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Behavioural evolution in penguins does not reflect phylogeny
Abstract
Over the past two decades, behavioural biologists and ecologists have made effective use of the comparative method, but have often stopped short of adopting an explicitly phylogenetic approach. We examined 68 behaviour and life history (BLH) traits of 15 penguin species to: (i) infer penguin phylogeny, (ii) assess homology of behavioural characters, and (iii) evaluate hypotheses about character evolution and ancestral states. Parsimony analysis of the BLH dataset found either two shortest trees (characters coded as unordered) or a single shortest tree (characters coded as a combination of unordered and Dollo). The BLH data had significant structure. Kishino–Hasegawa tests indicated that BLH trees were significantly different from most previous estimates of penguin phylogeny. The BLH phylogeny generated from Dollo characters appeared to be less accurate than the tree derived from the completely unordered dataset. Dividing BLH data into display and non-display traits resulted in no significant differences in level of homoplasy and no difference in the accuracy of phylogeny. Tests for homology of BLH traits were performed by mapping the characters onto a molecular tree. Assuming that independent gains are less likely than losses of character states, 65 of the 68 characters were likely to be homologous across taxa, and at least several characters appeared to have been stable since the origin of modern penguins around 30 Myr. Finally, the likely BLH traits of the most recent common ancestor of extant penguins were reconstructed from character states along the internal branch leading to the penguins. This analysis suggested that the “proto-penguin” probably had a similar life history to current temperate penguins but few ritualized behaviours. A southern, cool-temperate origin of penguins is suggested.
... Penguins are a monophyletic group in which many species are found breeding sympatrically at several sites in the sub-Antarctic islands (Forcada, Trathan, Reid, Murphy, & Croxall, 2006;Lynch, Fagan, Naveen, Trivelpiece, & Trivelpiece, 2012;Niemandt et al., 2016;Paterson, Wallis, Kennedy, & Gray, 2014;Trathan, Croxall, & Murphy, 1996). The co-occurrence of these closely related species has long raised questions regarding what degree of competition takes place and how these species successfully coexist (Lynnes, Reid, Croxall, & Trathan, 2002;Trivelpiece & Volkman, 1979;White & Conroy, 1975). ...
Penguins are a monophyletic group in which many species are found breeding sympatrically, raising questions regarding how these species coexist successfully. Here, the isotopic niche of three sympatric pygoscelid penguin species was investigated at Powell Island, South Orkney Islands, during two breeding seasons (austral summers 2013–2014 and 2015–2016). Measurements of carbon (δ13C) and nitrogen (δ15N) stable isotope ratios were obtained from blood (adults) or feather (chicks) samples collected from Adélie Pygoscelis adeliae, chinstrap P. antarctica, and gentoo P. papua penguins. Isotopic niche regions (a proxy for the realized trophic niches) were computed to provide estimates of the trophic niche width of the studied species during the breeding season. The isotopic niche regions of adults of all three species were similar, but gentoo chicks had noticeably wider isotopic niches than the chicks of the other two species. Moderate to strong overlap in isotopic niche among species was found during each breeding season and for both age groups, suggesting that the potential for competition for shared food sources was similar during the two study years, although the actual level of competition could not be determined owing to the lack of data on resource abundance. Clear interannual shifts in isotopic niche were seen in all three species, though of lower amplitude for adult chinstrap penguins. These shifts were due to variation in carbon, but not nitrogen, isotopic ratios, which could indicate either a change in isotopic signature of their prey or a switch to an alternative food web. The main conclusions of this study are that (1) there is a partial overlap in the isotopic niches of these three congeneric species and that (2) they responded similarly to changes that likely occurred at the base of their food chain between the 2 years of the study.
... However, in at least some cases behavioral characters exhibit analytical rates of homoplasy not statistically different from those generated from morphological datasets (de Quieroz and Wimberger 1993), and metrics that measure phylogenetic signal have found utility even in demonstrably labile behavioral datasets (Blomberg et al. 2003). Whether used as prima facie evidence in phylogenetic inference or not, various methods, including parsimony optimization, have been applied to study the evolution of behavioral characters in a phylogenetic context (e.g., Tullberg et al. 2002;Agnarsson et al. 2006;Paterson et al. 2014;Odom et al. 2015). One such method, the extant phylogenetic bracket (Witmer 1995), is especially useful when fossil taxa for which soft-tissue, physiological, life-history, and behavioral features are generally not directly preserved. ...
Actualistic observations form the basis of many taphonomic studies in paleontology. However, surveys limited by environment or taxon may not be applicable far beyond the bounds of the initial observations. Even when multiple studies exploring the potential variety within a taphonomic process exist, quantitative methods for comparing these datasets in order to identify larger scale patterns have been understudied. This research uses modern bite marks collected from 21 of the 23 generally recognized species of extant Crocodylia to explore statistical and phylogenetic methods of synthesizing taphonomic datasets. Bite marks were identified, and specimens were then coded for presence or absence of different mark morphotypes. Attempts to find statistical correlation between trace types, marking animal vital statistics, and sample collection protocol were unsuccessful. Mapping bite mark character states on a eusuchian phylogeny successfully predicted the presence of known diagnostic, bisected marks in extinct taxa. Predictions for clades that may have created multiple subscores, striated marks, and extensive crushing were also generated. Inclusion of fossil bite marks which have been positively associated with extinct species allow this method to be projected beyond the crown group. The results of this study indicate that phylogenies can and should be further explored for use as predictive tools in a taphonomic framework.
... A substantial literature exists on the co-evolution of the phylogenies of seabird ectoparasites and those of their hosts (e.g. Paterson, Gray & Wallis, 1993;Paterson, Wallis & Gray, 1995), with some authors going so far as to say the correlation is more relevant than behaviour (Paterson et al., 2014). Grey-faced and white-headed petrels are recorded hosts for six species of mallophagid, whereas just five are recorded for great-winged petrel (Table S3). ...
Although described as a distinct species in 1869, for more than a century now New Zealand's grey-faced petrel (Pterodroma macroptera gouldi Hutton, 1869) has been regarded as a subspecies of the great-winged petrel (P. macroptera A. Smith, 1840). However, several authors have recently questioned whether the taxon once again deserves full species status. Here, we demonstrate that the grey-faced petrel is sufficiently distinct in multiple facets of its biology (including mitochondrial DNA, plumage variation, morphometrics, osteology, vocalizations, external parasites, and feeding and breeding biology) to warrant reinstatement of full species status under the morphological, phylogenetic, and potentially biological species concepts. Moreover, we present new evidence from mitochondrial DNA (cytochrome c oxidase subunit 1 and cytochrome b) that suggests the great-winged petrel is actually more closely related to the white-headed petrel (Pterodroma lessonii Garnot, 1826) than to the grey-faced petrel. The reclassification of grey-faced petrel to full species status raises the degree of seabird endemism in New Zealand to 43%, emphasising the status of the archipelago as a hotspot for seabird diversity.
... This reluctance to shift motor strategy even when an alternative could provide a better outcome seems a barrier to our understanding of how behavior patterns evolve. Behavioral solutions developed in the past seem to provide inertia against change, with solutions from ancestors having resonance in the descendents (e.g., Henzi & Barrett, 2005; Thierry, 2007), although care must be taken in not assuming the same degree of conservatism in all behaviors and all lineages (Patterson, Wallis, Kennedy, & Gray, 2013). The problem of concern in this chapter is that of how biases, once they become embedded as part of the rule structure that organizes behavior, can be overcome. ...
Even though behavior tends to occur as a continuous stream of action in most situations, there are discernible regularities that lead observers to label and measure discrete “behavior patterns.” Yet such discrete actions may not be how the nervous system organizes motor output. Sometimes, the perceived regularities are emergent properties of the interaction among the brain, the body within which that brain is situated, and the context within which the action takes place. Moreover, the brain's contribution may be limited to providing a perceptual rule, where some feature of the environment is kept constant. However, a problem arises when more than one behavior can solve a perceptual problem. It is theorized that, in such situations, the nervous system has a built-in bias to preferentially use one behavioral solution over another. Some possible motor biases are explored. How regularity in motor output is achieved greatly constrains how behavior can evolve.
Here, we propose, prove mathematically and discuss maximum and minimum measures of maximum parsimony evolution across 12 discrete phylogenetic character types, classified across 4467 morphological and molecular datasets. Covered character types are: constant, binary symmetric, multistate unordered (non‐additive) symmetric, multistate linear ordered symmetric, multistate non‐linear ordered symmetric, binary irreversible, multistate irreversible, binary Dollo, multistate Dollo, multistate custom symmetric, binary custom asymmetric and multistate custom asymmetric characters. We summarize published solutions and provide and prove a range of new formulae for the algebraic calculation of minimum ( m ), maximum ( g ) and maximum possible ( g max ) character cost for applicable character types. Algorithms for exhaustive calculation of m , g and g max applicable to all classified character types (within computational limits on the numbers of taxa and states) are also provided. The general algorithmic solution for minimum steps ( m ) is identical to a minimum spanning tree on the state graph or minimum weight spanning arborescence on the state digraph. Algorithmic solutions for character g and g max are based on matrix mathematics equivalent to optimization on the star tree, respectively for given state frequencies and all possible state frequencies meeting specified numbers of taxa and states. We show that maximizing possible cost ( g max ) with given transition costs can be equivalent to maximizing, across all possible state frequency combinations, the lowest implied cost of state transitions if any one state is ancestral on the star tree, via the solution of systems of linear equations. The methods we present, implemented in the Claddis R package, extend to a comprehensive range, the fundamental character types for which homoplasy may be measured under parsimony using m , g and g max , including extra cost ( h ), consistency index ( ci ), retention index ( ri ) or indices based thereon.
The phylogenetic relationships among the four tribes of corbiculate bees (Euglossini, Bombini, Meliponini, and Apini) are controversial. There is substantial incongruence between morphological and molecular data, and the single origin of eusociality is questionable. The use of behavioral characters by previous workers has been restricted to some typological definitions, such as solitary and eusocial. Here, I expand the term "social" to 42 characters and present a tree based only on behavioral characters. The reconstructed relationships were similar to those observed in morphological and "total evidence" analyses, i.e., Euglossini + (Bombini + (Meliponini + Apini)), all of which support a single origin of eusociality. (C) 2002 The Willi Hennig Society.
Abstract The elaborate songs of songbirds are frequent models for investigating the evolution of animal signals. However, few previous studies have attempted to reconstruct historical changes in song evolution using a phylogenetic comparative approach. In particular, no comparative studies of bird song have used a large number of vocal characters and a well-supported, independently derived phylogeny. We identified 32 features in the complex vocal displays of male oropendolas (genera Psarocolius, Gymnostinops, and Ocyalus) that are relatively invariant within taxa and mapped these characters onto a robust molecular phylogeny of the group. Our analysis revealed that many aspects of oropendola song are surprisingly evolutionarily conservative and thus are potentially useful characters for reconstructing historical patterns. Of the characters that varied among taxa, nearly two thirds (19 of 29) showed no evidence of evolutionary convergence or reversal when mapped onto the tree, which was reflected in a high overall consistency index (CI = 0.78) and retention index (RI = 0.88). Some reconstructed patterns provided evidence of selection on these signals. For example, rapid divergence of the songs of the Montezuma oropendola, Gymnostinops montezuma, from those of closely related taxa suggests the recent influence of strong sexual selection. In general, our results provide insights into the mode of vocal evolution in songbirds and suggest that complex vocalizations can provide information about phylogeny. Based on this evidence, we use song characters to estimate the phylogenetic affinities of three oropendola taxa for which molecular data are not yet available.
Absolute criteria for evaluating cladistic analyses are useful, not only because cladistic algorithms impose structure, but also because applications of cladistic results demand some assessment of the degree of corroboration of the cladogram. Here, a means of quantitative evaluation is presented based on tree length. The length of the most-parsimonious tree reflects the degree to which the observed characters co-vary such that a single tree topology can explain shared character states among the taxa. This "cladistic covariation" can be quantified by comparing the length of the most parsimonious tree for the observed data set to that found for data sets with random covariation of characters. A random data set is defined as one in which the original number of characters and their character states are maintained, but for each character, the states are randomly reassigned to the taxa. The cladistic permutation tail probability, PTP, is defined as the estimate of the proportion of times that a tree can be found as short or shorter than the original tree. Significant cladistic covariation exists if the PTP is less than a prescribed value, for example, 0.05. In case studies based on molecular and morphological data sets, application of the PTP shows that: 1. (1) In the comparison of four different molecular data sets for orders of mammals, the sequence data set for alpha hemoglobin does not have significant cladistic covariation, while that for alpha crystallin is highly significant. However, when each data set was reduced to the 11 common taxa in order to standardize comparison, reduced levels of cladistic covariation, with no clear superiority of the alpha crystallin data, were found. Morphological data for these 11 taxa had a highly significant PTP, producing a tree roughly congruent with those for the three molecular sets with marginal or significant PTP values. Merging of all data sets, with the exclusion of the poorly structured alpha hemoglobin data, produced a data set with a significant PTP, and provides an estimate of the phylogenetic relationships among these 11 orders of mammals. 2. (2) In an analysis of lactalbumin and lysozyme DNA sequence data for four taxa, purine-pyrimidine coding yields a data set with significant cladistic covariation, while other codings fail. The data for codon position 3 taken alone exhibit the strongest cladistic covariation. 3. (3) A data set based on flavonoids in taxa of Polygonum initially yields a significant PTP; however, deletion of identically scored taxa leaves no significant cladistic covariation. 4. (4) For mitochondrial DNA data on population genome types for four species of the crested newt, there is significant cladistic covariation for the set of all genome types, and among the five mtDNA genome types within one of the species. However, a conditional PTP test that assumes species monophyly shows that no significant cladistic covariation exists among the fur species for these data. 5. (5) In an application of the test to a group of freshwater insects, as preliminary to biological monitoring, individual subsets of the taxonomic data representing larval, pupal, and adult stages had non-significant PTPs, while the complete data set showed significant cladistic structure.
Attempts to identify traits that might be considered convergent adaptations to life in the marine environment and to show how divergent life history patterns are optimal for different reasons. Seabirds invest a greater total amount of energy and protein into the offspring than pinnipeds, but this comes at the cost of making more trips to sea. Pinnipeds forage in a manner more consistent with the predictions of central place foraging theory and exhibit a greater ability to compensate to the shortened breeding season typical of high latitude environments. -from Author
ABsTRACT.--Phylogenetic patterns can be used to generate detailed historical hypotheses about the evolution of character systems, including parental-care behavior. Phylogenetic analysis of 60 taxa using parental-care data (15 characters) in combination with anatomical data (69 characters) shows that biparental care is primitive for birds, and that biparental incubation arose from an ancestral condition in which neither parent incubated. Components of parental care may be decoupled, such that incubation and feeding of nestlings is biparental, but postfledging care is uniparental; the pattern of decoupling varies across taxa. Behavioral plasticity, environmentally induced variation, and quantitative variation in characters that are coded qualitatively are some of the problems inherent in such an analysis. Nevertheless, historical analysis offers a good vehicle for obtaining rigorous hypotheses about the coevo-lution of avian character systems. Received 5 July 1991, accepted 29 October 1991. THE USE of phylogenetic pattern to elucidate questions about other evolutionary and ecolog-ical patterns or processes has become increas-ingly common in the last 15 years. Several rig-orous attempts to incorporate behavioral data into phylogenetic analyses or to compare such data with existing phylogenies have been made in the last five years. Clutton-Brock and Harvey (1977) recognized the pitfalls of comparing closely related species to study the origin of ecological phenomena, and this has been fur-ther emphasized by Clutton- outlined a method for using cladistic hypotheses of rela-tionship to test hypotheses of adaptation, and this kind of approach has been fruitfully ap-plied in a number of recent studies (Prum 1990, B. N. Danforth unpubl. data, R. J. Smith unpubl.
We reviewed the tremendous architectural diversity of ovenbird (Furnariidae) nests based on literature, museum collections, and new field observations. With few exceptions, furnariids exhibited low intraspecific variation for the nest characters hypothesized, with the majority of variation being hierarchically distributed among taxa. We hypothesized nest homologies for 168 species in 41 genera (ca. 70% of all species and genera) and coded them as 24 derived characters. Forty-eight most-parsimonious trees (41 steps, CI = 0.98, RC = 0.97) resulted from a parsimony analysis of the equally weighted characters using PAUP, with the Dendrocolaptidae and Formicarioidea as successive outgroups. The strict-consensus topology based on these trees contained 15 clades representing both traditional taxa and novel phylogenetic groupings. Comparisons with the outgroups demonstrate that cavity nesting is plesiomorphic to the furnariids. In the two lineages where the primitive cavity nest has been lost, novel nest structures have evolved to enclose the nest contents: the clay oven of Furnarius and the domed vegetative nest of the synallaxine clade. Although our phylogenetic hypothesis should be considered as a heuristic prediction to be tested subsequently by additional character evidence, this first cladistic analysis of the furnariids demonstrates the general utility of nest characters in reconstruction of avian relationships, and it provides a test of monophyly for several furnariid taxa.
Croizat’s radical attack on traditional modes of biological inquiry has produced strongly polarised responses. Instead of arguing for or against the panbiogeographic approach I attempt to analyse some basic assumptions shared by both its defenders and critics. Although their emphasis is quite different both sides rely on two central oppositions. In evolutionary arguments selective explanations are opposed to those emphasising phylogenetic constraint (orthogenesis). In biogeographic arguments ecological explanations are opposed to historical explanations. I discuss how the legendary opposition between nature and nurture was resolved by reformulating the way in which causation was viewed. I suggest that the selection/constraint opposition shares many features in common with the nature/nurture opposition and argue that it can be resolved in a similar manner. Conflicts between ecological and historical explanations in biogeography are explored by contrasting Diamond’s analysis of New Guinea bird distributions with that of Croizat’s. By again drawing on the resolution of the nature/nurture dispute a way of synthesising ecological and historical explanations is outlined.
Recent literature suggests that a phylogenetic orientation may lead to unique perspectives on homology. An analysis of absence of avian hind limb flexors is used to explore this perspective. This approach suggests that synapomorphy is not equivalent to homology but rather that synapomorphies are hypotheses of homology (otherwise phylogenetic analysis is tautological). Homologies should be viewed as relationships among historical individuals, and we can only discover interorganismic homologies by phylogenetic analysis. Intraorganismic homologies, however, must be studied by other methods, primarily ontogenetic ones. Viewing homologies as (1) genetically and developmentally individualized entities or (2) sets of historical individuals may not always be particularly helpful from an operational perspective, but this is irrelevant because homologies are real entities that must be discovered, yet the process by which they are sought, as in all science, is one that does not yield truths, only hypotheses.
This article considers the statistical issues relevant to the comparative method in evolutionary biology. A generalized Linear model (GLM) is presented for the analysis of comparative data, which can be used to address questions regarding the relationship between traits or between traits and environments, the rate of phenotypic evolution, the degree of phylogenetic effect, and the ancestral state of a character. Our approach thus emphasizes the similarity among evolutionary questions asked in comparative studies. We then discuss ways of specifying the sources of error involved in a comparative study (e.g., measurement error, error due to evolution along a phylogeny, error due to misspecification of a phylogeny) and show how the impact of these sources of error can be taken into account in a comparative analysis. In contrast to most existing phylogenetic comparative methods, our procedure offers substantial flexibility in the choice of microevolutionary assumptions underlying the statistical analysis, allowing researchers to choose assumptions that are most appropriate for their particular set of data and evolutionary question. In developing the approach, we also propose novel ways of incorporating within-species variation and/or measurement error into phylogenetic analyses, of estimating ancestral states, and of considering both continuous (quantitative) and categorical (qualitative or ''state'') characters in the same analysis.
It is widely believed that behavior is more evolutionarily labile and/or more difficult to characterize than morphology, and thus that behavioral characters are not as useful as morphological characters for estimating phylogenetic relationships. To examine the relative utility of behavior and morphology for estimating phylogeny, we compared levels of homoplasy for morphological and behavioral characters that have been used in systematic studies. In an analysis of 22 data sets that contained both morphological and behavioral characters we found no significant difference between mean consistency indices (CIs, which measure homoplasy) within data sets for the two types of characters. In a second analysis we compared overall CIs for 8 data sets comprised entirely of behavioral characters with overall CIs for 32 morphological data sets and found no significant difference between the two types of data sets. For both analyses, 95% confidence limits on the difference between the two types of characters indicate that, even if given the benefit of the doubt, morphological characters could not have substantially higher mean CIs than behavioral characters. These results do not support the idea that behavioral characters are less useful than morphological characters for the estimation of phylogeny.
Whether or not behavior accurately reflects evolutionary relationships (phylogeny) has been hotly debated by ethologists and comparative psychologists. Previous studies attempting to resolve this question have generally lacked a quantitative, phylogenetic approach. In this study we used behavior and life-history (BLH) information (72 characters) to generate phylogenetic trees for 18 seabird species (albatrosses, petrels, and penguins). We compared these trees with trees obtained from isozyme electrophoretic analysis of blood proteins (15 loci and 98 electromorphs) and partial mitochondrial 12S ribosomal DNA sequences (381 base pairs). Cladistic analysis of the BLH data set generated three MP trees (tree length = 243, CI = 0.52, RI = 0.57) with significant cladistic structure. The BLH characters were classified into four types (foraging, agonistic, reproductive, and life history) and levels of homoplasy for each type were measured. No significant differences were found among these categories. The BLH trees were shown to be significantly more congruent with the electrophoretic and 12S sequence trees than expected by chance. This indicates that seabird BLH data contains phylogenetic signal. Areas of incongruence between BLH trees and a phylogeny generated by combining the data sets were predicted to result from ecological constraints that did not covary with phylogeny. These predictions were supported by the results of a concentrated changes test. This study found that this BLH data set was no more homoplasious than molecular data and that BLH trees were significantly congruent with molecular trees.
Phylogenetic systematics offers ethologists a method for uncovering macroevolutionary patterns of behavioral diversity, which can be used to generate predictions for experimental investigation of microevolutionary processes. I used phylogenetic methods to disentangle the contributions of various selection pressures in the evolution of sexually dimorphic nuptial coloration in gasterosteid fishes in general, and Gasterosteus aculeatus in particular. Based upon the phylogenetic relationships between breeding colors and breeding behaviors in the gasterosteids, three predictions relevant to a discussion of nuptial coloration in G. aculeatus were proposed. The origin and elaboration of nuptial coloration was (1) weakly associated with male/male interactions (intrasexual selection), (2) moderately associated with parental care (natural selection) and (3) strongly associated with male/female interactions (intersexual selection). I tested the hypothesized macroevolutionary relationships between male breeding color and behavior in two sets of experiments studying changes in male nuptial coloration across the breeding cycle and female choice based on intensity of male color. The results of these experiments at the microevolutionary level of analysis corroborated the macroevolutionary predictions and confirmed the original intuition of previous researchers: male body color intensity is an important component of female mate choice in G. aculeatus. This study demonstrates that a phylogenetic hypothesis can generate predictions that are experimentally testable and falsifiable.
Theories of ecological diversification make predictions about the timing and ordering of character state changes through history. These theories are testable by "reconstructing" ancestor states using phylogenetic trees and measurements of contemporary species. Here we use maximum likelihood to estimate and evaluate the accuracy of ancestor reconstructions. We present likelihoods of discrete ancestor states and derive probability distributions for continuous ancestral traits. The methods are applied to several examples: diets of ancestral Darwin's finches; origin of inquilinism in gall wasps; microhabitat partitioning and body size evolution in scrubwrens; digestive enzyme evolution in artiodactyl mammals; origin of a sexually selected male trait, the sword, in platies and swordtails; and evolution of specialization in Anolis lizards. When changes between discrete character states are rare, the maximum-likelihood results are similar to parsimony estimates. In this case the accuracy of estimates is often high, with the exception of some nodes deep in the tree. If change is frequent then reconstructions are highly uncertain, especially of distant ancestors. Ancestor states for continuous traits are typically highly uncertain. We conclude that measures of uncertainty are useful and should always be provided, despite simplistic assumptions about the probabilistic models that underlie them. If uncertainty is too high, reconstruction should be abandoned in favor of approaches that fit different models of trait evolution to species data and phylogenetic trees, taking into account the range of ancestor states permitted by the data.
Ethological studies in the 1940s and 1950s, most notably those of Lorenz and Tinbergen, emphasized a historical perspective. By the 1970s, the notion that behavioral traits are too plastic to retain historical information became prevalent, and evolutionary approaches in behavioral studies were largely abandoned. However, several recent studies have demonstrated that behavioral characters are remarkably consistent with phylogenies obtained from other data and not particularly prone to homoplasy. In this study, I coded descriptions of courtship display behaviors in stork species (Aves: Ciconiiformes: Ciconiidae) as a matrix of discrete characters. I mapped each behavioral character onto a phylogeny based on DNA-DNA hybridization distances to test the homology of individual characters. Generally, displays occurring early in courtship were congruent with phylogenetic relationships and showed little homoplasy, while displays occurring late in courtship were more homoplastic. I also performed a phylogenetic analysis of the behavioral data matrix using maximum parsimony. The strict consensus of the 24 most-parsimonious trees was congruent with the DNA-DNA hybridization tree in all nodes having greater than 70% bootstrap support.
A homolog is a part of the phenotype that is homologous to equivalent parts in other species. A biological homology concept is expected to explain three properties of homologs: 1) the conservation of those features that are used to define a homolog, 2) the individualization of the homolog with regard to the rest of the body, and 3) the uniqueness of homologs, i.e., their specificity for monophyletic groups. The main obstacle to describing a mechanistic basis for homology is the variability of the developmental pathways of undoubtedly homologous characters. However, not all aspects of the developmental pathway are of equal importance. The only organizational features of the developmental system that matter are those that have been historically acquired and cause developmental constraints on the further evolutionary modification of the characters. Two main factors contribute to historically acquired developmental constraints: generative rules of pattern formation and ontogenetic networks. In particular, hierarchical and cyclical inductive networks have the required properties to explain homology. How common such networks are is an open empirical question. The development and variation of pectoral fin hooks in blenniid fishes is presented as a model for the study of a simple ontogenetic network.
Although traits of related species are likely to be similar due to common ancestry, mating signals are an exception. In singing insects, for example, song similarity has been documented only for allopatric or allochronic species pairs, and even then, not often. Where song similarity does occur, it has been logically attributed to the inheritance of ancestral traits rather than convergence. It is quite common for related, sympatric insect species to differ dramatically in calling song, which is predicted by evolutionary theory to maximize intraspecific mating success. Given that there are a limited number of ways to make sounds on anatomically similar organs and given that there would be no selective pressure for songs to differ in widely separated geographic areas, convergence in songs among related species living on different continents might be expected. Here we present the first well-documented case of such convergence, in a group of sibling, cryptic species characterized by substrate-borne vibrational mating songs. In this example from green lacewings of the carnea group of the genus Chrysoperla, a variety of statistical tests shows that one species in North America and another in Asia possess songs that are strikingly similar to each other. DNA data demonstrate that the species involved belong to divergent speciose lineages, and behavioral data demonstrate that the convergent songs are readily accepted by members of both species.
The narrow-gutted but lofty archipelago of New Zealand, consisting of two large and many smaller islands, rises from a system of relatively shallow submarine rises and plateaux between the South West Pacific Ocean and the Tasman Sea. The oceanic ridges, the basins and trenches that separate and flank them, and the chains of islands that surmount them are part of a complex system that extends north to Melanesia, thence westward to the Indonesian archipelago and northward to Japan and beyond as the geographic manifestation of the Circum-Pacific Mobile Belt, a zone in which geological processes and consequent geographic changes appear to have been exceptionally rapid throughout the span of geological time. The submarine rises (Fig. 1) represent vast areas for which geological knowledge is as yet extremely poor, but which cannot be ignored in interpreting the history of the region and have, indeed, been taken into account in most attempts to determine its geological history. Recent advances in marine geophysics have led to the widely supported theories of sea-floor spreading and plate-tectonics, and to a general acceptance of the concept of continental drift. Ideas of geological history have thus been polarised in terms of a unifying model, but in applying this model to the history of New Zealand there remain many degrees of freedom and no unique solution.
Although traits of related species are likely to be similar due to common ancestry, mating signals are an exception. In singing insects, for example, song similarity has been documented only for allopatric or allochronic species pairs, and even then, not often. Where song similarity does occur, it has been logically attributed to the inheritance of ancestral traits rather than convergence. It is quite common for related, sympatric insect species to differ dramatically in calling song, which is predicted by evolutionary theory to maximize intraspecific mating success. Given that there are a limited number of ways to make sounds on anatomically similar organs and given that there would be no selective pressure for songs to differ in widely separated geographic areas, convergence in songs among related species living on different continents might be expected. Here we present the first well-documented case of such convergence, in a group of sibling, cryptic species characterized by substrate-borne vibrational mating songs. In this example from green lacewings of the carnea group of the genus Chrysoperla, a variety of statistical tests shows that one species in North America and another in Asia possess songs that are strikingly similar to each other. DNA data demonstrate that the species involved belong to divergent speciose lineages, and behavioral data demonstrate that the convergent songs are readily accepted by members of both species.
The estimation procedure utilizes a compatibility analysis between enzyme data sets of the most parsimonious trees constructed from the restriction enzyme. Next, a non-parametric test is given for comparing alternative phylogenies. A 2nd non-parametric test is developed for testing the molecular clock hypothesis. To illustrate the power of these procedures, data derived from the mitochondrial DNA and globin DNA of man and the apes are analyzed. Although previous analyses of these data led to the speculation that 10 times more information would be required to resolve the evolutionary relationships between man with chimps and gorillas, this algorithm resolved these relationships at the 5% level of significance. The molecular clock hypothesis was rejected at the 1% level. The implications of this phylogenetic inference when coupled with other types of data lead to the conclusion that knuckle-walking - not bipedalism - is the evolutionary novelty in mode of locomotion in the primates and that many other hominid features are primitive whereas their African ape counterparts are derived.-from Author
We studied restriction-fragment length polymorphisms (RELPs) in mitochondrial DNA for 13 species of African francolins (Francolinus spp.) and the Japanese Quail (Coturnix c. japonica). Phylogenetic analyses of RFLPs for these 14 species and of morphological and behavioral characters for the 41 francolin species and other perdicine taxa do not confirm the monophyly of Francolinus as currently recognized. Analyses of morpho-behavioral characters suggest that Francolinus consists of at least four major assemblages: the five Asiatic species; two groups of African quail-like species; and the African partridge-like species. Within these assemblages, analyses of RFLPs and/or morpho-behavioral characters support the monophyly of six of eight species groups attributed to Francolinus. Assuming the monophyly of currently recognized supraspecific groups of galliform birds, morphometric analyses of galliform skeletons correctly classified 90-99% of specimens to family, subfamily and tribe, as well as 95% of the francolin specimens to genus. Genetic distances derived from RFLP data imply that African francolins diverged from their sister taxa at or before the mid-late Miocene, and that all species studied diverged from their sister-species during the Pliocene or early Pleistocene.
Ethological studies in the 1940s and 1950s, most notably those of Lorenz and Tinbergen, emphasized a historical perspective. By the 1970s, the notion that behavioral traits are too plastic to retain historical information became prevalent, and evolutionary approaches in behavioral studies were largely abandoned. However, several recent studies have demonstrated that behavioral characters are remarkably consistent with phylogenies obtained from other data and not particularly prone to homoplasy. In this study, I coded descriptions of courtship display behaviors in stork species (Aves: Ciconiiformes: Ciconiidae) as a matrix of discrete characters. I mapped each behavioral character onto a phylogeny based on DNA-DNA hybridization distances to test the homology of individual characters. Generally, displays occurring early in courtship were congruent with phylogenetic relationships and showed little homoplasy, while displays occurring late in courtship were more homoplastic. I also performed a phylogenetic analysis of the behavioral data matrix using maximum parsimony. The strict consensus of the 24 most-parsimonious trees was congruent with the DNA-DNA hybridization tree in all nodes having greater than 70% bootstrap support.
It is widely believed that behavior is more evolutionarily labile and/or more difficult to characterize than morphology, and thus that behavioral characters are not as useful as morphological characters for estimating phylogenetic relationships. To examine the relative utility of behavior and morphology for estimating phylogeny, we compared levels of homoplasy for morphological and behavioral characters that have been used in systematic studies. In an analysis of 22 data sets that contained both morphological and behavioral characters we found no significant difference between mean consistency indices (CIs, which measure homoplasy) within data sets for the two types of characters. In a second analysis we compared overall CIs for 8 data sets comprised entirely of behavioral characters with overall CIs for 32 morphological data sets and found no significant difference between the two types of data sets. For both analyses, 95% confidence limits on the difference between the two types of characters indicate that, even if given the benefit of the doubt, morphological characters could not have substantially higher mean CIs than behavioral characters. These results do not support the idea that behavioral characters are less useful than morphological characters for the estimation of phylogeny.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
Genealogical relationships among the five genera of stickleback (Gasterosteidae: Smegmomorpha) fishes were reconstructed based upon 47 behavioral and 89 morphological characters. Phylogenetic systematic analysis of the combined data set produced a single MPT (consistency index of 82.54%) structured (Spinachia (Apeltes-((Pungitius + Culaea)(Gasterosteus aculeatus + G. wheatlandi)))). When analyzed separately, both behavioral and morphological data sets produced the same tree topology, although the consistency index (excluding uninformative characters) was higher for the behavioral (90.70%) than for the morphological (68.13%) characters. This total evidence analysis provides a highly robust estimate of phylogeny for the sticklebacks and reiterates the importance of discarding a priori biases about behavioral evolution when selecting characters for phylogenetic analysis.
We performed a phylogenetic and biogeographic study of the riflebird genus Ptiloris (Paradisaeidae). Our analysis, which includes sonograms documenting intertaxonomic variation in male advertisement vocalizations and a reassessment of key morphological characters, indicates that (1) the genus includes two distinct clades (P. victoriae/paradiseus and the P. magnificus complex); (2) within the P. magnificus complex, the eastern Papuan form may constitute a heretofore unrecognized sibling species; and (3) the biogeography of Ptiloris provides additional confirming evidence that the Torres Strait has been of minor importance as a biogeographic barrier for Australo-Papuan rainforest birds.
This handbook provides identification information and illustrations for the birds of the Antarctic and sub-Antarctic and synopses) of their biology and distribution. It is intended to help scientists, ship's crewmen, and travelers identify most of the birds observed at sea and on land south of 55°S latitude and around the cold sub-Antarctic islands near the Antarctic convergence and the temper ate sub-Antarctic islands of the central Atlantic and Indian oceans. The handbook is divided into three sections. The in troduction discusses the antarctic and subantarctic en vironments, provides general information on how to iden tify and study birds, and discusses their zoogeography and conservation. The main body of the book consists of species accounts, illustrations, and distribution maps. The third section discusses the environment of each of the ma jor landmasses briefly and gives tabular checklists that show the bird observer what species to expect on the con tinent and various islands or in different sections of the ocean. Special features of this handbook are an extensive bibliography, an analysis of this bibliography by family group and geography, a variant name list that gives English and Latin synonyms as well as French and Spanish vernacular names, and twelve color plates.
"The merits of this work are many. A rigorous integration of phylogenetic hypotheses into studies of adaptation, adaptive radiation, and coevolution is absolutely necessary and can change dramatically our collective 'gestalt' about much in evolutionary biology. The authors advance and illustrate this thesis beautifully. The writing is often lucid, the examples are plentiful and diverse, and the juxtaposition of examples from different biological systems argues forcefully for the validity of the thesis. Many new insights are offered here, and the work is usually accessible to both the practiced phylogeneticist and the naive ecologist."—Joseph Travis, Florida State University
"[Phylogeny, Ecology, and Behavior] presents its arguments forcefully and cogently, with ample . . .support. Brooks and McLennan conclude as they began, with the comment that evolution is a result, not a process, and that it is the result of an interaction of a variety of processes, environmental and historical. Evolutionary explanations must consider all these components, else they are incomplete. As Darwin's explanations of descent with modification integrated genealogical and ecological information, so must workers now incorporate historical and nonhistorical, and biological and nonbiological, processes in their evolutionary perspective."—Marvalee H. Wake, Bioscience
Published claims that Macaroni Penguins Eudyptes chrysolophus and Rockhopper Penguins E. chrysocome lay clutches of one or three eggs are reviewed in the light of current knowledge of the breeding biology of these and other species of penguins. It is concluded that the clutch size of these Eudyptes penguins is invariably two eggs, that one-egg clutches are usually reported because the loss of the A egg was not noticed and that most three-egg clutches are the result of adoption of eggs.
We assessed the utility of congruence and multiple data sets to test species relationships and the accuracy of phylogenetic methods. The ongoing controversy about whether to combine data sets for phylogenetic analysis was evaluated against the naturalness of different types of data (as commonly recognized by systematists) and character independence. We defend the recommendation that independent data sets (defined in terms of process partitions; sensu bull et al., 1993, Syst. Biol. 42:384-397) should rarely be combined but should be kept separate for phylogenetic analysis because their independence increases the significance of corroboration. Trees of natural taxa, well supported by many independent lines of evidence, should be used in the same way as the known phylogenies of simulations and of certain laboratory and domesticated groups, i.e., as standards for evaluating the accuracy of different phylogenetic methods. Although compromised by their imperfect reliabilities, such tests using well-supported trees of wild taxa provide important reality checks on the conclusions of the other two approaches by encompassing more of the complexity and diversity of natural systems and their evolutionary processes. In this way, a combination of testing with the well-supported trees of natural groups, with simulations, and with those laboratory and domesticated taxa with known phylogenies is most likely to prove effective in establishing the strengths, weaknesses, and assumptions of different phylogenetic methods.
Whether or not behavior accurately reflects evolutionary relationships (phylogeny) has been hotly debated by ethologists and comparative psychologists. Previous studies attempting to resolve this question have generally lacked a quantitative, phylogenetic approach. In this study we used behavior and life-history (BLH) information (72 characters) to generate phylogenetic trees for 18 seabird species (albatrosses, petrels, and penguins). We compared these trees with trees obtained from isozyme electrophoretic analysis of blood proteins (15 loci and 98 electromorphs) and partial mitochondrial 12S ribosomal DNA sequences (381 base pairs). Cladistic analysis of the BLH data set generated three MP trees (tree length = 243, CI = 0.52, RI = 0.57) with significant cladistic structure. The BLH characters were classified into four types (foraging, agonistic, reproductive, and life history) and levels of homoplasy for each type were measured. No significant differences were found among these categories, The BLH trees were shown to be significantly more congruent with the electrophoretic and 12S sequence trees than expected by chance. This indicates that seabird BLH data contains phylogenetic signal. Areas of incongruence between BLH trees and a phylogeny generated by combining the data sets were predicted to result from ecological constraints that did not covary with phylogeny. These predictions were supported by the results of a concentrated changes test. This study found that this BLH data set was no more homoplasious than molecular data and that BLH trees were significantly congruent with molecular trees.
In recent years, two approaches have emerged for the analysis of character evolution: the largely statistical “convergence” approach and the mainly cladistic “homology” approach. I discuss the strengths and weaknesses of these approaches as they apply to phylogenetic analyses of life-history variation in birds. Using examples from analyses of character variation in swallows, I suggest that the phylogenetic approach yields distinctive insights into the selective role of the environment and other characters of the organism on the evolution of life-history traits. This view thus has the potential of bringing together micro- and macro-evolutionary views of life-history evolution.
Cave-dwelling Eudyptula minor nested in tight colonial aggregations and had no physical barriers between nest sites. Burrow-dwellers nested as solitary pairs or in loose aggregations and were physically isolated from one another. Cave-dwellers had significantly higher rates and complexity of agonistic interaction. Males were involved in a higher proportion of agonistic interactions than females. Burrow-dwellers used the overt behaviour Attack significantly more than cave-dwellers and also used the most dangerous fighting method more commonly than cave-dwellers. Burrow-dwellers also bit and fought significantly longer than cave-dwellers. An egg transfer experiment between cave and burrow colonies indicated that chicks may not be genetically bound to the use of a habitat-specific repertoire of agonistic behaviours. Plastic agonistic behaviour may allow immediate and potentially adaptive phenotypic change in response to environmental heterogeneity. The large cave repertoire may reduce the chances of any one interaction ending with overt aggression. The smaller burrow-dweller repertoire may be sufficient to defend the physical enclosed burrow nest sites. -from Author
Until the Early Cretaceous, when the first phases of rifting that preceded the eventual breakup of the Australasian sector of Gondwana occurred, New Zealand and Antarctica had a close geographic relationship on the eastern margin of Gondwana.
After having been influenced by proximity to the Permian ice sheet, during which New Zealand and Antarctica shared elements of the Glossopteris flora, the eastern margin of Gondwana entered into a phase of predominantly cool-temperate climates that lasted throughout the Triassic.
In the Jurassic, rotation of Gondwana had moved New Zealand and Antarctica into mid-latitudes and they experienced warm-temperate climates. These climatic conditions facilitated the dispersion of Tethyan marine faunas into the SW Pacific sector of eastern Gondwana. New Zealand and Antarctica shared many elements of the Tethyan marine faunas.
Starting in Middle Jurassic times, earth movements of the Rangitata Orogeny began to create considerable areas of land in the New Zealand region. Eventually, by Late Jurassic and Early Cretaceous times, a large landmass had been developed, extending northwards towards New Caledonia, eastwards to the Chatham Islands, westwards to the Lord Howe Rise and southwards to the edge of the Campbell Plateau. Creation of this landmass, and its associated land and marine links, together with the equable warm-temperate conditions then apparent over large areas of Gondwana, provided favourable conditions for the dispersal of Gondwana elements into New Zealand. Ancestral stocks of at least some of New Zealand’s archaic endemic terrestrial biota may have dispersed from eastern Gondwana into New Zealand at this time.
In the Early Cretaceous, land connections to eastern Gondwana became weakened by the onset of rifting along the future sites of the Tasman Sea and Southern Ocean. The eastern edge of Gondwana was rotated into high latitudes, and cool-temperate climates returned to New Zealand and Antarctica. Cool-temperate Austral marine faunas developed and were shared between New Zealand, Antarctica and southern South America. Land links between New Zealand and West Antarctica provided access for early angiosperms. However, all land links between New Zealand and Australia/Antarctica were broken after 85 Ma. From this time onwards the ancestral Tasman Sea and Southern Ocean became effective barriers to overland dispersal between eastern Gondwana and New Zealand. Subsequently, all terrestrial colonists had to arrive by either flying, swimming or floating. Many birds did so, but no terrestrial snakes and no mammals, except bats.
Although land connections had been broken in the Late Cretaceous, shallow-water marine links continued to exist between New Zealand and West Antarctica until the late Paleocene (the Weddellian Province). However, as New Zealand moved progressively northwards and the Southern Ocean widened, such southern links gave way in latest Paleocene and early Eocene times to northern (Australian and Malayo-Pacific) links.
Higher level phylogenetic analyses rarely include behavioural data, predominantly because such groups seldom have complex behaviours that are susceptible to analysis. Even when broad groups do share a complex behaviour, there is skepticism about the appropriateness of using behavioural traits in higher level phylogenetic analyses. The Integripalpia is a suborder of caddisflies and is an appropriate group to investigate the use of behaviour in higher level analyses because all larvae use a complex suite of behaviours to build portable cases. A thorough investigation of case-building in 10 families (19 exemplar genera) yielded 24 behavioural characters. A parsimony analysis produced 87 equally parsimonious trees (length = 56 steps, consistency index (CI) = 0.84, retention index (RI) = 0.88) that supported the monophyly of the integripalpian families, except for the Limnephilidae. Interfamilial relationships, although resolved, were not well supported with behaviour. Certain interfamilial relationships have also been difficult to establish reliably with morphological information, indicating a need for more characters (e.g., molecular) at this taxonomic level. This study indicates that if taxa share a complex behaviour (e.g., case building), then regardless of taxonomic level, one is likely to find shared derived behavioural characters that are useful for phylogeny reconstruction.
Phylogenetic systematic methods were applied in an investigation of the evolution of lek display behavior in the Neotropical manakins (Aves: Pipridae). Results of a previous investigation of the syringeal morphology of manakins were used as a behaviorally independent estimate of the phylogeny of the family. Three phylogenetic analyses of 44 display characters wer
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In 1991 de Pinna (Cladistics 7: 367–394) coined the term primary homology as the putative homology statements prior to tree reconstruction. However, some confusion still exists regarding the conjectural nature of homology and to the analysis of DNA sequences. By dividing de Pinna's term primary homology into topographical identity and character state identity, we emphasize the sequential refinement of putative homology statements. We discuss the problem of transformational versus taxic homology and explain the application of our terms to DNA sequence data.
The problem of assigning optimal character states to the hypothetical ancestors of an evolutionary tree under the Wagner parsimony criterion is examined. A proof is provided for the correctness of Farris's well-known, but previously unproven, algorithm for solving this problem. However, the solution is not, in general, unique, and Farris's method obtains only a subset (generally only one) of the possible solutions. Algorithms that discover other solutions and that resolve ambiguities through the imposition of ancillary criteria are developed and discussed. A method for determining the optimal length of a given tree without actually assigning character states to hypothetical ancestors is described.
Theories of ecological diversification make predictions about the timing and ordering of character state changes through history. These theories are testable by "reconstructing" ancestor states using phylogenetic trees and measurements of contemporary species. Here we use maximum likelihood to estimate and evaluate the accuracy of ancestor reconstructions. We present likelihoods of discrete ancestor states and derive probability distributions for continuous ancestral traits. The methods are applied to several examples: diets of ancestral Darwin's finches; origin of inquilinism in gall wasps; microhabitat partitioning and body size evolution in scrubwrens; digestive enzyme evolution in artiodactyl mammals; origin of a sexually selected male trait, the sword, in platies and swordtails; and evolution of specialization in Anolis lizards. When changes between discrete character states are rare, the maximum likelihood results are similar to parsimony estimates. In this case the accuracy of estimates is often high, with the exception of some nodes deep in the tree. If change is frequent then reconstructions are highly uncertain, especially of distant ancestors. Ancestor states for continuous traits are typically highly uncertain. We conclude that measures of uncertainty are useful and should always be provided, despite simplistic assumptions about the probabilistic models that underlie them. If uncertainty is too high, reconstruction should be abandoned in favor of approaches that fit different models of trait evolution to species data and phylogenetic trees, taking into account the range of ancestor states permitted by the data.
A homolog is a part of the phenotype that is homologous to equivalent parts in other species. A biological homology concept is expected to explain three properties of homologs: 1) the conservation of those features that are used to define a homolog, 2) the individualization of the homolog with regard to the rest of the body, and 3) the uniqueness of homologs, i.e., their specificity for monophyletic groups. The main obstacle to describing a mechanistic basis for homology is the variability of the developmental pathways of undoubtedly homologous characters. However, not all aspects of the developmental pathway are of equal importance. The only organizational features of the developmental system that matter are those that have been historically acquired and cause developmental constraints on the further evolutionary modification of the characters. Two main factors contribute to historically acquired developmental constraints: generative rules of pattern formation and ontogenetic networks. In particular, hierarchical and cyclical inductive networks have the required properties to explain homology. How common such networks are is an open empirical question. The development and variation of pectoral fin hooks in blenniid fishes is presented as a model for the study of a simple ontogenetic network.