Body temperatures of Namib Desert tenebrionid beetles: their relationship in laboratory and field

Abstract The body temperatures of six apterous species of Namib Desert tenebrionid beetles were measured continuously with indwelling thermocouples under laboratory conditions and in the field. The range of body temperatures selected was within the upper half of their 'tolerated range', which we defined as the temperatures lying between measured critical thermal maximum and critical thermal minimum. In the field, individuals also maintained their body temperatures within the upper half of the 'tolerated range'. These beetles maintained higher body temperatures than those recorded for any other ectothermic insect. Three of the six species maintained lower body temperatures in the field than they selected in the laboratory. The other three species showed no significant difference between field and laboratory body temperatures. We conclude that these beetles are not forced by biotic or abiotic factors to adopt thermal niches which present them with physiological difficulties.     

小尺度下柠条林破碎化生境对地表甲虫多样性的影响

一般认为,景观斑块面积和破碎化对物种丰富度和分布格局有重要的影响。在宁夏中部荒漠地区,天然柠条林和人工柠条林地交错排列,形成点、片、带状等大小不等的斑块性分布,表现为典型的破碎化斑块格局生境特征。本文采用巴氏罐诱法调查了在小尺度下荒漠景观人工柠条林破碎化生境不同斑块内地表甲虫的物种多样性。结果共获得10科20属29种地表甲虫,其中拟步甲科昆虫占绝对优势,阿小鳖甲Microdera kraatzi alashanica Skopin、克小鳖甲Microdera kraatzi kraatzi(Reitter)为优势种。Rarefaction曲线显示较大面积的斑块有较多的物种多样性,但群落多样性指数各斑间块差异不显著。利用斑块面积对物种数-个体数进行回归分析表明,地表甲虫的物种多样性受斑块面积的影响,生境破碎化会导致地表甲虫多样性下降。     

Alternative forms of competition and predation dramatically affect habitat selection under foraging--predation-risk trade-offs

Habitat selection under foraging—predation-risk trade-offshas been a frequent topic of interest to theoretical behavioraland evolutionary ecologists. However, most habitat selectionmodels assume that individuals compete exploitatively for resourcesand that predation is either density independent or dilutedcompletely by competitor number, despite empirical evidencethat other forms of competition and predation also occur innature. I developed an individual-based model for studyingthe effects of alternative forms of competition and predationon the process of habitat selection under foraging—predation-risktrade-offs. To make the model more relevant to natural populations,I assumed that individuals vary continuously in traits relatedto competitive ability and vulnerability to predation and allowed resources and predators to be distributed across more than twohabitats. The results of my investigation demonstrate thatthe predicted pattern of habitat selection can be affecteddramatically by the form predation is assumed to take. Whenpredation is density dependent or frequency dependent, individuals will tend to be distributed across habitats according to theirabsolute vulnerability to predation. In contrast, when predationis density dependent or vulnerability dependent, individualswill tend to segregate by competitive ability. Whether oneassumes that individuals compete for resources via exploitationor interference also influences the predicted pattern of habitat selection. In general, interference competition results in amore even distribution of competitors across habitats.     

Density-dependent habitat selection in plants

Pea plants exhibit density-dependent habitat selection as they grow. We split the root of a young pea (Pisum sativum L.) so that half grew in one pot and half in an adjacent pot. The rest of the plant remained intact. This is a ‘fence-sitter plant’. Each root-half was exposed either to no competition in its pot or to competitor plants sharing its pot. There were one, two, three or five competitor plants. The total root biomass and the fitness (= dry weight of fruit) of the fence-sitter decreased only slightly and insignificantly in response to increased density of the competitor plants. The fitness of the competing plants decreased with density. The fence-sitter shifted its root system from the pot with competition to that free of competition in proportion to the number of competitors. The fence-sitter apparently invested in each of its two roots so that the ratio between the roots was similar to the ratio between the resources in the pots. This result is analogous to the habitat-matching rule of the ideal free distribution of populations (Fretwell, 1972). We suggest that plants invest in each of their roots until the uptake rate per unit root biomass is equal for all roots. This revised version was published online in July 2006 with corrections to the Cover Date.     

贵州草海越冬黑颈鹤觅食栖息地选择的初步研究

本文报道了利用Friedman非参数统计方法,研究越冬黑颈鹤(Grus nigricollis)在贵州草海对其觅食栖息地选择的结果。在草海的黑颈鹤越冬觅食栖息地可分为5种——莎草草甸、浅水沼泽、草地、玉米地和蔬菜地。黑颈鹤对莎草草甸的选择性最高,对玉米地的选择性最差,对其余类型栖息地的选择随地点的不同而不同。草海的人为活动很多,是影响黑颈鹤利用和选择栖息地的一个重要因素。本文也对改善黑颈鹤越冬栖息地的管理和保护提出了一些建议。     

Measuring the benefit of habitat selection

We used a behavioral bioassay to estimate the advantages thattwo species of gerbils (Gerbillus allenbyi and G. pyramidum)experienced by preferring a semistabilized dune habitat overa stabilized sand habitat. We used the magnitude of foragingeffort by the gerbils to signal the difference between thetwo habitats. When they were foraging as much in stabilizedsand as in semistabilized dune, we inferred that these habitatswere providing equivalent rewards. We performed a series ofexperiments in two 1-ha field enclosures, each containing similarproportions of stabilized sand and semistabilized dune. Eachenclosure contained a population of only one of the species.By varying the amount of seeds added (either 0.5, 1, 2, or 3g of seeds in 18 seed trays) to each habitat and monitoringthe behavior of the gerbils, we were able to fit a curve thatreflected the change in habitat preference as a function ofseed addition rate. We were also able to show how much seedaddition had to be added to bring the two habitats into equaluse. Each species required only 13 g/ha/night to entirely offsetthe advantage of the semistabilized dune.     

Effects of habitat size and isolation on species immigration–extinction dynamics and community nestedness in a desert river system

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Maintenance of two genetic entities by habitat selection

Summary In the laboratory, the two species of copepodsLepeophtheirus thompsoni andLepeophtheirus europaensis, ectoparasites of flatfishes, can meet and mate on at least one host species. In the wild however, these two species are found isolated on their sympatric hosts. Habitat selection theoretically represents a powerful enough mechanism to explain the maintenance of genetic heterogeneity in the wide sense. In this paper, the host colonization process is studied for both parasite species. It is shown that each parasite can develop and reach adult age on each host species. However,L. thompsoni is highly selective; it almost totally refuses to colonize hosts other than its natural one.Lepeophtheirus europaensis, on the contrary, readily infests turbot and brill in single-host experiments, but strongly prefers the brill when it has a choice. It appears that these two genetic entities are sympatrically maintained due to strong habitat selection. Such a pattern could theoretically only occur in a soft-selection context (density dependence). This point is discussed with respect to the different patterns in host use found in the geographical distribution of these parasites.     

荒漠景观拟步甲科昆虫多样性及其对生境的指示作用

本研究在宁夏灵武白芨滩国家级自然保护区北部荒漠景观下的磁窑堡地区,选取典型荒漠草原区(DS)、人工柠条林种植区(CCL1,CCL2)及流沙人工封育区(SDE1,SDE2,SDE3)3类生境共6块样地,于2010年4月至8月以巴氏罐诱法采集该地区拟步甲科昆虫,研究该地区拟步甲科昆虫多样性及其对生境的指示作用。结果共获得标本3406头,计12属22种,其中东鳖甲属Anatolica和小鳖甲属Microdera为优势属,波氏东鳖甲Anatolica potanini Reitter、阿小鳖甲Microdera kraatz alashanica Skopin、小丽东鳖甲Anatolica amoenula Reitter和尖尾东鳖甲Anatolic amucronata Reitter为优势种。在流沙人工封育区的SDE1样地物种丰富度指数、均匀度指数和多样性指数最高,SDE3样地个体数量和优势度指数最高;人工柠条林种植区的CCL1样地个体数量最低。聚类分析结果表明流沙人工封育区与荒漠草原区、人工柠条种植区在物种组成及生境相似性方面差异较大。利用指示值法确定不同生境的特征指示种类,结果显示波氏东鳖甲、谢氏宽漠王Mantichorula semenowiReitter、尖尾东鳖甲可作为流沙封育区特征指示种;突角漠甲Trigonocnera pseudopimelia(Reitter)、中华砚甲Cyphogenia chinensis(Faldermann)和纳氏东鳖甲AnatolicanuretiSchusteretReymond可作为荒漠草原特征指示种;蒙古漠王Platyope mongolica Faldermann、多毛宽漠甲Sternoplax setosa setosa(Bates)、克小鳖甲Microdera kraatzi(Reitter)是生境恢复的监测种;小丽东鳖甲Anatolica amoenula Reitter等种类可作为生境退化的监测种。     

Availability of water controls Crassulacean acid metabolism in succulents of the Richtersveld (Namib desert,South Africa)

Features of Crassulacean acid metabolism (CAM) were studied in a variety of different succulents in response to climatic conditions between March 1977 and October 1983 in the southern Namib desert (Richtersveld). A screening in 1977 and 1978 revealed that nearly all investigated succulents performed a CAM, but overnight accumulation of malate declined gradually with decreasing soil water potential, tissue osmotic potential, and leaf water content. This was further substantiated by an extended period of insufficient rainfall in 1979 and 1980 which damaged the evergreen CAM succulents between 80 and 100%. In most of the species still living, neither CO₂-gas exchange nor diurnal acid fluctuation, indicative of CAM, could be detected unless an abundant rainfall restored both CAM features. Plants persisted in a stage of latent life.Water supply is one necessary prerequisite for CAM in the Richtersveld. But even well-watered plants with CAM were sensitive to short-term water stress caused by high water-vapour partialpressure deficit (VPD) in the night, which reduced or prevented CO₂ uptake and resulted in a linear relation between overnight accumulated malate and VPD. The results do not support the opinion that, for the Namib succulents, CAM is an adaptive mechanism to water stress since long-term and short-term water stress stopped nocturnal malate synthesis, but instead lead to the conclusion that nocuturnal CO₂ fixation is only performed when the water status of the plant can be improved simultaneously.Abbreviations CAM Crassulacean acid metabolism - VPD water vapour pressure deficit Dedicated to Professor H. Ziegler on the occasion of his 60th birthday     

Irreversible habitat specialization does not constrain diversification in hypersaline water beetles

Specialization to extreme environments is often considered an evolutionary dead‐end, leading to irreversible adaptations and reduced evolvability. There is, however, mixed evidence for this macroevolutionary pattern, and limited data from speciose lineages. Here, we tested the effect of habitat specialization to hypersaline waters in the diversification rates of aquatic beetles of the genus Ochthebius (Coleoptera, Hydraenidae), using a molecular phylogeny with more than 50% of the 546 recognized species, including representatives of all but one of the nine recognized subgenera and 17 species groups. Phylogenies were built combining mitochondrial and nuclear genes, with the addition of 42 mitochondrial genomes. Using Bayesian methods of character reconstruction, we show that hypersaline tolerance is an irreversible ecological specialization that arose multiple times. Two lineages of Ochthebius experienced a significant increase in diversification rates, one of them inhabiting hypersaline waters, but there was no overall correlation with habitat or any significant decrease in diversification rates despite the irreversibility of hypersaline tolerance. Our study tested for the first time the impact of hypersaline habitat specialization on diversification rates, finding no support for it to be an evolutionary dead‐end. On the contrary, multiple and ancient lineages fully adapted to these extreme osmotic conditions have persisted and diversified over a long evolutionary timescale.     

The importance of spatial scale in habitat selection by European beaver

We evaluated habitat selection by European beaver Castor fiber L. across a spatial gradient from local (within the family territory) to a broad, ecoregional scale. Based on aerial photography, we assessed the habitat composition of 150 beaver territories along the main water bodies of the Vistula River delta (northern Poland) and compared these data with 183 randomly selected sites not occupied by the species. The beavers preferred habitats with high availability of woody plants, including shrubs, and avoided anthropogenically modified habitats, such as arable lands. Within a single family territory, we observed decreasing woody plant cover with increasing distance from a colony centre, which suggests that beaver habitat preferences depend on the assessment of both the abundance and spatial distribution of preferred habitat elements. We tested the importance of spatial scale in beaver habitat selection with principal coordinates of neighbour matrices analysis, which showed that the geographical scale explained 46.7% of the variation in habitat composition, while the local beaver density explained only 10.3% of this variability. We found two main spatial gradients that were related to the broad spatial scale: first, the most important gradient was related to the largest distances between beaver sites and was independent of woody plant cover and the local beaver site density. The second most important gradient appeared more locally and was associated with these variables. Our results indicate that European beaver habitat selection was affected by different scale‐related phenomena related 1) to central place foraging behaviour, which resulted in the clumped distribution of woody plants within the territory, and 2) local population density and woody plant cover. Finally, 3) habitat selection occurs independently across the largest spatial scale studied (e.g. between watersheds), which was probably due to the limited natal dispersal range of the animals.     

Comparative phylogeography of tenebrionid beetles in the Aegean archipelago: the effect of dispersal ability and habitat preference

Comparative phylogeographical studies in island archipelagos can reveal lineage-specific differential responses to the geological and climatic history. We analysed patterns of genetic diversity in six codistributed lineages of darkling beetles (Tenebrionidae) in the central Aegean archipelago which differ in wing development and habitat preferences. A total of 600 specimens from 30 islands and eight adjacent mainland regions were sequenced for mitochondrial cytochrome oxidase I and nuclear Muscular protein 20. Individual gene genealogies were assessed for the presence of groups that obey an independent coalescent process using a mixed Yule coalescent model. The six focal taxa differed greatly in the number of coalescent groups and depth of lineage subdivision, which was closely mirrored by the degree of geographical structuring. The most severe subdivision at both mitochondrial DNA and nuclear DNA level was found in flightless lineages associated with presumed stable compact-soil habitats (phrygana, maquis), in contrast to sand-obligate lineages inhabiting ephemeral coastal areas that displayed greater homogeneity across the archipelago. A winged lineage, although associated with stable habitats, showed no significant phylogenetic or geographical structuring. Patterns of nucleotide diversity and local genetic differentiation, as measured using Φ_ST and hierarchical amova , were consistent with high levels of ongoing gene flow in the winged taxon; frequent local extinction and island recolonisation for flightless sand-obligate taxa; and very low gene flow and geographical structure largely defined by the palaeogeographical history of the region in flightless compact-soil taxa. These results show that differences in dispersal rate, mediated by habitat persistence, greatly influence the levels of phylogeographical subdivision in lineages that are otherwise subjected to the same geological events and palaeoclimatic changes.     

豚草发生地土壤昆虫群落结构及动态

采用分层取样法、大类群分类法和数理统计方法,研究了普通豚草Ambrosia artemisiifolia和三裂叶豚草A．trifida发生地土壤昆虫的群落结构及动态。共采集土壤昆虫46个科（成虫与幼虫分开单列）,隶属于12个目,其中弹尾目个体数占绝对优势（占土壤昆虫总个体数的80.879%）,其次是膜翅目（8.765%）、鞘翅目幼虫（3.951%）和双翅目虫（3.093%）,其余合计占3.312%。在46个科中,优势类群有3科：棘跳虫科（Onychiuridae）（占土壤昆虫总个体数的32.756%）、鳞跳虫科（Tomoceridae）（18.869%）和节跳虫科（Isotomidae）（18.178%）。土壤昆虫个体数和类群数,随着豚草生长季的推进而增加,8月达到最高值。豚草降低了土壤昆虫的个体数量和群落多样性,根据逐月方差分析和多重比较,8月和9月对照样地土壤昆虫个体数量和多样性指数均显著高于豚草样地。豚草表现出对某些土壤昆虫类群的驱避作用,步甲科（Carabidae）成虫、拟花萤科（Dasytidae）幼虫、叶蝉科（Jassidae）、阎甲科（Histeridae）和管蓟马科（Phloeothripidae）在豚草整个生长季从未在豚草样地中出现。豚草对于土壤昆虫的抑制作用在豚草纯群落和植物生长旺季更为明显。土壤昆虫的垂直分布具有明显的表聚性。     

Water balance and osmoregulation in a free-ranging tenebrionid beetle, Onymacris unguicularis, of the Namib Desert

Onymacris unguicularis, a fog-basking tenebrionid beetle of the Namib Desert, has mean water influx rates of 49.9 mg H₂O/g.d and mean efflux rates of 41.3 mg H₂O/g.d with mean mass gain being 10.7 mg/g.d. If only steady-state conditions are considered (no mass change), and passive vapour input subtracted, drinking accounts for 50% of water input. Active beetles must drink in order to maintain water balance, while inactive beetles can maintain water balance either eating seeds or by simply metabolizing fat. Little change is observed in ratios of haemolymph and total body water to dry mass when fogs occur, while significant changes in haemolymph osmotic pressure are associated with fog occurrence.For short periods, O. unguicularis can tolerate dehydration with only slight changes in the ratio of total body water and haemolymph to dry body mass and to haemolymph osmotic pressure. For longer periods however, active beetles must have access to fog water for water balance maintenance. This is probably necessary for reproduction.     

Adaptation and habitat selection in the eco-evolutionary process

The struggle for existence occurs through the vital rates of population growth. This basic fact demonstrates the tight connection between ecology and evolution that defines the emerging field of eco-evolutionary dynamics. An effective synthesis of the interdependencies between ecology and evolution is grounded in six principles. The mechanics of evolution specifies the origin and rules governing traits and evolutionary strategies. Traits and evolutionary strategies achieve their selective value through their functional relationships with fitness. Function depends on the underlying structure of variation and the temporal, spatial and organizational scales of evolution. An understanding of how changes in traits and strategies occur requires conjoining ecological and evolutionary dynamics. Adaptation merges these five pillars to achieve a comprehensive understanding of ecological and evolutionary change. I demonstrate the value of this world-view with reference to the theory and practice of habitat selection. The theory allows us to assess evolutionarily stable strategies and states of habitat selection, and to draw the adaptive landscapes for habitat-selecting species. The landscapes can then be used to forecast future evolution under a variety of climate change and other scenarios.     

Soil fungal diversity and assembly along a xeric stress gradient in the central Namib Desert

The Namib Desert of south-western Africa is one of the oldest deserts in the world and possesses unique geographical, biological and climatic features. While research through the last decade has generated a comprehensive survey of the prokaryotic communities in Namib Desert soils, little is yet known about the diversity and function of edaphic fungal communities, and even less of their responses to aridity. In this study, we have characterized soil fungal community diversity across the longitudinal xeric gradient across the Namib desert (for convenience, divided into the western fog zone, the central low-rainfall zone and the eastern high-rainfall zone), using internal transcribed sequence (ITS) metabarcoding. Ascomycota, Basidiomycota and Chytridiomycota consistently dominated the Namib Desert edaphic fungal communities and a core mycobiome composed of only 15 taxa, dominated by members of the class Dothideomycetes (Ascomycota), was identified. However, fungal community structures were significantly different in the fog, low-rainfall and high-rainfall zones. Furthermore, Namib Desert gravel plain fungal community assembly was driven by both deterministic and stochastic processes; the latter dominating in the all three xeric zones. We also present data that suggest that the inland limit of fog penetration represents an ecological barrier to fungal dispersal across the Namib Desert.     

Two gerbils of the Negev: A long-term investigation of optimal habitat selection and its consequences

Optimal foraging theory has entered a new phase. It is not so much tested as used. It helps behavioural ecologists discover the nature of the information in an animals brain. It helps population ecologists reveal coefficients of interaction and their patterns of density-dependent variation. And it helps community ecologists examine niche relationships. In our studies on two species of Negev desert gerbil, we have taken advantage of the second and third of these functions. Both these gerbils prefer semi-stabilized dune habitat, and both altered their selective use of this habitat and stabilized sand according to experimental changes we made in their populations. Their changes in selectivity agree with a type of optimal foraging theory called isoleg theory. Isoleg theories provide examples of dipswitch theories – bundles of articulated qualitative predictions – that are easier to falsify than single qualitative predictions. By linking behaviour to population dynamics through isoleg theory, we were able to use the behaviour of the gerbils to reveal the shapes of their competitive isoclines. These have the peculiar non-linear shapes predicted by optimal foraging theory. Finally, when owl predation threatens, the behaviour of Gerbillus allenbyi reveals the shape of their victim isocline. As has long been predicted by predation theory and laboratory experiments, it is unimodal.