Robert N. Reed

Severe mammal declines coincide with proliferation of invasive Burmese pythons in Everglades National Park

Invasive species represent a significant threat to global biodiversity and a substantial economic burden. Burmese pythons, giant constricting snakes native to Asia, now are found throughout much of southern Florida, including all of Everglades National Park (ENP). Pythons have increased dramatically in both abundance and geographic range since 2000 and consume a wide variety of mammals and birds. Here we report severe apparent declines in mammal populations that coincide temporally and spatially with the proliferation of pythons in ENP. Before 2000, mammals were encountered frequently during nocturnal road surveys within ENP. In contrast, road surveys totaling 56,971 km from 2003–2011 documented a 99.3% decrease in the frequency of raccoon observations, decreases of 98.9% and 87.5% for opossum and bobcat observations, respectively, and failed to detect rabbits. Road surveys also revealed that these species are more common in areas where pythons have been discovered only recently and are most abundant outside the pythons current introduced range. These findings suggest that predation by pythons has resulted in dramatic declines in mammals within ENP and that introduced apex predators, such as giant constrictors, can exert significant top-down pressure on prey populations. Severe declines in easily observed and/or common mammals, such as raccoons and bobcats, bode poorly for species of conservation concern, which often are more difficult to sample and occur at lower densities.

What parts of the US mainland are climatically suitable for invasive alien pythons spreading from Everglades National Park

The Burmese Python (Python molurus bivittatus) is now well established in southern Florida and spreading northward. The factors likely to limit this spread are unknown, but presumably include climate or are correlated with climate. We compiled monthly rainfall and temperature statistics from 149 stations located near the edge of the python’s native range in Asia (Pakistan east to China and south to Indonesia). The southern and eastern native range limits extend to saltwater, leaving unresolved the species’ climatic tolerances in those areas. The northern and western limits are associated with cold and aridity respectively. We plotted mean monthly rainfall against mean monthly temperature for the 149 native range weather stations to identify the climate conditions inhabited by pythons in their native range, and mapped areas of the coterminous United States with the same climate today and projected for the year 2100. We accounted for both dry-season aestivation and winter hibernation (under two scenarios of hibernation duration). The potential distribution was relatively insensitive to choice of scenario for hibernation duration. US areas climatically matched at present ranged up the coasts and across the south from Delaware to Oregon, and included most of California, Texas, Oklahoma, Arkansas, Louisiana, Mississippi, Alabama, Florida, Georgia, and South and North Carolina. By the year 2100, projected areas of potential suitable climate extend northward beyond the current limit to include parts of the states of Washington, Colorado, Illinois, Indiana, Ohio, West Virginia, Pennsylvania, New Jersey, and New York. Thus a substantial portion of the mainland US is potentially vulnerable to this ostensibly tropical invader.

Environmental DNA (eDNA) Sampling Improves Occurrence and Detection Estimates of Invasive Burmese Pythons

Environmental DNA (eDNA) methods are used to detect DNA that is shed into the aquatic environment by cryptic or low density species. Applied in eDNA studies, occupancy models can be used to estimate occurrence and detection probabilities and thereby account for imperfect detection. However, occupancy terminology has been applied inconsistently in eDNA studies, and many have calculated occurrence probabilities while not considering the effects of imperfect detection. Low detection of invasive giant constrictors using visual surveys and traps has hampered the estimation of occupancy and detection estimates needed for population management in southern Florida, USA. Giant constrictor snakes pose a threat to native species and the ecological restoration of the Florida Everglades. To assist with detection, we developed species-specific eDNA assays using quantitative PCR (qPCR) for the Burmese python (Python molurus bivittatus), Northern African python (P. sebae), boa constrictor (Boa constrictor), and the green (Eunectes murinus) and yellow anaconda (E. notaeus). Burmese pythons, Northern African pythons, and boa constrictors are established and reproducing, while the green and yellow anaconda have the potential to become established. We validated the python and boa constrictor assays using laboratory trials and tested all species in 21 field locations distributed in eight southern Florida regions. Burmese python eDNA was detected in 37 of 63 field sampling events; however, the other species were not detected. Although eDNA was heterogeneously distributed in the environment, occupancy models were able to provide the first estimates of detection probabilities, which were greater than 91%. Burmese python eDNA was detected along the leading northern edge of the known population boundary. The development of informative detection tools and eDNA occupancy models can improve conservation efforts in southern Florida and support more extensive studies of invasive constrictors. Generic sampling design and terminology are proposed to standardize and clarify interpretations of eDNA-based occupancy models.

Marsh rabbit mortalities tie pythons to the precipitous decline of mammals in the Everglades

To address the ongoing debate over the impact of invasive species on native terrestrial wildlife, we conducted a large-scale experiment to test the hypothesis that invasive Burmese pythons (Python molurus bivittatus) were a cause of the precipitous decline of mammals in Everglades National Park (ENP). Evidence linking pythons to mammal declines has been indirect and there are reasons to question whether pythons, or any predator, could have caused the precipitous declines seen across a range of mammalian functional groups. Experimentally manipulating marsh rabbits, we found that pythons accounted for 77% of rabbit mortalities within 11 months of their translocation to ENP and that python predation appeared to preclude the persistence of rabbit populations in ENP. On control sites, outside of the park, no rabbits were killed by pythons and 71% of attributable marsh rabbit mortalities were classified as mammal predations. Burmese pythons pose a serious threat to the faunal communities and ecological functioning of the Greater Everglades Ecosystem, which will probably spread as python populations expand their range.

A field test of attractant traps for invasive Burmese pythons (Python molurus bivittatus) in southern Florida

Context. Invasive Burmese pythons (Python molurus bivittatus) are established over thousands of square kilometres of southern Florida, USA, and consume a wide range of native vertebrates. Few tools are available to control the python population, and none of the available tools have been validated in the field to assess capture success as a proportion of pythons available to be captured. Aims. Our primary aim was to conduct a trap trial for capturing invasive pythons in an area east of Everglades National Park,wheremanypythonshadbeencapturedinpreviousyears,toassesstheefficacyoftrapsforpopulationcontrol.Wealso aimed to compare results of visual surveys with trap capture rates, to determine capture rates of non-target species, and to assess capture rates as a proportion of resident pythons in the study area. Methods.Weconductedamedium-scale(6053trapnights)experimentusingtwotypesofattractanttrapsbaitedwithlive ratsintheFrogPondareaeastofEvergladesNationalPark.Wealsoconductedstandardisedandopportunisticvisualsurveys inthetrappingarea.Followingthetrap trial,theareawasdischarrowedtoexposepythonsandallowcalculationof anindex of the number of resident pythons. Key results. We captured three pythons and 69 individuals of various rodent, amphibian, and reptile species in traps. Eleven pythons were discovered during disc harrowing operations, as were large numbers of rodents. Conclusions. The trap trial captured a relatively small proportion of the pythons that appeared to be present in the study area, although previous research suggests that trap capture rates improve with additional testing of alternative trap designs. Potential negative impacts to non-target species were minimal. Low python capture rates may have been associated with extremely high local prey abundances during the trap experiment. Implications. Results of this trial illustrate many of the challenges in implementing and interpreting results from tests of control tools for large cryptic predators such as Burmese pythons.

Genetic analysis of a novel invasion of Puerto Rico by an exotic constricting snake

The tropical island Puerto Rico is potentially vulnerable to invasion by some species of exotic snakes; however, until now no established populations had been reported. Here we report and genetically characterize the nascent invasion of Puerto Rico by an exotic constricting snake of the family Boidae (Boa constrictor) using mtDNA and microsatellite data. Over 150 individual B. constrictor have been removed from Mayagüez municipality since May 2011, and our results from the genetic analysis of 32 individuals suggest that this population was recently founded by individuals of one subspecies from a genetic lineage common to zoo and breeding collections, but that the potential propagule pool consists of two subspecies. We also suggest that anthropogenic long-distance dispersal within the island of Puerto Rico may be occurring from the established population, with implications for further establishment across the island. This study represents the first report of the naturalization of an invasive species of boid snake in Puerto Rico and will be important in determining mitigation strategies for this invasion as well as providing a basis for comparison to other on-going studies of invasive snakes.

Ecological correlates of invasion impact for Burmese pythons in Florida

An invasive population of Burmese pythons (Python molurus bivittatus) is established across several thousand square kilometers of southern Florida and appears to have caused precipitous population declines among several species of native mammals. Why has this giant snake had such great success as an invasive species when many established reptiles have failed to spread? We scored the Burmese python for each of 15 literature-based attributes relative to predefined comparison groups from a diverse range of taxa and provide a review of the natural history and ecology of Burmese pythons relevant to each attribute. We focused on attributes linked to spread and magnitude of impacts rather than establishment success. Our results suggest that attributes related to body size and generalism appeared to be particularly applicable to the Burmese pythons success in Florida. The attributes with the highest scores were: high reproductive potential, low vulnerability to predation, large adult body size, large offspring size and high dietary breadth. However, attributes of ectotherms in general and pythons in particular (including predatory mode, energetic efficiency and social interactions) might have also contributed to invasion success. Although establishment risk assessments are an important initial step in prevention of new establishments, evaluating species in terms of their potential for spreading widely and negatively impacting ecosystems might become part of the means by which resource managers prioritize control efforts in environments with large numbers of introduced species.

Tupinambis merianae as nest predators of crocodilians and turtles in Florida, USA

Tupinambismerianae, is a large, omnivorous tegu lizard native to South America. Two populations of tegus are established in the state of Florida, USA, but impacts to native species are poorly documented. During summer 2013, we placed automated cameras overlooking one American alligator (Alligator mississippiensis) nest, which also contained a clutch of Florida red-bellied cooter (Pseudemys nelsoni) eggs, and one American crocodile (Crocodylus acutus) nest at a site in southeastern Florida where tegus are established. We documented tegu activity and predation on alligator and turtle eggs at the alligator nest, and tegu activity at the crocodile nest. Our finding that one of the first two crocodilian nests to be monitored was depredated by tegus suggests that tegus should be further evaluated as a threat to nesting reptiles in Florida.

Divergence in morphology, but not habitat use, despite low genetic differentiation among insular populations of the lizard Anolis lemurinus in Honduras

Studies of recently isolated populations are useful because observed differences can often be attributed to current environmental variation. Two populations of the lizard Anolis lemurinus have been isolated on the islands of Cayo Menor and Cayo Mayor in the Cayos Cochinos Archipelago of Honduras for less than 15000 y. We measured 12 morphometric and 10 habitat-use variables on 220 lizards across these islands in 2 y, 2008 and 2009. The goals of our study were (1) to explore patterns of sexual dimorphism, and (2) to test the hypothesis that differences in environment among islands may have driven divergence in morphology and habitat use despite genetic homogeneity among populations. Although we found no differences among sexes in habitat use, males had narrower pelvic girdles and longer toe pads on both islands. Between islands, males differed in morphology, but neither males nor females differed in habitat use. Our data suggest that either recent selection has operated differentially on males despite low genetic differentiation, or that they display phenotypic plasticity in response to environmental variation. We suggest that patterns may be driven by variation in intrapopulation density or differences in predator diversity among islands.

Invasive Brown Treesnake Movements at Road Edges Indicate Road-Crossing Avoidance

Abstract Roads have significant impacts on the dispersal of wildlife. Although this poses a threat to the abundance and diversity of desirable flora and fauna, it also affords some opportunity for enhancing control of invasive species. Roads are the most common terrain features that may affect the rate of landscape-scale movements of invasive Brown Treesnakes (Boiga irregularis) throughout Guam. We radio tracked 45 free-ranging Brown Treesnakes in close proximity to two roads in Guam and recorded instances where daily relocations of snakes spanned roads. Then we reconstructed observed movement histories with randomized turning angles, which served as a useful null hypothesis for assessing the effect of roads or road edge habitat on Brown Treesnake movement patterns. Random walk simulations demonstrated that Brown Treesnakes crossed these roads at a rate far lower than would be expected if snake movement was random with respect to roads and road edge habitat. We discuss two alternative hypotheses for these results: 1) habitat gaps posed by roads physically or behaviorally restrict snake movement; or 2) road edges provide preferred foraging habitat from which snakes are reluctant to depart. Because roads often form the boundaries of jurisdictional and management units, the effects of roads on the movement of invasive Brown Treesnakes will influence the prospects for success of future landscape-level suppression efforts.