scholarly journals The scaling of expansive energy under the Red Queen predicts Cope’s Rule

2019 ◽  
Author(s):  
Indrė Žliobaitė ◽  
Mikael Fortelius
Keyword(s):  
Body Size ◽  
Population Growth ◽  
Body Mass ◽  
Limiting Factor ◽  
Large Species ◽  
Small Species ◽  
Evolutionary Advantage ◽  
Zero Sum ◽  
The Red Queen ◽  
Cope’S Rule

AbstractThe Red Queen’s hypothesis portrays evolution as a never-ending competition for expansive energy, where one species’ gain is another species’ loss. The Red Queen is neutral with respect to body size, implying that neither small nor large species have a universal competitive advantage. The maximum population growth in ecology; however, clearly depends on body size – the smaller the species, the shorter the generation length, and the faster it can expand. Here we ask whether, and if so how, the Red Queen’s hypothesis can accommodate a spectrum of body sizes. We theoretically analyse scaling of expansive energy with body mass and demonstrate that in the Red Queen’s zero-sum game for resources, neither small nor large species have a universal evolutionary advantage. We argue that smaller species have an evolutionary advantage only when resources in the environment are not fully occupied, such as after mass extinctions or following key innovations allowing expansion into freed up or previously unoccupied resource space. Under such circumstances, we claim, generation length is the main limiting factor for population growth. When competition for resources is weak, smaller species can indeed expand faster, but to sustain this growth they also need more resources. In the Red Queen’s realm, where resources are fully occupied and the only way for expansion is to outcompete other species, acquisition of expansive energy becomes the limiting factor and small species lose their physiological advantage. A gradual transition from unlimited resources to a zero-sum game offers a direct mechanistic explanation for observed body mass trends in the fossil record, known as Cope’s Rule. When the system is far from the limit of resources and competition is not maximally intense, small species take up ecological space faster. When the system approaches the limits of its carrying capacity and competition tightens, small species lose their evolutionary advantage and we observe a wider range of successful body masses, and, as a result, an increase in the average body mass within lineages.

The flight of petrels and albatrosses (procellariiformes), observed in South Georgia and its vicinity

1982 ◽  
Vol 300 (1098) ◽  
pp. 75-106 ◽  
Keyword(s):  
Body Mass ◽  
The Body ◽  
Large Species ◽  
Small Species ◽  
Classical Dynamic ◽  
South Georgia ◽  
Mass Comparison ◽  
Gliding Flight ◽  
Field Season ◽  
Simple Power Function

Nine procellariiform species, covering a range of body mass exceeding 200: 1, were studied during a visit to Bird Island, South Georgia, with the British Antarctic Survey, in the 1979-1980 field season. Speed measurements were made by ornithodolite of birds slope-soaring over land, birds flying over the sea but observed from land, and birds observed from a ship. In the second group, which showed the least anomalies, lift coefficients corresponding to mean airspeeds were about 1 for albatrosses, decreasing to about 0.3 for the smallest petrels. All species increased speed when flying against the wind. The small species proceeded by flap-gliding, while the large ones flapped infrequently, and only in light winds. The small species flew lower than the larger ones, but this may be related to the fact that most of the observations were of birds flying into wind. The albatrosses ( Diomedea, Phoebetria ) and giant petrels ( Macronectes ) were found to have a ‘shoulder lock’, consisting of a tendon sheet associated with the pectoralis muscle, which restrained the wing from elevation above the horizontal. This arrangement was not seen in the smaller species, and was interpreted as an adaptation reducing the energy cost of gliding flight. The main soaring method in the large species appeared to be slope-soaring along waves. Windward ‘pullups’ suggestive of the classical ‘dynamic soaring’ technique were seen in large and medium-sized species. However, the calculated strength of the wind gradient would have been insufficient to maintain airspeed to the heights observed, and it was concluded that most of the energy for the pullups must come from kinetic energy, acquired by gliding along a wave in slope lift. Gliding downwind through the wind gradient should significantly increase the glide ratio, but this was not observed. Slope-soaring along moving waves in zero wind was recorded. The data were used to derive estimates of the average speeds that the different species should be able to maintain on foraging expeditions. Estimates of the rate of energy consumption were also made, taking into account the greater dependence on flapping in the smaller species, and on soaring in the larger ones. The distance travelled in consuming fuel equivalent to a given fraction of the body mass would seem to be very strongly dependent on mass. Comparison of the largest species ( Diomedea exulans ) with the smallest ( Oceanites oceanicus ) suggests that ‘range’, defined in this way, varies as the 0.60 power of the mass, although the relation is more complex than a simple power function.

scholarly journals The type of forest edge governs the spatial distribution of different-sized ground beetles

2020 ◽  
Vol 66 (Suppl.) ◽  
pp. 69-96
Author(s):  
Tibor Magura ◽  
Gábor L. Lövei
Keyword(s):  
Body Size ◽  
Human Disturbance ◽  
Forest Edge ◽  
Ground Beetles ◽  
The Body ◽  
Large Species ◽  
Forest Edges ◽  
Small Species ◽  
Natural Processes ◽  
Species Tolerance

Worldwide human-induced habitat fragmentation intensifies the emergence of forest edges. In addition to these edges, there are edges evolved by natural processes. Edge-maintaining processes (natural vs. anthropogenic) fundamentally determine edge responses, and thus edge functions. Species with various traits show fundamentally different edge response, therefore the trait-based approach is essential in edge studies. We evaluated the edge effect on the body size of ground beetles in forest edges with various maintaining processes. Our results, based on 30 published papers and 221 species, showed that natural forest edges were impenetrable for small species, preventing their dispersal into the forest interiors, while both the medium and the large species penetrated across these edges and dispersed into the forest interiors. Anthropogenic edges maintained by continued human disturbance (agriculture, forestry, urbanisation) were permeable for ground beetles of all size, allowing them to invade the forest interiors. Overwintering type (overwintering as adults or as larvae) was associated with body size, since almost two-thirds of the small species, while slightly more than a third of both the medium and the large species were adult overwintering. Based on this, size-dependent permeability of natural edges may be related to overwintering type, which basically determines species tolerance to human disturbance.

scholarly journals Body size of female calves and natality rates of known-aged females in two adjacent Alaskan caribou herds, and implications for management

Rangifer ◽  
2003 ◽  
Vol 23 (5) ◽  
pp. 203 ◽  
Author(s):  
Patrick Valkenburg ◽  
Robert W. Tobey ◽  
Robert W. Tobey ◽  
Bruce W. Dale ◽  
Bruce W. Dale ◽  
...  
Keyword(s):  
Body Size ◽  
Population Growth ◽  
Body Mass ◽  
Real Option ◽  
Rangifer Tarandus ◽  
Herd Size ◽  
Adult Females ◽  
Potential Population ◽  
Newborn Calves ◽  
Natality Rate

We studied body mass of female calves and natality rate of adult females in two adjacent Interior Alaskan caribou (Rangifer tarandus granti) herds during 1991-2001. Mass of newborn calves was similar in both herds, but Delta calves gained significantly more mass over summer than Nelchina calves. In contrast, Nelchina calves consistently maintained their mass during winter while Delta calves lost mass. Metatarsus length was similar in both herds in 4-month-old and 10-month-old calves, and it increased over winter in both herds. Natality rates of females >3 years old were consistently higher in the Delta Herd than in the Nelchina Herd, primarily because natality in 3- to 5-year-old Nelchina females was low. Although body mass of Delta Herd calves consistently declined over winter, we concluded that nutrition was not significantly limiting herd growth. Managers are more likely to maximize harvest by maintaining the Delta Herd near its present size (i.e., 3500), or allowing it to increase only slightly. The only real option for increasing harvestable surpluses of caribou in the Delta Herd is reducing predation during calving and summer. In contrast, we conclude that summer nutrition significantly limits potential population growth and body mass in the Nelchina Herd, and managers are more likely to maximize harvest by maintaining herd size at or below 30 000 than by allowing the herd to grow to near historical highs (i.e., 60 000-70 000).

Flight and size constraints: hovering performance of large hummingbirds under maximal loading.

1997 ◽  
Vol 200 (21) ◽  
pp. 2757-2763 ◽  
Author(s):  
P Chai ◽  
D Millard
Keyword(s):  
Body Mass ◽  
Power Output ◽  
Food Resource ◽  
The United States ◽  
Large Species ◽  
Small Species ◽  
Size Classes ◽  
Maximal Load ◽  
Mechanical Power Output ◽  
Burst Performance

As the smallest birds, hummingbirds are the only birds capable of prolonged hovering. This suggests that hovering locomotion scales unfavourably with size. Is the hovering performance of larger hummingbird species more constrained by size than that of smaller ones? Maximal load-lifting capacities of the two largest species of hummingbirds found in the United States, the blue-throated (Lampornis clemenciae, 8.4 g) and magnificent (Eugenes fulgens, 7.4 g) hummingbird, as well as the two other local small species, the black-chinned (Archilochus alexandri, 3.0 g) and rufus (Selasphorus rufus, 3.3 g) hummingbird, were determined under conditions of short-burst performance. The power reserves of hummingbirds are substantial relative to normal hovering performance. The two large species lifted maximal loads close to twice their body mass for a very brief duration of over 0.4 s. The small species lifted maximal loads approximately equal to their own mass with a longer duration of over 0.6 s. For the two large species under maximal loading, estimates of burst muscle mass-specific mechanical power output assuming perfect elastic energy storage averaged 309 W kg-1, compared with 75 W kg-1 during free hovering without loading. For the two small species, these values were 228 W kg-1 and 88 W kg-1, respectively. The differences in aerodynamic force production and power output between the large and small size classes occur despite their similar wing stroke velocity. This indicates that, during burst performance in these hummingbirds, the larger ones had a higher load-lifting capacity and generated more muscle power. In spite of the twofold difference in body mass, both large and small hummingbirds have evolved to become potent aerial competitors in order to exploit their common food resource, nectar. Both size classes have evolved to cope with the multi-dimensional effects of size constraining their aerodynamics, muscle mechanics, metabolism and ecology.

scholarly journals All Sizes Fit the Red Queen

Paleobiology ◽  
2020 ◽  
Vol 46 (4) ◽  
pp. 478-494
Author(s):  
Indrė Žliobaitė ◽  
Mikael Fortelius
Keyword(s):  
Body Size ◽  
Energy Use ◽  
Large Species ◽  
Red Queen ◽  
Metabolic Scaling ◽  
Physiological Time ◽  
Time Modeling ◽  
Wide Range ◽  
Body Sizes ◽  
The Red Queen

AbstractThe Red Queen's hypothesis portrays evolution as a never-ending competition for expansive energy, where one species’ gain is another species’ loss. The Red Queen is neutral with respect to body size, implying that neither small nor large species have a universal competitive advantage. Here we ask whether, and if so how, the Red Queen's hypothesis really can accommodate differences in body size. The maximum population growth in ecology clearly depends on body size—the smaller the species, the shorter the generation length, and the faster it can expand given sufficient opportunity. On the other hand, large species are more efficient in energy use due to metabolic scaling and can maintain more biomass with the same energy. The advantage of shorter generation makes a wide range of body sizes competitive, yet large species do not take over. We analytically show that individuals consume energy and reproduce in physiological time, but need to compete for energy in real time. The Red Queen, through adaptive evolution of populations, balances the pressures of real and physiological time. Modeling competition for energy as a proportional prize contest from economics, we further show that Red Queen's zero-sum game can generate unimodal hat-like patterns of species rise and decline that can be neutral in relation to body size.

Gape and energy limitation determine a humped relationship between trophic position and body size

2015 ◽  
Vol 72 (2) ◽  
pp. 198-205 ◽  
Author(s):  
Angel Manuel Segura ◽  
Valentina Franco-Trecu ◽  
Paula Franco-Fraguas ◽  
Matías Arim
Keyword(s):  
Body Size ◽  
Body Mass ◽  
Nitrogen Isotopes ◽  
Trophic Position ◽  
Negative Slope ◽  
Large Species ◽  
Metabolic Demand ◽  
Southwestern Atlantic Ocean ◽  
Body Sizes ◽  
Energy Limitation

We found a segmented pattern, increasing for small sizes and decreasing for larger sizes, in the relationship between trophic position and body size. This pattern provides support for a recently developed theoretical model whose derivation was based on consumers’ metabolic requirements and on basic assumptions about feeding relationships. We combined original and published information about stable nitrogen isotopes, a proxy of trophic position, for a broad range of animal body sizes (10−3–105 kg) inhabiting the southwestern Atlantic Ocean. Linear, polynomic, and piecewise segmented models were fit to species trophic position and body mass. The segmented model had the best fit, presenting a positive slope (β1 = 0.33 ± 0.08) for small organisms (<200 kg) and a negative slope (β2 = −1.93 ± 0.16) for larger ones. This suggests that there are morphological restrictions to prey consumption in smaller organisms and energetic constraints to trophic position in larger ones. Furthermore, the predator–prey body mass ratio (BMR = 1.31; 95% CI = 0.9–2.40) estimated here is similar to previous reports of direct observations (BMR = 1.64 and 1.82). However, the trophic position of larger organisms decreases at a faster rate (β2 = −1.93) than expected by metabolic demand (β2expected = −0.16 to −0.82), suggesting that additional processes should be considered. Our results suggest that large species could be more vulnerable to global change than previously thought.

scholarly journals Morphological determinants of jumping performance in the Iberian green frog

2019 ◽  
Vol 66 (4) ◽  
pp. 417-424
Author(s):  
Gregorio Moreno-Rueda ◽  
Abelardo Requena-Blanco ◽  
Francisco J Zamora-Camacho ◽  
Mar Comas ◽  
Guillem Pascual
Keyword(s):  
Body Size ◽  
Body Mass ◽  
Fluctuating Asymmetry ◽  
Hind Limb ◽  
Limb Length ◽  
Age Classes ◽  
Jumping Performance ◽  
Hind Limbs ◽  
Selective Forces ◽  
Pelophylax Perezi

Abstract Predation is one of the main selective forces in nature, frequently selecting potential prey for developing escape strategies. Escape ability is typically influenced by several morphological parameters, such as morphology of the locomotor appendices, muscular capacity, body mass, or fluctuating asymmetry, and may differ between sexes and age classes. In this study, we tested the relationship among these variables and jumping performance in 712 Iberian green frogs Pelophylax perezi from an urban population. The results suggest that the main determinant of jumping capacity was body size (explaining 48% of variance). Larger frogs jumped farther, but jumping performance reached an asymptote for the largest frogs. Once controlled by structural body size, the heaviest frogs jumped shorter distances, suggesting a trade-off between fat storage and jumping performance. Relative hind limb length also determined a small but significant percentage of variance (2.4%) in jumping performance—that is, the longer the hind limbs, the greater the jumping capacity. Juveniles had relatively shorter and less muscular hind limbs than adults (for a given body size), and their jumping performance was poorer. In our study population, the hind limbs of the frogs were very symmetrical, and we found no effect of fluctuating asymmetry on jumping performance. Therefore, our study provides evidence that jumping performance in frogs is not only affected by body size, but also by body mass and hind limb length, and differ between age classes.

scholarly journals Ontogeny of body size and shape of Antarctic and subantarctic fur seals

2007 ◽  
Vol 85 (12) ◽  
pp. 1275-1285 ◽  
Author(s):  
Sebastián P. Luque ◽  
Edward H. Miller ◽  
John P.Y. Arnould ◽  
Magaly Chambellant ◽  
Christophe Guinet
Keyword(s):  
Body Size ◽  
Body Mass ◽  
Morphological Traits ◽  
Body Shape ◽  
Size And Shape ◽  
Fur Seals ◽  
Fur Seal ◽  
Adult Body ◽  
Weaning Age ◽  
Lactation Duration

Pre- and post-weaning functional demands on body size and shape of mammals are often in conflict, especially in species where weaning involves a change of habitat. Compared with long lactations, brief lactations are expected to be associated with fast rates of development and attainment of adult traits. We describe allometry and growth for several morphological traits in two closely related fur seal species with large differences in lactation duration at a sympatric site. Longitudinal data were collected from Antarctic ( Arctocephalus gazella (Peters, 1875); 120 d lactation) and subantarctic ( Arctocephalus tropicalis (Gray, 1872); 300 d lactation) fur seals. Body mass was similar in neonates of both species, but A. gazella neonates were longer, less voluminous, and had larger foreflippers. The species were similar in rate of preweaning growth in body mass, but growth rates of linear variables were faster for A. gazella pups. Consequently, neonatal differences in body shape increased over lactation, and A. gazella pups approached adult body shape faster than did A. tropicalis pups. Our results indicate that preweaning growth is associated with significant changes in body shape, involving the acquisition of a longer, more slender body with larger foreflippers in A. gazella. These differences suggest that A. gazella pups are physically more mature at approximately 100 d of age (close to weaning age) than A. tropicalis pups of the same age.

scholarly journals Intraspecific Variation in Maximum Ingested Food Size and Body Mass in Varecia rubra and Propithecus coquereli

2011 ◽  
Vol 2011 ◽  
pp. 1-8 ◽  
Author(s):  
Adam Hartstone-Rose ◽  
Jonathan M. G. Perry
Keyword(s):  
Body Size ◽  
Body Mass ◽  
Individual Variation ◽  
Size Variation ◽  
Maximum Size ◽  
Least Squares Regression ◽  
Scaling Relationship ◽  
Varecia Rubra ◽  
Food Size ◽  
The Relationship

In a recent study, we quantified the scaling of ingested food size (Vb )—the maximum size at which an animal consistently ingests food whole—and found that Vb scaled isometrically between species of captive strepsirrhines. The current study examines the relationship between Vb and body size within species with a focus on the frugivorous Varecia rubra and the folivorous Propithecus coquereli. We found no overlap in Vb between the species (all V. rubra ingested larger pieces of food relative to those eaten by P. coquereli), and least-squares regression of Vb and three different measures of body mass showed no scaling relationship within each species. We believe that this lack of relationship results from the relatively narrow intraspecific body size variation and seemingly patternless individual variation in Vb within species and take this study as further evidence that general scaling questions are best examined interspecifically rather than intraspecifically.

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