Wild Burmese python nest site selection, thermogenesis, and brooding behaviors in the Greater Everglades Ecosystem

Abstract

Invasive Burmese pythons (Python bivittatus) are extremely cryptic animals. Although their conservation status in their native range is Vulnerable, in the Greater Everglades Ecosystem (Florida, USA) they have become a dominant destructive force and usually are immediately removed whenever found. This poses a paradox where removals are occurring, yet the study and understanding of python ecology is needed to inform removal methods. An important component of life history includes the nesting season, but little is known about python nest site selection, nest brooding, thermogenesis, or hatching success in the wild. Here, we present the first complete and most detailed report of oviposition and brooding for this biologically significant time period of a female Burmese python. We describe anthropogenic nest site selection, document the extent of shivering thermogenesis, describe brooding behaviors, and provide photo‐documentation of complete hatching of the largest python clutch on record.

Oviparous (egg-laying) female reptiles influence individual reproductive success through a variety of factors (i.e., maternal effects) including selection of oviposition sites (nests) that incorporate favorable cover and microclimatic conditions (Bernardo, 1996; Refsnider & Janzen, 2010). Ectotherms lack metabolic heat production; therefore, those microclimatic conditions can significantly influence individual fitness as thermal regimes that eggs experience during incubation directly affect offspring survival and phenotypes (e.g., body size, length-to-mass ratio, growth rate, predator avoidance, feeding behaviors, and sex; Dunham et al., 1989; Josimovich et al., 2021; Shine et al., 1997). However, highly adaptable, generalist species are capable of overcoming suboptimal environmental conditions through behavioral thermoregulation (Bodensteiner et al., 2023; Shine, 2004). Examples of these types of behaviors can be seen in reptiles such as the Argentine black and white tegu lizard (overwintering thermal stability; Currylow et al., 2021) and Burmese python (Python bivittatus; nest brooding and shivering thermogenesis; Snow et al., 2010). Both of these reptiles are highly successful and ecologically destructive invasive species in southern Florida, USA (Guzy et al., 2023; Offner et al., 2021). Though difficult to document in situ, these species' success within their novel environment is thought to be attributable to behavioral and thermal plasticity supporting individual survival and increased fecundity (Goetz et al., 2021; Shine et al., 1997; Stahlschmidt & DeNardo, 2010). For example, unlike most reptiles, Burmese pythons provide maternal care via nest brooding and clutch defense (e.g., Currylow, McCollister, et al., 2022) and may produce heat for eggs via shivering thermogenesis (Snow et al., 2010). However, the prevalence or effectiveness of such behaviors is almost entirely unknown. Here, we present a rare view of the complete sequence of the secretive nesting behaviors of a large, wild Burmese python (Figure 1, Table 1; Video S1) using radiotelemetry, wildlife cameras, and temperature dataloggers. Better understanding of the fundamental aspects of this crucial time period, particularly for the largest/most fecund individuals, can help inform the development of novel control tools.

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TABLE 1 Select behavioral timeline with temperatures of photo-documented brooding Burmese python (Python bivittatus) on a clutch of eggs within a concrete pipe (Video S1) located within the Greater Everglades Ecosystem, Florida, USA.

Date	Temperature (°C)	Video timing	Activity description
Clutch	Pipe
2022-05-23	…	…	…	Python tracked to concrete pipe. Python had oviposited (Figure 1A).
2022-06-02	32.6	24.4	00:09	Biologists deployed dataloggers and camera facing brooding female prior day; eyeshine visible (Figure 1B). Video S1 shows python shivering and shifting coils.
2022-06-05	33.2	24.2	00:34	Virginia opossum (Didelphis virginiana) entered pipe and promptly leaves (Figure 1C).
2022-06-17	32.9	26.7	00:43	Brooding female with head lowered (Figure 1D). Video S1 shows python shivering.
2022-06-18	32.8	26.3	00:51	Brooding female yawns (Figure 1E). Video S1 shows python shivering and shifting.
2022-06-20	30.0	25.9	01:06	Smaller Virginia opossum entered pipe and promptly leaves (Figure 1F).
2022-06-26	26.1	25.1	01:26	Brooding female yawns from within coils (Figure 1G). Video S1 python shivering.
2022-06-27	30.7	27.5	…	Biologists moved camera inside and installed mesh screens to seal pipe (Figure 1H).
	30.7	29.1	01:43	Disturbed by biologist activity, python moved away from clutch (Figure 1I).
	28.3	26.1	01:52	Evening. Python remained near mesh at the end of pipe away from clutch.
2022-06-29	26.8	26.8	01:58	Python remains off clutch but rests head on left side of clutch facing camera.
2022-07-02	30.5	29.5	02:21	Python returned to clutch and resumed brooding/shivering. One egg was displaced between python and camera, while clutch proximal dataloggers are encompassed in coils (Figure 1, bottom; Video S1).
2022-07-03	30.9	26.5	…	Python moves head while investigating; appears to notice displaced egg in Video S1.
2022-07-07	31.3	26.3	02:43	Python's head close to camera, appearing to investigate camera (Figure 1J).
2022-07-09	31.5	29.5	02:49	Brooding female yawns toward camera (Figure 1K).
2022-07-10	31.3	31.4	02:50	Brooding female adjusts coils and reveals eggs beneath (Figure 1L).
2022-07-13	31.2	27.0	02:57	Python begins to uncoil from clutch.
	29.3	28.6		Python moves off clutch within pipe (Figure 1M).
2022-07-14	28.0	26.3	03:08	Biologists temporarily open mesh netting, allowing python to leave.
2022-07-19	30.3	30.1	03:22	Clutch temperatures increase with activity. First egg hatches (Figure 1N).
2022-07-22	28.7	26.1	03:35	Many eggs hatch. Clutch temperatures increase to >2°C with activity. Hatchlings explore pipe (Figure 1O; Video S1).
	28.5	26.9	03:38	Hatchlings continue to explore prior to biologist collecting them (Figure 1P).
2022-07-26	28.5	25.9	03:48	Dataloggers stop collecting temperature data. Memory storage full (Figure 1, bottom).
2022-08-02	…	…	…	Biologists retrieve equipment and remaining eggs/eggshells.

As part of ongoing studies involving invasive reptiles in the Greater Everglades Ecosystem of Florida, USA, we radiotrack wild Burmese pythons throughout the year using VHF telemetry. Tracking outside of the breeding season (which is ~100 days from December to March; Currylow, Falk, et al., 2022) provides opportunities to collect data on poorly known aspects of free-ranging, wild python ecology, including nesting (Figure 1). On 28 January 2022, we captured and began tracking an adult female python (73.6 kg and 442 cm snout–vent length [SVL]). By 23 May 2022, we found that the python had oviposited inside a reinforced concrete pipe (diameter: 46 cm [18 inches], length: approximately 6 m; Figure 1A,B). Though reports of this species using anthropogenically altered sites have been documented (Hanslowe et al., 2016; Lord et al., 2023; Smith et al., 2024), this species is considered to generally select oviposition sites in thick underbrush or debris piles in protected forest floor duff (e.g., Currylow, McCollister, et al., 2022; Sandfoss et al., 2024; Smith et al., 2024). Those natural nesting materials offer varying thermal habitats (Hilton et al., 2004; Magnusson et al., 1985) that have the potential to self-maintain higher clutch temperatures as the vegetation materials naturally decompose (Magnusson, 1979; Tansey, 1973). However, we report that this nest was located within an abandoned building materials storage area in Big Cypress National Preserve. Adjacent to the openly stored nest pipe were other concrete pipes of varying sizes (38–91 cm diam), corrugated metal pipes (59–91 cm diam), and black corrugated plastic drainpipes (61–91 cm diam). Two large gravel rock piles formed a quarry approximately 30 m north northwest of the pipes. After we located and observed the python, we left the area to minimize disruption and the chance the brooding female would abandon the clutch.

On 1 June 2022, we deployed three pairs of temperature dataloggers (each pair logged temperatures at 20- and 40-min intervals). We placed one pair within the python's coils (using a 1.6-m flexible plumbing grabber claw tool affixed to a 2-m fishing pole; “in clutch”) and two pairs on the bottom of the pipe next to the animal's coils (epoxied to a wooden stake at 0 cm [clutch proximal] and 40 cm [clutch distal]). We also deployed a wildlife surveillance camera (set to take a photograph every minute) outside the pipe, facing in. We infrequently changed the SD card in the camera to minimize disturbance to the brooding python.

Beginning 2 June 2022, images from the wildlife camera indicated steady micro-adjustment movements by the python, consistent with shivering thermogenesis (Video S1). Together with temperature data (McBride et al., 2025), we confirmed that this brooding female was exhibiting shivering thermogenesis and incubated the clutch at temperatures >5°C above ambient (33.0 vs. 27.7°C; Figure 1, bottom). While brooding (23 May to 13 July 2022), the python continued to shift her coils and yawned frequently (Figure 1, top; Video S1). Interestingly, it has been argued that yawning behavior in snakes is not spontaneous as that in other animals but is a function of preparation for or recovery from feeding (stretching, readjusting, and dislocating the jaw to accommodate large prey consumption; e.g., Gallup, 2011); however, this does not appear to be the case here as pythons fast during incubation (Murthy, 2010). Though brooding behavior is presumed to help protect the vulnerable clutch from nest depredation, only a single instance of this has been previously confirmed (Currylow, McCollister, et al., 2022). Here, the camera twice captured potential nest predators, Virginia opossums (Didelphis virginiana), entering the pipe, apparently detecting the python, and exiting within the same minute (Figure 1C,F, Table 1; Video S1).

Previous data on wild Burmese pythons in Florida indicate that hatching commences approximately 60 days after oviposition (Currylow, Falk, et al., 2022), but the brooding female may depart the nest just prior to hatching (Hanslowe et al., 2016; Wolf et al., 2016). On 27 June 2022, in anticipation of the python's departure from the nest and to ensure that no hatchlings would be inadvertently released into the environment, we moved the camera inside the pipe and enclosed it with the brooding female using two layers of heavy-duty nylon mesh, which were ratchet-strapped around the pipe openings (Figure 1H). It was presumably this activity that caused the python to move off her clutch (Figure 1I) but remain within the enclosed pipe. Subsequent wildlife camera photographs revealed that, for several days, the python remained adjacent to but not coiled around her clutch. Throughout that time, nest temperatures dropped from approximately 31.0°C to as low as 24.5°C (Figure 1, bottom). On 2 July, the python returned to coil around her clutch and resumed brooding, and the nest temperatures slowly rose ca. 4°C (26.8–30.8°C; Figure 1, bottom; Table 1). During this readjustment, the female's coils encompassed the pair of “clutch-proximal” temperature dataloggers and one egg was displaced from the clutch, remaining outside the python's coils (Figure 1J–L and bottom graph [red lines]).

On 13 July 2022, 51 days after oviposition, the python completely moved off the clutch of eggs (Figure 1M). On 14 July, we removed the mesh to allow the telemetered python to leave the nesting location on her own. Approximately 8 h later, we returned to find that the python had left the pipe, and we re-installed the mesh. Clutch temperatures fell to align with ambient. On 19 July, 6 days after the python ceased brooding, the first eggs pipped, the clutch began to hatch (Figure 1N–P; Video S1), and clutch temperatures began to raise ~2°C above ambient with hatching activity (Figure 1, bottom). We physically checked on the nest and removed hatchlings one to two times daily until no more hatched. Though the majority of the clutch hatched between 19 and 22 July, a few individuals were delayed over several days, the last emerging as late as 13 days after the first (on 1 August; Table 1; McBride et al., 2025). We removed the entire nest of eggshells on 2 August, checked for hatching completeness, and enumerated the clutch.

The nest contained 96 eggs: the largest Burmese python nest verified to date (Currylow, Evers, et al., 2023). Of those, 83 hatched at an equal sex ratio (41 female, 41 male, 1 unrecorded). Though this species can be prolific, average clutch sizes of wild Burmese pythons in Florida are much lower than those noted here, at approximately 49 eggs per clutch (Currylow, Falk, et al., 2022). Further, we have observed several instances of oviposited but inviable eggs (some smaller and/or misshapen; 3 of 74 and 12 of 62 in Josimovich et al., 2021; 9 of 40, 9 of 79, and 2 of 39 in Currylow, Falk, et al., 2022; 2 of 25 in Hanslowe et al., 2016; 2 of 46 in Snow et al., 2007; 13 of 96 in Currylow, Evers, et al., 2023). There is evidence of maladaptation as some eggs are not oviposited but retained within the reproductive tract in this invasive population (Anderson et al., 2022).

This report is only the second known of a wild, invasive Burmese python using an anthropogenic structure to oviposit a clutch, and both nested in concrete culvert pipes (see Hanslowe et al., 2016). Despite the area surrounding our python's nest comprising several types and sizes of pipes (Figure 1A), she selected a concrete pipe. It is possible that the protection, lighting, and microclimate of concrete could be more favorable than those of surrounding substrates for a brooding female. Pipe temperatures 40 cm from our python's clutch were relatively stable, only varying from 25 to 29°C (Figure 1, bottom). When the python was brooding, nest temperatures were maintained relatively stable around 32°C. However, nesting behaviors did not come without cost to the female's body condition. Likely due to both egg production and thermogenesis, the python lost 27.9% of her pre-nesting body mass (from 67.8 to 48.9 kg). Reproductive data indicate that Burmese pythons may nest every 2–3 years (Currylow, Falk, et al., 2022), and it may take that long for a female to physiologically recover (Madsen & Shine, 1999; Stahlschmidt et al., 2012; Stahlschmidt & DeNardo, 2010). Concordantly, the python in this study recovered only 77% (to 52.1 kg in December) of pre-breeding season mass by years' end, despite having been observed with a food bolus later in the year.

Brooding pythons rely solely on stored bodily reserves (Spotila & Standora, 1985), and thus energy expenditure is high, but as a result, increased temperatures can have lasting, positive effects on the survival and fitness of hatchlings (Booth, 2006; De Jong et al., 2023; Shine et al., 1997). While concealing a nest in vegetation for clutch incubation could have a negligible reduction in thermally induced fitness compared to brooding (Aubret et al., 2005), an unattended clutch would leave the nest vulnerable to depredation (Currylow, McCollister, et al., 2022; DeNardo et al., 2012; Spotila & Standora, 1985). Although nesting in a concrete pipe may offer limited thermal benefit (but refer to Massey & Hutchings, 2021), it does provide a secure and defendable cover site. Indeed, we documented two instances of potential nest predators deterred from accessing the clutch (Figure 1C,F). Further, nest defense can be energetically costly, but the protection offered by a concrete pipe could reduce energy expenditures as the female only has to monitor for threats in two directions and, in turn, may physiologically focus energy on heat production.

The capacity of this species to select and behaviorally acclimate to a variety of habitats and thermal environments, including anthropogenically disturbed areas (e.g., Hanslowe et al., 2016; Lord et al., 2023; Smith et al., 2024), may influence the extent of the invasion front. Though pythons have been established in Florida for several decades, they are so difficult to study in the wild (Guzy et al., 2023) that nearly all of what we understand about wild Burmese python nest site selection, oviposition, clutch size, brooding behaviors (including shivering thermogenesis), hatching success, and depredation is informed through publication of single observations such as this (e.g., op. cit., Crawford et al., 2023; Currylow, Fitzgerald, et al., 2023; Evers et al., 2024; McCollister et al., 2021). Here, we provide a complete compilation of naturally occurring, wild python nesting data throughout the entirety of the nesting period. Detailed observations like these of multiple physiological and environmental variables throughout complete biologically significant timelines provide a novel window into the reproductive capacity, adaptability, and invasion potential of this large but cryptic animal.

ACKNOWLEDGMENTS

Funding for this work was provided by US Geological Survey (USGS) Greater Everglades Priority Ecosystem Science (GEPES) Program, USGS Biological Threats and Invasive Species Research Program, and USGS Fort Collins Science Center. We thank T. Dean, B. Falk, T. Pernas, and S. Schulze of the National Park Service (NPS) and N. Aumen of the USGS for facilitation of this project. We thank F. Ridgley and veterinary staff at Zoo Miami for surgically implanting radiotransmitters (Holohil AI-2, Carp, Ontario, Canada) and animal support; D. Fliehler, B. Gross, M. Murray, K. Treichel, J. Torres, T. Evers, K. Woytek, E. Ribas, S. Thrasher, P. Crawford, T. Hough, NPS staff, volunteers, South Florida Water Management District personnel, and Florida Fish and Wildlife Conservation Commission personnel that assisted with associated projects leading to the collection of these data; and two anonymous reviewers for helpful feedback on earlier versions of this manuscript. This work is authorized under USGS FORT IACUC approval 2020-06, NPS IACUC approval SER_BICY_McCollister_Python_2019.A3, FL_BICY_Currylow_JuvPython_2020.A2, and amendments. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the US government.

CONFLICT OF INTEREST STATEMENT

The authors declare no conflicts of interest.

DATA AVAILABILITY STATEMENT

Data (McBride et al., 2025) are available from the U.S. Geological Survey ScienceBase-Catalog: .

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