Historical effects on local variation in walnut-feeding behavior by the Japanese wood mouse, Apodemus speciosus

The Japanese wood mouse Apodemus speciosus eats large, hard-walled walnuts of Juglans ailanthifolia immediately after finding or after hoarding them. However, not all individuals can efficiently eat the nuts. In this study, to examine local variation in the ability to eat walnuts, feeding behavior was compared among nine wood mouse populations, four from mainland Honshu, where the walnut tree is distributed, and five from the Izu Islands (30-100 km south of Honshu), which lack the tree species. To avoid the effects of pre-capture experience with walnuts, mice from areas lacking the walnut trees were used for testing, even in the Honshu sites. Most mice from Honshu were able to eat walnuts after a 14-day training period, whereas most insular mice could not, with the exception of mice on Kouzushima Island. An analysis of the population genetic structures of these mice based on sequences of the mitochondrial control (D-loop) region revealed that the four insular populations are genetically distinct from the mainland populations, whereas the Kouzushima population remains genetically similar to the mainland populations. The relatively recent colonization of Kouzushima may explain why mice from this island were able to feed on walnuts despite the lack of walnut trees on the island. Thus, walnut-feeding ability appears to have some innate basis in the Japanese wood mouse, and this trait would be selected for in a walnut-available environment as it would better enable mice to survive during food shortages.     

Population differences and learning effects in walnut feeding technique by the Japanese squirrel

Feeding efficiency is an important factor explaining diet selection in seed-eating animals. Behavioral adaptation to process and consume the seed is advantageous to improve their feeding rate. The Japanese walnut (Juglans ailanthifolia), which has a nutritious kernel surrounded by a hard shell, is an important food for the Japanese squirrel (Sciurus lis) in lowland mixed-species forests in Japan. In the present study, population differences in feeding technique were examined. In all squirrels captured from the Takao population (with wild Japanese walnuts in their habitat), the optimal feeding technique for quickly opening the walnut shell was observed. However, only 8% of the squirrels from the Fuji population (without walnuts in their habitat) processed the walnuts by the optimal feeding technique. By a repetitive provision of walnuts for 50 days, 14% of squirrels from the Fuji population acquired the optimal feeding technique. Learning effects did not differ significantly between solitary (without a demonstrator) and social conditions (with a demonstrator). The effects of age on learning the feeding technique were examined using individuals from a zoo population that had been naïve for Japanese walnuts. All five individuals aged 6 months old, and half the individuals aged 1–2 years old, ate walnuts by the optimal feeding technique after a 2-month learning period, but none of the individuals older than 3 years did. Consequently, the walnut feeding techniques of the Japanese squirrel are not entirely genetically fixed behavior and can be improved by learning at a young age.     

Incidence of Escherichia coli in Black Walnut Meats

Examination of commercially shelled black walnut meats showed inconsistent numbers of total aerobic bacteria, coliforms, and Escherichia coli; variation occurred among different meat sizes and within each meat size. The incidence of E. coli on meats of commercially hulled black walnuts depended on the physical condition of the nuts. Apparently tightly sealed ones contained only a few or none, whereas those with visibly separated sutures and spoiled meats yielded the most. This contamination was in part correlated to a hulling operation. Large numbers of E. coli on the husk of the walnuts contaminated the hulling water, subsequently also contaminating the meats by way of separated sutures. Chlorination of the hulling wash water was ineffective. Attempts were made to decontaminate the walnut meats without subsequent deleterious changes in flavor or texture. A treatment in coconut oil at 100 C followed by removal of excess surface oil by centrifugation was best.     

Walnut Consumption Increases Satiation but Has No Effect on Insulin Resistance or the Metabolic Profile Over a 4‐day Period

Obesity and diabetes have been associated with increased consumption of highly processed foods, and reduced consumption of whole grains and nuts. It has been proposed, mainly on the basis of observational studies, that nuts may provide superior satiation, may lead to reduced calorie consumption, and may decrease the risk of type 2 diabetes; but evidence from randomized, interventional studies is lacking. A total of 20 men and women with the metabolic syndrome participated in a randomized, double‐blind, crossover study of walnut consumption. Subjects had two 4‐day admissions to the clinical research center where they were fed an isocaloric diet. In addition, they consumed shakes for breakfast containing either walnuts or placebo (shakes were standardized for calories, carbohydrate, and fat content). Appetite, insulin resistance, and metabolic parameters were measured. We found an increased level of satiety (overall P value = 0.0079) and sense of fullness (P = 0.05) in prelunch questionnaires following the walnut breakfast as compared to the placebo breakfast, with the walnut effect achieving significance on day 3 and 4 (P = 0.02 and P = 0.03). We did not find any change in resting energy expenditure, hormones known to mediate satiety, or insulin resistance when comparing the walnut vs. placebo diet. Walnut consumption over 4 days increased satiety by day 3. Long‐term studies are needed to confirm the physiologic role of walnuts, the duration of time needed for these effects to occur, and to elucidate the underlying mechanisms.     

Coated nuts as an enrichment device to elicit tool use in tufted capuchins (Cebus apella)

The present paper describes a simple technique that hardens the shell of nuts and makes the use of a tool to crack them open more compelling. Walnuts were coated with a dough of sawdust and nontoxic white glue in different combinations; they were tested for hardness by using machines normally used to test different kinds of wood. Data on relative hardness for uncoated walnuts and walnuts coated with dough of two different combinations are presented. The coated walnuts were significantly harder to break than the uncoated ones, whereas no significant difference was found when comparing the hardness of two types of coated walnuts. Furthermore, observations on a captive group of tufted capuchins (Cebus apella) are described. The monkeys needed significantly more time to break open the coated walnuts. Early results show that coated walnuts may favor acquistion of tool use skills in a juvenile capuchin.     

Feeding ecology of rooks (Corvus frugilegus) on the Heretaunga Plains,Hawke's Bay,New Zealand

Abstract

The food resources of rooks were measured and related to changes in the birds' feeding behaviour during 1968-1970. In all, 381 birds were individually marked with wing tags and colour-coded leg bands and released; their activities were recorded for set periods. Rooks preferred to eat invertebrates and walnuts, but also took seed of cereals, pulses, and stock feeds. They spent at least 40% of their feeding time (75-80% in spring) eating invertebrates, 30-60% finding or hiding walnuts and sometimes acorns in autumn and winter, and 45% eating seed during hot, dry weather in summer. Up to 6% of feeding time was spent eating newly sown or ripening seed. Throughout winter rooks frequently searched for hidden nuts, which they ate or moved to new positions. The proportion of time they spent feeding and the kind of food taken varied seasonally and, in summer and winter, hourly; there were only slight differences between morning, mid-day, and afternoon periods. Rooks spent 65% of the time feeding in autumn, but only 25-30% in spring. Conversely, they spent longer at nest sites from late autumn, peaking at 60% in spring, then less time at nests as they spent longer searching for food. The size of flocks altered with the food being eaten and therefore with season and time of day. Small flocks were widespread where favoured food was plentiful in autumn, winter, and spring, but flocks were larger where food was unevenly abundant in summer and sometimes in winter. During early autumn each bird usually ate and stored walnuts in a selected area of 1-2 km3, returning there later in the season to recover stored nuts. The areas formed only part of the total range used by the study population, and were usually shared with other birds. Tests in aviaries showed that rooks preferred earthworms and walnuts to acorns, maize, and wheat (in that order), and field tests confirmed that rooks preferred walnuts to maize. The numbers of each food item required to sustain a rook for a daylight hour were estimated. Summer is the most difficult season for rooks to find food.     

Effective nut dispersal by magpies (<Emphasis Type="Italic">Pica pica</Emphasis> L.) in a Mediterranean agroecosystem

Scatter-hoarding animals such as corvids play a crucial role in the dispersal of nut-producing tree species. This interaction is well known for some corvids, but remains elusive for other species such as the magpie (Pica pica), an abundant corvid in agroecosystems and open landscapes of the Palearctic region. In addition, the establishment of the individual dispersed seeds—a prerequisite for determining seed-dispersal effectiveness—has never before been documented for the interaction between corvids and nut-producing trees. We analyzed walnut dispersal by magpies in an agroecosystem in southern Spain. We used several complementary approaches, including video recording nut removal from feeders, measuring dispersal distance using radio tracking (with radio transmitters placed inside nuts), and monitoring the fate of dispersed nuts to the time of seedling emergence. Magpies were shown to be highly active nut dispersers. The dispersal distance averaged 39.6 ± 4.5 m and ranged from 4.1 to 158.5 m. Some 90% of the removed walnuts were cached later, and most of these (98%) were buried in the soil or hidden under plant material. By the time of seedling emergence, ca. 33% of nuts remained at the caching location. Finally, 12% of the cached nuts germinated and 4% yielded an emerged seedling, facilitating the transition to the next regeneration stage. The results demonstrate for the first time that magpies can be an effective scatter-hoarding disperser of a nut-producing tree species, suggesting that this bird species may play a key role in the regeneration and expansion of broadleaf forests in Eurasia.     

Role of host fruit color in the behavior of the walnut fly Rhagoletis juglandis

Observations and sticky-trap tests were used to assess the effect of fruit color on the behavior of adult male and female Rhagoletis juglandis Cresson (Diptera: Tephritidae), a tephritid that infests husks of Arizona walnut in southeastern Arizona. In the first experiment, during which flies were observed foraging among walnut models suspended from small walnut trees, models were painted green to appear ripe and uninfested or yellow with brown patches to appear ripe and infested. Flies used for this first experiment were also of two types: prior to observations, one group of flies had access to real walnuts for 1.5 days (prior experience) while the other group of flies was held without real walnut fruits (no prior experience). Regardless of prior experience with real walnut fruits, female flies landed on green models more than yellow/brown models. Experienced males also were more likely to land on green models than on yellow/brown models. More interactions also occurred on green models, because there were more landings.In the field behavioral assay, flies from a natural population given a choice of green, yellow, and yellow/brown models landed most often on green models, and all interactions and oviposition attempts occurred on green models. Flies also distinguished models by color in field sticky trap assays.These results suggest that female response to ripeness cues is innate, while males develop a preference for green based on their encounter rate with females.     

Geographic variation in nut size of<Emphasis Type="Italic"> Castanopsis</Emphasis> species in Japan

The environmental conditions affecting the geographic variation in nut size of the Castanopsis species in Japan were examined. The mother trees having a relatively large nut tended to dominate in the geographically marginal area of the distribution range in Japan, i.e. the Japan Sea, Kanto, and Ryukyu areas, whereas in the central area, i.e. the Pacific area, mother trees having a relatively small nut occurred. We attempted an adaptive explanation, proposing that mother trees with large nuts have higher inclusive fitness than those with small nuts in unfavorable habitats. The Japan Sea area features large amounts of snow fall and low temperatures. The Ryukyu area is insular and is characterized by high temperatures. Within the Pacific area, mother trees with relatively large nuts were abundant at both higher and lower parts of the altitudinal range. We infer that environmental conditions in marginal areas, in terms of both horizontal and vertical distribution, favor mother trees having large nuts. However, in the marginal Kanto area, the cause of the absence of mother trees having relatively small nuts was independent of the environmental conditions. The geographic variation in nut size of Castanopsis species is not always explained by environmental factors.     

Impact of site‐induced mouse caching and transport behaviour on regeneration in Castanea crenata

Abstract. Dispersal and retrieval site selection by mice, transport distance, cache depth, and emergence and survival of seedlings of Castanea crenata (Japanese chestnut) were investigated by a magnet‐locating experiment in two habitat conditions (gap vs. forest understorey). Magnets were inserted into nuts (n= 450) and the nuts placed in the edge of forest gaps. Although wood mice (Apodemus speciosus and A. argenteus) initially buried nuts singly in shallow surface caches near the nut source, by the following spring these cached nuts were retrieved and re‐cached in larger, deeper caches farther from the source, particularly in forest understories, probably to reduce the threat of pilferage. All the nuts cached in the forest understories were consumed, but 4 seedlings emerged in gaps, apparently because of lower foraging activity in the gaps by the mice. Seed size was not correlated with cache depth or cache site selection. With increasing seed size, transport distance increased, particularly in gaps, possibly due to a greater potential energy gain (relative to handling cost to the cacher), or to attempts to prevent density‐ or mass‐dependent loss of caches by other foragers. Variable seed dispersal behaviour based on variation in seed size may influence the chances of colonization and distribution of the light‐demanding Castanea trees in mosaic landscapes and may play an important role in community organization and dynamics.     

DNA markers identify hybrids between butternut (Juglans cinerea L.) and Japanese walnut (Juglans ailantifolia Carr.)

Butternut (Juglans cinerea L.) is a temperate deciduous hardwood native to the eastern USA and southern Canada valued for its nuts and wood. Butternut’s survival is threatened by butternut canker, a disease caused by the exotic fungus Sirococcus clavigignenti-juglandacearum Nair, Kostichka & Kuntz. Field observations indicate that trees commonly called buartnut (a hybrid of butternut and its close congener Japanese walnut (Juglans ailantifolia × J. cinerea)) may be more resistant to butternut canker than is either parental species. Hybrids are difficult to distinguish morphologically from butternuts, and scientists have expressed concern over the possibility of range-wide genetic invasion by Japanese walnut via hybridization with butternut. We used pair-wise combinations of 40 random primers to screen bulked DNA pools of butternut, Japanese walnut, and buartnuts to identify genomic regions unique to Japanese walnut. We ultimately identified one ITS region marker, one chloroplast marker, one mitochondrial marker, and six nuclear markers. The utility of the markers for identifying hybrids was tested and verified using more than 190 genotypes. The markers will be used to identify buartnut hybrids based on the presence of introgressed genomic fragments inherited from Japanese walnut. We confirmed that hybrids have a complex genetic history and present features of the parental species in all possible combinations. These results will assist in the identification and testing of (non-hybrid) butternut for breeding and reintroduction of the species to its former habitats.     

Predator exaptations and defensive adaptations in evolutionary balance: No defence is perfect

Summary The lubber grasshopper,Romalea guttata, is large, aposematic, and extremely toxic. In feeding trials with 21 bird and lizard species, none were able to consume this chemically defended prey. Predators that attempted to eat lubbers, often gagged, regurgitated, and sometimes died. Loggerhead shrikes,Lanius ludovicianus, regularly impale this toxic prey in peninsular Florida. They, like other bird species, are unable to consume fresh lubbers. However, our tests show that they are able to consume lubbers if the prey are allowed to age for 1–2 days. This suggests that lubber toxins degrade following death and that shrike impaling behaviour serves as a preadaptation for overcoming the toxic defences of this large and abundant prey. These results also imply that counter adaptations against chemical defences need not involve major morphological or metabolic specializations, but that simple behavioural traits can enable a predator to utilize toxic prey.     

Vice-anvil use in nut processing by two Corvus species

Woodpeckers and certain passerine species secure encased food in the environment in various ways to facilitate the extraction of the contents with their bills. They do this by securing the food items in locations such as crevices and holes, newly defined in this paper as ‘vice-anvils’. Here I report that free-living New Caledonian crows (Corvus moneduloides) and rooks (Corvus frugilegus, in New Zealand) also use vice-anvils to process candlenuts and walnuts, respectively. New Caledonian crows placed candlenut sections in vice-anvils to aid kernel extraction, after the candlenuts had been dropped onto an anvil to break them open. In contrast, rooks used vice-anvils to secure walnuts while they broke the shell with their bills. Long-term use by rooks of a vice-anvil in a tree had produced a ‘purpose-made’ nut-cracking site. My findings extend the persistent use of specific vice-anvils to Corvus species and further demonstrate their innovative and flexible foraging behaviour.     

Aspergillus on tree nuts: incidence and associations

California exports tree nuts to countries where they face stringent standards for aflatoxin contamination. Trade concerns have stimulated efforts to eliminate aflatoxins and Aspergillus flavus from almonds, pistachios and walnuts. Incidence of fungi on tree nuts and associations among fungi on tree nuts were studied. Eleven hundred pistachios, almonds, walnuts and brazil nuts without visible insect damage were plated on salt agar and observed for growth of fungi. Samples came both from California nut orchards and from supermarkets. To distinguish internal fungal colonization of nuts from superficial colonization, half the nuts were surface-sterilized before plating. The most common genera found were Aspergillus , Rhizopus and Penicillium . Each species of nut had a distinct mycoflora. Populations of most fungi were reduced by surface sterilization in all except brazil nuts, suggesting that they were present as superficial inoculum on (rather than in) the nuts. In general, strongly positive associations were observed among species of Aspergillus ; nuts infected by one species were likely to be colonized by other species as well. Presence of Penicillium was negatively associated with A. niger and Rhizopus in some cases. Results suggest that harvest or postharvest handling has a major influence on nut mycoflora, and that nuts with fungi are usually colonized by several fungi rather than by single species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Genetic variation in walnuts (Juglans regia and J. sigillata; Juglandaceae): Species distinctions, human impacts, and the conservation of agrobiodiversity in Yunnan, China

Walnuts are a major crop of many countries and mostly cultivated in large-scale plantations with few cultivars. Landraces provide important genetic reservoirs; thus, understanding factors influencing the geographic distribution of genetic variation in crop resources is a fundamental goal of agrobiodiversity conservation. Here, we investigated the role of human settlements and kinship on genetic variation and population structure of two walnut species: Juglans regia, an introduced species widely cultivated for its nuts, and J. sigillata, a native species cultivated locally in Yunnan. The objectives of this study were to characterize sympatric populations of J. regia and J. sigillata using 14 molecular markers and evaluate the role of Tibetan villages and kin groups (related households) on genotypic variation and population structure of J. regia and J. sigillata. Our results based on 220 walnut trees from six Tibetan villages show that although J. regia and J. sigillata are morphologically distinct, the two species are indistinguishable based on microsatellite data. Despite the lack of interspecific differences, AMOVAs partitioned among villages (5.41%, P = 0.0068) and kin groups within villages (3.34%, P = 0.0068) showed significant genetic variation. These findings suggest that village environments and familial relationships are factors contributing to the geographic structure of genetic variation in Tibetan walnuts.     

Geographic variation in walnut seed size correlates with hoarding behaviour of two rodent species

Seeds of the Japanese walnut, Juglans ailanthifolia, are usually scatter-hoarded by two rodent species, the Japanese squirrel Sciurus lis and the field mouse Apodemus speciosus, but only by the latter in several areas where S. lis is absent. We examined seed-size-mediated interactions of these three species across a wide geographic range. Field tracking of walnuts with miniature radio-transmitters revealed that squirrels hoarded larger seeds more frequently and at greater distances than smaller seeds. In contrast, mice hoarded smaller seeds more frequently and transported them farther than larger seeds. These seed dispersers could affect the evolution of seed size because seeds hoarded at sites farther from source trees are known to survive better until germination and as seedlings. We expect that larger seeds may be advantageous in regeneration if the main seed dispersers are squirrels, whereas smaller seeds may be advantageous if mice are the dominant dispersers. These predictions were supported by the fact that seed size was smaller on islands inhabited only by mice and at the edge of the squirrel distribution, compared to areas where mice and squirrels are both common.     

Genetic diversity of superior Persian walnut genotypes in Azadshahr,Iran

Persian walnut (Juglans regia L.) is known to have originated in central and eastern Asia. Remnants of these wild populations can still be found in the Hyrcanian forest in north-eastern Iran. In this study, 102 individual walnut trees from four geographic populations in the Azadshahr province (Vamenan, Kashidar, Rudbar and Saidabad) were sampled. We characterized individual trees using 28 standard morphological traits. The range of traits varied widely for some economically important characteristics including nut weight (6.1–19.79 g), kernel weight (2.9–9.4 g), and kernel fill percentage (26.51–60.34%). After morphological evaluation, 39 superior individuals based on nut quality and kernel fill percentage were selected for further genetic analysis. Individual superior trees were genotyped using 10 simple sequence repeat markers (SSR) and genetic diversity. Number of alleles per locus ranged from 3 (WGA005) to 12 (WGA054). Clustering analysis of 10 SSR loci divided the genotypes into three main groups. PCoA analysis clearly sorted genotypes into one of four distinctive groups which aligned with the cluster analysis. All analyses showed that individuals from Saidabad were genetically distinct. Likewise, results indicated that the high level of genetic diversity in Azadshahr region walnuts may provide a diverse source for superior walnuts in walnut breeding programs.     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.     

Interannual Variation in Nut Abundance Is Related to Agonistic Interactions of Foraging Female Japanese Macaques (Macaca fuscata)

The importance of dominance status to foraging and ultimately survival or reproductive success in wild primates is known; however, few studies have addressed these variables simultaneously. We investigated foraging and social behavior among 17 adult female Japanese macaques (Macaca fuscata) on Kinkazan Island, northern Japan, from September to November in 2 consecutive years (2004 and 2005) to determine whether interannual variation in food availability was related to variation in agonistic interactions over food resources and the feeding behavior of individuals of different dominance rank. We compared energy obtained with daily energy requirements and also examined the effect of variation in feeding behavior on female survival and reproductive success. Fruiting conditions differed considerably between the 2 yr: of four nut-producing species, the nuts of only Torreya nucifera fruited in 2004, whereas all four species, particularly Fagus crenata, produced nuts in abundance in 2005. The abundance and average crown size of trees of Torreya nucifera were smaller than those of Fagus crenata, and there was a higher frequency of agonistic interactions during 2004, when dominant, but not subordinate, individuals were able to satisfy daily energy requirements from nut feeding alone through longer nut feeding bouts. In contrast, all macaques, regardless of their dominance rank, were able to satisfy their energy requirements by feeding on nuts in 2005. Subordinate macaques appeared to counter their disadvantage in 2004 by moving and searching for food more and maintaining larger interindividual distances. Several lower-ranking females died during the food-scarce season of 2004, and only one dominant female gave birth the following birth season. In contrast, none of the adult females died during the food-scarce season of 2005, and 12 females gave birth the following birth season. These findings suggest that an interaction between dominance rank and interannual variation in food availability are related to macaque behavior, survival, and reproduction.