Cardiovascular responses and the role of the neurohumoral cardiac regulation during digestion in the herbivorous lizard Iguana iguana.

Carnivorous reptiles exhibit an intense metabolic increment during digestion, which is accompanied by several cardiovascular adjustments responsible for meeting the physiological demands of the gastrointestinal system. Postprandial tachycardia, a well-documented phenomenon in these animals, is mediated by the withdrawal of vagal tone associated with the chronotropic effects of non-adrenergic and non-cholinergic (NANC) factors. However, herbivorous reptiles exhibit a modest metabolic increment during digestion and there is no information about postprandial cardiovascular adjustments. Considering the significant impact of feeding characteristics on physiological responses, we investigated cardiovascular and metabolic responses, as well as the neurohumoral mechanisms of cardiac control, in the herbivorous lizard Iguana iguana during digestion. We measured oxygen consumption rate (O2), heart rate (fH), mean arterial blood pressure (MAP), myocardial activity, cardiac autonomic tone, fH/MAP variability and baroreflex efficiency in both fasting and digesting animals before and after parasympathetic blockade with atropine followed by double autonomic blockade with atropine and propranolol. Our results revealed that the peak of O2 in iguanas was reached 24 h after feeding, accompanied by an increase in myocardial activity and a subtle tachycardia mediated exclusively by a reduction in cardiac parasympathetic activity. This represents the first reported case of postprandial tachycardia in digesting reptiles without the involvement of NANC factors. Furthermore, this withdrawal of vagal stimulation during digestion may reduce the regulatory range for short-term fH adjustments, subsequently intensifying the blood pressure variability as a consequence of limiting baroreflex efficiency.

Ontogenetic scaling of the baroreflex function in the green iguana (Iguana iguana).

Large body mass (Mb) in vertebrates is associated with longer pulse intervals between heartbeats (PI) and thicker arterial walls. Longer PI increases the time for diastolic pressure decay, possibly resulting in loss of cardiac energy as "oscillatory power," whereas thicker arterial walls may affect the transmission of impulses and sensing of pressure fluctuations thus impairing baroreflex function. We aimed to investigate the effect of growth on the relative cardiac energy loss and baroreflex function. We predicted that 1) the relative use of cardiac energy should be preserved with increased time constant for pressure decay (τ = vascular resistance × compliance) and 2) if arterial circumferential distensibility does not change, baroreflex function should be unaltered with Mb. To test these hypotheses, we used green iguanas (Iguana iguana) weighing from 0.03 to 1.34 kg (43-fold increment in Mb). PI (P = 0.037) and τ (P = 0.035) increased with Mb, whereas the oscillatory power fraction (P = 0.245) was unrelated to it. Thus, the concomitant alterations of τ and PI allowed the conservation of cardiac energy in larger lizards. Larger animals had thicker arterial walls (P = 0.0007) and greater relative collagen content (P = 0.022). Area compliance scaled positively to Mb (P = 0.045), though circumferential distensibility (P = 0.155) and elastic modulus (P = 0.762) were unaltered. In addition, baroreflex sensitivity, measured by both the pharmacological (P = 0.152) and sequence methods (P = 0.088), and the baroreflex effectiveness index (P = 0.306) were also unrelated to Mb. Therefore, changes in arterial morphology did not affect circumferential distensibility and presumably sensing of pressure fluctuation, and the cardiovagal baroreflex is preserved across different Mb.

Conservation of a flagship species: Health assessment of the pink land iguana, Conolophus marthae.

The pink land iguana, Conolophus marthae, is one of four species of iguanas (three terrestrial and one marine) in the Galápagos Islands, and the only one listed as critically endangered by the IUCN. The species can only be found on the north-west slopes of the highest volcano on Isabela Island and was first described to science in 2009. As part of a population telemetry study, a health assessment was authorized by the Galápagos National Park. Wild adult iguanas were captured on Wolf Volcano in September 2019 and April 2021 to record morphological and physiological parameters including body temperature, heart rate, intraocular pressures, tear formation, and infrared iris images. Blood samples were also collected and analyzed. An i-STAT portable blood analyzer was used to obtain values for base excess in the extracellular fluid compartment (BEecf), glucose (Glu), hematocrit (HctPCV), hemoglobin (Hb), ionized calcium (iCa), partial pressure of carbon dioxide (pCO2), partial pressure of oxygen (pO2), percent oxygen saturation (sO2%), pH, potassium (K), and sodium (Na). When possible, data were compared to previously published and available data for the other Galápagos iguanas. The results reported here provide baseline values that will be useful in detecting changes in health status among pink land iguanas affected by climate change, invasive species, anthropogenic threats, or natural disturbances. The collected data also provide an invaluable resource for conservation scientists planning to implement conservation strategies, like translocations, that may temporarily alter these baseline values.

Cranial anatomy of the Galápagos marine iguana Amblyrhynchus cristatus (Squamata: Iguanidae).

Amblyrhynchus cristatus, the marine iguana, is unique among the ~7,000 species of living limbed lizards as it has successfully evolved adaptations that allow it to live in both terrestrial and marine environments. This species is endemic to the Galápagos Archipelago and has evolved a specialized feeding behavior, consuming primarily the algae that grow on the rocky seafloor. The intriguing questions arising around the evolution of the marine iguana concerns the use of exaptations of terrestrial features for aquatic and specifically marine adaptations. However, the lack of fundamental information about its anatomy currently prevents us from understanding how it became adapted to such a peculiar lifestyle in comparison to all other iguanids. The goal of this study is to provide the first ever description of the skull, mandible, and hyoid of Amblyrhynchus. We examined several specimens of marine iguana, including skeletal, wet, and ct-scanned material, and individuals at different ontogenetic stages. We also analyzed specimens of all other modern iguanid genera (Conolophus, Iguana, Ctenosaura, Cyclura, Dipsosaurus, Brachylophus, Sauromalus) in order to make comparisons between Amblyrhynchus and its closest relatives. We were able to identify several autapomorphic features that distinguish the marine iguana from all other iguanids. These unique morphologies are mostly associated with the modified configuration of the snout (nasal chamber), increased muscle attachments in the temporal-postorbital region of the skull, and dentition. Since Amblyrhynchus is the only nonophidian squamate currently able to exploit the ocean at least for some vital functions (i.e., feeding), we used comparisons to fossil marine lizards (e.g., mosasaurids) to discuss some of these unique traits. The new cranial features described for Amblyrhynchus may represent a source of novel morphological characters for use in future phylogenetic analyses of iguanian (or squamate) relationships, which will then serve as the foundation for the exploration of evolutionary patterns and processes that led to the development of such unique adaptations.

Thermoregulation of Liolaemus aparicioi (Iguania: Liolaemidae) along a 1000 m elevational gradient in La Paz Valley, La Paz, Bolivia.

Lizard species have diverse behavioral and physiological responses to thermo-environmental conditions, which allow them to inhabit a broad range of latitudes and elevations. Because the availability of suitable thermal resources is limited and more variable at high-elevation environments than at lower elevations, we expect high-elevation lizards to be constrained in their thermoregulation relative to lizards at lower elevations by the fewer available thermal resources to reach optimal temperatures (colder environment). We studied the thermal biology of an endemic and Critically Endangered lizard, Liolaemus aparicioi, to assess its thermal responses along a 1000 m elevational gradient in La Paz Valley from May to August of 2015 (dry season). We took field body and microhabitat temperatures at capture sites (substrate and air above ground), and body size (snout-vent length and mass) of individuals at Taypichullo (3000 m asl), Gran Jardín de la Revolución Municipal Park (3500 m asl), and Taucachi (4000 m asl) localities. Operative temperatures were taken from calibrated models deployed in different available microhabitats. Preferred temperatures and thermal tolerance limits were determined in laboratory settings for lizards from each locality. Field body, microhabitat, and operative temperatures decreased with increasing elevation and differed between sexes. Lizards at the high elevation locality had the lowest thermoregulatory efficiency as compared with the mid and lower elevation localities. In laboratory measurements, while the preferred temperatures varied between sexes, pooled preferred temperatures and thermal tolerances were similar in all localities. Although thermal resources at high elevation can limit thermoregulatory possibilities in L. aparicioi, behavioral microhabitat use, time allocated to thermoregulation, and physiological adjustments seem to be possible strategies to counteract thermal costs along elevational gradients.

Influence of light/dark cycle and orexins on breathing control in green iguanas (Iguana iguana).

Light/dark cycle affects the physiology of vertebrates and hypothalamic orexin neurons (ORX) are involved in this function. The breathing pattern of the green iguana changes from continuous to episodic across the light/dark phases. Since the stimulatory actions of ORX on breathing are most important during arousal, we hypothesized that ORX regulates changes of breathing pattern in iguanas. Thus, we: (1) Localized ORX neurons with immunohistochemistry; (2) Quantified cyclic changes in plasma orexin-A levels by ELISA; (3) Compared breathing pattern at rest and during hypoxia and hypercarbia; (4) Evaluated the participation of the ORX receptors in ventilation with intracerebroventricular microinjections of ORX antagonists during light and dark phases. We show that the ORX neurons of I. iguana are located in the periventricular hypothalamic nucleus. Orexin-A peaks during the light/active phase and breathing parallels these cyclic changes: ventilation is higher during the light phase than during the dark phase. However, inactivation of ORX-receptors does not affect the breathing pattern. Iguanas increase ventilation during hypoxia only during the light phase. Conversely, CO2 promotes post-hypercarbic hyperpnea during both phases. We conclude that ORXs potentiate the post-hypercarbic (but not the hypoxic)-drive to breathe and are not involved in light/dark changes in the breathing pattern.

Health assessment of Conolophus subcristatus, Conolophus pallidus, and C. subcristatus X Amblyrhynchus cristatus hybrid (Galápagos land iguanas).

The land iguanas, Conolophus pallidus and Conolophus subcristatu are large and charismatic lizards endemic to the Galápagos archipelago, but little information exists on their normal health parameters. The former is restricted to Santa Fe island, while C. subcristatus inhabits the islands of the central and western region of the archipelago. Both species are classified as vulnerable by the IUCN Red List of Threatened Species. As part of a population health assessment authorized by the Galápagos National Park, wild adult iguanas from three islands (North Seymour, South Plazas, and Santa Fe) were captured in July 2018. Data from a single C. subcristatus X Amblyrhynchus cristatus hybrid captured on South Plazas is also included. We analyzed blood samples drawn from 52 healthy wild adult land iguanas captured on three islands. An iSTAT portable blood analyzer was used to obtain values for pH, lactate, pO2, pCO2, HCO3-, sO2%, hematocrit, packed cell volume (PCV), hemoglobin Na, K, iCa, and glucose. Standard laboratory hematology techniques were employed for PCV determination; resulting values were also compared to the hematocrit values generated by the iSTAT. Body temperature, heart rate, respiratory rate, and body measurements were also recorded and compared to previously published data for the marine iguana (Amblyrhynchus cristatus), which shares a common ancestor with the land iguana. The data reported here provide preliminary baseline values that may be useful in comparisons between captive and wild populations, between wild populations, and in detecting changes in health status among Galápagos land iguanas affected by anthropogenic threats, climate change, or natural disturbances.

Evaluation of the sequence method as a tool to assess spontaneous baroreflex in reptiles.

The sequence method is an alternative to the traditional pharmacological approach (i.e., the Oxford technique) used to calculate baroreflex gain (G) in mammals. Although the sequence method assesses baroreflex by measuring spontaneous events of blood pressure regulation, the pharmacological method relies on the injection of vasoactive drugs that impact the baroreflex mechanism itself. The sequence method might be relevant for dynamic measurement of baroreflex modulation but it was never validated for vertebrates with low heart rate. Hence, we tested the sequence method in three species of reptiles and compared the results with those provided by the classic pharmacological method. G was similar between both methods and values correlated when parameters for the sequence method were set at delay 0 or 1 (i.e., the baroreflex system responds immediately to blood pressure changes or after 1 heartbeat). Calculation of the baroreflex effectiveness index was adequate at a minimum of 300 cycles and a delay of 1 for the three species. Therefore, the sequence method has been validated to investigate baroreflex regulation in reptiles, enabling studies during dynamic alterations in homeostasis.

Can reptiles perceive visual illusions? Delboeuf illusion in red-footed tortoise (Chelonoidis carbonaria) and bearded dragon (Pogona vitticeps).

Optical illusions have been widely used to compare visual perception among vertebrates because they can reveal how the system is able to adapt to visual input. Sensitivity to visual illusions has never been studied in reptiles. Here, we investigated whether red-footed tortoises, Chelonoidis carbonaria, and bearded dragons, Pogona vitticeps, perceive the Delboeuf illusion. This illusion involves the misperception of the size of a target circle depending upon the context in which it is presented. We adopted the same size discrimination for both species to compare their performance. Animals were presented with two different types of trials. In control trials, they received two different-sized food portions on two plates of the same size. In test trials, they received two same-sized food portions but presented on two different-sized plates. If they perceived the illusion in the same way as humans, we expected them to select the food portion presented on the smaller plate. The tortoises exhibited poor performance in the control trials, which prevented us from drawing any conclusions about their perception of the Delboeuf illusion. In contrast, the bearded dragons selected the larger amount of food in control trials. In test trials, they selected the portion presented on the smaller plate significantly more often than chance, suggesting a human-like sensitivity to the Delboeuf illusion. Our study provides the first evidence of the perception of a visual illusion in a reptile species, suggesting that rather than simply detecting visual input, they interpret sensory information captured by photoreceptors. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

The Design of an Energy Harvesting Wireless Sensor Node for Tracking Pink Iguanas.

The design of wireless sensor nodes for animal tracking is a multidisciplinary activity that presents several research challenges both from a technical and a biological point of view. A monitoring device has to be designed accounting for all system requirements including the specific characteristics of animals and environment. In this work we present some aspects of the design of a wireless sensor node to track and monitor the pink iguana of the Galápagos: a recently discovered species living in remote locations at the Galápagos Islands. The few individuals of this species live in a relatively small area that lacks of any available communication infrastructure. We present and discuss the energy harvesting architecture and the related energy management logic. We also discuss the impact of packaging on the sensor performance and the consequences of the limited available energy on the GPS tracking.

Reverse-engineering the locomotion of a stem amniote.

Reconstructing the locomotion of extinct vertebrates offers insights into their palaeobiology and helps to conceptualize major transitions in vertebrate evolution1-4. However, estimating the locomotor behaviour of a fossil species remains a challenge because of the limited information preserved and the lack of a direct correspondence between form and function5,6. The evolution of advanced locomotion on land-that is, locomotion that is more erect, balanced and mechanically power-saving than is assumed of anamniote early tetrapods-has previously been linked to the terrestrialization and diversification of amniote lineages7. To our knowledge, no reconstructions of the locomotor characteristics of stem amniotes based on multiple quantitative methods have previously been attempted: previous methods have relied on anatomical features alone, ambiguous locomotor information preserved in ichnofossils or unspecific modelling of locomotor dynamics. Here we quantitatively examine plausible gaits of the stem amniote Orobates pabsti, a species that is known from a complete body fossil preserved in association with trackways8. We reconstruct likely gaits that match the footprints, and investigate whether Orobates exhibited locomotor characteristics that have previously been linked to the diversification of crown amniotes. Our integrative methodology uses constraints derived from biomechanically relevant metrics, which also apply to extant tetrapods. The framework uses in vivo assessment of locomotor mechanics in four extant species to guide an anatomically informed kinematic simulation of Orobates, as well as dynamic simulations and robotics to filter the parameter space for plausible gaits. The approach was validated using two extant species that have different morphologies, gaits and footprints. Our metrics indicate that Orobates exhibited more advanced locomotion than has previously been assumed for earlier tetrapods7,9, which suggests that advanced terrestrial locomotion preceded the diversification of crown amniotes. We provide an accompanying website for the exploration of the filters that constrain our simulations, which will allow revision of our approach using new data, assumptions or methods.

Non-integumentary melanosomes can bias reconstructions of the colours of fossil vertebrates.

The soft tissues of many fossil vertebrates preserve evidence of melanosomes-micron-scale organelles that inform on integumentary coloration and communication strategies. In extant vertebrates, however, melanosomes also occur in internal tissues. Hence, fossil melanosomes may not derive solely from the integument and its appendages. Here, by analyzing extant and fossil frogs, we show that non-integumentary melanosomes have high fossilization potential, vastly outnumber those from the skin, and potentially dominate the melanosome films preserved in some fossil vertebrates. Our decay experiments show that non-integumentary melanosomes usually remain in situ provided that carcasses are undisturbed. Micron-scale study of fossils, however, demonstrates that non-integumentary melanosomes can redistribute through parts of the body if carcasses are disturbed by currents. Collectively, these data indicate that fossil melanosomes do not always relate to integumentary coloration. Integumentary and non-integumentary melanosomes can be discriminated using melanosome geometry and distribution. This is essential to accurate reconstructions of the integumentary colours of fossil vertebrates.

Among-Individual Variation in Desert Iguanas (Squamata: Dipsosaurus dorsalis): Endurance Capacity Is Positively Related to Home Range Size.

Among species of lizards, endurance capacity measured on a motorized treadmill is positively related to daily movement distance and time spent moving, but few studies have addressed such relationships at the level of individual variation within a sex and age category in a single population. Both endurance capacity and home range size show substantial individual variation in lizards, rendering them suitable for such studies. We predicted that these traits would be positively related because endurance capacity is one of the factors that has the potential to limit home range size. We measured the endurance capacity and home range size of adult male desert iguanas (Dipsosaurus dorsalis). Lizards were field captured for measurements of endurance, and home range data were gathered using visual identification of previously marked individuals. Endurance was significantly repeatable between replicate trials, conducted 1-17 d apart ([Formula: see text] for log-transformed values, [Formula: see text], [Formula: see text]). The log of the higher of two endurance trials was positively but not significantly related to log body mass. The log of home range area was positively but not significantly related to log body mass, the number of sightings, or the time span from first to last sighting. As predicted, log endurance was positively correlated with log home range area ([Formula: see text], [Formula: see text], one-tailed [Formula: see text]; for body-mass residual endurance values: [Formula: see text], one-tailed [Formula: see text]). These results suggest that endurance capacity may have a permissive effect on home range size. Alternatively, individuals with larger home ranges may experience training effects (phenotypic plasticity) that increase their endurance.

Predictability in evolution: Adaptation of the Bonaire anole (Anolis bonairensis) to an extreme environment.

The extent to which evolution is deterministic (predictable), or random, is a fundamental question in evolution. This case study attempts to determine the extent to which interspecific divergence can be predicted from intraspecific trends related species. The mountainous Lesser Antilles are occupied by one or two anole species with very substantial intraspecific differences in the quantitative traits between xeric and rainforest habitats. These ecologically determined differences tend to be in parallel in each island species. A related species (Anolis bonairensis) lives on the far more xeric island of Bonaire, and this study tests the extent to which its interspecific divergence in hue and pattern traits can be predicted from the parallel intraspecific variation exhibited in Lesser Antillean anoles. Regression against a multivariate climate variable suggests that the hue and pattern of the Bonaire anole are consistently predicted from the ecologically determined intraspecific variation of its Lesser Antillean relatives. However, this predictability may be less consistent with other character systems, for example, scalation.

Integrating gastrocnemius force-length properties, in vivo activation and operating lengths reveals how Anolis deal with ecological challenges.

A central question in biology is how animals successfully behave under complex natural conditions. Although changes in locomotor behaviour, motor control and force production in relation to incline are commonly examined, a wide range of other factors, including a range of perch diameters, pervades arboreal habitats. Moving on different substrate diameters requires considerable alteration of body and limb posture, probably causing significant shifts in the lengths of the muscle-tendon units powering locomotion. Thus, how substrate shape impacts in vivo muscle function remains an important but neglected question in ecophysiology. Here, we used high-speed videography, electromyography, in situ contractile experiments and morphology to examine gastrocnemius muscle function during arboreal locomotion in the Cuban knight anole, Anolis equestris The gastrocnemius contributes more to the propulsive effort on broad surfaces than on narrow surfaces. Surprisingly, substrate inclination affected the relationship between the maximum potential force and fibre recruitment; the trade-off that was present between these variables on horizontal surfaces became a positive relationship on inclined surfaces. Finally, the biarticular nature of the gastrocnemius allows it to generate force isometrically, regardless of substrate diameter and incline, despite the fact that the tendons are incapable of stretching during cyclical locomotion. Our results emphasize the importance of considering ecology and muscle function together, and the necessity of examining both mechanical and physiological properties of muscles to understand how animals move in their environment.

Rates of oxygen uptake increase independently of changes in heart rate in late stages of development and at hatching in the green iguana, Iguana iguana.

Oxygen consumption (VO2), heart rate (fH), heart mass (Mh) and body mass (Mb) were measured during embryonic incubation and in hatchlings of green iguana (Iguana iguana). Mean fH and VO2 were unvarying in early stage embryos. VO2 increased exponentially during the later stages of embryonic development, doubling by the end of incubation, while fH was constant, resulting in a 2.7-fold increase in oxygen pulse. Compared to late stage embryos, the mean inactive level of VO2 in hatchlings was 1.7 fold higher, while fH was reduced by half resulting in a further 3.6 fold increase in oxygen pulse. There was an overall negative correlation between mean fH and VO2 when data from hatchlings was included. Thus, predicting metabolic rate as VO2 from measurements of fH is not possible in embryonic reptiles. Convective transport of oxygen to supply metabolism during embryonic incubation was more reliably indicated as an index of cardiac output (COi) derived from the product of fH and Mh. However, a thorough analysis of factors determining rates of oxygen supply during development and eclosion in reptiles will require cannulation of blood vessels that proved impossible in the present study, to determine oxygen carrying capacity by the blood and arteriovenous oxygen content difference (A-V diff), plus patterns of blood flow.

The effects of multiple obstacles on the locomotor behavior and performance of a terrestrial lizard.

Negotiation of variable terrain is important for many small terrestrial vertebrates. Variation in the running surface resulting from obstacles (woody debris, vegetation, rocks) can alter escape paths and running performance. The ability to navigate obstacles likely influences survivorship through predator evasion success and other key ecological tasks (finding mates, acquiring food). Earlier work established that running posture and sprint performance are altered when organisms face an obstacle, and yet studies involving multiple obstacles are limited. Indeed, some habitats are cluttered with obstacles, whereas others are not. For many species, obstacle density may be important in predator escape and/or colonization potential by conspecifics. This study examines how multiple obstacles influence running behavior and locomotor posture in lizards. We predict that an increasing number of obstacles will increase the frequency of pausing and decrease sprint velocity. Furthermore, bipedal running over multiple obstacles is predicted to maintain greater mean sprint velocity compared with quadrupedal running, thereby revealing a potential advantage of bipedalism. Lizards were filmed running through a racetrack with zero, one or two obstacles. Bipedal running posture over one obstacle was significantly faster than quadrupedal posture. Bipedal running trials contained fewer total strides than quadrupedal ones. But on addition of a second obstacle, the number of bipedal strides decreased. Increasing obstacle number led to slower and more intermittent locomotion. Bipedalism provided clear advantages for one obstacle, but was not associated with further benefits for an additional obstacle. Hence, bipedalism helps mitigate obstacle negotiation, but not when numerous obstacles are encountered in succession.

Galápagos land iguana (Conolophus subcristatus) as a seed disperser.

The role of the most common land iguana (Conolophus subcristatus) in the Galápagos Islands as an effective seed disperser is explored in this study. A total of 5705 seeds of 32 plant species were identified from 160 scats, 4545 of which (80%) appeared visually undamaged. Germination trials of 849 seeds from 29 species revealed that at least 10 species remained viable after passing through the iguana's gut, although only a small proportion of those seeds (4%) germinated. In any case, we argue that C. subcristatus exerts an important role on the 7 Galapagos islands where it occurs because of its abundance and capacity to ingest and disperse seeds at long distances. Our results strongly suggest that the Galápagos C. subcristatus plays an important role as a seed disperser of not only of native species but also some introduced plants in the Galápagos Islands.

XROMM analysis of rib kinematics during lung ventilation in the green iguana, Iguana iguana.

The three-dimensional rotations of ribs during breathing are typically described as bucket-handle rotation about a dorsoventrally oriented axis, pump-handle rotation about a mediolateral axis, and caliper rotation about a rostrocaudal axis. In amniotes with double-headed ribs, rib motion is constrained primarily to one degree-of-freedom (DOF) rotation about an axis connecting the two rib articulations. However, in Squamata, the ribs are single headed and the hemispherical costovertebral joints permit rotations with three DOF. In this study, we used X-ray reconstruction of moving morphology (XROMM ) to quantify rib rotation during deep breathing in four green iguanas. We found that rib rotation was strongly dominated by bucket-handle rotation, thus exhibiting nearly hinge-like motion, despite the potential for more complex motions. The vertebral and sternal segments of each rib did not deform measurably during breathing, but they did move relative to each other at a thin, cartilaginous intracostal joint. While standing still and breathing deeply, four individual iguanas showed variability in their rib postures, with two breathing around a highly inflated posture, and two breathing around a posture with the ribs folded halfway back. Bucket-handle rotations showed clear rostrocaudal gradients, with rotation increasing from the third cervical to the first or second dorsal rib, and then decreasing again caudally, a pattern that is consistent with the intercostal muscles in the rostral intercostal spaces being the primary drivers of inspiration. The constrained, primarily bucket-handle rotations observed here during breathing do not help to explain the evolution of permissive, hemispherical costovertebral joints in squamates from the more constrained, double-headed rib articulations of other amniotes.