Evolution Of Large Body Size Research Articles

Cymbospondylus (Ichthyopterygia) from the Early Triassic of Svalbard and the early evolution of large body size in ichthyosaurs

Ichthyosaurs were a highly successful group of marine reptiles in the Mesozoic. The ichthyosaur radiation is part of the recovery from the Permian-Triassic mass extinction. In the Early Triassic, this group underwent extensive global radiation, filling ecological niches for the first time that were later occupied by various other lineages of marine amniotes. However, the evolution of body size in ichthyosaurs is not fully understood, as most large-bodied taxa originate from the Middle Triassic and later, and are mostly known from only a few specimens. In this study, we describe three articulated posterior dorsal vertebrae (IGPB R660) of the ichthyosaur Cymbospondylus sp. from the latest Olenekian Keyserlingites subrobustus zone of the Vikinghøgda Formation of the Agardhdalen area, eastern Spitsbergen, Svalbard. We numerically estimated the total body length of IGPB R660 from dorsal vertebral centrum length using a comparative dataset of other species of the genus and two different allometric analyses. This approach yields total length estimates of 7.5 m and 9.5 m for the individual, respectively, the highest for any unambiguous Early Triassic ichthyosaur find. Earlier, higher estimates of 11 m were based on taxonomically and stratigraphically inconclusive material but do not appear unreasonable based on evidence provided in this paper. Our study underscores both the rapid ecosystem recovery after a major mass extinction and extremely rapid increases in body size in ichthyosaurs after their adaptation to a secondarily aquatic lifestyle.

Forelimb muscle activation patterns in American alligators: Insights into the evolution of limb posture and powered flight in archosaurs.

The evolution of archosaurs provides an important context for understanding the mechanisms behind major functional transformations in vertebrates, such as shifts from sprawling to erect limb posture and the acquisition of powered flight. While comparative anatomy and ichnology of extinct archosaurs have offered insights into musculoskeletal and gait changes associated with locomotor transitions, reconstructing the evolution of motor control requires data from extant species. However, the scarcity of electromyography (EMG) data from the forelimb, especially of crocodylians, has hindered understanding of neuromuscular evolution in archosaurs. Here, we present EMG data for nine forelimb muscles from American alligators during terrestrial locomotion. Our aim was to investigate the modulation of motor control across different limb postures and examine variations in motor control across phylogeny and locomotor modes. Among the nine muscles examined, m. pectoralis, the largest forelimb muscle and primary shoulder adductor, exhibited significantly smaller mean EMG amplitudes for steps in which the shoulder was more adducted (i.e., upright). This suggests that using a more adducted limb posture helps to reduce forelimb muscle force and work during stance. As larger alligators use a more adducted shoulder and hip posture, the sprawling to erect postural transition that occurred in the Triassic could be either the cause or consequence of the evolution of larger body size in archosaurs. Comparisons of EMG burst phases among tetrapods revealed that a bird and turtle, which have experienced major musculoskeletal transformations, displayed distinctive burst phases in comparison to those from an alligator and lizard. These results support the notion that major shifts in body plan and locomotor modes among sauropsid lineages were associated with significant changes in muscle activation patterns.

Evolutionary loss of complexity in animal signals: cause and consequence.

We identified hypotheses for the cause and consequences of the loss of complexity in animal signals and tested these using a genus of visually communicating lizards, the Southeast Asian Draco lizards. Males of some species have lost the headbob component from their display, which is otherwise central to the communication of this genus. These males instead display a large, colorful dewlap to defend territories and attract mates. This dewlap initially evolved to augment the headbob component of the display, but has become the exclusive system of communication. We tested whether the loss of headbobs was caused by relaxed selection, habitat-dependent constraints, or size-specific energetic constraints on display movement. We then examined whether the consequences of this loss have been mitigated by increased signaling effort or complexity in the color of the dewlap. It appears the increased cost of display movement resulting from the evolution of large body size might have contributed to the loss of headbobs and has been somewhat compensated for by the evolution of greater complexity in dewlap color. However, this evolutionary shift is unlikely to have maintained the complexity previously present in the communication system, resulting in an apparent detrimental loss of information potential.

Anatomical mechanism for protecting the airway in the largest animals on earth

Separation of respiratory and digestive tracts in the mammalian pharynx is critical for survival. Food must be kept out of the respiratory tract, and air must be directed into the respiratory tract when breathing.1 Cetaceans have the additional problem of feeding while underwater. Lunge-feeding baleen whales (rorquals) open the mouth while swimming at high speeds to engulf a volume of prey-laden water as large as their own body2 and experience tremendous forces as water floods the mouth. How the respiratory tract is protected in the pharynx during engulfment and while swallowing a massive slurry of tiny living prey remains unknown, despite its importance to survival. By dissecting adult and fetal fin whales, we determined that a large musculo-fatty structure passively seals the oropharyngeal channel. This "oral plug" is not observed in other animals, and its position indicates it must be shifted to allow swallowing; it is a part of the soft palate and can only shift posteriorly and dorsally. Elevation of the oral plug allows food transfer to the pharynx and protects the upper airways from food entry. The laryngeal inlet in the floor of the pharynx is sealed by laryngeal cartilages, and the muscular laryngeal sac moves upward into the laryngeal cavity, completely occluding the airway. The pharynx is dedicated to the digestive tract during swallowing, with no connection between upper and lower airways. These adaptations to facilitate swallowing were a critical development in the evolution of large body size in these, the largest animals on earth.

Early giant reveals faster evolution of large body size in ichthyosaurs than in cetaceans.

Body sizes of marine amniotes span six orders of magnitude, yet the factors that governed the evolution of this diversity are largely unknown. High primary production of modern oceans is considered a prerequisite for the emergence of cetacean giants, but that condition cannot explain gigantism in Triassic ichthyosaurs. We describe the new giant ichthyosaur Cymbospondylus youngorum sp. nov. with a 2-meter-long skull from the Middle Triassic Fossil Hill Fauna of Nevada, USA, underscoring rapid size evolution despite the absence of many modern primary producers. Surprisingly, the Fossil Hill Fauna rivaled the composition of modern marine mammal faunas in terms of size range, and energy-flux models suggest that Middle Triassic marine food webs were able to support several large-bodied ichthyosaurs at high trophic levels, shortly after ichthyosaur origins.

Mitogenomic analysis of extant condor species provides insight into the molecular evolution of vultures

The evolution of large vultures linked to mountainous habitats was accompanied by extreme physiological and behavioral specializations for energetically efficient flights. However, little is known on the genetic traits associated with the evolution of these obligate soaring scavengers. Mitochondrial DNA plays a vital role in regulating oxidative stress and energy production, and hence may be an important target of selection for flight performance. Herein, we characterized the first mitogenomes of the Andean and California condors, the world’s heaviest flying birds and the only living representatives of the Vultur and Gymnogyps genus. We reconstructed the phylogenetic relationships and evaluated possible footprints of convergent evolution associated to the life-history traits and distributional range of vultures. Our phylogenomic analyses supported the independent evolution of vultures, with the origin of Cathartidae in the early Paleogene (~ 61 Mya), and estimated the radiation of extant condors during the late Miocene (~ 11 Mya). Selection analyses indicated that vultures exhibit signals of relaxation of purifying selection relative to other accipitrimorph raptors, possibly indicating the degeneration of flapping flight ability. Overall, our results suggest that the extreme specialization of vultures for efficient soaring flight has compensated the evolution of large body sizes mitigating the selection pressure on mtDNA.

Social selection is density dependent but makes little contribution to total selection in New Zealand giraffe weevils.

Social selection occurs when traits of interaction partners influence an individual's fitness and can alter total selection strength. However, we have little idea of what factors influence social selection's strength. Further, social selection only contributes to overall selection when there is phenotypic assortment, but simultaneous estimates of social selection and phenotypic assortment are rare. Here, we estimated social selection on body size in a wild population of New Zealand giraffe weevils (Lasiorhynchus barbicornis). We measured phenotypic assortment by body size and tested whether social selection varied with sex ratio, density and interacted with the body size of the focal individual. Social selection was limited and unaffected by sex ratio or the size of the focal individual. However, at high densities social selection was negative for both sexes, consistent with size-based competitive interactions for access to mates. Phenotypic assortment was always close to zero, indicating negative social selection at high densities will not impede the evolution of larger body sizes. Despite its predicted importance, social selection may only influence evolutionary change in specific contexts, leaving direct selection to drive evolutionary change.

An island-hopping bird reveals how founder events shape genome-wide divergence.

When populations colonize new areas, both strong selection and strong drift can be experienced due to novel environments and small founding populations, respectively. Empirical studies have predominantly focused on the phenotype when assessing the role of selection, and limited neutral-loci when assessing founder-induced loss of diversity. Consequently, the extent to which processes interact to influence evolutionary trajectories is difficult to assess. Genomic-level approaches provide the opportunity to simultaneously consider these processes. Here, we examine the roles of selection and drift in shaping genomic diversity and divergence in historically documented sequential island colonizations by the silvereye (Zosterops lateralis). We provide the first empirical demonstration of the rapid appearance of highly diverged genomic regions following population founding, the position of which are highly idiosyncratic. As these regions rarely contained loci putatively under selection, it is most likely that these differences arise via the stochastic nature of the founding process. However, selection is required to explain rapid evolution of larger body size in insular silvereyes. Reconciling our genomic data with these phenotypic patterns suggests there may be many genomic routes to the island phenotype, which vary across populations. Finally, we show that accelerated divergence associated with multiple founding steps is the product of genome-wide rather than localized differences, and that diversity erodes due to loss of rare alleles. However, even multiple founder events do not result in divergence and diversity levels seen in evolutionary older subspecies, and therefore do not provide a shortcut to speciation as proposed by founder-effect speciation models.

A new parapithecine (Primates: Anthropoidea) from the early Oligocene of Libya supports parallel evolution of large body size among parapithecids

Parapithecines are an extinct subfamily of stem anthropoid primates previously known only from the Jebel Qatrani Formation in Egypt. Here, we describe isolated teeth pertaining to Simonsius harujensis sp. nov., a relatively small-bodied parapithecine from strata near Zallah Oasis in the Sirt Basin of central Libya that is estimated to date to ∼31 Ma on the basis of mammalian biostratigraphy. The dental morphology of S. harujensis sp. nov. is generally intermediate between that of the closely related parapithecines Parapithecus fraasi and Simonsius grangeri, highlighting some of the anatomical features distinguishing the latter taxa and providing further support for their generic separation. A phylogenetic analysis using parsimony methods was performed on a character-taxon matrix incorporating data from the new Libyan parapithecine, virtually all other parapithecids and the proteopithecid Proteopithecus sylviae. Results of this analysis suggest that parapithecids comprise a basal clade consisting of three species of Biretia and a more derived clade including Parapithecinae (Parapithecus and Simonsius) and Qatraniinae (Qatrania, Ucayalipithecus, and Apidium). Body mass estimates for parapithecids were calculated on the basis of regression equations generated to predict body mass from the occlusal area of upper and lower cheek teeth in extant anthropoids. The relatively small body mass of S. harujensis sp. nov. and its reconstructed phylogenetic position as the sister group of S. grangeri, which is the largest known parapithecid, support the convergent acquisition of body mass larger than 500 g among multiple clades of early Oligocene African anthropoids. The new Libyan parapithecine augments previously reported evidence supporting a substantial degree of faunal provincialism across northern Africa/Arabia during the early Oligocene.

A new large-bodied Pliocene seal with unusual cutting teeth.

Today, monachine seals display the largest body sizes in pinnipeds. However, the evolution of larger body sizes has been difficult to assess due to the murky taxonomic status of fossil seals, including fossils referred to Callophoca obscura, a species thought to be present on both sides of the North Atlantic during the Neogene. Several studies have recently called into question the taxonomic validity of these fossils, especially those from the USA, as the fragmentary lectotype specimen from Belgium is of dubious diagnostic value. We find that the lectotype isolated humerus of C. obscura is too uninformative; thus, we designate C. obscura as a nomen dubium. More complete cranial and postcranial specimens from the Pliocene Yorktown Formation are described as a new taxon, Sarcodectes magnus. The cranial specimens display adaptations towards an enhanced ability to cut or chew prey that are unique within Phocidae, and estimates indicate S. magnus to be around 2.83 m in length. A parsimony phylogenetic analysis found S. magnus is a crown monachine. An ancestral state estimation of body length indicates that monachines did not have a remarkable size increase until the evolution of the lobodontins and miroungins.

Bone histology of varanopids (Synapsida) from Richards Spur, Oklahoma, sheds light on growth patterns and lifestyle in early terrestrial colonizers.

Varanopids were a group of small to medium-sized synapsids whose fossil record spans the Carboniferous through middle Permian. Although their phylogenetic relationships have received some interest in recent years, little is known about other aspects of their palaeobiology, including their skeletal growth, allometry and habitat preference. Here, we describe varanopid long bone histology based on a sample of well-preserved femora from the lower Permian Richards Spur fissure fill locality, Comanche County, Oklahoma, USA. The sample includes five femora from at least two varanopid taxa-Mycterosaurus and the large varanodontine Varanops brevirostris-and four additional mycterosaurine femora not diagnosed to genus. Prior work on femoral bone compactness provided a baseline to make lifestyle inferences and evaluate whether varanopids were ancestrally terrestrial. Moreover, the large availability of specimens spanning different sizes made possible an assessment of size-related ontogenetic histovariability. All specimens revealed moderately dense cortical bone tissues composed of sparsely vascularized parallel-fibred and lamellar bone with radially arranged rows of longitudinal canals (mostly simple), and many preserved regularly spaced growth marks (annuli and lines of arrested growth) as in modern varanids. We show that bone histology has the potential to explain how ballast was shed and the skeleton lightened for terrestrial mobility in ancestral synapsids and their basal amniote kin, as well as how adjustments in postnatal growth influenced the evolution of larger body sizes in the terrestrial frontier. This article is part of the theme issue 'Vertebrate palaeophysiology'.

Differential reproductive investment in co-occurring oviparous and viviparous common lizards (Zootoca vivipara) and implications for life-history trade-offs with viviparity

Live-bearing reproduction (viviparity) has evolved from egg-laying (oviparity) independently many times and most abundantly in squamate reptiles. Studying life-history trade-offs between the two reproductive modes is an inherently difficult task, as most transitions to viviparity are evolutionarily old and/or are confounded by environmental effects. The common lizard (Zootoca vivipara) is one of very few known reproductively bimodal species, in which some populations are oviparous and others viviparous. Oviparous and viviparous populations can occur in sympatry in the same environment, making this a unique system for investigating alternative life-history trade-offs between oviparous and viviparous reproduction. We find that viviparous females exhibit larger body size, smaller clutch sizes, a larger reproductive investment, and a higher hatching success rate than oviparous females. We find that offspring size and weight from viviparous females was lower compared to offspring from oviparous females, which may reflect space constraints during pregnancy. We suggest that viviparity in common lizards is associated with increased reproductive burden for viviparous females and speculate that this promoted the evolution of larger body size to create more physical space for developing embryos. In the context of life-history trade-offs in the evolution of viviparity, we suggest that the extent of correlation between reproductive traits, or differences between reproductive modes, may also depend on the time since the transition occurred.

Large neotheropods from the Upper Triassic of North America and the early evolution of large theropod body sizes

AbstractLarge body sizes among nonavian theropod dinosaurs is a major feature in the evolution of this clade, with theropods reaching greater sizes than any other terrestrial carnivores. However, the early evolution of large body sizes among theropods is obscured by an incomplete fossil record, with the largest Triassic theropods represented by only a few individuals of uncertain ontogenetic stage. Here I describe two neotheropod specimens from the Upper Triassic Bull Canyon Formation of New Mexico and place them in a broader comparative context of early theropod anatomy. These specimens possess morphologies indicative of ontogenetic immaturity (e.g., absence of femoral bone scars, lack of co-ossification between the astragalus and calcaneum), and phylogenetic analyses recover these specimens as early-diverging neotheropods in a polytomy with other early neotheropods at the base of the clade. Ancestral state reconstruction for body size suggests that the ancestral theropod condition was small (~240 mm femur length), but the ancestral neotheropod was larger (~300–340 mm femur length), with coelophysoids experiencing secondary body size reduction, although this is highly dependent on the phylogenetic position of a few key taxa. Theropods evolved large body sizes before the Triassic–Jurassic extinction, as hypothesized in most other ancestral state reconstructions of theropod body sizes, but remained rare relative to smaller theropods until the Jurassic.

A reexamination of Milosaurus mccordi, and the evolution of large body size in Carboniferous synapsids

ABSTRACTMilosaurus mccordi was described in 1970 as a large, pelycosaurian-grade synapsid from the latest Carboniferous of Illinois but has since received little attention. Here, the holotype and referred material of Milosaurus are reexamined and incorporated into a phylogenetic analysis. Milosaurus is resolved within Haptodontiformes, sharing with this clade a posterodorsally expanded ischium and a calcaneum with greater length than width. The more plesiomorphic ischium indicates a basal position within this clade. Most of the referred material shows very little overlap with the holotype and so most was not included in the cladistic analysis. However, what was originally described as a dorsal rib is here judged to be a femur that shares a highly distinctive morphology with the holotype. With its mass estimated as 41 kg, Milosaurus represents one of the largest Carboniferous synapsids. Large size evolved at least twice independently in Haptodontiformes during the Pennsylvanian: once in Sphenacodontidae and once in Edaphosauridae. Using phylogenetic comparative methods, Milosaurus is found to have evolved a large size independently of these two clades. As one of the largest Carboniferous amniotes, and the outgroup to one of the most diverse Paleozoic synapsid lineages, Milosaurus represents a crucial taxon for understanding early terrestrial ecosystems.

A Zombie LIF Gene in Elephants Is Upregulated by TP53 to Induce Apoptosis in Response to DNA Damage

Large-bodied organisms have more cells that can potentially turn cancerous than small-bodied organisms, imposing an increased risk of developing cancer. This expectation predicts a positive correlation between body size and cancer risk; however, there is no correlation between body size and cancer risk across species ("Peto's paradox"). Here, we show that elephants and their extinct relatives (proboscideans) may have resolved Peto's paradox in part through refunctionalizing a leukemia inhibitory factor pseudogene (LIF6) with pro-apoptotic functions. LIF6 is transcriptionally upregulated by TP53 in response to DNA damage and translocates to the mitochondria where it induces apoptosis. Phylogenetic analyses of living and extinct proboscidean LIF6 genes indicates that its TP53 response element evolved coincident with the evolution of large body sizes in the proboscidean stem lineage. These results suggest that refunctionalizing of a pro-apoptotic LIF pseudogene may have been permissive (although not sufficient) for the evolution of large body sizes in proboscideans.

Niche modeling reveals lack of broad-scale habitat partitioning in extinct horses of North America

The classic traits of modern horses (Equidae) – large body size, high-crowned teeth (hypsodonty), and a single toe (monodactyly) – are often considered adaptations to grassland environments. However, extinct horses that varied in these three traits overlapped geographically for millions of years during the Miocene and Pliocene. It has been hypothesized that co-occurring horse species partitioned habitats, with large-bodied, hypsodont, and monodactyl equids dominating open grasslands, while equids with different combinations of traits lived in more wooded areas. We tested for the presence of broad-scale habitat partitioning by compiling a large database of North American horse fossil occurrences with data on trait state (body size, hypsodonty index, and toe number) and paleoenvironment (derived from paleovegetation records). Null modeling of niche overlap in each of the North American Land Mammal Ages of the Miocene and Pliocene revealed that taxonomic and trait-based groups show no differences in habitat occupancy. Cluster dendrograms visualizing niche overlap showed that some ecological guilds shared derived traits, but derived traits were not associated exclusively with grassland habitats. Trait values are not predicted by the proportion of grassland habitats in time bins. Further, the three traits show no co-evolution across the equid tree when corrected for phylogenetic relatedness. Together, these results suggest that the evolution of large body size, hypsodonty, and monodactyly in equids was not due to a shared selective regime in response to expanding grassland habitats; instead, these traits may have evolved separately, likely due to a variety of small-scale selective pressures acting across the variety of habitats present in the Miocene and Pliocene.

A Paleocene penguin from New Zealand substantiates multiple origins of gigantism in fossil Sphenisciformes

One of the notable features of penguin evolution is the occurrence of very large species in the early Cenozoic, whose body size greatly exceeded that of the largest extant penguins. Here we describe a new giant species from the late Paleocene of New Zealand that documents the very early evolution of large body size in penguins. Kumimanu biceae, n. gen. et sp. is larger than all other fossil penguins that have substantial skeletal portions preserved. Several plesiomorphic features place the new species outside a clade including all post-Paleocene giant penguins. It is phylogenetically separated from giant Eocene and Oligocene penguin species by various smaller taxa, which indicates multiple origins of giant size in penguin evolution. That a penguin rivaling the largest previously known species existed in the Paleocene suggests that gigantism in penguins arose shortly after these birds became flightless divers. Our study therefore strengthens previous suggestions that the absence of very large penguins today is likely due to the Oligo-Miocene radiation of marine mammals.

TP53 copy number expansion is associated with the evolution of increased body size and an enhanced DNA damage response in elephants.

A major constraint on the evolution of large body sizes in animals is an increased risk of developing cancer. There is no correlation, however, between body size and cancer risk. This lack of correlation is often referred to as 'Peto's Paradox'. Here, we show that the elephant genome encodes 20 copies of the tumor suppressor gene TP53 and that the increase in TP53 copy number occurred coincident with the evolution of large body sizes, the evolution of extreme sensitivity to genotoxic stress, and a hyperactive TP53 signaling pathway in the elephant (Proboscidean) lineage. Furthermore, we show that several of the TP53 retrogenes (TP53RTGs) are transcribed and likely translated. While TP53RTGs do not appear to directly function as transcription factors, they do contribute to the enhanced sensitivity of elephant cells to DNA damage and the induction of apoptosis by regulating activity of the TP53 signaling pathway. These results suggest that an increase in the copy number of TP53 may have played a direct role in the evolution of very large body sizes and the resolution of Peto's paradox in Proboscideans.

Systematics of the Rubidgeinae (Therapsida: Gorgonopsia).

The subfamily Rubidgeinae, containing the largest known African gorgonopsians, is thoroughly revised. Rubidgeinae is diagnosed by the absence of a blade-like parasphenoid rostrum and reduction or absence of the preparietal. Seven rubidgeine species from the Karoo Basin of South Africa are recognized as valid: Aelurognathus tigriceps, Clelandina rubidgei, Dinogorgon rubidgei, Leontosaurus vanderhorsti, Rubidgea atrox, Smilesaurus ferox, and Sycosaurus laticeps. Rubidgeines are also present in other African basins: A. tigriceps and S. laticeps occur in the Upper Madumabisa Mudstone Formation of Zambia, and D. rubidgei, R. atrox, and the endemic species Ruhuhucerberus haughtoni comb. nov. and Sycosaurus nowaki comb. nov. occur in the Usili Formation of Tanzania. Aelurognathus nyasaensis from the Chiweta Beds of Malawi also represents a rubidgeine, but of uncertain generic referral pending further preparation. No rubidgeine material is known outside of Africa: the purported Russian rubidgeine Leogorgon klimovensis is not clearly referable to this group and may not be diagnosable. Phylogenetic analysis of rubidgeines reveals strong support for a clade (Rubidgeini) of advanced rubidgeines including Clelandina, Dinogorgon, Leontosaurus, and Rubidgea. Support for Smilesaurus as a rubidgeine is weak; it may, as previous authors have suggested, represent an independent evolution of large body size from an Arctops-like ancestor. Temporally, rubidgeines are restricted to the Late Permian, first appearing in the Tropidostoma Assemblage Zone and reaching highest diversity in the Cistecephalus and Daptocephalus assemblage zones of the Beaufort Group.

Material affects attack rates on dummy caterpillars in tropical forest where arthropod predators dominate: an experiment using clay and dough dummies with green colourants on various plant species

AbstractPredation can be one of the key factors that determine abundance in insect herbivore communities, and drive evolution of body size, and anti‐predator traits, including crypsis. Population dynamics and selection pressures will depend on the identity of dominant predators in the system, and these may vary substantially among habitats. Arthropods emerge as chief predators on caterpillars in the understorey of non‐montane tropical forest, whereas birds dominate elsewhere. In a tropical forest in Uganda, Africa, we evaluated marks on dummy caterpillars that differed in size, material (clay vs. dough), colourant, and plant species on which dummy caterpillars were exposed. We included live caterpillars to estimate the extent to which studies using artificial caterpillars reflect actual levels of predation. Ants and wasps were the most important damagers of dummy caterpillars, whereas bug and beetle damage was very rare, and no bird or small mammal damage was observed. Daily attack rates did not differ significantly from apparent mortality of live caterpillars (daily mortality = 12.1%), but dummy caterpillars made from dough were attacked more frequently (daily attack rate = 18.4%) than those from clay (daily attack rate = 6.9%). Caterpillars of different colour and size, and caterpillars exposed on different plant species had the same chances to be predated. This is in contrast to results from temperate area studies where birds dominate and are not affected by dummy caterpillar material, but prefer larger caterpillars. Our results are consistent with dominant predators on tropical forest caterpillars being invertebrates that are more chemically than visually oriented, so that: (1) material used for dummy caterpillars is important, (2) background matching is relatively unimportant, and (3) being large may have less of a cost. These patterns in predation might facilitate polyphagy and evolution of large body size in tropical Lepidoptera.