Diet selection and digestion in Iguana iguana: the importance of age and nutrient requirements

Summary The green iguana, Iguana iguana, is herbivorous throughout life, and depends on a microbial fermentation system in the hindgut to degrade plant fiber. Because the metabolic rates of lizards are proportional to body mass raised to the 0.80 power, hatchling iguanas have 2X, and juveniles 1.4X, greater relative energy requirements (kJxg body mass^-1xday^-1) than full-grown adults. Growing animals also need a higher protein intake, for contruction of body tissues, than do mature animals. This study investigated how growing iguanas achieve a relatively greater nutrient intake than adults. Hatchling and juvenile iguanas do not have higher relative capacities of the digestive tract than mature iguanas, nor do they digest plant materials more effectively. Instead, growing iguanas select diets higher in digestible protein, and digest the same food 1.3X to 2X more rapidly, than adults. Young iguanas may accomplish their shorter food transit times by maintaining higher body temperatures.     

Diet selection of African elephant over time shows changing optimization currency

Multiple factors determine diet selection of herbivores. However, in many diet studies selection of single nutrients is studied or optimization models are developed using only one currency. In this paper, we use linear programming to explain diet selection by African elephant based on plant availability and nutrient and deterrent content over time. Our results indicate that elephant at our study area maximized intake of phosphorus throughout the year, possibly in response to the deficiency of this nutrient in the region. After adjusting the model to incorporate the effects of this deficiency, elephant were found to maximize nitrogen intake during the wet season and energy during the dry season. We reason that the increased energy requirements during the dry season can be explained by seasonal changes in water availability and forage abundance. As forage abundance decrease into the dry season, elephant struggle to satisfy their large absolute food requirements. Adding to this restriction is the simultaneous decrease in plant and surface water availability, which force the elephant to seek out scarce surface water sources at high energy costs. During the wet season when food becomes more abundant and energy requirements are satisfied easier, elephant aim to maximize nitrogen intake for growth and reproduction. Our study contributes to the emerging theory on understanding foraging for multiple resources.     

Environmental constraints on the foraging behaviour of a dwarf antelope (Madoqua kirkii)

Dik-diks (Madoqua sp.) inhabit semi-arid regions and experience very different conditions of food availability and quality between wet and dry seasons. By comparing the behaviour of dik-diks between these two seasons, we identified environmental constraints affecting their feeding strategies. In both seasons foraging time was limited by high mid day temperatures. In the wet season a high intake rate compensates for the loss in foraging time, but in the dry season water and protein become limiting. To meet minimum daily water requirements in the dry season dik-diks fed on plant species that they avoided during the wet season. Analysis at the plant species level showed higher species selectivity in the wet season than in the dry season. In a multiple regression analysis food species preferences were best explained by relative abundance and water content in the dry season, and by dry matter content in the wet season. In the wet season the daily dry-matter intake of dik-diks in the field was only about 10% higher than the theoretically predicted minimum for a ruminant of this body weight, while protein and water intake were about 3 times as high. This suggests that the most limiting dietary component in the wet season is energy. In the dry season the daily intake of all dietary components is lower than the theoretical minimum required, and also lower than the values suggested by laboratory studies of dik-diks. This dry season deficit is presumably met from body reserves. Dry season water intake was approximately 30% of the intake observed in laboratory studies indicating that dikdiks are even better adapted to arid conditions than suggested by physiological experiments.     

Diet and nutrition in wild mongoose lemurs (Eulemur mongoz) and their implications for the evolution of female dominance and small group size in lemurs

Data collected on the feeding behavior, food intake, and chemical analyses of plant foods were used to document seasonal variation in diet and nutrition in Eulemur mongoz in northwestern Madagascar. E. mongoz conforms to the general Eulemur dietary pattern, with a predominantly frugivorous diet supplemented mainly by leaves, flowers, and nectar. Phytochemical analysis revealed high water contents in all the main plant foods; mature fruit and flowers contained the most water-soluble carbohydrates; immature leaves were richest in protein and essential amino acids; the limiting amino acids in all plant foods were methionine and cystine; ash (mineral) content was highest in petioles and mature leaves; crude lipid content was highest in seeds; and crude fiber content was indistinguishable between immature and mature fruit and leaves. High-fiber foods were eaten during both seasons; the wet season diet was dominated by high-energy foods (mature fruit, nectar, and seeds), while the dry season diet contained foods high in energy (mature fruit and flowers) and high in protein (immature leaves) and minerals (mature leaves and petioles). However, nutrient intake did not vary between seasons, implying that nutrient requirements are met throughout the year. These results suggest we draw more conservative conclusions when interpreting dietary variability in the absence of chemical analysis, and also draw into question the idea that nutritional stress is a factor in the timing of reproduction in lemurs and, by extension, is linked to the prevalence of female dominance and small group size in lemurs.     

Food limitation and demography of a migratory antelope,the white-eared kob

Summary Although it is commonly presumed that many populations of large-herbivores are limited by food availability, supporting evidence is scarce. This hypothesis was investigated in a population of over 800,000 white-eared kob in the Boma National Park region of the southern Sudan. Food availability, body condition, and mortality rates of adults and calves were measured during the dry seasons of 1982 and 1983. Sampled age distributions from both the live population and carcasses were used to calculate age-specific rates of mortality. In 1982, food supplies during the dry season were augmented by substantial rainfall, which produced regrowth of grass in areas that ordinarily had little green forage. As a result, fat reserves declined little, and rates of adult mortality showed no increasing trend. Total adult mortality was 5%. In 1983, there was no rainfall during the dry season and food intake was insufficient to meet the estimated energy requirements of kob. As a result, fat reserves declined and adult mortality rates increased fourfold. Total adult mortality was 10% (equivalent to the recruitment rate of yearling into the population). Calf mortality during the dry season was similar in both years (50%), based on field estimates of mortality rates and calf/female ratios. Lactation throughout the dry season possibly provided a buffer for calves against variations in food availability. The age structure of the live population in 1983 suggests that a drought in 1980 reduced kob numbers by 40%. These results suggest that adult survival is influenced strongly by the availability of food during the dry season. However, the duration of the dry season also plays an important role. During the dry season, declining fat reserves make an increasing proportion of the population vulnerable to mortality. As a result, even moderate droughts may lead to substantial changes in population numbers.     

Ontogenetic changes in food intake and digestion rate of the herbivorous marine iguana (Amblyrhynchus cristatus,Bell)

Young reptiles have higher relative energy demands than adults, but the proposed ontogenetic changes in diet to fulfil these demands were not found in the algae-eating Galápagos marine iguanas on Santa Fé. Feeding and digestion rates were investigated to analyse how young achieve higher energy intake. Daily food intake of free ranging marine iguana hatchlings (6–11 months old) was about one third that of adults, but relative intake (g dry mass · g^–1 wet mass · day^–1) was four times higher in the hatchlings. During feeding experiments, relative daily food intake of hatchling marine iguanas was approximately three times higher than that of adults (0.042 vs 0.013 g dry mass · g^–0.8 wet mass · day^–1), and mean gut passage time was two times shorter (5 vs 10 days). The hatchlings also maintained high body temperatures (36.7° C) even under relatively cool day-time air temperatures of 32° C. Apparent digestibility of algal food measured both during feeding trials and by Mn²⁺ AAS (atomic absorption spectrometry) for free-ranging iguanas was 70%, independent of body size and temperature. The red algae prevailing in the diet were high in protein (30% dry mass) and energy (12.1 kJ/g dry mass). Diving iguanas had higher rates of energy intake than intertidal foragers, but daily intake was less. Maintenance of high body temperature enabled hatchlings to achieve high digestion rates and, combined with high relative intake, thus achieve sufficient energy intake for rapid growth despite higher mass specific metabolic rates. Estimates of biomass of marine iguanas and their algal food are given for a section of coastline on Santa Fé.     

Spatial and temporal patterns of home range use by olive baboons (Papio anubis) in eastern Laikipia,Kenya

The study investigated the relationship between topography, seasonal food availability and home range use by a troop of olive baboons (Papio anubis) in a dry savannah environment. Valleys were the most utilized parts of the home range in dry seasons while old boma sites were more frequently used in the wet season. Herb‐layer food biomass decreased significantly in dry seasons. The flowers of both Acacia tortilis and Acacia etbaica were available only in dry seasons with valleys having the highest biomass. The biomass of Acacia nilotica seeds was higher in valleys than on other locations. Leaves of the shrub Lycium europaeum were available on all locations in the wet season but valleys had the highest biomass of this food type. Herb‐layer baboon food biomass at ridge tops, slopes, valleys and old boma sites showed similar trends in productivity between seasons. Baboons tracked food along the soil catenas by feeding at ridge tops and old boma sites where foods were available during the wet season and exploiting food sources at the valleys during the dry seasons. Food availability was related to the catena effect and baboons used the catenas in the home range in a predictable manner.     

Nutritional quality of gorilla diets: consequences of age,sex, and season

We tested the effects of age, sex, and season on the nutritional strategies of a group of mountain gorillas (Gorilla beringei) in the Bwindi Impenetrable National Park, Uganda. Through observations of food intake of individual gorillas and nutritional analyses of dietary components over different seasons and environments, we estimated nutrient intake and evaluated diet adequacy. Our results suggest that the nutritional costs of reproduction and growth affect nutrient intake; growing juveniles and adult females ate more food and more protein per kilogram of metabolic body mass than did silverbacks. The diets of silverback males, adult females, and juveniles contained similar concentrations of protein, fiber, and sugar, indicating that adult females and juveniles did not select higher protein foods than silverbacks but rather consumed more dry matter to ingest more protein. Juveniles consumed more minerals (Ca, P, Mg, K, Fe, Zn, Mn, Mo) per kilogram of body mass than adult females and silverback males, and juveniles consumed diets with higher concentrations of phosphorous, iron, and zinc, indicating that the foods they ate contained higher concentrations of these minerals. Seasonally, the amount of food consumed on a dry weight basis did not vary, but with increased frugivory, dietary concentrations of protein and fiber decreased and those of water-soluble carbohydrates increased. Energy intake did not change over the year. With the exception of sodium, gorillas ate diets that exceeded human nutrient requirements. A better understanding of the relative importance of food quantity and quality for different age–sex classes provides insights into the ways in which gorillas may be limited by food resources when faced with environmental heterogeneity.     

Nutritional ecology of a browsing ruminant, the kudu (Tragelaphus strepsiceros), through the seasonal cycle

The activities and food selection of four hand-reared kudus were recorded in a large fenced enclosure containing natural savanna vegetation in the Nylsvley Nature Reserve, South Africa. Leaves of selected species were analysed chemically for crude protein, fibre constituents, phosphorus, condensed tannin and total polyphenols. Available protein and metabolizable energy were estimated allowing for potential antinutritional effects of tannins.
Leaves of palatable deciduous woody plants and herbaceous forbs formed the main dietary constituents during the late wet season. Foliage from palatable evergreens and robust forbs were added to the diet during the dry season. Towards the end of the dry season unpalatable species of evergreens were eaten. At the start of the growing season new leaves of otherwise unpalatable woody species formed the staple food source, together with fruits of Strychnos spp. Correspondingly, protein and digestible dry matter concentrations in the diet declined to reach a low at the end of the dry season.
Total daily food intake increased to compensate for reduced dietary quality during the dry season, until little edible foliage remained. While the estimated daily intake of protein remained well above maintenance requirements, the estimated metabolizable energy intake fell below requirements during the late dry season. Phosphorus intake may have been submaintenance in the dry season. Nutritional balance was dependent on the availability of particular vegetation components to serve as nutritional stepping stones during crucial times of the year. These included forbs during the late wet season, palatable evergreens in the dry season, and Strychnos fruits plus early-flushing woody plants during the dry season-wet season transition.     

Diet and Geophagy Across a Western Amazonian Parrot Assemblage

We identified species‐ and community‐level dietary characteristics for a species‐rich Amazonian parrot assemblage to determine relationships among dietary metrics and use of geophagy sites. Previous studies suggest that soil is consumed at geophagy sites in this region mainly to supplement dietary sodium. We accumulated 1400 feeding records for 16 parrot species over 2 yr and found that seeds, flowers, and fruit pulp featured prominently in diets, while bark, insects, and lichen were consumed in small quantities. Food availability across 1819 trees was measured, and we found that flower availability was highest in the dry season and fruit production peaked in the wet season, but that phenology patterns of the 20 most commonly foraged plant species suggest no serious food bottlenecks. Partitioning of available food resources among the 13 most commonly encountered parrots is suggested by an ordination analysis (DCA), which placed the large macaws (Ara) with the Amazona parrots at the ‘primary forest’ end of a dietary resource axis and four smaller species at the ‘successional forest’ end of the axis. Parrot species associated with successional forest also consumed less plant species overall. Furthermore, these parrot species consuming successional forest resources had higher claylick visitation rates than those consuming primary forest resources suggesting they derive the greatest benefits from soil consumption.     

Nutritional Correlates of Population Density Across Habitats and Logging Intensities in Redtail Monkeys (Cercopithecus ascanius)1

Food resources are arguably the most common limiting factor for most species, yet little is known about how food quantity and quality interact to determine the size and distribution of wildlife populations. Frugivores may be limited by food quality since fruits, unlike leaves, are typically low in protein and minerals. Understanding the factors influencing frugivore population dynamics is particularly important due to rapid habitat loss and the need to construct informed management plans. Therefore, this study used redtail monkeys in Kibale National Park, Uganda as a model frugivore to (1) compare nutrient intake, reproduction, and habitat use patterns for three redtail groups in an unlogged and heavily logged area; and (2) examine relationships between nutrient intake and redtail population densities across five habitats. Feeding time and intake of crude protein, lipids, and minerals were higher in diets consumed by groups in unlogged areas than heavily logged areas. Intake of both copper and sodium intake was below NRC requirement levels suggested for macaques. Food availability in the unlogged areas was 3.5 times higher than the heavily logged area. Across five study sites, absorbable copper and the ratio of copper to caloric intake in redtail diets were significantly related to redtail population density. These results suggest that food quality is an important factor determining frugivore population density in anthropogenically disturbed frugivore habitats. Copper deficiency, which has been associated with population declines of wild herbivores, has the potential to be a key factor limiting densities of frugivores in tropical habitats.     

Dietary generalists and nutritional specialists: Feeding strategies of adult female blue monkeys (Cercopithecus mitis) in the Kakamega Forest,Kenya

Generalist primates eat many food types and shift their diet with changes in food availability. Variation in foods eaten may not, however, match variation in nutrient intake. We examined dietary variation in a generalist‐feeder, the blue monkey (Cercopithecus mitis), to see how dietary food intake related to variation in available food and nutrient intake. We used 371 all‐day focal follows from 24 adult females (three groups) in a wild rainforest population to quantify daily diet over 9 months. We measured food availability using vegetation surveys and phenology monitoring. We analyzed >700 food and fecal samples for macronutrient content. Subjects included 445 food items (species‐specific plant parts and insect morphotypes) in their diet. Variation in fruit consumption (percentage of diet and total kcal) tracked variation in availability, suggesting fruit was a preferred food type. Fruits also constituted the majority of the diet (by calories) and some fruit species were eaten more than expected based on relative availability. In contrast, few species of young leaves were eaten more than expected. Also, subjects ate fewer young leaves (based on calories consumed) when fruit or young leaves were more available, suggesting that young leaves served as fallback foods. Despite the broad range of foods in the diet, group differences in fiber digestibility, and variation that reflected food availability, subjects and groups converged on similar nutrient intakes (grand mean ± SD: 637.1 ± 104.7 kcal overall energy intake, 293.3 ± 46.9 kcal nonstructural carbohydrate, 147.8 ± 72.4 kcal lipid, 107.8 ± 12.9 kcal available protein, and 88.1 ± 17.5 kcal structural carbohydrate; N = 24 subjects). Thus, blue monkeys appear to be food composition generalists and nutrient intake specialists, using flexible feeding strategies to regulate nutrient intake. Findings highlight the importance of simultaneously examining dietary composition at both levels of foods and nutrients to understand primate feeding ecology.     

Nutritional estimate of populations of some wild free-ranging African ungulates in grassland (Nechisar national park, Ethiopia) in dry season

The amount and nutritive value of forage plants, diet composition, digestibility of dry matter and nutrients were recorded for zebra. Grant's gazelle, Swayne's hartebeest and hippopotamus in November-December 1991 Besides, daily egest of feces, the level of food and nutrient consumption, energy and protein requirements were recorded for zebra and Grant's gazelle The digestibility of pasture forage was determined as a ratio of lignin concentration in food to the concentration m feces (lignin tracer technique), a daily intake was calculated on the basis of the daily feces egest Protein percentage m the diet of zebra and hartebeest consuming dry parts of grasses did not exceed 5% Gazelle diet consists of green parts of plants and included 18% of protein The digestibility of dry matter in nonruminants (zebra, hippopotamus) was 40-45%, in hartebeest - 50%, in gazelle - 60% Due to the abundance of dry grasses (3 7 ton ha^-1) the daily food consumption of zebra was high - 7 2 kg ind^-1 (dry weight), the metabolizable energy intake (ME) being 51 MJ Adult gazelles consumed 15-25 kg ind^-1 of food and 14-24 MJ of ME The energy requirements of adult males and non-lactating females of zebras and gazelles (48 and 13 MJ respectively) were met, the energy balance berig negative for lactating animals The daily protein requirement was not met in zebra (392-704 g md^-1 vs 134 g ind^-1 of intake) and in lactating gazelles (250 g ind^-1 vs 197 g md^-1) Non-lactating gazelles consume sufficient amount of both energy and protein due to the high feeding selectivity of the species and thanks to the abundance of burnt areas with young green after-grass m the dry period     

Nutritional correlates of the “lean season”: Effects of seasonality and frugivory on the nutritional ecology of diademed sifakas

Primate field studies often identify “lean seasons,” when preferred foods are scarce, and lower‐quality, abundant foods (fallback foods) are consumed. Here, we quantify the nutritional implications of these terms for two diademed sifaka groups (Propithecus diadema) in Madagascar, using detailed feeding observations and chemical analyses of foods. In particular, we sought to understand 1) how macronutrient and energy intakes vary seasonally, including whether these intakes respond in similar or divergent ways; 2) how the amount of food ingested varies seasonally (including whether changes in amount eaten may compensate for altered food quality); and 3) correlations between these variables and the degree of frugivory. In the lean season, sifakas shifted to non‐fruit foods (leaves and flowers), which tended to be high in protein while low in other macronutrients and energy, but the average composition of the most used foods in each season was similar. They also showed dramatic decreases in feeding time, food ingested, and consequently, daily intake of macronutrients and energy. The degree of frugivory in the daily diet was a strong positive predictor of feeding time, amount ingested and all macronutrient and energy intakes, though season had an independent effect. These results suggest that factors restricting how much food can be eaten (e.g., handling time, availability, or intrinsic characteristics like fiber and plant secondary metabolites) can be more important than the nutritional composition of foods themselves in determining nutritional outcomes—a finding with relevance for understanding seasonal changes in behavior, life history strategies, competitive regimes, and conservation planning. Am J Phys Anthropol 153:78–91, 2014. © 2013 Wiley Periodicals, Inc.     

The relationship between dietary intake and the number of teeth in elderly Japanese subjects

Objective: This study used a precise weighing method to assess whether tooth loss was related to nutrient intake in elderly Japanese subjects. Material and methods: Fifty‐seven subjects aged 74 years were randomly selected from a longitudinal interdisciplinary study of ageing. Complete 3‐day food intake data were obtained by a precise weighing method. The dietary intakes of energy and nutrients were calculated based on the Standard Tables of Food Composition in Japan (5th ed.). A clinical evaluation of the number of teeth present was carried out. Multiple regression standardised coefficients for each nutrient was estimated based on a continuous scale adjusted for gender, smoking habits, and educational level. After dividing the subjects into two groups according to the number of teeth present (0–19, 20+), the difference in the intake of nutrients and the amount of food consumed per day was evaluated. Results: The number of teeth present had a significant relationship with the intake of several nutrients. In particular, total protein, animal protein, sodium, vitamin D, vitamin B₁, vitamin B₆, niacin, and pantothenic acid were significantly associated with the number of teeth present and with the two groups (0–19, 20+). The intake of vegetables and fish, shellfish, and their products was significantly lower among subjects with fewer teeth. Conclusion: This study suggests that there was a significant relationship between nutrient intake, such as minerals and vitamins from food, and tooth loss.     

Diet selection in the green ringtail possum (Pseudochirops archeri): A specialist folivore in a diverse forest

Abstract This study investigated the feeding ecology of the green ringtail possum, Pseudochirops archeri (Pseudocheiridae) in a tropical rainforest with 94 plant species in the canopy. Over 50% of tree use was from only four tree species, Aleurites rockinghamensis, Ficus fraseri, Arytera divaricata and Ficus copiosa. These species were used significantly more frequently than would be expected if tree species were selected randomly in proportion to their relative abundance in the forest. Conversely, 88 other tree species present were used less frequently than expected. Possums also favoured particular individual trees within some of the preferred tree species. In 91% of feeding observations, possums consumed mature leaves only. The availability of young leaves, flowers and fruit varied throughout the year, with a peak in availability of these resources during the early wet season. By primarily selecting mature leaves, green ringtail possums reduce their dependence on seasonally variable resources. We suggest that green ringtail possums should be considered as specialist folivores, focusing on only a few of the tree species available, possibly due to advantages associated with limiting the number of plant secondary metabolites in their diet. Furthermore, they favour certain individual trees within species, perhaps due to intraspecific variation in plant secondary metabolites or nutrient content, behaviour that has been well established in eucalypt folivores. We conclude that green ringtail possums are highly specialized in their feeding ecology, limiting their diet to a small number of continuously available food items.     

Nutrition as a major facet of reptile conservation

The importance of nutrition has not received much recognition in conservation biology. However, captive breeding is possible only if nutritional requirements of animals are met, and effective habitat management requires an evaluation of nutritional resources. Three examples involving reptile conservation are presented. The formulation and testing of experimental meal-type diets proved essential for the large-scale rearing of green iguanas (Iguana iguana) in Panama and Costa Rica, thousands of which have been released into the wild. Survival and growth of captive land iguanas (Conolophus subcristatus) in the Galapagos Islands was markedly improved by development of a complete feed based on locally available ingredients; this was essential to continuation of the conservation program in which juvenile iguanas were repatriated to islands where populations had previously been exterminated. Research on the desert tortoise (Gopherus agassizii) in the Mojave Desert has identified nutritional constraints that may limit utilization of potential food plants. Thus, nutritional status of wild tortoises may depend more on availability of plant species of high nutritional quality than on overall amounts of annual vegetation. Federal and local agencies involved in the conservation and management of tortoise habitat have recognized the need to fund research on tortoise nutrition. We contend that nutrition should be given a central role in conservation programs for reptiles and other animals. (This article is a US Government work and, as such, is in the public domain in the United States of America.) © 1996 Wiley-Liss, Inc.     

Seasonal intake responses in the nectar-feeding bat Glossophaga soricina

Food intake in nectar-feeding animals is affected by food quality, their energetic demands, and the environmental conditions they face. These animals increase their food intake in response to a decrease in food quality, a behavior named “intake response”. However, their capacity to achieve compensatory feeding, in which they maintain a constant flux of energy, could be constrained by physiological processes. Here we evaluated how both a seasonal change in environmental conditions and physiological constraints affected the food ingestion in the bat Glossophaga soricina. We measured food intake rate during both the wet/warm and dry/cool seasons at sucrose solutions ranging from 146 to 1,022 mmol L⁻¹. We expected that food intake and metabolic demands would be greater during the dry/cool season. Bats ingested ~20% more food in the dry/cool than in the wet/warm season. Regardless of season, bats were unable to achieve a constant flux of energy when facing the different sugar concentrations that we used in our experiments. This suggests that the rate of food intake is physiologically constrained in G. soricina. Using the digestive capacity of bats we modeled their food intake. The analytic model we used predicts that digestive limitations to ingest energy should have an important effect on the ecology of this species.     

Preliminary observations on the relationship of calcium ingestion to vitamin D status in the green iguana (Iguana iguana)

We hypothesized the vitamin D-deficient green iguanas with depleted calcium stores would seek to augment calcium intake by self-selection of a high calcium source. Eight green iguanas were offered free-choice ground oystershell in addition to their regular diet. Of these, two had not been exposed to ultraviolet (UV-B) radiation for > 5 years and were demonstrated to be vitamin D-deficient by low circulating levels of the principal vitamin D metabolite, calcidiol (25-hydroxy-cholecalciferol). The six others had been exposed to a UV-B emitting bulb for the previous 3 years and had high circulating calcidiol levels. Average daily food intake (expressed as dry matter per kg body mass) did not differ between the Low-D and High-D iguanas. The daily oystershell intake of the Low-D iguanas (0.02–0.03 g/kg) was lower than that of the High-D iguanas (0.06–0.70 g/kg), leading to a significant difference in calcium intake. The failure of iguanas to increase calcium intake in response to vitamin D-deficiency was puzzling and suggests that vitamin D, as a steroid hormone, may play some role in the expression of calcium appetite. Zoo Biol 16:201–207, 1997. © 1997 Wiley-Liss, Inc.