An adaptive benefit of facultative coprophagy in the German cockroach Blattella germanica

1. A possible adaptive benefit of coprophagy was investigated in nymphs of the German cockroach Blattella germanica (L.). 2. Newly ecdysed first instars, given no source of food other than conspecific faeces, survived significantly longer than first instars deprived of faeces. The faeces of adult males and females may be of different quality, however, because nymphs given female faeces were more likely to moult into the second stadium than nymphs given male faeces. 3. In contrast to first instars, second instars provided adult faeces survived only slightly longer than starved counterparts. Faecal feeding is therefore stage‐specific, as is the benefit derived from it. 4. The relationship between the nutrient composition of faeces and the survival of nymphs was also examined. First instars fed the faeces of adults that had been maintained on a high (50%) protein diet, died more slowly than first instars fed the faeces of adults that had been maintained on medium (22.5%) and low (5%) protein diets. Chemical analysis of faeces showed that the concentration of protein in adult faeces increased with the level of protein in the diet. Moreover, food choice assays showed that first instars, unlike adults, ingested more of the high‐protein diets. 5. These data support the idea that coprophagy is a stage‐specific adaptive behaviour that permits first instars to moult into the second stadium with minimal foraging.     

The metabolism of niacytin in the rat. Studies of the excretion of nicotinic acid metabolites

1. Nicotinic acid-deficient rats were given a dose of niacytin or an equivalent one of free nicotinic acid or hydrolysed niacytin. 2. The excretion of N'-methylnicotinamide and of tertiary nicotinoyl derivatives in urine showed that niacytin was not metabolized as free nicotinic acid, although hydrolysed niacytin was equivalent to free nicotinic acid. 3. Little or none of the niacytin dose was recovered as tertiary nicotinoyl derivatives in faeces. This result was not affected by fitting rats with tail-cups to prevent coprophagy. 4. At the high doses used niacytin restored the growth rate of the deficient animals because of a small degree of hydrolysis of the bound nicotinic acid.     

The energetics of coprophagy: a theoretical analysis

Lagomorphs, many rodents and some other small mammals eat their faeces during the part of the day when they are not foraging for fresh food. One of the possible benefits of this habit of coprophagy is that it may enable them to extract more energy from their food. A computer model is used to assess the likely benefits and explore their relationships to food, feeding rate and gut morphology. The predicted benefits are much larger for hindgut fermenters than for foregut fermenters, and especially large for hindgut fermenters with relatively small fermentation chambers. They are larger for poor foods (with lower proportions of cell contents) than for richer ones. At low feeding rates the energetic advantage of coprophagy may disappear if the faeces from food eaten during one feeding period emerge largely during the next, but this can be avoided by adjusting the rate of passage of gut contents during the intervening rest period.     

Effects of coprophagy on serum urea and the weight of the gastrointestinal tract of fed or fasted rats

Coprophagy can be minimized by fitting rat cages with metal grids which allow faecal pellets to pass through to the floor of the cage. When bedding was omitted overnight, the extent of coprophagy could be estimated from the weight of the droppings on the cage floor or the weight of the gastrointestinal (GI) tract removed from rats housed with or without grids. The effect of coprophagy was also demonstrated by the elevation of serum urea nitrogen in rats that consumed faeces. Therefore, precautions against coprophagy, or their absence, should be specified in all experimental protocols and reports.     

Evidence of coprophagy in freshwater zooplankton

1. Vertical transport of nutrients in sedimenting faecal material is greatly reduced by coprophageous organisms. Unfortunately, nearly all work on faecal production, sedimentation and coprophagy has dealt with copepods in marine ecosystems. Here, we report the first evidence of coprophagy in freshwater zooplankton from oligotrophic and eutrophic lakes. We used ¹⁴C‐labelled algae and faecal material to estimate the rates of algal clearance and coprophagy. 2. Measured feeding rates per individual on faecal material were similar (Daphnia pulex, D. rosea, Leptodiaptomus tyrelli) or even higher (D. lumholtzi) than filtering rates on phytoplankton. This finding does not necessarily implicate active selection of faeces over algae because: (i) we did not use the same food concentrations for faeces and algae, and (ii) grazers of slightly different sizes were used in each test. 3. Weight‐specific clearance rates of L. tyrelli and Holopedium gibberum on faecal matter (0.084–0.089 mL μg^?1 h^?1) were higher than in the daphniids (0.026 mL μg^?1 h^?1). 4. The data indicate that coprophagy in freshwater ecosystems is an important mechanism of nutrient recycling, and this process should be taken into account when studying nutrient fluxes within lakes and reservoirs.     

Reduction of cyanobacterial toxins through coprophagy in Mytilus edulis

An experiment was conducted to follow the fate of the cyanobacterial toxin, nodularin, produced by Nodularia spumigena through ingestion by Mytilus edulis and re-ingestion of faecal material (coprophagy). Mussels were fed with cultures of N. spumigena, and the faeces that were produced were fed to other mussels not previously exposed to N. spumigena. Concentrations of nodularin were measured in the food (N. spumigena), the mussels and in the faeces in order to make a toxin budget. High concentrations of nodularin were found in the mussels and their faeces after 48 h incubation with N. spumigena. When the toxic faeces were fed to new mussels, the toxin content of faeces was reduced from 95 μg nod g⁻¹ dry weight (DW) to 1 μg nod g⁻¹ DW through the process of coprophagy. Hence, when toxic faeces were fed to mussels, the nodularin concentration of the resulting faecal material was reduced by 99%. Pseudofaeces were produced when the mussels were grazing on N. spumigena, but not when grazing on faeces. The pseudofaeces contained high concentrations of nodularin and apparently intact N. spumigena cells. However, these cells were growth-inhibited and their potential contribution to seeding a bloom is probably limited. Our data indicate that a large fraction of ingested nodularin in M. edulis is egested with the faeces, and that the concentration of nodularin in the faeces is reduced when faeces are re-ingested.     

Demonstrating coprophagy with passage markers? The example of the plains viscacha (Lagostomus maximus)

Coprophagy, or the ingestion of a certain fraction of the faeces, is a well-known strategy of small herbivores. However, the question of whether a particular species actually uses this cryptic strategy or not is not easily answered experimentally. In this study we introduce the use of ingesta passage markers as a tool to verify coprophagy in species where individual observation might be difficult. In two captive adult plains viscachas (Lagostomus maximus), we demonstrate recurrent excretion peaks for both a fluid (cobald-EDTA) and a particle (chromium-mordanted fibre) marker, the most parsimonious explanation for which is faeces re-ingestion. In addition, a literature review of graphical presentations of passage marker excretion revealed such recurrent marker peaks in a large number of studies; however, these observations were rarely explicitly contributed to a coprophagic digestion strategy. The widespread use of semi-logarithmic plots or cumulative marker excretion curves in displaying passage studies makes recurrent marker excretion peaks less evident. We conclude that passage markers offer a comparatively easy way of assessing the incidence of coprophagy. Due to the reported absence of coprophagy in rabbits, a species well-known for using this strategy, on high-protein, low-fibre diets, it is recommended that investigations of the occurrence of coprophagy should be made with animals fed challenging, high-fibre diets.     

Digestive physiology of wild capybara

The capybara (Hydrochaeris hydrochaeris) is a hindgut fermenter whose digestive efficiency is comparable to that of ruminants on similar diets. It is an interesting case for study because it is the largest caecum fermenter and uses coprophagy as part of its digestive strategy. It practices coprophagy in the early morning and forages and defaecates in the evening. Its anatomy is well known but the limited information available about its digestive physiology has been obtained from captive animals. In this work we studied the capybara's digestive physiology, using microbial and chemical information from samples taken from wild capybaras in the early wet season in the morning (0600–0700 hours), noon (1200–1300 hours) and evening (1800–1900 hours), key points in the digestive cycle. Bacteria (cellulolytic and non-cellulolytic) and protozoa were present in high concentrations in the caecum and colon. There were no significant differences in nitrogen concentrations between digestive tract compartments in the coprophagy period (0600 hours), but in the other two periods nitrogen concentrations were significantly higher in the caecum than in the stomach and colon. This is suggestive of selective retention of microbial cells with fluid digesta in the caecum and of cecotrophy (the production of two distinctly different kinds of faeces — one kind called cecotrophes formed from caecal contents and ingested). The capybara hindgut (caecum and colon) with its content, was heavier during the dry season (period of poor diet quality) than in the wet season, but there were no significant seasonal differences between the stomach or small intestine and their contents. This suggests changes in the capacity of the hindgut, the site of microbial fermentation, related to seasonal variation in resource quality.Abbreviations NDF neutral-detergent fibre - SCFA Short-chain fatty acid(s)     

The role of coprophagy in the feeding strategies of terrestrial isopods

Summary The hypothesis that faeces recycling in isopods evolved as an adaption to facilitate maintenance of an adequate copper balance in terrestrial environments is examined. Experimental observations on the consumption, absorption and growth rates of Porcellio scaber fed Betula pendula leaf litter varying in copper content and extent of microbial decay are reported. Preventing the isopods from reingesting their faeces caused a reduction in the growth rates of experimental animals fed their natural low copper diet but also of those fed copper enriched diets. When the availability of copper in the primary food was increased consumption of the litter decreased and growth rates were significantly reduced. These results suggest that copper is not normally a critically limiting nutrient for terrestrial isopods.When the primary diet was supplemented with shredded carrot, faeces deprivation did not cause a decrease in growth rates. These experimental animals gained weight significantly faster than controls fed decaying leaf litter alone.Faeces formed a significantly greater proportion of the diet when the animals were fed freshly fallen rather than decayed litter.We conclude a) that enhanced microbial activity in the faeces increases their nutrient status in such a way that some coprophagy is necessary in order to optimize overall nutrient uptake, and b) that theability to vary the extent to which faeces are recycled in response to differences in food quality is important in that it introduces greater flexibility into the feeding strategies of these generalist macro-decomposers.     

Interspecific coprophagia by wild red foxes: DNA metabarcoding reveals a potentially widespread form of commensalism among animals

Vertebrate animals are known to consume other species'' faeces, yet the role of such coprophagy in species dynamics remains unknown, not least due to the methodological challenges of documenting it. In a large‐scale metabarcoding study of red fox and pine marten scats, we document a high occurrence of domestic dog DNA in red fox scats and investigate if it can be attributed to interspecific coprophagia. We tested whether experimental artifacts or other sources of DNA could account for dog DNA, regressed dog occurrence in the diet of fox against that of the fox’s main prey, short‐tailed field voles, and consider whether predation or scavenging could explain the presence of dog DNA. Additionally, we determined the calorific value of dog faeces through calorimetric explosion. The high occurrence of dog DNA in the diet of fox, the timing of its increase, and the negative relationship between dog and the fox''s main prey, point to dog faeces as the source of DNA in fox scats. Dog faeces being highly calorific, we found that foxes, but not pine martens, regularly exploit them, seemingly as an alternative resource to fluctuating prey. Scattered accounts from the literature may suggest that interspecific coprophagia is a potentially frequent and widespread form of interaction among vertebrates. However, further work should address its prevalence in other systems and the implications for ecological communities. Tools such as metabarcoding offer a way forward.     

Analyses of constituents of feces and the effect of a vitamin B12 fortified diet on coprophagy in the mouse

In order to investigate coprophagy from the viewpoint of nutrition, fecal constituents were analyzed in freeze-dried samples. Feces were collected from 7:00 to 11:00 and from 19:00 to 23:00. Inorganic elements and crude fibers per unit weight were 3-4 times more concentrated in feces than in basal diet, whereas, crude proteins, crude fats and nitrogen-free extract showed various degrees of reduction. There were no differences in these tendencies with sampling time. As for some B vitamins, feces collected from 7:00 to 11:00 contained 22-92% more vitamins than feces collected from 19:00 to 23:00. In comparison with the dietary concentration, vitamin B12 was increased by 124-197 times (520-730 micrograms/100 g) in feces collected between 7:00 and 11:00. Folic acid in feces collected between 7:00 and 11:00 was 10 times greater than that in the diet. On the basis of the findings on vitamins, the effect of a vitamin B12 fortified diet (1,350 micrograms/100 g) on coprophagy was examined. Mean frequency of coprophagy per animal per day was 9.6 when animals were fed on the basal diet, whereas the frequency was immediately and significantly (p less than 0.05 approximately p less than 0.01) reduced to 4.7 after the diet had been replaced by the fortified one. However, coprophagy was not completely inhibited by vitamin B12 fortification. This indicates that some nutrient(s) in feces other than vitamin B12 might be of use to the host, and that otherwise, coprophagy might be a basically habitual form of behavior. Furthermore, under the fortified diet, the frequency of coprophagy increased gradually.(ABSTRACT TRUNCATED AT 250 WORDS)     

Effects of prevention of coprophagy on pregnant mice--is coprophagy beneficial on a balanced diet?

The effects of prevention of coprophagy on reproductive performance were examined in ICR mice. Females were treated with restrainers in order to prevent them from ingesting their feces from day 1 through day 17 of pregnancy. The restrained animals fed a commercial diet did not show any clear adverse effects. In contrast, restrained dams fed a purified diet deficient in vitamin B12 exhibited stillbirths (14%) and abortions (7%). Restrained dams fed a diet lacking in vitamin B12 and folic acid also experienced frequent abortions (27%). In addition, six out of 14 restrained dams (43%) aborted when fed a vitamin B complex-deficient diet. Sham-restrained animals, fed the vitamin B complex deficient-diet, but able to ingest their feces trapped by smaller-mesh floors, escaped these adverse effects. Sham-restrained animals fed the commercial diet, however, showed only a slight improvement in their reproductive performance. In conclusion, coprophagy has nutritional significance as long as the diet is lacking at least B vitamins, especially vitamin B12 and folic acid, whereas it almost entirely loses its nutritional significance when the mouse has access to a balanced diet such as the one made available to the laboratory mice in the present study.     

Coprophagy in iron-deficient rats: II. 2 novel methods of prevention

2 novel methods of coprophagy prevention were used in iron-deficient rats to measure their effect on haemoglobin regeneration during the feeding of ferrous sulphate or several soya proteins. Neck collars produced considerable weight loss and depression in food intake compared to aluminium anal cups which were tolerated well. Both methods of coprophagy prevention reduced iron availability from a range of soya proteins and ferrous sulphate and it is suggested that the practice of coprophagy in iron-deficient and possibly iron-replete rats may overestimate iron (and possibly other mineral) absorption (retention) studies.     

Stability of the Lactobacillus population in feces and stomach contents of rats prevented from coprophagy

Lev, Meir (Albert Einstein College of Medicine, New York, N.Y.), Raymond H. Alexander, and Stanley M. Levenson. Stability of the Lactobacillus population in feces and stomach contents of rats prevented from coprophagy. J. Bacteriol. 92: 13-16. 1966.-Lactobacilli were enumerated in the feces of rats prevented from coprophagy by tail-cupping. No differences were found when numbers of these organisms were compared with lactobacilli in feces of control rats, without tail-cups. High and similar numbers of lactobacilli were found in the stomachs of rats with and without tail-cups. The effect of coprophagy on fecal lactobacilli was therefore negligible.     

The relationship between forage material and levels of coprophagy in captive chimpanzees (Pan troglodytes)

Although coprophagy is practiced in the wild by chimpanzees (Pan troglodytes), it occurs more frequently and under more varied circumstances in captivity. This study was designed to determine if different forage materials and amount of residual undigested grain particles found in the feces might cause an increase in coprophagous behavior in those animals which already exhibited the behavior. A possible effect of availability of seed pits and fibrous leaves for “wadge” making, a typical chimpanzee behavior, on levels of coprophagy was also considered. Observations for coprophagous behavior were conducted on 65 juvenile, adolescent, and adult chimpanzees. Coprophagy levels were significantly lower with popcorn than either chicken scratch or sweet feed. A significant increase in coprophagy was noted for all weeks of forage types when tested against the wadge weeks. Residual grain content analysis showed no significant difference in coprophagous behavior between any of the testing conditions. Decreasing levels of coprophagous behaviors may be assisted by the provision of wadge materials. © 1992 Wiley-Liss, Inc.     

Cannibalism and Coprophagy Are Modes of Transmission of Blastocrithidia triatomae (Trypanosomatidae) between Triatomines

Transovarial transmission was not detectable among Blastocrithidia triatomae- infected Triatoma infestans . Rather, B. triatomae was transmitted directiy between triatomines by cannibalism and coprophagy. Cannibalism conditions that excluded coprophagy always resulted in an infection of Dipetalogaster maxima . The efficiency of transmission was not influenced by the blood source—mice or chickens—fed to the infected donor bugs although chicken blood lyses the epimastigotes of the stomach population. Triatoma infestans was infected by coprophagy only if fed, not if unfed. Blastocrithidia triatomae in dry feces was taken up only if the feces were redissolved in fresh feces. Infections also appeared in groups of bugs fed on chickens previously used for feeding infected bugs.     

Cannibalism and coprophagy are modes of transmission of Blastocrithidia triatomae (Trypanosomatidae) between triatomines

Transovarial transmission was not detectable among Blastocrithidia triatomae-infected Triatoma infestans. Rather, B. triatomae was transmitted directly between triatomines by cannibalism and coprophagy. Cannibalism conditions that excluded coprophagy always resulted in an infection of Dipetalogaster maxima. The efficiency of transmission was not influenced by the blood source--mice or chickens--fed to the infected donor bugs although chicken blood lyses the epimastigotes of the stomach population. Triatoma infestans was infected by coprophagy only if fed, not if unfed. Blastocrithidia triatomae in dry feces was taken up only if the feces were redissolved in fresh feces. Infections also appeared in groups of bugs fed on chickens previously used for feeding infected bugs.     

Ecology of the coprophagous moth Trisyntopa neossophila Edwards (Lepidoptera: Oecophoridae)

Trisyntopa neossophila Edwards (Lepidoptera: Oecophoridae) is an unusual moth whose breeding cycle is closely synchronised with a termite mound nesting parrot of northern Australia; the hooded parrot ( Psephotus dissimilis Collet). T. neossophila is one of three coprophagous, nest dwelling moths in the genus Trisyntopa Lower 1918. True coprophagy is rare in the Lepidoptera, although some species occasionally consume faeces to gain rare nutrients. We present observations of the life cycle of T. neossophila , a moth that lays its eggs in the nest of a hooded parrot, so that larvae hatch in synchrony with the hatching of the parrot's eggs. The larvae spend their larval period in the nest and exclusively consume the excrement of the nestling parrots. When the parrot chicks fledge, the larvae move to the walls of the nest cavity to pupate, emerging the following wet season to repeat the process during the next parrot breeding season.     

Variables influencing the origins of diverse abnormal behaviors in a large sample of captive chimpanzees (Pan troglodytes)

The developmental origin of abnormal behaviors is generally associated with early rearing environments that lack sufficient physical and sensory stimulation. However, other factors should also be considered. A large sample of captive chimpanzees (128 males and 140 females) was surveyed for the presence or absence of 18 abnormal behaviors. Origin variables included the subject's source (zoo, pet, performer, or laboratory), rearing (mother‐ or hand‐reared), and sex. Animals were assessed while held at the Primate Foundation of Arizona, University of Texas M. D. Anderson Cancer Center, or White Sands Research Center. There was a confound among origin variables; more hand‐reared animals than expected were from laboratories. Logistic regression tested the relationship of rearing and source, with sex as a secondary predictor variable, to each of the abnormal behaviors. There was no clear association between any abnormal behavior and source. However, for coprophagy, relative to animals from the laboratory, zoo animals tended to show a higher prevalence, while performers tended to show a lower prevalence (when rearing and sex were controlled). Rocking and self‐sucking were significantly more likely in hand‐reared animals. Coprophagy and depilation of self were significantly more likely in mother‐reared animals. When rearing and source were statistically controlled, the only significant sex difference was a higher prevalence of coprophagy in females and a higher prevalence of rocking in males. In a second, smaller sample of 25 males and 33 females from Southwest Foundation for Biomedical Research, no significant sex association was found for coprophagy, urophagy, rocking, or self‐depilation. In this second sample, coprophagy was also significantly more likely in mother‐reared than hand‐reared subjects. The association of some abnormal behaviors with mother‐rearing suggests that some form of social learning may be involved in the origin of some of these behavior patterns. This indicates that some abnormal behaviors may not be always be indicative of reduced psychological well‐being in captive chimpanzees. Am. J. Primatol. 48:15–29, 1999. © 1999 Wiley‐Liss, Inc.     

Lifelong coprophagy in male mice

Changes in coprophagy with age were investigated in male ICR mice during their life span. Sucklings showed coprophagy at 17 to 18 days old, i.e., a few days after they began to excrete feces autonomously. The number of fecal pellets ingested peaked at 5 to 6 weeks old (13 pellets/day) and gradually decreased, thereafter (2.1 pellets at 78 weeks old, 1.5 pellets at 104 weeks old). The diurnal pattern of coprophagy also changed with age. Growing mice showed vigorous coprophagous activity in both light and dark phases, whereas animals over 30 weeks old exhibited less activity in both phases, especially in the dark phase. Feces proved to be abundant in vitamin B12 and folic acid throughout the life span. These results suggest that the frequency of coprophagy changes in association with the nutritional requirements of mice during the process of growth or aging.