Energetics of New Zealand's temperate parrots

Abstract

As in most parrots (Order: Psittaciformes) studied, New Zealand species have, independent of body mass, high basal rates of metabolism, low thermal conductances, and precisely regulated body temperatures. An analysis of covariance showed that basal rate in parrots correlates with body mass and thermal climate; temperate species have basal rates that are 21% higher than those of tropical species; and New Zealand's parrot have basal rates 32% higher than expected from body mass. Present information suggests that basal rate in parrots appears to be correlated neither with water availability in the environment nor with food habits. High basal rates in parrots are associated with large pectoral muscle masses. The very small pectoral muscle masses of the kakapo, Strigops habroptilus, predict that this flightless parrot has a low basal rate.     

Flightless rails endemic to islands have lower energy expenditures and clutch sizes than flighted rails on islands and continents

Data are presented on the standard energetics of six flighted and five flightless species of rails (Aves: Rallidae). The factors influencing these data and those from three additional species available from the literature, one of which was flightless, are examined. Basal rate of metabolism correlates with body mass, residency on islands or continents, volant condition, pectoral muscle mass, and food habits, but not with climate. The greatest capacity (96.2%) to account for the variation in basal rate of metabolism in 15 populations that belong to the 14 species occurs when body mass, volant condition, and food habits are combined. Then flighted species have basal rates that average 1.38 times those of flightless species and herbivorous rails have basal rates that are 1.37 times those of omnivorous species, which means that, independent of body mass, flighted gallinules have basal rates that are 1.9 times those of flightless, omnivorous rails. Distribution, pectoral muscle mass, and flight ability cannot be combined in the same analysis because they code for similar information. The evolution of a flightless condition in rails requires the absence of eutherian predators, but has occurred in the presence of marsupial predators. Each of the six studied flightless rails independently evolved a flightless condition and a low basal rate, whereas the evolution of herbivory and an associated high basal rate evolved at least twice in these species. Flightless rails on islands have clutch sizes that are only about one-half those of flighted rails living on continents, the reduction in clutch size correlating with a reduction in basal rate of metabolism. Thermal conductance in rails is correlated with body mass and food habits: herbivorous rails had conductances that were 1.43 times those of omnivores, i.e., conductances are highest in species with the highest basal rates.     

A test of the wintering strategy hypothesis with teal (Anas crecca) ringed in the Camargue, southern France.

Body mass changes in wintering ducks have been considered as the consequence of their wintering strategy, i.e. low mass after autumn migration, active foraging leading to body mass restoration, then pairing leading to low body mass again. This pattern is supported by weights of hunter-killed birds, but this is not enough to rule out potential alternative hypotheses linked to the existence of different populations with different body masses. The use of data collected at ringing and subsequent recaptures of known birds demonstrates that teal (Anas crecca) gain and lose body mass from September to March.     

Avian influenza viruses and paramyxoviruses in wintering and resident ducks in Texas

Cloacal swabs were collected from teal (Anas crecca, Anas cyanoptera, Anas discors), mottled duck (Anas fulvigula) and northern pintail (Anas acuta) in Brazoria County, Texas, USA, during February 2001, mottled ducks during August 2001, and blue-winged teal (A. discors) during February 2002. Prevalence of avian influenza virus (AIV) infections during each sampling period were 11, 0, and 15%, respectively. The hemagglutinin (H) subtypes H2 and H7 were detected in both years, while the H8 subtype was detected in 2001 and the H1 subtype was detected in 2002. Avian paramyxovirus type 1 (APMV-1) was isolated from 13% of mottled ducks sampled in August 2001 and 30.7% of teal in February 2002. The season of isolation of both viruses and the majority of the AIV subtypes detected in this study are not typical based on previous reports of these viruses from North American ducks.     

Experimental duck plague in blue-winged teal and Canada geese

Ten adult blue-winged teal (Anas discors) and six Canada goose (Branta canadensis) goslings were inoculated with liver tissue from a natural case of duck plague in a wild mallard (Anas platyrhynchos). Four additional teal were placed in contact with the inoculated ducks. Inoculated teal died 63.5-68 hr after inoculation; two of the contact teal died 161-162 hr after exposure. Three of the goslings died 119-133 hr after inoculation, the others were killed when moribund 90-133 hr postinfection. The clinical course of disease was extremely rapid in both species. Signs were limited to sudden onset of profound weakness, ataxia, tremors and terminal convulsions. The only consistent gross lesion in the teal was a small dark spleen; half the teal also had inconspicuous foci of epithelial necrosis in the distal esophagus and in the cloaca. Goslings had more severe lesions, with focal hepatic necrosis visible in all, and mucosal necrosis over the intestinal lymphoid tissue, and intestinal hemorrhage in three birds each. Microscopic lesions in both species were similar to those reported in mallards. Duck plague might be overlooked at necropsy of blue-winged teal, because of the paucity of gross lesions.     

The metabolism of New Guinean pteropodid bats

Rate of metabolism and body temperature were measured in eight species of pteropodid bats that live in Papua New Guinea. These data and those from 13 other species available in the literature are examined to determine the factors with which the energetics of pteropodids are correlated. Lowland populations of species that weigh < 35 g generally have low basal rates and often become torpid, whereas their highland populations had standard or high basal rates and were precise thermoregulators, as were all adult pteropodids that weighed more than 60 g. In large pteropodids belonging to the genera Dobsonia and Pteropus, females are smaller and consequently have lower total basal rates than males. Compared to species found on continents and large islands, species limited in distribution to small islands have lower basal rates, both because of a smaller mass and a reduction of metabolism independent of body mass. These trends are most marked in females that belong to small-island species, adjustments that may facilitate reproduction and survival on small oceanic islands with a limited resource base. Minimal thermal conductances are usually low in species that roost exposed in trees, whereas large species that roost in caves have either standard or high conductances.     

The comparative energetics of 'caviomorph' rodents.

The energetics of 11 species of New World hystricognath ('caviomorph') rodents are presented and compared with data from the literature on 19 additional species. Log(10) body mass alone accounts for 94% of the variation in the log(10) basal rate of metabolism in caviomorphs. The residual variation in basal rate is correlated with the stratum on which species live: arboreal species have low basal rates; terrestrial and fossorial species have intermediate basal rates; and aquatic species have high basal rates. When stratum is not included in the analysis, folivores, especially those that are arboreal, have lower basal rates than species with other food habits when combined with log(10) body mass. Small island endemics, all of which are folivores, have basal rates that are 61% of continental species. Log(10) basal rate correlates with family affiliation when combined with log(10) mass, but only if no other factor is included. Therefore, caviomorphs with low basal rates are arboreal, folivorous, live on small islands and belong to the Capromyidae, whereas other character combinations are associated with higher basal rates. These observations demonstrate that the basal rates of caviomorphs reflect many factor interactions. No differences in basal rate were found to reflect climate. Log(10) mass, the only factor to correlate with conductance, accounts for 82% of the variation in log(10) minimal thermal conductance. Mean interspecific body temperature was 36.9 degrees C; it was lowest in aquatic and fossorial species.     

The comparative energetics of rigid endothermy: the Arvicolidae

Rate of metabolism and body temperature were examined in 24 species of arvicolid rodents, for 15 of which data are presented here, to determine the factors that influence their level of energy expenditure. Arvicolids are characterized by a high, precisely-regulated body temperature, a high basal rate of metabolism by general mammalian standards, and a standard thermal conductance, except at large masses, when low conductances can occur. No evidence of entrance into torpor is known for any arvicolid; its absence is associated with especially high basal rates at masses smaller than 52 g. Arvicolids that live in cold climates, i.e. at high altitudes and latitudes, have higher basal rates than species that live in other environments. Basal rates, however, appear to be independent of food habits, presumably because of the small mass of most species and possibly because all species are herbivorous. A small size in combination with herbivory has permitted arvicolids to maintain continuous endothermy, which in turn has permitted them to exploit cool-to-cold environments by means of a high rate of production. The use of daily or seasonal torpor might well have prevented arvicolids from attaining the pivotal position in energy transfer that they presently occupy in many high-latitude communities.     

Fish–duck interactions in boreal lakes in Finland as reflected by abundance correlations

We studied the hypothesis that fish play an important role in lake use by ducks (pairs and broods) in boreal lakes. The study was based on densities of different duck and fish species in 28 boreal lakes in southern Finland. We focused on the three most common duck species (mallard Anas platyrhynchos, green-winged teal A. crecca and common goldeneye Bucephala clangula) and on the three most common fish species (perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius) in the region. We considered both competitive and predatory interactions between ducks and fish, the perch and roach being potential competitors with ducks and the pike a potential predator of ducks. We found a negative association between green-winged teal brood density and total fish density, the other duck species having only a weak association with total fish density. When the three fish species were considered separately, a negative association, suggesting food competition, was found between perch, green-winged teal and goldeneye, whereas the role of roach as a food competitor seemed to be of minor importance. We did not find any clear signs of predatory effects of pike on ducks. Our results suggest that food competition is a more important factor than pike predation in affecting lake use by ducks in oligotrophic boreal environments in southern Finland.     

Ecological studies on Cyathocotyle bushiensis (Digenea) and Sphaeridiotrema globulus (Digenea), possible pathogens of dabbling ducks in southern Québec

Over the past 20 yr, recurrent late-summer mortality of dabbling ducks in southern Québec has been attributed to the digenean parasite Cyathocotyle bushiensis. This study attempted to determine whether this parasite was still implicated in the epizootics, and examined aspects of the ecology of the parasite in the definitive host. Comparison of prevalence and mean intensity of infection of all species of Digenea between salvaged carcasses and hunter-shot ducks revealed that C. bushiensis and a second digenean Sphaeridiotrema globulus were both significantly more prevalent in salvaged ducks. Mean intensity of infection was consistently higher for S. globulus in the salvaged carcasses than in hunter-shot ducks. These data provide strong circumstantial support for the hypothesis that C. bushiensis continues to contribute to the annual mortality, but also strongly suggest that S. globulus may be involved. Comparison of the relative numbers of each species of duck between the salvaged carcass sample and the hunter-shot sample revealed that blue-winged teal (Anas discors) occurred significantly more frequently in the salvaged carcass sample whereas wood ducks (Aix sponsa) were common in the hunter-shot sample but were never found in the salvaged carcass sample. A comparison of susceptibility to experimental infection with C. bushiensis revealed that mallards (Anas platyrhynchos), black ducks (Anas rubripes), blue-winged teal, pintail (Anas acuta) and lesser scaup (Aythya affinis) had similar susceptibilities but that wood ducks were significantly less susceptible to infection. Experimental infections also showed that age of mallards significantly influenced the number of parasites obtained 7 days postinfection; ducklings were most heavily infected. Significant differences were detected in the gross pathology caused by C. bushiensis among duck species. Data obtained from sentinel ducks placed in the field for 24 hr revealed that transmission of both C. bushiensis and S. globulus occurs at least between mid-May and mid-September and that large, potentially lethal, infections can be acquired in as little as 24 hr. Finally evidence is presented to suggest that ducks acquire resistance to challenge infections of C. bushiensis.     

Interspecific interactions and co-existence in dabbling ducks: observations and an experiment

We studied the possible role of resource limitation and interspecific competition in assemblages of dabbling ducks on breeding lakes in Finland and Sweden with observational and experimental data. After initial vegetation mapping and yearly censuses of ducks in 1985–1990, we collected observational data in 1991–1994 from 28 lakes with natural populations of mallard Anas platyrhynchos and teal A. crecca. Mallard and teal co-occur over vast areas in the Holarctic and they are the only breeding dabbling ducks on many oligotrophic lakes. Both species are migratory in our study regions, teal arriving later in spring than mallards. Log-linear model analysis of observational presence/absence data revealed a positive, not a negative, association between the species. This association was independent of habitat diversity as well as of lake size. Mallard-teal interaction was also studied in a cross-over introduction experiment in 32 other lakes in two years. Wing-clipped mallards were introduced to breeding lakes before the arrival of teal to induce resource limitation and interspecific competition, hypothesized to reduce lake use by teal. The density of mallard pairs on experimental lakes was 2.9–8.0 times higher than on controls, but there was no negative response by teal to the treatment. This is the first combined observational-experimental demonstration of lack of interspecific competition in waterfowl. Our results indicate that heterospecific attraction may affect species co-existence in dabbling ducks. Received: 28 October 1996 / Accepted: 13 January 1997     

Presence of fish affects lake use and breeding success in ducks

Several previous studies indicate that presence of fish has negative effects on waterbirds breeding on lakes, owing either to competition for common invertebrate prey or fish predation on ducklings/chicks. However, others have reported results to the contrary and it remains unresolved what factors trigger, inhibit, and modulate fish–waterbird interactions. The present study was designed to test the effect of fish presence per se, with a minimum of variation in possibly confounding environmental variables. Thus, after stratifying for area, depth, altitude, pH, and total phosphorus we compared 13 lakes with and 12 without fish (mainly pike Esox lucius and perch Perca fluviatilis) with respect to (i) general species richness of waterbirds, (ii) species-specific utilization and breeding success of two dabbling ducks (mallard Anas platyrhynchos and teal Anas crecca) and a diving duck (goldeneye Bucephala clangula). General species richness of waterbirds was higher on fishless lakes. Overall use (bird days) and brood number of teal and goldeneye were higher on fishless lakes. The latter also had more benthic and free-swimming prey invertebrates compared to lakes with fish. Mallard use, mallard brood number, and abundance of emerging insects did not differ between lake groups. Generalized linear models including fish presence as factor and considering seven environmental variables as covariates, confirmed that all waterbird variables except mallard days and broods were negatively correlated to fish presence. There was also a residual positive relationship of lake area on general species richness, teal days, and teal broods. Our data demonstrate a stronger effect of fish presence on diving ducks and small surface feeding ducks than on large surface-feeding ducks. We argue that observed patterns were caused by fish predation on ducks rather than by fish–duck competition for common prey.     

Efficacy of a type C botulism vaccine in green-winged teal

We tested the efficacy of a single dose of Botumink toxoid for protecting wild green-winged teal (Anas crecca) during botulism epizootics caused by Clostridium botulinum type C. We challenged control and immunized ducks with four different doses of type C botulinum toxin to determine the LD50 for this species and to evaluate vaccine protection. Fewer immunized ducks were affected with botulism than control ducks, indicating that a single dose of Botumink toxoid could increase the survival of ducks during epizootics. However, the frequency of immunized ducks with signs of botulism increased with the challenge dose of botulinum toxin. Even at doses of botulinum toxin approximately 2 to 4 green-winged teal LD50, about 50% of the immunized ducks were affected. We believe an improved vaccine or a better delivery system is required to justify immunization of wild birds for experimental survival studies.     

Gastrointestinal helminths of the Mexican duck, Anas platyrhynchos diazi Ridgway, from north central Mexico and southwestern United States

Twenty-five species of helminths, recovered from the gastrointestinal tracts of 129 Mexican ducks from Mexico and the United States, were all new host records. The species included: Echinoparyphium recurvatum, Echinostoma revolutum, Hypoderaeum conoideum, Notocotylus attenuatus, Prosthogonimus cuneatus, Zygocotyle lunata, Anomotaenia ciliata, Cloacotaenia megalops, Diorchis bulbodes, Diorchis sp., Drepanidotaenia lanceolata, Echinocotyle rosseteri, Fimbriaria fasciolaris, Fimbriarioides sp., Hymenolepis sp. 1, Hymenolepis sp. 2, Sobolevicanthus gracilis. Corynosoma constrictum, Polymorphus minutus, Amidostomum acutum, Echinuria sp., Epomidiostomum crami, Hystrichis varispinosus, Rusguniella arctica, and Tetrameres sp. Fimbriarioides sp. occurred predominantly in ducks from south-central Chihuahua, Mexico. The distributions for the other 24 species of the helminths were not significantly different among the four collecting areas. The helminth fauna for the 32 complete specimens and 97 intestinal tracts was distributed relatively evenly among the hosts with a calculated mean evenness of 0.77 +/- 0.15 and 0.89 +/- 0.03 respectively. The parasite fauna was more similar to those of the black duck Anas rubripes Brewster of eastern North America (53%), the mallard, Anas platyrhynchos (L.) (49%), and the mottled duck, Anas fulvigula Ridgway, from Florida (45%), than to the green-winged teal, Anas crecca (L.) (36%), the gadwall, Anas strepera (L.) (30%), and the American wigeon, Anas americana Gmelin (26%), collected in the Southwest.     

Analysis of factors that influence energy expenditure in honeyeaters (Meliphagidae)

The energy expenditure of the tūī (Prosthemadera novaeseelandiae), a meliphagid endemic to New Zealand, was measured and compared with 20 species of honeyeaters (family Meliphagidae) to determine whether its expenditure is influenced either by life in a moist, temperate climate or an island residence. Body mass in the honeyeaters accounted for 91.5% of the variation in basal rate. The combination of body mass, climate and the maximal limit to an altitudinal distribution explained 98.6% of the variation in basal rate with tropical, low-altitude species having the highest mass-independent rate. The basal rates of meliphagids in tropical highlands are similar to those in temperate lowlands, which may reflect similar food supplies. The tūī mass-independent expenditure appears to reflect an active lifestyle in a temperate climate with no evidence that an island residence influenced its rate, whereas sedentary birds on New Zealand have responded to island life with a depressed basal rate. An effective analysis of the variation in energy expenditure requires the inclusion of the ecological and behavioural characteristics that distinguish species.     

Physiological convergence amongst ant-eating and termite-eating mammals

Ant- and termite-eating are among the few food habits common to monotremes, marsupials, and eutherians. Data are reported on the rate of metabolism and temperature regulation of 14 species of mammals having these food habits, including two monotremes, one marsupial and 11 eutherians. Small mammals with these habits have comparatively high body temperatures and high basal rates of metabolism, but ant- and termite-eaters that weigh more than 1 kg generally have low body temperatures and low basal rates of metabolism. The higher basal rates in small species ensure effective temperature regulation. Low body temperatures in large species principally result from low rates of metabolism. Rates of metabolism are low in these mammals because they use a food that has a limited availability and a low energy density, the density being further decreased in large species by the ingestion of non-nutritive material during feeding. Burrowing habits in some large species also contribute to low rates of metabolism. The combination of body size, food habits, and presence or absence of burrowing behaviour can account for all but about 6% of the range in basal rate in ant- and termite-eaters. Ants and termites, because of their locally clumped distributions, permit a larger mass in terrestrial predators than do other invertebrate prey. The reason why so many "primitive" mammals feed on ants and termites is that, once evolved, mammals with these habits are nearly impossible to displace ecologically, because much of ecological replacement is associated with high rates of reproduction, which are themselves correlated with high rates of metabolism in eutherians. Consequently, the ecological replacement of ant- and termite-eaters is inhibited, because this food habit does not permit high rates of metabolism, except at small masses.     

Ecological factors affect the level and scaling of avian BMR

The basal rate of metabolism (BMR) in 533 species of birds, when examined with ANCOVA, principally correlates with body mass, most of the residual variation correlating with food habits, climate, habitat, a volant or flightless condition, use or not of torpor, and a highland or lowland distribution. Avian BMR also correlates with migratory habits, if climate and a montane distribution is excluded from the analysis, and with an occurrence on small islands if a flightless condition and migration are excluded. Residual variation correlates with membership in avian orders and families principally because these groups are behaviorally and ecologically distinctive. However, the distinction between passerines and other birds remains a significant correlate of avian BMR, even after six ecological factors are included, with other birds having BMRs that averaged 74% of the passerine mean. This combination of factors accounts for 97.7% of the variation in avian BMR. Yet, migratory species that belong to Anseriformes, Charadriiformes, Pelecaniformes, and Procellariiformes and breed in temperate or polar environments have mass-independent basal rates equal to those found in passerines. In contrast, penguins belong to an order of polar, aquatic birds that have basal rates lower than passerines because their flightless condition depresses basal rate. Passerines dominate temperate, terrestrial environments and the four orders of aquatic birds dominate temperate and polar aquatic environments because their high BMRs facilitate reproduction and migration. The low BMRs of tropical passerines may reflect a sedentary lifestyle as much as a life in a tropical climate. Birds have BMRs that are 30-40% greater than mammals because of the commitment of birds to an expensive and expansive form of flight.     

A refined model of body mass and population density in flightless birds reconciles extreme bimodal population estimates for extinct moa

Flightless birds were once the largest and heaviest terrestrial fauna on many archipelagos around the world. Robust approaches for estimating their population parameters are essential for understanding prehistoric insular ecosystems and extinction processes. Body mass and population density are negatively related for extant flightless bird species, providing a method for quantifying densities and population sizes of extinct flightless species. Here we assemble an updated global data set of body mass and population densities for extant flightless birds and estimate the relationship between these variables. We use generalised least squares models that account for phylogenetic relatedness and incorporate the effects of limiting factors (e.g. habitat suitability) on population density. We demonstrate the applicability of this allometric relationship to extinct species by estimating densities for each of the nine species of moa (Dinornithiformes) and generating a combined spatially explicit map of total moa density across New Zealand. To compare our density estimates with those previously published, we summed individual species' abundances to generate a mean national density of 2.02–9.66 birds km⁻² for low- and high-density scenarios, respectively. Our results reconcile the extreme bimodality of previous estimates (< 2 birds km⁻² and > 10 birds km⁻²) and are comparable to contemporary densities of large herbivorous wild mammals introduced into New Zealand about 150 yr ago. The revised moa density has little effect on the harvest rates required to bring about extinction within 150–200 yr, indicating that rapid extinction was an inevitable response to human hunting, irrespective of the initial population of moa.     

A preliminary report on the effects of parasitism by the leeches Theromyzon cooperi and Placobdella garoui on the physiology of the redbilled teal Anas erythrorhyncha

The effects of parasitism by leeches Theromyzon cooperi and Placobdella garoui on the redbilled teal Anas erythrorhyncha were investigated. Leeches with initial body mass ranging from 5.2 to 28.4 mg were allowed to parasitize redbilled teal clamped in Perspex containers. Theromyzon cooperi increased its body mass after parasitism with a mean of 10.5-fold compared to 6.9-fold for P. garoui. Changes were also observed in values of various haematological and biochemical variables of the redbilled teal monitored during this investigation.     

Basal rate of metabolism and temperature regulation of two desert herbivorous octodontid rodents: Octomys mimax and Tympanoctomys barrerae

Summary We studied the energetics of two herbivorous desert rodents from South America. The two species had slightly lower basal metabolic rates, lower thermal conductances, and higher temperature differentials than those expected from their body mass. Mass-independent basal rates of metabolism were higher than those reported for seed-eating desert rodents from North America. Our observations support the hypothesis that desert rodents that eat foods with high water content have higher mass-independent metabolic rates than seed-eating desert rodents.