Science.gov

Sample records for antbird hylophylax poecilinotus

  1. A putative RA-like region in the brain of the scale-backed antbird, Willisornis poecilinotus (Furnariides, Suboscines, Passeriformes, Thamnophilidae)

    PubMed Central

    de Lima, Jamily L.R.; Soares, Fabricio A.; Remedios, Ana C.S.; Thom, Gregory; Wirthlin, Morgan; Aleixo, Alexandre; Schneider, Maria Paula C.; Mello, Claudio V.; Schneider, Patricia N.

    2015-01-01

    The memorization and production of song in songbirds share important parallels with the process of speech acquisition in humans. In songbirds, these processes are dependent on a group of specialized telencephalic nuclei known as the song system: HVC (used as a proper name), RA (robust nucleus of arcopallium), LMAN (lateral magnocellular nucleus of the nidopallium) and striatal Area X. A recent study suggested that the arcopallium of the Sayornis phoebe, a non vocal learner suboscine species, contains a nucleus with some properties similar to those of songbird RA, suggesting that the song system may have been present in the last common ancestor of these groups. Here we report morphological and gene expression evidence that a region with some properties similar to RA is present in another suboscine, the Amazonian endemic Willisornis poecilinotus. Specifically, a discrete domain with a distinct Nissl staining pattern and that expresses the RA marker RGS4 was found in the arcopallium where the oscine RA is localized. Our findings, combined with the previous report on the S. phoebe, suggest that an arcopallial region with some RA-like properties was present in the ancestor of both Suboscines infraorders Tyranni and Furnarii, and is possibly an ancestral feature of Passeriformes. PMID:26500428

  2. Species loss, delayed numerical responses, and functional compensation in an antbird guild.

    PubMed

    Touchton, Janeene M; Smith, James N M

    2011-05-01

    When a community loses species through fragmentation, its total food consumption may drop. Compensatory responses of remaining species, whereby survivors assume roles of extinct competitors, may reduce the impact of species loss through numerical or functional responses. We measured compensatory responses in two remaining antbird species on Barro Colorado Island, Panama, four decades after the loss of their dominant competitor, the Ocellated Antbird, Phaenostictus mcleannani. We compared current abundances and behavior of these two species on Barro Colorado to those reported before the island lost Ocellated Antbirds, and to those in a nearby mainland population where all three species still exist as a space-for-time substitution. The smaller, more subordinate Spotted Antbird, Hylophylax naevioides, responded far more strongly than the larger Bicolored Antbird, Gymnopithys leucaspis, which is functionally more like the Ocellated Antbird. Islandwide density of Spotted Antbirds has more than doubled since the loss of Ocellated Antbirds. Moreover, Spotted Antbirds now spend so much more of their time following ant swarms that their metabolic biomass at these swarms has more than tripled since Ocellated Antbirds disappeared. These responses in Spotted Antbirds were apparently delayed by >20 years. Bicolored Antbirds have not increased substantially in islandwide density or metabolic biomass at ant swarms. We hypothesize that behavioral flexibility, as shown by Spotted Antbirds on Barro Colorado Island, is a major factor governing the extent to which fragmented ecosystems can buffer the impacts of species loss. PMID:21661573

  3. The effect of local dominance and reciprocal tolerance on feeding aggregations of ocellated antbirds.

    PubMed

    Chaves-Campos, Johel; Araya-Ajoy, Yi-Men; Lizana-Moreno, Claudia A; Rabenold, Kerry N

    2009-11-22

    We studied ocellated antbirds (Phaenostictus mcleannani) to test the hypothesis that reciprocal tolerance between dominant individuals can favour feeding in aggregations. Mated pairs hold large non-exclusive feeding ranges, but roost and nest in a small portion of this range ('roosting area'); adjacent roosting neighbours are unrelated. Ocellated antbirds congregate to feed on arthropods fleeing from nomadic swarms of army ants that move across the ranges of many pairs. We used playback experiments to simulate acoustic challenges, and results showed that males responded aggressively to other males only in their roosting areas. Responses to adjacent neighbours were less aggressive than to non-neighbours (i.e. the 'dear enemy' effect). Prey intake rates were higher when birds fed in their own roosting area or in that of adjacent neighbours compared with more distant sites. Males tolerated adjacent neighbours at swarm fronts where prey are most dense, but more distant neighbours were displaced. Despite small samples for some analyses, our results suggest that reciprocal tolerance between adjacent unrelated neighbours can ameliorate intraspecific competition within ephemeral feeding aggregations. PMID:19710061

  4. Phylogenetic conservatism and antiquity of a tropical specialization: army-ant-following in the typical antbirds (Thamnophilidae).

    PubMed

    Brumfield, Robb T; Tello, Jose G; Cheviron, Z A; Carling, Matthew D; Crochet, Nanette; Rosenberg, Kenneth V

    2007-10-01

    One of the most novel foraging strategies in Neotropical birds is army-ant-following, in which birds prey upon arthropods and small vertebrates flushed from the forest floor by swarm raids of the army-ant Eciton burchellii. This specialization is most developed in the typical antbirds (Thamnophilidae) which are divisible into three specialization categories: (1) those that forage at swarms opportunistically as army-ants move through their territories (occasional followers), (2) those that follow swarms beyond their territories but also forage independently of swarms (regular followers), and (3) those that appear incapable of foraging independently of swarms (obligate followers). Although army-ant-following is one of the great spectacles of tropical forests, basic questions about its evolution remain unaddressed. Using a strongly resolved molecular phylogeny of the typical antbirds, we found that army-ant-following is phylogenetically conserved, with regular following having evolved only three times, and that the most likely evolutionary progression was from least (occasional) to more (regular) to most (obligate) specialized, with no reversals from the obligate state. Despite the dependence of the specialists on a single ant species, molecular dating indicates that army-ant-following has persisted in antbirds since the late Miocene. These results provide the first characterization of army-ant-following as an ancient and phylogenetically conserved specialization. PMID:17768072

  5. High-Pitched Notes during Vocal Contests Signal Genetic Diversity in Ocellated Antbirds

    PubMed Central

    Araya-Ajoy, Yi-men; Chaves-Campos, Johel; Kalko, Elisabeth K. V.; DeWoody, J. Andrew

    2009-01-01

    Animals use honest signals to assess the quality of competitors during aggressive interactions. Current theory predicts that honest signals should be costly to produce and thus reveal some aspects of the phenotypic or genetic quality of the sender. In songbirds, research indicates that biomechanical constraints make the production of some acoustic features costly. Furthermore, recent studies have found that vocal features are related to genetic diversity. We linked these two lines of research by evaluating if constrained acoustic features reveal male genetic diversity during aggressive interactions in ocellated antbirds (Phaenostictus mcleannani). We recorded the aggressive vocalizations of radiotagged males at La Selva Biological Station in Costa Rica, and found significant variation in the highest frequency produced among individuals. Moreover, we detected a negative relationship between the frequency of the highest pitched note and vocalization duration, suggesting that high pitched notes might constrain the duration of vocalizations through biomechanical and/or energetic limitations. When we experimentally exposed wild radiotagged males to simulated acoustic challenges, the birds increased the pitch of their vocalization. We also found that individuals with higher genetic diversity (as measured by zygosity across 9 microsatellite loci) produced notes of higher pitch during aggressive interactions. Overall, our results suggest that the ability to produce high pitched notes is an honest indicator of male genetic diversity in male-male aggressive interactions. PMID:19956580

  6. Rivers, refuges and population divergence of fire-eye antbirds (Pyriglena) in the Amazon Basin.

    PubMed

    Maldonado-Coelho, M; Blake, J G; Silveira, L F; Batalha-Filho, H; Ricklefs, R E

    2013-05-01

    The identification of ecological and evolutionary mechanisms that might account for the elevated biotic diversity in tropical forests is a central theme in evolutionary biology. This issue is especially relevant in the Neotropical region, where biological diversity is the highest in the world, but where few studies have been conducted to test factors causing population differentiation and speciation. We used mtDNA sequence data to examine the genetic structure within white-backed fire-eye (Pyriglena leuconota) populations along the Tocantins River valley in the south-eastern Amazon Basin, and we confront the predictions of the river and the Pleistocene refuge hypotheses with patterns of genetic variation observed in these populations. We also investigated whether these patterns reflect the recently detected shift in the course of the Tocantins River. We sampled a total of 32 individuals east of, and 52 individuals west of, the Tocantins River. Coalescent simulations and phylogeographical and population genetics analytical approaches revealed that mtDNA variation observed for fire-eye populations provides little support for the hypothesis that populations were isolated in glacial forest refuges. Instead, our data strongly support a key prediction of the river hypothesis. Our study shows that the Tocantins River has probably been the historical barrier promoting population divergence in fire-eye antbirds. Our results have important implications for a better understanding of the importance of large Amazonian rivers in vertebrate diversification in the Neotropics. PMID:23442128

  7. Ecological opportunity leads to the emergence of an alternative behavioural phenotype in a tropical bird.

    PubMed

    Touchton, Janeene M; Wikelski, Martin

    2015-07-01

    Loss of a dominant competitor can open ecological opportunities. Ecological opportunities are considered prerequisites for adaptive radiations. Nonetheless, initiation of diversification in response to ecological opportunity is seldom observed, so we know little about the stages by which behavioural variation either increases or coalesces into distinct phenotypes. Here, a natural experiment showed that in a tropical island's guild of army ant-following birds, a new behavioural phenotype emerged in subordinate spotted antbirds (Hylophylax naevioides) after the socially dominant ocellated antbird (Phaenostictus mcleannani) died out. Individuals with this behavioural phenotype are less territorial; instead, they roam in search of ant swarms where they feed in locations from which dominant competitors formerly excluded them. Roaming individuals fledge more young than territorial individuals. We conclude that ecological opportunity arising from species loss may enhance the success of alternative behavioural phenotypes and can favour further intraspecific diversification in life-history traits in surviving species. PMID:25640464

  8. A neotropical forest bird can measure the slight changes in tropical photoperiod

    PubMed Central

    Hau, M.; Wikelski, M.; Wingfield, J. C.

    1998-01-01

    Many tropical birds breed seasonally, but it is largely unknown which environmental cues they use to time reproduction. Changes in tropical photoperiod have been regarded as too small to be used as a proximate environmental cue. This hypothesis, however, has never been rigorously tested. Here, we report on experimental evidence that photoperiodic changes characteristic of tropical latitudes stimulate reproductive activity in a neotropical bird from the forest understory. In the central Republic of Panam (9 degrees N), photoperiod varies annually between 12 hours (December) and 13 hours (June). Free-living spotted antbirds (Hylophylax n. naevioides) had regressed gonads in December, but increased gonads ahead of the rainy (the breeding) season in May. Captive spotted antbirds exposed to a 'long' photoperiod of 13 hours increased gonadal size eight-fold and song activity six-fold over that of control birds remaining on a simulated 'short' photoperiod of 12 hours of daylight. Moreover, even a photoperiod of 12 hours 17 minutes was sufficient to stimulate gonadal growth in photostimulated birds over that of controls. The dramatic changes in gonadal development were not accompanied by similar changes in hormone titres such as luteinizing hormone and testosterone as expected from temperate zone birds. We propose a more general role of the tropical photoperiod in the regulation of seasonal events in tropical organisms, or in temperate zone species migrating to the tropics.

  9. Captive Rearing Experiments Confirm Song Development without Learning in a Tracheophone Suboscine Bird

    PubMed Central

    Touchton, Janeene M.; Seddon, Nathalie; Tobias, Joseph A.

    2014-01-01

    The origin of vocal learning in animals has long been the subject of debate, but progress has been limited by uncertainty regarding the distribution of learning mechanisms across the tree of life, even for model systems such as birdsong. In particular, the importance of learning is well known in oscine songbirds, but disputed in suboscines. Members of this diverse group (∼1150 species) are generally assumed not to learn their songs, but empirical evidence is scarce, with previous studies restricted to the bronchophone (non-tracheophone) clade. Here, we conduct the first experimental study of song development in a tracheophone suboscine bird by rearing spotted antbird (Hylophylax naevioides) chicks in soundproofed aviaries. Individuals were raised either in silence with no tutor or exposed to standardized playback of a heterospecific tutor. All individuals surviving to maturity took a minimum of 79 days to produce a crystallized version of adult song, which in all cases was indistinguishable from wild song types of their own species. These first insights into song development in tracheophone suboscines suggest that adult songs are innate rather than learnt. Given that empirical evidence for song learning in suboscines is restricted to polygamous and lek-mating species, whereas tracheophone suboscines are mainly monogamous with long-term social bonds, our results are consistent with the view that sexual selection promotes song learning in birds. PMID:24788343

  10. Signal design and perception in Hypocnemis antbirds: evidence for convergent evolution via social selection.

    PubMed

    Tobias, Joseph A; Seddon, Nathalie

    2009-12-01

    Natural selection is known to produce convergent phenotypes through mimicry or ecological adaptation. It has also been proposed that social selection--i.e., selection exerted by social competition--may drive convergent evolution in signals mediating interspecific communication, yet this idea remains controversial. Here, we use color spectrophotometry, acoustic analyses, and playback experiments to assess the hypothesis of adaptive signal convergence in two competing nonsister taxa, Hypocnemis peruviana and H. subflava (Aves: Thamnophilidae). We show that the structure of territorial songs in males overlaps in sympatry, with some evidence of convergent character displacement. Conversely, nonterritorial vocal and visual signals in males are strikingly diagnostic, in line with 6.8% divergence in mtDNA sequences. The same pattern of variation applies to females. Finally, we show that songs in both sexes elicit strong territorial responses within and between species, whereas songs of a third, allopatric and more closely related species (H. striata) are structurally divergent and elicit weaker responses. Taken together, our results provide compelling evidence that social selection can act across species boundaries to drive convergent or parallel evolution in taxa competing for space and resources. PMID:19659594

  11. Phylogeny and phylogenetic classification of the antbirds, ovenbirds, woodcreepers, and allies (Aves: Passeriformes: Infraorder Furnariides)

    USGS Publications Warehouse

    Moyle, R.G.; Chesser, R.T.; Brumfield, R.T.; Tello, J.G.; Marchese, D.J.; Cracraft, J.

    2009-01-01

    The infraorder Furnariides is a diverse group of suboscine passerine birds comprising a substantial component of the Neotropical avifauna. The included species encompass a broad array of morphologies and behaviours, making them appealing for evolutionary studies, but the size of the group (ca. 600 species) has limited well-sampled higher-level phylogenetic studies. Using DNA sequence data from the nuclear RAG-1 and RAG-2 exons, we undertook a phylogenetic analysis of the Furnariides sampling 124 (more than 88%) of the genera. Basal relationships among family-level taxa differed depending on phylogenetic method, but all topologies had little nodal support, mirroring the results from earlier studies in which discerning relationships at the base of the radiation was also difficult. In contrast, branch support for family-rank taxa and for many relationships within those clades was generally high. Our results support the Melanopareidae and Grallariidae as distinct from the Rhinocryptidae and Formicariidae, respectively. Within the Furnariides our data contradict some recent phylogenetic hypotheses and suggest that further study is needed to resolve these discrepancies. Of the few genera represented by multiple species, several were not monophyletic, indicating that additional systematic work remains within furnariine families and must include dense taxon sampling. We use this study as a basis for proposing a new phylogenetic classification for the group and in the process erect new family-group names for clades having high branch support across methods. ?? 2009 The Willi Hennig Society.

  12. The spatial distribution of avian relatives: do obligate army-ant-following birds roost and feed near family members?

    PubMed

    Chaves-Campos, Johel; DeWoody, J Andrew

    2008-06-01

    The ocellated antbird (Phaenostictus mcleannani) feeds in groups and therefore is an informative species in which to study the biological factors that modulate avian group living. These birds congregate at swarms of army ants to capture fleeing prey, and previous observations suggest that males may be philopatric, feed with close relatives, and defend communal feeding ranges. We assessed whether kin selection could be an important factor maintaining group formation in a population of ocellated antbirds inhabiting continuous forest at La Selva Biological Station, Costa Rica, using radiotelemetry and 15 novel microsatellite markers. We predicted that the roosting areas of closely related adult males should overlap and that adult males feeding simultaneously at the same swarm should be highly related. We banded and genotyped 65 individuals (>or= 88% of the population) and radiotagged 30 of them. The results generally did not conform to our predictions. Little overlap occurred among the roosting areas of same-sex individuals, and nearest roosting neighbours (either same or opposite sex) were generally unrelated. A small proportion of male dyads suggested short-distance dispersal, but in general the distribution of genotypes within the study area approached randomness. We found little evidence of natal philopatry in either sex. Less than half of the feeding groups sampled included highly related males; most consisted of unrelated individuals. Hence, we found limited potential for kin selection to favour group living and suggest that other factors, particularly direct benefits (e.g. food intake), are probably more important than indirect effects (nepotism). PMID:18565030

  13. A new coccidian, Isospora parnaitatiaiensis n. sp. (Apicomplexa, Eimeriidae), from the white-shouldered fire-eye Pyriglena leucoptera (Passeriformes, Thamnophilidae) from South America.

    PubMed

    da Silva, Lidiane Maria; Rodrigues, Mariana Borges; Lopes, Bruno do Bomfim; Berto, Bruno Pereira; Luz, Hermes Ribeiro; Ferreira, Ildemar; Lopes, Carlos Wilson Gomes

    2016-02-01

    A new coccidian species (Protozoa: Apicomplexa: Isospora) parasitizing the white-shouldered fire-eye Pyriglena leucoptera (Vieillot, 1818) is described in the Parque Nacional do Itatiaia. This park is a protected area in southeastern Brazil with a high degree of vulnerability, representing a "conservation island" of biodiversity. Isospora parnaitatiaiensis n. sp. has oocysts that are ellipsoidal, 23.8 × 19.4 μm, with smooth, bilayered wall, ~1.1 μm thick. Micropyle and oocyst residuum are absent, but one or two polar granules are present. Sporocysts are ellipsoidal, 14.6 × 9.3 μm. The Stieda body is nipple- to knob-like and sub-Stieda body rounded to rectangular. Sporocyst residuum is present, usually as a cluster of numerous granules. Sporozoites are vermiform with two refractile bodies and a nucleus. This is the second isosporoid coccidian described from antbirds (Thamnophilidae). PMID:26508009

  14. Forest Fragmentation and Selective Logging Have Inconsistent Effects on Multiple Animal-Mediated Ecosystem Processes in a Tropical Forest

    PubMed Central

    Schleuning, Matthias; Farwig, Nina; Peters, Marcell K.; Bergsdorf, Thomas; Bleher, Bärbel; Brandl, Roland; Dalitz, Helmut; Fischer, Georg; Freund, Wolfram; Gikungu, Mary W.; Hagen, Melanie; Garcia, Francisco Hita; Kagezi, Godfrey H.; Kaib, Manfred; Kraemer, Manfred; Lung, Tobias; Schaab, Gertrud; Templin, Mathias; Uster, Dana; Wägele, J. Wolfgang; Böhning-Gaese, Katrin

    2011-01-01

    Forest fragmentation and selective logging are two main drivers of global environmental change and modify biodiversity and environmental conditions in many tropical forests. The consequences of these changes for the functioning of tropical forest ecosystems have rarely been explored in a comprehensive approach. In a Kenyan rainforest, we studied six animal-mediated ecosystem processes and recorded species richness and community composition of all animal taxa involved in these processes. We used linear models and a formal meta-analysis to test whether forest fragmentation and selective logging affected ecosystem processes and biodiversity and used structural equation models to disentangle direct from biodiversity-related indirect effects of human disturbance on multiple ecosystem processes. Fragmentation increased decomposition and reduced antbird predation, while selective logging consistently increased pollination, seed dispersal and army-ant raiding. Fragmentation modified species richness or community composition of five taxa, whereas selective logging did not affect any component of biodiversity. Changes in the abundance of functionally important species were related to lower predation by antbirds and higher decomposition rates in small forest fragments. The positive effects of selective logging on bee pollination, bird seed dispersal and army-ant raiding were direct, i.e. not related to changes in biodiversity, and were probably due to behavioural changes of these highly mobile animal taxa. We conclude that animal-mediated ecosystem processes respond in distinct ways to different types of human disturbance in Kakamega Forest. Our findings suggest that forest fragmentation affects ecosystem processes indirectly by changes in biodiversity, whereas selective logging influences processes directly by modifying local environmental conditions and resource distributions. The positive to neutral effects of selective logging on ecosystem processes show that the

  15. Year-round resource defence and the evolution of male and female song in suboscine birds: social armaments are mutual ornaments.

    PubMed

    Tobias, J A; Gamarra-Toledo, V; García-Olaechea, D; Pulgarín, P C; Seddon, N

    2011-10-01

    The evolution of sexually monomorphic (i.e. mutual) ornamentation has attracted growing attention as a 'blind-spot' in evolutionary biology. The popular consensus is that female ornaments are subject to the same modes of sexual selection as males: intrasexual competition and mate choice. However, it remains unclear how these forces interact within and between sexes, or whether they fully capture selection on female traits. One possibility is that the 'armament-ornament' model - which proposes that traits used primarily in male-male contests are also co-opted by females as indicators of male quality - can be extended to explain signal evolution in both sexes. We examine this idea by testing the function of acoustic signals in two species of duetting antbirds. Behavioural observations and playback experiments suggest that male and female songs function primarily as armaments in competitive interactions. Removal experiments reveal that song is also a classic ornament used by unpaired males and females to advertise for mates. These results indicate that 'armament-ornament' processes may operate in reciprocal format, potentially explaining widespread mutual ornamentation in species with elevated intrasexual competition for resources. In addition, given that songs mediate competition between species outside the breeding season, our findings suggest that processes shaping monomorphic ornaments extend beyond the traditional definitions of sexual selection and are best understood in the broader framework of social selection. PMID:21707816

  16. When David beats Goliath: the advantage of large size in interspecific aggressive contests declines over evolutionary time.

    PubMed

    Martin, Paul R; Ghalambor, Cameron K

    2014-01-01

    Body size has long been recognized to play a key role in shaping species interactions. For example, while small species thrive in a diversity of environments, they typically lose aggressive contests for resources with larger species. However, numerous examples exist of smaller species dominating larger species during aggressive interactions, suggesting that the evolution of traits can allow species to overcome the competitive disadvantage of small size. If these traits accumulate as lineages diverge, then the advantage of large size in interspecific aggressive interactions should decline with increased evolutionary distance. We tested this hypothesis using data on the outcomes of 23,362 aggressive interactions among 246 bird species pairs involving vultures at carcasses, hummingbirds at nectar sources, and antbirds and woodcreepers at army ant swarms. We found the advantage of large size declined as species became more evolutionarily divergent, and smaller species were more likely to dominate aggressive contests when interacting with more distantly-related species. These results appear to be caused by both the evolution of traits in smaller species that enhanced their abilities in aggressive contests, and the evolution of traits in larger species that were adaptive for other functions, but compromised their abilities to compete aggressively. Specific traits that may provide advantages to small species in aggressive interactions included well-developed leg musculature and talons, enhanced flight acceleration and maneuverability, novel fighting behaviors, and traits associated with aggression, such as testosterone and muscle development. Traits that may have hindered larger species in aggressive interactions included the evolution of morphologies for tree trunk foraging that compromised performance in aggressive contests away from trunks, and the evolution of migration. Overall, our results suggest that fundamental trade-offs, such as those associated with body size

  17. When David Beats Goliath: The Advantage of Large Size in Interspecific Aggressive Contests Declines over Evolutionary Time

    PubMed Central

    Martin, Paul R.; Ghalambor, Cameron K.

    2014-01-01

    Body size has long been recognized to play a key role in shaping species interactions. For example, while small species thrive in a diversity of environments, they typically lose aggressive contests for resources with larger species. However, numerous examples exist of smaller species dominating larger species during aggressive interactions, suggesting that the evolution of traits can allow species to overcome the competitive disadvantage of small size. If these traits accumulate as lineages diverge, then the advantage of large size in interspecific aggressive interactions should decline with increased evolutionary distance. We tested this hypothesis using data on the outcomes of 23,362 aggressive interactions among 246 bird species pairs involving vultures at carcasses, hummingbirds at nectar sources, and antbirds and woodcreepers at army ant swarms. We found the advantage of large size declined as species became more evolutionarily divergent, and smaller species were more likely to dominate aggressive contests when interacting with more distantly-related species. These results appear to be caused by both the evolution of traits in smaller species that enhanced their abilities in aggressive contests, and the evolution of traits in larger species that were adaptive for other functions, but compromised their abilities to compete aggressively. Specific traits that may provide advantages to small species in aggressive interactions included well-developed leg musculature and talons, enhanced flight acceleration and maneuverability, novel fighting behaviors, and traits associated with aggression, such as testosterone and muscle development. Traits that may have hindered larger species in aggressive interactions included the evolution of morphologies for tree trunk foraging that compromised performance in aggressive contests away from trunks, and the evolution of migration. Overall, our results suggest that fundamental trade-offs, such as those associated with body size