Improving supplementary feeding in species conservation

Supplementary feeding is often a knee‐jerk reaction to population declines, and its application is not critically evaluated, leading to polarized views among managers on its usefulness. Here, we advocate a more strategic approach to supplementary feeding so that the choice to use it is clearly justified over, or in combination with, other management actions and the predicted consequences are then critically assessed following implementation. We propose combining methods from a set of specialist disciplines that will allow critical evaluation of the need, benefit, and risks of food supplementation. Through the use of nutritional ecology, population ecology, and structured decision making, conservation managers can make better choices about what and how to feed by estimating consequences on population recovery across a range of possible actions. This structured approach also informs targeted monitoring and more clearly allows supplementary feeding to be integrated in recovery plans and reduces the risk of inefficient decisions. In New Zealand, managers of the endangered Hihi (Notiomystis cincta) often rely on supplementary feeding to support reintroduced populations. On Kapiti island the reintroduced Hihi population has responded well to food supplementation, but the logistics of providing an increasing demand recently outstretched management capacity. To decide whether and how the feeding regime should be revised, managers used a structured decision making approach informed by population responses to alternative feeding regimes. The decision was made to reduce the spatial distribution of feeders and invest saved time in increasing volume of food delivered into a smaller core area. The approach used allowed a transparent and defendable management decision in regard to supplementary feeding, reflecting the multiple objectives of managers and their priorities.     

The moral residue of conservation

Should conservationists use lethal management to control introduced wildlife populations? Should they kill individual animals to protect endangered species? Are trade-offs that prioritize some values at the expense of others morally appropriate? These sorts of ethical questions are common in conservation. In debating such questions, conservationists often seem to presume 1 of 2 possible answers: the act in question is right or it is wrong. But morality in conservation is considerably more complex than this simple binary suggests. A robust conservation ethic requires a vocabulary that gives voice to the uncertainty and unease that arise when what seems to be the best available course of action also seems to involve a measure of wrongdoing. The philosophical literature on moral residue and moral dilemmas supplies this vocabulary. Moral dilemmas arise when one must neglect certain moral requirements to fulfill others. Under such circumstances, even the best possible decision leaves a moral residue, which is experienced emotionally as some form of grief. Examples of conservation scenarios that leave a moral residue include management of introduced rabbits in Australia, trophy hunting in Africa, and forest management trade-offs in the Pacific Northwest. Moral residue is integral to the moral experience of conservationists today, and grief is an appropriate response to many decisions conservationists must make. Article impact statement: Defensible conservation decisions may neglect moral requirements, leaving a moral residue; conservationists should respond with grief.     

Inbreeding and Loss of Genetic Variation in a Reintroduced Population of Mauritius Kestrel

Abstract:  Many populations have recovered from severe bottlenecks either naturally or through intensive conservation management. In the past, however, few conservation programs have monitored the genetic health of recovering populations. We conducted a conservation genetic assessment of a small, reintroduced population of Mauritius Kestrel ( Falco punctatus ) to determine whether genetic deterioration has occurred since its reintroduction. We used pedigree analysis that partially accounted for individuals of unknown origin to document that (1) inbreeding occurred frequently (2.6% increase per generation; N _eI= 18.9), (2) 25% of breeding pairs were composed of either closely or moderately related individuals, (3) genetic diversity has been lost from the population (1.6% loss per generation; N _eV= 32.1) less rapidly than the corresponding increase in inbreeding, and (4) ignoring the contribution of unknown individuals to a pedigree will bias the metrics derived from that pedigree, ultimately obscuring the prevailing genetic dynamics. The rates of inbreeding and loss of genetic variation in the subpopulation of Mauritius Kestrel we examined were extreme and among the highest yet documented in a wild vertebrate population. Thus, genetic deterioration may affect this population's long-term viability. Remedial conservation strategies are needed to reduce the impact of inbreeding and loss of genetic variation in this species. We suggest that schemes to monitor genetic variation after reintroduction should be an integral component of endangered species recovery programs.     

Spatiotemporal Patterns and Risk Factors for Lead Exposure in Endangered California Condors during 15 Years of Reintroduction

Large‐scale poisoning events are common to scavenging bird species that forage communally, many of which are in decline. To reduce the threat of poisoning and compensate for other persistent threats, management, including supplemental feeding, is ongoing for many reintroduced and endangered vulture populations. Through a longitudinal study of lead exposure in California condors (Gymnogyps californianus), we illustrate the conservation challenges inherent in reintroduction of an endangered species to the wild when pervasive threats have not been eliminated. We evaluated population‐wide patterns in blood lead levels from 1997 to 2011 and assessed a broad range of putative demographic, behavioral, and environmental risk factors for elevated lead exposure among reintroduced California condors in California (United States). We also assessed the effectiveness of lead ammunition regulations within the condor's range in California by comparing condor blood lead levels before and after implementation of the regulations. Lead exposure was a pervasive threat to California condors despite recent regulations limiting lead ammunition use. In addition, condor lead levels significantly increased as age and independence from intensive management increased, including increasing time spent away from managed release sites, and decreasing reliance on food provisions. Greater independence among an increasing number of reintroduced condors has therefore elevated the population's risk of lead exposure and limited the effectiveness of lead reduction efforts to date. Our findings highlight the challenges of restoring endangered vulture populations as they mature and become less reliant on management actions necessary to compensate for persistent threats. Patrones Espaciotemporales y Factores de Riesgo por Exposición a Plomo en Cóndores de California Durante 15 Años de Reintroducción     

Whooping Crane Recovery: A Case Study in Public and Private Cooperation in the Conservation of Endangered Species

I describe three theoretical approaches to endangered species conservation: public-sector models, private-sector models, and mixed models. Criteria to consider in evaluating these models are scientific, economic, legal, ethical, and administrative. The history of conservation efforts on behalf of the endangered Whooping Crane ( Grus americana ) is used as an example of a mixed-model approach (one that involves both public- and private-sector organizations and individuals) to the conservation of endangered species. Evaluation of Whooping Crane conservation efforts, using the specified criteria, suggests that this mixed model gets relatively high marks in all areas. My recommendations for endangered species conservation activities include (1) ensuring that all potential sources of scientific expertise and "caring" are included, (2) developing a robust national funding mechanism for endangered species conservation efforts, and (3) creating streamlined administrative structures and operating procedures tailored to the unique needs of each endangered species conservation team. Broader implications of the Whooping Crane story for conservation biology as a profession and for conservation biologists as individuals include (1) being inclusive rather than exclusive in conservation planning and implementation, (2) recognizing the power of individual initiative and commitment, (3) collecting basic data over an extended period of time, and (4) emphasizing the importance of communication, cooperation, and coordination.     

Diel vertical migrations and nocturnal feeding of a dense coastal krill scattering layer (Thysanoessa raschi and Meganyctiphanes norvegica) in stratified surface waters

To study the nocturnal feeding of euphausiids during vertical migrations and its impact on the phytoplankton, a phytoplankton-rich water mass (drogue marked) drifting over a dense krill scattering layer (acoustic 104kHz) in the lower St. Lawrence estuary was monitored over 46 h in July 1982. Phytoplankton >20 m was abundant and mostly concentrated at the bottom of the photic layer above the pycnocline. Less than 42% of the particulate carbon was due to phytoplankton. The krill scattering layer was about 2 to 3 km in width, elongated along the 100-m bathymetric contour, and absent when the bottom was shallower than 50 m. Its upper day depth was 50 m. At deeper depths, its vertical distribution frequently changed from unimodal to polymodal shapes and viceversa, often with large concentrations of zooplankton just above the bottom. Typical vertical migrations were observed on both days. At night the scattering layer had a lower scattering strength. Most of it was below the thermocline but net catches showed that large concentrations of euphausiids (up to 57 individuals m^-3) crossed it. Stomach pigment content of Thysanoessa raschi was generally low, but mean stomach fullness was always high. They were more opportunistic than herbivorous. From stomach fullness and the presence of a food bolus in mouth parts, feeding in surface waters appeared to be intensive, but gut content indicated that food was not processed there. It is therefore suggested that individuals underwent vertical interchanges across the thermocline while feeding during the night. Meganyctiphanes norvegica had significant herbivorous activity during the night. The grazing pressure impact of the scattering layer on phytoplankton was negligible.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Ecological versatility and the decline of coral feeding fishes following climate driven coral mortality

Coral reefs are under threat due to climate-mediated coral mortality, which affects some reef coral genera more severely than others. The impact this has on coral reef fish is receiving increasing attention, with one focal area assessing impacts on fish that feed directly on live coral. It appears that the more specialised a species of corallivore, the more susceptible it is to coral declines. However data are sparse for the Indian Ocean, and little is known about why some corals are preferentially fed upon over others. Here I assess feeding specialisation in three species of coral feeding butterflyfish in the Chagos Archipelago, central Indian Ocean, assess the food quality of the coral genera they target and document patterns of decline in the Seychelles following a severe coral mortality event. Cheatodon trifascialis was the most specialised coral feeder, preferentially selecting for Acropora corals, however, when Acropora was scarce, individuals showed considerable feeding plasticity, particularly for the dominant Pocillopora corals. C. trifasciatus also preferentially fed on Acropora corals, but fed on a much more diverse suite of corals and also displayed some selectivity for Porites. C. auriga is a facultative corallivore and consumed ∼55% live coral, which lies within the wide range of coral dependence reported for this species. C:N ratio analysis indicated Lobophyllia and Acropora have the highest food quality, with Pocillopora having the lowest, which conforms with diet selection of corallivores and helps explain preferential feeding. Obligate specialist feeders displayed the greatest declines through coral mortality in the Seychelles with obligate generalists also declining substantially, but facultative feeders showing little change. Clearly a greater understanding of the species most vulnerable to disturbance, their habitat requirements and the functional roles they play will greatly assist biodiversity conservation in a changing climate.     

Intake and coversion of food in the fish Limanda limanda exposed to different temperatures

In the flat fish Limanda limanda L., feeding rate and conversion efficiency were studied as functions of body weight, sex, temperature and food quality. When offered herring meat at 13 °C (series I), females (live weights 1 to 150 g) consume more food than males; the magnitude of this difference is body weight-dependent. With increasing wieght, both females and males consume less food per unit body weight per day. Variations in daily ration are considerable; the range of deviation from mean feeding rate is about 60% for males and 40% for females. The range of deviation does not vary significantly among females and males of different body weights. At the same temperature level (13 °C; series II), females consume almost the same, or even less, cod meat than males. Among individuals of series I and II, there is a little difference in the feeding rate; however, herring-fed individuals obtain about 2 times more energy than cod-fed individuals. Each gram wet weight of herring meat yields 2001, each gram cod meat 1137, calories. Small individuals completely cease to feed at 3°C; they feed little at 8 °C. Larger females consume maximum amounts at 8 °C. Small individuals consume maximum amounts at higher temperatures. Thus, with increasing body weight (age), the temperature for maximum feeding shifts downwards. Feeding with cod or herring meat results in considerable changes in composition and calorific content of L. Limanda. The magnitude of these changes depends both on temperature and food quality. Food conversion efficiency values of herring-fed individuals are about 1 1/2 times higher than of cod-fed individuals. In series I and II, females are more efficient converters than males. In individuals weighing more than 50 g, conversion efficiency decreases in the order: 8°, 13°, 18° C; in smaller individuals this order is 13°, 18°, 8 °C. Conversion rate is about 2 to 5 times faster in individuals fed herring meat than those receiving cod meat. Conversion rate decreases in the order 13°, 8°, 18 °C in males, and in the order 18°, 13°, 8 °C in females; females of more than 80 g are exceptional in that they reach the maximum at 8 °C. From the data on food intake and food conversion, the biologically useful energy available for metabolism has been calculated for each test individual kept at 13° and 18 °C. At these temperature levels, the weight exponents are about 0.6; the a value or metabolic level for the 18 °C series is about 2 times higher than that at 13 °C. Thus, temperature affects metabolic rate but not the exponential value. The exponential value for the body weight-metabolism relation at 13 °C is for dab fed herring meat 0.9; the a value amounts to about half that for dab fed cod meat. Food quality, unlike temperature, alters not only the exponential value but also metabolic rate.     

Energetic state and the performance of dawn chorus in silvereyes (Zosterops lateralis)

Stochastic dynamic programming (SDP) models predict that males singing to attract a mate should concentrate singing in what has been termed the dawn chorus. This is because male birds should have a variable surplus of fat in the morning that can be used to fuel singing, with the amount of fat available dependent upon such factors as his quality, foraging success and risk of predation. In this manner, the dawn chorus can act as an indicator of male quality in the context of female mate choice. We test a key prediction of SDP models of singing behaviour that males with greater fat levels should sing more. We conducted an experiment where we recorded the dawn chorus of male silvereyes (Zosterops lateralis) on three consecutive days. Each male received supplementary food on the second day, which enabled us to sample his dawn chorus before, during and after food supplementation. We also collected data on the effect of supplementary food on the body mass of silvereyes. As predicted by SDP models, we found that silvereyes sang for a greater proportion of the time after receiving supplementary food. Supplementary food also had a significant effect on the complexity of a male song, indicating that males not only increased the quantity of their song but also the quality of their song when they received extra food. As the provision of supplementary food significantly increased the mass of fed birds, our results support a causal link between male energy reserves and his ability to perform the dawn chorus.     

Transformation of food in the fish Megalops cyprinoides

Comparative studies on feeding, digestion, absorption and conversion have been made in the fish Megalops cyprinoides fed with mosquito fish, Gambusia affinis, and prawn, Metapenaeus monoceros, (Pandian, 1967). Feeding as well as digestion rates are faster in individuals fed with Gambusia than in those fed with prawn. The extent of the differences in food intake is more pronounced in smaller individuals than in larger ones. The prawn exoskeleton prolongs the duration of digestion and thus further reduces rate and amount of food intake. Changes in food quality do not affect absorption efficiency. Both, rate and efficiency of food conversion, are higher in M. cyprinoides fed with Gambusia than with prawn.     

Diagnosing Units of Conservation Management

Species-oriented conservation programs attempt to analyze and maintain intra-specific variation in order to maximally preserve biological diversity. The "evolutionarily significant unit" has become an operational term for a group of organisms that should be the minimial unit for conservation management. No generally accepted definition for this term exists that would be the basis for the evaluation of these units in practical conservation situations. Currently, taxonomic decisions in species conservation are mostly based on the biological species concept. But the universal application of criteria of reproductive isolation or phenetic similarity to delimit conservation units is problematical. We favor a definition for evolutionarily significant units based on patterns of variation. In the theoretical framework of the phylogenetic species concept, conservation units are delimited by characters that diagnose clusters of individuals or populations to the exclusion of other such clusters. Characters are used for cladistic analysis to infer hypotheses of the phylogenetic relationships of individuals, and differentiated populations are diagnosed using population aggregation analysis. Characters can be based on genetic, morphological, ecological, or behavioral information, provided they are inferred to be heritable. The use of cladistics and population aggregation analysis has the potential to make the evaluation of evoluntionarily significant units objective and testable, an important consideration in politically controversial cases. Our cladistic approach is demonstrated by the evaluation of potential conservation units in the endangered tiger beetles Cicindela dorsalis and C. puritana .     

The Conservation Relevance of Epidemiological Research into Carnivore Viral Diseases in the Serengeti

Abstract:  Recent outbreaks of rabies and canine distemper in wildlife populations of the Serengeti show that infectious disease constitutes a significant cause of mortality that can result in regional extirpation of endangered species even within large, well-protected areas. Nevertheless, effective management of an infectious disease depends critically on understanding the epidemiological dynamics of the causative pathogen. Pathogens with short infection cycles cannot persist in small populations in the absence of a more permanent reservoir of infection. Development of appropriate interventions requires detailed data on transmission pathways between reservoirs and wildlife populations of conservation concern. Relevant data can be derived from long-term population monitoring, epidemic and case-surveillance patterns, genetic analyses of rapidly evolving pathogens, serological surveys, and intervention studies. We examined studies of carnivore diseases in the Serengeti. Epidemiological research contributes to wildlife conservation policy in terms of management of endangered populations and the integration of wildlife conservation with public health interventions. Long-term, integrative, cross-species research is essential for formulation of effective policy for disease control and optimization of ecosystem health.     

Begging calls and parental feeding decisions in tree swallows (Tachycineta bicolor)

We conducted playback experiments to examine how parent tree swallows (Tachycineta bicolor) use nestling begging calls to distribute feedings to individuals within broods. In a first study, we used a paired-choice test to determine if parents discriminated between the taped begging calls of nestlings deprived of food and those of nestlings that had been recently fed. Our results showed that parents directed their first feeding attempt towards model nestlings near speakers playing deprived calls significantly more often than to models near speakers playing fed calls. They also made more feeding attempts overall to models with deprived calls. In the second study, we varied call rate and amplitude to examine which call features parents might use to discriminate begging calls. Parents directed significantly more first feeding attempts and more feeding attempts overall towards non-begging nestlings near speakers playing high call rates than to nestlings near speakers playing low call rates. They did not, however, discriminate between calls differing in amplitude. Previous studies have shown that parent birds use begging calls to regulate overall feeding rates to the brood. Our results suggest that parent tree swallows also use begging calls when feeding individual nestlings and, in particular, prefer calls associated with increased levels of nestling hunger. Received: 14 February 2000 / Revised: 6 October 2000 / Accepted: 16 October 2000     

Predictors of food-related aggression in wild Assamese macaques and the role of conflict avoidance

The quality, availability and distribution of food resources and their influence on types and levels of feeding competition play a central role in ecological models of female social structure in mammals. Here, we investigate the impact of social and ecological factors on rates of food-related aggression and the use of potential conflict avoidance mechanisms in wild Assamese macaques (Macaca assamensis) in northeastern Thailand. These questions were addressed by examining feeding competition at the level of individual food patches which provides detailed information on resource characteristics, aggression and feeding party composition. Results suggest that the frequency of aggression in food patches increases with increasing feeding group size and decreasing patch size but is not affected by ecological variables representing resource value (abundance and nutritional value of food items in a patch). Interestingly, females appear to employ several mechanisms to avoid direct conflicts, including the use of alternative feeding sites within food patches, delayed arrival at feeding sites, storing food in cheek pouches and co-feeding with individuals they share strong social bonds with. Conflict avoidance may be partially responsible for the absence of a rank-related skew in female energy intake in this population and may explain other cases where empirical data do not fit the predictions of socio-ecological theory. Our findings also inform the debate about the mechanisms generating fitness benefits from strong social bonds by suggesting that by increasing feeding tolerance, social bonds may enhance resource acquisition, reduce the risk of injuries and lower levels of agonism-related stress.     

When do helpers help? Food availability and helping in the moorhen,Gallinula chloropus

Summary The importance of food availability as a constraint upon cooperative behaviour was examined by providing supplementary food on some breeding territories of a population of moorhens (Gallinula chloropus). First brood juveniles on territories provided with food were more likely to feed their younger second brood siblings. Supplementary feeding improved the physical condition of juveniles above that of unfed conspecifics, and birds in better condition were more likely to be helpers. Helpers on fed territories also provided more food to siblings than did their counterparts on unfed territories, enabling parent birds to reduce their work rates. Results are discussed in relation to the benefits which may accrue to helpers and the energetic costs of cooperation which may act as proximate constraints upon potential helpers.     

Food regulates reproduction differently in different habitats: experimental evidence in the Goshawk

Food supplementation experiments have been widely used to get detailed insight into how food supply contributes to the reproductive performance of wild animals. Surprisingly, even though food seldom is distributed evenly in space, variation in local habitat quality has usually not been controlled for in food supplementation studies. With results from a two-year feeding experiment involving a habitat-sensitive avian top predator, the Northern Goshawk Accipiter gentilis, we show that treatment effects on goshawk reproductive performance are habitat dependent. Extra food reduced nestling mortality in low-quality territories where prime habitat (forest) is scarce, but not in high-quality territories where prime habitat is abundant. Consequently, brood size did not differ between treatment categories in heavily forested territories, but fledgling numbers differed between unfed and fed goshawk pairs breeding in territories where forest is scarce. However, because extra food was not superabundant, this artificial increase in offspring number induced a dramatic decrease in nestling condition in low-quality territories. Treatment effects were detected even after controlling statistically for other potentially confounding effects (year, territory identity) and strongly covaried with territory-specific abundances of the most important summer prey species. These results highlight the importance of acknowledging the effect that small-scale variation in habitat quality and availability of natural food may have on the results of food supplementation experiments. In order to assess the generality of food supplementation effects, the integration of habitat heterogeneity and variation in food abundance is thus needed, especially among species in which small-scale variation in habitat quality influences demographic patterns.     

Assessing Suitability for Conservation Action: Prioritizing Interpond Linkages for the California Tiger Salamander

Abstract:  Conservation organizations and public agencies are interested in identifying and prioritizing areas for conservation action, often acquisition or easements. Typically, this requires the use of uncertain data and vaguely defined decision criteria. I developed a decision support system to address these uncertainty issues and assist in evaluating conservation opportunities for the endangered California tiger salamander ( Ambystoma californiense ) in Santa Barbara, California. Functionally defined planning units were used to aggregate data on land suitability, land cover change, salamander presence, and movement risk along potential linkages between breeding ponds. I used a fuzzy-logic-based inference engine to evaluate the planning units and rank the relative suitability of interpond linkages for conservation action. The sensitivity of the rankings was considered with respect to uncertainty in salamander occurrence data and the relationship between land-cover-change threats and site suitability. All linkages were substantially degraded, but five areas were consistently identified with high relative suitability for conservation action despite differences in assumptions and uncertainty in biological data. The combination of functionally defined planning units and a fuzzy-logic-based decision support system provides a general framework for considering the suitability of sites for conservation action.     

Effects of a Proposed Ex Situ Conservation Program on In Situ Conservation of the Babirusa, an Endangered Suid

Abstract: Detailed data are rarely available to show how interventions such as captive breeding programs can create an uncontrolled demand for live specimens of endangered species. We present a case study of the effect of a planned, internationally recognized captive breeding program on trade in the endangered babirusa wild pig from July to December 1998. Although the program had not yet begun, international interest in the captive breeding of babirusas gave hunters and dealers the false impression that there was a potentially lucrative and officially sanctioned national and international demand for any live babirusas they might catch. Swift action by the Indonesian authorities halted this trade, but the study provides a warning about the damage that can be caused to the conservation of a species if management programs are instituted without a full understanding of the practicalities of its conservation, particularly interactions between the species and local people.     

Structuring Decisions for Managing Threatened and Endangered Species in a Changing Climate

The management of endangered species under climate change is a challenging and often controversial task that incorporates input from a variety of different environmental, economic, social, and political interests. Yet many listing and recovery decisions for endangered species unfold on an ad hoc basis without reference to decision‐aiding approaches that can improve the quality of management choices. Unlike many treatments of this issue, which consider endangered species management a science‐based problem, we suggest that a clear decision‐making process is equally necessary. In the face of new threats due to climate change, managers’ choices about endangered species require closely linked analyses and deliberations that identify key objectives and develop measurable attributes, generate and compare management alternatives, estimate expected consequences and key sources of uncertainty, and clarify trade‐offs across different dimensions of value. Several recent cases of endangered species conservation decisions illustrate our proposed decision‐focused approach, including Gulf of Maine Atlantic salmon (Salmo salar) recovery framework development, Cultus Lake sockeye salmon (Oncorhynchus nerka) management, and Upper Columbia River white sturgeon (Acipenser transmontanus) recovery planning. Estructuración de Decisiones para Manejar Especies Amenazadas y en Peligro en un Clima Cambiante     

Problems with mitigation translocation of herpetofauna

Mitigation translocation of nuisance animals is a commonly used management practice aimed at resolution of human–animal conflict by removal and release of an individual animal. Long considered a reasonable undertaking, especially by the general public, it is now known that translocated subjects are negatively affected by the practice. Mitigation translocation is typically undertaken with individual adult organisms and has a much lower success rate than the more widely practiced conservation translocation of threatened and endangered species. Nonetheless, the public and many conservation practitioners believe that because population‐level conservation translocations have been successful that mitigation translocation can be satisfactorily applied to a wide variety of human‐wildlife conflict situations. We reviewed mitigation translocations of reptiles, including our own work with 3 long‐lived species (Gila monsters [Heloderma suspectum], Sonoran desert tortoises [Gopherus morafkai], and western diamond‐backed rattlesnakes [Crotalus atrox]). Overall, mitigation translocation had a low success rate when judged either by effects on individuals (in all studies reviewed they exhibited increased movement or increased mortality) or by the success of the resolution of the human–animal conflict (translocated individuals often returned to the capture site). Careful planning and identification of knowledge gaps are critical to increasing success rates in mitigation translocations in the face of increasing pressure to find solutions for species threatened by diverse anthropogenic factors, including climate change and exurban and energy development. Problemas con la Mitigación por Traslocación de Herpetofauna