Effects of body size,seed density,and soil characteristics on rates of seed harvest by heteromyid rodents

Summary Although it is well established that coexisting heteromyid rodent species forage in different microhabitats, we do not yet know the basis for divergent microhabitat choice. One possibility is that seed harvest rates differ among microhabitats, and each species forages where it can extract seeds most efficiently. Microhabitats vary in several factors that could affect heteromyid foraging efficiency, including seed density, soil organic content and particle size distribution. We have explored the effect of each of these variables on harvest rates of several species feeding from petri dishes containing known densities of millet seeds embedded in soil of known particle size and density. Results indicate that the number of seeds harvested per second increases uniformly with seed density and soil density and decreases with soil particle size. Body size affects these relationships: larger animals have higher harvest rates for a given set of conditions and experience a greater relative change in harvest rate for a give change in conditions. This implies that heteromyids can be expected to exhibit species-specific microhabitat preferences while foraging in nature.     

Cheek pouch capacity in heteromyid rodents

Summary Diplacus aurantiacus produces a full canopy of leaves during the rainy winter and spring. As the drought begins in summer, all but the terminal leaves are lost. The leaves present during the growth period have a comparatively low specific weight and a high content of water, protein, and non-structural carbohydrate on a weight basis. Leaves of this type have a high carbon-gain per unit dry matter investment.The larvae of Euphydryas chalcedona utilize Diplacus as their principal food source. Following the first winter rains, the shrub starts to grow and the larvae of Euphydryas break diapause and begin actively feeding. Adults are produced which lay eggs that hatch into prediapause larvae. During the end of the growth period of the shrub, as the quality and quantity of Diplacus leaves decline, the prediapause larvae have a brief period of active feeding and growth and then enter diapause. Diplacus produces a leaf surface resin which inhibits the growth of Euphydryas larvae. It is present in the highest amounts on those few leaves that remain on the shrub during the drought period.The type and pattern of herbivore defense in Diplacus fits the model described for apparent plants.     

Mechanisms of seed harvest by heteromyid rodents: soil texture effects on harvest rate and seed size selection

Summary Several lines of evidence show that soil texture plays an important role in the distribution of desert-dwelling heteromyid rodents. This is not surprising, since texture influences the energetic cost of digging burrows and of scratching at the soil surface to harvest buried seeds. Texture also may influence the efficiency with which seeds can be separated from the soil particles with which they are mixed. To explore mechanisms of particle separation by foraging heteromyids we measured seed harvest rates and size selection in the laboratory for a variety of seed sizes and soil textures. Harvest rate declined with increasing soil coarseness, and the preference for seeds of intermediate size that was apparent in fine soil disappeared when seeds were mixed with soil slightly coarser than the preferred seed size. In addition, there was evidence that particle separation efficiency is sensitive to the relative sizes of seeds and soil. A discontinity in the function relating harvest rate to soil texture occurred at finer soil textures for small seeds than for large seeds, suggesting that harvest techniques change once soil particle diameter equals or exceeds that of seeds. These results are consistent with the hypothesis that heteromyids use a combination of gravity-and rake-sorting mechanisms for particle separation.     

Microhabitat selection in two species of heteromyid rodents

Summary An experiment was conducted to determine the microhabitat preferences of two heteromyid rodents, Dipodomys ordi and Perognathus flavus. This experiment used marked seeds and the atomic absorption spectrophotometer in order to study the environment as a mosiac of microhabitats. The results of our analysis indicate that these two heteromyids are microhabitat selectors. The preferences of the rodents are D. ordi: grass habitat 0.0%, near grass habitat 22.5%, open habitat 77.4% and P. flavus: grass habitat 46.2%, near grass habitat 32.2%, open habitat 21.4%. The overlap between the two species is only 0.43.     

Cheek pouch capacities and loading rates of heteromyid rodents

Rodents of the family Heteromyidae are proficient gatherers and hoarders of seeds. A major component of their adaptive specialization for harvesting and transporting seeds is their spacious, fur-lined cheek pouches. Precise measurements of cheek pouch capacities are essential if ecologists are to understand the foraging ecology, possible constraints on locomotion patterns, and competitive relationships of heteromyid rodents. To measure the size of these cheek pouches and the rate at which animals load seeds into their pouches during seed harvest, we attracted 56 individuals representing ten species of heteromyid rodents to bait stations in the field and allowed them to fill their cheek pouches with seeds several times while we observed and timed the events with the aid of night-vision equipment. The largest load taken by each individual was used as an estimate of its cheek pouch capacity. At the end of observations, each subject was captured and its mass and other data gathered. The allometric relationship between cheek pouch capacity and body mass for ten species of heteromyids was significant [pouch capacity (ml) = 0.148 body mass (g)^0.992, r ²=0.91, P<0.0001]. The regression coefficient is ≈1.0, which indicates that the volume of the cheek pouches scales in direct proportion to body size. When the data were subdivided into quadrupeds (Perognathus and Chaetodipus) and bipeds (Dipodomys) (n=5 for each), the relationships between pouch capacity and body mass were significant, but the two regressions were not significantly different from each other. When all loads (full and partial) were considered, subjects filled their cheek pouches an average of 93 ± 10% of pouch capacity (n=185). Cheek pouch capacities from published studies of artificially filled pouches of heteromyids in the laboratory averaged about 40% below the field measurements obtained here. The allometric relationship between mean loading rate and body mass was also significant [seeds/s=1.067 bodymass (g)^0.830, r ²=0.85,P=0.0011), but when quadrupeds and bipeds were considered separately, the relationships were not significant. Seed densities and bulk densities were used to calculate packing coefficients for seed species, which, when used in conjunction with the allometric relationship between cheek pouch capacity and body size, can be used to estimate the maximum load carried by a heteromyid. Except for the very largest kangaroo rat species, a full pouch load of Indian ricegrass seeds represents less than the daily energy requirements of an active heteromyid. Received: 3 March 1997 / Accepted: 15 July 1997     

A functional-morphometric analysis of forelimbs in bipedal and quadrupedal heteromyid rodents

The rodent family Heteromyidae contains bipedal hoppers and quadrupedal runners. The possibility that bipedalism is associated with forelimb specialization for nonlocomotory functions, such as burrowing and seed-gathering, motivated a static functional-morphometric and interspecific allometric analysis of 18 metric characters of the forelimb skeleton. A principal-components analysis, across 28 species in six genera, showed that lengths of proximal (scapula, humerus) and distal (ulna, radius, metacarpal) elements were negatively allometric, and widths were positively allometric. Quadrupedal and bipedal species groups showed qualitatively similar allometric patterns, except that scapula width anterior to the spine was positively allometric in quadrupeds and negatively allometric in bipeds; scapula width posterior to the spine was positively allometric in bipeds and isometric in quadrupeds; and olecranon length was isometric in bipeds and positively allometric in quadrupeds. Most morphometric characters varied significantly among species within genera, even when effects of size variation were reduced by reconstructing all species to a common general size (as indicated by their score on the first principal component). These shape differences caused species to vary in the mechanical advantage of the forelimb, of possible importance for digging and seed-harvesting performance. Relative to quadrupeds, bipedal species tended to have greater mechanical advantage for proximal forelimb elements and smaller mechanical advantage for distal forelimb elements, but only the distal pattern remained in reconstructed forms, and no functional character was significantly different when tested over variation among genera nested within locomotion type. Cluster analysis confirmed that forelimb characters related to digging or seed-harvest are not coincident with mode of locomotion. Forelimb characters were, however, associated with digging or seed-harvest performance. Mechanical advantage of the proximal forelimb was positively related to an index of the compaction of soils with which 26 desert-dwelling species are associated, and also to relative use of heavy vs. light soils by nine species in the laboratory. Across 10 species, deviations in seed-harvest rate from expected allometric values were negatively correlated with mechanical advantage of the distal forelimb.     

The effects of owl predation on the foraging behavior of heteromyid rodents

Summary Researchers have documented microhabitat partitioning among the heteromyid rodents of the deserts of North America that may result from microhabitat specific predation rates; large/bipedal species predominate in the open/risky microhabitat and small/quadrupedal species predominate in the bush/safer microhabitat. Here, we provide direct experimental evidence on the role of predatory risk in affecting the foraging behavior of three species of heteromyid rodents: Arizona pocket mouse (Perognathus amplus; small/quadrupedal), Bailey's pocket mouse (P. baileyi; large/quadrupedal), and Merriam's kangaroo rat (Dipodomys merriami; large/bipedal). Both kangaroo rats and pocket mice are behaviorally flexible and able to adjust their foraging behavior to nightly changes in predatory risk. Under low levels of perceived predatory risk the kangaroo rat foraged relatively more in the open microhabitat than the two pocket mouse species. In response to the presence of barn owls, however, all three species shifted their habitat use towards the bush microhabitat. In response to direct measures of predatory risk, i.e. the actual presence of owls, all three species reduced foraging and left resource patches at higher giving up densities of seeds. In response to indirect indicators of predatory risk, i.e. illumination, there was a tendency for all three species to reduce foraging. The differences in morphology between pocket mice and kangaroo rats do appear to influence their behavioral responses to predatory risk.     

华山松种子大小对啮齿动物贮藏行为的影响及其时空变化

华山松(Pinus armandii)是广泛分布在中国中西部海拔1000～3300m的山地、种子较大(约300mg)的松属植物。为探究华山松种子大小对啮齿动物贮藏行为的影响,于2006年和2007年在滇西北3个不同地点进行种子标记和追踪实验。结果表明,在所有年份和地点,啮齿动物都倾向于贮藏更多的大种子和取食更多的小种子;啮齿动物贮藏大种子的数量,以及平均距离和最大距离均显著高于小种子。3个地点具有不同的啮齿动物群落结构,从而对种子命运产生显著影响。种子命运在两个年份间也存在显著差异。     

Effects of seed size of wind-dispersed pines (Pinus) on secondary seed dispersal and the caching behavior of rodents

Seeds and nuts dispersed by scatter-hoarding animals are relatively large compared to propagules dispersed by other means. Possible selective forces in the evolution of large seed size include the selectivity of foraging animals and the ways that food-storing animals treat seeds and nuts after harvest. Treatment by rodents, primarily yellow pine chipmunks ( Tamias amoenus ), of four species of pine seeds that vary in size was studied in the Carson Range of western Nevada. The pines, lodgepole pine ( Pinus contorta , 8.7 mg), ponderosa pine ( P. ponderosa , 55 mg), Jeffrey pine ( P. jeffreyi , 157 mg), and sugar pine ( P. lambertiana , 213 mg), produce winged seeds that are initially wind-dispersed but are gathered by rodents and cached in the soil. Radioactive scandium-46 was used to follow the fates of seeds of all fours species placed around three source trees during autumn 1998 to 2000. Rodents gathered the seeds of all four species, but they took fewer of the lodgepole pine seeds and only six lodgepole seed caches (n=2106 total caches) were found during the three years. Among the other three species, number of seeds per cache decreased with increasing seed mass. However, the product of number of seeds per cache and seed mass was similar for all species. Sugar pine seeds were cached slightly deeper than ponderosa and Jeffrey pine seeds. For the species examined, seed size appeared to have had little effect on several other attributes, including mean dispersal distance, substrate choice, and microhabitat choice. Large size decreases wind dispersibility of pine seeds, but secondary dispersal by scatter-hoarding rodents compensates for poor wind dispersal so that total dispersibility of large-seeded pines is not compromised.     

Salivary glands of heteromyid rodents, with a summary of the literature on rodent submandibular gland morphology

The principal parenchymal elements of the submandibular glands of the heteromyid rodents Dipodomys merriami, Perognathus longimembris, Perognathus fallax, Perognathus penicillatus and Perognathus baileyi consist of acini, granular tubules and striated ducts. Acinar cells of the four species of Perognathus are aniline blue, PAS (magenta) and Alcian blue (pH 2.5) positive and metachromatic with toluidine blue and safranin. The granules of the tubule cells are orthochromatic and react with aniline blue, orange G, the PAS reagent (deep pink) and the tryptophan indicator, xanthydrol. Acinar and tubule cells of D. merriami exhibit similar reactions except for the Alcian blue stain. Acinar cells of D. merriami do not react with Alcian blue. Submandibular glands of D. merriami exhibit a sexual dimorphism of the granular tubules. There is little observable difference between the sexes in the species of Perognathus but the ratio of granular tubules to acinar elements, the degree of hypertrophy of the tubules, and the amount of mucosubstance and protein (granules) contained in their cells are different in the four species studied. Since these desert rodents have similar habitats and habits, the differences observed between the two heteromyid subfamilies studied, as well as among the four members of a single subfamily, suggest that these are inherent species variations rather than variations of adaptation to environment.     

楝树籽提取物对土壤供氮和氮肥利用率的影响

研究了楝树籽不同方式的提取物（NI,NⅡ）对普通铁质淋溶土壤,简有水耕人为土壤供N和N肥利用率的影响。NI可在一定程度上抑制两种土壤产生NO3－N,NⅡ在两种土壤上前期均能显著固定NH4^ -N并在后期释放出NH4^ -N。如果NI和NⅡ2种提取物同时与化肥N施入土壤,则可明显改善土壤的供N状况,使这与作物需N过程更吻合。盆栽实验证明,NⅡ可显著提高肥旱耕人为土壤上作物对N肥的利用率。     

Effects of summer and winter harvesting on element phytoextraction efficiency of Salix and Populus clones planted on contaminated soil

The clones of fast-growing trees (FGTs) were investigated for phytoextraction of soil contaminated with risk elements (REs), especially Cd, Pb, and Zn. As a main experimental factor, the potential effect of biomass harvesting time was assessed. The field experiment with two Salix clones (S1 – (Salix schwerinii × Salix viminalis) × S. viminalis, S2 – S. × smithiana) and two Populus clones (P1 – Populus maximowiczii × Populus nigra, P2 – P. nigra) was established in April 2009. Shoots of all clones were first harvested in February 2012. After two further growing seasons, the first half of the trees was harvested in September 2013 before leaf fall (summer harvest) and the second half in February 2014 (winter harvest). Remediation factors (RFs) for all clones and all REs (except Pb for clone S1) were higher in the summer harvest. The highest annual RFs for Cd and for Zn (1.34 and 0.67%, respectively) were found for clone S2 and were significantly higher than other clones. Although no increased mortality of trees harvested in the summer was detected in the following season, the effect of summer harvesting on the phytoextraction potential of FGTs clones should be investigated in long-term studies.     

Codivergence in heteromyid rodents (Rodentia: heteromyidae) and their sucking lice of the genus Fahrenholzia (Phthiraptera: anoplura)

Although most studies of codivergence rely primarily on topological comparisons of host and parasite phylogenies, temporal assessments are necessary to determine if divergence events in host and parasite trees occurred contemporaneously. A combination of cophylogenetic analyses and comparisons of branch lengths are used in this study to understand the host-parasite association between heteromyid rodents (Rodentia: Heteromyidae) and their sucking lice of the genus Fahrenholzia (Phthiraptera: Anoplura). Cophylogenetic comparisons based on nucleotide substitutions in the mitochondrial COI gene reveal a significant, but not perfect, pattern of cophylogeny between heteromyids and their sucking lice. Regression analyses show a significant functional relationship between the lengths of analogous branches in the host and parasite trees, indicating that divergence events in hosts and parasites were approximately contemporaneous. Thus, the topological similarity observed between heteromyids and their lice is the result of codivergence. These analyses also show that the COI gene in lice is evolving two to three times faster than the same gene in their hosts (similar to the results of studies of other lice and their vertebrate hosts) and that divergence events in lice occurred shortly after host divergence. We recommend that future studies of codivergence include temporal comparisons and, when possible, use the same molecular marker(s) in hosts and parasites to achieve the greatest insight into the history of the host-parasite relationship.     

Foraging behaviour of harvesting ants determines seed removal and dispersal

Harvesting ants can affect the regeneration of plants through at least two different processes: seed removal and seed dispersal. We analyse the role of different foraging strategies of ants on patterns of seed removal and dispersal by three Messor species with considerable differences in their foraging systems. Messor capitatus workers rarely leave the nest in well-formed columns, while the other two species form foraging trails, with M. bouvieri forming temporary trails and M. barbarus foraging on a stable system of permanent foraging trails. Overall seed intake of M. capitatus colonies is considerably less than that of the two group-foraging species. There are also differences in the size of seeds collected: M. barbarus and M. capitatus harvest similar amounts of large and small seeds, while M. bouvieri harvests small seeds more intensely than large ones, due to the smaller size of the worker caste. The three Messor species differ in the percent of seed dropping of the different seed type and in the seed dispersal distance. Moreover, M. bouvieri and M. capitatus redistributed dropped seeds preferentially in bare soil and low sparse vegetation habitats, while M. barbarus redistributed seeds mainly in the high vegetation habitat. These results show that the foraging systems of these harvesting ants determine different patterns of seed removal and dispersal and, thus, affect the abundance and redistribution of seeds in the area.     

Tree-to-tree variation in seed size and its consequences for seed dispersal versus predation by rodents

Individual variation in seed size and seed production is high in many plant species. How does this variation affect seed-dispersing animals and, in turn, the fitness of individual plants? In this study, we first surveyed intraspecific variation in seed mass and production in a population of a Chinese white pine, Pinus armandii. For 134 target trees investigated in 2012, there was very high variation in seed size, with mean seed mass varying among trees almost tenfold, from 0.038 to 0.361 g. Furthermore, 30 of the 134 trees produced seeds 2 years later, and for these individuals there was a correlation in seed mass of 0.59 between years, implying consistent differences among individuals. For a subset of 67 trees, we monitored the foraging preferences of scatter-hoarding rodents on a total of 15,301 seeds: 8380 were ignored, 3184 were eaten in situ, 2651 were eaten after being cached, and 395 were successfully dispersed (cached and left intact). At the scale of individual seeds, seed mass affected almost every decision that rodents made to eat, remove, and cache individual seeds. At the level of individual trees, larger seeds had increased probabilities of both predation and successful dispersal: the effects of mean seed size on costs (predation) and benefits (caching) balanced out. Thus, despite seed size affecting rodent decisions, variation among trees in dispersal success associated with mean seed size was small once seeds were harvested. This might explain, at least in part, the maintenance of high variation in mean seed mass among tree individuals.     

Native seed preferences of shrub-steppe rodents,birds and ants: the relationships of seed attributes and seed use

Summary This study established the preferences of shrubsteppe granivores among seeds of 6 common sagebrushsteppe plants and related the preferences observed to physical and nutritional attributes of the seeds. Seeds of big sagebrush (Artemisia tridentata), cheatgrass (Bromus tectorum), Indian ricegrass (Oryzopsis hymenoides), western wheatgrass (Pascopyrum smithii), bitterbrush (Purshia tridentata) and green needlegrass (Stipa viridula) were placed in groups of petri dishes designed such that seed removal could be ascribed to either diurnal vertebrates, nocturnal vertebrates or ants. Though absolute quantities of seeds removed varied among the 3 granivore classes, calculations of preference based on weights of each seed species removed by each granivore class indicated that all 3 ranked the seeds similarly. Preference hierarchies of the 3 granivore classes were highly positively correlated with both calories per seed and % soluble carbohydrate of the seeds. The first correlation supports a basic prediction of optimal foraging theory —that foragers should maximize energy intake per unit time spent foraging. Both correlations emphasize the role of seed nutritional qualities in granivore seed selectivity in that soluble carbohydrate is a water-efficient energy source and its percentage is a good indicator of the digestible energy available in a food item. A corollary experiment comparing granivore use of an exotic seed (millet [Panicum miliaceum]) and a preferred native seed (Oryzopsis) demonstrated a distinct preference for the exotic. Since millet seeds are particularly high in % soluble carbohydrate, this result reinforced the apparent value of this nutritional attribute as a predictor of granivore seed preference. Among many seed resource characteristics upon which granivore seed selectivity might operate, our results indicate that individual species' nutritional composition may be particularly important. Thus, inferences about seed selectivity and resource partitioning among arid-land granivores should be interpreted with caution, especially those based on experiments using seed introductions, since the influence of seed nutritional attributes has not been widely acknowledged heretofore.     

Evidence for secondary seed dispersal by rodents in Panama

Summary The data presented show thatVirola nobilis (Myristicaceae), a bird/mammal-dispersed tree species in Panama, may also be dispersed by a terrestrial rodent, the agouti (Dasyprocta punctata). Using a thread-marking method, we observed that agoutis scatterhoardedV. nobilis seeds that they found both singly or in clumps. Seed removal and seed burial rates were strongly affected by features of forest habitats, such asV. nobilis tree richness (rich vs poor) and/or forest age (old vs young), but not by seed dispersal treatment (scattered vs clumped). Predation (mostly post-dispersal) of unburied seeds by weevils was independent of habitat and dispersal treatment. Seeds artificially buried in aVirola-rich area were more likely to escape predation and become established than unburied seeds under natural conditions. The food reward for agoutis is in the germinating seedlings. The seed dispersal syndrome ofV. nobilis involves long- and short-distance dispersers which both appear important for tree recruitment.     

啮齿动物和鸟类对东灵山地区辽东栎种子丢失的影响

20 0 0年 8月中旬至 10月上旬 ,在北京东灵山地区小龙门林场选取两块辽东栎分布近似而坡向不同的样地 ,并对样地内种子库与啮齿动物的种群数量变化进行了调查。结果表明 ,两块样地种子雨持续 4 0天左右 ,且种子下落趋势基本一致 ,高峰期都集中在 9月中旬 ,不同坡向的种子产量差异显著。在两样地随机各设置 2 4个种子方形收集器 (0 5m2 )和 2 0个地表样方 (1 0× 0 5m2 )调查种子产量。通过比较收集器内壳斗和种子数量 ,发现二者无显著差异 ,说明鸟类对林冠层种子丢失作用不明显 ;而收集器和地表样方种子数量差异显著 ,表明辽东栎种子库扩散主要由林中啮齿动物完成。标记重捕发现辽东栎林中啮齿动物群落包括大林姬鼠 ,社鼠 ,棕背鼠平和花鼠 ,其中大林姬鼠为优势种 ,占群落的 77.2 %。