation cover, prey availability, and anthropogenic factors. However, there is still a large knowledge gap regarding what determines jaguar occupancy, especially in the Amazon. This knowledge gap is even more pronounced for oceanic islands, which represent unique and very sensitive ecosystems, such as the Maracá-Jipioca Islands of the Northeastern Brazilian Amazon. Our study aimed to establish the spatial ecology of jaguars in this insular ecosystem and to provide information to facilitate sustainable management of the population there. We assessed how different factors (vegetation type and prey availability) potentially influence detectability and occupancy patterns of the jaguars of the Maracá-Jipioca Islands. We found that greater wetland (beach-sea and lagoon-channel) cover was the main driver of jaguar detectability and occupancy. We revealed factors driving the population and spatial ecology of jaguars in an insular system. Despite imminent threats to the region, the knowledge we present can inform the sustainable management of jaguars to ensure that the fundamental and unique ecosystem services provided by this top predator are maintained.

Effects of camera trap placement on photo rates of jaguars, their prey, and competitors in northwestern Costa Rica

AbstractThe use of camera‐trap methods, though important, has pitfalls and inconsistencies such as factors and interactions that may influence species photo rates. Jaguar (Panthera onca) camera‐trap studies place cameras at sites where jaguar detections are likely greater (e.g., sites with previous evidence such tracks, feces, and other presence signs), but do not account for potential placement biases. Our study evaluated methodological implications of a paired camera‐trap design at trail and off‐trail locations, and climate seasonality, on photo rates of jaguars, their prey, and competitors. From June 2016 to June 2017, camera traps were deployed at 58 different sites in a hexagon grid array of 3 km2 each in the Sector Santa Rosa (SSR) of Área de Conservación Guancaste in Costa Rica. Half of the cameras were located at a trail location that jaguars were likely to use, and the other half at an off‐trail location closest to each hexagon centroid. We estimated a photographic relative abundance index (RAI; no. of independent photos/100 trap nights) and used Generalized Linear Models (GLMs) to assess statistical evidence of trail location and climate seasonality. With a total effort of 19,408 trap nights, we recorded 12,678 independent photo events of 64 species of amphibians, reptiles, birds, and mammals. For the 3 bird species and 14 mammal species we focused on, most species‐specific RAIs were greater at trail locations (n = 8 species, including jaguars and pumas [Puma concolor]), some were greater at off‐trail locations (n = 2 species, including collared peccaries [Pecari tajacu]), and there was no discernable location effect for others (n = 7 species, including tapirs [Tapirus bairdii] and white‐tailed deer [Odocoileus virginianus]). For jaguars, sex ratio data showed a smaller proportion of males at off‐trail locations. Analysis of competitor and prey interaction data indicated temporal segregation at trail locations. Further jaguar camera‐trap studies should highlight camera placement as important source of bias that might influence results; hence, conservationists must be aware of this to avoid erroneous decisions.

Impending anthropogenic threats and protected area prioritization for jaguars in the Brazilian Amazon

Jaguars (Panthera onca) exert critical top-down control over large vertebrates across the Neotropics. Yet, this iconic species have been declining due to multiple threats, such as habitat loss and hunting, which are rapidly increasing across the New World tropics. Based on geospatial layers, we extracted socio-environmental variables for 447 protected areas across the Brazilian Amazon to identify those that merit short-term high-priority efforts to maximize jaguar persistence. Data were analyzed using descriptive statistics and comparisons of measures of central tendency. Our results reveal that areas containing the largest jaguar densities and the largest estimated population sizes are precisely among those confronting most anthropogenic threats. Jaguars are threatened in the world’s largest tropical forest biome by deforestation associated with anthropogenic fires, and the subsequent establishment of pastures. By contrasting the highest threats with the highest jaguar population sizes in a bivariate plot, we provide a shortlist of the top-10 protected areas that should be prioritized for immediate jaguar conservation efforts and 74 for short-term action. Many of these are located at the deforestation frontier or in important boundaries with neighboring countries (e.g., Peruvian, Colombian and Venezuelan Amazon). The predicament of a safe future for jaguars can only be ensured if protected areas persist and resist downgrading and downsizing due to both external anthropogenic threats and geopolitical pressures (e.g., infrastructure development and frail law enforcement).

Jaguar (Panthera onca) food resource use and its interaction with humans: scoping review

A scoping review was conducted to map, summarize, and understand the extent of evidence on jaguar (Panthera onca) food resource use and its interaction with humans. A total of 105 studies were identified in electronic databases: 57 analyzed jaguar food resource use, and 48 described jaguar interactions with humans. The studies were published in 51 journals, mainly in English, from 1993–2021 in 14/19 countries where jaguars live. The evidence demonstrated the variability and prey consumption of the jaguar; there are contrasting results regarding the diversity of prey consumed. The frequency of livestock predation was found to be related to the abundance and availability of natural prey and the distance of ranches from the jaguar habitat. The interaction of the jaguar with humans presented an interesting and contrasting picture, since the perception of the jaguar varied depending on the sociocultural context and the study site. The results showed a negative effect of human activities on the distribution and density of the feline, and there was evidence of poaching and attacks on humans, although the information was insufficient to determine the causes in depth. More research and dissemination are needed on these issues at the local level in the regions where the jaguar lives, as these are the areas where decisions and actions are needed to conserve the jaguar in coexistence with communities and livestock.

A review of environmental and anthropogenic variables used to model jaguar occurrence

Jaguars (Panthera onca) are a landscape species of conservation importance and our understanding of environmental and anthropogenic drivers of jaguar occurrence is necessary to improve conservation strategies. We reviewed available literature to simply describe environmental and anthropogenic variables used and found to be significant in occurrence modeling. We reviewed 95 documents published from 1980 to 2021 that focused on jaguar occurrence and that used 39 variable types (21 anthropogenic, 18 environmental) among different techniques, scales, and approaches. In general, these variables included both anthropogenic (roads, land use, human activities, and population) and environmental (climate, vegetation, ecological interactions, topographic, water, and others) factors. Twelve variables were identified as affecting jaguar occurrence overall, eleven at local scale and seven at broad scales (regional and continental). Focusing more specifically on the variables that correlate with occurrence should help researchers to make better predictions in areas without quantitative jaguar data.

Unveiling demographic and mating strategies of Panthera onca in the Pantanal, Brazil.

We conducted the first long-term and large-scale study of demographic characteristics and reproductive behavior in a wild jaguar (Panthera onca) population. Data were collected through a combination of direct observations and camera trapping on a study area that operates both as a cattle ranch and ecotourism destination. Jaguars exhibited two birth peaks: April/May and October/November, that are the end and the beginning of the wet season in the Pantanal, respectively. The average litter size was 1.43 ± 0.65. Single cubs made up a total of 65.7% of the births, and we found a slight predominance of females (1.15:1 ratio) in litters. The mean age at independence was 17.6 ± 0.98 months, with sex-biased dispersal, with all males (n = 27) leaving the natal home range and 63.6% of females exhibiting philopatry. The interbirth intervals were 21.8 ± 3.2 months and the mean age at first parturition was 31.8 ± 4.2 months. Our results estimated a lifetime reproductive success for female jaguars of 8.13 cubs. Our observations also indicate that female jaguars can display mating behavior during cub rearing or pregnancy, representing 41.4% of the consorts and copulations recorded. We speculate that this behavior has evolved as a defense against infanticide and physical harm to the female. To our knowledge, this is the first time that such behavior is described for this species. All aggressive interactions between females involved the presence of cubs, following the offspring-defense hypothesis, that lead to territoriality among females in mammals, regardless of food availability. In the face of growing threats to this apex predator, this work unveils several aspects of its natural history, representing a baseline for comparison with future research and providing critical information for population viability analysis and conservation planning in the long term.

Ecuadorian Children’s Willingness to Protect Endangered Species – Identifying Behavioral Predictors in a Biodiversity Hotspot

Abstract The loss of biodiversity is a problem that particularly affects biodiversity hotspots. Children play a crucial role in the conservation of endangered species. One important prerequisite for conservation behavior is the willingness to protect endangered animal species. The present study investigated the influence of several variables on Ecuadorian children’s willingness to protect domestic endangered animal species (N = 154; MAge = 8.57; SD = 0.55; 48.1% female). Gender, caring beliefs (a subdimension of the wildlife value orientation), dispositional empathy with endangered animals, and threat perception of the Andean condor (Vultur gryphus) toward humans were strong predictors of the willingness to protect them. Conversely, psychological distance, and threat perception of both the Andean bear (Tremarctos ornatus) and the jaguar (Panthera onca) had no significant influence on willingness to protect. However, their effects may be indirect. The results of this study are relevant for biodiversity conservation and educators in schools.

Detecção do vírus da panleucopenia felina (Carnivore protoparvovirus 1) em Panthera onca de vida livre no Brasil

ABSTRACT: The decline in the jaguar population confirms how much the species is vulnerable to extinction in Brazil. It also indicates the degradation of its natural habitat’s environmental integrity and quality. Studies claim that large felids are susceptible to feline panleukopenia virus (FPV) and are presumptively diagnosed clinically in Brazil. A free-living jaguar (Panthera onca) cub was found unconscious and rescued due to a possible hit-and-run in the savannah of Mato Grosso. During recovery, it exhibited clinical and hematological signs consistent with FPV infection. The PCR was positive for FPV, with 99.61% identity between the FPV sequences available in the GenBank database through the BLAST tool. Due to habitat restrictions, certain diseases threaten wild cats and habitat encroachment by domestic animals can alter the pattern of spread of pathogens. We highlight the importance of the molecular diagnosis and phylogenetic analysis of FPV to elucidate how it has reached wild felids.

Anaplastic mammary carcinoma in a captive jaguar (Panthera onca)

ABSTRACT A captive female of jaguar (Panthera onca), 12 years old, presenting an 8-day history of anorexia and lethargy, died in its enclosure in Manaus (Amazonas, Brazil). The necropsy revealed an abdominal multinodular ventral plaque into the mammary region. Additionally, in several locations of the subcutaneous tissue, there were multiple metastatic nodules of firm texture. Metastases were also observed in the lymph nodes, heart and lungs. The immunohistochemistry results revealed the cytoplasmic staining for cytokeratin on neoplastic cells, mild staining (+) for E-cadherin, moderate staining (++) for estrogen and progesterone receptors, and about 50 to 70% of staining for Ki-67. The neoplastic cells were negative for vimentin staining. Therefore, the histopathological examination led to a diagnosis of metastatic anaplastic mammary carcinoma, which is an unusual condition in wild felines.

Drivers of large carnivore density in non‐hunted, multi‐use landscapes

AbstractProtected areas serve as population strongholds for many large carnivores, with multi‐use landscapes along their borders forming the front‐lines of wildlife conservation. Understanding large carnivore population dynamics within working landscapes is difficult where anthropogenic mortality is high and unregulated. This study focused on working ranches, where killing jaguars (Panthera onca) and their prey was prohibited, to gain insight into jaguar population potential across multi‐use landscapes. Faced with forest fragmentation, presence of domestic livestock, and dynamic land‐use practices, we expected jaguar populations in working landscapes to be predominantly male and transient, with low cub production, and inflated population densities in remnant forest patches, versus protected areas where we expected native forest habitat and stable jaguar territories. Using camera traps and spatial‐capture recapture analyses, we observed that male jaguars demonstrated larger‐scale movements and were more detectable than females (0.07 ± 0.01 SE vs. 0.02 ± 0.01 SE) in both working and protected landscapes. Female jaguars in ranches traveled farther than females in parks. Carnivore density increased with forest cover and wild prey activity, decreased with domestic prey activity, and was marginally higher in ranches (4.08 individuals/100 km2 ± 0.73 SE) than in parks (3.59 individuals/100 km2 ± 0.64 SE). Females outnumbered males in both landscapes (2.20–2.60 females/100 km2vs. ~1.60 males/100 km2), although local male density reached up to 11.00 males/100 km2in ranches (vs. 3.50 males/100 km2in parks). While overall jaguar density was patchier in protected areas ( = 0.69 parks, 0.54 ranches), inter‐annual patchiness was higher within ranches (Moran'sI = 0.49–0.60 ranches, 0.69–0.70 parks), reflecting changes in cattle management. Despite major habitat alterations, working landscapes can support carnivore densities equivalent to (or exceeding that of) unmodified forest habitat, provided that wildlife‐tolerant ranching practices are maintained.

Efficient recovery of in vivo mature and immature oocytes from jaguars (Panthera onca) and pumas (Puma concolor) by Laparoscopic Ovum Pick-Up (LOPU)

The jaguar and the puma play an essential role in balancing the ecosystem. Developing reproductive biotechnologies for such species could permit the exchange of genetic material across in situ and ex situ populations, encompassing the concept of One Conservation. Therefore, the purpose of the current study was to compare different gonadotropin stimulation strategies in conjunction with laparoscopic ovum pick-up (LOPU) technology for the recovery of immature and mature oocytes from these species. Adult female pumas (n = 10) and jaguars (n = 11) underwent 12 and 16 LOPU treatments, respectively. Hormonal stimulation was performed with eCG (750 IU for pumas; 800 IU for jaguars; im), and they were randomized into two treatment groups: Group 1 (G1) received hCG (500 IU, im) approximately 85 h after eCG administration to promote oocyte maturation and recover in vivo matured oocytes; and Group 2 (G2) did not receive hCG to recover immature oocytes. Between the two treatments, a total of 416 and 160 viable oocytes were recovered, 205 and 80 from G1 and 211 and 80 from G2-treated females from pumas and jaguars, respectively. We found that LOPU is a safe technique for getting high-quality oocytes from wild cats, such as pumas and jaguars, and that it may be included in conservation efforts. In pumas and jaguars, protocols employing eCG for ovarian stimulation are effective, as is the use of hCG to stimulate in vivo oocyte maturation prior to collection.

Medium and large-sized mammals of a private protected wetland in the Cerrado-Amazon biological corridor, Brazil.

Brazil is the world's richest country in biodiversity, including mammal species. In the Brazilian Cerrado biome, mammalian diversity is vast, with about 251 species, 32 of them are endemic and 22 listed as threatened species. In this work, we investigated species diversity of medium- and large-sized mammals in the private protected area RPPN Pontal do Jaburu (RPPN-PJ) and its surroundings, which is a flooded area located in an important biological corridor in the Cerrado-Amazon ecotone zone, a priority area for biodiversity conservation in Brazil. We used camera-trapping, active search (night and day), and track survey during dry season (Apr - Aug 2016). We recorded 29 mammal species, being the Carnivora order the most representative with 11 species. Regarding threat status, 35.7% of the recorded species were listed as threatened in Brazil and 32.1% worldwide. We highlight the high relative frequency of threatened species records such as Tapirus terrestris, Panthera onca, Blastocerus dichotomus, Pteronura brasiliensis, Priodontes maximus, and other, as well as the presence of the newly described aquatic mammal species Inia araguaiaensis. We stress the importance of RPPN-PJ and its surroundings for mammal conservation, which include complex habitats (wetlands) located in an important ecotone zone.

Monitoring of Panthera onca should include long-term studies and genetic approaches

Onças-pintadas (Panthera onca Linnaeus, 1758) são predadores do topo de cadeias alimentares, e sua ocorrência é um bom indicador de qualidade ambiental. O estudo das onças-pintadas, incluindo seu comportamento, ecologia e conservação, tem atraído atenção significativa da comunidade científica. Realizamos uma análise cienciométrica sobre estudos da biologia da onça-pintada através do uso de armadilhas fotográficas nos biomas Cerrado e Pantanal. Nossa análise revelou que conflitos entre agricultores e pecuaristas são um impacto importante na ecologia do movimento dos felinos. Encontramos duas lacunas importantes: uma escassez de estudos com uma abordagem genética sobre a biologia das onças, e que incluem a ecologia do movimento da espécie nos biomas Cerrado e Pantanal. Ademais, na maioria dos estudos as onças foram monitoradas por períodos curtos, o que limita a qualidade dos dados obtidos.

Maternal behavioural analysis during a successful captive breeding of jaguars Panthera onca

The breeding of captive wild species can play a crucial role in their preservation. Jaguars, the largest felines in the Americas, face challenges in captivity that hinder their breeding success in zoos and conservation institutions. One major barrier to successful captive breeding is the difficulty of replicating the natural environment, including nutritional, social, and habitat needs. Hence, any insights into the breeding behaviour and parental care of jaguars can inform and enhance breeding strategies for these captive felines. In this study, we aimed to examine maternal behaviours during the pre- and post-partum periods in the first successful case of natural reproduction of captive jaguars in Brazil. The study took place at the Mata Ciliar Association in Jundiaí , Brazil, where infrared cameras were used to observe maternal behaviours without human interference. The behaviours were analysed using an ethogram of the mother and Generalized Linear Models (GLM) to examine her behavioural repertoire during three different phases of parental care. Our findings suggest that there were differences in the behaviours exhibited by the female jaguar in the pre- and post-partum periods. Furthermore, the mother's interactions with her jaguar cubs reduced her stereotyped behaviours, implying an improvement in her welfare. Finally, we offer suggestions and recommendations to facilitate future breeding efforts for captive jaguars.

Jaguar’s Predation and Human Shield, a Tapir Story

Despite the risks associated, some species choose to shield behind a predator to decrease predation risk by another predator. In this study, we demonstrate how Baird’s tapirs (Tapirus bairdii) use humans as a “shield” to reduce the risk of being preyed upon by jaguars (Panthera onca). We collected georeferenced photographic records of 23 tapirs (seven of them injured) sighted near human settlements (0 to 5 km) in the Calakmul region of Mexico from 2008 to 2019. Using multidimensional scale analysis, we determined which possible factors (tapir health status, injuries, distance to the settlement, as well as seasonality) are related to the decision of tapirs to approach human settlements. To support our claims of jaguars’ attacks, we described the pattern of injuries believed to have been inflicted by jaguars on tapirs, and we analysed photographs and videos of species of the genus Panthera attacking larger prey than themselves to establish a pattern of injuries and compare it to the injuries observed on tapirs. Our study shows that tapir sightings near human settlements are related to health deterioration, injuries by jaguars and seasonality. The injuries found on tapirs are similar to those caused by other big cats on large prey, providing strong support for jaguar-inflicted wounds. Further studies should investigate whether the increasing human presence in different habitats in the Neotropical region could be influencing the behaviour and distribution of prey and predators.

Does cattle ranching drive activity patterns of jaguar (Panthera onca) and puma (Puma concolor) in the Brazilian Pantanal?

We conducted a comparative study of jaguar and puma activity patterns within a mosaic of protected areas (AMR) and on a cattle ranch (CR) in Pantanal, Brazil, to better understand their activity patterns in these landscapes. We hypothesized that the activity patterns of the jaguar and puma would be biased to the nocturnal period within the cattle ranch but not in the protected areas. We used data from camera traps analyzed through a non-parametric kernel density approach to explore interspecific and intraspecific temporal relationships between these species at both sites. We obtained 71 jaguar and 29 puma independent records at AMR, and 85 jaguar and 26 puma independent records at CR. Activity patterns of jaguars and pumas differed between sites, both being cathemeral in AMR, but nocturnal at CR with moderate to high overlaps, concordant with our hypothesis. Overall, our data suggest that the cattle ranching is not incompatible with the existence of jaguars and pumas but does shape their pattern of activities.