Intake, ingesta retention, particle size distribution and digestibility in the hippopotamidae

Although several aspects of the digestive physiology of the hippopotamidae-non-ruminating foregut fermenters-have been described, ingesta kinetics and passage characteristics of these species are not well understood. The most outstanding feature of the hippo digestive physiology reported so far is the very long mean ingesta retention times (MRTs) measured by Foose [Foose, T., 1982. Trophic strategies of ruminant versus nonruminant ungulates. PhD dissertation, University of Chicago, Chicago.]. Since those data had been investigated with animals without water access, we intended to measure MRT in hippos which were allowed to enter water pools during the night. MRT parameters as well as dry matter (DM) digestibility were determined in four common (Hippopotamus amphibius) and four pygmy hippos (Hexaprotodon liberiensis) on two different diets each using cobalt ethylendiamintetraacetate (Co-EDTA) as a fluid, chromium (Cr)-mordanted fibre (<2 mm) as a particle and acid detergent lignin (ADL) as an internal digestibility marker. Four of the animals additionally received cerium (Ce)-mordanted fibres (2-10 mm) as particle markers. Total MRTs for fluids and particles ranged between 20-35 and 48-106 h in the common and between 13-39 and 32-107 h in the pygmy hippos. The difference between fluid and particle retention was greater than usually reported in ruminants. Excretion patterns of the markers differed from those usually observed in ruminants but resembled those reported for macropods (kangaroos), indicating a plug-flow reactor-like physiology in the hippo forestomach (FRST). This finding complements other described similarities between the macropod and the hippo forestomach. The measurements of larger particle retention profiles suggest that in the hippo, larger particles might be excreted either faster or at the same rate as smaller particles, indicating a general difference between ruminants and hippos with respect to differential particle retention. The digestive physiology of hippos is characterised by a generally low food intake, long ingesta retention times and dry matter digestibilities lower than reported in ruminants. Moderate digestibilities in spite of long retention times might be the result of the generally high average ingesta particle size in hippos. The comparatively easy management of pygmy hippos, together with the significant correlations between food intake, MRT and digestibility in the pygmy hippos of this study, recommends this species for further studies on the interplay of these parameters in herbivore digestive physiology.     

Digesta passage and fiber digestibility in captive white-faced sakis (Pithecia pithecia)

Wild white-faced sakis (Pithecia pithecia) ingest primarily seeds that provide a diet that is lipid-rich and moderately high in dietary fiber. Although little anatomical information is available on sakis, evidence from other vertebrate seed predators suggests that such a diet is correlated with adaptations in gut morphology or physiology. Milton [1984] reported a 20 hr transit time (TT=transit time or time of first appearance in feces) of a particulate marker for a single monk saki (Pithecia monachus). This suggests that TT for Pithecia sakis may be four to five times longer than what has been reported for soft-fruit-eating platyrrhines, such as Ateles and Cebus. During a captive study, we calculated an average TT of 14.7+/-0.4 hr (n=5 trials started in the evening) for a chromic oxide (Cr(2)O(3)) marker that follows liquid digesta and TTs of 14.5 hr (trial started the previous evening) and 23.0 hr (trial started the previous morning) for two trials using particulate markers. Mean retention time (MRT) for the liquid marker ranged from 15.3 hr to 37.7 hr in four trials that lasted longer than 90 hr. Marker recovery was incomplete for the particulate markers in these trials, and thus MRT could only be determined for the liquid phase marker. Three 5-day trials on a low-fiber, blended diet revealed high fiber fraction digestibilities (neutral detergent fiber (NDF)=77.4% and acid detergent fiber (ADF)=74.4%). Data collected for this study and nutritional data from wild sakis suggest that pitheciin seed predators may have a potential for fiber digestibility that is intermediate between ripe-pulp frugivores and folivores.     

Feed intake,digestion and passage of the proboscis monkey (Nasalis larvatus) in captivity

Feeding trials were conducted with a troop of six proboscis monkeys (Nasalis larvatus) at the New York Zoological Park to quantify aspects of digestive physiology. Diets consumed comprised, on an as-fed basis, 55% browse and green produce, 11% fruit, 9% commercial primate diets, 9% root vegetables, and 16% miscellaneous items. Total feed intake averaged 12% of body mass; dry matter intake was 3%. Dry matter and plant cell wall disappearance exceeded 80%. Passage marker studies (three 6-day trials) revealed: transit time=14 hr; mean passage time=49 hr; and 5–80% retention time=52 hr. Digestion coefficients and retention times were greater than expected based on body size. These results might reflect the low fiber content of the diets fed, or alternatively, provide evidence that proboscis monkeys are efficient digesters of cell wall constituents.     

Retention of solutes and different-sized particles in the digestive tract of the ostrich (Struthio camelus massaicus), and a comparison with mammals and reptiles

Ostriches (Struthio camelus) achieve digesta retention times, digesta particle size reduction and digestibilities equal to similar-sized herbivorous mammals, in contrast to some other avian herbivores. The sequence of digestive processes in their gastrointestinal tract, however, is still unexplored. Using two groups of four ostriches (mean body mass 75.1 ± 17.3 kg) kept on fresh alfalfa, we tested the effect of two intake levels (17 and 42 g dry matter kg(-0.75)d(-1)) on the mean retention time (MRT) of a solute and three different-sized (2, 10, 20 mm) particle markers, mean faecal particle size (MPS), and digestibility. Intake level did not affect MRT, but MPS (0.74 vs. 1.52 mm) and dry matter digestibility (81 vs. 78%). The solute marker (MRT 22-26 h) was excreted faster than the particle markers; there was no difference in the MRT of 10 and 20 mm particles (MRT 28-32 h), but 2mm particles were retained longer (MRT 39-40 h). Because the solute marker was not selectively retained, and wet-sieving of gut contents of slaughtered animals did not indicate smaller particles in the caeca, the long MRT of small particles is interpreted as intermittent excretion from the gizzard, potentially due to entrapment in small grit. The marker excretion pattern also showed intermittent peaks for all markers in five of the animals, which indicates non-continuous outflow from the gizzard. When adding our data to literature data on avian herbivores, a dichotomy is evident, with ostrich and hoatzin (Opisthocomus hoazin) displaying long MRTs, high digestibilities, and gut capacities similar to mammalian herbivores, and other avian herbivores such as grouse, geese or emus with shorter MRTs, lower fibre digestibilities and lower gut capacities. In the available data for all avian herbivores where food intake and MRTs were measured, this dichotomy and food intake level, but not body mass, was related to MRT, adding to the evidence that body mass itself may not be sole major determinant of digestive physiology. The most striking difference between mammalian and avian herbivores from the literature is the fundamentally lower methane production measured in the very few studies in birds including ostriches, which appears to be at the level of reptiles, in spite of general food intake levels of a magnitude as in mammals. Further studies in ostriches and other avian herbivores are required to understand the differences in digestive mechanisms between avian and mammalian herbivores.     

Binturong (Arctictis binturong) and Kinkajou (Potos flavus) Digestive Strategy: Implications for Interpreting Frugivory in Carnivora and Primates

Exclusive frugivory is rare. As a food resource, fruit is temporally and spatially patchy, low in protein, and variable in terms of energy yield from different carbohydrate types. Here, we evaluate the digestive physiology of two frugivorous Carnivora species (Potos flavus, Arctictis binturong) that converge with primates in a diversity of ecological and anatomical traits related to fruit consumption. We conducted feeding trials to determine mean digestive retention times (MRT) on captive animals at the Carnivore Preservation Trust (now Carolina Tiger Rescue), Pittsboro, NC. Fecal samples were collected on study subjects for in vitro analysis to determine methane, pH, and short chain fatty acid profiles; fiber was assayed using standard neutral detergent (NDF) and acid detergent (ADF) fiber methods. Results indicate that both carnivoran species have rapid digestive passage for mammals that consume a predominantly plant-based diet: A. binturong MRT = 6.5 hrs (0.3); P. flavus MRT = 2.5 hrs (1.6). In vitro experiments revealed no fermentation of structural polysaccharides – methane levels did not shift from 0 h to either 24 or 48 hours and no short chain fatty acids were detected. In both species, however, pH declined from one incubation period to another suggesting acidification and bacterial activity of microbes using soluble carbohydrates. A comparison with primates indicates that the study species are most similar in digestive retention times to Ateles – the most frugivorous anthropoid primate taxon.     

小兴安岭地区原麝冬季食性研究

2004 年2 ～3 月和2004 年11 月至2005 年1 月在黑龙江省通河县龙口林场,对原麝冬季的食性进行了研究。采用粪便显微组织学分析技术分析原麝冬季采食植物的种类组成和比例,并测定了主要取食的9 种木本植物及苔藓和蕨类植物的营养成分。研究结果表明,原麝冬季取食的植物共计46 科66 种,其中兴安杜鹃、光萼溲疏、五角槭、羊胡子苔草分别占32.7%、7.8%、5.81%、5.56%, 是原麝冬季主要取食的植物种类,占取食植物的51.9% 。木本植物枝叶是原麝冬季的主要食物,在冬季食物组成中所占百分比最高, 达75.0% 。对草本植物、蕨类、苔藓、地衣的取食量均较小。冬季原麝比较喜欢采食蛋白含量高的植物,采食部位主要为植物木质化较低的当年枝顶端部分。藓类主要出现在1 ～3 月份的食物中。     

The effects of ration level on food retention time in juvenile lemon sharks,Negaprion brevirostris

Synopsis Fecal production was monitored to observe the effects of meal size on retention time of food in the digestive tracts of lemon sharks, Negaprion brevirostris. Initial appearance of feces occurred more rapidly when ration level was increased. The onset of fecal production was negatively correlated with rate of intake. Production of feces continued for a longer period of time when meal size was increased. Retention time of food was directly related to feeding rate, suggesting that the rate of digestion was constant. The correlation between retention time and intake on a percentage body weight basis was greater than the correlation between retention time and intake on an energy density basis. The use of agar to bind food may have delayed digestion and prolonged food passage for sharks fed an experimental diet.     

Retention of fluid and particles in the digestive tract of the llama (Lama guanacoe f. glama)

The retention times of fluid and particles of different lengths were measured in the digestive tract of the llama. PEG was used as a fluid marker; hay particles were labelled with cerium or samarium and the samples were determined by neutron activation analysis. In compartments 1 and 2 the retention time of fluid (9.7 hr) was significantly shorter than that of particles having a length of 0.2-1.0 cm (19.2 hr) and 2.5-4.0 cm (25.0 hr). In the intestine slightly significant differences in the retention times of fluid (20.1 hr) and particles (26.2 hr) were seen. In the whole digestive tract retention times were 36.2 hr for fluid, 52.0 hr for smaller particles and 59.9 hr for larger particles.     

Digestive efficiency of the Hoatzin, Opisthocomus hoazin: a folivorous bird with foregut fermentation

The Hoatzin Opisthocomus hoazin is the only known bird with a well-developed foregut plant fermentation system; most fermentation takes place in the crop and caudal oesophagus. To test Hoatzin digestive efficiency, balance (total collection) trials with captive Hoatzins were made using two experimental diets of different composition and fibre content. Dry matter (DM) intakes were similar for the diets (mean = 62.8 g DM/kg body mass/ day). Average DM, organic matter and nitrogen digestibilities were not significantly different between diets, with average values of 72.9%, 75.0% and 78.3%, respectively. In vitro organic matter digestibilities by cow ruminal inoculum were very similar to organic matter digestibilities in live Hoatzins for both diets. Fibre digestibility was among the highest recorded for herbivorous birds. Cellulose and acid detergent fibre digestibilities were 58.8% and 52.7%, respectively. Neutral detergent fibre (NDF) digestibility differed among diets—the higher the NDF content of the diet, the higher the NDF digestibility. The NDF digestibilities were 37.9% and 70.9% for the two diets with NDF concentrations of 32.4% and 37.3%, respectively. Differences in NDF digestibility can be attributed to the different concentrations of hemicellulose in the experimental diets. The high overall digestibility by captive Hoatzins is higher than values previously reported for other avian herbivores but similar to those of foregut-fermenting mammals on similar diets. The unique digestive strategy of the Hoatzin maximizes digestion of cell wall and cell contents. The high digestive efficiency in the Hoatzin is not predicted by allometric models of fibre digestion as a function of body mass. Other nutritional benefits, such as detoxification of plant secondary compounds and microbial synthesis of essential amino acids and vitamins, may explain the evolution of foregut fermentation in this avian folivore.     

More efficient mastication allows increasing intake without compromising digestibility or necessitating a larger gut: Comparative feeding trials in banteng (Bos javanicus) and pygmy hippopotamus (Hexaprotodon liberiensis)

The digestion of plant material in mammalian herbivores basically depends on the chemical and structural composition of the diet, the mean particle size to which the forage is processed, and the ingesta retention time. These different factors can be influenced by the animal, and they can presumably compensate for each other. The pygmy hippopotamus, a non-ruminating foregut fermenter, has longer mean retention times than ruminants; however hippos do not achieve higher (fibre) digestibilities on comparable diets, which could be due to ineffective mastication. We performed feeding trials with six pygmy hippos (Hexaprotodon liberiensis) and six banteng cattle (Bos javanicus) on a grass diet. As predicted, both species achieved similar dry matter, organic matter, crude protein and gross energy digestibilities. However, neutral and acid detergent fibre digestibility was lower in pygmy hippos. Apparently, in these species, fibre digestibility was more influenced by particle size, which was larger in pygmy hippos compared to banteng, than by retention time. In spite of their higher relative food intake, the banteng in this study did not have greater relative gut fills than the hippos. Ruminants traditionally appear intake-limited when compared to equids, because feed particles above a certain size cannot leave the rumen. But when compared to nonruminating foregut fermenters, rumination seems to free foregut fermenters from an intrinsic food intake limitation. The higher energy intakes and metabolic rates in wild cattle compared to hippos could have life-history consequences, such as a higher relative reproductive rate.     

Retention of solute and particle markers in the digestive tract of captive Somali wild asses (<Emphasis Type="Italic">Equus africanus somaliensis</Emphasis>)

In contrast to the domestic horse, whose digestive physiology has been thoroughly investigated, knowledge on the digestive physiology of wild equids is scarce. Comparisons between the domestic horse and the domestic donkey suggest that wild asses might achieve higher digestibilities. This could derive from longer retention times or a greater difference in the mean retention time (MRT) of particles vs. fluid (the selectivity factor (SF)). Here, we measured MRT of a solute (fluid; MRT_solute) and a particle (<2 mm; MRT_particle) marker in five captive male Somali wild asses (Equus africanus somaliensis) fed a diet of 95% grass hay. At a mean dry matter intake of 94 ± 3 g kg^?0.75 day^?1, MRT_solute was 33.3 ± 5.4 h and MRT_particle 39.6 ± 3.9 h, resulting in a SF of 1.21 ± 0.14. For their food intake, Somali wild asses appeared to have slightly higher MRT_particle than expected based on domestic equid data, in contrast to Grevy zebras (Equus grevyi), potentially indicating higher capacities of the digestive tract. However, considering data on domestic horses, donkeys, and zebra, there was no evident difference in the SF of wild equids compared to domestic ones. Together with an absence of reported anatomical differences in the digestive tract of wild and domestic equids, the data suggest a general similarity in the digestive physiology of equid species that contrasts with the diversity in the digestive physiology of ruminants, and that might be one contributing factor to a lack of sympatric, niche-differentiated equid species.     

The influence of natural diet composition, food intake level, and body size on ingesta passage in primates

An important component of digestive physiology involves ingesta mean retention time (MRT), which describes the time available for digestion. At least three different variables have been proposed to influence MRT in herbivorous mammals: body mass, diet type, and food intake (dry matter intake, DMI). To investigate which of these parameters influences MRT in primates, we collated data for 19 species from trials where both MRT and DMI were measured in captivity, and acquired data on the composition of the natural diet from the literature. We ran comparative tests using both raw species values and phylogenetically independent contrasts. MRT was not significantly associated with body mass, but there was a significant correlation between MRT and relative DMI (rDMI, g/kg(0.75)/d). MRT was also significantly correlated with diet type indices. Thus, both rDMI and diet type were better predictors of MRT than body mass. The rDMI-MRT relationship suggests that primate digestive differentiation occurs along a continuum between an "efficiency" (low intake, long MRT, high fiber digestibility) and an "intake" (high intake, short MRT, low fiber digestibility) strategy. Whereas simple-stomached (hindgut fermenting) species can be found along the whole continuum, foregut fermenters appear limited to the "efficiency" approach.     

Variation in digestive performance between geographically disjunct populations of Atlantic salmon: countergradient in passage time and digestion rate

European Atlantic salmon (Salmo salar) populations inhabit rivers from northern Portugal to northern Norway across a wide spectrum of environmental variability. To address whether single physical factors might lead to genetic divergence of isolated populations, we compared the digestive performances total digestibility, relative nitrogen digestibility, passage time, and digestion rate (g dry matter · h^–1) — of northern (Scotland) and southern (Asturias, northern Spain) populations at three temperature regimes (5, 12, and 20° C). Total dry matter digestibilities increased directly with temperature but were similar for both populations at each of the three trials. Relative nitrogen digestibility did not differ between populations nor among temperature regimes. In contrast, passage time was significantly longer for low-than for high-latitude fish at both 5 and 20° C. When the percentage of food digested and the passage time were integrated as digestion rates (food digested per unit time), a significant population × temperature interaction consistent with a genotype × environment interaction was detected in addition to the population and temperature effects. This implies that not only is the digestive performance of the high-latitude population higher throughout the range of temperatures examined, but moreover the difference is reinforced at high temperatures, where the digestion rate of high-latitude fish was 1.6 times greater. Taken together, these two results provide preliminary evidence for countergradient variation in digestive rates of salmonids in response to variation in growth opportunity. The data support our previous work on the same two populations showing differences in growth rates, and underlie one of the possible mechnisms leading to more rapid growth of the high-latitude fish when both populations are reared in a common environment.     

Seasonal energy utilization in bamboo by the red panda (Ailurus fulgens)

Seasonal energy intake and digestion of bamboo by red pandas were measured in energy balance trials at the field station at an altitude of 3,100 m in Yele Natural Reserve, Mianning County of Sichuan Province, Peoples Republic of China. Two‐week trials were conducted at monthly intervals by feeding animals with only natural bamboo browse, Bashania spanostachya, from May to November 1995. Results demonstrated that energy intake, digestible energy, and digestibility differed significantly among seasons and diets. Energy digestibilities were high in summer‐autumn, intermediate in spring, and low in winter. These variations correlated positively with the nutrient contents of bamboo. The daily metabolic energy requirement measured varied from 2,603.3 kJ in the spring to 3,139.8 kJ in the summer‐autumn to 2,740.8 kJ in the winter. The results also showed that shoots were more easily digested than leaves and exhibited the highest digestibility. Restricted by the capacity of digestive tracts, red pandas poorly utilized the energy provided by bamboo as indicated by the lower energy digestibility (averaging 26.47% for leaves and 44.08% for shoots among the seasons). To fulfill their daily energy requirements, red pandas ingested a large amount of energy from bamboo leaves, which varied from 10,145.8 kJ in the spring, 12,045.1 kJ in the summer‐autumn, and 12,276.9 kJ in the winter to maximize the rate of energy intake. Zoo Biol 19:27–33, 2000. © 2000 Wiley‐Liss, Inc.     

A comparative study of digestion in North Atlantic seabirds

We present data on digestive efficiencies and gut retention times of eight North Atlantic seabird species, fed on two fish species – lesser sandeel Ammodytes marinus and whiting Merlangius merlangus – which commonly occur in the diet of wild seabirds. In an interspecific comparison, there was a positive relationship between retention time and digestive efficiency, which we suggest represents a trade-off between conflicting benefits of efficient digestion and rapid digestion. Analysis of excretion curves revealed that retention time of digesta in the stomach was more important than passage time of digesta through the intestine in determining whole gut retention time. Differences in stomach retention time of lesser sandeel and whiting explained the longer overall retention time of the latter diet. Stomach retention time and whole gut retention time were greater in species with relatively large stomachs, while intestine passage time was correlated with relative intestine length. Species which typically eat a wide range of food types, including low quality items, tended to have slow and efficient digestion and heavy stomachs, whereas species which specialise on readily digestible and energy dense food types had the opposite digestion strategy.     

Comparison of gastrointestinal transit times between chickens from D+ and D- genetic lines selected for divergent digestion efficiency

D+ (high digestion efficiency) and D- (low digestion efficiency) genetic chicken lines selected for divergent digestion efficiency were compared in this experiment. Gizzard functions were tested in terms of digesta mean retention time and reactions to high dilution of a corn diet with 15% coarse sunflower hulls. The corn standard (S) and high fibre (F) experimental diets were given from 9 days of age to chickens from both lines. Besides the measurements of growth efficiencies (9 to 20 days), digestibilities (20 to 23 days) and gut anatomy (0, 9, 29, 42 and 63 days), two digestive transit studies were performed at 9 and 29 days of age. For the transit studies, the S and F diets were labelled with 0.5% TiO2 and 1% Cr-mordanted sunflower hulls. These diets were fed ad libitum during 3 days, and then the birds were euthanized. The digestive contents were analysed for the determination of marker concentrations and mean retention times (MRTs) in digestive compartments (crop + oesophagus, proventriculus + gizzard, duodenum + jejunum, ileum, rectum + cloaca and caeca) were determined. D+ birds were confirmed as better digesters than D- birds during the growth period, in association with larger gizzard and pancreas, and lighter small intestine in D+ than in D-birds. The MRT in the proventriculus-gizzard system, higher in D+ than in D- birds, was a major factor associated with differences between D+ and D- birds regarding digestion efficiencies and gut anatomy. Diet dilution with fibres reduced differences in digestion efficiencies and proventriculus-gizzard MRT between lines. Differences in gut anatomy between lines tended to disappear after 8 weeks of age. In conclusion, this study showed that MRT in the proventriculus-gizzard system was a major factor associated with genotype differences between the D+ and D- genetic chicken lines selected for divergent digestion efficiency, with longer MRT found in D+ than in D- birds.     

Digestion in the common marmoset (Callithrix jacchus), a gummivore–frugivore

Wild common marmosets (Callithrix jacchus) feed on fruits, insects, and gums, all of which provide different digestive challenges. Much of the ingested mass of fruits consists of seeds. In general, seeds represent indigestible bulk to marmosets and could inhibit feeding if they are not eliminated rapidly. In contrast, gums are β‐linked polysaccharides that require microbial fermentation. Their digestion would benefit from an extended residence time within the gut. Earlier research found that mean retention time (MRT) for a liquid digestive marker (cobalt EDTA) was significantly longer than MRT for a particulate marker (chromium‐mordanted fiber), suggesting that common marmosets preferentially retain liquid digesta. We conducted two four‐day‐long digestion trials on 13 individually housed adult common marmosets fed a single‐item, purified diet in order to examine the relations among MRT of cobalt EDTA and chromium‐mordanted fiber, food dry matter intake (DMI), and apparent digestibility of dry matter (ADDM). We compared the MRT values with the data from the previous study mentioned above and a study using polystyrene beads. There were no significant correlations among MRT, ADDM, or DMI, although increases in DMI between trials were associated with decreases in MRT. ADDM was consistent within individuals between trials; but the mean values ranged from 75.0 to 83.4% among individuals. We found no difference in MRT between the liquid (17.5±1.6 hr) and particulate (17.9±1.4 hr) markers. Although these values were not significantly different than found previously, the MRT for chromium‐mordanted fiber tended to be longer. This probably reflects the relatively small size of the chromium‐mordanted fiber particles used in this study. An inverse relationship between particle size and MRT was evident; the mean MRT of polysterene beads, the largest marker, was only 8.3±1.5 hr. Marmosets appear to retain liquids and small particles within the gut longer than large particles. Am. J. Primatol. 71:957–963, 2009. © 2009 Wiley‐Liss, Inc.     

The use of dosed and herbage n-alkanes as markers for the determination of digestive strategies of captive giraffes (Giraffa camelopardalis)

Selected aspects of digestion in captive giraffes were investigated in two trials with a type of marker that is new for digestive studies in non-domestic species. N-Alkanes were used as internal and external markers. In Trial 1, diet composition, intake, and digestibility were directly measured and estimated with the marker. Six giraffes were dosed once daily for 3 weeks with labeled pellets containing 3,800 ppm of each C₂₈, C₃₂, and C₃₆ alkanes at ˜100 mg/100 kg bodyweight. Intake of cabbage, browse, and a cattle pellet could accurately be estimated with the alkane method. For an oat/wheat mix and clover hay, however, there were large differences between directly measured intakes and those obtained using the alkane method, with the alkane method substantially underestimating. In Trial 2 mean retention times (MRTs) of fluid and particle phase were estimated. Three subadult giraffes were dosed once with cobalt-ethylenediamine tetraacetic acid (Co-EDTA) (260 mg) and alkanes C₂₉ (140 mg) and C₃₆ (370 mg). The average MRT for the fluid phase (Co-EDTA) was 31 hr and for the particle phase (alkanes) was 41 hr (C₃₆) and 43 h (C₂₉). This study showed that n-alkanes have potential as markers for investigating digestive strategies in non-domestic herbivores. Zoo Biol 17:295–309, 1998. © 1998 Wiley-Liss, Inc.     

Digesta retention time and recovery rates of ants and termites in Chinese pangolins (Manis pentadactyla)

Pangolins are myrmecophagous mammals whose biology and ecology remain poorly studied. Termite mandibles and ant head capsules are the two primary remains found in pangolin feces. Determining the retention time of insect cuticles is important for understanding the digestive physiology of pangolins, while determining the recovery rate of termites and ants in feces is required to estimate the number of these prey items that are consumed by pangolins. In this study, the authors conducted feeding trials with captive Chinese pangolins (Manis pentadactyla). Sixty grams of the fungus-growing termite Odontotermes formosanus (18,816 individuals) and 15–20 g of the yellow crazy ant Anoplolepis gracilipes (14,400–19,200 individuals) were fed to each pangolin. After feeding, pangolin feces were collected daily for 1 week. The authors also assessed the accuracy of using chromium (III) oxide powder (Cr₂O₃) as a proxy for determining gut passage time, as has been done in previous studies. The results showed that remaining termite mandibles and ant head capsules in feces peaked at 66 and 90 hr after feeding and their recovery rates were 0.35 ± 0.10 and 0.65 ± 0.04, respectively. In both feeding trials, the retention time of Cr₂O₃ was much shorter than that of the termite mandibles and ant head capsules, indicating that Cr₂O₃ is not an appropriate indicator for estimating food retention time of myrmecophagous animals. Our results revealed that the ant head capsules were preserved better in feces compared with the termite mandibles, suggesting that termites may be considerably underestimated in the feces of wild pangolins.     

Nutritional ecology of dimorphic herbivores: digestion and passage rates in Nubian ibex

We compared forage digestion and passage rates among three groups of Nubian ibex (Capra ibex nubiana) — mature males, non-lactating females, and lactating females — to test hypotheses relating intraspecific digestive ability to body mass and reproduction costs. We hypothesized that large males (60 kg) would exhibit longer forage retention times and more complete digestion of fermentable cell walls than adult females (23 kg). We tested these predictions by measuring digestion and retention of a grass hay and an alfalfa hay, forages that exhibited contrasting rates and extents of cell wall digestion. Consistent with predictions, males retained both forages longer than non-lactating females. However, by substantially increasing gut fill, lactating females increased both intake and retention time with respect to non-lactating females. Contrary to predictions, all three groups digested the grass (66% digestible) and alfalfa hay (63%) equally well. Alfalfa cell wall was less digestible than that of grass hay (60% vs 69% digestible), and retention time of alfalfa was consistently, but not statistically significantly, shorter. Fiber digestion was not correlated with retention time, emphasizing the ability of behavioral processes to modify digestion rate. We postulate that females achieved their greater digestion rate by masticating forages much more thoroughly than males.