Fundamental avian energetics: 2. The ability of birds to change heat loss and explanation of the mass exponent for basal metabolism in homeothermic animals

The maximum ability of birds to generate heat due to increasing metabolism, as a result of both activity and heat stress, was determined in relation to the evaporative and nonevaporative heat losses at various temperatures in passerines and nonpasserines at the beginning and at the end of thermoneutral zones. The minimum (h _min) and maximum (h _max) nonevaporative thermal conductances in both species change similarly depending on the body mass, and the slopes of regression lines in h _min and h _max are identical. At the same time, h _max is approximately four times higher than h _min. Experimental data obtained both in this study and by other authors show that the ratio h _max/h _min = 4 is constant for all homeothermic animals and appears to be a sensible compromise found by the evolution between an increase in activity and the minimum effectiveness profitable for life of the transfer of metabolic power into mechanical power (?? = 1/4) during its fulfillment. An increase in the ratio h _max/h _min, although it allows an animal to augment its daily activity, leads to a reduction in the effectiveness and is, therefore, not used by homeothermic animals. The abilities of birds and mammals to change their heat loss are determined by the ratio h _max/h _min = 4, which is an integrated indicator of the level of development of blood circulation and respiration systems and the degree of development of external covers, as well as the ability of both to change heat loss. In homeothermic animals, this ratio is associated with the body mass exponent in allometric dependences for basal metabolism and determines the efficiency of transfer of metabolic power into mechanical work.     

Heart mass and the maximum cardiac output of birds and mammals: implications for estimating the maximum aerobic power input of flying animals

Empirical studies of cardiovascular variables suggest that relative heart muscle mass (relative M_h) is a good indicator of the degree of adaptive specialization for prolonged locomotor activities, for both birds and mammals. Reasonable predictions for the maximum oxygen consumption of birds during flight can be obtained by assuming that avian heart muscle has the same maximum physiological and biomechanical performance as that of terrestrial mammals. Thus, data on M_h can be used to provide quantitative estimates for the maximum aerobic power input (aerobic P_i,max) available to animals during intense levels of locomotor activity. The maximum cardiac output of birds and mammals is calculated to scale with respect to M_h (g) as 213 M_h^0.88+-0.04 (ml min^-1), while aerobic P_i,max is estimated to scale approximately as 11 M_h^0.88+-0.09 (W). In general, estimated inter-species aerobic P_i,max, based on M_h for all bird species (excluding hummingbirds), is calculated to scale with respect to body mass (M_b in kg) as 81 M_b^0.82+-0.11 (W). Comparison of family means for M_h indicate that there is considerable diversity in aerobic capacity among birds and mammals, for example, among the medium to large species of birds the Tinamidae have the smallest relative M_h (0.25 per cent) while the Otidae have unusually large relative M_h (1.6 per cent). Hummingbirds have extremely large relative M_h (2.28 per cent), but exhibit significant sexual dimorphism in their scaling of M_h and flight muscle mass, so that when considering hummingbird flight performance it may be useful to control for sexual differences in morphology. The estimated scaling of aerobic P_i,max (based on M_h and M_b in g) for male and female hummingbirds is 0.51 M_b^{0.83 +/-0.07} and 0.44 M_b^{0.85+- 0.11} (W), respectively. Locomotory muscles are dynamic structures and it might be anticipated that where additional energetic ''costs'' occur seasonally (e.g. due to migratory fattening or the development of large secondary sexual characteristics) then the relevant cardiac and locomotor musculature might also be regulated seasonally. This is an important consideration, both due to the intrinsic interest of studying muscular adaptation to changes in energy demand, but also as a confounding variable in the practical use of heart rate to estimate the energetics of animals. Haemoglobin concentration (or haematocrit) may also be a confounding variable. Thus, it is concluded that data on the cardiovascular and flight muscle morphology of animals provides essential information regarding the behavioural, ecological and physiological significance of the flight performance of animals.     

Sperm length variation as a predictor of extrapair paternity in passerine birds

Background

The rate of extrapair paternity is a commonly used index for the risk of sperm competition in birds, but paternity data exist for only a few percent of the approximately 10400 extant species. As paternity analyses require extensive field sampling and costly lab work, species coverage in this field will probably not improve much in the foreseeable future. Recent findings from passerine birds, which constitute the largest avian order (∼5 900 species), suggest that sperm phenotypes carry a signature of sperm competition. Here we examine how well standardized measures of sperm length variation can predict the rate of extrapair paternity in passerine birds.

Methodology/Principal Findings

We collected sperm samples from 55 passerine species in Canada and Europe for which extrapair paternity rates were already available from either the same (n = 24) or a different (n = 31) study population. We measured the total length of individual spermatozoa and found that both the coefficient of between-male variation (CV_bm) and within-male variation (CV_wm) in sperm length were strong predictors of the rate of extrapair paternity, explaining as much as 65% and 58%, respectively, of the variation in extrapair paternity among species. However, only the CV_bm predictor was independent of phylogeny, which implies that it can readily be converted into a currency of extrapair paternity without the need for phylogenetic correction.

Conclusion/Significance

We propose the CV_bm index as an alternative measure to extrapair paternity for passerine birds. Given the ease of sperm extraction from male birds in breeding condition, and a modest number of sampled males required for a robust estimate, this new index holds a great potential for mapping the risk of sperm competition across a wide range of passerine birds.     

Aerobic Biotransformation of N-Nitrosodimethylamine and N-Nitrodimethylamine by Benzene-, Butane-, Methane-, Propane-, and Toluene-Fed Cultures

N-Nitrosodimethylamine (NDMA) is an emerging contaminant of concern. N-nitrodimethylamine (DMNA) is a structural analog to NDMA. NDMA and DMNA have been found in drinking water, groundwater, and other media and are of concern due their toxicity. The authors evaluated biotransformation of NDMA and DMNA by cultures enriched from contaminated groundwater growing on benzene, butane, methane, propane, or toluene. Maximum specific growth rates of enriched cultures on butane (μ_max = 1.1 h^?1) and propane (μ_max = 0.65 h^?1) were 1 to 2 orders of magnitude higher than those presented in the literature. Growth rates of mixed cultures grown on benzene (μ_max = 1.3 h^?1), methane (μ_max = 0.09 h^?1), and toluene (μ_max = 0.99 h^?1) in these studies were similar to those presented in the literature. NDMA biotransformation rates for methane oxidizers (υ_max = 1.4 ng min^?1 mg^?1) and toluene oxidizers (υ_max = 2.3 ng min^?1 mg^?1) were comparable to those presented in the literature, whereas the biotransformation rate for propane oxidizers (υ_max = 0.37 ng min^?1 mg^?1) was lower. NDMA biotransformation rates for benzene oxidizers (υ_max = 1.02 ng min^?1 mg^?1) and butane oxidizers (υ_max = 1.2 ng min^?1 mg^?1) were comparable to those reported for other primary substrates. These studies showed that DMNA biotransformation rates for benzene (υ_max = 0.79 ng min^?1 mg^?1), butane (υ_max = 1.0 ng min^?1 mg^?1), methane (υ_max = 2.1 ng min^?1 mg^?1), propane (υ_max = 1.46 ng min^?1 mg^?1), and toluene (υ_max = 0.52 ng min^?1 mg^?1) oxidizers were all comparable. These studies highlight potential bioremediation methods for NDMA and DMNA in contaminated groundwater.     

Plasticity of the thermal requirements of exotherms and adaptation to environmental conditions

In exothermal organisms, temperature is an important determinant of the rate of ecophysiological processes, which monotonically increase between the minimum (t_{d min}) and maximum (t_{d max}) temperatures typical for each species. In insects, t_{d min} and t_{d max} are correlated and there is a approximately 20°C interval (thermal window W_T = t_{d max} − t_{d min}) between them over which insects can develop. We assumed that other exotherms have similar thermal windows because the thermal kinetics of their physiological processes are similar. In this study, we determined the thermal requirements for germination in plants. Seeds of 125 species of Central European wild herbaceous and crop plants were germinated at nine constant temperatures between 5 and 37°C, and the time to germination of 50% of the seeds D and rate of germination R (=1/D) were determined for each temperature and the Lactin model used to determine t_{d min}, t_{d max}, and W_T. The average width of the thermal windows for seeds was significantly wider (mean 24°C, 95% CI 22.7–24.2°C), varied more (between 14.5 and 37.5°C) and development occurred at lower temperatures than recorded for insects. The limiting temperatures for germination, t_{d min} and t_{d max}, were not coupled, so the width of the thermal window increased with both a decrease in t_{d min} and/or increase in t_{d max}. Variation in W_T was not associated with taxonomic affiliation, adult longevity, or domestication of the different species, but tends to vary with seed size. Plants are poor at regulating their temperature and cannot move to a more suitable location and as a consequence have to cope with wider ranges in temperatures than insects and possibly do this by having wider thermal windows. Synthesis: The study indicated specificity of W_T in different exotherm taxa and/or their development stages.     

Phenotypic flexibility of body composition associated with seasonal acclimatization in passerine birds

Improved winter cold tolerance is widespread among small birds overwintering in cold climates and is associated with improved shivering endurance and elevated summit metabolic rate (M_sum). Phenotypic flexibility resulting in elevated M_sum could result from either increased skeletal muscle mass (perhaps with support from similar adjustments in “nutritional organs”) and/or cellular metabolic intensity. We investigated seasonal changes in body composition of three species of passerine birds resident in cold winter climates, all of which show large seasonal variations in M_sum (>25%); white-breasted nuthatch (Sitta carolinensis), black-capped chickadee (Poecile atricapillus), and house sparrow (Passer domesticus). All three species displayed significant winter increases in pectoralis and heart masses, and supracoracoideus mass also increased in winter chickadees. Gizzard mass increased in winter for all three species, but masses of other nutritional organs did not vary consistently with season. These data suggest that winter increases in pectoralis and heart masses are important contributors to elevated thermogenic capacity and cold tolerance, but seasonal variation in nutritional organ masses, other than gizzard, which is likely associated with dietary changes, are not universally associated with seasonal phenotypes. The winter increases in pectoralis and heart masses are consistent with data from other small passerines showing marked seasonal changes in cold tolerance and support the Variable Maximum Model of seasonal phenotypic flexibility, where physiological adjustments that promote improved cold tolerance, also result in elevated M_sum.     

Energy expenditures for flight,aerodynamic quality,and colonization of forest habitats by birds

The power that the birds can use for flight (available power) and the power required for flight according to physical laws (requisite power) grow with an increase in body mass, the exponents of the corresponding functions being different. Small birds can follow different strategies, either improving the aerodynamic quality of the body (thereby saving the excess available power) or sacrifice aerodynamic quality in favor of morphological adaptation to factors other than the demands of flight proper, which provides the possibility of utilizing a wider range of ecological niches. A hypothesis is proposed that the high metabolic rate of passerine birds, compared to representatives of other bird orders, is an adaptation to maneuverable (i.e., relatively low-speed) flight necessary for successful colonization of forest habitats. The speed that birds of such size can develop according to the scaling theory is too high for nesting and foraging in tree crowns, and its reduction is possible in two ways: by increasing air drag or by changing the style of flight (by analogy with airplane vs. helicopter). The first way is feasible, but a high air drag due to morphological modifications (e.g., in the size of the tail or characteristics of the wing) interferes with the possibility of long-distance migration flight, as energy expenditures for it will exceed the energy potential of the bird. This is why migratory nonpasserine birds, which have used this strategy, are practically absent in forests of the temperate zone. Therefore, more promising is the second way involving transition to a new flight style and, in a certain sense, to a new morphophysiological organization. Passerines have achieved this by changing their flight style so that the wing actively generates forces (lift and thrust) only in downstroke. Such a flight requires more energy, and, to provide it in sufficient amounts, passerine birds have increased their basal metabolic rate (BMR). Thus, both their flight energy expenditures and BMR are higher than in nonpasserines. Remarkably, among approximately 8660 extant bird species known today, more than half (about 5100 species) belong to the order Passeriformes. Such a ratio, unknown in any other vertebrate class, is evidence that passerines have gained a considerable biological advantage over all other birds due to their increased BMR.     

Duplication of accelerated evolution and growth hormone gene in passerine birds

We report the discovery of a duplication of the growth hormone (GH) gene in a major group of birds, the passerines (Aves: Passeriformes). Phylogenetic analysis of 1.3-kb partial DNA sequences of GH genes for 24 species of passerines and numerous outgroups indicates that the duplication occurred in the ancestral lineage of extant passerines. Both duplicates and their open-reading frames are preserved throughout the passerine clade, and both duplicates are expressed in the zebra finch brain, suggesting that both are likely to be functional. The estimated rates of amino acid evolution are more than 10-fold higher in passerine GH genes than in those of their closest nonpasserine relatives. In addition, although the 84 codons sequenced are generally highly conserved for both passerines and nonpasserines, comparisons of the nonsynonymous/synonymous substitution ratios and the rate of predicted amino acid changes indicate that the 2 gene duplicates are evolving under different selective pressures and may be functionally divergent. The evidence of differential selection, coupled with the preservation of both gene copies in all major lineages since the origin of passerines, suggests that the duplication may be of adaptive significance, with possible implications for the explosive diversification of the passerine clade.     

Moult-related reduction of aerobic scope in passerine birds

It is well established that the nutrient and energy requirements of birds increase substantially during moult, but it is not known if these increased demands affect their aerobic capacity. We quantified the absolute aerobic scope of house and Spanish sparrows, Passer domesticus and P. hispaniolensis, respectively, before and during sequential stages of their moult period. The absolute aerobic scope (AAS) is the difference between maximum metabolic rate (MMR) during peak locomotor activities and minimum resting metabolic rate (RMR_min), thus representing the amount of aerobic power above that committed to maintenance needs available for other activities. As expected, RMR_min increased over the moult period by up to 40 and 63% in house and Spanish sparrows, respectively. Surprisingly, the maximum metabolic rates also decreased during moult in both species, declining as much as 25 and 38% compared with pre-moult values of house and Spanish sparrows, respectively. The concurrent changes in RMR_min and MMR during moult resulted in significant decreases in AAS, being up to 32 and 47% lower than pre-moult levels of house and Spanish sparrows, respectively, during moult stages having substantial feather replacement. We argue that the combination of reduced flight efficiency due to loss of wing feathers and reduced aerobic capacity places moulting birds at greater risk of predation. Such performance constraints likely contribute to most birds temporally separating moult from annual events requiring peak physiological capacity such as breeding and migration.

     

Within‐generation variation of critical thermal limits in adult Mediterranean and Natal fruit flies Ceratitis capitata and Ceratitis rosa: thermal history affects short‐term responses to temperature

Insect thermal tolerance shows a range of responses to thermal history depending on the duration and severity of exposure. However, few studies have investigated these effects under relatively modest temperature variation or the interactions between short‐ and longer‐term exposures. In the present study, using a full‐factorial design, 1 week‐long acclimation responses of critical thermal minimum (CT_min) and critical thermal maximum (CT_max) to temperatures of 20, 25 and 30 °C are investigated, as well as their interactions with short‐term (2 h) sub‐lethal temperature exposures to these same conditions (20, 25 and 30 °C), in two fruit fly species Ceratitis capitata (Wiedemann) and Ceratitis rosa Karsch from South Africa. Flies generally improve heat tolerance with high temperature acclimation and resist low temperatures better after acclimation to cooler conditions. However, in several cases, significant interaction effects are evident for CT_max and CT_min between short‐ and long‐term temperature treatments. Furthermore, to better comprehend the flies' responses to natural microclimate conditions, the effects of variation in heating and cooling rates on CT_max and CT_min are explored. Slower heating rates result in higher CT_max, whereas slower cooling rates elicit lower CT_min, although more variation is detected in CT_min than in CT_max (approximately 1.2 versus 0.5 °C). Critical thermal limits estimated under conditions that most closely approximate natural diurnal temperature fluctuations (rate: 0.06 °C min^?1) indicate a CT_max of approximately 42 °C and a CT_min of approximately 6 °C for these species in the wild, although some variation between these species has been found previously in CT_max. In conclusion, the results suggest critical thermal limits of adult fruit flies are moderated by temperature variation at both short and long time scales and may comprise both reversible and irreversible components.     

Thermal plasticity in the invasive south American tomato pinworm Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae)

South American tomato pinworm, Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) is a devastating invasive global insect pest of tomato, Solanum lycopersicum (Solanaceae). In nature, pests face multiple overlapping environmental stressors, which may significantly influence survival. To cope with rapidly changing environments, insects often employ a suite of mechanisms at both acute and chronic time-scales, thereby improving fitness at sub-optimal thermal environments. For T. absoluta, physiological responses to transient thermal variability remain under explored. Moreso, environmental effects and physiological responses may differ across insect life stages and this can have implications for population dynamics. Against this background, we investigated short and long term plastic responses to temperature of T. absoluta larvae (4th instar) and adults (24–48 h old) from field populations. We measured traits of temperature tolerance vis critical thermal limits [critical thermal minima (CT_min) and maxima (CT_max)], heat knockdown time (HKDT), chill coma recovery time (CCRT) and supercooling points (SCP). Our results showed that at the larval stage, Rapid Cold Hardening (RCH) significantly improved CT_min and HKDT but impaired SCP and CCRT. Heat hardening in larvae impaired CT_min, CCRT, SCP, CT_max but not HKDT. In adults, both heat and cold hardening generally impaired CT_min and CT_max, but had no effects on HKDT, SCP and CCRT. Low temperature acclimation significantly improved CT_min and HKDT while marginally compromising CCRT and CT_max, whereas high temperature acclimation had no significant effects on any traits except for HKDT in larvae. Similarly, low and high temperature acclimation had no effects on CT_min, SCPs and CT_max, while high temperature acclimation significantly compromised adult CCRT. Our results show that larvae are more thermally plastic than adults and can shift their thermal tolerance in short and long timescales. The larval plasticity reported here could be advantageous in new envirnments, suggesting an asymmetrical ecological role of larva relative to adults in facilitating T. absoluta invasion.     

Potential Impacts of Future Warming and Land Use Changes on Intra-Urban Heat Exposure in Houston,Texas

Extreme heat events in the United States are projected to become more frequent and intense as a result of climate change. We investigated the individual and combined effects of land use and warming on the spatial and temporal distribution of daily minimum temperature (T_min) and daily maximum heat index (HI_max) during summer in Houston, Texas. Present-day (2010) and near-future (2040) parcel-level land use scenarios were embedded within 1-km resolution land surface model (LSM) simulations. For each land use scenario, LSM simulations were conducted for climatic scenarios representative of both the present-day and near-future periods. LSM simulations assuming present-day climate but 2040 land use patterns led to spatially heterogeneous temperature changes characterized by warmer conditions over most areas, with summer average increases of up to 1.5°C (T_min) and 7.3°C (HI_max) in some newly developed suburban areas compared to simulations using 2010 land use patterns. LSM simulations assuming present-day land use but a 1°C temperature increase above the urban canopy (consistent with warming projections for 2040) yielded more spatially homogeneous metropolitan-wide average increases of about 1°C (T_min) and 2.5°C (HI_max), respectively. LSM simulations assuming both land use and warming for 2040 led to summer average increases of up to 2.5°C (T_min) and 8.3°C (HI_max), with the largest increases in areas projected to be converted to residential, industrial and mixed-use types. Our results suggest that urbanization and climate change may significantly increase the average number of summer days that exceed current threshold temperatures for initiating a heat advisory for metropolitan Houston, potentially increasing population exposure to extreme heat.     

Reverse lateralization of visual discriminative abilities in the European starling

Previous experiments on visual feature discrimination abilities have consistently shown a right-eye system lateralization in pigeons, Columba livia, and young domestic chickens, Gallus gallus domesticus, both nonpasserine species. Recently, however, it has been shown that photoreceptor distribution in the left and right retinas are asymmetrical in the European starling, Sturnus vulgaris, a passerine species. Single cone receptors are significantly more abundant in the left retina, which suggests that starlings should perform visual discrimination tasks more proficiently with the left eye, in contrast to previous findings with nonpasserines. We tested this hypothesis using the technique of monocular occlusion. In the first experiment, starlings were tested on a simultaneous visual discrimination task in three conditions: binocular (both eyes), left monocular (left eye only) and right monocular (right eye only). Subjects in the binocular and left-monocular conditions achieved significantly higher performance scores on the discrimination task than birds in the right-monocular condition. A second experiment found similar results, with birds in the left-monocular condition learning the discrimination task more than twice as quickly as those in the right-monocular condition. Subsequent tests with the alternative eye for both groups indicated no interocular transfer. These findings suggest that visual discriminative abilities in starlings are asymmetrical, and that they are lateralized in the opposite eye system than has been reported for all other species tested to date.     

Chewing lice from high‐altitude and migrating birds in Yunnan,China, with descriptions of two new species of Guimaraesiella

In total, 366 birds representing 55 species in 24 families and eight orders, were examined for chewing lice (Phthiraptera: Amblycera, Ischnocera) in two high‐altitude localities in Yunnan Province, China. In Ailaoshan, almost all of the birds examined were resident passeriforms, of which 36% were parasitized by chewing lice. In Jinshanyakou, most birds were on migration, and included both passerine and non‐passerine birds. Of the passerine birds caught in Jinshanyakou, only one bird (0.7%) was parasitized by chewing lice. The prevalence of Myrsidea and Brueelia‐complex lice on birds caught in Ailaoshan was higher than in previous reports. Of the chewing lice identifiable to species level, three represent new records for China: Actornithophilus hoplopteri (Mjöberg, 1910), Maculinirmus ljosalfar Gustafsson & Bush, 2017 and Quadraceps sinensis Timmermann, 1954. In total, 17 new host records are included, of which we describe two as new species in the Brueelia‐complex: Guimaraesiella (Cicchinella) ailaoshanensis sp. nov. ex Schoeniparus dubius dubius (Hume, 1874) and G. (C.) montisodalis sp. nov. ex Fulvetta manipurensis tonkinensis Delacour & Jabouille, 1930. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:9FC3D8EE‐2CED‐4DBE‐A1DB‐471B71260D27 .     

Insects, birds and lizards as pollinators of the largest-flowered Scrophularia of Europe and Macaronesia

Background and Aims

It has traditionally been considered that the flowers of Scrophularia are mainly pollinated by wasps. We studied the pollination system of four species which stand out for their large and showy flowers: S. sambucifolia and S. grandiflora (endemics of the western Mediterranean region), S. trifoliata (an endemic of the Tyrrhenian islands) and S. calliantha (an endemic of the Canary Islands). Our principal aim was to test whether these species were pollinated by birds or showed a mixed pollination system between insects and birds.

Methods

Censuses and captures of insects and birds were performed to obtain pollen load transported and deposited on the stigmas. Also, a qualitative and quantitative analysis of the flowers and inflorescences was carried out.

Key Results

Flowers were visited by Hymenoptera and by passerine birds. The Canarian species was the most visited by birds, especially by Phylloscopus canariensis, and its flowers were also accessed by juveniles of the lizard Gallotia stehlini. The most important birds in the other three species were Sylvia melanocephala and S. atricapilla. The most important insect-functional groups in the mixed pollination system were: honey-bees and wasps in S. sambucifolia; bumble-bees and wasps in S. grandiflora; wasps in S. trifoliata; and a small bee in S. calliantha.

Conclusions

The species studied show a mixed pollination system between insects and passerine birds. In S. calliantha there is, in addition, a third agent (juveniles of Gallotia stehlini). The participation of birds in this mixed pollination system presents varying degrees of importance because, while in S. calliantha they are the main pollinators, in the other species they interact to complement the insects which are the main pollinators. A review of different florae showed that the large showy floral morphotypes of Scrophularia are concentrated in the western and central Mediterranean region, Macaronesia and USA (New Mexico).     

Avoidance of headwinds or exploitation of ground effect—why do birds fly low?

ABSTRACT Birds often fly close to the ground or water. Wind shear theory predicts that wind speeds decline with proximity to the substratum, so birds might be expected to fly lower when flying upwind than when flying downwind. We tested this prediction and found that the wind shear equation is valid at heights below 4 m, with wind speed over a smooth surface ～40% lower at a height of 0.08 m than at 4 m. Birds that fly close enough to smooth substrata can also benefit energetically from ground effect, where vortices generated by their flight interact with the ground or water. This suggests that birds should use ground effect more when flying upwind than when flying downwind. We determined the percent time spent flying in ground effect by 21 species of passerine and non‐passerine birds flying in sheltered coastal aquatic and nearby terrestrial areas of County Cork, Ireland. We found that use of ground effect was uncommon for passerines, but common for a variety of non‐passerine waterbirds. However, phylogenetic analysis indicates no linkage between phylogeny and incidence of ground effect use and it is probable that incidence of use is determined by ecology rather than phylogeny. Great Cormorants (Phalacrocorax carbo) used ground effect most frequently over water (59.4% of time in flight). Over land, Barn Swallows (Hirundo rustica) used ground effect most often (19.8% of time). Phylogenetic contrasts regression analysis showed no significant relationship between use of ground effect and either wing aspect ratio or wing loading for 18 of our focal species, though simple linear regression analysis indicated that birds with greater wing loading used ground effect slightly (but significantly) more often. We found that 95% of Great Cormorants flying upwind used ground effect whereas only 35% did so when flying downwind. Few Black‐headed Gulls (Chroicocephalus ridibundus) used ground effect (probably because they fly high to locate prey), but still showed greater use when flying upwind (25%) than downwind (2.5%). When flying upwind in ground effect at the wind speeds encountered in our study, the velocity for minimum power (V_mp) for Great Cormorants was exceeded, suggesting theoretical benefits of about 14.3%. Our study indicates that several species exploit both wind shear and ground effect to minimize energy expenditure during commuting and foraging, but that others do not, because of either complexity of habitat morphology or the demands of their foraging ecology.     

Ontogeny of hypoxic modulation of cardiac performance and its allometry in the African clawed frog Xenopus laevis

The ontogeny of cardiac hypoxic responses, and how such responses may be modified by rearing environment, are poorly understood in amphibians. In this study, cardiac performance was investigated in Xenopus laevis from 2 to 25 days post-fertilization (dpf). Larvae were reared under either normoxia or moderate hypoxia (PO₂ = 110 mmHg), and each population was assessed in both normoxia and acute hypoxia. Heart rate (f _h ) of normoxic-reared larvae exhibited an early increase from 77 ± 1 beats min^?1 at 2 dpf to 153 ± 1 beats min^?1 at 4 dpf, followed by gradual decreases to 123 ± 3 beats min^?1 at 25 dpf. Stroke volume (SV), 6 ± 1 nl, and cardiac output (CO), 0.8 ± 0.1 μl min^?1, at 5 dpf both increased by more than 40-fold to 25 dpf with rapid larval growth (~30-fold increase in body mass). When exposed to acute hypoxia, normoxic-reared larvae increased f _h and CO between 5 and 25 dpf. Increased SV in acute hypoxia, produced by increased end-diastolic volume (EDV), only occurred before 10 dpf. Hypoxic-reared larvae showed decreased acute hypoxic responses of EDV, SV and CO at 7 and 10 dpf. Over the period of 2–25 dpf, cardiac scaling with mass showed scaling coefficients of ?0.04 (f _h ), 1.23 (SV) and 1.19 (CO), contrary to the cardiac scaling relationships described in birds and mammals. In addition, f _h scaling in hypoxic-reared larvae was altered to a shallower slope of ?0.01. Collectively, these results indicate that acute cardiac hypoxic responses develop before 5 dpf. Chronic hypoxia at a moderate level can not only modulate this cardiac reflex, but also changes cardiac scaling relationship with mass.     

Song structure variability in passerine birds: Random variation or direct informative changes

The variability of the fine song structure was studied in the wood warbler (Phylloscopus sibilatrix), willow warbler (Ph. trochilus), greenish warbler (Ph. throchiloides viridanus), chaffinch (Fringilla coelebs), and gray shrike thrush (Colluricincla harmonica). The increase in the number of similar elements per phrase is shown to be related to the conspecific and heterospecific male-male song interactions in these birds. The responses of wood warbler males to playback conspecific songs with standard and prolonged (in the number of similar elements) second phrases differ significantly. The initial phrases of willow warbler songs become significantly longer in the populations with a high density as compared to those in populations of low density. The increase in the variability of both phrase combinations and song duration is shown to be related also to the song interaction with conspecific males for the willow warbler, greenish warbler, chaffinch, and gray shrike thrush. The within-song type variations turned out to play an important role during male-male interactions of different species. The results obtained suggest that similar ways of song responses may be quite widespread among passerine birds and reflect the general principles of changes in the song structure during acoustic interactions.     

Application of leaf size and leafing intensity scaling across subtropical trees

Understanding the scaling between leaf size and leafing intensity (leaf number per stem size) is crucial for comprehending theories about the leaf costs and benefits in the leaf size–twig size spectrum. However, the scaling scope of leaf size versus leafing intensity changes along the twig leaf size variation in different leaf habit species remains elusive. Here, we hypothesize that the numerical value of scaling exponent for leaf mass versus leafing intensity in twig is governed by the minimum leaf mass versus maximum leaf mass (M _min versus M _max) and constrained to be ≤−1.0. We tested this hypothesis by analyzing the twigs of 123 species datasets compiled in the subtropical mountain forest. The standardized major axis regression (SMA) analyses showed the M _min scaled as the 1.19 power of M _max and the ‐α (−1.19) were not statistically different from the exponents of M _min versus leafing intensity in whole data. Across leaf habit groups, the M _max scaled negatively and isometrically with respect to leafing intensity. The pooled data''s scaling exponents ranged from −1.14 to −0.96 for M _min and M _max versus the leafing intensity based on stem volume (LIV). In the case of M _min and M _max versus the leafing intensity based on stem mass (LIM), the scaling exponents ranged from −1.24 to −1.04. Our hypothesis successfully predicts that the scaling relationship between leaf mass and leafing intensity is constrained to be ≤−1.0. More importantly, the lower limit to scaling of leaf mass and leafing intensity may be closely correlated with M _min versus M _max. Besides, constrained by the maximum leaf mass expansion, the broad scope range between leaf size and number may be insensitive to leaf habit groups in subtropical mountain forest.