The Ruminant sorting mechanism protects teeth from abrasives.

Dental wear due to ingestion of dust and grit has deleterious consequences. Herbivores that could not wash their food hence had to evolve particularly durable teeth, in parallel to the evolution of dental chewing surface complexity to increase chewing efficacy. The rumen sorting mechanism increases chewing efficacy beyond that reached by any other mammal and has been hypothesized to also offer an internal washing mechanism, which would be an outstanding example of an additional advantage by a physiological adaptation, but in vivo evidence is lacking so far. Here, we investigated four cannulated, live cows that received a diet to which sand was added. Silica in swallowed food and feces reflected experimental dietary sand contamination, whereas the regurgitate submitted to rumination remained close to the silica levels of the basal food. This helps explain how ruminants are able to tolerate high levels of dust or grit in their diet, with less high-crowned teeth than nonruminants in the same habitat. Palaeo-reconstructions based on dental morphology and dental wear traces need to take the ruminants' wear-protection mechanism into account. The inadvertent advantage likely contributed to the ruminants' current success in terms of species diversity.

Standard addition method for rapid, cultivation-independent quantification of Legionella pneumophila cells by qPCR in biotrickling filters.

Cultivation-independent molecular biological methods are essential to rapidly quantify pathogens like Legionella pneumophila (L. pneumophila) which is important to control aerosol-generating engineered water systems. A standard addition method was established to quantify L. pneumophila in the very complex matrix of process water and air of exhaust air purification systems in animal husbandry. Therefore, cryopreserved standards of viable L. pneumophila were spiked in air and water samples to calibrate the total bioanalytical process which includes cell lysis, DNA extraction, and qPCR. A standard addition algorithm was employed for qPCR to determine the initial concentration of L. pneumophila. In mineral water, the recovery rate of this approach (73%-134% within the concentration range of 100-5000 Legionella per mL) was in good agreement with numbers obtained from conventional genomic unit (GU) calibration with DNA standards. In air samples of biotrickling filters, in contrast, the conventional DNA standard approach resulted in a significant overestimation of up to 729%, whereas our standard addition gave a more realistic recovery of 131%. With this proof-of-principle study, we were able to show that the molecular biology-based standard addition approach is a suitable method to determine realistic concentrations of L. pneumophila in air and process water samples of biotrickling filter systems. Moreover, this quantification strategy is generally a promising method to quantify pathogens in challenging samples containing a complex microbiota and the classical GU approach used for qPCR leads to unreliable results.

The effect of size and density on the mean retention time of particles in reindeer (Rangifer tarandus).

Particle passage from the reticulorumen (RR) depends on particle density and size. A classic way of assessing these effects is the use of plastic markers of varying density and size that are recovered in the faeces. Here, we report results of an experiment where four fistulated reindeer (Rangifer tarandus, 96 ± 12 kg) were fed two different diets (browse, voluntary dry matter intake [DMI] 70 ± 10 g/kg0.75/d; or a pelleted diet, DMI 124 ± 52 g/kg0.75/d) and dosed via fistula with 8 different particle types combining densities of 1.03, 1.22 and 1.44 g/ml and sizes of 1, 10 and 20 mm. Generally, particles that passed the digestive tract intact (not ruminated) did so relatively early after marker dosing, and therefore had shorter mean retention times (MRT) than ruminated particles. On the higher intake, the overall mean retention time (MRT) of particles was shorter, but this was not an effect of shorter MRT for either intact or ruminated particles, but due to a higher proportion of intact particles at the higher intake. This supports the concept that ruminants do not adjust chewing behaviour depending on intake, but that a lower proportion of digesta is submitted to rumination due to pressure-driven escape from the forestomach at higher gut fills. Compared to cattle (Bos primigenius taurus), muskoxen (Ovibos moschatus) and moose (Alces alces) that had received the same markers, reindeer had a lower proportion of 1 mm particles that passed intact. Our results support the concept that the critical size threshold for particles leaving the ruminant forestomach is dependent on body size. While the results likely do not represent findings peculiar for reindeer, they indicate fundamental mechanisms operating in the forestomach of ruminants.

Comparative study of feeding and rumination behaviour of goats and sheep fed mixed grass hay of different chop length.

Rumination is reported to be more pronounced in sheep compared to goats. This study compared the feeding and rumination behaviour of small ruminants and consisted of two experiments (E1 and E2). In E1, four sheep and four goats were offered low-quality hay (NDFom: 692 g/kg dry matter [DM]), processed to two chop lengths (long hay [LH]: 35 mm; short hay [SH]: 7 mm) in a 2 × 2 factorial (2 species × 2 chop lengths), cross-over design. In E2, the same animals were offered moderate-quality hay (NDFom: 636 g/kg DM) processed as LH and SH. Hay was offered for ad libitum consumption. Feeding and rumination behaviour was evaluated using video recordings. Aspects of rumination like chewing frequency were evaluated for 30 min per day. Faecal samples were analysed for faecal-N and particle size. There was no species effect on feed intake and organic matter digestibility (faecal N as proxy); however, goats consumed more LH than SH in E1 and E2. There was an effect of species on rumination:eating duration (R:E) ratio (higher in sheep) in E1 but not in E2, where there was a tendency for a species effect on rumination duration. In E1 and E2, sheep had a higher R:E ratio for SH than for LH. For rumination behaviour, there was a species effect for number of daily boli, chewing frequency and chews per day (more in sheep) in E1 and E2. No effect of species was found for faecal particle size. Despite much concordance, feed comminution behaviour differed in some aspects between sheep and goats. In an evolutionary context, a shift of significance of rumination could be triggered by a higher amount of abrasives in natural diets of sheep, rendering a shift of chewing towards ruminally prewashed material a rewarding strategy.

Rabbits (Oryctolagus cuniculus) increase caecal calcium absorption at increasing dietary calcium levels.

Hindgut fermenting herbivores from different vertebrate taxa, including tortoises, and among mammals some afrotheria, perissodactyla incl. equids, several rodents as well as lagomorphs absorb more calcium (Ca) from the digesta than they require, and excrete the surplus via urine. Both proximate and ultimate causes are elusive. It was suggested that this mechanism might ensure phosphorus availability for the hindgut microbiome by removing potentially complex-building Ca from the digesta. Here we use Ussing chamber experiments to show that rabbits (Oryctolagus cuniculus) maintained on four different diets (six animals/diet) increase active Ca absorption at increasing Ca levels. This contradicts the common assumption that at higher dietary levels, where passive uptake should be more prevalent, active transport can relax and hence supports the deliberate removal hypothesis. In the rabbits, this absorption was distinctively higher in the caecum than in the duodenum, which is unexpected in mammals. Additional quantification of the presence of two proteins involved in active Ca absorption (calbindin-D9K CB; vitamin D receptor, VDR) showed higher presence with higher dietary Ca. However, their detailed distribution across the intestinal tract and the diet groups suggests that other factors not investigated in this study must play major roles in Ca absorption in rabbits. Investigating strategies of herbivores to mitigate potential negative effects of Ca in the digesta on microbial activity and growth might represent a promising area of future research.

Putting zoo animal cancer into perspective.

As part of a comparative research agenda that promises insights that help extend the human lifespan and combat cancer, cancer prevalence in zoo animals has received recent attention. Here, we want to draw attention to a principle of cancer research that was introduced into the zoo world as early on as 1933, but that seems to have gone somewhat forgotten: Cancer is mainly a disease of old age, and therefore studies aiming at identifying taxa that are particularly susceptible or resistant to cancer must control for whether the respective zoo populations are 'old.' In a comparative context, 'old age' cannot be measured in absolute terms (e.g., years), but only in relation to a species' maximum lifespan: Species that achieve, across zoos, a higher mean lifespan as a percent of their maximum lifespan are 'older.' When applying this metric to former as well as more recently published data on cancer prevalence, it appears that those species that become relatively old in zoos-in particular, the carnivores-have a relatively high cancer prevalence. Any improvement in animal husbandry-which reduces premature deaths-should, by default, lead to more cancer. Cancer in zoo animals, like any other old-age condition, might therefore be embraced as a proxy for good husbandry. Rather than following a sensationalist approach that dramatizes disease and death per se, zoos should be clear about what their husbandry goals are, what relative longevities they want to achieve for which species, and what old-age diseases they should therefore expect: in the end, one has to die of something.

Breaking the spell: Changes in the behavior of two zoo-kept tigers (Panthera tigris) after exposure to a distinct feast-and-fast feeding regime.

The behavior of zoo carnivores has received intense attention due to their propensity for locomotor stereotypies. We observed two adult male tiger (Panthera tigris) siblings kept together for the duration of 104 days by round-the-clock video observation. The period consisted of three baseline periods with the zoo's regular feeding regime of five feeding days per week interrupted by two individual fasting days, with feeding occurring in the evening (B1-B3 of 14 days each). These periods were interrupted by two intervention periods (I1: randomized feeding times, 28 days; I2: gorge-feeding with three 10-day fasting periods, 34 days). As expected, day and night-time behavior was different, with the majority of sleep occurring at night. Pacing, which was mainly considered anticipatory, significantly decreased from 88 ± 132 min/day during B1 to 20 ± 33 min/day during B3. Pacing did not increase during the fasting days of I2. Over the course of whole study, lying time decreased and nonpacing locomotion increased. A major difference was observed between gorge-feeding and the subsequent first fasting days: during gorge-feeding, tigers spent a large part of the day feeding and locomoting (and less sleeping); on the subsequent day, they locomoted about 4.5 h less and slept about 4.3 h more. We suggest that interrupting routines by fasting periods of several days may be effective for reducing regular anticipatory behavior and creates an across-day structure that may correspond to the evolved psychological disposition of large carnivores.

Shape, size, and quantity of ingested external abrasives influence dental microwear texture formation in guinea pigs.

Food processing wears down teeth, thus affecting tooth functionality and evolutionary success. Other than intrinsic silica phytoliths, extrinsic mineral dust/grit adhering to plants causes tooth wear in mammalian herbivores. Dental microwear texture analysis (DMTA) is widely applied to infer diet from microscopic dental wear traces. The relationship between external abrasives and dental microwear texture (DMT) formation remains elusive. Feeding experiments with sheep have shown negligible effects of dust-laden grass and browse, suggesting that intrinsic properties of plants are more important. Here, we explore the effect of clay- to sand-sized mineral abrasives (quartz, volcanic ash, loess, kaolin) on DMT in a controlled feeding experiment with guinea pigs. By adding 1, 4, 5, or 8% mineral abrasives to a pelleted base diet, we test for the effect of particle size, shape, and amount on DMT. Wear by fine-grained quartz (>5/<50 µm), loess, and kaolin is not significantly different from the abrasive-free control diet. Fine silt-sized quartz (∼5 µm) results in higher surface anisotropy and lower roughness (polishing effect). Coarse-grained volcanic ash leads to significantly higher complexity, while fine sands (130 to 166 µm) result in significantly higher roughness. Complexity and roughness values exceed those from feeding experiments with guinea pigs who received plants with different phytolith content. Our results highlight that large (>95-µm) external silicate abrasives lead to distinct microscopic wear with higher roughness and complexity than caused by mineral abrasive-free herbivorous diets. Hence, high loads of mineral dust and grit in natural diets might be identified by DMTA, also in the fossil record.

Is there a difference in ruminal fermentation control between cattle and sheep? A meta-analytical test of a hypothesis on differential particle and fluid retention.

Ruminant species differ in digestive physiology. The species-specific ratio of mean retention time of particles and fluid (MRTparticle/MRTfluid) in the reticulorumen has been interpreted as controlling ruminal fermentation: a higher ratio indicates of a more distinct 'washing' of particulate digesta by liquid. This should increase the harvest of microbes from the reticulorumen, and keep the microbiome in a state of more intense growth; at the same time, this should increase the metabolic losses of faecal nitrogen of microbial origin, leading to lower values for the apparent digestibility of crude protein (aD CP). A systematic difference has been hypothesized between cattle (higher ratio) and sheep (lower ratio), with a lower MRTfluid in cattle due to a higher saliva production. Here, we test these hypotheses in a meta-analysis, using only studies that investigated cattle and sheep simultaneously. The datasets included 12 studies on MRT (of which 11 contained information on feed intake), yielding 102 (or 89) individual data; and 26 studies on protein digestibility (of which 18 contained information on intake), yielding 349 individual data. Cattle had a higher MRTparticle/MRTfluid (2.1) than sheep (1.7), mainly due to longer MRTparticle; only if body mass was included in the model, MRTfluid was significantly shorter in cattle in the larger MRT dataset (and tended to be shorter in the slightly smaller dataset). Cattle had a significantly lower aD CP than sheep, while there was no such difference in overall (dry or organic matter) digestibility. The dataset confirms a shift in fermentation strategy towards microbial production in cattle. While this has been suggested for ruminants in general, cattle appear particularly far on an evolutionary trajectory of maximizing microbial yield from the forestomach. The application of more specific digestive physiology data (like endogenous losses) gained from sheep to cattle should be done bearing these differences in mind.

Fluid and particle retention in a small New World and a small Old World cervid, the southern pudu (Pudu puda) and Reeves's muntjac (Muntiacus reevesi).

Ruminants differ in the pattern how small particles and liquids pass through their gastrointestinal tract, and in particular their reticulorumen (RR). Based on that they may be classified into 'moose-type' and 'cattle-type' species (smaller and larger differences between particle and liquid passage, respectively). The ratio between the retention of particles and fluids is called the 'selectivity factor' (SF) and is a species-specific characteristic, studied in tragulids, giraffids and bovids, but not in many cervid species. Recently, it has been suggested that a high SF might also serve to wash digesta clean of external abrasives prior to regurgitation for rumination. In this study, we measured SF and passage kinetics (using a liquid marker and markers of different particle size, fed with the diet) in a capreoline deer, the southern pudu (Pudu puda, n = 5, 10.3 ± 2.9 kg, kept at two zoos) and a cervine deer, the Reeves's muntjac (Muntiacus reevesi, n = 6, 11.0 ± 1.7 kg, kept at a research facility). The relative daily dry matter intake (38 ± 3 g/kg0.75 for pudu and 76 ± 5 g/kg0.75 for muntjac) was higher, and the mean retention times (MRT) correspondingly shorter (e.g., MRT small particles in the total digestive tract 39 ± 8 h for pudu and 15 ± 2 h for muntjac), in the muntjac. The SF for small particles/liquid in the reticulorumen were, however, similar for both species, at 1.47 ± 0.21 for pudu and 1.66 ± 0.20 for muntjac, indicating a 'moose-type' physiology for both, irrespective of their different phylogenetic origin. To date, SF recorded in bovids attain distinctively higher values than the few reported for cervids. This situation reflects the degree of hypsodonty (tooth crown height) attained by these taxa, which is higher in bovids than in cervids. Together, constraints in hypsodonty as well as SF might limit cervids to more mesic habitats without distinct loads of external abrasives (such as dust or grit) on their food. In both species, some animals showed the typical ruminant pattern of a longer MRT for large than for small particle markers, but in some animals, this difference was not evident. This may be due to variable degrees of marker chewing during ingestion.

Individual differences in digesta retention and their relation to chewing in cattle-A pilot investigation.

While information on individual differences in digesta mean retention time (MRT) might be interesting when selecting phenotypes for digestive efficiency, MRT measurements are prohibitively labour-intensive for large-scale application. Therefore, more easily measured proxies of MRT might be helpful. We used the opportunity of an experiment applying saliva stimulant in cattle to investigate the effect of different individual chewing behaviour on fluid and particle MRT with a consistent diet. Four non-lactating cattle (670-850 kg body mass [BM]) were used in a 4 × 4 Latin square design, treated with the saliva stimulant pilocarpine in dosages of 0, 1, 2.5 and 5 mg/kg BM per day. The cattle were fed hay with dry matter intake (DMI) assigned according to their metabolic body weight. MRT in the whole gastrointestinal tract (GIT), the reticulorumen (RR) and the distal tract were measured using Co-EDTA, Cr-mordanted fibre and La-mordanted fibre as markers representing fluid, small particles (2 mm) and large particles (1 cm), respectively. The chewing behaviour was measured via noseband pressure sensor and expressed as chewing frequency (chews per time) and chewing intensity (chews per DMI), both for total chewing (ingestion plus rumination) and rumination chewing alone. The animals differed considerably in chewing behaviour and MRT measures. BM did not show a significant effect on chewing behaviour and MRT measures, though it tended to negatively correlated to total chewing intensity. Chewing intensity exerted a significant negative influence on MRT of fluid and particles in the RR, which was not the case for chewing frequency. Chewing frequency showed a significant relationship with MRT of large particles in the GIT. We suggest that chewing behaviour could influence MRT in two ways: (i) by affecting saliva production via the masticatory-salivary reflex and subsequently, the fluid inflow to the RR; (ii) by contributing to particle size reduction. Should the link between chewing behaviour and MRT be corroborated in larger studies, chewing measures, with their large interindividual variation, could emerge as an easy-to-measure proxy for MRT characteristics.

Effect of induced saliva flow on fluid retention time, ruminal microbial yield and methane emission in cattle.

Both in vitro and animal studies indicated that a higher dilution rate is related to a more efficient microbial synthesis and a lower methane (CH4 ) yield. The latter could be a consequence of the former, as an increase in microbial cell synthesis offers an alternative hydrogen sink competing with methanogenesis. To test this assumption in live animals, we applied a saliva stimulant, pilocarpine, to modify liquid flow rate in cattle. Four non-lactating cows (750 ± 71 kg) were fed forage only (restricted to constant intake) in a 4 × 4 Latin square design with oral doses of 0, 1, 2.5 and 5mg pilocarpine/kg body weight and day. We quantified feed and water intake, ruminal and total tract mean retention time (MRT) of solute and particle markers, ruminal microbial yield (via urinary purine bases or metabolic faecal nitrogen), CH4 emission, digestibility, chewing behaviour, reticular motility and rumen fluid parameters. The effect of induced saliva flow was evident by visibly increased salivation and water intake. Increasing the pilocarpine dosages resulted in a linearly decreased MRT of fluid and small particles (p < 0.001 and p< 0.05, respectively) and methane yield as related to digested DM (p < 0.05), the latter at a magnitude of 5%. No effect of treatment was found on ruminal microbial yield estimated via purine derivates. Metabolic faecal N as an indicator of microbial growth linearly correlated with pilocarpine dosages (p < 0.05). No significant relationship was found between pilocarpine dosages and large particle MRT, nutrient digestibility, ruminal pH and short-chain fatty acids. In conclusion, different from some in vitro studies, there was little indication of a reciprocal effect of CH4 and microbial biomass production in cows fed a forage-only diet.

A pilot investigation on the effect of induced saliva flow on digestive parameters in sheep, and a comparison with cattle.

Sheep with a relatively low methane yield were observed to have shorter fluid and particle mean retention times (MRT). Because the application of pilocarpine, a saliva stimulant, was successful in reducing retention times in ruminants in previous studies, we applied this substance to sheep, expecting a reduction in MRT and methane yield. Three non-pregnant sheep (74 ± 10 kg) were fed a hay-only diet in a 3 × 3 Latin square design with oral doses of 0, 2.5 and 5 mg pilocarpine/kg body weight and day. Measurements included feed and water intake, MRT of liquid and particulate phases in the reticulorumen (RR) and total gastrointestinal tract (GIT), ruminal microbial yield (via urinary purine bases and metabolic faecal nitrogen), total tract methane emission, apparent nutrient digestibility and rumen fluid parameters. Data were investigated for linear and quadratic effects using orthogonal polynomial contrasts. The MRT of liquid and small particles in the RR and total GIT, and the short-chain fatty acid concentration in rumen fluid, linearly declined with increasing pilocarpine dosage, while no quadratic relationship was detected. Intake of feed DM and water, apparent nutrient digestibility, methane yield and microbial yield were not affected by pilocarpine. When combining the sheep data with that of a similar experiment in cattle, we found that the MRT of the liquid phase was positively associated with estimated NDF digestibility and with methane production per digested NDF, but was not associated with microbial yield or the ratio of acetate to propionate. The ratio between MRT of the particulate and the liquid phase was smaller for sheep than that for cattle, and was not affected by treatment. Differences in this ratio might explain why species reacted differently to the saliva-inducing agent, which might help to explain the discrepancy between species in the effect of induced saliva flow on digestive parameters.

From left to right all through the night: Characteristics of lying rest in zoo elephants.

Despite increased research during the past years, many characteristics of resting behavior in elephants are still unknown. For example, there is only limited data suggesting elephants express longer lying bouts and increased total nightly lying durations on soft substrates as compared to hard surfaces. Additionally, it has not been investigated how frequently elephants change body sides between lying bouts. Here we present these characteristics based on observations of nighttime lying behavior in 10 zoo elephants (5 African Loxodonta africana and 5 Asian Elephas maximus elephants) living in five different European facilities. We found that elephants housed on soft substrates have significantly increased total lying durations per night and longer average lying bouts. Furthermore, at 70%-85% of all bouts, a consistently higher frequency of side change between lying bouts occurred on soft substrates, leading to an overall equal laterality in resting behavior. Deviations from this pattern became evident in elephants living on nonsand flooring or/and in nondominant individuals of nonfamily groups, respectively. Based on our findings, we consider elephants to normally have several lying bouts per night with frequent side changes, given an appropriate substrate and healthy social environment. We encourage elephant-keeping facilities to monitor these characteristics in their elephants' nighttime behavior to determine opportunities for further improvements and detect alterations putatively indicating social or health problems in individual elephants at an early stage.

The historical development of zoo elephant survivorship.

In the discussion about zoo elephant husbandry, the report of Clubb et al. (2008, Science 322: 1649) that zoo elephants had a "compromised survivorship" compared to certain non-zoo populations is a grave argument, and was possibly one of the triggers of a large variety of investigations into zoo elephant welfare, and changes in zoo elephant management. A side observation of that report was that whereas survivorship in African elephants (Loxodonta africana) improved since 1960, this was not the case in Asian elephants (Elephas maximus). We used historical data (based on the Species360 database) to revisit this aspect, including recent developments since 2008. Assessing the North American and European populations from 1910 until today, there were significant improvements of adult (≥10 years) survivorship in both species. For the period from 1960 until today, survivorship improvement was significant for African elephants and close to a significant improvement in Asian elephants; Asian elephants generally had a higher survivorship than Africans. Juvenile (<10 years) survivorship did not change significantly since 1960 and was higher in African elephants, most likely due to the effect of elephant herpes virus on Asian elephants. Current zoo elephant survivorship is higher than some, and lower than some other non-zoo populations. We discuss that in our view, the shape of the survivorship curve, and its change over time, are more relevant than comparisons with specific populations. Zoo elephant survivorship should be monitored continuously, and the expectation of a continuous trend towards improvement should be met.

Historical development of the survivorship of zoo rhinoceroses-A comparative historical analysis.

Zoo animal husbandry is a skill that should be developing constantly. In theory, this should lead to an improvement of zoo animal survivorship over time. Additionally, it has been suggested that species that are at a comparatively higher risk of extinction in their natural habitats (in situ) might also be more difficult to keep under zoo conditions (ex situ). Here, we assessed these questions for three zoo-managed rhinoceros species with different extinction risk status allocated by the IUCN: the "critically endangered" black rhino (Diceros bicornis), the "vulnerable" greater one-horned (GOH) rhino (Rhinoceros unicornis), and the "near threatened" white rhino (Ceratotherium simum). Comparing zoo animals ≥1 year of age, the black rhino had the lowest and the white rhino the highest survivorship, in congruence with their extinction risk status. Historically, the survivorship of both black and white rhino in zoos improved significantly over time, whereas that of GOH rhino stagnated. Juvenile mortality was generally low and decreased even further in black and white rhinos over time. Together with the development of population pyramids, this shows increasing competence of the global zoo community to sustain all three species. Compared to the continuously expanding zoo population of GOH and white rhinos, the zoo-managed black rhino population has stagnated in numbers in recent years. Zoos do not only contribute to conservation by propagating ex situ populations, but also by increasing species-specific husbandry skills. We recommend detailed research to understand specific factors responsible for the stagnation but also the general improvement of survivorship of zoo-managed rhinos.

SEASONAL FLUCTUATION OF FECAL CONSISTENCY IN ZOO-HOUSED MOOSE (ALCES ALCES).

Fecal consistency is routinely used as an indicator of gastrointestinal health and diet suitability in zoo animals. The underlying processes in ungulate fecal consistency are not often investigated. The authors assessed multiple factors on fecal consistency scores in six adult zoo moose (Alces alces) retrospectively for a 3-yr period. All animals were regularly tested for gastrointestinal parasites and received antiparasitics when noted. Neither the provision of dietary browse nor number of visitors had a significant effect. Seasonally varying intake of the staple pelleted diet was the main explanatory factor, with an additional effect from environmental temperature or season. Higher intakes and higher temperatures were linked to less well-formed feces. The authors hypothesize that during the summer hyperphagia to replete body reserves, the water re-absorption function of the large intestine apparently becomes overtaxed. Heat stress may have an additional, intake-independent effect. Although monitoring fecal consistency is a part of good health surveillance, physiologic seasonal effects, as reported here, need to be taken into consideration when interpreting the observations.

Diet, habitat and flight characteristics correlate with intestine length in birds.

A link between diet and avian intestinal anatomy is generally assumed. We collated the length of intestinal sections and body mass of 390 bird species and tested relationships with diet, climate and locomotion. There was a strong phylogenetic signal in all datasets. The total and small intestine scaled more-than-geometrically (95%CI of the scaling exponent > 0.33). The traditional dietary classification (faunivore, omnivore and herbivore) had no significant effect on total intestine (TI) length. Significant dietary proxies included %folivory, %frugi-nectarivory and categories (frugi-nectarivory, granivory, folivory, omnivory, insectivory and vertivory). Individual intestinal sections were affected by different dietary proxies. The best model indicates that higher consumption of fruit and nectar, drier habitats, and a high degree of flightedness are linked to shorter TI length. Notably, the length of the avian intestine depends on other biological factors as much as on diet. Given the weak dietary signal in our datasets, the diet intestinal length relationships lend themselves to narratives of flexibility (morphology is not destiny) rather than of distinct adaptations that facilitate using one character (intestine length) as proxy for another (diet). Birds have TIs of about 85% that of similar-sized mammals, corroborating systematic differences in intestinal macroanatomy between vertebrate clades.

Teeth out of proportion: Smaller horse and cattle breeds have comparatively larger teeth.

There are different descriptions of allometric relationships between important components of the mammalian skull. Craniofacial evolutionary allometry describes a pattern of increasing facial cranium in larger skulls. Another body of literature describes disproportionately larger teeth in smaller species or specimens, matching anecdotal observations with dental problems in dwarf breeds whose teeth appear "too large for their skulls." We test the scaling of tooth row length with body size and skull length in a data set comprising 114 domestic horses (representing 40 breeds) and in another data set of 316 domestic cattle (of >60 breeds). We demonstrate that smaller skulls have a relatively longer tooth row in both horses and cattle; larger specimens have relatively shorter tooth rows. Whereas in horses, larger skulls have a relatively longer diastema, the distance of the mesial maxillary premolar to the premaxilla was proportional to cranium length in cattle. While the reasons for these patterns remain to be detected, they support the hypothesis that tooth size might be less "evolvable," in terms of time required for changes, than body size. The pattern may affect (i) the selective breeding for dwarf breeds by setting minimum constraints for skull size, as described previously for domestic horses with the same data set; (ii) the susceptibility of small breeds for dental problems; and (iii) differences in chewing efficiency between breeds of different sizes. The findings support the existing concept that scaling of tooth to body size across taxa becomes more isometric the longer these taxa are separated in evolutionary time.

Macrowear effects of external quartz abrasives of different size and concentration in rabbits (Oryctolagus cuniculus).

External quartz abrasives are one of the driving forces of macrowear in herbivorous animals. We tested to what extent different sizes and concentrations influence their effect on tooth wear. We fed seven pelleted diets varying only in quartz concentration (0%, 4%, and 8%) and size (fine silt: ∼4 µm, coarse silt: ∼50 µm, fine sand: ∼130 µm) to rabbits (Oryctolagus cuniculus, n = 16) for 2 weeks each in a randomized serial experiment. Measurements to quantify wear and growth of incisors and the mandibular first cheek tooth, as well as heights of all other cheek teeth, were performed using calipers, endoscopic examination, and computed tomography scans before and after each feeding period. Tooth growth showed a compensatory correlation with wear. Absolute tooth height (ATH) and relative tooth height (RTH); relative to the 0% quartz "control" diet) was generally lower on the higher concentration and the larger size of abrasives. The effect was more pronounced on the maxillary teeth, on specific tooth positions and the right jaw side. When offered the choice between different sizes of abrasives, the rabbits favored the silt diets over the control and the fine sand diet; in a second choice experiment with different diets, they selected a pelleted diet with coarse-grained sand, however. This study confirms the dose- and size-dependent wear effects of external abrasives, and that hypselodont teeth show compensatory growth. The avoidance of wear did not seem a priority for animals with hypselodont teeth, since the rabbits did not avoid diets inducing a certain degree of wear.