Thermoregulation in the Papuan-New Guinean boid and colubrid snakes, Candoia carinata, Candoia aspera and Boiga irregularis

Thermoregulation was investigated in the Papuan-New Guinean boids, Candoia carinata and C. aspera , and the arboreal colubrid, Boiga irregularis , based upon radiant and solar heating with thermocouple implants in the head and mid-body. Significant differences were found between head and body thermal preferenda values with either grouped data or between specimens in C. carinata and B. irregularis , but not in C. aspera , although higher temperatures were tolerated by the body than the head in all species examined. Less diversity in thermal control methods was found in this study than in previous investigations on some Australian pythons and elapid snakes, suggesting that possibly there is less need for such diversity in tropical rainforest snakes, as compared to those living in more temperate or arid environments.     

Cold-induced mortality of invasive Burmese pythons in south Florida

A recent record cold spell in southern Florida (2–11 January 2010) provided an opportunity to evaluate responses of an established population of Burmese pythons (Python molurus bivittatus) to a prolonged period of unusually cold weather. We observed behavior, characterized thermal biology, determined fate of radio-telemetered (n = 10) and non-telemetered (n = 104) Burmese pythons, and analyzed habitat and environmental conditions experienced by pythons during and after a historic cold spell. Telemetered pythons had been implanted with radio-transmitters and temperature-recording data loggers prior to the cold snap. Only one of 10 telemetered pythons survived the cold snap, whereas 59 of 99 (60%) non-telemetered pythons for which we determined fate survived. Body temperatures of eight dead telemetered pythons fluctuated regularly prior to 9 January 2010, then declined substantially during the cold period (9–11 January) and exhibited no further evidence of active thermoregulation indicating they were likely dead. Unusually cold temperatures in January 2010 were clearly associated with mortality of Burmese pythons in the Everglades. Some radio-telemetered pythons appeared to exhibit maladaptive behavior during the cold spell, including attempting to bask instead of retreating to sheltered refugia. We discuss implications of our findings for persistence and spread of introduced Burmese pythons in the United States and for maximizing their rate of removal.     

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.     

Temporal and spatial complexity of maternal thermoregulation in tropical pythons

Parental care is a widespread adaptation that evolved independently in a broad range of taxa. Although the dynamics by which two parents meet the developmental needs of offspring are well studied in birds, we lack understanding about the temporal and spatial complexity of parental care in taxa exhibiting female-only care, the predominant mode of parental care. Thus, we examined the behavioral and physiological mechanisms by which female water pythons Liasis fuscus meet a widespread developmental need (thermoregulation) in a natural setting. Although female L. fuscus were not facultatively thermogenic, they did use behaviors on multiple spatial scales (e.g., shifts in egg-brooding postures and surface activity patterns) to balance the thermal needs of their offspring throughout reproduction (gravidity and egg brooding). Maternal behaviors in L. fuscus varied by stage within reproduction and were mediated by interindividual variation in body size and fecundity. Female pythons with relatively larger clutch sizes were cooler during egg brooding, suggesting a trade-off between reproductive quantity (size of clutch) and quality (developmental temperature). In nature, caregiving parents of all taxa must navigate both extrinsic factors (temporal and spatial complexity) and intrinsic factors (body size and fecundity) to meet the needs of their offspring. Our study used a comprehensive approach that can be used as a general template for future research examining the dynamics by which parents meet other developmental needs (e.g., predation risk or energy balance).     

Parasite spillover: indirect effects of invasive Burmese pythons

Identification of the origin of parasites of nonindigenous species (NIS) can be complex. NIS may introduce parasites from their native range and acquire parasites from within their invaded range. Determination of whether parasites are non‐native or native can be complicated when parasite genera occur within both the NIS’ native range and its introduced range. We explored potential for spillover and spillback of lung parasites infecting Burmese pythons (Python bivittatus) in their invasive range (Florida). We collected 498 indigenous snakes of 26 species and 805 Burmese pythons during 2004–2016 and examined them for lung parasites. We used morphology to identify three genera of pentastome parasites, Raillietiella, a cosmopolitan form, and Porocephalus and Kiricephalus, both New World forms. We sequenced these parasites at one mitochondrial and one nuclear locus and showed that each genus is represented by a single species, R. orientalis, P. crotali, and K. coarctatus. Pythons are host to R. orientalis and P. crotali, but not K. coarctatus; native snakes are host to all three species. Sequence data show that pythons introduced R. orientalis to North America, where this parasite now infects native snakes. Additionally, our data suggest that pythons are competent hosts to P. crotali, a widespread parasite native to North and South America that was previously hypothesized to infect only viperid snakes. Our results indicate invasive Burmese pythons have affected parasite‐host dynamics of native snakes in ways that are consistent with parasite spillover and demonstrate the potential for indirect effects during invasions. Additionally, we show that pythons have acquired a parasite native to their introduced range, which is the initial condition necessary for parasite spillback.     

Homing of invasive Burmese pythons in South Florida: evidence for map and compass senses in snakes

Navigational ability is a critical component of an animal''s spatial ecology and may influence the invasive potential of species. Burmese pythons (Python molurus bivittatus) are apex predators invasive to South Florida. We tracked the movements of 12 adult Burmese pythons in Everglades National Park, six of which were translocated 21–36 km from their capture locations. Translocated snakes oriented movement homeward relative to the capture location, and five of six snakes returned to within 5 km of the original capture location. Translocated snakes moved straighter and faster than control snakes and displayed movement path structure indicative of oriented movement. This study provides evidence that Burmese pythons have navigational map and compass senses and has implications for predictions of spatial spread and impacts as well as our understanding of reptile cognitive abilities.     

Black box and attics: Habitat selection and resource use by large threatened pythons in landscapes with contrasting human modification

Woodland and forest degradation, driven predominately by agricultural and pastoral production, is a crisis facing many species globally, in particular hollow‐dependent fauna. Large predatory species play important roles in both ecosystems and conservation strategies, but few studies have examined habitat selection of such species in intensively human‐modified landscapes. We quantified habitat selection and resource use by a large, top‐order and threatened snake (carpet python, Morelia spilota), between adjacent areas of high and low anthropogenic modification in inland Australia, a region that has undergone considerable alteration since European settlement. At the low‐impact site, snakes preferred tree hollows and a structurally complex understorey, whereas at the high‐impact site, snakes preferred homestead attics. Based on the decline of the species in this region, however, high‐impact landscapes may only support snakes when they are adjacent to low‐impact habitats. Invasive species comprised a large part of snake diets in both landscape types. Carpet pythons, with large home ranges and habitat requirements that overlap with many smaller threatened mammalian and avian fauna, are generally well liked and easily identifiable by rural landholders. Accordingly, they may play a key role in conservation strategies aimed at the protection of woodland and hollow‐dependent fauna in heavily modified landscapes of Australia's inland regions. However, invasive species, which tend to contribute to declines in native species inhabiting arid and semi‐arid Australia, are beneficial and important to pythons. Our study therefore highlights the diversity of effects that two major threats to biodiversity – habitat loss and invasive species – can have on different species within the same ecosystem.     

The use of ultrasonography to assess reproductive investment and output in pythons

Reproductive investment and output are integral fitness components, often incorporated into life‐history trade‐off models and important to population dynamics. The trade‐offs associated with reproduction can be dramatic in species such as snakes that make especially large investments into reproduction. Unfortunately, traditional methods used to determine reproductive investment and output are effective in many (but not all) situations. Thus, we used portable ultrasonography to serially estimate reproductive investment and reproductive output in three python species that exhibit significant variation in phylogeny, geographic range, body size, egg size, and clutch size: ball pythons (Python regius), Children's pythons (Antaresia childreni), and water pythons (Liasis fuscus). At each time point of measurement (range: 1–49 days pre‐oviposition), ultrasound estimates of viable clutch size were highly accurate in all three species. However, ultrasound estimates of mean viable egg mass, and thus viable clutch mass, significantly differed from the actual values (range: 23–73% error). Interestingly, this error was considerably smaller as females approached oviposition, suggesting that female pythons transfer a significant amount of water into their eggs during the week before oviposition. Thus, water balance during late‐stage egg development may be an integral part of reproductive success. The results obtained in the present study form the foundation for future assessments of reproductive investment, and also provide insight into the use of ultrasound technology to assist such efforts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 772–778.     

Invasive pythons,not anthropogenic stressors,explain the distribution of a keystone species

Untangling the causes of native species loss in human-modified systems is difficult and often controversial. Evaluating the impact of non-native species in these systems is particularly challenging, as additional human perturbations often precede or accompany introductions. One example is the ongoing debate over whether mammal declines within Everglades National Park (ENP) were caused by either the establishment of non-native Burmese pythons (Python molurus bivittatus) or the effects of other anthropogenic stressors. We examined the influence of both pythons and a host of alternative stressors—altered hydrology and habitat characteristics, mercury contamination and development—on the distribution of the marsh rabbit (Sylvilagus palustris), a once common mammal in ENP. Distance from the epicenter of the python invasion best explained marsh rabbit occurrence in suitable habitat patches, whereas none of the alternative stressors considered could explain marsh rabbit distribution. Estimates of the probability of marsh rabbit occurrence ranged from 0 at the python invasion epicenter to nearly 1.0 150 km from the invasion epicenter. These results support the hypothesis that invasive pythons shape the distribution of marsh rabbits in southern Florida. The loss of marsh rabbits and similar species will likely alter trophic interactions and ecosystem function within the Everglades, an internationally important hotspot of biodiversity. Further, our results suggest that non-native species can have profound impacts on mainland biodiversity.     

Thermoregulation in four Australian lizards of the genus Egernia (Sauria: Scincidae)

Mean thermal preferenda for the head and body were found to be similar in Egernia whitei, E. cunninghami, and E. bungana (range in T_H= 33.6^o -33.8^o C; range in T_B= 34.0^o-35.3^oC). However, those for E. majors were significantly lower (T_H= 31.1^o C, T_B=32.9^oC). Significant differences were found between mean head and body preferred temperatures in E. cunninghami and E. major. Higher temperatures were tolerated by the body than the head in all four species. Gaping was not observed to occur prior to cessation of respiration in any of the four species during radiant or solar heating. Based upon thermal preferenda and critical thermal maxima data, thermal safety margins for body temperature were calculated for E. whitei and E. cunninghami Postural orientation to the sun's rays increased and decreased heat gain. The seeking of shade was the most effective method employed by all four species for reducing head and body temperature.     

Perch color preference in juvenile green tree pythons,Chondropython viridis

Green tree pythons, Chondropython viridis, are polymorphic for color as juveniles, commonly being primarily yellow or brown until becoming mostly green at about 1 year of age. We tested the hypothesis that the different morphs arose as a result of selection for differential background matching, yellow morphs selecting light-colored backgrounds, and brown morphs selecting dark-colored backgrounds. Twelve yellow and eight brown morphs were placed repeatedly in individual testing enclosures and allowed to choose between black and white or yellow and brown halves of a t-perch. Trials showed that both color morphs preferred dark over light perches. We tentatively suggest that individuals chose dark-colored perches for purposes of concealment. © 1994 Wiley-Liss, Inc.     

Reptilian endothermy: a field study of thermoregulation by brooding diamond pythons

Miniature temperature-sensitive radiotransmitters were surgically implanted into free-ranging adult diamond pythons ( Morelia s. spilota ), which are medium-sized boid snakes of south-eastern Australia. Four female pythons oviposited during the study, and constructed incubation mounds. These apparently provided excellent insulation, and the snakes maintained high (approx. 31 C) and relatively constant body temperatures throughout the two-month incubation period. They apparently maintained these temperatures primarily by endogenous heat production (shivering thermogenesis), but also basked briefly on most mornings.
Brooding females maintained a body temperature differential above ambient of about 9C, occasionally up to 13C; their temperatures were significantly higher amd less variable than those of non-brooding females or males. The energetic cost of brooding must be high, but these costs may be outweighed by the benefits of rapid embryonic development and high embryonic survivorship.     

Betrayal: radio-tagged Burmese pythons reveal locations of conspecifics in Everglades National Park

The “Judas” technique is based on the idea that a radio-tagged individual can be used to “betray” conspecifics during the course of its routine social behavior. The Burmese python (Python bivittatus) is an invasive constrictor in southern Florida, and few methods are available for its control. Pythons are normally solitary, but from December–April in southern Florida, they form breeding aggregations containing up to 8 individuals, providing an opportunity to apply the technique. We radio-tracked 25 individual adult pythons of both sexes during the breeding season from 2007–2012. Our goals were to (1) characterize python movements and determine habitat selection for betrayal events, (2) quantify betrayal rates of Judas pythons, and (3) compare the efficacy of this tool with current tools for capturing pythons, both in terms of cost per python removed (CPP) and catch per unit effort (CPUE). In a total of 33 python-seasons, we had 8 betrayal events (24 %) in which a Judas python led us to new pythons. Betrayal events occurred more frequently in lowland forest (including tree islands) than would be expected by chance alone. These 8 events resulted in the capture of 14 new individuals (1–4 new pythons per event). Our effort comparison shows that while the Judas technique is more costly than road cruising surveys per python removed, the Judas technique yields more large, reproductive females and is effective at a time of year that road cruising is not, making it a potential complement to the status quo removal effort.     

Identifying plausible scenarios for the establishment of invasive Burmese pythons (<Emphasis Type="Italic">Python molurus</Emphasis>) in Southern Florida

Successful invasions of secretive alien species often go unrecognized until spread has exceeded the point where control or eradication is feasible. In such situations, understanding factors that contributed to establishment can be critical to preventing subsequent introductions of previously-successful invaders or ecologically similar species. The Burmese python (Python molurus bivittatus), a native to Southeast Asia, is abundant in the pet trade and is now well-established in southern Florida. Although there can be little argument that the ultimate source of Florida pythons was the pet industry, there has been limited consideration of biological support for scenarios that may have lead to their establishment. In this study we use information on python capture rates and biologically-derived population growth models to evaluate the plausibility of various scenarios for python establishment. Our results indicate that scenarios involving relatively recent establishment (post-1990) require large numbers (100–1,000) of founders or unrealistically high juvenile survivorship. Intentional simultaneous release of large numbers of pythons is unlikely and accidental release of large numbers of founders is inconsistent with the spatial and temporal pattern of pythons captures in the region. We conclude that the most parsimonious scenario for establishment of pythons in Florida involves the release of a relatively small number of founders prior to 1985. Our results demonstrate that for pythons and other species with low inherent detection probabilities, early action during incipient phases of an invasion is critical and understanding likely introduction scenarios is important for preventing similar situations from occurring elsewhere or with other species.     

A classification of pythons (Serpentes, Pythoninae)

The Central American Loxocemus and the pythons are assigned to sister subfamilies Loxoceminae and Pythoninae, within the weakly characterized family Boidae. They share the character of the left cerebral artery foramen being larger than the right. Within the Henophidia, paired subcaudal scales associate these subfamilies with Xenopeltis and the Uropeltidae.
Twenty-nine discrete variables were coded as 52 binary characters. Nine continuous (meristic) variables were examined. A method is described by which 18 binary characters were selected from seven of these variables, for inclusion in the data set. With Loxocemus and Xenopeltis as outgroups, the data were analysed by a compatibility method. The Pythoninae are resolved into tribes Pythonini and Moreliini on the basis of complementary synapomorphies. The Pythonini have only the genus Python , for the African and Asiatic species, including reticulatus and timoriensis. The Moreliini have all of the Australasian species, in two genera: Aspidites and Morelia.
A phylogenetic analysis of the two tribes is presented, with incomplete resolution of the Moreliini. The two tribes overlap in the areas of the Moluccas and Timor. It is postulated that: Loxocemus plus Pythoninae represent a Laurasian stock; south-east Asia was the primary centre of radiation of the Pythoninae; by the Miocene dispersal to Australasia had occurred, where there was a second radiation.     

Cold weather and the potential range of invasive Burmese pythons

The Burmese python (Python molurus bivittatus) is established in Everglades National Park and neighboring areas in south Florida. Beyond its substantial ecological impacts to native fauna in south Florida, concerns have been raised as to its potential to occupy other parts of the USA, even as far north as Washington, DC. During a recent period of cold weather, seven of nine captive Burmese pythons held in outdoor pens at our facility in north-central Florida died, or would have died absent our intervention. This cold-induced mortality occurred despite the presence of refugia with heat sources. Our findings cast doubt on the ability of free-ranging Burmese pythons to establish and persist beyond the subtropical environment of south Florida.     

Heavily exploited but poorly known: systematics and biogeography of commercially harvested pythons (Python curtus group) in Southeast Asia

More than 100000 blood pythons (brongersmai) and short-tailed pythons (curtus and breitensteini) are taken from Borneo and Sumatra each year for the commercial leather trade. Traditionally, all have been treated as a single polytypic species (Python curtus) , with three subspecies differing in colour, size and geographic distribution. Analyses of DNA sequences and morphological data clarify the phylogenetic relationships, taxonomy and biogeography of this group. The lineage is monophyletic, and each of the three subspecies differs from the other two both morphologically and genetically. Given the morphological and genetic distinctiveness of each taxon, we here elevate the three subspecies to full species status. Python brongersmai is the most distinctive in terms of colour (of the three, only brongersmai has colour-morphs that are red or orange), size (it grows to 2.6 m, vs. approx. 2.0 m for the other taxa), and scalation (e.g. brongersmai has <166 ventral scales, vs. <166 in the other taxa and has two supralabials over each orbit, vs. one supralabial for the other two taxa). In terms of cytochrome b mitochondrial DNA sequence data, brongersmai is almost as distant genetically from the short-tailed pythons (8.9% divergence) as is the reticulated python (P. reticulatus: 10.3% divergence). The other two taxa (P. breitensteini from Kalimantan and P. curtus from western and southern Sumatra) are closely related (3% divergence), despite their disjunct distribution (separated by P. brongersmai). Sea-level fluctuations provide a plausible biogeographic scenario to explain phylogenetic divergence within this lineage. Given the distinctiveness of the component taxa, and the ease with which even dried skins can be identified to species level (based on ventral counts), the managers of this important commercial resource should no longer treat the P. curtus group as a single biological taxon.     

Ecological divergence among sympatric colour morphs in blood pythons, Python brongersmai

Blood pythons in northeastern Sumatra display a series of discrete colour morphs, even among hatchlings within a single clutch. The first step towards understanding the maintenance of this polymorphism is to test the null hypothesis that colour variation in this species has no major biological correlates. Data on >2,000 blood pythons killed for the commercial leather industry enabled us to test, and reject, this hypothesis. The four colour morphs differed significantly in most of the traits that we measured, including temporal and spatial abundances, sex ratios, age structures, mean adult body sizes, body shapes (tail length and body mass relative to snout-vent length), energy stores, numbers of gut parasites, prey types, feeding frequencies and clutch sizes. The causal basis for these associations remains unclear, but is likely to involve three processes: direct effects of colour, linkages between genes for colour and other traits, and correlated spatial heterogeneity in colour, morphology and ecology. The colour polymorphism may be maintained by frequency-dependent selection and genotype-specific habitat selection, because these sedentary ambush predators are under strong selection for effective camouflage to hide them from both predators and potential prey. In support of this hypothesis, similar colour polymorphisms have evolved independently in several other snake taxa that rely upon ambush predation. Received: 18 December 1997 / Accepted: 23 March 1998     

Change of cardiac function, but not form, in postprandial pythons

Pythons are renowned for a rapid and pronounced postprandial growth of the heart that coincides with a several-fold elevation of cardiac output that lasts for several days. Here we investigate whether ventricular morphology is affected by digestive state in two species of pythons (Python regius and Python molurus) and we determine the cardiac right-to-left shunt during the postprandial period in P. regius. Both species experienced several-fold increases in metabolism and mass of the digestive organs by 24 and 48 h after ingestion of meals equivalent to 25% of body mass. Surprisingly there were no changes in ventricular mass or dimensions as we used a meal size and husbandry conditions similar to studies finding rapid and significant growth. Based on these data and literature we therefore suggest that postprandial cardiac growth should be regarded as a facultative rather than obligatory component of the renowned postprandial response. The cardiac right-to-left shunt, calculated on the basis of oxygen concentrations in the left and right atria and the dorsal aorta, was negligible in fasting P. regius, but increased to 10-15% during digestion. Such shunt levels are very low compared to other reptiles and does not support a recent proposal that shunts may facilitate digestion in reptiles.     

Parallel selective pressures drive convergent diversification of phenotypes in pythons and boas

Pythons and boas are globally distributed and distantly related radiations with remarkable phenotypic and ecological diversity. We tested whether pythons, boas and their relatives have evolved convergent phenotypes when they display similar ecology. We collected geometric morphometric data on head shape for 1073 specimens representing over 80% of species. We show that these two groups display strong and widespread convergence when they occupy equivalent ecological niches and that the history of phenotypic evolution strongly matches the history of ecological diversification, suggesting that both processes are strongly coupled. These results are consistent with replicated adaptive radiation in both groups. We argue that strong selective pressures related to habitat‐use have driven this convergence. Pythons and boas provide a new model system for the study of macro‐evolutionary patterns of morphological and ecological evolution and they do so at a deeper level of divergence and global scale than any well‐established adaptive radiation model systems.