Large-sized Prey Research Articles

Elementary trophic ecology of pacific salmons in the Bering Sea. Species and regional differences. Provision with food in different environments

The final issue in the series of studies on trophic ecology of pacific salmons in the Bering Sea conducted by planktonologists of Pacific Fish. Res. Center (TINRO) for NPAFC. Salmons dwell in the upper epipelagic layer and feed mainly in the daytime by large-sized zooplankton that is more abundant in the upper pelagic layer as compared with the lower pelagic layer in summer-fall season. The zooplankton and nekton species dominant in the salmon diet are determined on the base of 8,000 plankton samples and 45,000 samples of salmon stomachs. Two periods with different environments are revealed in the Bering Sea in the early 21 century: the relatively warm period of 2000-2006 and the relatively cold period of 2007-2012. The regime shift between them affected the plankton community and trophic ecology of salmon, though not by means of water cooling directly but presumably because of the water circulation change. Food composition is determined for each size group of salmons in both periods, by biostatistical areas and bathymetric ranges. Zooplankton grazing by salmons increased significantly in all areas during the cold period, whereas the nekton grazing decreased, but their total consumption did not change much. Transition to the next warm period since 2013 does not affect negatively to the salmon food base; their diet changes significantly but within the list of preferable species. Feeding intensity of pink, chum, and sockeye salmons was similar in any period, as well as in the 2014-2015, food deficit was never observed. The salmons show a great plasticity in feeding: in general, their diet corresponds to the structure of plankton community, though the portion of nekton increases with the age. Large-sized salmons consume a large-sized prey and need more time for its complete digestion, so there is necessary to apply reduction coefficients in calculation of their daily food rations. N​utritional needs of all salmon stocks are estimated for summer-fall season and offered as a guideline for evaluation their load on the food base and their food security for certain spatial and temporal circumstances.

Concealed by darkness: interactions between predatory bats and nocturnally migrating songbirds illuminated by DNA sequencing.

Recently, several species of aerial-hawking bats have been found to prey on migrating songbirds, but details on this behaviour and its relevance for bird migration are still unclear. We sequenced avian DNA in feather-containing scats of the bird-feeding bat Nyctalus lasiopterus from Spain collected during bird migration seasons. We found very high prey diversity, with 31 bird species from eight families of Passeriformes, almost all of which were nocturnally flying sub-Saharan migrants. Moreover, species using tree hollows or nest boxes in the study area during migration periods were not present in the bats' diet, indicating that birds are solely captured on the wing during night-time passage. Additional to a generalist feeding strategy, we found that bats selected medium-sized bird species, thereby assumingly optimizing their energetic cost-benefit balance and injury risk. Surprisingly, bats preyed upon birds half their own body mass. This shows that the 5% prey to predator body mass ratio traditionally assumed for aerial hunting bats does not apply to this hunting strategy or even underestimates these animals' behavioural and mechanical abilities. Considering the bats' generalist feeding strategy and their large prey size range, we suggest that nocturnalbat predation may have influenced the evolution of bird migration strategies and behaviour.

Spatial and temporal distribution of gerrid (Heteroptera) and predation on microcrustaceans from a tropical shallow lake.

Predator-prey interactions involving an aquatic insect and zooplanktonic prey of different sizes were investigated to quantify prey mortality exposed to predators. Laboratory experiments were undertaken with the young and adult gerrid Rheumatobates crassifemur to test predation and size selectivity on the cladocerans Daphnia gessneri, Ceriodaphnia richardi, and Bosmina tubicen. Population fluctuations and spatial distribution of the gerrid were also evaluated in a small and shallow Brazilian lake throughout 12 months in fortnightly samples. The insects were more abundant in the littoral (mean density 7.0 ± 1.2 ind.m-2) compared to the limnetic zone. The period with the highest densities was late January to June, in both zones. Predation by young instars on Daphnia and Ceriodaphnia was significant (mean ingestion rate of 1.3 ± 0.1 D. gessneri and 0.7 ± 0.1 C. richardi per predator per hour). Adult insect fed only the large-sized prey (mean ingestion rate of 1.0 ± 0.1 D. gessneri per predator per hour). Young gerrids have greater potential to prey on cladocerans than adults, and size selectivity occurred for both predators. Preference of adults by the larger prey is probably related to difficulties in manipulating smaller planktonic prey, such as Ceriodaphnia. Due to higher densities of insects in the littoral, higher predation on zooplankton in this zone is expected. This study does contribute to a better understanding of trophic interactions in tropical shallow lakes and is the first to investigate predation of a gerrid on cladocerans in laboratory experiments.

The Importance of Estuarine Production of Large Prey for the Growth of Juvenile Temperate Seabass (Lateolabrax japonicus)

The feeding ecology of juvenile temperate seabass (Lateolabrax japonicus) in relation to various prey environments was analyzed in the Yura River estuary from 2008 to 2012. Juveniles mainly fed on copepods and mysids, both in the Yura River estuary and in the adjacent coastal area. The dependency on the mysids increased as juveniles grew from 15 to 25 mm standard length (SL), and the main stomach contents were mysids for juveniles larger than 25 mm SL. The maximum size of food contents increased from 2 to 15 mm with the growth of juveniles from 15 to 25 mm SL. Most prey items smaller and larger than 2 mm were composed of copepods and mysids, respectively. The mean size of ingested food items increased with the size of the juveniles, while the mean number of prey items in each stomach decreased with growth after juveniles reached 22 mm SL. As a result, the stomach content index increased with the size of juveniles. These results indicate that the shift to larger-sized prey items (from copepods to mysids) is important for juveniles to obtain energy efficiently. Mysids were larger and more abundant in the estuaries than those in the coastal area. Therefore, the estuary provided better foraging conditions to juvenile temperate seabass than the coastal area.

Variation in parental care in the spectacled tyrant Hymenops perspicillatus is associated with increased nest predation in grassland fragments

Predation risk on birds is often an important source of natural selection that shapes parental care and may promote behavioral changes. Parents can often estimate certain risks and adjust their behavior to reduce the likelihood of nest predation. The fragmentation of habitats is one of the main consequences of loss of habitats, and in general, for birds breeding in smaller patches, their daily nest-survival rate is lower due to increased nest predation. Since nest survival is an estimate of predation risk in the environment, we evaluated the daily survival rate (DSR) for nests of spectacled tyrants (Hymenops perspicillatus) and parental care behavior on fragmented and unfragmented grasslands. We conducted nest searching and monitoring during the 2012–2013 breeding season in small patches and in a continuous patch of grassland. In addition, parental activity was recorded using video monitoring. We found a lower DSR for the spectacled tyrant in fragmented grasslands, associated with increased nest predation risk; females showed a variation in parental care. This variation was evidenced by larger incubation bouts and lower visitation rate during the incubation period, and by a lower food delivery rate to nestlings, compensated by larger prey sizes. The results show that fragmentation not only reduces the fitness of individuals and impacts adversely on population, but individuals are also subjected to a strong selection pressure, and their reproductive success may depend to some extent on the ability of parents to estimate at least certain predation risk and adjust their behavior in this regard.

Prey-Specific Growth Responses of Freshwater Flagellate Communities Induced by Morphologically Distinct Bacteria from the Genus Limnohabitans.

Because their large growth potential is counterbalanced with grazing by heterotrophic nanoflagellates (HNF), bacteria of the genus Limnohabitans, which are common in many freshwater habitats, represent a valuable model for examining bacterial carbon flow to the grazer food chain. We conducted experiments with natural HNF communities taken from two distinct habitats, the meso-eutrophic Římov Reservoir and the oligo-mesotrophic Lake Cep (South Bohemia). HNF communities from each habitat at distinct seasonal phases, a late April algal bloom and a late May clear water phase, were each fed 3 Limnohabitans strains of differing cell sizes. Water samples were prefiltered (5 μm) to release natural HNF communities from zooplankton control and then amended with the Limnohabitans strains L. planktonicus II-D5 (medium sized, rod shaped), Limnohabitans sp. strain T6-5 (thin, long, curved rod), and Limnohabitans sp. strain 2KL-3 (large solenoid). Using temporal sampling and prey treatment, we determined HNF growth parameters such as doubling time, growth efficiency, and length of lag phase prior starting to exponential growth. All three Limnohabitans strains supported HNF growth but in significant prey-, site-, and season-dependent fashions. For instance, addition of the moderately large T6-5 strain yielded very rapid HNF growth with a short lag phase. In contrast, the curved morphology and larger cell size of strain 2KL-3 made this prey somewhat protected against grazing by smaller HNF, resulting in slower HNF growth and longer lag phases. These trends were particularly pronounced during the late May clear-water phase, which was dominated by smaller HNF cells. This may indicate a longer "adaptation time" for the flagellate communities toward the large prey size offered.

A model of hunter-gatherer skeletal element transport: The effect of prey body size, carriers, and distance

Zooarchaeologists frequently use the relative abundance of skeletal elements in faunal assemblages in conjunction with foraging theory models to infer subsistence decisions made by prehistoric hunter-gatherers. However, foraging models applied to ethnoarchaeological cases have had variable success linking skeletal transport decisions with foraging predictions. Here, we approach this issue with the well-known Hadza data to statistically model the skeletal element transport decisions in response to distance from the residential hub and the number of carriers available for carcass transport. We compare our modeling approach to the traditional skeletal element utility curves from Binford's work with the Nunamiut, and to the more recently proposed Shannon evenness measure. Our approach, based on standard yet powerful statistical modeling techniques, can help researchers gain increased insight into the prey part transport responses of hunter-gatherers. Our analyses treat individual prey skeletal elements by body size as the response variable. The results of this analysis suggest that utility curves, and the Shannon evenness approach as a proxy for utility curves, are problematic for making statements about prehistoric foraging from zooarchaeological data. Transport distance does not explain a significant portion of small prey (size class 2) skeletal element transport variation. However, distance explains a great deal of transport variation in large prey (size classes 4 and 5). Inferences from skeletal element profiles should be made relative to prey body size and the discard probability of individual elements. Understanding the influence of these variables allows construction of a framework for testing archaeological element profiles against ethnographically derived transport models.

Remarks on Correlations and Implications of the Mandibular Structure and Diet in Some Seals (Mammalia, Phocidae)

Abstract Remarks on Correlations and Implications of the Mandibular Structure and Diet in Some Seals (Mammalia, Phocidae). Koretsky, I. A., Rahmat, S. J., Peters, N. — The diverse representatives of Recent seals within the three extant subfamilies (Cystophorinae, Phocinae, Monachinae) of the family Phocidae exhibit dietary variations among species, feeding on invertebrates and a diversity of prey depending on the season and availability. To explain this variability, an introductory morphological examination of the mandibular structure of Recent seals from each subfamily was performed, focusing on: heights of the mandibular ramus and condyloid process; measurements of the masseteric fossa; and attachments of muscles of mastication. After measuring the condyloid angles (the inclination of the condyloid process in relation to the axis of the alveolar row) among the examined species, a correlation between the size ofthe angle, function, and diet was recognized. Seals with a tall mandibular ramus and greater condyloid angle (Cystophorinae) feed on larger-sized prey, while seals with a shorter ramus and lesser condyloid angle feed on small (Monachinae) to medium-sized (Phocinae) prey, regardless of the overall size of the seal. This study focused on the mandibular morphology of some living and fossil representatives of Phocidae, providing an association between functional and ecological interpretations of modern seals in general and extrapolating this knowledge for fossil dietary preference.

Population structure and biology of shortfin mako, Isurus oxyrinchus, in the south-west Indian Ocean

The population structure, reproductive biology, age and growth, and diet of shortfin makos caught by pelagic longliners (2005–10) and bather protection nets (1978–2010) in the south-west Indian Ocean were investigated. The mean fork length (FL) of makos measured by observers on longliners targeting tuna, swordfish and sharks was similar, and decreased from east to west, with the smallest individuals occurring near the Agulhas Bank edge, June to November. Nearly all makos caught by longliners were immature, with equal sex ratio. Makos caught by bather protection nets were significantly larger, males were more frequent, and 93% of males and 55% of females were mature. Age was assessed from band counts of sectioned vertebrae, and a von Bertalanffy growth model fitted to sex-pooled length-at-age data predicted a birth size (L0) of 90 cm, maximum FL (L∞) of 285 cm and growth coefficient (k) of 0.113 y–1. Males matured at 190 cm FL, aged 7 years, and females at 250 cm, aged 15 years. Litter sizes ranged from nine to 14 pups, and the presence of gravid females in bather protection nets suggested that some pupping occurred in shelf waters. Teleosts (mainly Trachurus capensis) occurred in 84% of stomachs collected on longliners, whereas elasmobranchs (63.5%) were most common in samples collected from bather protection nets, followed by teleosts (43.1%) and cephalopods (36.5%). Larger prey size may be a factor that attracts large makos to coastal waters.

Skeletal element distributions of the large herbivores from the Lingjing site, Henan Province, China

More than ten thousands of bone fragments were recovered from the Lingjing site, Henan Province during 2005 and 2006. In this paper, through the quantification and statistical analyses of the skeletal elements of the two predominant species in this assemblage, aurochs (Bos primigenius) and horse (Equus caballus), the differential influences and weights of a variety of taphonomic agencies in the formation of the assemblage are assessed respectively. Compared to the natural agencies, hominid hunting and the subsequent disarticulation, slaughtering, and their transport of the bone elements of the prey species are the main factors accounting for the formation of the present assemblage. More importantly, this study initiatively identifies hominid’s differential treatment of the bones of aurochs and horse in the Paleolithic record of East Asia and demonstrably suggests that hominids at the site have already practiced sophisticated hunting techniques and subsistence strategies and may be quite familiar with the ecological and anatomical characteristics and nutritional values of the large-sized prey animals and can accordingly take different processing and handling strategies at the hunting site.

Food-niche Differences Between Two Syntopic Scops-Owls on Okinawa Island, Japan

Two similar-sized scops-owl species, the Elegant Scops-Owl (Otus elegans) and the Japanese Scops-Owl (O. semitorques) occur syntopically in a forest on Okinawa Island, Japan. We investigated diets of these owls to evaluate the breadth of their food niches and examine potential mechanisms that might differentiate their food niches. Elegant Scops-Owls fed predominantly on orthopterans, whereas Japanese Scops-Owls took prey from many taxonomic groups. Although we observed dietary overlap, food niches were differentiated by prey size and habitat. Elegant Scops-Owls mostly foraged on arboreal prey, whereas Japanese Scops-Owls took both arboreal and terrestrial prey. Elegant Scops-Owls specialized on small prey, whereas Japanese Scops-Owls preyed equally on small- to large-sized prey. Foraging habitat of these owls may reflect differences in prey locomotion, and differences in prey sizes may correspond to body-size differences between the two species. Food-niche differences may relax competition between these species.

Trophic Niche Width, Shift, and Specialization of Dascyllus aruanus in Toliara Lagoon, Madagascar

Abstract Intrapopulation diet specializations may result from the use of different dietary items or foraging tactics by individuals within a single population. The damselfish, Dascyllus aruanus, is a highly site-attached coral reef fish living in size hierarchies among branched corals. The trophic niche width and feeding specialization of this species were explored using stable isotopes and stomach content analyses. Intra-group niche variation was mainly related to fish size. Within social groups, D. aruanus gradually shifted its foraging tactics according to size; smaller fish fed on benthic prey such as isopods and copepods, and the larger fish foraged in the water column on planktonic copepods and larger-sized prey. Group density was found to explain some variation in trophic niche characteristics; greater specialization on prey size was observed in the colony having the highest density. All members of the largest colony foraged more frequently in the water column. Knowing that planktonic copepods are ...

Barn and Long-Eared Owl Diets: A Comparative Study from Central Pennsylvannia and a Key for Identification of Prey Items

Two sympatric owl species, Tyto alba (Barn Owl) and Asio otus (Longeared Owl), are found in Lewisburg, PA. These species are of similar size, have similar hunting styles, and most likely use similar prey items. In studying the diets of these two owl species, we were able to determine their prey selection, the types and relative abundance of prey in the area, and how these sympatric predators may be partitioning their resources. A total of 80 pellets (40 from Barn Owls and 40 from Long-eared Owls) were analyzed to understand their diets. Pellets were collected from two locations in Lewisburg, PA approximately 2 km apart. Cranial and postcranial remains within the owl pellets were used to identify species by comparing them against our reference collections and keys. Typically, the Barn Owl is present in Lewisburg from late spring through fall and the Long-eared Owl from late fall through spring. Barn Owl pellets contained a greater number of larger-sized prey than Long-eared Owl pellets. Both owl species rely heavily upon rodents as their primary food source, which make up over 90% of their diets. Temporal differences and differences in prey selectivity are most likely the major factors allowing these species to share their common resources.

Reproduction, grazing, and development of the large subarctic calanoid Eucalanus bungii: is the spring diatom bloom the key to controlling their recruitment?

The response of large calanoid, Eucalanus bungii, to environmental fluctuation, particularly in relation to the spring diatom bloom in the Oyashio region, western subarctic Pacific Ocean, was examined by investigating egg production, grazing, development and starvation tolerance. Mean in situ egg production rate increased with ambient chlorophyll-a concentration, ranging from 0 to 47 eggs female−1 d−1, while no diurnal synchronous spawning behavior was observed. Under the spring bloom condition, E. bungii showed prey preference for less mobile and larger-sized prey (≥30 μm ESD) and bloom-forming diatom Thalassiosira spp. accounted for >80% of ingested carbon. In the laboratory, E. bungii was successfully reared from newly hatched nauplii to adult with the diatom, Thalassiosira nordenskioldi, as a food resource. Nauplii newly hatched from eggs reached the adult stage in ca. 150 days (5°C) with a sigmoidal developmental pattern and no sexual difference in development pattern. Starvation experiments indicated that the starved copepodids (C1–C4) became more vulnerable to high temperature with the progression of developmental stage, suggesting that the post-bloom condition with low food availability and increased temperature is harsh for their copepodids. The results of this study in conjunction with previous findings suggest that E. bungii is well adapted to utilize large-sized phytoplankton, such as a bloom-forming diatoms and, therefore, their recruitment processes, including egg production, development and mortality would be strongly affected by the duration and intensity of the spring bloom.

Análise comparativa da dieta de quatro espécies de cucos (Aves: Cuculidae), no sul do Brasil

We studied stomachs contents of 50 specimens of four common cuckoo species (Cuculidae) from southern Brazil: Guira guira (Gmelin, 1788) (n = 21), Coccyzus melacoryphus (Vieillot, 1817) (n = 8), Crotophaga ani (Linnaeus, 1758) (n = 11), and Piaya cayana (Linnaeus, 1766) (n = 10). We measured stomach volume and prey were identified and quantified based in their dimensions. The size of the prey was associated with bill width and body mass. Diet items were analyzed and identified to the lowest possible taxonomic level, based on literature, scientific collection data, and specialists. The diet of the four species was composed by 100% animal species, with invertebrate dominance. Only in G. guira stomachs the presence of vertebrates was observed more than once (amphibian, reptile, and bird). C. ani, which had similar diet, ingested a large number of Arachnida and Orthoptera. Piaya cayana ate mostly Hemiptera. The predominance of caterpillars (Lepidoptera), suggested in the literature was not observed in the diet of C. melacoryphus. There was a statistically significant difference in bill width and body mass among the bird species, and there was no correlation between bill width and the proportion of medium and large-sized prey.

Experimental Observations of Early Mesolithic Points in North-East Italy

We present an analysis of Sauveterrian microlithic backed points from sites located in the Trentino region of the north-eastern Italian Alps: Pradestel, Lago delle Buse & Colbricon. Experimental arrows were produced, hafted and shot into an animal target. The results of this study indicate that these points may have been “ineffective” for the hunting of medium to large-sized prey, such as ibex, red deer, bear, or wild boar. We propose that Sauveterrian microliths can rather be correlated with the hunting of small forest prey such as roe deer, marmot and other animals with thick fur. An alternative hypothesis for the hunting of large game is also proposed.

Influence of prey body characteristics and performance on predator selection

At the time of settlement to the reef environment, coral reef fishes differ in a number of characteristics that may influence their survival during a predatory encounter. This study investigated the selective nature of predation by both a multi-species predator pool, and a single common predator (Pseudochromis fuscus), on the reef fish, Pomacentrus amboinensis. The study focused on the early post-settlement period of P. amboinensis, when mortality, and hence selection, is known to be highest. Correlations between nine different measures of body condition/performance were examined at the time of settlement, in order to elucidate the relationships between different traits. Single-predator (P. fuscus) choice trials were conducted in 57.4-l aquaria with respect to three different prey characteristics [standard length (SL), body weight and burst swimming speed], whilst multi-species trials were conducted on open patch reefs, manipulating prey body weight only. Relationships between the nine measures of condition/performance were generally poor, with the strongest correlations occurring between the morphological measures and within the performance measures. During aquaria trials, P. fuscus was found to be selective with respect to prey SL only, with larger individuals being selected significantly more often. Multi-species predator communities, however, were selective with respect to prey body weight, with heavier individuals being selected significantly more often than their lighter counterparts. Our results suggest that under controlled conditions, body length may be the most important prey characteristic influencing prey survival during predatory encounters with P. fuscus. In such cases, larger prey size may actually be a distinct disadvantage to survival. However, these relationships appear to be more complex under natural conditions, where the expression of prey characteristics, the selectivity fields of a number of different predators, their relative abundance, and the action of external environmental characteristics, may all influence which individuals survive.

Evolution of prey holding behaviour and large male body size in Ninox owls (Strigidae)

The hawk owl genus Ninox is unique among raptorial birds in that it includes three species in which males are substantially larger than females. This is a reversal of the normal pattern observed in both diurnal and nocturnal raptorial birds in which females are larger. Interestingly, these three Ninox species also are both the largest of the 22 species in the genus and the only species that exhibit the striking behaviour of ‘prey holding’ in which large (> 600 g) mammalian or avian prey is captured at night and held with body parts intact, and draped below a roost for the entire day without being consumed. Because explanations of the evolution of large male size suggest that it results from competition among males, the adaptive significance of prey holding was studied in a wild population of powerful owl Ninox strenua. Prey holding is largely confined to breeding males and its occurrence varies significantly across the breeding cycle, being most frequent during incubation and brooding. The study did not clearly resolve whether prey holding is a form of food storage or territorial display; however, both functions can select for large male body size and therefore play a significant role in the evolution of nonreversed size dimorphism. Although female-only incubation and brooding is typical of Ninox owls and other owl species, prey holding appears to occur only in the large Ninox species because of the unique combination of large body size, large prey size, separate sex roles, and obligate cavity nesting.

Does a Gigantic Insular Snake Grow Faster or Live Longer to be Gigantic? Evidence from a Long-term Field Study

Insular snake populations show bimodal size distributions with giants evolving on islands with large prey size and dwarfs on islands with small prey size. However, there is virtually no published study that examined growth trajectories of wild individual snakes to test whether giant and dwarf populations of a species differ in their growth rates and life spans. In order to address this issue, we compared the growth history of individually marked snakes in three insular and one mainland populations of Elaphe quadrivirgata. Our long-term field study, which started in 1982, suggests that gigantic snakes on Tadanae-jima, the Izu islands, take twice as long (30 to 40 years) to attain maximum snout-vent length as snakes from two non-gigantic insular populations. Our study also clearly demonstrated that snake gigantism is achieved by continuous growth throughout ontogeny rather than extensive growth early in their life. Contrary to expectations, young snakes from the gigantic populations grew more slowly than those of the other populations. We discuss potential proximate mechanisms that cause these different growth trajectories, particularly focusing on the exploitation of large sized avian eggs by older aged snakes, which would be a key factor enabling gigantism of the snake.

Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation

Faunal remains from spoil heaps of two burrows inhabited by small carnivores (fox and badger) are analysed from a taphonomical point of view. This analysis provides characteristics for bone accumulation produced by small carnivores and will be a powerful tool for deciphering site formation about the occupational alternation of small carnivores and humans. Identified species were grouped by size classes. Faunal spectrum is composed by varied species of microfauna (70%), mesofauna (30%). Macromammal remains are under represented (less than 1%) and come from scavenged carcasses. Predators and consumed species are compared on the basis of the skeletal part representation, age classes and recording of predation marks (gnawed and digested bones). Skeletal part representation shows that all taxons exhibit a low-representation of axial skeleton and autopodial bones. Predators show a high representation of hind limb bones and a low representation of the fore limb bones whereas consumed species exhibit a reverse pattern. Mortality curve analysis provides an attritional profile for carnivores and helps for the establishment of the season of occupation of the burrow. Adults largely dominate consumed species. Moreover, predation marks are found in 1% of the carnivores’ bones and from 15% to more than 40% on prey bones. The large-sized prey bones only wear gnawed marks (20%) and anthropic marks (10–20%) whereas microfaunal remains exhibits more digested marks (40%) than gnawing stigmata (5%). Medium-sized animals bones wear both marks and with the same proportions (10–15%).