Belize

Tropical forests are important reservoirs of biodiversity 1 , but the processes that maintain this diversity remain poorly understood 2 . The Janzen-Connell hypothesis 3,4 suggests that specialized natural enemies such as insect herbivores and fungal pathogens maintain high diversity by elevating mortality when plant species occur at high density (negative density dependence; NDD). NDD has been detected widely in tropical forests , but the prediction that NDD caused by insects and pathogens has a community-wide role in maintaining tropical plant diversity remains untested. We show experimentally that changes in plant diversity and species composition are caused by fungal pathogens and insect herbivores. Effective plant species richness increased across the seed-to-seedling transition, corresponding to large changes in species composition 5 . Treating seeds and young seedlings with fungicides significantly reduced the diversity of the seedling assemblage, consistent with the Janzen-Connell hypothesis. Although suppressing insect herbivores using insecticides did not alter species diversity, it greatly increased seedling recruitment and caused a marked shift in seedling species composition. Overall, seedling recruitment was significantly reduced at high conspecific seed densities and this NDD was greatest for the species that were most abundant as seeds. Suppressing fungi reduced the negative effects of density on recruitment, confirming that the diversityenhancing effect of fungi is mediated by NDD. Our study provides an overall test of the Janzen-Connell hypothesis and demonstrates the crucial role that insects and pathogens have both in structuring tropical plant communities and in maintaining their remarkable diversity.

Carcharhinid sharks can make up a large fraction of the top predators inhabiting tropical marine ecosystems and have declined in many regions due to intense fishing pressure. There is some support for the hypothesis that carcharhinid species that complete their life-cycle within coral reef ecosystems, hereafter referred to as ''reef sharks'', are more abundant inside no-take marine reserves due to a reduction in fishing pressure (i.e., they benefit from marine reserves). Key predictions of this hypothesis are that (a) individual reef sharks exhibit high site-fidelity to these protected areas and (b) their relative abundance will generally be higher in these areas compared to fished reefs. To test this hypothesis for the first time in Caribbean coral reef ecosystems we combined acoustic monitoring and baited remote underwater video (BRUV) surveys to measure reef shark site-fidelity and relative abundance, respectively. We focused on the Caribbean reef shark (Carcharhinus perezi), the most common reef shark in the Western Atlantic, at Glover's Reef Marine Reserve (GRMR), Belize. Acoustically tagged sharks (N = 34) were detected throughout the year at this location and exhibited strong site-fidelity. Shark presence or absence on 200 BRUVs deployed at GRMR and three other sites (another reserve site and two fished reefs) showed that the factor ''marine reserve'' had a significant positive effect on reef shark presence. We rejected environmental factors or site-environment interactions as predominant drivers of this pattern. These results are consistent with the hypothesis that marine reserves can benefit reef shark populations and we suggest new hypotheses to determine the underlying mechanism(s) involved: reduced fishing mortality or enhanced prey availability.

The Maya archaeological site of Ucanal is located in Peten, Guatemala, close to the contemporary border with Belize. In pre-Columbian times, the site also sat at the frontiers of some of the largest political centers, Naranjo in Peten, Guatemala, and Caracol, in Belize. Entangled between these dominant centers and with ties to peoples in the Upper Belize Valley, the Petexbatun region in Guatemala, northern Yucatan, and elsewhere, Ucanal was a critical convergence zone of political and cultural interaction. This paper synthesizes archaeological research by the Proyecto Arqueológico Ucanal to underscore the ways in which this provincial polity, identified epigraphically as K'anwitznal, maneuvered within and between different cultural affiliations and political networks. We find that the site's role as a political frontier during the Late Classic period was more of a bridge than an edge. During the later Terminal Classic period, the K'anwitznal kingdom gained independence, but continued to serve as a critical convergence of influences and interaction spheres from throughout the Maya area and beyond.

Often understudied by archaeologists, ground stone tools (GST) were ubiquitous in the ancient Maya world. Their applications ranged from household tools to ceremonial equipment and beyond. Little attention has been focused on chemically sourcing the raw stone material used in GST production, largely because these tools were fashioned out of igneous or sedimentary rock, which can present characterization challenges. And, although portable X-ray fluorescence (pXRF) has been applied widely to source obsidian, the utility of pXRF for geochemically sourcing other kinds of stone remains underexplored. We present a small-scale application of pXRF for determining granite provenance within a section of the Middle Belize Valley in Belize, Central America. Belize is an ideal location to test chemical sourcing studies of granite because there are only three tightly restricted and chemically distinct sources of granite in the country, from which the overwhelming majority of granite for ancient tool production derived. The method described here demonstrates that successful and accurate geological characterizations can be made on granite GST. This cutting-edge sourcing technique has the potential to be more widely applied in other regions to reveal deeper connections between the sources of GST production and sites of consumption across space and through time.

Excavations at Crawford Bank in Crooked Tree, Belize, exposed a lithic deposit with no associated ceramics. The deposit primarily consists of chipped chert and chalcedony tools and debitage, as well as a small number of worked slate fragments. Most of the chert likely sources to the Northern Belize Chert-bearing Zone (NBCZ). The recovery of two Archaic period formal tools – a Lowe point and a constricted uniface/trimmed macroblade – suggests a pre-Maya occupation. Use-wear analysis of both tools and debitage demonstrates a wide range of uses with a focus on wood and hard contact materials. The use-wear patterns demonstrate a heavy reliance on ad hoc/expedient technology for the completion of different tasks involving wood by preceramic peoples. The Crawford Bank site likely represents one or more short-term, task-orientated
preceramic occupation(s) for the extraction and use of the available resources of the local wetland environment, most notably logwood.

There is a great need to develop efficient, noninvasive genetic sampling methods to study wild populations of multiple, co-occurring, threatened felids. This is especially important for molecular scatology studies occurring in challenging tropical environments where DNA degrades quickly and the quality of faecal samples varies greatly. We optimized 14 polymorphic microsatellite loci for jaguars (Panthera onca), pumas (Puma concolor) and ocelots (Leopardus pardalis) and assessed their utility for cross-species amplification. Additionally, we tested their reliability for species and individual identification using DNA from faeces of wild felids detected by a scat detector dog across Belize in Central America. All microsatellite loci were successfully amplified in the three target species, were polymorphic with average expected heterozygosities of H E = 0.60 AE 0.18 (SD) for jaguars, H E = 0.65 AE 0.21 (SD) for pumas and H E = 0.70 AE 0.13 (SD) for ocelots and had an overall PCR amplification success of 61%. We used this nuclear DNA primer set to successfully identify species and individuals from 49% of 1053 field-collected scat samples. This set of optimized microsatellite multiplexes represents a powerful tool for future efforts to conduct noninvasive studies on multiple, wild Neotropical felids.

Unmanned aerial systems (UASs) are powerful tools for research and monitoring of wildlife. However, the effects of these systems on most marine mammals are largely unknown, preventing the establishment of guidelines that will minimize animal disturbance. In this study, we evaluated the behavioral responses of coastal bottlenose dolphins (Tursiops truncatus) and Antillean manatees (Trichechus manatus manatus) to small multi-rotor UAS flight. From 2015 to 2017, we piloted 211 flights using DJI quadcopters (Phantom II Vision +, 3 Professional and 4) to approach and follow animals over shallow-water habitats in Belize. The quadcopters were equipped with high-resolution cameras to observe dolphins during 138 of these flights, and manatees during 73 flights. Aerial video observations of animal behavior were coded and paired with flight data to determine whether animal activity and/or the UAS's flight patterns caused behavioral changes in exposed animals. Dolphins responded to UAS flight at altitudes of 11–30 m and responded primarily when they were alone or in small groups. Single dolphins and one pair responded to the UAS by orienting upward and turning toward the aircraft to observe it, before quickly returning to their pre-response activity. A higher number of manatees responded to the UAS, exhibiting strong disturbance in response to the aircraft from 6 to 104 m. Manatees changed their behavior by fleeing the area and sometimes this elicited the same response in nearby animals. If pursued post-response, manatees repeatedly responded to overhead flight by evading the aircraft's path. These findings suggest that the invasiveness of UAS varies across individuals, species, and taxa. We conclude that careful exploratory research is needed to determine the impact of multi-rotor UAS flight on diverse species, and to develop best practices aimed at reducing the disturbance to wildlife that may result from their use.

Located on the Caribbean coast of Central America and flanked by the second longest barrier reef in the world, Belize is a nation reliant upon marine resources. Each year, the country's predominantly small-scale fisheries generate an estimated US$22 million in revenue – 1.8%of GDP – and employ 3000 people. However, the nation's fishing communities are facing an unprecedented challenge. Existing threats posed by declining fish stocks have been exacerbated by the introduction of the invasive alien red lionfish Pterois volitans in 2008. This Indo-Pacific predator has the potential to cause significant losses to the recruitment of native fish, in turn disrupting coral reef community dynamics in the Belize Barrier Reef Reserve System, a UNESCO World Heritage Site. Howefver, targeted lionfish fishing may offer a cost effective means to control the invasion, while also creating an alternative livelihood solution and improving food security among Belize's coastal fishing communities. This study summarises the recent history of the lionfish invasion in Belize, describes the multi-sector approach being used to address the invasion, and presents preliminary research summarising the characteristics of invasive alien lionfish in Belize. Data from Belize's nascent 'lionfishery' are also presented, demonstrating that demand for lionfish is outweighing supply – largely as a result of awareness-raising initiatives – and highlighting the strong potential for replication of this approach elsewhere in the Caribbean. The study concludes by discussing the barriers and potential solutions to this market-based approach to invasive species management.

Understanding factors that strengthen community support for ecotourism and conservation is key to a successful ecotourism project. This study of Five Blues Lake National Park in central Belize explored attitudes towards ecotourism and conservation of local people living adjacent to and co-managing a national park. The authors conducted attitudinal surveys of residents of Saint Margarets village in 1995 and in 2003. The data suggested high levels of support for ecotourism development and continued protection of the land and conservation efforts. Despite this support, by 2003, most ecotourism activities and park management had been curtailed. Although the inception of co-managed ecotourism was received well and some level of interest existed, over time, there was a decline in community participation and ecotourism activities. This is attributable to factors including resident apathy, low community
participation, diminished financial resources, insufficient local and national management capacity, and perceptions of few personal benefits by residents. Furthermore, the often claimed theoretical benefits of ecotourism in developing communities were not fully realised in this case. As demonstrated by this study, local
involvement of residents in development and implementation of ecotourism initiatives, including necessary capacity-building, may help ensure sustainable management and success of long-term ecotourism and conservation efforts.

Seed predation and seed dispersal can have strong eVects on early life history stages of plants. These processes have often been studied as individual eVects, but the degree to which their relative importance co-varies with seed predator abundance and how this inXuences seed germination rates is poorly understood. Therefore, we used a combination of observations and Weld experiments to determine the degree to which germination rates of the palm Astrocaryum mexicanum varied with abundance of a small mammal seed predator/disperser, Heteromys desmarestianus, in a lowland tropical forest. Patterns of abundance of the two species were strongly related; density of H. desmarestianus was low in sites with low density of A. mexicanum and vice versa. Rates of predation and dispersal of A. mexicanum seeds depended on abundance of H. desmarestianus; sites with high densities of H. desmarestianus had the highest rates of seed predation and lowest rates of seed germination, but a greater total number of seeds were dispersed and there was greater density of seedlings, saplings, and adults of A. mexicanum in these sites. When abundance of H. desmarestianus was experimentally reduced, rates of seed predation decreased, but so did dispersal of A. mexicanum seeds. Critically, rates of germination of dispersed seeds were 5 times greater than undispersed seeds. The results suggest that the relationship between A. mexicanum and H. desmarestianus is a conditional mutualism that results in a strong local eVect on the abundance of each species. However, the magnitude and direction of these eVects are determined by the relative strength of opposing, but related, mechanisms. A. mexicanum nuts provide H. desmarestianus with a critical food resource, and while seed predation on A. mexicanum nuts by H. desmarestianus is very intense, A. mexicanum ultimately beneWts because of the relatively high germination rates of its seeds that are dispersed by H. desmarestianus.

This study investigated the relationship of ethnicity, parental education, gender, and parental attachment to multiple dimensions of self-image among 285 (161 female and 124 male) late adolescent Belizean students. Student ratings of self-image were unrelated to paternal education and student ethnicity. For maternal education, ethnic identity was higher for students whose mothers were high school graduates than for those whose mothers had completed college or a graduate degree. Female students reported more positive self-images than males in vocational attitudes. Favourable self-images were associated with secure ratings of parental attachment. r