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Ingestion of bones, stones, and soil by desert tortoises
Abstract
We used field observations and radiographs t document that desert tortoises (Gopherus agassizii) ingest soil and stones. The ingestion of soil and stones can be a common albeit rarely observed behavior of desert tortoises. We also describe the ingestion of bones by wild tortoises. Desert tortoises readily eat weathered bones, and some individuals persist in eating bone even when they are disturbed. Eating soil, stones, and bones could be important in supplementing the herbivorous diet of tortoises, and thus the relative use of these behaviors in different localities could serve as indicators of nutritive stress. Other possible benefits from the ingestion of soil, stones, and bones include the maintenance of gut pH, nullification of platen secondary compounds, control of intestinal parasites, or maintenance of beak shape (in a fashion analogous to the behavior of birds). Bones and other mineral resources seem to be important to tortoises. However, the mechanistic explanations of eating soil, stones and bones are currently unknown. Research into the mineral physiology of tortoises is necessary to determine the causes of the ingestion of soil, bones, and stones and to provide guidance for management.
... Among chelonians, most reports of geophagy are limited to terrestrial taxa, especially female tortoises (Testudinidae; e.g. Esque and Peters 1994;Walde et al. 2007; Moore and Dornburg 2014;Sullivan and Cahill 2019) and hatchlings of the painted turtle, Chrysemys picta (Packard et al. 2001;Costanzo et al. 2003;Packard 2003a, 2006). Despite reports of geophagy and its significance in animals, there exists no study comparing the intra-or interspecific frequency of this behaviour in chelonians, especially in early life stages. ...
... For example, several thousand hours of field research over 11 years, comprising several hundred tortoises of both sexes, resulted in only six observations of female Gopherus agassizii exhibiting geophagy (Marlow and Tollestrup 1982). Geophagic behaviour was directly observed in < 1% of wild G. agassizii, although radiographs revealed stones and soil in 67% of individuals (Esque and Peters 1994). Observations of geophagy for semiaquatic and aquatic chelonians are nearly absent in the literature, excluding the terrestrial hatchling life stage (Table S1). ...
... On the contrary, a growing number of behavioural observations report juvenile and adult tortoises, namely females, actively forage for and consume gastroliths (e.g. Esque and Peters 1994;Stitt and Davis 2003;Sullivan and Cahill 2019; Table S1). There is no clear indication whether hatchling turtles incidentally and/or intentionally consume their nesting substrate. ...
Geophagy is the consumption of hard objects with no caloric value (e.g. soil, sand, sediment) called gastroliths. This behaviour is widespread in animals, and among reptiles, geophagy has been reported in crocodilians and lizards and occasionally in chelonians. In this study, we calculated geophagy rates in snapping turtle (Chelydra serpentina) and painted turtle (Chrysemys picta) hatchlings under various incubation protocols, ranging from highly artificial to semi-natural conditions. Among multiple experiments where eggs were incubated atop the nesting substrate, 66% of painted turtle and 58–93% of snapping turtle hatchlings exhibited geophagy within 24 h of hatching. Snapping turtle hatchlings that experienced a simulated natural nest emergence had an 85–100% rate of gastrolith consumption. Hatchling snapping turtles from shallow simulated nests emerged earlier and had higher rates of geophagy compared with those from deeper nests. Given the high frequency, short time period (24~72 h) and variety of incubation protocols under which geophagy occurred, we suggest that this behaviour is intentional. We discuss multiple hypothesis for the adaptive and functional significance of geophagic behaviour with respect to chelonians, synthesize existing literature on chelonian geophagy and highlight the possible implications of hatchling turtle geophagy for ex situ captive breeding and head-starting programs. Given that hatchlings readily consume their incubation medium, caretakers should carefully consider the substrate their animals are exposed to. Future research should address how widespread geophagy is among hatchling turtles and the possible role of this behaviour for hatchling ecology and health, including effects on the gut microbiome.
Significance statement
Animals regularly consume non-caloric foods, such as rocks and soils. This behaviour is exhibited by hatchling turtles, but why? Our literature review suggests that the ingestion of non-caloric foods by turtles is important for nutrition and, in particular, this behaviour may help establish gut health in hatchlings. Observational and experimental study demonstrated that nest characteristics in-part account for why hatchling ‘turtles eat dirt’. This has applications for hatchling ecology as well as captive rearing conservation programs, an increasingly common strategy for the conservation of these globally imperilled animals. This work complements several recent studies and review articles about geophagy in other major vertebrate groups (birds, mammals) and provides a comprehensive summary on the current state of knowledge of this behaviour for turtles.
... The ingestion of pebbles or rocks, dubbed gastroliths (Wings 2007), as physiological aids is a well-known phenomenon that occurs in various vertebrate clades including birds (Smith and Rastall 1911;Franson et al. 2001;Clausen and Pütz 2003;Beaune et al. 2009), crocodilians (Cott 1961;Brazaitis 1969;Delany and Abercrombie 1986;Fitch-Snyder and Lance 1993), mammals (Nord¯y 1995;Drehmer and Oliveira 2003), and dinosaurs (Wieland 1906;Sanders et al. 2001;Wings and Sander 2007;Cerda 2008). Ingestion of gastroliths provides support for a variety of physiological challenges spanning digestion (Sokol 1971; Thomas et al. 1977), parasite removal (Esque and Peters 1994;Robinson et al. 2008), stomach cleaning (Wings 2007), buoyancy modification (Taylor 1993), and the supplementation of nutritional deficiencies (Kreulen 1985;Panichev et al. 2013). Given the range of gastrolith uses, documenting and testing for factors that promote their ingestion represents an important step toward understanding the basic ecophysiology of organisms that ingest them. ...
... Several ornithological investigations have noted increased ingestion of snail shells, calcareous grit, small bones and seashells by egg-laying female birds (MacLean 1974;Ankey and Scott 1980;Graveland et al. 1994;Brenninkmeijer et al. 1997). Likewise, desert tortoises (Gopherus agassizi) and leopard tortoises (Geochelone pardalis) have also been noted to eat stones and bones (Milton 1992;Esque and Peters 1994). This behavior has been linked to calcium supplementation for eggshell deposition: Marlow and Tollestrup (1982) observed desert tortoises digging to forage on a leached caliche (calcium carbonate) bed during nesting season. ...
Instances of vertebrates supplementing their diets through the ingestion of gastroliths have been
observed or documented in numerous organisms, including birds, crocodiles and dinosaurs. Although
not uncommon, this behavior can be motivated by a wide range of factors that include
endoparasite removal, stomach cleaning, or calcium supplementation during eggshell formation.
As plant-based diets are often mineral deficient, the use of gastroliths as calcium supplements
may play an important role in the reproductive biology of egg-laying herbivorous reptiles.
Whereas gravid females of some species of herbivorous reptiles have been shown to seek out calcareous
gastroliths, the occurrence or prevalence of gastroliths in gravid females remains unknown
for many species. Using field observations and radiographs, we studied patterns of
gastrolith ingestion by adult female gopher tortoises (Gopherus polyphemus), a herbivorous
species of testudinoid turtles. Gastroliths documented included fossil seashells, calcareous stones
and carnivore feces containing vertebrate bones. The prevalence of gastroliths was significantly
different between gravid and non-gravid tortoises. We suggest this behavior is a means of supplementing
calcium intake during eggshell formation and may play a part in the female reproductive
biology of this species.
... The two species of tortoise had a similar niche breadth for use of vegetation types, and there was considerable overlap between them (0 = 0.76; Hailey and Coulson 1995). Gross features of the habitat thus cannot account for the large home range of G. pardalis, but minor features, such as sites where soil is ingested for minerals (Esque and Peters 1994), may be important. Movements of desert tortoises for the purpose of mineral ingestion have been reported by Marlow and Tollestrup (1982), though these were much shorter than the kilometre-scale movements reported here. ...
... For personal use only. nians also appear to have been for calcium (Esque and Peters 1994), but this is unlikely in G. pardalis. The level of calcium in the soil at the site of ingestion was not notably high, and other sources of calcium would be available in the area. ...
Movements of the tortoises Geochelone pardalis (mean body mass 4.0 kg) and Kinixys spekii (0.62 kg) were studied by thread-trailing. The mean daily movement distance of G. pardalis was 435 m, and the short-term home-range area was 26 ha. The values for K. spekii were 172 m and 1.9 ha, respectively. The area used by G. pardalis was significantly larger than would be produced by scaling up the movements of K. spekii in the same geometric pattern. Home-range areas were about the same size (K. spekii) or several times as large (G. pardalis) as those predicted for mammals of equal body mass. Ingestion of soil for sodium may be a factor determining the large home range of the purely herbivorous G. pardalis. These tortoises frequented a small area of strongly sodic soil from which they dispersed over long distances.
... Three ISO 25178 parameters were calculated (Table 4): the root mean square height (Sq), the degree of bias of the shape of the roughness (Ssk), and the developed surface ratio (Sdr). Previous studies have shown the pertinence of these parameters to characterize different degrees of polish on archaeological artefacts [11,39,64]. In addition, they present the advantage of being largely independent from one another. ...
Upper Palaeolithic sites in southwestern France attributed to the Upper Gravettian and the Solutrean yielded sub spherical gravels with a highly shiny appearance that have intrigued researchers since the 1930s. In this work, we analyze specimens from five sites, including the recently excavated Solutrean site of Landry, to establish whether their presence in archaeological layers and peculiar aspect are due to natural processes or human agency. We study the spatial distribution of gravels at Landry and submit archaeological gravels from the five sites, natural formations, Landry sediment sieving, and polishing experiments with a rotary tumbling machine to morphometric, colorimetric, microscopic, and textural analyses. Our results indicate the lustrous gravels found at the five sites result from deliberate selection and suggest their shiny appearance is the consequence of human agency, possibly resulting from prolonged contact with a soft material such as animal skin. Ethnographic accounts indicate that these gravels may have been used for magico-religious ritual purposes (charms, sorcery, divination etc.), in games, as elements of musical instruments, and as items serving other social and personal purposes. We argue that these objects reflect a cultural innovation emerged during the Gravettian and continued into the Solutrean.
... Likewise, we believe that the current diversity of terrestrial chelonians is the result of a history of environmental changes that led to a different plant composition in the landscape. It is believed that moments of great vegetation change may raise the extinction rates for chelonians, considering that most terrestrial species are completely or predominantly herbivores, feeding on leaves, fruits or flowers, though some can complement their diet opportunistically with animal-derived food sources, such as invertebrates, bones and feces (Esque & Peters, 1994;Pritchard, 1979;Rafferty, 2011). The present work aimed on a fossil-based analysis of the diversification process of tortoises, exposing the dynamics of speciation and extinction in order to find peaks or lows and exploring its correlates. ...
Great environmental changes may affect the survival capability of a variety of organisms. Testudinidae is the most diverse family of terrestrial chelonians within the whole order (Testudines). Interestingly, however, the number of extinct species overcome the extant ones. In order to understand better how the diversification process of this family occurred, this work used the PyRate software, which estimates both the preservation and diversification processes in a continuous time interval. For such, the software used a list of fossil occurrences obtained from the Paleobiology Database whereas the extant species list was obtained from Catalogue of Life. This way, the software was able to infer the probability of the ancestral clade having resulted in these species during its evolutionary history. The analyses generated graphs containing the diversification, extinction and speciation curves and their respective associated 95% credibility intervals. A great rise in the extinction rate was observed starting 6 million years ago. This rise is believed to be related to the drop of atmospheric CO2 all over the globe at the end of the Miocene, about 8-6 million years ago. This event led to a turnover of the vegetation composition on the warmer areas of the planet, with plants that used C3 metabolism giving way to C4 plants. In terms of landscape, grasses and herbal vegetation, such as savannas, started dominating. As for other animal groups, those herbivores with grazing habits were more successful than those used to only browsing or that did not have enough flexibility of choice.
... early hominids) appear to be at a lower trophic level from these ratios (Reynard, Henderson & Hedges, 2010). Osteophagous herbivorous taxa (Esque & Peters, 1994;Hutson, Burke & Haynes, 2013) will presumably appear to be at a higher trophic level. ...
Birds are some of the most diverse organisms on Earth, with species inhabiting a wide variety of niches across every major biome. As such, birds are vital to our understanding of modern ecosystems. Unfortunately, our understanding of the evolutionary history of modern ecosystems is hampered by knowledge gaps in the origin of modern bird diversity and ecosystem ecology. A crucial part of addressing these shortcomings is improving our understanding of the earliest birds, the non‐avian avialans (i.e. non‐crown birds), particularly of their diet. The diet of non‐avian avialans has been a matter of debate, in large part because of the ambiguous qualitative approaches that have been used to reconstruct it. Here we review methods for determining diet in modern and fossil avians (i.e. crown birds) as well as non‐avian theropods, and comment on their usefulness when applied to non‐avian avialans. We use this to propose a set of comparable, quantitative approaches to ascertain fossil bird diet and on this basis provide a consensus of what we currently know about fossil bird diet. While no single approach can precisely predict diet in birds, each can exclude some diets and narrow the dietary possibilities. We recommend combining (i) dental microwear, (ii) landmark‐based muscular reconstruction, (iii) stable isotope geochemistry, (iv) body mass estimations, (v) traditional and/or geometric morphometric analysis, (vi) lever modelling, and (vii) finite element analysis to reconstruct fossil bird diet accurately. Our review provides specific methodologies to implement each approach and discusses complications future researchers should keep in mind. We note that current forms of assessment of dental mesowear, skull traditional morphometrics, geometric morphometrics, and certain stable isotope systems have yet to be proven effective at discerning fossil bird diet. On this basis we report the current state of knowledge of non‐avian avialan diet which remains very incomplete. The ancestral dietary condition in non‐avian avialans remains unclear due to scarce data and contradictory evidence in Archaeopteryx. Among early non‐avian pygostylians, Confuciusornis has finite element analysis and mechanical advantage evidence pointing to herbivory, whilst Sapeornis only has mechanical advantage evidence indicating granivory, agreeing with fossilised ingested material known for this taxon. The enantiornithine ornithothoracine Shenqiornis has mechanical advantage and pedal morphometric evidence pointing to carnivory. In the hongshanornithid ornithuromorph Hongshanornis only mechanical advantage evidence indicates granivory, but this agrees with evidence of gastrolith ingestion in this taxon. Mechanical advantage and ingested fish support carnivory in the songlingornithid ornithuromorph Yanornis. Due to the sparsity of robust dietary assignments, no clear trends in non‐avian avialan dietary evolution have yet emerged. Dietary diversity seems to increase through time, but this is a preservational bias associated with a predominance of data from the Early Cretaceous Jehol Lagerstätte. With this new framework and our synthesis of the current knowledge of non‐avian avialan diet, we expect dietary knowledge and evolutionary trends to become much clearer in the coming years, especially as fossils from other locations and climates are found. This will allow for a deeper and more robust understanding of the role birds played in Mesozoic ecosystems and how this developed into their pivotal role in modern ecosystems. Video abstract
... Consumptive taste associations for several Ca salts have been observed among diverse groups of animals including amphibians, birds, and mammals (Gabriel et al., 2009;Niknafs & Roura, 2018;Tordoff, 2001;Tordoff et al., 2008). Calcium taste is likely to be under stronger selection in herbivores than predators, due to the generally lower Ca content of plant biomass (Figure 2; (Walde et al., 2007), and Ca carbonate stones (Esque & Peters, 1994). Though it remains unclear if these foraging behaviors are informed by gustatory sensing of Ca or some other mechanism, the discovery of Ca taste and associated taste receptors in a variety of species suggests a potential role for gustation (Gabriel et al., 2009;Niknafs & Roura, 2018;Tordoff, 2001;Tordoff et al., 2008). ...
A major conceptual gap in taste biology is the lack of a general framework for understanding the evolution of different taste modalities among animal species. We turn to two complementary nutritional frameworks, biological stoichiometry theory and nutritional geometry, to develop hypotheses for the evolution of different taste modalities in animals. We describe how the attractive tastes of Na‐, Ca‐, P‐, N‐, and C‐containing compounds are consistent with principles of both frameworks based on their shared focus on nutritional imbalances and consumer homeostasis. Specifically, we suggest that the evolution of multiple nutritive taste modalities can be predicted by identifying individual elements that are typically more concentrated in the tissues of animals than plants. Additionally, we discuss how consumer homeostasis can inform our understanding of why some taste compounds (i.e., Na, Ca, and P salts) can be either attractive or aversive depending on concentration. We also discuss how these complementary frameworks can help to explain the evolutionary history of different taste modalities and improve our understanding of the mechanisms that lead to loss of taste capabilities in some animal lineages. The ideas presented here will stimulate research that bridges the fields of evolutionary biology, sensory biology, and ecology. This paper describes how the complementary nutritional frameworks, biological stoichiometry theory and nutritional geometry, can inform our understanding of the evolution of multiple taste modalities in animals. We describe how the attractive tastes of Na‐, Ca‐, P‐, N‐, and C‐containing compounds are consistent with principles of both frameworks based on their shared focus on nutritional imbalances and consumer homeostasis.
... Other possible vertebrate actors in the Mygatt-Moore assemblage include turtles, salamanders and rhynchocephalians (Foster et al., 2018). In modern observational studies, turtles have been noted to use their keratinous beaks to pry off chips of weathered bone for ingestion, rather than percussively gnawing on the bone surface (Esque & Peters, 1994), which is inconsistent with the morphology of M. bohemicus. As for amphibians, the anterior tooth morphology of salamanders is not compatible with the protruding incisors needed to create this trace, and adult rhynchocephalians are too large to be a potential actor. ...
A survey of 2,368 vertebrate fossils from the Upper Jurassic Mygatt-Moore Quarry (MMQ) (Morrison Formation, Brushy Basin Member) in western Colorado revealed 2,161 bone surface modifications on 884 specimens. This is the largest, site-wide bone surface modification survey of any Jurassic locality. Traces made by invertebrate actors were common in the assemblage, second in observed frequency after vertebrate bite marks. Invertebrate traces are found on 16.174% of the total surveyed material and comprise 20.148% of all identified traces. Six distinct invertebrate trace types were identified, including pits and furrows, rosettes, two types of bioglyph scrapes, bore holes and chambers. A minimum of four trace makers are indicated by the types, sizes and morphologies of the traces. Potential trace makers are inferred to be dermestid or clerid beetles, gastropods, an unknown necrophagous insect, and an unknown osteophagus insect. Of these, only gastropods are preserved at the site as body fossils. The remaining potential trace makers are part of the hidden paleodiversity from the North American Late Jurassic Period, revealed only through this ichnologic and taphonomic analysis. Site taphonomy suggests variable, but generally slow burial rates that range from months up to 6 years, while invertebrate traces on exposed elements indicate a minimum residence time of five months for carcasses with even few preserved invertebrate traces. These traces provide insight into the paleoecology, paleoclimate, and site formation of the MMQ, especially with regards to residence times of the skeletal remains on the paleolandscape. Comprehensive taphonomic studies, like this survey, are useful in exploring patterns of paleoecology and site formation, but they are also rare in Mesozoic assemblages. Additional work is required to determine if 16.174% is typical of bulk-collected fossils from Jurassic ecosystems in North America, or if the MMQ represents an unusual locality.
... Les animaux -Des études éthologiques sur des espèces actuelles et passées ont démontré que la géophagie et la lithophagie sont un comportement assez commun chez les animaux, comme par exemple les crocodiles, les pinnipèdes et un grand nombre d'herbivores et d'oiseaux (Tryon, 2010 ;Wings, 2004 ;Esque et Peters, 1994 ;Scott, 1982 ;Hladik et Gueguen, 1974). Cette consommation aurait plusieurs vertus et fonctionnalités dans lesquelles chaque espèce y trouve son compte. ...
In this work we analyze shiny gravels from the newly excavated Solutrean site of Landry and from Fourneau du Diable, Abri Casserole, Laugerie-Haute and Pech de la Boissière to establish whether they are natural or anthropogenic in origin. We study the spatial distribution of gravels at Landry and a variety of sedimentary contexts, and submit archaeological gravels, gravels from Landry sieving, and experimentally polished gravels to morphometric, colorimetric and roughness analyses. Spatial analysis reveals that shiny gravels are concentrated at Landry in the areas in which the remainder of the archaeological material was recovered, which supports the anthropogenic argument. Roughness analysis demonstrate that the shine present on the archaeological specimens is not observed on natural gravel and show a degree of similarity with gravels experimentally polished in a tumbler covered with skin. Our results are consistent with the hypothesis that shiny gravels are the result of deliberate selection and their shiny appearance is the consequence of long-term contact with animal skin, by their use as ornaments or elements of musical instruments. We propose that these small shiny stones were a part of a cultural innovation prompted by Gravettian hunter-gatherers and developed by those of the Solutrean.
... These episodes, known as geophagy, have only been observed in sexually mature females during the nesting season, a time when females may experience calcium stress due to egg production (Marlow and Tollestrup 1982). Osteophagy, or the consumption of bones, has also been observed in G. agassizii and may be similarly related to calcium, phosphorus, or other mineral deficiencies in the diet (Esque and Peters 1994). ...
Of the 356 species of turtles worldwide, approximately 61% are threatened or already extinct. Turtles are among the most threatened of the major groups of vertebrates, in general, more so than birds, mammals, fishes or even the much besieged amphibians. Reasons for the dire situation of turtles worldwide include the familiar list of impacts to other species including habitat destruction, unsustainable overexploitation for pets and food, and climate change (many turtles have environmental sex determination). Two notable characteristics of pre-Anthropocene turtles were their massive population sizes and correspondingly high biomasses, the latter among the highest values (over 855 kilograms per hectare) ever reported for animals. As a result of their numerical dominance, turtles have played important roles as significant bioturbators of soils, infaunal miners of sea floors, dispersers and germination enhancers of seeds, nutrient cyclers, and consumers. The collapse of turtle populations on a global scale has greatly diminished their ecological roles.
... Some vertebrates consume their own or other individuals' feces, as exemplified by woolly mammoths (van Geel et al., 2011), and herbivore feces might be mistaken for carnivore/omnivore feces too. Some herbivorous animals are known to ingest bones, perhaps for supplementation of minerals, as for example tortoises (Esque and Peters, 1994). In such a case, the bulk of the coprolite material would be from a herbivore, which would obviously influence the interpretation. ...
Residues of twenty-five coprolite fragments collected from the Upper Permian of Vyazniki (European Russia) were studied in detail. The phosphatic composition, general shape and size, and bone inclusions of these specimens indicate that medium to large-sized carnivores, such as therocephalian therapsids or early archosauriforms, were the most likely coprolite producers. The contents of the examined fossils (i.e. scale, bone and tooth fragments, mineral grains, and microbial structures) do not differ significantly among the samples, implying fairly comparable feeding habits of their producers. Fragments of large tooth crowns in two of the analyzed samples imply that either (1) the coprolite producer swallowed the cranial elements of its prey or (2) the coprolite producer broke and swallowed its own tooth while feeding (such tooth damage is known in archosaurs that have tooth replacement, e.g. crocodiles and dinosaurs). Indeed, the most complete tooth fragment in these fossils is serrated, most likely belonging to an early archosauriform known from skeletal records from the Late Permian of Vyazniki. Another coprolite fragment contains the etched tooth of a lungfish, while putative actinopterygian fish remains (scales and small fragments of bones) are abundant in some samples. Mineral particles (mostly quartz grains, feldspars and mica) may have been swallowed accidentally. The preserved microbial colonies (mineralized fossil fungi and bacteria or their pseudomorphs), manifested in the coprolites as Fe-rich mineral structures, seem to have developed on the expelled feces rather than on the items before they were swallowed.
... Results from this study have implications for assessing risks associated with ecological health and with potential mobilization of natural ClO 4 to groundwater. Direct ingestion of soil by livestock and wildlife (e.g., Mayland et al., 1975; Beyer et al., 1994; Esque and Peters, 1994) and transfer from soil through plants to higher organisms can contribute to ClO Rao et al., 2007 ), the depthnormalized average for shallow soil (48,300 mg ha -1 m -1 ) is 4 times that for the deeper profile (11,970 mg ha -1 m -1 ). ...
... George, Utah Field Office. His continued association with Bruce resulted in a fruitful collaboration on the book "Biology of North American Tortoises", in which Todd contributed to three chapters (Bury et al. 1994a;Esque and Peters 1994;. Later, he contributed to another book on Morafka's Desert Tortoise in the Sonoran Desert (Esque et al. 2002). ...
The U.S. Department of the Interior (DOI) has a long and distinguished history of employing herpetologists to conduct basic and applied research to better manage amphibian and reptile populations on public lands and even outside the boundaries of the United States. This history extends back over 125 years with roots in the U.S. Biological Survey, the Fish and Wildlife Service, the Bureau of Land Management, the National Park Service and later, the National Biological Service. In more recent times, the DOI employed more professional herpetologists than any single organization in the world, especially in the U.S. Geological Survey. In 1938, Henry Fitch was the first Interior scientist hired who conducted substantial herpetological research. William and Lucille Stickel of the Fish and Wildlife Service conducted herpetological research throughout the period from the 1940s-1980s but most DOI herpetologists were hired from 1975-80 with another hiring spike from 2000-2005. The former spike was congruent with early versions of the Endangered Species Act while the latter reflected growing recognition of global amphibian decline and the creation of the Amphibian Research and Monitoring Initiative in DOI. Collectively, these herpetologists produced hundreds of books, scientific publications and other scholarly publications, many of which are classics in the literature. In addition, many have served as officers and on the boards of numerous scientific societies particularly those specializing in amphibian and reptile research. The DOI shows a continuing commitment to funding herpetological research by hiring young scientists to replace the aging ranks of herpetologists who started their careers in the 1970s. This commitment is critical given the global decline of both amphibians and reptiles, including those found on public lands in the United States.© 2012. Herpetological Conservation and Biology. All Rights Reserved.
... In addition, a mandibular ramus of Clevosaurus was found associated with the gastroliths (PEO, personal observation), which led Barrett (2000) to argue that 'prosauropods' were omnivorous. Possible alternative explanations are that the small reptile bone was deliberately ingested for nutrients (Esque and Peters 1994;White 2011), or that the bone was accidentally associated with the dinosaurian remains (Fedak 2007). ...
The Fundy basin in Nova Scotia and New Brunswick is the largest exposed rift basin of the Newark Supergroup and also extends beneath the Bay of Fundy. Its strata can be divided into four tectonostratigraphic sequences (TS). TS I is represented by the probably Permian Honeycomb Point Formation and possibly the Lepreau Formation. TS II includes the Wolfville Formation with the probably Middle Triassic Economy Member and the early Late Triassic Evangeline Member. These members have yielded markedly different assemblages of continental tetrapods. TS III comprises most of the Blomidon Formation, which is Norian to Rhaetian in age. The Blomidon Formation has yielded few skeletal remains of tetrapods to date but many tetrapod tracks. TS IV includes the late Rhaetian top of the Blomidon Formation and the McCoy Brook Formation, which overlies the North Mountain Basalt and is latest Rhaetian and earliest Jurassic (Hettangian) in age. The McCoy Brook Formation has yielded a diversity of continental tetrapods and lacks any of the characteristic Late Triassic forms. Recent work has correlated the Global Boundary Stratotype Section and Point (GSSP) for the base of the Jurassic (Hettangian) to a level above the North Mountain Basalt. Thus most of the tetrapod fossils from the McCoy Brook Formation are latest Rhaetian in age, but the higher horizon with skeletal remains of sauropodomorph dinosaurs may be earliest Hettangian in age. The Fundy basin preserves the only known, stratigraphically tightly constrained record of the profound biotic changes in continental ecosystems across the Triassic-Jurassic transition.
... In addition, a mandibular ramus of Clevosaurus was found associated with the gastroliths (PEO, personal observation), which led Barrett (2000) to argue that 'prosauropods' were omnivorous. Possible alternative explanations are that the small reptile bone was deliberately ingested for nutrients (Esque and Peters 1994;White 2011), or that the bone was accidentally associated with the dinosaurian remains (Fedak 2007). ...
The Fundy basin in Nova Scotia and New Brunswick is the largest exposed rift basin of the Newark Supergroup and also extends beneath the Bay of Fundy. Its strata can be divided into four tectonostratigraphic sequences (TS). TS I is represented by the probably Permian Honeycomb Point Formation and possibly the Lepreau Formation. TS II includes the Wolfville Formation with the probably Middle Triassic Economy Member and the early Late Triassic Evangeline Member. These members have yielded markedly different assemblages of continental tetrapods. TS III comprises most of the Blomidon Formation, which is Norian to Rhaetian in age. The Blomidon Formation has yielded few skeletal remains of tetrapods to date but many tetrapod tracks. TS IV includes the late Rhaetian top of the Blomidon Formation and the McCoy Brook Formation, which overlies the North Mountain Basalt and is latest Rhaetian and earliest Jurassic (Hettangian) in age. The McCoy Brook Formation has yielded a diversity of continental tetrapods and lacks any of the characteristic Late Triassic forms. Recent work has correlated the Global Boundary Stratotype Section and Point (GSSP) for the base of the Jurassic (Hettangian) to a level above the North Mountain Basalt. Thus most of the tetrapod fossils from the McCoy Brook Formation are latest Rhaetian in age, but the higher horizon with skeletal remains of sauropodomorph dinosaurs may be earliest Hettangian in age. The Fundy basin preserves the only known, stratigraphically tightly constrained record of the profound biotic changes in continental ecosystems across the Triassic-Jurassic transition.
... Results from this study have implications for assessing risks associated with ecological health and with potential mobilization of natural ClO 4 to groundwater. Direct ingestion of soil by livestock and wildlife (e.g., Mayland et al., 1975;Beyer et al., 1994;Esque and Peters, 1994) and transfer from soil through plants to higher organisms can contribute to ClO 4 exposure in a desert landscape. The creosote bush canopy represents a previously unrecognized ClO 4 reservoir (1310 mg ha -1 ). ...
Perchlorate (ClO) is a contaminant that occurs naturally throughout the world, but little is known about its distribution and interactions in terrestrial ecosystems. The objectives of this Amargosa Desert, Nevada study were to determine (i) the local-scale distribution of shallow-soil (0-30 cm) ClO with respect to shrub proximity (far and near) in three geomorphic settings (shoulder slope, footslope, and valley floor); (ii) the importance of soil, plant, and terrain variables on the hillslope-distribution of shallow-soil and creosote bush [ (Sessé & Moc. ex DC.) Coville] ClO; and (iii) atmospheric (wet plus dry, including dust) deposition of ClO in relation to soil and plant reservoirs and cycling. Soil ClO ranged from 0.3 to 5.0 μg kg. Within settings, valley floor ClO was 17× less near shrubs due in part to enhanced leaching, whereas shoulder and footslope values were ∼2× greater near shrubs. Hillslope regression models (soil, = 0.42; leaf, = 0.74) identified topographic and soil effects on ClO deposition, transport, and cycling. Selective plant uptake, bioaccumulation, and soil enrichment were evidenced by leaf ClO concentrations and Cl/ClO molar ratios that were ∼8000× greater and 40× less, respectively, than soil values. Atmospheric deposition ClO flux was 343 mg ha yr, ∼10× that for published southwestern wet-deposition fluxes. Creosote bush canopy ClO (1310 mg ha) was identified as a previously unrecognized but important and active reservoir. Nitrate δO analyses of atmospheric deposition and soil supported the leaf-cycled-ClO input hypothesis. This study provides basic data on ClO distribution and cycling that are pertinent to the assessment of environmental impacts in desert ecosystems and broadly transferable to anthropogenically contaminated systems.
Copyright © by the American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America, Inc.
... La géophagie est également remarquée chez les oiseaux, comme par exemple en Nouvelle-Guinée , sans que sa fonction ait été clairement identifiée par les auteurs (Diamond et al. 1999 ). Ce type de comportement a déjà été observé et rapporté chez les tortues en captivité et plus rarement dans la nature (Sokol 1971, Kramer 1973, Cheylan 1981, Mc Donald & Mushinsky 1988, Esque & Peters 1994, Huot-Daubremont 1996). En Afrique, Stigmochelys pardalis consomme préférentiellement des sols riches en sodium (Hailey & Coulson 1996Sokol 1971, Cheylan 1981, Huot Daubremont 1996 ). ...
Geophagy by Hermann’s Tortoise, Testudo hermanni Gmelin, 1789 (Chelonii, Testudinidae)
in the region of the Maures (Var, France). Eating some ground elements, a behaviour
called « geophagy », is observed mainly in captive tortoises but also in the wild. For the first time, this
study reports a quantitative analysis of such absorption by free ranging Hermann’s tortoises, Testudo
hermanni in department of Var (France). The analysis of digestive tract shows an important quantity of
soil elements, with sometimes a few large particles, which makes us wonder about the purpose of this
behaviour.
... Research on desert tortoises has focused on 2 nutrient wisdom hypotheses (Minnich 1977(Minnich , 1979Medica 1977, 1986;Jarchow 1984;Oftedal and Allen 1996;Oftedal 2002;Oftedal and others 2002). The first hypothesis centers on balancing the need for calcium, phosphorus and magnesium for bones, shell and eggs (Esque and Peters 1994). Additionally, calcium is required for maintenance of osmotic balance in cells, blood coagulation, nerve transmission and muscular activity (Despopoulos and Silbernagl 1991). ...
... Rugiero, M. Capula, L. Luiselli, personal observation). Tortoise egg shells were eaten during autumn by semiwild T. hermanni in France (Huot-Daubremont, 1999) as well by Gopherus agassizii in the United States (Esque and Peters, 1994), but we found no evidence of this at our study area, although such egg shells are typically available (L. Rugiero, M. Capula, L. Luiselli, personal observation). ...
Herbivory is the dominant feeding strategy in tortoises, and dietary shifts are common in response to changes in resource availability. We conducted the first large-scale study of the diets of wild Hermann’s Tortoises (Testudo hermanni hermanni) and found that the study population in central Italy was strictly herbivorous. The tortoises ate primarily legume leaves and grasses in the spring, and switched to
flowers and unripe fruit of Ruscus aculeatus as these became available in the autumn. There were no
significant differences between the diets of males and females. Although tortoise diets included both rare and abundant plant species, they consumed abundant plant species in a higher proportion than those species occurred in the study area. However, some rare plants made up relatively large fractions of the diet, and one of the few nonnative plants (Conyza canadensis) at the study area was eaten frequently by tortoises in all seasons, despite its relative rarity. Ruscus aculeatus berries may be particularly valuable to tortoises that are about to enter hibernation; hence, T. hermanni habitat should be managed to maintain this important plant species.
... Rugiero, M. Capula, L. Luiselli, personal observation). Tortoise egg shells were eaten during autumn by semiwild T. hermanni in France (Huot-Daubremont, 1999) as well by Gopherus agassizii in the United States (Esque and Peters, 1994), but we found no evidence of this at our study area, although such egg shells are typically available (L. Rugiero, M. Capula, L. Luiselli, personal observation). ...
Herbivory is the dominant feeding strategy in tortoises, and dietary shifts are common in response to changes in resource availability. We conducted the first large-scale study of the diets of wild Hermann's Tortoises (Testudo hermanni hermanni) and found that the study population in central Italy was strictly herbivorous. The tortoises ate primarily legume leaves and grasses in the spring, and switched to flowers and unripe fruit of Ruscus aculeatus as these became available in the autumn. There were no significant differences between the diets of males and females. Although tortoise diets included both rare and abundant plant species, they consumed abundant plant species in a higher proportion than those species occurred in the study area. However, some rare plants made up relatively large fractions of the diet, and one of the few nonnative plants (Conyza canadensis) at the study area was eaten frequently by tortoises in all seasons, despite its relative rarity. Ruscus aculeatus berries may be particularly valuable to tortoises that are about to enter hibernation; hence, T. hermanni habitat should be managed to maintain this important plant species.
... Interestingly, the most common vertebrate remains we observed were carapacial bones of other box turtles. Osteophagy is widespread among terrestrial chelonians (Walde et al. 2007), and bones are an excellent source of calcium and phosphorus (Esque and Peters 1994)—nutrients that are poorly represented in the shallow, leached sandy soils typical of pine rockland forest (Snyder et al. 1990). The large size of most carapacial bones indicated they were from adult turtles and almost certainly consumed as carrion; however, one scat contained costal bones from a hatchling or small juvenile. ...
We studied the wet-season food habits of adult Terrapene carolina bauri (Florida Box Turtles) on National Key Deer Wildlife Refuge, Big Pine Key, FL. Feces were collected from 112 Box Turtles (64 females, 48 males) and analyzed to determine diet. Based on percent occurrence (number of samples in which a particular dietary item occurs divided by the sample size), terrestrial gastropods and fleshy fruits were the principal foods consumed. Lesser amounts of leafy vegetation were recovered, whereas insects and other invertebrates, vertebrates (most likely consumed as carrion), and fungi composed a minor portion of the diet. The occurrence of deer feces in a single scat constitutes the first report of coprophagy by box turtles. Our fecal analysis provided little evidence of dietary specialization by either sex, which is consistent with earlier descriptions of box turtles as generalist omnivores. Male Florida Box Turtles were significantly larger than females, but there was no indication that larger body size in males provides access to an expanded resource base; the near-complete dietary overlap between the sexes suggests they consume the same foods.
... The animals are known to depend on Opuntia for water during prolonged drought (Fritts, 1983). It is possible that tortoises and other arid-dwelling reptiles that lack su¤cient mobility to search for free water are adapted to cope with physical and chemical substances of plants by various means including: (a) excretion of some of the substances (oxalates) in crystalline form as uric acid crystals rather than urea (Milton, 1992), which requires some water for its elimination (Pritchard, 1967); (b) ingesting supplementary minerals to nullify the toxic e¡ects of secondary compounds in food plants (Esque & Peters, 1994), and (c) consuming small amounts of the various items to avoid lethal doses. The succulent diet of leopard tortoises in the Serengeti may have implications for the ranging patterns of tortoises there. ...
Sighting frequency and food habits of the leopard tortoise (Geochelone pardalis) were studied in northern Tanzania from October 1993 to June 1996. Sighting frequency varied significantly between protected (0.22 mhr−1) and unprotected sites (0.59 mhr−1), and between Arusha (0.27 mhr−1) and Serengeti sites (0.56 mhr−1).
The tortoise diet comprised mostly plants (97.8%) and rarely inorganic matter (2.2%). A total of 47 plant species from 21 families was eaten. Forbs made up 74.5% of the plant items and monocots the remainder. Succulents contributed 51.0% of the plant items eaten while grasses and legumes accounted for 16.8% and 13.5%, respectively. Some plant items were eaten more frequently than their occurrence in the habitats.
On a étudié la fréquence des observations et les habitudes alimentaires de la tortue léopard (Geochelone pardalis) dans le nord de la Tanzanie d'octobre 1993 à juin 1996. La fréquence des observations variait significativement entre les sites protégés (0,22 m/h) et les sites non protégés (0,59 m/h), et entre les sites d'Arusha (0,27 m/h) et ceux du Serengeti (0,56 m/h).
Le régime alimentaire des tortues comprenait surtout des plantes (97,8%) et, rarement, des matières inorganiques (2,2%). On a rapporté un total de 47 espèces végétales appartenant à 21 familles. Les monocotylédones représentent 74,5% des éléments végétaux et les monocotylédones, le reste. Les plantes succulentes comptent pour 51,0% des plantes broutées tandis que les plantes grasses et les légumineuses représentent respectivement 16,8% et 13,5%. Certaines plantes étaient broutées plus souvent qu'elles n'étaient présentes dans les habitats.
... It has been proposed that during egg production , female tortoises may become deficient in calcium, as can birds (Klasing, 1998). Adult female tortoises have been observed to forage on calcium-rich material such as caliche soil and bone (Marlow and Tollestrup, 1982; Esque and Peters, 1994 ), suggesting a high dietary requirements for this mineral. Texas Tortoises (pherus berlandieri) have a high reproductive rate relative to the other Gopherus, and it has been speculated that the high mortality of females in this species could be a result of bone loss caused by this increased calcium output (Hellgren et al., 2000). ...
Exotic plants can make up a major component of the diet for some Desert Tortoises (Gopherus agassizii) in the Mojave Desert. If introduced plants differ nutritionally from native plants, they may influence the growth and welfare of young tortoises. Minerals available from a native grass (Achnatherum hymenoides), an introduced grass (Schismus barbatus), a native forb (Malacothrix glabrata), and an introduced forb (Erodium cicutarium) were measured for juvenile Desert Tortoises voluntarily eating single-species diets. We offered tortoises weighed amounts of chopped foods daily for similar to 130 days (dry grasses; summer diet) or similar to 90 days (green forbs; spring diet). Orts and feces were collected daily and dried to constant mass, and calcium, phosphorus, and magnesium contents of food and feces were measured and used to calculate nutrient digestibilities. Overall, tortoises gained more minerals from forbs than from grasses. Tortoises lost small but significant amounts of phosphorus when eating grasses, which may have contributed to observed decreases in tortoise mass and shell volume on grass diets. There were few nutritional differences between native and exotic forbs or between native and exotic grasses. Comparisons of nutrient availability to estimated requirements for growth by juveniles and for egg production by adult females suggest that phosphorus is more limiting than calcium or magnesium and that calcium may pose a significant osmotic challenge for excretion in this desert species. Management practices that promote availability of forbs could increase growth rates and shell ossification, which would enhance predator resistance of juvenile tortoises.
... Research on desert tortoises has focused on 2 nutrient wisdom hypotheses (Minnich 1977(Minnich , 1979Medica 1977, 1986;Jarchow 1984;Oftedal and Allen 1996;Oftedal 2002;Oftedal and others 2002). The first hypothesis centers on balancing the need for calcium, phosphorus and magnesium for bones, shell and eggs (Esque and Peters 1994). Additionally, calcium is required for maintenance of osmotic balance in cells, blood coagulation, nerve transmission and muscular activity (Despopoulos and Silbernagl 1991). ...
Many of the threats to the persistence of populations of sensitive species have physiological or pathological mechanisms, and those mechanisms are best understood through the inherently integrative discipline of physiological ecology. The desert tortoise was listed under the Endangered Species Act largely due to a newly recognized upper respiratory disease thought to cause mortality in individuals and severe declines in populations. Numerous hypotheses about the threats to the persistence of desert tortoise populations involve acquisition of nutrients, and its connection to stress and disease. The nutritional wisdom hypothesis posits that animals should forage not for particular food items, but instead, for particular nutrients such as calcium and phosphorus used in building bones. The optimal foraging hypothesis suggests that, in circumstances of resource abundance, tortoises should forage as dietary specialists as a means of maximizing intake of resources. The optimal digestion hypothesis suggests that tortoises should process ingesta in ways that regulate assimilation rate. Finally, the cost-of-switching hypothesis suggests that herbivores, like the desert tortoise, should avoid switching food types to avoid negatively affecting the microbe community responsible for fermenting plants into energy and nutrients. Combining hypotheses into a resource acquisition theory leads to novel predictions that are generally supported by data presented here. Testing hypotheses, and synthesizing test results into a theory, provides a robust scientific alternative to the popular use of untested hypotheses and unanalyzed data to assert the needs of species. The scientific approach should focus on hypotheses concerning anthropogenic modifications of the environment that impact physiological processes ultimately important to population phenomena. We show how measurements of such impacts as nutrient starvation, can cause physiological stress, and that the endocrine mechanisms involved with stress can result in disease. Finally, our new syntheses evince a new hypothesis. Free molecules of the stress hormone corticosterone can inhibit immunity, and the abundance of "free corticosterone" in the blood (thought to be the active form of the hormone) is regulated when the corticosterone molecules combine with binding globulins. The sex hormone, testosterone, combines with the same binding globulin. High levels of testosterone, naturally occurring in the breeding season, may be further enhanced in populations at high densities, and the resulting excess testosterone may compete with binding globulins, thereby releasing corticosterone and reducing immunity to disease. This sequence could result in physiological and pathological phenomena leading to population cycles with a period that would be essentially impossible to observe in desert tortoise. Such cycles could obscure population fluctuations of anthropogenic origin.
Upper Paleolithic sites in southwestern France attributed to the Upper Gravettian and the Solutrean yielded sub spherical gravels with a highly shiny appearance that have intrigued researchers since the 1930s. In this work, we analyze specimens from five sites, including the recently excavated Solutrean site of Landry, to establish whether their presence in archaeological layers and peculiar aspect are due to natural processes or human agency. We study the spatial distribution of gravels at Landry and submit archaeological gravels from the five sites, natural formations, Landry sediment sieving, and polishing experiments with a rotary tumbling machine to morphometric, colorimetric and textural analyses. Spatial analysis reveals that shiny gravels are concentrated at Landry in the areas in which the remainder of the archaeological material was recovered, which supports their anthropogenic origin. Morphometric and colorimetric analyses demonstrate archaeological lustrous gravels are homogenous in size and color. Textural analysis of the gravel surfaces demonstrates that the shine on the archaeological specimens is not observed on natural gravels and gravels found at Landry outside anthropogenic concentrations. They show strong similarity with gravels polished by putting them in a tumbling machine with the barrel covered by skin. Our results suggest that the lustrous gravels found at the five sites result from deliberate selection and their shiny appearance is the probable consequence of prolonged contact with a soft material such as animal skin. Ethnographic accounts indicate that these gravels may have been used for magico-religious ritual purposes (charms, sorcery, divination etc.), in games, as elements of musical instruments, and as items serving other social and personal purposes. We argue that these objects reflect a cultural innovation emerged during the Gravettian and continued into the Solutrean.
Tortoises (land turtles) are familiar animals and are generally assumed to be strict herbivores. Their ecological roles are most obvious in giant tortoise species which, due to their size and local abundance, play major roles as keystone species and ecosystem engineers1, 2, 3. In the Galápagos and Seychelles islands these species are known to play major roles as the islands’ largest herbivores, with exceptionally high biomass and consuming up to 11% of primary production¹. In addition they act as ecosystem engineers, dispersing seeds, breaking vegetation and eroding rocks². However, as slow-moving poikilotherms most people assume their behaviour to be simple. Here we present video evidence of a Seychelles giant tortoise (Aldabrachelys gigantea) attacking a tern chick and pursuing it along a log. Finally the tortoise killed the chick and was observed to eat it. Other tortoises in the same area have been seen making similar attacks, although those were not fully documented. We believe that the exceptional combination of a tree-nesting tern colony with a resident giant tortoise population has created conditions leading to systematic hunting of birds by several individual tortoises; an entirely novel behavioural strategy for any tortoise species.
Video abstract
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A major conceptual gap in taste biology is the lack of a general framework for understanding the evolution of different taste modalities among animal species. We turn to two complementary nutritional frameworks, biological stoichiometry theory and nutritional geometry, to develop hypotheses for the evolution of different taste modalities in animals. We describe how the attractive tastes of Na, Ca, P, N and C containing compounds are consistent with principles of both frameworks based on their shared focus on nutritional imbalances and consumer homeostasis. Specifically, we suggest that the evolution of multiple nutritive taste modalities can be predicted by identifying individual elements that are typically more concentrated in the tissues of animals than plants. Additionally, we discuss how consumer homeostasis can inform our understanding of why some taste compounds (i.e., Na, Ca and P salts) can be either attractive or aversive depending on concentration. We also discuss how these complementary frameworks can help to explain the phylogenetic distribution of different taste modalities and improve our understanding of the mechanisms that lead to loss of taste capabilities in some animal lineages. The ideas presented here will stimulate research that bridges the fields of evolutionary biology, sensory biology and ecology.
Report of the first observation of scavenging behaviour in the population of Testudo hermanni boettgeri that
has been monitored for six years in the village Kunovica near the city of Niš in Serbia. On 31 May 2015 at
10:18 a.m., the adult tortoise was observed while eating a dead European green lizard (Lacerta viridis).
La Tortue d’Hermann est l’unique chélonien terrestre de la faune française, on distingue actuellement deux sous espèces : Testudo hermanni boettgeri localisée dans les Balkans et Testudo hermanni hermanni situé sur la frange méditerranéenne occidentale. L’aire de répartition de l’espèce est particulièrement fragmentée, notamment dans sa partie occidentale. La population de Testudo hermanni hermanni varoise est la plus importante population de France continentale et une des plus importantes d’Europe occidentale. Les menaces à l’origine de la disparition de l’espèce portent à la fois sur son habitat et sur les individus qui composent les populations sauvages. Sa conservation passe avant tout par le maintien des habitats et des populations existantes (IUCN, 1995 ; Tenhumberg et al., 2004) ; la gestion de ses habitats en vue de leur optimisation et la limitation des facteurs de mortalité. Chaque année, une cinquantaine de spécimens de Tortue d’Hermann issus d’opérations de sauvetage ou trouvés blessés, sont ramenés par des particuliers et divers organismes à la SOPTOM (le seul organisme reconnu comme centre de soins et d’élevage de la Tortue d’Hermann). La plupart ne peuvent pas être relâchés sur leur lieu d’origine ni même gardés à la SOPTOM. Toutefois, nous disposons d’une occasion unique de réinsérer un petit nombre d’individus sauvages dans le milieu naturel de façon expérimentale, afin de mieux connaître leur devenir. Il est unanimement reconnu que si une population se trouve sur un site voué à être détruit, tous les individus qui ne sont pas transloqués et qui sont donc tués, représentent une perte de matériel biologique/génétique irrémédiable. La translocation est donc dans ce cas une opération qui peut s’imposer (Burke, 1991). Un des objectifs à long terme du Plan National d’Actions de l’espèce est l’accroissement de l’aire actuelle de répartition et la diminution de la fragmentation (Cheylan et al., 2008). Une fois les espaces naturels nécessaires protégés, des méthodes de réinsertions pourraient permettre par exemple de renforcer des populations dans le cas de zones précédemment incendiées. Néanmoins, les techniques et conditions favorables à la réussite de l’opération sont mal connues chez les (Germano & Bishop, 2009) et cette espèce. Les relâchers dans le Var n’ont jamais bénéficié de suivis rigoureux contrairement à certains réalisés en Espagne où une comparaison reste limitée en raison de problématiques locales différentes (ex : habitat différent). Il faut donc au préalable avoir réalisé des progrès importants sur la sélection et la préparation des spécimens, les méthodes pour les insérer dans le milieu en optimisant leurs chances de survie, les méthodes de gestion du milieu pour le maintenir en état d’accueillir durablement dans de bonnes conditions une population, ainsi que sur les suivis postopératoires qui doivent permettre des retours d’expérience tangibles et validés scientifiquement. Faisant suite au programme européen FEDER « Des tortues et des Hommes », le programme LIFE+ Tortue d’Hermann intitulé « Vers une gestion intégrée favorable à la tortue terrestre dans le Var Création d’outils pour les gestionnaires d’espaces naturels en Europe » a débuté en janvier 2010 et se terminera en décembre 2014. Déclinaison du Plan National d’Action en faveur de la Tortue d’Hermann, il vise la protection de l’espèce via la mise en oeuvre de 34 actions réparties autour de trois axes : l’amélioration et la restauration des habitats, la sensibilisation et l’information, le transfert d’expérience. Le présent cahier des charges a été réalisé dans le cadre de ce programme LIFE+. Il s’inscrit pleinement dans les objectifs du Plan National d’Action. Bien que dans le passé divers auteurs aient mentionné que les projets de translocations de reptiles et amphibiens ont peu de succès (Burke 1991 ; Dodd & Seigel, 1991 ; Guyot, 1996 ; Reinert 1991) ; plus récemment, d’autres affirment qu’au contraire le succès de telles opérations est bien supérieur à ce qui était pensé (Germano & Bishop, 2009 ; Griffiths & Pavajeau, 2008). Les principales causes d’échec sont associées au phénomène de « homing », aux fortes dispersions et à un habitat inadapté (Germano & Bishop, 2009 ; Tuberville, 2008). Une opération de cette envergure représente une épreuve pour les individus puisqu’ils subissent des perturbations stressantes inévitables (manipulation, transport, libération dans un environnement inconnu). Celles-ci peuvent avoir des effets négatifs sur le long terme comme l’augmentation du stress et la réduction de la survie (Letty et al., 2007 ; Chipman et al., 2008). Il est donc nécessaire de déterminer quels sont les principaux facteurs impliqués dans l’échec ou la réussite d’une réinsertion en comparant différentes méthodologies de relâcher (Armstrong & Seddon, 2007 ; Sutherland et al., 2010). Le recours à l’expérimentation permet de comparer directement entre elles les alternatives possibles dans le cadre d’une translocation, et d’en déduire la solution maximisant les chances de réussite (Marchandeau et al., 2000). De nombreuses études permettent d’avoir une meilleure idée du déterminisme général de la réussite d’une translocation, laquelle reste spécifique et demeure complexe étant donné le grand nombre de facteurs impliqués. Il est ainsi possible de favoriser la réussite d’une telle opération en sélectionnant par exemple le type d’individus, le site de relâcher, ou en gérant le milieu d’accueil et mettant en oeuvre des méthodes de relâcher adéquates (Letty et al., 2007). L’intérêt de cette réinsertion expérimentale (ou relâcher d’individus sauvages) est de déterminer les facteurs qui sont favorables à la sédentarisation des animaux relâchés et à plus long terme, leur survie et leur reproduction. Pour cela, deux sites ont été retenus (le Plateau du Lambert dans le Massif des Maures et le Vallon de Saint- Daumas en Plaine des Maures) afin de tester l’influence de deux paramètres sur la dispersion, la sédentarisation, la survie et les paramètres physiologiques après relâcher : la saison et le passage ou non par un enclos d’acclimatation. Le choix des sites, des spécimens et des méthodes pour effectuer cette réinsertion nécessite la mise au point de protocoles visant à maximiser les chances de succès de ces opérations. Il est donc important d’établir comment choisir les sites ayant vocation à accueillir des individus, de définir des protocoles précis et de les réactualiser à la lumière de l’expérience acquise (Sutherland et al., 2010). Le développement des outils sanitaires et génétiques indispensables pour garantir la qualité des animaux (Cunningham, 1996 ; Chipman et al., 2008), le développement de la « biotélémétrie » depuis les années 90’, et l’expérience acquise depuis plus de 15 ans par la SOPTOM sur les plans sanitaires et scientifiques, permettent aujourd’hui d’envisager de telles opérations dans des conditions compatibles avec les exigences internationales en la matière. Les différentes études (utilisation des habitats, modèles de mouvement, stratégies thermorégulatrices, réactions physiologiques) permettront de fournir des informations d’une grande valeur quant aux réponses des individus à un nouvel environnement. La première phase de cette opération concerne le choix des sites, des spécimens, des méthodes et des protocoles visant à maximiser les chances de son succès. Un des deux sites expérimentaux concerne la Réserve Naturelle Nationale de la Plaine des Maures, l’autre est localisé en forêt domaniale. Une autorisation préfectorale ou une déclaration préalable est requise après avis de plusieurs instances (CNPN, Comité consultatif et Conseil scientifique de la Réserve Naturelle Nationale de la Plaine des Maures). Pour mener à bien ces opérations, différentes validations administratives sont nécessaires (autorisation de transport, demande de dérogation pour la capture etc.). Une étude de faisabilité centrée sur la biologie de l’espèce a donc été entreprise (origine des individus, génétique, etc.). Parallèlement, l’environnement du relâcher (qualité des habitats, causes de déclins, protection à long terme de la zone) a été évalué. La seconde phase de l’évaluation concerne la phase de relâcher à proprement parler et la phase de suivi (comportemental et démographique) de l’ensemble des individus. Ce suivi est nécessaire à long terme, tout comme la protection des habitats. Enfin, les résultats de l’opération doivent être diffusés et publiés afin d’ouvrir des voies d’expérimentation (IUCN, 1998 ; Mihoub, 2009 ; Sutherland et al., 2010). Lors du montage du projet, il est indispensable de s’assurer de l’absence ou non de l’espèce. Enfin, les espèces transloquées constituent dans de nombreux cas des symboles spectaculaires pour le grand public. Ces types de programmes représentent une conservation active et constructive permettant de mieux faire accepter d’autres mesures souvent plus restrictives. Ils constituent un moyen de sensibilisation et d’éducation visant à la conservation du patrimoine naturel.
FORWARD The goal of this book is threefold. First, it is to provide a detailed reference for those involved in resource management of the amphibians and reptiles of Joshua Tree National Park. Second, it will provide a source of information to the rangers and interpreters enabling them to answer questions from the general public on our amphibian and reptile resources.Third, it is hoped this will be a valuable reference source for all the friends of Joshua Tree National Park who are seriously interested in this aspect of the park's fauna. A word should be said about the maps accompanying each species discussion.There is a generalized map, courtesy of ICE, UC Davis, showing the overall range of the animal in California.This map pretty much ignores local range phenomena.There is a map showing the localities of all museum specimens and verified sightings of each species. Although this type of map is often favored by scientists for its exactness, it is highly biased by collecting techniques. For example, it indicates the most favored habitat of snakes is roads, an obvious artifact of observation techniques. This map also shows my prediction of the actual range of the species in the park. I have probably erred on the conservative side in showing the range of most species. Future observations in less traveled regions of the park will help refine our knowledge of most species' distribution.
Tooth marks on sandstone in an area of the Cerrado Biome are reported, indicating geophagy. The tooth marks were found on reddish sandstones cropping out in a pasture environment with typical components of the Cerrado Biome, in the Municipality of Campina Verde, Triângulo Mineiro region (west Minas Gerais State, Brazil). Studies have shown that the soil of the Cerrado is acid, with a low concentration of nutrients and minerals (also present in the plants living on this environment), which usually produce an alimentary deficiency in herbivorous animals. Therefore, these tooth marks indicate geophagy, in order to extract extra minerals from these sandstone levels, which have a high concentration of calcium carbonate and iron. The tooth marks consist of two parallel concave grooves and a medial prominent crest which results from the action of the incisors of mammals. Although the identification of the gnawing species for these sets of tooth marks are estimates at best, after wide comparisons we tentatively suggest that the tooth marks are most likely the result of the action of the incisors of rodents, such as Dasyprocta or Coendou. INTRODUCTION The ingestion of soil or geophagy is a regular activity of wild animals, especially those with frugivorous and
Turtles provide excellent models for studies of life history strategies, but terrestrial species are underrepresented in these analyses. We present a life table and demographic parameters of an inland population of Gopherus berlandieri to contribute to the study of evolution of turtle life histories. Data were gathered during a mark-recapture and radiotelemetry study in the Tamaulipan Biotic Province in southern Texas, USA. A total of 835 individuals were captured, measured, and their ages estimated. Females matured at 131 mm carapace length at an average of 5 yr of age (range: 4-8 yr). Clutch size, as determined by ultrasound, averaged 2.07 ± 0.15 eggs (n = 49). No nests were found, and we estimated clutch frequency with a quadratic model to be 1.34 cultches· female-1· year-1. Survival was estimated from age frequency regression and telemetry. Annual male survival (0.828, 0.834) differed (P 0.05) among years, and population estimates did not differ among years; therefore, we constructed a life table under the assumption of a stationary population (r = 0.0). Under this assumption, survival from nest to age 4 yr must be at least 0.245, with hatchling survival of 0.528 to maintain a stationary population. Gopherus berlandieri matured at an earlier age, had smaller clutch sizes, and exhibited lower rates of female survival than other Gopherus species. We propose a physiological mechanism for lower female survival that implies trade-offs among egg size, subsequent hatchling survival, and female health. We maintain that high hatchling survival is necessary for population persistence. In an evolutionary context, we theorize that the selective advantages of small size and the life history strategies of G. berlandieri have been and are critical to its persistence.
La población de tortugas del desierto (Gopherus agassizii) ha experimentado un decremento significativo en las últimas décadas. Por consiguiente existe un gran número de restos de tortuga en el terreno. Documentamos un fenómeno de osteofagia por parte de las tortugas del desierto, en particular el consumo de huesos de las tortugas muertas. Parece ser que las tortugas del desierto tienen un apetito por sustancias de alto contenido en calcio. Que tengamos conocimiento, ésta es la primera vez que se documenta el consumo de desechos esqueléticos conespecíficos.
1. Mixed diets are common in vertebrates, and may be explained by several hypotheses including the availability and nutrient composition of different types of food, the presence of toxins, associative effects in digestion or non‐nutritional factors such as foraging efficiency or predation. This study investigated diet mixing in the omnivorous tortoise Kinixys spekii , which consumes fungi, vascular plants and invertebrates in the field. Tortoises feeding ad libitum on these three diets had a higher intake of digestible energy from fungi than from leaves or millipedes (69, 42 and 31 kJ kg –1 day –1 respectively).
2. Tortoises offered pairwise combinations of foods chose mostly that giving the highest rate of digestible energy intake; 73:27 fungi:leaves, 92:8 fungi:millipedes, and 91:9 leaves:millipedes, by wet mass. Nevertheless, some of the food giving a lower energy intake was eaten.
3. Tortoises feeding ad libitum on a mixed diet (90:10 leaves:millipedes by wet mass) had similar food intake rates and gut retention times to those feeding on leaves. The digestibility of energy was lower for the mixed diet than for diets of leaves or millipedes; a significant negative associative effect.
4. The rate of intake of dry mass did not differ between the three pure diets, the selected diets, or the mixed diet, though this was about 50% higher at 30 °C than outdoors, and represents a limitation due to digestion or appetite. Intake of other foods decreased the intake of dry mass of the preferred food, fungi, by an equal amount.
5. The three foods had similar levels of protein as a proportion of dry mass, partly because of the high ash content of millipedes. The digestibility of protein was higher in millipedes, but the rate of intake of total and of digestible protein was in the same rank order as that of energy (fungi > leaves > millipedes). Although the quantity of protein provided no nutritional explanation for inclusion of lower energy foods in the diet, the quality of protein may differ between food types.
6. Herbivorous tortoises are known to face limitations from calcium and sodium balance. Fungi had a lower level of calcium and a lower ratio of sodium:potassium than leaves; millipedes had values higher than leaves. Foods that are non‐optimal in terms of energy are probably included in the diet for micronutrients such as minerals or essential amino acids.
This paper studies a class of source coding problems that combines elements of the CEO problem with the multiple description problem. In this setting, noisy versions of one remote source are observed by two nodes with encoders (which is similar to the CEO problem). However, it differs from the CEO problem in that each node must generate multiple descriptions of the source. This problem is of interest in multiple scenarios in efficient communication over networks. In this paper, an achievable region and an outer bound are presented for this problem, which is shown to be sum rate optimal for a class of distortion constraints.
Twenty-four juvenile leopard tortoises were divided into four groups of six; one group was fed a basic low-calcium feed for six months, and the other three groups were fed the same basic diet supplemented with one, three and nine times the amount of calcium recommended as a supplement to the diet of reptiles. The animals' bone mineral content and bone mineral density were estimated by dual energy x-ray absorptiometry, and blood samples were taken at the start and at the conclusion of the study. One tortoise from each group was examined postmortem. There was a clear depletion of calcium in the body of the tortoises receiving no calcium supplement, and the shell of the tortoises receiving the recommended calcium supplement did not calcify to the extent expected. The tortoises that received three times the recommended calcium supplementation had the highest growth rate and were thriving. However, metastatic calcifications were observed postmortem in the two groups that were given the highest doses of calcium.
Geophagia and osteophagia were a common feature of the feeding routine of the southern giraffe (Giraffa camelopardalis giraffa) during the months from April to November. Osteophagia occurred at approximately the same frequency in adults and sub-adults. However, geophagia was primarily exhibited by sub-adult giraffe. The acts of geophagia and osteophagia were carried out by an identical set of search, accusative, and gratification behaviors. Kidney stones and associated cortical and medullary lesions were observed in 29 percent of the 75 adult and sub-adult giraffe sampled. The frequency of osteophagia and geophagia indicated that these forms of pica were important nutritional supplements. The occurrence of pica or renal calculi can be used as an indicator of nutritional stress and in the assessment of habitat deficiencies.
Thesis (Ph. D.)--Syracuse University. Vita. Includes bibliographical references (leaves 390-409). Microfilm. s
Two groups of porcupines (Hystrix africaeaustralis) were maintained for a period of 23 weeks on a diet containing high levels of Ca2* and PO4. Parallel to this another two groups were maintained on a diet containing low levels of these minerals. All groups were provided with defatted bovine bones during weeks 18 to 23 of the experiment. Plasma Ca2+ and P04 levels were determined at intervals throughout the experiment. The amount of osteophagia that occurred was determined by weighing the bones at two-weekly intervals. No significant differences in the plasma Ca2+ and P04 levels of the porcupines on the high diet and those on the low diet were detected. Osteophagia by porcupines on the low diet was twice that of porcupines on the high diet.Twee groepe ystervarke (Hystrix africaeaustralis) is vir 23weke 'n dieet bevattende hog vlakke Ca2+ en PO4 gevoer en twee ander groepe 'n dieet bevattende hoë vlakke Ca2+ en PO4. Tydens weke 18 tot 23 van die eksperiment is alle groepe van ontvette beesbene voorsien. Plasma Ca?+ en PO4 is op verskeie stadiums in die verloop van die <5 eksperiment vasgestel. Die hoeveelheid osteofagie wat plaasgevind het is elke twee weke bepaal deur die bene te weeg. Geen betekenisvolle verskille in die plasma Ga2+-of 3 PCVvlakke van die ystervarke op die hoë dieet en die op die lae dieet is gevind nie. Ystervarke op die lae dieet het twee keer soveel osteofagie getoon as ystervarke op die hoë dieet.
Leopard tortoise Geochelone pardalis faeces collected in rocky habitats in the southern Karoo contained at least 75 species of grasses, succulents and forbs belonging to 26 plant families. Soft, green plants were broken down by digestion but twigs, thorns and fibrous naterials were not digested. Flowers, fruits and seeds made up 67% of 356 identified plant fragments. Germination trials demonstrated that leopard tortoises could disperse viable seeds of Aizoaceae, Chenopodiaceae, Crassulaceae, Cyperaceae, Fabaceae, Poaceae and Scrophulariaceae.
Ants and beetles were the primary foods of 60 eastern fence lizards and 25 desert spiny lizards during spring and summer in southwestern Colorado. Lepidopterous larvae and grasshoppers comprised most of the food of 49 western whiptails. Although there was some seasonal change in the diet of all species, there was little qualitative difference in diet between sexes or age groups. Sand grains and small pebbles found in stomachs may act as abrasives which macerate hard exoskeletons. Female fence and desert spiny lizards had more food in their stomachs during the spring months than did males, probably to meet the metabolic requirements of egg-laying. Assuming that these species fill the stomach twice daily, that stomach capacity is twice that of the mean weight of the stomach contents, an assimilation efficiency of 2/3, and an energy value of 5363 cal/g, the energy assimilated by an adult eastern fence lizard (15 g), western whiptail (22 g), and desert spiny lizard (30 g) is estimated as 0.83, 1.57, and 2.17 kcal/day (55, 71, and 72 cal/g/day), respectively.
The nutritional significance of invertebrate foods in the diet of breeding hen waterfowl during the period of egg formation is discussed. Proximate, elemental, and amino acid analyses of the principal foods consumed by hen pintails (Anas acuta) during the nesting season indicate the animal foods selected are rich sources of protein and calcium, whereas plant foods tested were low in protein, particularly in the essential amino acids lysine and methionine. Calcium content in major plant foods was far below needs considered adequate for egg production. Studies with penned, hand-reared pintails indicated the inadequacy of wheat as the dietary staple during breeding. Egg production among pairs on the control diet differed significantly from that of the wheat plus oystershell (P < 0.05) and wheat (P < 0.01) diets. Pairs fed wheat during the breeding period experienced nearly complete reproductive failure. The poor reproductive response among hens on the diet of wheat plus oystershell suggested that calcium availability was not the only factor limiting productivity.
A group of Presbytis rubicunda (red leaf) monkeys was observed for 13 months in the lowland dipterocarp forest at Sepilok, Sabah, northern Borneo. The main items in the animals' annual diet were young leaves, seeds, whole fruits, and flowers. Large, dry seeds made up over 85 percent of the diet in 2 months, but young leaves, fruits, and flowers were generally eaten when large seeds were unavailable. Soil eating was observed on nine occasions. The soil was always collected by breaking lumps off termitaria and never from the forest floor. Analytical comparison of termite-mound and forest-floor soils showed that both had similar clay contents, but the termitaria soil was of higher pH and had higher levels of the main cationic nutrients and lower levels of labile aluminium. The geophagy was observed at times when different items predominated in the diet. It was noted when the dietary items were highly digestible and contained few tannins, indicating that geophagy need not be primarily for tannin absorption, although it might have helped to absorb toxins. The main benefits are more likely to be the alleviation of digestive disorders, such as forestomach acidosis, and the supplementation of mineral nutrient uptake in a generally oligotrophic environment. There is no reason why geophagy should have a single function; it may serve different functions at different times.
Small rodent interest in calcium, sodium and phosphorus was investigated by means of impregnated wooden sticks placed on abandoned fields and clearcuts in areas with cyclic and non-cyclic populations during two-three years, covering one rodent peak and one crash. Sodium was generally attractive but with pronounced variations in attack rates related to rodent densities; calcium was attacked only in very cyclic populations; and at least the specific compound of P offered was repellent. Sodium was taken in locally varying amounts on abandoned fields and mainly in summer, while sodium consumption on clearcuts was closely related to densities and cyclicity and was mainly taken in winter. Sodium consumption correlated best with the abundance of the folivorous Microtus agrestis but increased more rapidly than total rodent numbers. In experiments, sodium consumption by M. agrestis was much higher when the animals were given growing plants than seed-based food, indicating a possible role of sodium in detoxification of plant secondary metabolites. Granivorous rodent species did not consume sodium when kept on their normal food. Mature or senescent vegetation as food did not cause M. agrestis to consume more sodium than still-growing vegetation, so sodium concentrations in plant food do not appear to be limiting. Neither did supply of growing vegetation or sodium affect growth rates of young M. agrestis. Sodium consumption increased at habitat fertilization with sodium and at growing vegetation, which indicated some intricate positive feedback mechanism. Sodium and calcium availability appear as two potentially limiting factors at rodent peak densities.
Scute annuli, produced during the first 20-25 yr of initial, rapid growth, can be used to age young desert tortoises (Gopherus agassizii). Regression analysis did not show a difference between age and the number of scute rings counted on the carapace for 13 desert tortoises from the Nevada Test Site, although ring counts on most individuals were 1-2 rings less than age. The number of scute rings exactly matched the ages of six desert tortoises raised in captivity. No significant differences were found between the estimated length of the plastron based on selected scute rings and the actual length of the plastron as recorded 7-12 yr previous to my handling of individual tortoises from the Nevada Test Site, which supports the accuracy of the ring counts. The number of bone rings did not differ significantly from the number of scute rings for 16 preserved desert tortoises. Growth histories of individual tortoises can be determined from scute annuli.
During 20 months of field work in Wilpattu National Park in Sri Lanka (8°30, N, 80°l'E) I recorded 102 instances of osteophagia by Axis deer (Axis axis). I found no seasonal variation in the occurence of osteophagia. All age-sex classes ate bones and antlers, but among adult males, velvet antlered ones did it significantly more often than hard antlered males. Also, among velvet males, the ones with fully grown antlers, i.e. antlers in the process of mineralization, ate bones or antlers significantly more often than other velvet males. Since antler growth is not seasonal in Wilpattu these results cannot be explained by a seasonal variation in the mineral content of the diet. This is therefore a clear demonstration that osteophagia is done in response to a greater mineral need during antler growth and mineralization.
We assigned ages of 1-4 yr to 15 desert tortoises originally marked in Rock Valley, Nevada, between 1963 and 1965, at which time their plastron lengths ranged from 47-74 mm. Continued measurements of these tortoises enabled us to estimate mean body sizes of tortoises from 1-26 yr of age. Growth of males and females over this period did not differ significantly. Tortoises grew to plastron lengths of 100 mm in 6-7 yr, to 130 mm in 10-11 yr, to around 150 mm in 13-14 yr and were 215 mm long at estimated ages of 24 yr. Four females 23-24 yr old were X-rayed in 1985; three had clutches of 4-5 eggs. If Rock Valley female tortoises are sexually mature at the same body size as those in eastern San Bernardino County, California, sexual maturity is attained at an age of 17-20 yr.
Scat analysis and foraging observations were used to determine the diet of a gopher tortoise, Gopherus polyphemus, population in a sandhill community in west-central Florida. Vegetation was assessed to determine plant availability. The bulk of the gopher tortoise diet was composed of dominant plants in the habitat, although the tortoises were selective with respect to most plant genera. Tortoises ingested many plant taxa, with Poaceae, Asteraceae, Fabaceae, Pinaceae, and Fagaceae being the most frequent components of the diet. Sixty-eight genera from 26 families were identified in the diet. In the scat, the most common genus identified was Aristida and the most common family was Poaceae. Other common genera were Pinus, Quercus, Galactia, Cnidoscolus, Tillandsia, Pityopsis, and Richardia. Insects and charcoal were often present in scats. Comparing the overall diet indicated no difference between sexes or age groups. Comparing diet with respect to individual plant taxa indicated that young tortoises ingested significantly less Poaceae and fewer plants with external defense mechanisms (Rubus, Cnidoscolus) than did adults. Non-plant items may be a significant source of nutrients. Jacobs' Electivity Index indicated that the tortoises preferred Galactia and Tephrosia, Cnidoscolus, Pinus, Quercus, Vaccinium, Richardia, and Rubiaceae. Some of the most common plants in the habitat (Poaceae, Aristida, Asteraceae), were selected at or below the proportions at which they were available. Feinsinger's Proportional Similarity Index indicated the foraging habits of the gopher tortoise lie midway between a specialist and generalist.
Six female desert tortoises, Gopherus agassizii, were observed to consume the soil of a ‘limelayer’ on an exposed site, or to excavate through the overburden where the layer was not exposed. The calcium content of soil from the layer mined was significantly higher than that of adjacent surface soil. All geophagous episodes involved reproductively mature females during the nesting season, a period in which the females were in potential calcium stress. To our knowledge this is the first report of mining and long-term exploitation of natural mineral deposits by a lower vertebrate and may represent behaviour at least as complicated as that seen in many ungulates.
Thesis (Ph. D.)--University of Chicago, 1985. Includes bibliographical references (p. 317-328). Photocopy.
THE anthropologist, when studying excavated bone remains, often finds it
difficult to distinguish specimens worked by hominids from those damaged
by other processes. Although damage by carnivores and rodents is widely
recognised as a potential source of confusion1, the role of
herbivores (though well known to cattle and deer herders) is less well
documented. In this communication, which originates from a visit to the
Hardangervidda Mountains, Norway, in 1971, I describe some bones and
antlers gnawed by Norwegian reindeer and Scottish red deer. It is clear
that bones and antlers so gnawed can resemble human artefacts and have
sometimes been mistaken for them.
Radant provided workspace, logistical support, or advice. M. Cof feen generously provided unpublished data from radiographs
- H Hohman
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Hohman, H. Martin, C. Pregler, and especially R.
B. Bury and C. R. Tracy provided helpful com
ments and reviews of the manuscript. R. Benton,
T. A. Duck, D. Pietrzak, and R. Radant provided
workspace, logistical support, or advice. M. Cof
feen generously provided unpublished data from
radiographs. R. B. Bury, P. A. Medica, and F. B.
Care and nutrition of pet reptiles
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Ecology and management of the desert tortoise (Gopherus agassizii) in Califor nia
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Luckenbach, R. A 1982. Ecology and management of
the desert tortoise (Gopherus agassizii) in Califor
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