Integration of pecking,filter feeding and drinking mechanisms in waterfowl

This paper is one of several contributions in a series, illustrating the application of a specific deductive methodology to explain diversity of form. The methodology facilitates the explanation of feeding morphologies in various ducks as a transformation of the mallard's feeding design maximized for specific proportions of performance that are contributed by pecking and filter feeding mechanisms.The earlier described anatomy and formal analyses of the three mechanisms in the mallard served as the initial conditions used in simulation models. Four elements of the feeding system were chosen that play a major role in all three mechanisms. For each element, the main parameter was selected: storage capacity of the rostral mouth cavity, transport capacity of the rostral mouth tube, storage capacity of the caudal mouth cavity and transport capacity of the caudal mouth tube. The boundary conditions for the simulation were determined from internal organismic constraints. The total food uptake of the mallard was regarded as the function to be maximized. This object function is the summation of the food uptake by one second of pecking and one second of filter feeding. The drinking mechanism was shown not to interfere, since that mechanism operates sufficiently whenever the pumping mechanism works properly.The object function, made up by the pecking and filter feeding performances was graphed. From these graphs a morphospace was developed: the region within which modifications of the mouth design are feasible. This procedure allowed examination of the general hypothesis that different modifications of one design for a complex multi-role system are explainable from differences in proportions of the functional performance contributed by each of the roles.Two predictions were evaluated more specifically: 1) If filter feeding performance must increase for a specific change in total food uptake, the volume of the rostral mouth cavity must increase; this requires widening and lengthening of the rostral maxillar portion and also a phase shift in jaw and lingual motion patterns, increasing the stroke volume. 2) If pecking performance must increase, the transport capacity of the rostral mouth tube must increase; this requires shortening of the maxillar mid portion. These two predictions regarding change in mouth morphology were borne out by shovelers and tufted ducks, respectively.The deductive analysis of the feeding system's morphospace allowed the explanation of the wide diversity of forms in the duck mouth designs as emerging from the great radiation capacity of the ancestral duck's mouth design: it is a versatile, potentially multi-role system in which pecking, drinking and filter feeding mechanisms are mutually very tolerant and, by epiphenomena, includes the elements of a grazing mechanism. In addition, the deduced morphospace was shown to explain phenotypic plasticity, since it explains the occurrence of two morphotypes that develop due to habitat discrimination: one type that has been forced to filter feed, and one type that feeds by pecking.Address reprint requests to Dr. G.A. Zweers     

Ecology predicts large-scale patterns of phylogenetic diversification in birds

One of the most striking patterns in evolutionary biology is that clades may differ greatly in the number of species they contain. Numerous hypotheses have been put forward to explain this phenomenon, and several have been tested using phylogenetic methods. Remarkably, however, all such tests performed to date have been characterized by modest explanatory power, which has generated an interest in explanations stressing the importance of random processes. Here we make use of phylogenetic methods to test whether ecological variables, typically ignored in previous models, may explain phylogenetic tree imbalance in birds. We show that diversification rate possesses an intermediate phylogenetic signal across families. Using phylogenetic comparative methods, we then build a multipredictor model that explains more than 50% of the variation in diversification rate among clades. High annual dispersal is identified as the strongest predictor of high rates of diversification. In addition, high diversification rate is strongly associated with feeding generalization. In all but one instance, these key findings remain qualitatively unchanged when we use an alternative phylogeny and methodology and when small clades, containing five species or less, are excluded. Taken together, these results suggest that large-scale patterns in avian diversification can be explained by variation in intrinsic biology.     

Neural sites and pathways regulating food intake in birds: a comparative analysis to mammalian systems

The paper reviews hypotheses explaining the regulation of food intake in mammals that have addressed specific anatomical structures in the brain. An hypothesis, poikilostasis, is introduced to describe multiple, homeostatic states whereby the regulation of metabolism and feeding occur in birds. Examples are given for both wild and domestic avian species, illustrating dynamic shifts in homeostasis responsible for the changes in body weights that are seen during the course of an annual cycle or by a particular strain of bird. The following neural structures are reviewed as each has been shown to affect food intake in birds or in mammals: ventromedial hypothalamic nucleus (n.), lateral hypothalamic area, paraventricular hypothalamic n., n. tractus solitarius and area postrema, amygdala, parabrachial n., arcuate n. and bed n. of the stria terminalis. Two neural pathways are described which have been proposed to regulate feeding. The trigeminal sensorimotor pathway is the most complete neural pathway characterized for this behavior and encompasses the mechanics of pecking, grasping and mandibulating food particles from the tip of the bill to the back of the buccal cavity. A second pathway, the visceral forebrain system (VFS), affects feeding by regulating metabolism and the balance of the autonomic nervous system. Wild, migratory birds are shown to exhibit marked changes in body weight which are hypothesized to occur due to shifts in balance between the sympathetic and parasympathetic nervous systems. Domestic avian species, selected for a rapid growth rate, are shown to display a dominance of the parasympathetic nervous system. The VFS is the neural system proposed to effect poikilostasis by altering the steady state of the autonomic nervous system in aves and perhaps is applicable to other classes of vertebrates as well.     

Review: Key tweaks to the chicken's beak: the versatile use of the beak by avian species and potential approaches for improvements in poultry production

The avian beak is a multipurpose organ playing a vital role in a variety of functions, including feeding, drinking, playing, grasping objects, mating, nesting, preening and defence against predators and parasites. With regards to poultry production, the beak is the first point of contact between the bird and feed. The beak is also manipulated to prevent unwanted behaviour such as feather pecking, toe pecking and cannibalism in poultry as well as head/neck injuries to breeder hens during mating. Thus, investigating the beak morphometry of poultry in relation to feeding and other behaviours may lead to novel insights for poultry breeding, management and feeding strategies. Beak morphometry data may be captured by advanced imaging techniques coupled with the use of geometric morphometric techniques. This emerging technology may be utilized to study the effects of beak shape on many critical management issues including heat stress, parasite management, pecking and feeding behaviour. In addition, existing literature identifies several genes related to beak development in chickens and other avian species. Use of morphometric assessments to develop phenotypic data on beak shape and detailed studies on beak-related behaviours in chickens may help in improving management and welfare of commercial poultry.     

Density-dependent diversification in North American wood warblers

Evidence from both molecular phylogenies and the fossil record suggests that rates of species diversification often decline through time during evolutionary radiations. One proposed explanation for this pattern is ecological opportunity, whereby an initial abundance of resources and lack of potential competitors facilitate rapid diversification. This model predicts density-dependent declines in diversification rates, but has not been formally tested in any species-level radiation. Here we develop a new conceptual framework that distinguishes density dependence from alternative processes that also produce temporally declining diversification, and we demonstrate this approach using a new phylogeny of North American Dendroica wood warblers. We show that explosive lineage accumulation early in the history of this avian radiation is best explained by a density-dependent diversification process. Our results suggest that the tempo of wood warbler diversification was mediated by ecological interactions among species and that lineage and ecological diversification in this group are coupled, as predicted under the ecological opportunity model.     

The evolution of the modern avian digestive system: insights from paravian fossils from the Yanliao and Jehol biotas

The avian digestive system, like other aspects of avian biology, is highly modified relative to other reptiles. Together these modifications have imparted the great success of Neornithes, the most diverse clade of amniotes alive today. It is important to understand when and how aspects of the modern avian digestive system evolved among neornithine ancestors in order to elucidate the evolutionary success of this important clade and to understand the biology of stem birds and their closest dinosaurian relatives: Mesozoic Paraves. Although direct preservation of the soft tissue of the digestive system has not yet been reported, ingested remains and their anatomical location preserved in articulated fossils hint at the structure of the digestive system and its abilities. Almost all data concerning direct evidence of diet in Paraves comes from either the Upper Jurassic Yanliao Biota or the Lower Cretaceous Jehol Biota, both of which are known from deposits in north-eastern China. Here, the sum of the data gleaned from the thousands of exceptionally well-preserved fossils of paravians is interpreted with regards to the structure and evolution of the highly modified avian digestive system and feeding apparatus. This information suggests intrinsic differences between closely related stem lineages implying either strong homoplasy or that diet in each lineage of non-ornithuromorph birds was highly specialized. Regardless, modern digestive capabilities appear to be limited to the Ornithuromorpha, although the complete set of derived feeding related characters is restricted to the Neornithes.     

To feed or to breed: morphological constraints of mouthbrooding in coral reef cardinalfishes

Functionally coupled biomechanical systems are widespread in nature and are viewed as major constraints on evolutionary diversification, yet there have been few attempts to explore the implications of performing multiple functions within a single anatomical structure. Paternally mouthbrooding cardinalfishes present an ideal system to investigate the constraints of functional coupling as the oral jaws of male fishes are directly responsible for both feeding and reproductive functions. To test the effects of (i) mouthbrooding on feeding and (ii) feeding on reproductive potential we compared the feeding apparatus between sexes of nine species of cardinalfish and compared brood characteristics among species from different trophic groups, respectively. Mouthbrooding was strongly associated with the morphology of the feeding apparatus in males. Male cardinalfishes possessed longer heads, snouts and jaws than female conspecifics irrespective of body size, trophic group or evolutionary history. Conversely, reproductive potential also appeared to be related to trophic morphology. Piscivorous cardinalfishes produced larger, but fewer eggs, and had smaller brood volumes than species from the two invertebrate feeding groups. These interrelationships suggest that feeding and reproduction in the mouth of cardinalfishes may be tightly coupled. If so this may, in part, have contributed to the limited morphological diversification exhibited by cardinalfishes.     

Revisiting Amazonian phylogeography: insights into diversification hypotheses and novel perspectives

The Amazon Basin harbors one of the richest biotas on Earth, such that a number of diversification hypotheses have been formulated to explain patterns of Amazonian biodiversity and biogeography. For nearly two decades, phylogeographic approaches have been applied to better understand the underlying causes of genetic differentiation and geographic structure among Amazonian organisms. Although this research program has made progress in elucidating several aspects of species diversification in the region, recent methodological and theoretical developments in the discipline of phylogeography will provide new perspectives through more robust hypothesis testing. Herein, we outline central aspects of Amazonian geology and landscape evolution as well as climate and vegetation dynamics through the Neogene and Quaternary to contextualize the historical settings considered by major hypotheses of diversification. We address each of these hypotheses by reviewing key phylogeographic papers and by expanding their respective predictions. We also propose future directions for devising and testing hypotheses. Specifically, combining the exploratory power of phylogeography with the statistical rigor of coalescent methods will greatly expand analytical inferences on the evolutionary history of Amazonian biota. Incorporation of non-genetic data from Earth science disciplines into the phylogeographic approach is key to a better understanding of the influence of climatic and geophysical events on patterns of Amazonian biodiversity and biogeography. In addition, achieving such an integrative enterprise must involve overcoming issues such as limited geographic and taxonomic sampling. These future challenges likely will be accomplished by a combination of extensive collaborative research and incentives for conducting basic inventories.     

A model for the evolutionary dynamics of cross-feeding polymorphisms in microorganisms

 Understanding mechanisms of evolutionary diversification is central to evolutionary biology. Microbes constitute promising model systems for observing processes of diversification directly in the laboratory. One of the main existing paradigms for microbial diversification is the evolution of cross-feeding polymorphisms, in which a strain specializing on a primary resource coexists with a cross-feeding strain that specializes on a waste product resulting from consumption of the primary resource. Here I propose a theoretical model for the evolutionary dynamics through which cross-feeding polymorphisms can gradually emerge from a single ancestral strain. The model is based on the framework of adaptive dynamics, which has proved to be very useful for studying adaptive processes of divergence under sympatric conditions. In particular, the phenomenon of evolutionary branching serves as a general paradigm for diversification. I show that evolutionary branching naturally occurs in evolutionary models of cross-feeding if (1) there is a trade-off between uptake efficiencies on the primary and secondary resources, and (2) this trade-off has positive curvature. The model also suggests that the evolution of cross-feeding should be more likely in chemostat cultures than in serial batch cultures, which conforms with empirical observations. Overall, the model provides a theoretical metaphor for the evolution of cross-feeding polymorphisms. Received: February 19, 2002 / Accepted: May 8, 2002     

Continuously stable strategies as evolutionary branching points

Evolutionary branching points are a paradigmatic feature of adaptive dynamics, because they are potential starting points for adaptive diversification. The antithesis to evolutionary branching points are continuously stable strategies (CSS's), which are convergent stable and evolutionarily stable equilibrium points of the adaptive dynamics and hence are thought to represent endpoints of adaptive processes. However, this assessment is based on situations in which the invasion fitness function determining the adaptive dynamics have non-zero second derivatives at CSS. Here we show that the scope of evolutionary branching can increase if the invasion fitness function vanishes to higher than first order at CSS. Using classical models for frequency-dependent competition, we show that if the invasion fitness vanishes to higher orders, a CSS may be the starting point for evolutionary branching. Thus, when invasion fitness functions vanish to higher than first order at equilibrium points of the adaptive dynamics, evolutionary diversification can occur even after convergence to an evolutionarily stable strategy.     

The prior probabilities of phylogenetic trees

Bayesian methods have become among the most popular methods in phylogenetics, but theoretical opposition to this methodology remains. After providing an introduction to Bayesian theory in this context, I attempt to tackle the problem mentioned most often in the literature: the “problem of the priors”—how to assign prior probabilities to tree hypotheses. I first argue that a recent objection—that an appropriate assignment of priors is impossible—is based on a misunderstanding of what ignorance and bias are. I then consider different methods of assigning prior probabilities to trees. I argue that priors need to be derived from an understanding of how distinct taxa have evolved and that the appropriate evolutionary model is captured by the Yule birth–death process. This process leads to a well-known statistical distribution over trees. Though further modifications may be necessary to model more complex aspects of the branching process, they must be modifications to parameters in an underlying Yule model. Ignoring these Yule priors commits a fallacy leading to mistaken inferences both about the trees themselves and about macroevolutionary processes more generally.     

Concepts of an architectonic approach to transformation morphology

This paper is about a general methodology for pattern transformation. Patterns are network representations of the relations among structures and functions within an organism. Transformation refers to any realistic or abstract transformation relevant to biology, e.g. ontogeny, evolution and phenotypic clines. The main aim of the paper is a methodology for analyzing the range of effects on a pattern due to perturbing one or more of its structures and/or functions (transformation morphology). Concepts relevant to such an analysis of pattern transformation are reviewed and several new ones introduced: pattern unit; direct and indirect functional demands; compatibility and trade-off; integrating, adding and decoupling; functional effectiveness; spatial, profile and other architectonic constraints; domains of structure-function relations; goal and process adaptability; multiple pathways. The paper is written from the the perspective of architectonic morphology, viz. functional morphology focusing on the relation between anatomical coherence and the compatibility of functions. The advantages and disadvantages of inductive and deductive approaches are discussed.     

Disruptive selection as a driver of evolutionary branching and caste evolution in social insects

Theory suggests that evolutionary branching via disruptive selection may be a relatively common and powerful force driving phenotypic divergence. Here, we extend this theory to social insects, which have novel social axes of phenotypic diversification. Our model, built around turtle ant (Cephalotes) biology, is used to explore whether disruptive selection can drive the evolutionary branching of divergent colony phenotypes that include a novel soldier caste. Soldier evolution is a recurrent theme in social insect diversification that is exemplified in the turtle ants. We show that phenotypic mutants can gain competitive advantages that induce disruptive selection and subsequent branching. A soldier caste does not generally appear before branching, but can evolve from subsequent competition. The soldier caste then evolves in association with specialized resource preferences that maximize defensive performance. Overall, our model indicates that resource specialization may occur in the absence of morphological specialization, but that when morphological specialization evolves, it is always in association with resource specialization. This evolutionary coupling of ecological and morphological specialization is consistent with recent empirical evidence, but contrary to predictions of classical caste theory. Our model provides a new theoretical understanding of the ecology of caste evolution that explicitly considers the process of adaptive phenotypic divergence and diversification.     

Functional implications of species differences in the size and morphology of the isthmo optic nucleus (ION) in birds

In birds, there is a retinofugal projection from the brain to the retina originating from the isthmo optic nucleus (ION) in the midbrain. Despite a large number of anatomical, physiological and histochemical studies, the function of this retinofugal system remains unclear. Several functions have been proposed including: gaze stabilization, pecking behavior, dark adaptation, shifting attention, and detection of aerial predators. This nucleus varies in size and organization among some species, but the relative size and morphology of the ION has not been systematically studied. Here, we present a comparison of the relative size and morphology of the ION in 81 species of birds, representing 17 different orders. Our results show that several orders of birds, besides those previously reported, have a large, well-organized ION, including: hummingbirds, woodpeckers, coots and allies, and kingfishers. At the other end of the spectrum, parrots, herons, waterfowl, owls and diurnal raptors have relatively small ION volumes. ION also appears to be absent or unrecognizable is several taxa, including one of the basal avian groups, the tinamous, which suggests that the ION may have evolved only in the more modern group of birds, Neognathae. Finally, we demonstrate that evolutionary changes in the relative size and the cytoarchitectonic organization of ION have occurred largely independent of phylogeny. The large relative size of the ION in orders with very different lifestyles and feeding behaviors suggest there is no clear association with pecking behavior or predator detection. Instead, our results suggest that the ION is more complex and enlarged in birds that have eyes that are emmetropic in some parts of the visual field and myopic in others. We therefore posit that the ION is involved in switching attention between two parts of the retina i.e. from an emmetropic to a myopic part of the retina.     

New inferences from tree shape: numbers of missing taxa and population growth rates

The relative positions of branching events in a phylogeny contain information about evolutionary and population dynamic processes. We provide new summary statistics of branching event times and describe how these statistics can be used to infer rates of species diversification from interspecies trees or rates of population growth from intraspecies trees. We also introduce a phylogenetic method for estimating the level of taxon sampling in a clade. Different evolutionary models and different sampling regimes can produce similar patterns of branching events, so it is important to consider explicitly the model assumptions involved when making evolutionary inferences. Results of an analysis of the phylogeny of the mosquito-borne flaviviruses suggest that there could be several thousand currently unidentified viruses in this clade.     

EVOLUTIONARY LOSS OF LARVAL FEEDING: DEVELOPMENT,FORM AND FUNCTION IN A FACULTATIVELY FEEDING LARVA,BRISASTER LATIFRONS

Species with large eggs and nonfeeding larvae have evolved many times from ancestors with smaller eggs and feeding larvae in numerous groups of aquatic invertebrates and amphibians. This change in reproductive allocation and larval form is often accompanied by dramatic changes in development. Little is known of this transformation because the intermediate form (a facultatively feeding larva) is rare. Knowledge of facultatively feeding larvae may help explain the conditions under which nonfeeding larvae evolve. Two hypotheses concerning the evolutionary loss of larval feeding are as follows: (1) large eggs evolve before modifications in larval development, and (2) the intermediate form (facultatively feeding larva) is evolutionarily short-lived. I show that larvae of a heart urchin, Brisaster latifrons, are capable of feeding but do not require food to complete larval development. Food for larvae appears to have little effect on larval growth and development. The development, form, and suspension feeding mechanism of these larvae are similar to those of obligate-feeding larvae of other echinoids. Feeding rates of Brisaster larvae are similar to cooccurring, obligate-feeding echinoid larvae but are low relative to the large size of Brisaster larvae. The comparison shows that in Brisaster large egg size, independence from larval food, and relatively low feeding rate have evolved before the heterochronies and modified developmental mechanisms common in nonfeeding echinoid larvae. If it is general, the result suggests that hypotheses concerning the origin of nonfeeding larval development should be based on ecological factors that affect natural selection for large eggs, rather than on the evolution of heterochronies and developmental novelties in particular clades. I also discuss alternative hypotheses concerning the evolutionary persistence of facultative larval feeding as a reproductive strategy. These hypotheses could be tested against a phylogenetic hypothesis.     

Independent evolutionary reductions of the phallus in basal birds

Despite a long history of anatomical studies in birds, the genitalia of most avian species remain undescribed. Birds are the only vertebrate taxon with internal fertilization where an intromittent phallus has been lost in most species. Studying the anatomical transitions of the avian phallus in those species where it is still present, allows us to test evolutionary hypotheses of why the phallus was lost in the ancestor of modern birds. As part of an anatomical survey of the evolution of avian phallus morphology, we have examined some avian species whose genitalia have not been described. Previously, there were only two known events of phallus reduction in birds: one transition from intromittent to non‐intromittent in the Galliformes, and a complete loss of phallic structures in the ancestor of Neoaves. Here we report three additional cases of phallus reduction in birds: a transition from intromittent to non‐intromittent phallus in Tinamiformes (Crypturellus, Tinamidae), the presence of a non‐intromittent phallus in Alectura (Megapodidae), and a complete loss of the phallus in Leipoa (Megapodidae). In addition, we report on the unique morphology of the Crypturellus non‐intromittent phallus. These new records of phallus reduction highlight the dynamic nature of phallus evolution in birds. Our findings provide evidence against the hypothesis that the phallus in birds is maintained to insure paternity in taxa with exclusive male parental care, since both groups where we report phallus reduction provide predominately male‐only care.     

Bird evolution in the Eocene: climate change in Europe and a Danish fossil fauna

The pattern of the evolutionary radiation of modern birds (Neornithes) has been debated for more than 10 years. However, the early fossil record of birds from the Paleogene, in particular, the Lower Eocene, has only recently begun to be used in a phylogenetic context to address the dynamics of this major vertebrate radiation. The Cretaceous-Paleogene (K-P) extinction event dominates our understanding of early modern bird evolution, but climate change throughout the Eocene is known to have also played a major role. The Paleocene and Lower Eocene was a time of avian diversification as a result of favourable global climatic conditions. Deteriorations in climate beginning in the Middle Eocene appear to be responsible for the demise of previously widespread avian lineages like Lithornithiformes and Gastornithidae. Other groups, such as Galliformes display replacement of some lineages by others, probably related to adaptations to a drier climate. Finally, the combination of slowly deteriorating climatic conditions from the Middle Eocene onwards, appears to have slowed the evolutionary rate in Europe, as avian faunas did not differentiate markedly until the Oligocene. Taking biotic factors in tandem with the known Paleogene fossil record of Neornithes has recently begun to illuminate this evolutionary event. Well-preserved fossil taxa are required in combination with ever-improving phylogenetic hypotheses for the inter-relationships of modern birds founded on morphological characters. One key avifauna of this age, synthesised for the first time herein, is the Lower Eocene Fur Formation of Denmark. The Fur birds represent some of the best preserved (often in three dimensions and with soft tissues) known fossil records for major clades of modern birds. Clear phylogenetic assessment of these fossils will prove critical for future calibration of the neornithine evolutionary timescale. Some early diverging clades were clearly present in the Paleocene as evidenced directly by new fossil material alongside the phylogenetically constrained Lower Eocene taxa. A later Oligocene radiation of clades other than Passeriformes is not supported by available fossil data.