小果野芭蕉种子散布和不同时空尺度上种子被捕食格局

 以分布在云南西双版纳地区的大型先锋草本植物小果野芭蕉(Musa acuminata)为研究材料,研究其种子初次散布过程和不同时空尺度上种子被 捕食格局。小果野芭蕉的成熟果实有75%在夜间被取食和传播,在白天消失的果实则占25%。蝙蝠是其最主要的种子传播者,鸟类在其种子传播 过程中也起到一定的作用。人工摆放种子试验结果显示小果野芭蕉种子的主要转移者是小型啮齿类(鼠类)和蚁类：在开放处理下3 d后转移率为 86%,排除蚁类(鼠类可进入)处理下种子转移率为69%以及排除鼠类(蚂蚁可进入)处理下种子被转移率为56%。季节、地点和生境均显著影响人工 摆放种子被转移强度：雨季显著高于旱季(p<0.001), 野芭蕉生境显著高于与其相连的自然森林和荒地(p<0.001),在人为干扰较少的补蚌自然 保护区显著低于西双版纳热带植物园和新山,而后两者之间并无显著差异(p>0.05)。同时,地点和生境以及季节、地点和生境都有显著的交互 作用。与相邻的森林和荒地相比,野芭蕉群落中种子被鼠类捕食的强度最大且受蚁类二次转移的比例最少,森林和荒地中种子被鼠类捕食的强 度相对较小且蚁类对种子的二次转移比例较高,从而更好地帮助种子逃避鼠类捕食。因此,依赖于食果动物(主要是蝙蝠, 也包括鸟类)的初次 散布是小果野芭蕉种子逃避捕食的关键。     

Seed Ecology of Woody Species in a Caribbean Dry Forest

Peak flowering activity among woody species in the tropical dry forests of St. John, U.S. Virgin Islands, coincided with the brief spring rainy season but continued at moderate levels for six months, abating with the autumn rains. Fruit maturation showed a major peak in the long winter dry season and a minor crest during the summer dry season. Seeds of wind-dispersed species disseminated mainly during the winter dry season, while animal dispersal of seeds (74% of all woody species) followed the bimodal pattern (for wet and dry seasons) described for the community as a whole. Under shadehouse conditions, most dry forest tree species germinated well (> 80%) and emerged promptly (within four weeks of planting) and synchronously (90% emergence within a four-week interval). Nine of 29 species tested in the shadehouse manifested dormancy of at least six weeks. Seed germinability varied among tree species, and the viability of most species began to decline following six months of dry storage. Few species retained high germinability after nine months of dry storage. The species composition of soil seed banks did not correspond closely with above-ground communities on three forested sites of varying stand age. In the youngest stand (35 years old), dominated by the weedy, arborescent legume Leucaena leucocephala, the soil seed bank was also dominated by this species, but no seeds of any other tree species were found in the soil samples. Seeds of native trees were scarcely encountered (only one indigenous species) in soil seed bank samples of three forest sites. Local seed rain from less disturbed forest may not be sufficient for prompt recovery of the dry forest community on degraded sites.     

Interaction between a threatened endemic plant (<Emphasis Type="Italic">Anchusa crispa</Emphasis>) and the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Seed dispersal mutualisms are essential to ensure the survival of diverse plant species and communities worldwide. Here, we investigated whether the invasive Argentine ant can replace native ants by fulfilling their functional role in the seed dispersal of the rare and threatened endemic myrmecochorous plant, Anchusa crispa, in Corsica (France). Our study addressed the potential of Linepithema humile to disperse elaiosome-bearing seeds of A. crispa, examining L. humile’s effects on (1) the composition of communities of ants removing seeds, (2) the number of seed removals, (3) seed preference, (4) the distance of seed dispersion, and (5) seed germination. We caught seven native species at the control site, but only the Argentine ant at invaded sites. L humile removed A. crispa seeds in greater numbers than did native ants, respectively 66 and 23%, probably due to their higher worker density. The invader was similar to native ants with respect to distance of seed transport. Finally, rates of seed germination were not significantly different between seeds previously in contact with either Argentine ants or not. Taken all together, these results suggest that the Argentine ant is unlikely to pose a threat to A. crispa population. These results have important implications for the management of this rare and threatened endemic plant and provide an example of non-negative interactions between invasive and native species.     

西双版纳热带森林土壤种子库的季节变化

通过萌发实验法对西双版纳地区的一类热带季节雨林 (番龙眼、千果榄仁 )和 2类次生林(白背桐林、中平树林 )的土壤种子库的季节变化进行了探讨。结果表明 :该地区的土壤种子库动态具有明显的季节性。季节雨林的土壤种子库储量相对稳定 ,土壤上层 ( 0～ 2 cm )的种子储量在雨季末期较大。2类次生林土壤种子库的变化则相反 ,土壤种子库中的种子种类在旱季末期较雨季末期多 ,土壤上层的种子储量在旱季末期较大。各样地均有一些种类只出现在旱季末期或雨季末期。种子在土壤种子库的动态与植物的繁殖物候和所处的环境紧密相关 ,不同种类植物的土壤种子库由于植物本身的生物学特性、传播方式和所处环境的影响而表现出不同的动态模式     

Seed rain from forest fragments into tropical pastures in Los Tuxtlas,Mexico

Deforestation has resulted in the fragmentation of forests. Remnant fragments are widely assumed to be sources of seeds for forest regeneration in abandoned pastures. The seed rain in 12 pastures at Los Tuxtlas, Mexico, their relationship with riparian vegetation adjacent to them, and their closeness to the reserve is described. For all the species found in the seed rain, we classified them by typical habitat (pasture or rain forest), life form, dispersal syndrome, and plant strategy (pioneer or non-pioneer species). In addition, germinability was evaluated for all seeds. We also assessed the correlation in composition of the seed rain and the riparian vegetation. Only 11% of the total species that occur in the Los Tuxtlas region reach pastures via dispersal from forest fragments. However, nearly 80% of the species in the seed rain did not come from the fruiting individuals in the adjacent riparian vegetation. The proportion of dispersal syndromes, life forms and plant strategies of the species in the seed rain was similar to those observed in the rain forest. On average, the germinability of forest species was less than 30%.The forest species richness was similar in the seed rain in pastures and inside the forest, but the pasture seed rain contained fewer seeds per species. Pastures have a high potential for natural regeneration because of seed dispersal from adjacent forest. However, the forest that regenerates in pastures close to the reserve is expected to contain different species than the forest regenerating far away from it.     

Water dispersal as an additional pathway to invasions by the primarily wind-dispersed tree <Emphasis Type="Italic">Ailanthus altissima</Emphasis>

Long-distance dispersal is a key process in biological invasions. Previous research has emphasized the role of nonstandard dispersal vectors, but consequences of a change in dispersal vector for the establishment of invasive plant species have received less attention. We analyzed how water-mediated dispersal rather than the more expected wind-mediated dispersal can affect the establishment of the invasive tree Ailanthus altissima in riparian corridors by changing the germination rate and velocity and by providing the option of a new pathway of vegetative propagation. We analyzed the potential of different types of propagules (fruits that have floated or been submerged, current- and second-year stem fragments) to establish new individuals after contact with water for 0, 3, 10, and 20 days. Length and type of seed contact with water led to divergent germination responses. Seeds that had floated for 3 days had an increased level of seed germination (87%), while a 20-day stay in water water-curbed germination to 32% compared to 53% in control. After floatation, the maximum number of emerged seedlings was achieved more than 3 weeks earlier than in all other treatments. In general, the germination was enhanced in floating compared to submerged fruits. Experiments with stem fragments revealed the option of a novel pathway for long-distance dispersal in river corridors: Except for stem fragments that floated for 20 days, 33–75% of buried stem fragments produced adventitious shoots, 10% also set roots. The results suggest that both generative and vegetative propagules of A. altissima can be dispersed at regional scales in river corridors. Hence, water as an additional dispersal vector is expected to enhance invasions by species with wind-dispersed seeds. Our findings suggest the importance of control of initial colonizations in riparian habitats and emphasize the need to include consequences of secondary dispersal when modeling the spread of invasive species.     

Size traits and site conditions determine changes in seed bank structure caused by grazing exclusion in semiarid annual plant communities

1. Contrasting patterns of change in the seed bank of natural grasslands are frequently found in response to grazing by domestic herbivores. Here, we studied the hypotheses that a) patterns of change in seed bank density and composition in response to grazing depend on spatial variation in resource availability and productivity, and b) that variation among species in patterns of seed bank response to grazing is linked to differences in species size traits (i.e. size of plant, dispersal unit and seed). 2. Effects of sheep grazing exclusion on the seed bank were followed during five years in a semiarid Mediterranean annual plant community in Israel. Seed bank density and composition were measured in autumn, before the rainy season, inside and outside fenced exclosures in four neighboring topographic sites differing in vegetation characteristics, soil resources and primary productivity: Wadi (dry stream terraces, high productive site), Hilltop, South‐ and North‐facing slopes (less productive sites). 3. Topographic sites differed in seed density (range ca 2500–18000 seed m^?2) and in seed bank response to grazing exclusion. Fencing increased seed density by 78, 51 and 18% in the Wadi, South‐ and North‐facing slopes, respectively, but had no effect in the Hilltop. At the species level, grazing exclusion interacted with site conditions in determining species seed bank density, with larger or opposite changes in the high productive Wadi compared to the other less productive sites. 4. Changes in seed bank structure after grazing exclusion were strongly related to species size traits. Grazing exclusion favored species with large size traits in all sites, while seed density of tiny species decreased strongly in the high productive Wadi. Species with medium and small size traits showed lesser or no responses. 5. The size of plants, dispersal units and seeds were strongly correlated to each other, thus confounding the evaluation of the relative importance of each trait in the response of species to grazing and site conditions. We propose that the relative importance of plant size vs seed size in the response to grazing changes with productivity level.     

Dormancy as exaptation to protect mimetic seeds against deterioration before dispersal

Background and Aims

Mimetic seeds simulate the appearance of fleshy fruits and arilled seeds without producing nutritive tissues as a reward for seed dispersers. In this strategy of seed dispersal, seeds may remain attached to the mother plant for long periods after maturity, increasing their availability to naïve seed dispersers. The hypothesis that seed coat impermeability in many tropical Fabaceae with mimetic seeds serves as an exaptation to protect the seeds from deterioration and rotting while awaiting dispersal was investigated.

Methods

Seed coat impermeability was evaluated in five mimetic-seeded species of tropical Fabaceae in south-eastern Brazil (Abarema langsdorffii, Abrus precatorius, Adenanthera pavonina, Erythrina velutina and Ormosia arborea) and in Erythrina speciosa, a ‘basal’ species in its genus, which has monochromatic brown seeds and no mimetic displays. Seed hardness was evaluated as a defence against accelerated ageing (humid chamber at 41 °C for 144 h). Seed development and physiological potential of O. arborea was evaluated and the effect of holding mature seeds in pods on the mother plant in the field for a period of 1 year under humid tropical conditions was compared with seeds stored under controlled conditions (15 °C and 40 % relative air humidity).

Key Results

All five mimetic-seeded species, and E. speciosa, showed strong coat impermeability, which protected the seeds against deterioration in accelerated ageing. Most O. arborea seeds only became dormant 2 months after pod dehiscence. Germination of seeds after 1 year on the plant in a humid tropical climate was 56 %, compared with 80 % for seeds stored in controlled conditions (15 °C, 45 % relative humidity). Seedling shoot length after 1 year did not differ between seed sources.

Conclusions

Dormancy acts in mimetic-seeded species as an exaptation to reduce seed deterioration, allowing an increase in their effective dispersal period and mitigating the losses incurred by low removal rates by naïve avian frugivores.     

Seed bank dynamics and tree‐fall gaps in a northwestern Mexican Quercus‐Pinus forest

Questions: How does the seed bank respond to different types of tree‐fall gaps and seasonal variations? How does the soil seed bank influence recovery of the standing vegetation in the mature forest and tree‐fall gaps? Location: 1800 — 2020 m a.s.l., Quercus‐Pinus forest, Baja California Sur, Mexico. Methods: Seed size, species composition and germination were estimated under different environmental conditions during dry and rainy seasons: a mature forest plot and gaps created by dead standing trees, snapped‐of f trees and uprooted trees. The soil seed bank was investigated using direct propagule emergence under laboratory conditions, from soil cores obtained during both seasons. Results: 21 species, 20 genera and 14 families constitute the seed bank of this forest community. Fabaceae, Asteraceae, Euphorbiaceae and Lamiaceae were the most frequently represented families in the seed bank. Floristic composition and species richness varied according to the different modes of tree death. Species composition of seed banks and standing vegetation had very low similarity coefficients and were statistically different. Seed bank sizes varied between 164 and 362 ind.m^‐2 in the mature forest plot for the dry and rainy seasons, respectively, while soil seed bank sizes for gaps ranged between 23–208 ind.m^‐2 forthe dry season and between 81–282 ind.m^‐2 for the rainy season. Conclusions: Seed bank sizes and germination response were always higher in the rainy season under all the environmental conditions analysed. Results suggest that timing responses to gap formation of the soil seed bank could be more delayed in this temperate forest than expected.     

Hydrodynamics and seed dispersal in the lower Amazon

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Secondary seed dispersal in Montrichardia arborescens (L.) schott dominated wetlands in Laguna Grande, Venezuela

Laguna Grande, Monagas State, Venezuela, is a shallow, V-shaped lake created by the confluence of two rivers. Montrichardia arborescens (L.) Schott. dominated wetlands cover most of the north and south arms and the littoral zone of the main body of the lake. The vegetation and seed banks of Montrichardia wetland sites were sampled in the north arm, south arm and main body five times from the end of the dry season in 1991 to beginning of the rainy season in 1992. The composition of the vegetation was similar and changed very little at all three sites during the course of the study. These wetlands had 53 species. Besides M. arborescens, other common species were Hamelia patens Jacq., Mikania cordifolia (L.) Wild., Sarcostemma clausum (Jacq.) Roem. & Schult., and Vitis caribaea L. In both the vegetation and seed banks, species richness was highest during the dry season. Altogether, the seed banks contained the seeds of 61 species of which 35 were also found in the vegetation. Seeds of three tree species were found in the seed banks that did not grow anywhere in the lake. In the seed bank, seeds of Cyperus odoratus L., Eleocharis interstincta (Vahl.) R&S, Ludwigia hyssopifolia (G. Don.) Excell, L. lithospermifolia (Mich.) Hara, Polygonum acuminatum H.B.K., and Sacciolepis striata (L.) Nash were the most abundant. Mean total seed density over the entire study was 6,500, 3,800, and 6,000 seeds/m² in the north arm, south arm, and main basin, respectively. Seed production and dispersal occur primarily during the dry season, and the highest seed densities at all sites were found in the dry season when there was no or little standing water. The lowest seed densities at all sites were found during the rainy season during which seed densities declined over 80% at the north and south arm sites. In the main body of the lake, however, seed densities during the rainy season, although lower than during the dry season, actually increased significantly from 3,600 seeds/m² in August 1991 to 6,000 seeds/m² in October 1991. A significant decrease in seed density in either the north or south arms or both and a significant increase in the main body site during the rainy season occurred for 5 of the 8 species whose seeds were the most abundant, for all life-form guilds, except hydrophytes and for the entire seed bank. Secondary dispersal by water currents during the rainy season appears to be transporting seeds from the north and south arms into the seed bank of the main body of the lake.     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Potential of endozoochorous seed dispersal by sheep in calcareous grasslands: correlations with seed traits

Questions: What is the potential of sheep to serve as seed dispersers via ingestion and defecation in calcareous grasslands? Is the presence of viable seeds from dung correlated with specific seed traits? Location: Calcareous grasslands, South Limburg, the Netherlands/Belgium. Methods: Dung samples (n=24) from sheep were collected between September 2006 and November 2007 from five sites with Mesobromion plant communities, and communities of Nardo‐Galion saxatilis. Germinability and identity of seeds in the dung samples were ascertained from germination of seedlings under glasshouse conditions. Seed traits of species with viable seeds in dung were compared with those present in the local species pool. Results: Seventy‐two plant species from 23 plant families had viable seeds in sheep dung. The plant families encountered most frequently were Gramineae and Compositae. The most abundant and frequently recorded plant species in dung samples was Urtica dioica, accounting for >80% of the total number of seeds. Mean seed density in sheep dung was 0.8 seeds g^?1 dry matter. Seeds with low seed mass and a high seed longevity index were over‐represented in dung. Viable seeds >2.5 mg were infrequent in the dung samples. Conclusions: We conclude that sheep are potentially important dispersers of plant species in Dutch calcareous grasslands. Although smaller seeds were relatively abundant in sheep dung, it cannot be excluded that this was mainly caused by differences in seed abundance.     

Understory spider diversity in two remnants of tropical montane cloud forest in Chiapas, Mexico

We evaluated the spider diversity of a tropical montane cloud forest understory in two nearby sites with different degree of human disturbance at the Biosphere Reserve Volcán Tacaná, Chiapas, Mexico. The study was conducted over a 24 days period distributed in 6 months in 2009, covering dry and rainy seasons. A total of 8,370 spiders (1,208 adults and 7,162 juveniles) were collected. Determined specimens (7,747) represented 112 species and morphospecies, 71 genera and 22 families. The results showed that human disturbance has an influence on spider communities: species richness was significantly higher in the preserved site as regards to the disturbed site. Despite their proximity, the composition of spider communities showed only a moderate similarity between the two sites. No differences in abundance were found among sites when considering the whole sample, but sites differed clearly when seasons were analyzed separately. The rainy season had a negative effect on the abundance of spiders in the preserved site. Although the spider community structure was very similar between sites, there was a trend towards a greater species evenness in the preserved site for the whole sampling period and for the dry season.     

Endozoochorous seed dispersal by free-ranging herbivores in an abandoned landscape

Seed dispersal is a crucial process for the dynamics and maintenance of plant populations. Free-ranging animals are effective dispersal vectors because they can move between similar habitats and transport seeds into favourable environments. Dung samples from two species of common free-ranging mammals—deer and wild boar—were used to study endozoochorous dispersal of seeds in a military training area in western Bohemia. The area was abandoned after WWII, and the military training area was established in 1953. The vegetation consists of shrublands and dry grasslands. Data on the local species pool of grassland herbs and forbs were collected to compare the characteristics of dispersed versus non-dispersed plants. Deer and wild boar dispersed 84 plant species; however, species composition of seedlings emerging from dung samples showed significant differences between dispersal vectors and notable change across the growing season. 80% of all seedlings extracted from the dung samples belonged to stinging nettle, Urtica dioica. From trait analyses, seeds of endozoochorous plants had a higher longevity index in the soil seed bank than non-endozoochorous plants and more often possessed a mucilaginous surface. Our results show that deer and boar are successful, though not substitutable dispersers.     

Individual variation in resource use by opossums leading to nested fruit consumption

Despite recent findings on the ecological relevance of within population diet variation far less attention has been devoted to the role diet variation for ecological services. Seed dispersal is a key ecological service, affecting plant fitness and regeneration based on foraging by fruit‐eating vertebrates. Here we used a network approach, widely used to understand how seed‐dispersal is organized at the species level, to gain insights into the patterns that emerge at the individual‐level. We studied the individual fruit consumption behavior of a South American didelphid Didelphis albiventris, during the cool–dry and warm–wet seasons. In species–species networks the heterogeneity in specialization levels generates patterns such as nestedness and asymmetry. Because generalist populations may be comprised of specialized individuals, we hypo thesized that network structural properties, such as nestedness, should also emerge at the individual level. We detected variation in fruit consumption that was not related to resource availability, ontogenetic or sexual factors or sampling biases. Such variation resulted in the structural patterns often found in species–species seed‐dispersal networks: low connectance, a high degree of nestedness and the absence of modules. Moreover structure varied between the warm–wet and cool–dry seasons, presumably as a consequence of seasonal fluctuation in fruit availability. Our findings suggest individuals may differ in selectivity causing asymmetries in seed dispersal efficiency within the population. In this sense the realized dispersal would differ from the expected dispersal estimated from their average dispersal potential. Additionally the results suggest possible frequency‐dependent effects on seed dispersal that might affect individual plant performance and plant community composition.     

Soil seed bank dynamics of fire-prone wooded grassland, woodland and dry forest ecosystems in Ethiopia

In order to reveal the role of soil seed banks in vegetation recovery after fire in savanna, the spatial distribution and temporal changes in the soil seed banks of regularly burning savanna in Gambella, western Ethiopia, was studied. The seedling emergence technique was employed to determine the species composition and density of the soil seed bank of six sites ranging in fire severity from wooded grassland with frequent fires over woodland with intermediate fire frequency to forest with absence of fires. Species composition and density of seeds in the soil were compared between seasons, depths and sites with different types of standing vegetation. Fourteen plant species were recorded in the soil seed bank from the grassland and woodland sites and 6 from the dry forests; 60 % of the taxa in the soil seed bank were annuals and 40 % were perennials. The soil seed banks were largely dominated by graminoids and 48–97 % of the soil seed bank in the grasslands and woodlands was of a single grass species, Hyparrhenia confinis , which was absent from the dry forests. The soil seed pools ranged from less than 100 to 4700 seeds per m² depending upon the season. The soil seed bank of graminoids was nearly empty after the onset of the rainy season whereas seeds of broadleaved herbs and woody species able to germinate were still found after this time. Floristic composition, representation of life forms and density of seeds in soil did not correspond closely with that of the standing vegetation, but within graminoids there was a strong similarity between the soil seed bank and the standing vegetation. The current fire regime of Ethiopian savanna woodlands appears to maintain the dominance of graminoids over broadleaved herbs and woody plants both as seeds in the soil and in the standing vegetation.     

西双版纳热带季节雨林145个树种繁殖体特征

分析了西双版纳热带季节雨林20公顷动态监测样地中145个树种的繁殖体特征。结果表明：雨季散布的种类最多（71.03%）,在雾凉季及干热季散布的种类相对较少;果实类型以核果,蒴果,浆果居多,黑色、黄色、褐色果实最为常见;动物散布的种类最多（56.55%）,其次为机械散布,独立散布,风力散布;大种子树种（种子最大长度〉10mm）较占优势（53.10%）;树种单粒种子重量从2.3×10^-5～22.29g不等,但76.55%的种子的重量集中在0.01～10g之间;干热季散布的种子平均重量显著高于雨季和雾凉季;独立散布的种子重量最大,风力散布的种子重量最小,而机械弹射的种子重量和动物散布的种子重量之间差异不显著;树种的成年个体最大胸径与种子重量之间存在显著正相关。     

Genetic evidence for predominantly hydrochoric gene flow in the invasive riparian plant Impatiens glandulifera (Himalayan balsam)

Background and Aims

Riparian systems are prone to invasion by alien plant species. The spread of invasive riparian plants may be facilitated by hydrochory, the transport of seeds by water, but while ecological studies have highlighted the possible role of upstream source populations in the establishment and persistence of stands of invasive riparian plant species, population genetic studies have as yet not fully addressed the potential role of hydrochoric dispersal in such systems.

Methods

A population genetics approach based on a replicated bifurcate sampling design is used to test hypotheses consistent with patterns of unidirectional, linear gene flow expected under hydrochoric dispersal of the invasive riparian plant Impatiens glandulifera in two contrasting river systems.

Key results

A significant increase in levels of genetic diversity downstream was observed, consistent with the accumulation of propagules from upstream source populations, and strong evidence was found for organization of this diversity between different tributaries, reflecting the dendritic organization of the river systems studied.

Conclusions

These findings indicate that hydrochory, rather than anthropogenic dispersal, is primarily responsible for the spread of I. glandulifera in these river systems, and this is relevant to potential approaches to the control of invasive riparian plant species.     

Experimental assessment of plant seed retention times in fur of cattle and horse

Epizoochorous dispersal of plant seeds is an important long-distance dispersal mechanism. Yet little is known about retention times of seeds in animal furs and hence about potential dispersal distances of the seeds. Here, we used marked seeds of 12 plant species to determine seed depletion curves on Galloway cattle and Haflinger horse in three vegetation types (forest, tall herbage vegetation and meadow), in both dry and rainy weather conditions. In the long fur of Galloway cattle, seeds were retained significantly longer than in the short fur of Haflinger horse. In general, seed retention times were not considerably affected by the structure of the surrounding vegetation. The impact of the weather was negligible, only affecting the retention of some plant species. Negative exponential functions were fitted to the seed depletion curves. Using the parameters of curve estimations in the different conditions of animal species and vegetation structure, half-life seed retention times of up to 13 h for Galloway cattle and up to more than 4 h for Haflinger horse could be calculated, with corresponding potential half-life dispersal distances in the order of magnitude of tens of metres to a few kilometres. Different seed traits correlated with seed retention times in the long cattle fur and in the short horse fur, respectively.