Reptile responses to fire and the risk of post-disturbance sampling bias

Altered fire regimes are a driver of biodiversity decline. To plan effective management, we need to know how species are influenced by fire and to develop theory describing fire responses. Animal responses to fire are usually measured using methods that rely on animal activity, but animal activity may vary with time since fire, potentially biasing results. Using a novel approach for detecting bias in the pit-fall trap method, we found that leaf-litter dependent reptiles were more active up to 6 weeks after fire, giving a misleading impression of abundance. This effect was not discovered when modelling detectability with zero-inflated binomial models. Two species without detection bias showed early-successional responses to time since fire, consistent with a habitat-accommodation succession model. However, a habitat specialist did not have the predicted low abundance after fire due to increased post-fire movement and non-linear recovery of a key habitat component. Interactions between fire and other processes therefore must be better understood to predict reptile responses to changing fire-regimes. We conclude that there is substantial bias when trapping reptiles after fire, with species that are otherwise hard to detect appearing to be abundant. Studies that use a survey method based on animal activity such as bird calls or animal movements, likely face a similar risk of bias when comparing recently-disturbed with control sites.

Why do some species have geographically varying responses to fire history?

A capacity to predict the effects of fire on biota is critical for conservation in fire-prone regions as it assists managers to anticipate the outcomes of different approaches to fire management. The task is complicated because species' responses to fire can vary geographically. This poses challenges, both for conceptual understanding of post-fire succession and fire management. We examine two hypotheses for why species may display geographically varying responses to fire. 1) Species' post-fire responses are driven by vegetation structure, but vegetation - fire relationships vary spatially (the 'dynamic vegetation' hypothesis). 2) Regional variation in ecological conditions leads species to select different post-fire ages as habitat (the 'dynamic habitat' hypothesis). Our case study uses data on lizards at 280 sites in a ~ 100 000 km2 region of south-eastern Australia. We compared the predictive capacity of models based on 1) habitat associations, with models based on 2) fire history and vegetation type, and 3) fire history alone, for four species of lizards. Habitat association models generally out-performed fire history models in terms of predictive capacity. For two species, habitat association models provided good discrimination capacity even though the species showed geographically varying post-fire responses. Our results support the dynamic vegetation hypothesis, that spatial variation in relationships between fire and vegetation structure results in regional variation in fauna-fire relationships. These observations explain how the widely recognised 'habitat accommodation' model of animal succession can be conceptually accurate yet predictively weak. © 2014 The Authors.

Successional specialization in a reptile community cautions against widespread planned burning and complete fire suppression

Summary: Conservation of biodiversity in fire-prone regions depends on understanding responses to fire in animal communities and the mechanisms governing these responses. We collated data from an Australian semi-arid woodland reptile community (4796 individuals captured over 6 years) to: (i) determine the ability of commonly used shorter-term (2 years) surveys to detect reptile responses to time since fire (TSF) and (ii) investigate whether ecological traits of species reliably predicted their responses to fire. Of 16 reptile species analysed, four had responses to TSF consistent with shorter-term surveys and three showed no response to TSF. Nine species had responses to TSF not detected in previous studies using smaller but substantial subsets of the same data. Among the 13 affected species, times of peak abundance ranged from 1 to 50 years after fire. Nocturnal, burrowing species tended to be early successional and leaf-litter dwellers to be late successional, but these were only weak trends. Synthesis and applications. We found only limited support for a generalizable, trait-based model of succession in reptiles. However, our study revealed that the majority of common reptile species in our study region specialize on a post-fire successional stage and may therefore become threatened if homogeneous fire regimes predominate. Our study highlights the importance of interpreting results from time- or sample-limited fire studies of reptiles with the knowledge that many ecological responses may not have been detected. In such cases, an adaptive or precautionary approach to fire management may be necessary. We found only limited support for a generalizable, trait-based model of succession in reptiles. However, our study revealed that the majority of common reptile species in our study region specialize on a post-fire successional stage and may therefore become threatened if homogeneous fire regimes predominate. Our study highlights the importance of interpreting results from time- or sample-limited fire studies of reptiles with the knowledge that many ecological responses may not have been detected. In such cases, an adaptive or precautionary approach to fire management may be necessary.

Small reptile community responses to rotational logging

Timber harvesting is a common global disturbance that has important effects on the ability of forests to provide ecosystems services and retain biodiversity. Using predictive frameworks to examine biodiversity responses to logging could assist in retaining natural forest values. The intermediate disturbance hypothesis (IDH) and the habitat accommodation model (HAM) potentially offer frameworks for explaining different coarse scale community responses to logging. We used a 60. year post-logging chronosequence to investigate small reptile community responses to age post-logging in temperate forests using three metrics (species richness, evenness and relative abundance). First, we evaluated if variation in these metrics adhered to prior predictions, including the IDH. Second, we evaluated how age post-logging influence community responses through fine scale vegetation elements. Third, we evaluated support for the HAM by measuring compositional change (species turnover) of small reptile community to age post-logging. Reptile relative abundance exhibited a curvilinear relationship to age since logging, contradicting our prior prediction of sustained increase. Species richness and evenness were unrelated to age since logging thus providing no support to IDH and other prior predictions. Relative abundance and richness did not relate to any vegetation characteristic tested. These metrics were also unrelated to logging method. Community composition was marginally significantly influenced by age since logging, thus supporting the HAM. Our results suggest that forest reptiles exposed to logging exhibit variable changes depending on the community metric in question, and that different approaches, including those based on species traits, are needed to improve evaluating disturbance related biodiversity responses.

Integration of autonomic and local mechanisms in regulating cardiovascular responses to heating and cooling in a reptile (Crocodylus porosus)

Reptiles change heart rate and blood flow patterns in response to heating and cooling, thereby decreasing the behavioural cost of thermoregulation. We tested the hypothesis that locally produced vasoactive substances, nitric oxide and prostaglandins, mediate the cardiovascular response of reptiles to heat. Heart rate and blood pressure were measured in eight crocodiles (Crocodylus porosus) during heating and cooling and while sequentially inhibiting nitric-oxide synthase and cyclooxygenase enzymes. Heart rate and blood pressure were significantly higher during heating than during cooling in all treatments. Power spectral density of heart rate and blood pressure increased significantly during heating and cooling compared to the preceding period of thermal equilibrium. Spectral density of heart rate in the high frequency band (0.19-0.70 Hz) was significantly greater during cooling in the saline treatment compared to when nitric-oxide synthase and cyclooxygenase enzymes were inhibited. Cross spectral analysis showed that changes in blood pressure preceded heart rate changes at low frequencies (

Mechanisms of the impact of a weed (grader grass, Themeda quadrivalvis) on reptile assemblage structure in a tropical savannah

Invasive grasses are among the worst threats to native biodiversity, but the mechanisms causing negative effects are poorly understood. To investigate the impact of an invasive grass on reptiles, we compared the reptile assemblages that used native kangaroo grass (Themeda triandra), and black spear grass (Heteropogon contortus), to those using habitats invaded by grader grass (Themeda quadrivalvis). There were significantly more reptile species, in greater abundances, in native kangaroo and black spear grass than in invasive grader grass. To understand the sources of negative responses of reptile assemblages to the weed, we compared habitat characteristics, temperatures within grass clumps, food availability and predator abundance among these three grass habitats. Environmental temperatures in grass, invertebrate food availability, and avian predator abundances did not differ among the habitats, and there were fewer reptiles that fed on other reptiles in the invaded than in the native grass sites. Thus, native grass sites did not provide better available thermal environments within the grass, food, or opportunities for predator avoidance. We suggest that habitat structure was the critical factor driving weed avoidance by reptiles in this system, and recommend that the maintenance of heterogeneous habitat structure, including clumping native grasses, with interspersed bare ground, and leaf litter are critical to reptile biodiversity.

Mechanisms of the impact of a weed (grader grass, Themeda quadrivalvis) on reptile assemblage structure in a tropical savannah

Invasive grasses are among the worst threats to native biodiversity, but the mechanisms causing negative effects are poorly understood. To investigate the impact of an invasive grass on reptiles, we compared the reptile assemblages that used native kangaroo grass (Themeda triandra), and black spear grass (Heteropogon contortus), to those using habitats invaded by grader grass (Themeda quadrivalvis). There were significantly more reptile species, in greater abundances, in native kangaroo and black spear grass than in invasive grader grass. To understand the sources of negative responses of reptile assemblages to the weed, we compared habitat characteristics, temperatures within grass clumps, food availability and predator abundance among these three grass habitats. Environmental temperatures in grass, invertebrate food availability, and avian predator abundances did not differ among the habitats, and there were fewer reptiles that fed on other reptiles in the invaded than in the native grass sites. Thus, native grass sites did not provide better available thermal environments within the grass, food, or opportunities for predator avoidance. We suggest that habitat structure was the critical factor driving weed avoidance by reptiles in this system, and recommend that the maintenance of heterogeneous habitat structure, including clumping native grasses, with interspersed bare ground, and leaf litter are critical to reptile biodiversity.

Evaluating the role of fire disturbance in structuring small reptile communities in temperate forests

Fire is an integral disturbance shaping forest community dynamics over large scales. However, understanding the relationship between fire induced habitat disturbance and biodiversity remain equivocal. Ecological theories including the intermediate disturbance hypothesis (IDH) and the habitat accommodation model (HAM) offer predictive frameworks that could explain faunal responses to fire disturbances. We used an 80 year post-fire chronosequence to investigate small reptile community responses to fires in temperate forests across 74 sites. First, we evaluated if changes in species richness, abundance and evenness post-fire followed trends of prior predictions, including the IDH. Second, using competing models of fine scale habitat elements we evaluated the specific ways which fire influenced small reptiles. Third, we evaluated support for the HAM by examining compositional changes of reptile community post-fire. Relative abundance was positively correlated to age post-fire while richness and evenness showed no associations. The abundance trend was as expected based on the prior prediction of sustained population increase post-disturbance, but the trend for richness contradicted the prediction of highest diversity at intermediate levels of disturbance (according to IDH). Abundance changes were driven mainly by changes in overstorey, ground layer, and shelter, while richness and evenness did not associate with any vegetation parameter. Community composition was not strongly correlated to age since fire, thus support for the HAM was weak. Overall, in this ecosystem, frequent fire disturbances can be detrimental to small reptiles. Future studies utilizing approaches based on species traits could enhance our understanding of biodiversity patterns post-disturbance.

Loss of the Ability to Control Right-to-Left Shunt Does Not Influence the Metabolic Responses to Temperature Change or Long-Term Fasting in the South American Rattlesnake Crotalus durissus

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Prostaglandins are important in thermoregulation of a reptile (Pogona vitticeps)

The effectiveness of behavioural thermoregulation in reptiles is amplified by cardiovascular responses, particularly by differential rates of heart beat in response to heating and cooling (heart-rate hysteresis). Heart-rate hysteresis is ecologically important in most lineages of ectothermic reptile' and we demonstrate that heart-rate hysteresis in the lizard Pogona vitticeps is mediated by prostaglandins. In a control treatment (administration of saline), heart rates during heating were significantly faster than during cooling at any given body temperature. When cyclooxygenase 1 and 2 enzymes were inhibited, heart rates during heating were not significantly different from those during cooling. Administration of agonists showed that thromboxane B-2 did not have a significant effect on heart rate, but prostacyclin and prostaglandin F-2alpha caused a significant increase (3.5 and 13.6 beats min(-1), respectively) in heart rate compared with control treatments. We speculate that heart-rate hysteresis evolved as a thermoregulatory mechanism that may ultimately be controlled by neurally induced stimulation of nitric oxide production, or maybe via photolytically induced production of vitamin D.

Diving behaviour of a reptile (Crocodylus johnstoni) in the wild: Interactions with heart rate and body temperature

The differences in physical properties of air and water pose unique behavioural and physiological demands on semiaquatic animals. The aim of this study was to describe the diving behaviour of the freshwater crocodile Crocodylus johnstoni in the wild and to assess the relationships between diving, body temperature, and heart rate. Time-depth recorders, temperature-sensitive radio transmitters, and heart rate transmitters were deployed on each of six C. johnstoni (4.0-26.5 kg), and data were obtained from five animals. Crocodiles showed the greatest diving activity in the morning (0600-1200 hours) and were least active at night, remaining at the water surface. Surprisingly, activity pattern was asynchronous with thermoregulation, and activity was correlated to light rather than to body temperature. Nonetheless, crocodiles thermoregulated and showed a typical heart rate hysteresis pattern (heart rate during heating greater than heart rate during cooling) in response to heating and cooling. Additionally, dive length decreased with increasing body temperature. Maximum diving length was 119.6 min, but the greatest proportion of diving time was spent on relatively short (

Response of a shrubland mammal and reptile community to a history of landscape-scale wildfire

Fire plays a strong role in structuring fauna communities and the habitat available to them in fire-prone regions. Human-mediated increases in fire frequency and intensity threaten many animal species and understanding how these species respond to fire history and its associated effect on vegetation is essential to effective biodiversity management. We used a shrubland mammal and reptile community in semiarid south-western Australia as a model to investigate interactions between fire history, habitat structure and fauna habitat use. Of the 15 species analysed, five were most abundant in recently burnt habitat (8–13 years since last fire), four were most abundant in long unburnt areas (25–50 years) and six showed no response to fire history. Fauna responses to fire history were divergent both within and across taxonomic groups. Fire management that homogenises large areas of habitat through either fire exclusion or frequent burning may threaten species due to these diverse requirements, so careful management of fire may be needed to maximise habitat suitability across the landscape. When establishing fire management plans, we recommend that land managers exercise caution in adopting species-specific information from different locations and broad vegetation types. Information on animal responses to fire is best gained through experimental and adaptive management approaches at the local level.

Lizard responses to forest fire and timber harvesting: complementary insights from species and community approaches

Understanding the relationship between community composition and ecosystem function is essential for managing forests with complex disturbance regimes. Studies of animal responses to fire and timber harvesting in forest ecosystems typically focus on a single level of community diversity. Measures of species abundance and diversity at the community level, along with measures of functional diversity that incorporate information on species traits, provide opportunities for complementary insights into biodiversity responses to disturbances. We quantified community and functional responses of a temperate forest lizard community to fire and rotational logging using metrics including species-specific abundance, community abundance, species richness and evenness, as well as trait-based measures of functional diversity. We used non-linear regression models to examine the relationships between reptile data and time since fire and timber harvesting, using sites arrayed along a 30-years post-disturbance chronosequence. We modelled responses separately in two major vegetation types: coastal Banksia woodland and lowland eucalypt forests. Species and community measures offered different insights into the role of fire and logging. Species responses to disturbance differed between disturbance type and vegetation type. Four species exhibited significant population responses to either fire or timber harvesting, while the rest were unaffected by either disturbance. At the community level, species richness and community abundance increased significantly with time since fire in woodland vegetation. In forest vegetation, community abundance decreased with time since fire. Surprisingly, community evenness and functional diversity did not show marked responses to fire or timber harvesting. This is likely a result of trait homogeneity and the asynchrony in species responses to disturbance. We advocate using multiple measures of community composition - incorporating species-specific information, community metrics and functional traits - to ensure a more holistic understanding of disturbance ecology in forest landscapes.

Ecosystem responses to fire : identifying cross-taxa contrasts and complementarities to inform management strategies

Changes in fire frequency, extent, and intensity mean that understanding the effects of fire on plants and animals is a primary concern for ecologists and land managers. Given the potentially conflicting fire responses of species both within and across taxonomic groups, prescribing fire regimes based on the response of one or only a few species may have negative consequences for other species. Here, we integrate data collected from a series of independent but complementary studies spanning a 75 + year chronosequence in a semi-arid shrubland ecosystem in south-western Australia to consider how fire management can best promote biodiversity both within and across taxonomic groups (plants, birds, small mammals, and reptiles). Younger fire ages (6–14 years) contained sparse shrubs, large areas of bare ground, and lacked a distinct litter layer and canopy. The oldest vegetation (60–85 years) had a distinct canopy, a well-developed litter layer and cryptogamic crust, higher variability in patch width, and more woody debris. Plant species richness and diversity decreased with time since fire, whereas bird species richness and diversity increased with time since fire, and mammal and reptile species richness and diversity showed no trend. The composition of all four taxonomic groups varied according to time since fire and the presence of 11 species was confined above or below certain fire-age thresholds. Our results support the need to maintain a mix of both younger and older fire ages across the landscape to maximise species diversity, and highlight the particular importance of older fire ages for many species. Future fire management for biodiversity conservation will benefit from identifying and reconciling cross-taxa contrasts and complementarities.

Physiological and symptomatic responses to cycling and walking in intermittent claudication

To shed light on the potential efficacy of cycling as a testing modality in the treatment of intermittent claudication (IC), this study compared physiological and symptomatic responses to graded walking and cycling tests in claudicants. Sixteen subjects with peripheral arterial disease (resting ankle: brachial index (ABI) < 0.9) and IC completed a maximal graded treadmill walking (T) and cycle (C) test after three familiarization tests on each mode. During each test, symptoms, oxygen uptake (VO2), minute ventilation (VE), respiratory exchange ratio (RER) and heart rate (HR) were measured, and for 10 min after each test the brachial and ankle systolic pressures were recorded. All but one subject experienced calf pain as the primary limiting symptom during T; whereas the symptoms were more varied during C and included thigh pain, calf pain and dyspnoea. Although maximal exercise time was significantly longer on C than T (690 +/- 67 vs. 495 +/- 57 s), peak VO2, peak VE and peak heart rate during C and T were not different; whereas peak RER was higher during C. These responses during C and T were also positively correlated (P < 0.05) with each other, with the exception of RER. The postexercise systolic pressures were also not different between C and T. However, the peak decline in ankle pressures from resting values after C and T were not correlated with each other. These data demonstrate that cycling and walking induce a similar level of metabolic and cardiovascular strain, but that the primary limiting symptoms and haemodynamic response in an individual's extremity, measured after exercise, can differ substantially between these two modes.