西双版纳勐宋山区热带山地雨林的群落学研究

首次报道了云南西双版纳勐宋的热带山地雨林群落 ,该群落植物区系组成特殊 ,森林上层乔木组成中以古老的单室茱萸科 (Mastixiaceae)和紫树科 (Nyssaceae)植物为优势 ,亦富有木兰科 (Magnoliaceae)的长蕊木兰、云南拟单性木兰 ,红花木莲等系统发育上较原始的植物以及一些典型东南亚热带山地雨林群落的代表植物。通过对该热带山地雨林群落组成与结构的调查研究 ,认为该热带山地雨林可区分为沟谷和山坡 2个类型。沟谷群落类型以八蕊单室茱萸和大萼楠为乔木层优势种 ,可定义为八蕊单室茱萸—大萼楠林。山坡群落类型以云南拟单性木兰和云南裸花为优势 ,可定义为云南拟单性木兰—云南裸花林。     

Nonrandom co-occurrence patterns of rainforest chameleons

Null‐model analysis of co‐occurrence patterns is a powerful tool to identify ‘structure’ in community ecology data sets. We evaluated the community structure of chameleons in rainforest regions of Nigeria and Cameroon using available data in the literature, including peer‐reviewed articles and unpublished environmental reports to industries. We performed Monte Carlo simulations (5000 iterations, using the sequential swap algorithm) under several model assumptions to derive co‐occurrence patterns among species. Food and spatial (habitat) segregation patterns in both lowland rainforest and montane forest were investigated. We subjected four indices of co‐occurrence patterns (C‐ratio, number of checkerboard species pairs, number of species combinations, and V‐score) to randomization procedures. Overall, the chameleon communities do not show random organization, but instead exhibit precise deterministic patterns. In lowland rainforest, chameleon communities are assembled deterministically along the food niche resource axis, but not along the habitat niche resource axis. The opposite holds for chameleon communities in montane rainforest. We predict that these patterns can be generalized to other regions of tropical Africa, thus helping to determine the general structure of chameleon communities in tropical African forests.     

How resilient are Andean montane forest bird communities to habitat degradation?

The threatened forest habitats of the tropical Andes are reportedly being modified and destroyed 30% faster than their lowland tropical counterparts, but impacts on the hyper-diverse resident avifauna have received little systematic study. We present a baseline analysis of the effects of habitat modification on birds in a lower montane forest landscape in Ecuador, comparing avian community composition in landscape elements subjected to different levels of human modification: primary forest, secondary forest, edge habitat and agricultural land. We use data from a point count survey of 300 counts at 150 sites to test whether community composition and density of birds with different reported habitat preferences and foraging strategies change among landscape elements. Species richness and diversity were lowest in agricultural land, but on some measures, equally low in primary forest. Richness and diversity peaked in secondary forest and edge habitat, but ordination and density analysis revealed clear differences in their species composition. While secondary forest contained mostly forest-preferring species, edge habitat harboured a mix of forest and open-land birds. There was a clearly structured gradient in species composition across landscape elements, with densities of habitat specialists, foraging guilds and families varying considerably from primary forest to agricultural land. Agricultural land was characterised by an assemblage of widespread, abundant species very different from that in core forest habitats. As such, while the majority of montane forest birds appear resilient to a certain level of habitat modification, they cannot persist, and are displaced, where forest has been cleared outright. We argue that, for Andean montane forests, preservation of mature secondary forest offers flexibility in supplementing preserved primary forest areas to provide sufficient habitat for the persistence of this incredibly diverse but severely threatened bird community.     

五指山热带雨林黑桫椤种群及其所在群落特征

黑桫椤(Alsophila podophylla)属国家二级保护植物.作者在海南岛五指山山地雨林和低地雨林分别设立样地对黑桫椤种群及其所在群落进行了调查,分析了热带雨林黑桫椤种群的年龄结构、空间分布格局、数量分布以及群落结构和物种多样性.结果表明,山地雨林和低地雨林中黑桫椤种群分别为稳定型种群和增长型种群,都是完全集群分布.低地雨林的黑桫椤平均密度是山地雨林的1.7倍.土壤pH值对黑桫椤种群密度有负的影响,全磷含量对其有正的影响.山地雨林林地土壤pH值大而全磷含量低,低地雨林刚好相反.土壤酸碱度和全磷含量的差异是导致两地黑桫椤数量差异的原因之一.pH值对黑桫椤平均密度的影响效应是全磷含量的2.5倍,说明pH值是影响黑桫椤种群密度的主要因素.黑桫椤所在群落垂直结构分为3个乔木亚层、2个灌木亚层及草本层.山地雨林黑桫椤所在群落的Simpson指数和Shannon-Wiener指数分别为0.957和5.305,均匀度为0.811;低地雨林黑桫椤所在群落的Simpson指数和Shannon-Wiener指数分别为0.937和5.484,均匀度为0.804.总之,五指山山地雨林和低地雨林的黑桫椤种群年龄结构、分布格局和数量分布都有一定差异,这些差异是由生境差异造成的.热带雨林黑桫椤所在群落结构复杂,物种多样性丰富.     

Are Montane Forests Demographic Sinks for Bornean White‐bearded Gibbons Hylobates albibarbis?

I estimated habitat-specific population densities for a population of Bornean white-bearded gibbons Hylobates albibarbis inhabiting seven distinct forest types at Gunung Palung National Park, West Kalimantan, Indonesia. Population densities in montane forests (0.44 individuals/km²) were almost ten times lower than those in the next best habitat (upland granite forest; 4.2 individuals/km²) and far lower than those in lowland forest types. Demographic data on 33 gibbon groups living across the seven forest types showed that reproduction was substantially depressed in montane forests compared to high-quality lowland habitats. A simple model suggests that montane forests are demographic sinks for gibbons at Gunung Palung. Follow-up data from observations of montane groups 5 yr after the initial observation period support this result. As high-quality lowland forests (source habitat for gibbons) are being disproportionately lost in and around Gunung Palung National Park due to illegal logging and conversion to oil palm plantations, an increasing percentage of the remaining forest in the park comprises sink habitat for gibbons. This result has disquieting implications for the long-term viability of gibbon populations at Gunung Palung. In addition, as montane forests are generally low-quality habitat for most rainforest vertebrates, and since lowland forests are being lost at alarming rates across the tropics, source-sink population dynamics similar to those I describe here may characterize populations of other tropical vertebrate species.     

Fine-scale habitat structure complexity determines insectivorous bird diversity in a tropical forest

Habitat complexity in reforested stands has been acknowledged as a key factor that influences habitat use by birds, being especially critical for habitat disturbance-sensitive species such as tropical understory insectivorous birds. Most studies regarding the relationship between forest structure and species diversity were conducted at the landscape scale, but different diversity patterns may emerge at a finer scale (i.e., within a habitat patch). We examined a tropical reforested area (State of Caldas, Colombia), hypothesizing that insectivorous bird richness, abundance, and foraging guild abundance would increase as intra-habitat complexity increases. We established 40 monitoring plots within a reforested area, measured their structural features, and determined their relationships with species richness, total abundance, and foraging guild abundance, using Generalized Additive Models. We found that the increasing variation in basal area, stem diameter, and number of stems was positively correlated with species richness, total abundance, and foraging guild abundance. Relationships between richness or abundance and structural features were not lineal, but showing curvilinear responses and thresholds. Our results show that heterogeneity on basal area, stem diameter, and the number of stems was more correlated to insectivorous bird richness and abundance than the average of those structural features. Promoting structural variation on reforested areas by planting species with different growth rates may contribute to increase the richness and abundance of a tropical vulnerable group of species such as the understory insectivorous birds.     

西双版纳热带雨林和热带山地常绿阔叶林附生苔藓的组成与多样性格局

为探讨西双版纳热带雨林和热带山地常绿阔叶林附生苔藓的组成、多样性格局及其维持机制,该文对两种植被的树干附生苔藓植物进行了调查与分析.所选择的样地位于西双版纳勐腊县,每种植被类型选择10个20 m×20 m的样方,每个样方选择约10棵样树.结果表明:(1)本调查共记录到附生苔藓20科39属60种,其中热带雨林附生苔藓有1...     

An altitudinal comparison of caterpillar (Lepidoptera) assemblages on Ficus trees in Papua New Guinea

Aim This analysis of caterpillar (Lepidoptera) beta‐diversity between tropical lowlands and highlands attempts to separate the effects of between‐site (1) turnover of herbivore species on particular host plants, (2) changes in host use by herbivores, and (3) turnover of plant species on changes in herbivore assemblages. Location Two rain forest areas 130 km and 1700 altitudinal metres apart were studied in Papua New Guinea: one in the lowlands (100 m a.s.l.) on the northern coast of the island and one in the central New Guinean cordillera at 1800 m a.s.l. Methods The analysis is based on caterpillar feeding records obtained by quantitative sampling and rearing of caterpillars from four Ficus species studied in the mountains and 21 Ficus species and 62 plant species from other genera and families studied in the lowlands, including three Ficus species studied in both areas. Results Only 17% of species feeding on Ficus in the highlands also occurred in the lowlands. These species represented 1–46% of individuals in caterpillar assemblages on particular Ficus hosts. Widespread species included both Ficus specialists and generalists feeding on numerous plant families. Some of the Ficus specialists changed their preferred host species with altitude. High species turnover was not explained by changes in the species composition of host plants with altitude as lowland and montane assemblages feeding on the same Ficus species showed high turnover. Despite the rarity of widespread caterpillars, the lowland and montane Ficus assemblages were remarkably similar in their dominance structure, species richness, host specificity, generic composition and familial composition. Main conclusions Ficus‐feeding Lepidoptera assemblages between tropical lowlands and highlands are characterized by substantial species turnover not explained by altitudinal changes in the composition of the vegetation. Further, species‐rich plant genera can support caterpillar assemblages with relatively low beta‐diversity compared with species‐poor genera as caterpillars can switch their host preferences from one congeneric host species to another along an altitudinal gradient. Closely related plant species can thus represent a broad, continuously distributed resource along such gradients.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

西双版纳热带季节雨林的研究

本文以多个样方的资料分析,从群落综合特征上研究了西双版纳热带季节雨林的特点和类型、物种组成、生活型谱、叶级谱等。根据上层标志树种、群落生态外貌特征和生境把西双版纳的热带季节雨林区分为低丘雨林和沟谷雨林二个群系组,各包括若干群系。低丘雨林中面积最大和最有代表性的群系是大药树、龙果林,而沟谷雨林中最有代表性的群系是番龙眼、千果榄仁林。二者相比,低丘雨林的物种多样性要小一些,附生植物相对少一些,小、矮高位芽植物和小叶、落叶树种比例稍高,在生态上向季雨林和热带山地的常绿阔叶林过渡,有更强的地方代表性。沟谷雨林则更接近典型的湿热带雨林。     

Altitudinal zonation of montane Quercus forests along two transects in Chirripó National Park,Costa Rica

Abiotic and vegetation data were collected along two altitudinal transects through mature montane Quercus forests on the Pacific and Atlantic slopes of Costa Rica's Chirripó Massif. Between 2000 and 3200 m asl twenty-four 0.05 ha forest plots were selected at altitudinal intervals of 100 m, and eight soil profiles were described at intervals of 200 m. A TWINSPAN classification aided in the determination of eight zonal forest communities on the basis of their floristic composition. They are grouped in two sets of four: (i) the palm-rich lauraceous-fagaceous Lower Montane Mollinedia-Quercus Forests (2000–2600 m asl) and (ii) the bamboo-rich myrsinaceous-fagaceous Upper Montane Schefflera-Quercus Forests (2500–3200 m asl), respectively. Vegetation changes seem correlated with two major climatic gradients: (i) a temperature gradient (altitude), and (ii) a moisture gradient (wet Atlantic vs. moist Pacific slope). Most soils are Andepts, and residual, colluvial or derived from volcanic material. Humus layers are thicker on the wetter Atlantic slope. A total of 431 vascular plant species consisted of 86 pteridophytes, 1 gymnosperm, 296 dicots and 48 monocots. Species richness, canopy height and stem diameter decrease with increasing altitude, while the canopy surface becomes more flattend. A comparison with other studies shows that Chirripó's montane Quercus forests fit within the environmental ranges known from altitudinal zonations elsewhere in the Tropics.Abbreviations asl above sea level - dbh diameter at breast height - LM Lower Montane - Mt. Mountain - TWINSPAN two way indicator species analysis - UM Upper Montane - VU code referring to soil profiles as presented in Van Uffelen (1991) This paper is dedicated to the memory of Alwyn H. Gentry, an outstanding and inspiring tropical botanist who tragically died in a plane crash in the mountains of Ecuador on August 3 1993, when surveying possible boundaries for a new tropical cloud forest reserve.     

The Assemblage of Birds Foraging in Native West Indian Pine (Pinus occidentalis) Forests of the Dominican Republic during the Nonbreeding Season1

We studied avian resource use in a native West Indian pine (Pinus occidentalis) forest in the Cordillera Central of the Dominican Republic during the nonbreeding season. The forest is characterized by a fairly open pine canopy and a dense mixed-broadleaf and pine understory. We used a principal components analysis of 23 foraging characters for 23 bird species, including foraging height, the proportional use of 5 different foraging methods, 3 horizontal positions and foliage densities, and 11 foraging substrates. Five principal components accounted for 74 percent of the total variance of the assemblage variables and resulted in the delineation of at least 5 foraging guilds. 78 percent of bird species had mean foraging heights of 5.0–10.0 m, corresponding to the region of overlap of pine and broadleaf vegetation. As a result, the diversity of foraging substrates and maneuvers used by birds, rather than foraging height, appears to be the primary means by which birds that exploit this habitat separate ecologically. Migrant wood warblers (Parulinae) are probably able to integrate into the community because of little diet overlap between residents and migrants, and the fairly specialized nature of their largely insectivorous foraging habits. Avian foraging in this habitat may reflect the effects of frequent disturbance such as hurricanes.     

Structural comparison of tropical montane rain forests along latitudinal and altitudinal gradients in south and east Asia

Geographical patterns of altitudinal zonation, floristic composition, and structural features of tropical montane rain forests were examined along latitudinal gradients in south and east Asia. On equatorial mountains, the tropical montane rain forests occur above 1000 m. Toward middle latitudes, they come farther down and reach sea level at c. 35° N. Thus, the forests are equivalent to the subtropical rain forests of the latitudinal, horizontal zonation series. They exhibit gradual changes in floristic composition and structure along both altitudinal and latitudinal gradients. On equatorial mountains, they are divided into three types, i.e. tropical lower montane, upper montane, and subalpine forests. The three tree regeneration types, having emergent, sporadic and inverse-J type stem-diameter class frequency distributions, coexist in the lower montane forests, but the upper and subalpine forests display only the inverse-J type species with a few species of the sporadic type. Toward the northern latitudinal limit, the distinction between the three tropical montane forest zones in equatorial mountains becomes less clear. This can be explained by temperature conditions: on equatorial mountains, a temperature sum of 85° C months which controls the upper limit of the lower montane forests, and a coldest month mean temperature of-1° C which controls the evergreen broad-leaved trees, appear at c. 2500 and c. 4000 m respectively. The altitudinal range between 2500 m and 3800 m, which is the upper forest limit, is covered by upper montane and subalpine forests. On the other hand, at the latitudinal northern limit, the tropical upper montane and subalpine forests cannot exist because the above mentioned two temperature conditions occur at nearly the same point. Thus, at the northern latitudinal limit of the tropical montane forests, the three zones of equatorial mountains amalgamate into a single subtropical lowland forest community. This is due to the seasonal temperature climate in middle latitudes in, e.g., central Japan and central China.A part of this paper was presented as an oral presentation at the Vth International Congress of Ecology, Yokohama 23–30.8.1990.     

Foraging ecology of an assemblage of birds in lowland rainforest in northern Queensland

This paper presents an analysis of the foraging ecologies and hence means of partitioning resources amongst the twenty-eight most conspicuous species of birds in lowland tropical rainforest at Lacey's Creek, North Queensland. The height and site of foraging and the type of behaviour used by the species were compared. The height at which species foraged appeared to be more important for separating species than the site of foraging or the behaviour used. There appeared to be a greater proportion of species with generalized foraging behaviour at Lacey's Creek than in bird communities in the neotropics. Frugivorous species had very similar foraging ecologies but there were differences in the species of fruit eaten by some species. There were seasonal shifts in foraging by some species. It is postulated that the relative shortage of terrestrial species at Lacey's Creek compared with highland forest in North Queensland is due to the harsh dry season in the lowlands. The structure of this bird community appeared to be strongly affected by seasonality of climate.     

Functional diversity of epiphytes in two tropical lowland rainforests, French Guiana: using bryophyte life-forms to detect areas of high biodiversity

Recent studies have described a new tropical lowland forest type in the Guianas, the tropical lowland cloud forest. It is characterized by an enriched epiphytic species diversity particularly for bryophytes compared to common lowland rainforest, and is facilitated by frequent early morning fog events in valley locations. While the increase in epiphytic species diversity in lowland cloud forests has been documented, uncertainties remain as to (1) how this small scale variation in water supply is shaping the functional diversity of epiphytic components in lowland forests, and (2) whether information on functional group composition of epiphytes might aid in discerning these cloud forests from the common lowland rainforest. We compare the distribution of functional groups of epiphytes across height zones in lowland cloud forest and lowland rain forest of French Guiana in terms of biomass, cover as well as the composition of bryophyte life-forms. Both forests differed in functional composition of epiphytes in the canopy, in particular in the mid and outer canopy, with the cloud forest having a higher biomass and cover of bryophytes and vascular epiphytes as well as a richer bryophyte life-form composition. Bryophyte life-forms characteristic for cloud forests such as tail, weft and pendants were almost lacking in the canopies of common rain forest whereas they were frequent in lowland cloud forests. We suggest that ground-based evaluation of bryophyte life-form composition is a straightforward approach for identifying lowland cloud forest areas for conservation, which represent biodiversity hotspots in tropical lowland forests.     

The effect of altitude,patch size and disturbance on species richness and density of lianas in montane forest patches

The species richness and density of lianas (woody vines) in tropical forests is determined by various abiotic and biotic factors. Factors such as altitude, forest patch size and the degree of forest disturbance are known to exert strong influences on liana species richness and density. We investigated how liana species richness and density were concurrently influenced by altitude (1700–2360 m), forest patch size, forest patch location (edge or interior) and disturbance intensity in the tropical montane evergreen forests, of the Nilgiri and Palni hills, Western Ghats, southern India. All woody lianas (≥1 cm dbh) were enumerated in plots of 30 × 30 m in small, medium and large forest patches, which were located along an altitudinal gradient ranging from 1700 to 2360 m. A total of 1980 individual lianas were recorded, belonging to 45 species, 32 genera and 21 families, from a total sampling area of 13.86 ha (across 154 plots). Liana species richness and density decreased significantly with increasing altitude and increased with increasing forest patch size. Within forest patches, the proportion of forest edge or interior habitat influenced liana distribution and succession especially when compared across the patch size categories. Liana species richness and density also varied along the altitudinal gradient when examined using eco-physiological guilds (i.e. shade tolerance, dispersal mode and climbing mechanism). The species richness and density of lianas within these ecological guilds responded negatively to increasing altitude and positively to increasing patch size and additionally displayed differing sensitivities to forest disturbance. Importantly, the degree of forest disturbance significantly altered the relationship between liana species richness and density to increasing altitude and patches size, and as such is likely the primary influence on liana response to montane forest succession. Our findings suggest that managing forest disturbance in the examined montane forests would assist in conserving local liana diversity across the examined altitudinal range.     

“西双版纳热带山地雨林的植物多样性研究”的一些问题讨论

该文针对“西双版纳热带山地雨林的植物多样性研究”论文中存在的一些问题进行了讨论。原文所依据的6个调查样地, 从其分布海拔、生境、群落的生态外貌特征、植物区系组成及单位面积植物种数的统计上反映出它们并非都属于同样的植被类型,即原文所称的热带山地雨林,而可能分别属于《云南植被》中所应用的热带季节雨林的次生林群落(样地I和II)、季风常绿阔叶林群落(样地V和VI)及类似于苔藓常绿阔叶林(样地III和IV)的群落类型。由于这些样地代表了不同的植被类型, 导致在对这些样地的植物多样性特征的比较上出现较大差异。对原文在资料分析和与其它森林群落植物多样性的比较上存在的一些问题也作了讨论。原文的研究结果显示了西双版纳不同海拔高度上森林植物群落的物种多样性特征,但并不能全部运用于该地区的热带山地雨林这一特定植被类型。     

滇南勐宋热带山地雨林的物种多样性与生态学特征

 研究了鲜为人知的滇南勐宋地区的原始山地雨林植被, 根据分布生境、群落结构和种类组成特征,可将该山地雨林区分为沟谷和山坡两个类型, 分别定义为八蕊单室茱萸(Mastixia euonymoides)-大萼楠(Phoebe megacalyx)林和云南拟单性木兰(Parachmeria yunnanensis)-云南裸花(Gymnanthes remota)林。该山地雨林的外貌仍以单叶、革质、全缘、中叶为主的常绿中、小高位芽植物组成为特征,层间木质藤本植物仍较丰富,草本高位芽植物和附生植物丰富,但板根和茎花现象少见,属于热带山地垂直带上低山雨林或山地雨林植被类型。与该地区的典型热带季节雨林和赤道热带雨林相比, 勐宋的山地雨林群落中的大、中高位芽植物和藤本高位芽植物比例相对减少, 小、矮高位芽植物和草本高位芽植物比例相对增加,单叶、革质、非全缘叶和小叶比例相对增加,板根现象少见。与中国热带北缘-南亚热带地区（季风）常绿阔叶林比较, 勐宋的山地雨林有较多的附生植物和草本高位芽植物,相对较少的小高位芽植物和矮高位芽植物,小叶比例亦较少,非全缘叶和革质叶比例相对较低。故勐宋山地雨林是滇南热带北缘山地的一种较湿润生境的植被类型, 与所谓的季风常绿阔叶林不同。在物种多样性上,勐宋热带山地雨林在单位面积植物种数上并不比该地区的热带季节雨林低, 物种多样性指数与低丘季节雨林相当,比沟谷季节雨林低, 明显高于季风常绿阔叶林。     

ADAPTIVE ADVANTAGES OF MIXED-SPECIES FEEDING FLOCKS AMONG SEED-EATING FINCHES IN COSTA RICA

The feeding ecology of three Costa Rican finches occurring in mixed flocks, Tiaris olivacea, Sporophila aurita corvina and Sporophila torquella , was investigated by measuring both behavioural and ecological variables. Observations on foraging height, rate of hopping and pecking rate, as well as the identity, proximity and number of nearest neighbours were recorded. In addition the duration of all feeding and perching episodes were timed. Comparisons were also made between the abundances of food items (grass seeds) consumed and those potentially available for consumption.
The analysis of variance of the feeding behaviour revealed that the presence or absence of neighbouring birds, whether of the same or different species, influenced the duration of feeding bouts more significantly than did either differences in habitat or species-characteristic behaviour. In addition the dietary comparisons revealed overlap in both species and size of seed consumed. Such similarities suggest that these species are not partitioning fields in the classical sense.
We propose that the increase in the duration of the feeding bout associated with the presence of mixed species aggregations leads to increased feeding efficiency and is the result of intra- and inter-specific social learning. Certainly flocking is often advantageous, since searching in a group facilitates finding clumped resources; mixed species flocking, by increasing exposure to a diversity of foraging places and patterns, can further augment feeding efficiency.