Energetic cost of hovering flight in nectar-feeding bats (Phyllostomidae: Glossophaginae) and its scaling in moths, birds and bats

Three groups of specialist nectar-feeders covering a continuous size range from insects, birds and bats have evolved the ability for hovering flight. Among birds and bats these groups generally comprise small species, suggesting a relationship between hovering ability and size. In this study we established the scaling relationship of hovering power with body mass for nectar-feeding glossophagine bats (Phyllostomidae). Employing both standard and fast-response respirometry, we determined rates of gas exchange in Hylonycteris underwoodi (7 g) and Choeronycteris mexicana (13–18 g) during hover-feeding flights at an artificial flower that served as a respirometric mask to estimate metabolic power input. The O₂ uptake rate (V˙ _o2) in ml g⁻¹ h⁻¹ (and derived power input) was 27.3 (1.12 W or 160 W kg⁻¹) in 7-g Hylonycteris and 27.3 (2.63 W or 160 W kg⁻¹) in 16.5-g Choeronycteris and thus consistent with measurements in 11.9-g Glossophagasoricina (158 W kg⁻¹, Winter 1998). V˙ _o2 at the onset of hovering was also used to estimate power during forward flight, because after a transition from level forward to hovering flight gas exchange rates initially still reflect forward flight rates. V˙ _o2 during short hovering events (<1.5 s) was 19.0 ml g⁻¹ h⁻¹ (1.8 W) in 16-g Choeronycteris, which was not significantly different from a previous, indirect estimate of the cost of level forward flight (2.1 W, Winter and von Helversen 1998). Our estimates suggest that power input during hovering flight P _h (W) increased with body mass M (kg) within 13–18-g Choeronycteris (n = 4) as P _h  = 3544 (±2057 SE) M ^{1.76 (±0.21 SE)} and between different glossophagine bat species (n = 3) as P _h  = 128 (±2.4 SE) M ^{0.95 (±0.034 SE)}. The slopes of three scaling functions for flight power (hovering, level forward flight at intermediate speed and submaximal flight power) indicate that: 1. The relationship between flight power to flight speed may change with body mass in the 6–30-g bats from a J- towards a U-shaped curve. 2. A metabolic constraint (hovering flight power equal maximal flight power) may influence the upper size limit of 30–35 g for this group of flower specialists. Mass-specific power input (W kg⁻¹) during hovering flight appeared constant with regard to body size (for the mass ranges considered), but differed significantly (P < 0.001) between groups. Group means were 393 W kg⁻¹ (sphingid moths), 261 W kg⁻¹ (hummingbirds) and 159 W kg⁻¹ (glossophagine bats). Thus, glossophagine bats expend the least metabolic power per unit of body mass supported during hovering flight. At a metabolic power input of 1.1 W a glossophagine bat can generate the lift forces necessary for balancing 7 g against gravitation, whereas a hummingbird can support 4 g and a sphingid moth only 3 g of body mass with the same amount of metabolic energy. These differences in power input were not fully explained by differences in induced power output estimated from Rankine-Froude momentum-jet theory. Accepted: 10 November 1998     

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.     

Collecting history of the hummingbird genera Chaetocercus Gray, 1855 and Lophornis Lesson, 1829

We examined the Chaetocercus and Lophornis holdings of 21 European, US and South American avian collections. Only skin specimens with exact (to the year) collecting dates and reliable locality data on their labels, which could be checked personally, were analysed. We located 466 Chaetocercus and 373 Lophornis specimens collected between the years 1859 and 1999. The vast majority of these records are old ones, dated between 1880 and 1960: 93% of the Chaetocercus and 82% of the Lophornis study skins. Less than 1% of all specimens was obtained after 1990. Potential explanations for our findings, such as declining population levels of these hummingbird genera, historically changing collecting activities regarding commercial dealers and collecting techniques, and legal collecting restrictions are discussed. Our findings reflect a general trend of decreasing collecting activities worldwide.Communicated by F. Bairlein     

Hummingbirds as net rate maximisers

The behaviour of foraging hummingbirds can be accounted for by assuming the maximisation of net rate. The currency, net energy per unit volume consumed suggested by Montgomerie et al. (1984), is shown to be an unnecessary means of interpreting behaviour when compared alongside an extended net rate model. The reason for the failure of previous net rate models to account for observed hummingbird feeding times lies in their negleet of the associated costs outside of the foraging period.     

Nectar robbing in Ipomopsis aggregata : effects on pollinator behavior and plant fitness

Hummingbirds foraging in alpine meadows of central Colorado, United States, face a heterogeneous distribution of nectar rewards. This study investigated how variability in nectar resources caused by nectar-robbing bumblebees affected the foraging behavior of hummingbird pollinators and, subsequently, the reproductive success of a host plant (Ipomopsis aggregata). We presented hummingbirds with experimental arrays of I. aggregata and measured hummingbird foraging behavior as a function of known levels of nectar robbing. Hummingbirds visited significantly fewer plants with heavy nectar robbing (over 80% of available flowers robbed) and visited fewer flowers on those plants. These changes in hummingbird foraging behavior resulted in decreased percent fruit set as well as decreased total seed set in heavily robbed plants. These results indicate that hummingbird avoidance of nectar-robbed plants and flowers reduces plant fitness components. In addition, our results suggest that the mutualisms between pollinators and host plants may be affected by other species, such as nectar robbers. Received: 22 April 1998 / Accepted: 12 May 1998     

A new dimension to hummingbird-flower relationships

The close correspondence between the bills of hummingbirds and the lengths of the flowers they feed from has been interpreted as an example of coadaptation. Observations of birds feeding at flowers longer and shorter than their bills, however, and the lack of experimental evidence for any feeding advantage to short bills, seem to contradict this interpretation. I address this problem by considering a little-studied dimension of floral morphology: corolla diameter. In laboratory experiments on female ruby-throated hummingbirds (Archilochus colubris), probing abilities (maximum extraction depths) increased with increasing corolla diameter. Handling times increased with decreasing corolla diameter, resulting in handling time equivalents, i.e., flowers having the same handling times but different lengths and diameters. Longer-billed birds had greater maximum extraction depths and shorter handling times than shorter-billed birds at all corolla diameters greater than the width of the bill. In contrast, shorter-billed birds made fewer errors inserting their bills into narrow flowers. Hence, differences in bill lengths apparently are associated with trade-offs in foraging abilities, whereby longer-billed birds are able to feed at long flowers and may do so more quickly, whereas shorter-billed birds are able to feed more successfully at narrow flowers.     

New specimens of the early Oligocene Old World hummingbird Eurotrochilus inexpectatus

New specimens of the early Oligocene (32 million years ago) stem group hummingbird Eurotrochilus inexpectatus Mayr, 2004, are described from the type locality Frauenweiler in Southern Germany. One of these is the second slab of the holotype, whose existence has been hitherto unknown. The new fossils add significantly to our knowledge of the morphology of E. inexpectatus and allow more detailed comparisons with other stem and crown group Trochilidae. A new apomorphy of the clade (Eurotrochilus + crown group Trochilidae) is described. With species successively more closely related to the crown group, the temporal sequence of stem group hummingbirds in Europe indicates an Old World origin of Pan-Trochilidae. E. inexpectatus is the earliest long-beaked stem group hummingbird with hovering capabilities, and absence of modern-type hummingbirds before the early Oligocene may provide a maximum age for the origin of hummingbird-pollinated plants and the colonization of the New World by modern-type stem group hummingbirds.     

Phylogenetic uncertainty revisited: Implications for ecological analyses

Ecologists and biogeographers usually rely on a single phylogenetic tree to study evolutionary processes that affect macroecological patterns. This approach ignores the fact that each phylogenetic tree is a hypothesis about the evolutionary history of a clade, and cannot be directly observed in nature. Also, trees often leave out many extant species, or include missing species as polytomies because of a lack of information on the relationship among taxa. Still, researchers usually do not quantify the effects of phylogenetic uncertainty in ecological analyses. We propose here a novel analytical strategy to maximize the use of incomplete phylogenetic information, while simultaneously accounting for several sources of phylogenetic uncertainty that may distort statistical inferences about evolutionary processes. We illustrate the approach using a clade‐wide analysis of the hummingbirds, evaluating how different sources of uncertainty affect several phylogenetic comparative analyses of trait evolution and biogeographic patterns. Although no statistical approximation can fully substitute for a complete and robust phylogeny, the method we describe and illustrate enables researchers to broaden the number of clades for which studies informed by evolutionary relationships are possible, while allowing the estimation and control of statistical error that arises from phylogenetic uncertainty. Software tools to carry out the necessary computations are offered.     

Is natural selection promoting sexual dimorphism in the Green-backed Firecrown Hummingbird ( Sephanoides Sephaniodes )?

In many hummingbird species there is an opposite pattern of sexual dimorphism in bill length and other morphometric measures of body size. These differences seem to be closely related with differences in foraging ecology directly associated with a different resource exploitation strategy. The aim of this study was to assess if natural selection is acting on wing length and bill size in hummingbird males and females with different resource exploitation strategies (i.e., territorial males and non-territorial females). If competition for resources promotes sexual dimorphism as a selective pressure, males should be subjected to negative directional selection pressure for wing length and no selection pressure over bill size, while females should undergo positive directional selection pressure for both bill size and wing length. The morphometric data we collected suggests that there is no selection for wing length and bill size in male hummingbirds. In contrast, our females exhibited positive directional selection for both wing length and bill size. Although we cannot reject sexual selection acting on sexually dimorphic traits, this study suggests that natural selection may promote sexual dimorphism in traits that are closely related with hummingbird foraging ecology and resource exploitation strategies.