Projected changes in elevational distribution and flight performance of montane Neotropical hummingbirds in response to climate change

The hovering flight of hummingbirds is one of the most energetically demanding forms of animal locomotion and is influenced by both atmospheric oxygen availability and air density. Montane Neotropical hummingbirds are expected to shift altitudinally upwards in response to climate change to track their ancestral climatic regime, which is predicted to influence their flight performance. In this study, we use the climate envelope approach to estimate upward elevational shifts for five Andean hummingbird species under two climate change scenarios. We then use field‐based data on hummingbird flight mechanics to estimate the resulting impact of climate change on aerodynamic performance in hovering flight. Our results show that in addition to significant habitat loss and fragmentation, projected upwards elevational shifts vary between 300 and 700 m, depending on climate change scenario and original mean elevation of the target species. Biomechanical analysis indicates that such upwards elevational shifts would yield a～2–5° increase in wing stroke amplitude with no substantial effect on wingbeat frequency. Overall, the physiological impact of elevational shifts of <1000 m in response to climate change is likely to be small relative to other factors such as habitat loss, changes in floristic composition, and increased interspecific competition.     

Effects of flight behaviour on body temperature and kinematics during inter-male mate competition in the solitary desert bee Centris pallida

Abstract. Body temperatures and kinematics are measured for male Centris pallida bees engaged in a variety of flight behaviours (hovering, patrolling, pursuit) at a nest aggregation site in the Sonoran Desert. The aim of the study is to test for evidence of thermoregulatory variation in convective heat loss and metabolic heat production and to assess the mechanisms of acceleration and forward flight in field conditions. Patrolling males have slightly (1–3 °C) cooler body temperatures than hoverers, despite similar wingbeat frequencies and larger body masses, suggesting that convective heat loss is likely to be greater during patrolling flight than during hovering. Comparisons of thorax and head temperature as a function of air temperature (T_a) indicate that C. pallida males are thermoregulating the head by increasing heat transfer from the thorax to the head at cool T_a. During patrolling flight and hovering, wingbeat frequency significantly decreases as T_a increases, indicating that variation in metabolic heat production contributes to thermal stability during these behaviours, as has been previously demonstrated for this species during flight in a metabolic chamber. However, wingbeat frequency during brief (1–2 s) pursuits is significantly higher than during other flight behaviours and independent of T_a. Unlike most other hovering insects, C. pallida males hover with extremely inclined stroke plane angles and nearly horizontal body angles, suggesting that its ability to vary flight speed depends on changes in wingbeat frequency and other kinematic mechanisms that are not yet described.     

Structure of the vortex wake in hovering Anna's hummingbirds (Calypte anna)

Hummingbirds are specialized hoverers for which the vortex wake has been described as a series of single vortex rings shed primarily during the downstroke. Recent findings in bats and birds, as well as in a recent study on Anna''s hummingbirds, suggest that each wing may shed a discrete vortex ring, yielding a bilaterally paired wake. Here, we describe the presence of two discrete rings in the wake of hovering Anna''s hummingbirds, and also infer force production through a wingbeat with contributions to weight support. Using flow visualization, we found separate vortices at the tip and root of each wing, with 15% stronger circulation at the wingtip than at the root during the downstroke. The upstroke wake is more complex, with near-continuous shedding of vorticity, and circulation of approximately equal magnitude at tip and root. Force estimates suggest that the downstroke contributes 66% of required weight support, whereas the upstroke generates 35%. We also identified a secondary vortex structure yielding 8–26% of weight support. Lift production in Anna''s hummingbirds is more evenly distributed between the stroke phases than previously estimated for Rufous hummingbirds, in accordance with the generally symmetric down- and upstrokes that characterize hovering in these birds.     

Leading edge vortex in a slow-flying passerine

Most hovering animals, such as insects and hummingbirds, enhance lift by producing leading edge vortices (LEVs) and by using both the downstroke and upstroke for lift production. By contrast, most hovering passerine birds primarily use the downstroke to generate lift. To compensate for the nearly inactive upstroke, weight support during the downstroke needs to be relatively higher in passerines when compared with, e.g. hummingbirds. Here we show, by capturing the airflow around the wing of a freely flying pied flycatcher, that passerines may use LEVs during the downstroke to increase lift. The LEV contributes up to 49 per cent to weight support, which is three times higher than in hummingbirds, suggesting that avian hoverers compensate for the nearly inactive upstroke by generating stronger LEVs. Contrary to other animals, the LEV strength in the flycatcher is lowest near the wing tip, instead of highest. This is correlated with a spanwise reduction of the wing's angle-of-attack, partly owing to upward bending of primary feathers. We suggest that this helps to delay bursting and shedding of the particularly strong LEV in passerines.     

Hovering and intermittent flight in birds

Two styles of bird locomotion, hovering and intermittent flight, have great potential to inform future development of autonomous flying vehicles. Hummingbirds are the smallest flying vertebrates, and they are the only birds that can sustain hovering. Their ability to hover is due to their small size, high wingbeat frequency, relatively large margin of mass-specific power available for flight and a suite of anatomical features that include proportionally massive major flight muscles (pectoralis and supracoracoideus) and wing anatomy that enables them to leave their wings extended yet turned over (supinated) during upstroke so that they can generate lift to support their weight. Hummingbirds generate three times more lift during downstroke compared with upstroke, with the disparity due to wing twist during upstroke. Much like insects, hummingbirds exploit unsteady mechanisms during hovering including delayed stall during wing translation that is manifest as a leading-edge vortex (LEV) on the wing and rotational circulation at the end of each half stroke. Intermittent flight is common in small- and medium-sized birds and consists of pauses during which the wings are flexed (bound) or extended (glide). Flap-bounding appears to be an energy-saving style when flying relatively fast, with the production of lift by the body and tail critical to this saving. Flap-gliding is thought to be less costly than continuous flapping during flight at most speeds. Some species are known to shift from flap-gliding at slow speeds to flap-bounding at fast speeds, but there is an upper size limit for the ability to bound (~0.3 kg) and small birds with rounded wings do not use intermittent glides.     

Hummingbirds fuel hovering flight with newly ingested sugar

We sought to characterize the ability of hummingbirds to fuel their energetically expensive hovering flight using dietary sugar by a combination of respirometry and stable carbon isotope techniques. Broadtailed hummingbirds (Selasphorus platycercus) were maintained on a diet containing beet sugar with an isotopic composition characteristic of C3 plants. Hummingbirds were fasted and then offered a solution containing cane sugar with an isotopic composition characteristic of C4 plants. By monitoring the rates of CO2 production and O2 consumption, as well as the stable carbon isotope composition of expired CO2, we were able to estimate the relative contributions of carbohydrate and fat, as well as the absolute rate at which dietary sucrose was oxidized during hovering. The combination of respirometry and carbon isotope analysis revealed that hummingbirds initially oxidized endogenous fat following a fast and then progressively oxidized proportionately more carbohydrates. The contribution from dietary sources increased with each feeding bout, and by 20 min after the first meal, dietary sugar supported approximately 74% of hovering metabolism. The ability of hummingbirds to satisfy the energetic requirements of hovering flight mainly with recently ingested sugar is unique among vertebrates. Our finding provides an example of evolutionary convergence in physiological and biochemical traits among unrelated nectar-feeding animals.     

The smallest avian genomes are found in hummingbirds

It has often been suggested that the genome sizes of birds are constrained relative to other tetrapods owing to the high metabolic demands of powered flight and the link between nuclear DNA content and red blood cell size. This hypothesis predicts that hummingbirds, which engage in energy-intensive hovering flight, will display especially constrained genomes even relative to other birds. We report genome size measurements for 37 species of hummingbirds that confirm this prediction. Our results suggest that genome size was reduced before the divergence of extant hummingbird lineages, and that only minimal additional reduction occurred during hummingbird diversification. Unlike in some other avian taxa, the small amount of variation observed within hummingbirds is not explained by variation in respiratory and flight-related parameters. Unexpectedly, genome size appears to have increased in four unrelated hummingbird species whose distributions are centred on humid forests of the upper-tropical elevational zone on the eastern slope of the Andes. This suggests that the secondary expansion of the genome may have been mediated by biogeographical and demographic effects.     

Wingbeat frequency, temperature and body size in bees and flies

ABSTRACT. This paper describes the relationship between ambient temperature and wingbeat frequency in bees and flies of different sizes, and presents new data from insects in free fight. The slope of the relationship changes with the size of the insect, and was different for insects in hovering flight compared with individuals of the same species in forward flight.     

The Effects of Moisture on Acetylene Reduction by Mats of Blue-green Algae in Sub-tropical Grassland

During the dry winter months desiccation is the main factorpreventing nitrogen fixation by mats of blue-green algae inkikuyu lawns. In the wetter summer months a few days of sunshinewithout rain will dry out the algal mats and depress nitrogenfixation unless the algae are protected by swards of grass.Dry mats resume nitrogen fixation within an hour of becomingdamp and attain their normal rate of fixation within 5 h. Maximumfixation rates occur at 100 per cent relative humidity and withsoil moisture contents of 22 to 42 per cent. Dry algal matsare unable to take up sufficient water as vapour (even in saturatingconditions) to permit resumption of acetylene reduction. Algal-dominatedsoils show greater water retention than adjacent bare soils.     

The Aerodynamics of Flapping Animal Flight

Our understanding of the aerodynamics of flapping animal flightis largely based on the quasi-steady assumption: the instantaneousaerodynamic forces on a flapping wing are assumed to be identicalwith those which the wing would experience in steady motionat the same instantaneous speed and angle of attack. Researchup to a decade ago showed that the assumption was sufficientto explain the flight of the vast majority of animals, but didnot rule out the possibility that alternative aerodynamic mechanismswere employed instead. Results are presented here for four hoveringanimals for which the quasi-steady explanation fails. Theseanimals apparently use lift mechanisms that rely on vorticesshed during the rotational motion of the wing at either endof the wingbeat. The postulated rotational lift mechanisms shouldalso apply to other hovering animals, even though the quasi-steadyassumption could explain their flight. Measurements of the wingforces produced by locusts cast doubt on the validity of thequasi-steady assumption for fast forward flight as well.     

Of hummingbirds and helicopters: hovering costs, competitive ability, and foraging strategies

Wing morphology and flight kinematics profoundly influence foraging costs and the overall behavioral ecology of hummingbirds. By analogy with helicopters, previous energetic studies have applied the momentum theory of aircraft propellers to estimate hovering costs from wing disc loading (WDL), a parameter incorporating wingspan (or length) and body mass. Variation in WDL has been used to elucidate differences either among hummingbird species in nectar-foraging strategies (e.g., territoriality, traplining) and dominance relations or among gender-age categories within species. We first demonstrate that WDL, as typically calculated, is an unreliable predictor of hovering (induced power) costs; predictive power is increased when calculations use wing length instead of wingspan and when actual wing stroke amplitudes are incorporated. We next evaluate the hypotheses that foraging strategy and competitive ability are functions of WDL, using our data in combination with those of published sources. Variation in hummingbird behavior cannot be easily classified using WDL and instead is correlated with a diversity of morphological and physiological traits. Evaluating selection pressures on hummingbird wings will require moving beyond wing and body mass measurements to include the assessment of the aerodynamic forces, power requirements, and power reserves of hovering, forward flight, and maneuvering. However, the WDL-helicopter dynamics model has been instrumental in calling attention to the importance of comparative wing morphology and related aerodynamics for understanding the behavioral ecology of hummingbirds.     

Sugar flux through the flight muscles of hovering vertebrate nectarivores: a review

In most vertebrates, uptake and oxidation of circulating sugars by locomotor muscles rises with increasing exercise intensity. However, uptake rate by muscle plateaus at moderate aerobic exercise intensities and intracellular fuels dominate at oxygen consumption rates of 50 % of maximum or more. Further, uptake and oxidation of circulating fructose by muscle is negligible. In contrast, hummingbirds and nectar bats are capable of fueling expensive hovering flight exclusively, or nearly completely, with dietary sugar. In addition, hummingbirds and nectar bats appear capable of fueling hovering flight completely with fructose. Three crucial steps are believed to be rate limiting to muscle uptake of circulating glucose or fructose in vertebrates: (1) delivery to muscle; (2) transport into muscle through glucose transporter proteins (GLUTs); and (3) phosphorylation of glucose by hexokinase (HK) within the muscle. In this review, we summarize what is known about the functional upregulation of exogenous sugar flux at each of these steps in hummingbirds and nectar bats. High cardiac output, capillary density, and blood sugar levels in hummingbirds and bats enhance sugar delivery to muscles (step 1). Hummingbird and nectar bat flight muscle fibers have relatively small cross-sectional areas and thus relatively high surface areas across which transport can occur (step 2). Maximum HK activities in each species are enough for carbohydrate flux through glycolysis to satisfy 100 % of hovering oxidative demand (step 3). However, qualitative patterns of GLUT expression in the muscle (step 2) raise more questions than they answer regarding sugar transport in hummingbirds and suggest major differences in the regulation of sugar flux compared to nectar bats. Behavioral and physiological similarities among hummingbirds, nectar bats, and other vertebrates suggest enhanced capacities for exogenous fuel use during exercise may be more wide spread than previously appreciated. Further, how the capacity for uptake and phosphorylation of circulating fructose is enhanced remains a tantalizing unknown.     

Surpassing Mt. Everest: extreme flight performance of alpine bumble-bees

Animal flight at altitude involves substantial aerodynamic and physiological challenges. Hovering at high elevations is particularly demanding from the dual perspectives of lift and power output; nevertheless, some volant insects reside and fly at elevations in excess of 4000 m. Here, we demonstrate that alpine bumble-bees possess substantial aerodynamic reserves, and can sustain hovering flight under hypobaria at effective elevations in excess of 9000 m, i.e. higher than Mt. Everest. Modulation of stroke amplitude and not wingbeat frequency is the primary means of compensation for overcoming the aerodynamic challenge. The presence of such excess capacity in a high-altitude bumble-bee is surprising and suggests intermittent behavioural demands for extreme flight performance supplemental to routine foraging.     

Methodological procedures for detecting antibiotic producers among cultures of the family Micromonosporaceae]

Data on intensification of the search for active cultures among Micromonospora are presented. It was shown that the frequency of detecting the antibiotic-producing cultures among Micromonospora under conditions of fermentation on the corn-glucose medium inoculated with agar blocks amounted to 35 per cent. The use of nutrient media of different composition for growing submerged inoculum of Micromonospora demonstrated that the rate of its growth reached maximum on the peastarch medium. The use of this medium for growing submerged seed material for fermentation in the corn-glucose medium increased the frequency of detecting active cultures from 35 to 43.1 per cent. The assay of Micromonospora antibiotic activity twice, i.e. in 96 and 240 hours of the fermentation process increased the frequency of detecting active cultures up to 57.1 per cent and revealing greater variety of antibiotics. Fermentation of Micromonospora cultures simultaneously on 6 different nutrient media inoculated with submerged seed mycelium and assay of the activity for 2 times, i. e. in 96 and 240 hours allowed a detection of up to 76.2 per cent of active strains out of the total number of the isolates.     

Why hummingbirds hover and honeyeaters perch

Evidence is presented in support of the suggestion that a hovering bird is able to move between flowers more quickly than one that is perching. This advantage to hovering may be offset, however, by the higher energetic costs of hovering as compared with perching. This trade-off is evaluated in two field situations, one for perching honeyeaters and the other for hovering hummingbirds. In each case it is estimated that the birds employ the foraging mode (hovering versus perching) that results in the greatest net rate of energy gain.     

Resource value affects territorial defense by Broad-tailed and Rufous hummingbirds

ABSTRACT.   Territorial behavior of Broad-tailed ( Selasphorous platycercus ) and Rufous ( Selasphorous rufus ) hummingbirds in Colorado was measured at sites with feeders containing10%, 20%, and 30% sucrose solutions, respectively. The presence or absence of territory holders, number of intruders, and intensity of defense were measured at the three levels of energy availability. Migrating Rufous Hummingbirds displaced Broad-tailed Hummingbirds from territories they had defended during the breeding season; Broad-tailed Hummingbirds then defended only lower quality sites. Both Broad-tailed and Rufous hummingbirds employed more energetically expensive behaviors when defending high quality sites, with longer chases more often supplemented with chip calls and hovering. Other investigators have suggested that chip calls and hovering are precursors to a chase. However, I found that chasing was the default response to the presence of an intruder. Chip calls and hovering were added to intensify a chase. In the few cases where chip calls were uttered or hovering occurred without a chase, Rufous Hummingbirds were more likely to exhibit this behavior than Broad-tailed Hummingbirds.     

喀斯特弃耕地草本植物根茎叶经济型谱间协调及其对极端降雨的响应

植物功能性状之间的关系是反映植物在资源获取和资源保守策略之间权衡的有力指标。喀斯特是全球最脆弱的陆地生态系统之一,其中的植被长期遭受干旱的胁迫,对极端天气和极端降水高度敏感。在降雨极端化的背景下,喀斯特地区植物生态策略的变化还不得而知。以西南喀斯特弃耕地演替初期群落中12种草本植物为对象,通过保持年降水量不变,人为控制单次降水量以及降水频率,以自然降雨为对照组(CK),设置中雨频率增强处理组(T10),大雨频率增强处理组(T25)和暴雨频率增强处理组(T50)3个处理组。模拟在全球变化导致的降雨极端化情境下,研究植物根茎叶功能性状的响应特征及经济型谱之间的关系。主要结果如下:(1)根茎叶的功能性状对降雨极端化的响应不同,其中根的理论导水率随着降雨极端化增强而增加,而茎和叶的理论导水率仅在T25增加,根茎叶三种器官中对降雨极端化最敏感的是茎;(2)根茎叶各器官均有自身稳定的"快-慢"经济型谱。降雨极端化仅T50的植物会选择介于资源获取和资源保守策略之间稳定的经济型谱;(3)不同植物器官之间的经济型谱的协调并不一致,其中茎根经济型谱之间的相关系数在三个处理组中均变大,而叶根经济型谱之间的相关系数仅在T25变大,叶茎经济型谱之间的相关系数仅在T10变大。总之,喀斯特弃耕地演替初期群落中草本植物的根茎叶有自身稳定的经济型谱,降雨极端化导致了以根茎叶经济型谱所反映的植物地上和地下资源保存和获取策略的解耦;本研究中的植物最容易发生功能性状变异的器官是茎。本研究提高了对植物不同器官各功能性状之间关系和植物地上与地下部分关联性的理解,揭示了植物不同器官的"快-慢"植物经济型谱以及由此反映的植物适应能力的差异和功能策略的多样性,促进了对植物资源获取与环境响应策略的深入认识,为预测未来气候变化背景下喀斯特弃耕地草本植物的适应趋势和响应策略提供了新见解。     

Respiratory evaporative water loss during hovering and forward flight in hummingbirds

Hummingbirds represent an end point for small body size and water flux in vertebrates. We explored the role evaporative water loss (EWL) plays in management of their large water pool and its use in dissipating metabolic heat. We measured respiratory evaporative water loss (REWL) in hovering hummingbirds in the field (6 species) and over a range of speeds in a wind tunnel (1 species) using an open-circuit mask respirometry system. Hovering REWL during the active period was positively correlated with operative temperature (T_e) likely due to some combination of an increase in the vapor-pressure deficit, increase in lung ventilation rate, and reduced importance of dry heat transfer at higher T_e. In rufous hummingbirds (Selasphorus rufus; 3.3 g) REWL during forward flight at 6 and 10 m/s was less than half the value for hovering. The proportion of total dissipated heat (TDH) accounted for by REWL during hovering at T_e > 40 °C was < 40% in most species. During forward flight in S. rufus the proportion of TDH accounted for by REWL was ~ 35% less than for hovering. REWL in hummingbirds is a relatively small component of the water budget compared with other bird species (< 20%) so cutaneous evaporative water loss and dry heat transfer must contribute significantly to thermal balance in hummingbirds.     

Assessment of volatilization loss of ammonia from surface-applied urea on forest soil by N15 recovery

Summary Using vertically isolated micro-plots the isotopic recovery technique was tested for assessing the extent of ammonia volatilization loss from N¹⁵-labelled urea applied on the forest floor (Pinus silvestris L.). The size of the ammonia loss was obtained as a difference between the amounts of labelled urea N added and the amounts of labelled N recovered in the soil profile after 13, 31, and 39 days's exposure, respectively. Urea materials of two different pellet size were used: granulated small pellets (280 pellets per gram) and tablets (2.06 g each). The nitrogen application rate was 200 kg N per hectare. The recovery data for 13 days' exposure indicated a volatilization loss, which for the small pellet urea was 24.9 per cent and for the tabletted urea 12.1 per cent. The corresponding figures for the 31 days' exposure, during which the total amount of precipitation was 14 mm, were 15.1 and 26.9 per cent, respectively. The pattern of labelled N distribution in the soil profile examined showed that during the period of exposure in question a leaching loss of labelled N was rather unlikely. It was demonstrated, furthermore, that nitrogen from the tabletted urea had diffused to a greater depth of the soil than that from the small-pellet urea. Nitrogen from the small-pellet urea was to a large extent recovered in the litter layer. On exposure to heavy rain the tabletted urea was subjected to the highest leaching loss. An addition of 10 per cent (w/w) of metaphosphoric acid or sublimed sulphur to the tabletted urea did not result in any further reduction of the volatilization loss. The merits and limits of the isotopic recovery technique are discussed.     

Comparative energetics of the giant hummingbird (Patagona gigas)

Hummingbirds (family Trochilidae) represent an extreme outcome in vertebrate physiological design and are the only birds capable of sustained hovering. The giant hummingbird (Patagona gigas) is the largest trochilid, with a mass of ~20 g, and is found over an altitudinal range from 0 to 4,500 m above sea level. We report here measurements of daily, basal, and hovering rates of oxygen consumption in the giant hummingbird; compare these values with data from smaller hummingbirds; and assess overall metabolic and allometric limits to trochilid body size. The sustained metabolic scope (i.e., the ratio of daily energy expenditure to basal metabolic rate) in the giant hummingbird is higher than that in smaller hummingbirds but lies below a proposed theoretical maximum value for endotherms. Scaling exponents in the allometric relationships for different modes of energetic expenditure were comparable, suggesting that the giant hummingbird, although a clear outlier in terms of body size, does not obviously deviate from metabolic relationships derived from other trochilid taxa.