Size and number of the hypoglossal nerve foramina in the avian skull and their potential neuroanatomical significance

The cranial openings of nervus hypoglossus, the 12th cranial nerve, are for the first time studied across a broad range of higher avian taxa. This nerve plays an important role in the innervation of the syrinx and exits the skull through a variable number of foramina. Most previous authors described 2–3 foramina nervi hypoglossi (FNH) for neornithine birds, but the number, size, and arrangement of FNH is actually more variable than what is apparent from the literature. In the case of three foramina, there is usually a pair of caudal foramina and a rostral one, but even in closely related taxa, a great variability of the FNH pattern may exist. Many taxa of Neognathae exhibit a quadruple of symmetrically arranged FNH, in others four foramina are strung together in a line. A few taxa show more than four FNH, although in these cases the additional foramina are very small. Of particular interest is the occurrence of a very large caudal FNH in Trochilidae and many species of Passeriformes. This large foramen is suggestive of a correlation with the highly developed vocal capabilities of these birds and may transmit fibers from the tracheosyringeal portion of nucleus nervi hypoglossi, in which case it would be an osteological correlate of vocal capabilities. However, targeted neuroanatomical studies are required to determine how individual hypoglossal foramina relate to hypoglossal roots and their branches, and which of them receive fibers supplying axial, lingual, and syringeal muscles.     

Phylogenetic interrelationships of living and extinct Tinamidae,volant palaeognathous birds from the New World

Tinamous, one of the earliest diverging living avian lineages, consists of a Neotropical clade of palaeognathous birds with a fossil record limited to the early Miocene–Quaternary of southern South America. Here, we conduct a comprehensive, morphology‐based phylogenetic study of the interrelationships among extinct and living species of tinamous. Morphological data of fossil species are included in a matrix of 157 osteological and myological characters of 56 terminal taxa. The monophyly of most recognized genera is supported by the results of the analysis. The cladistic analysis also recovers the traditional subdivision between those tinamous specialized for open areas (Nothurinae) and those inhabiting forested environments (Tinaminae). Temporal calibration of the resultant phylogeny indicates that such a basal divergence had already taken place in the early Miocene, some 17 million years ago. The placement of the fossil species within the open‐area (Nothurinae) and the forest‐dwelling (Tinaminae) tinamous is also consistent with the palaeoenvironmental conditions inferred from the associated fauna. © 2014 The Linnean Society of London     

Hindlimb morphology and locomotor performance in waders: an evolutionary approach

Locomotion performance (measured as stride frequency and stride length) was studied in 16 species of waders. Differences in hindlimb morphology (osteology and myology) were analysed among species. Evolutionary changes in both locomotion and morphological variables were analysed using comparative methods revealing the existence of some ecomorphological patterns relating these two sets of characters. Evolutionary changes in stride frequency were correlated with changes in the muscles M. iliotibialis cranialis, M. iliotibiales lateralis and M. gastrocnemius, whereas changes in stride length showed correlated evolution with changes in the length of distal segments of the leg. We identify two different evolutionary strategies in locomotion of waders. One is a change in distal leg segments (skeletal system), an adaptive modification that increases stride length; the second is a change in the skeletal-muscular system, providing an increase in muscular performance (force or speed of contraction) in several muscles, and is an adaptation that increases stride frequency.     

The fossil record and limb disparity of enantiornithines, the dominant flying birds of the Cretaceous

Morphometric and stratigraphic analyses that encompass the known fossil record of enantiornithine birds (Enantiornithes) are presented. These predominantly flighted taxa were the dominant birds of the second half of the Mesozoic; the enantiornithine lineage is known to have lasted for at least 60 million years (Ma), up until the end of the Cretaceous. Analyses of fossil record dynamics show that enantiornithine 'collectorship' since the 1980s approaches an exponential distribution, indicating that an asymptote in proportion of specimens has yet to be achieved. Data demonstrate that the fossil record of enantiornithines is complete enough for the extraction of biological patterns. Comparison of the available fossil specimens with a large data set of modern bird (Neornithes) limb proportions also illustrates that the known forelimb proportions of enantiornithines fall within the range of extant taxa; thus these birds likely encompassed the range of flight styles of extant birds. In contrast, most enantiornithines had hindlimb proportions that differ from any extant taxa. To explore this, ternary diagrams are used to graph enantiornithine limb variation and to identify some morphological oddities ( Otogornis , Gobipteryx ); taxa not directly comparable to modern birds. These exceptions are interesting – although anatomically uniform, and similar to extant avians in their wing proportions, some fossil enantiornithines likely had flight styles not seen among their living counterparts.     

The evolutionary radiation of modern birds (Neornithes): reconciling molecules, morphology and the fossil record

The pattern, timing and extent of the evolutionary radiation of anatomically modern birds (Neornithes) remains contentious: dramatically different timescales for this major event in vertebrate evolution have been recovered by the 'clock-like' modelling of molecular sequence data and from evidence extracted from the known fossil record. Because current synthesis would lead us to believe that fossil and nonfossil evidence conflict with regard to the neornithine timescale, especially at its base, it is high time that available data are reconciled to determine more exactly the evolutionary radiation of modern birds. In this review we highlight current understanding of the early fossil history of Neornithes in conjunction with available phylogenetic resolution for the major extant clades, as well as recent advancements in genetic methods that have constrained time estimates for major evolutionary divergences. Although the use of molecular approaches for timing the radiation of Neornithes is emphasized, the tenet of this review remains the fossil record of the major neornithine subdivisions and better-preserved taxa. Fossils allowing clear phylogenetic constraint of taxa are central to future work in the production of accurate molecular calibrations of the neornithine evolutionary timescale.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 153–177.     

Phylogenetic relationships of the paraphyletic ‘caprimulgiform’ birds (nightjars and allies)

In the past years, paraphyly of the traditional ‘Caprimulgiformes’ (nightjars and allies) with respect to Apodiformes (swifts and hummingbirds) has been well established, but the relationships between the five ‘caprimulgiform’ family‐level taxa remain controversial. These crepuscular or nocturnal birds differ in numerous anatomical features, and here an analysis of 69 morphological characters is performed. Except for the position of the Nyctibiidae (potoos), the topology of the single most‐parsimonious tree agrees with the results of a recently published large‐scale ‘phylogenomic’ study. Whereas molecular data support a clade including Nyctibiidae and the Steatornithidae (oilbird), potoos were shown to be the sister taxon of Caprimulgidae (nightjars) in the present analysis. A sister group relationship between Nyctibiidae and Caprimulgidae is strongly supported, both in terms of bootstrap robustness and the number of synapomorphies, and it is detailed that the morphological data are more likely to reflect the true relationships of these birds. A classification is proposed, and the term Strisores is introduced for a clade including all ‘Caprimulgiformes’ and Apodiformes. It is most parsimonious to assume a single origin of dark activity in the stem lineage of Strisores and a reversal to diurnal activity in Apodiformes. However, a fourfold origin of dark‐activity in the stem lineages of Steatornithidae, Podargidae, Aegothelidae and the Caprimulgidae/Nyctibiidae cannot be conclusively excluded with the data at hand.     

Comparative morphology of the avian maxillary bone (os maxillare) based on an examination of macerated juvenile skeletons

For the first time, isolated maxillary bones of juvenile neornithine birds are examined and compared. Contrary to the anatomical terminology currently employed, the avian maxillare exhibits five rather than four processes. In addition to the praemaxillary, jugal, nasal, and maxillopalatine processes, all palaeognathous and many neognathous birds also have a palatine process. The occurrence of these processes is, however, variable across different clades and only few taxa exhibit a pentaradiate maxillare with all five processes. Within Neognathae, a great morphological variability exists in the shape of the maxillopalatine process, which is more easily studied in juvenile individuals, in which the bones of the beak and palate are not co-ossified. In some Neognathae, a caudally facing recess is situated in the junction of the maxillopalatine and jugal processes, which is likely to be homologous to the pneumatic recess of palaeognathous birds. Several derived morphologies of potential phylogenetic significance for the characterization of neognathous clades are identified and major morphological transformations in the lineage leading towards modern birds are highlighted.     

Osteology and myology of the cephalic region and pectoral girdle of the Chinese catfish Cranoglanis bouderius,with a discussion on the autapomorphies and phylogenetic relationships of the Cranoglanididae (Teleostei: Siluriformes)

The cephalic and pectoral girdle structures of the Chinese catfish Cranoglanis bouderius are described and compared with those of other catfishes as the foundation for an analysis on the Cranoglanididae autapomorphies and also for a discussion on the phylogenetic relationships between the cranoglanidids and the other catfishes. Our observations and comparisons indicate that cranoglanidids are defined, at least, by four autapomorphies, namely: 1) the cartilages associated with the mandibular barbels are broad, somewhat circular; 2) epioccipital with a well-developed posterodorsal process, which presents a large, deep, circular posterior concavity; 3) a well-defined, deep, anteroposteriorly elongated concavity formed by both the frontal and the lateral ethmoid to receive the anteromedial surface of the metapterygoid; 4) the adductor mandibulae A3" is dorsally divided into two bundles and partially inserted on the posterior portion of the primordial ligament. With respect to the phylogenetic relationships of the Cranoglanididae, this study strongly suggests that these fishes are probably closely related to the Ariidae and the Claroteidae.     

Osteology and myology of the cephalic region and pectoral girdle of Erethistes pusillus, comparison with other erethistids, and comments on the synapomorphies and phylogenetic relationships of the Erethistidae (Teleostei: Siluriformes)

The cephalic and pectoral girdle structures of the erethistid Erethistes pusillus (Erethistinae) are described and compared with those of another species of the subfamily Erethistinae, namely Hara filamentosa , and of the single species of the subfamily Continae, Conta conta , as well as of several other catfishes, as the foundation for a discussion on the synapomorphies and phylogenetic position of the Erethistidae. The observations and comparisons support de Pinna's phylogenetic hypothesis, according to which the Erethistidae is the sister‐group of the Aspredinidae, with the clade formed by these two families being the sister‐group of the Sisoridae sensu stricto . In addition, the observations and comparisons pointed out a new, additional character to diagnose the family Erethistidae, namely: mesocoracoid arch deeply bifurcated dorsally.     

Diversity of feeding adaptations in certain columbid birds: A functional morphological approach

With gradual increasing complexity in higher vertebrate structure and function, the birds as a class have acquired very high degree of feeding adaptations for diverse food-niches. A comparative functional morphological study of the feeding apparatus of 6 species of columbid birds showing diversification in their food-habits reveals that some correlations exist between the form-function complexes of the feeding apparatus and the extent of diversity of food-habits shown by these birds. Among the species of columbid birds selected for the Study,Columba andStreptopelia are ground feeders and predominantly grain-eaters, although quite often they invade diversified food-niches.Treron andDucula, on the other hand, are almost exclusively fruit-eaters, plucking and swallowing fruits from the lofty tree branches, WhileColumba andStreptopelia show better kinesis of their jaws for ground-pecking,Treron. andDucula possess wider gape as well as stronger grasp of their bill for plucking off, grasping and swallowing large-sized fruits. Consequently, the size and pinnateness of the jaw muscles in these fruit-pigeons have developed far greater than those observed inColumba andStreptopelia. Further, inTreron andDucula, the thick and broad ‘venter externus’ slip of the M. pterygoideus ensures complete closure of the bill and possibly prevent any excess lateral expansion of the mandibular rami. Similar correlations have also been observed between the tongue features of columbid birds and the diversity of their feeding adaptation.     

Body mass and foraging ecology predict evolutionary patterns of skeletal pneumaticity in the diverse "waterbird" clade

Extensive skeletal pneumaticity (air-filled bone) is a distinguishing feature of birds. The proportion of the skeleton that is pneumatized varies considerably among the >10,000 living species, with notable patterns including increases in larger bodied forms, and reductions in birds employing underwater pursuit diving as a foraging strategy. I assess the relationship between skeletal pneumaticity and body mass and foraging ecology, using a dataset of the diverse "waterbird" clade that encompasses a broad range of trait variation. Inferred changes in pneumaticity and body mass are congruent across different estimates of phylogeny, whereas pursuit diving has evolved independently between two and five times. Phylogenetic regressions detected positive relationships between body mass and pneumaticity, and negative relationships between pursuit diving and pneumaticity, whether independent variables are considered in isolation or jointly. Results are generally consistent across different estimates of topology and branch lengths. "Predictive" analyses reveal that several pursuit divers (loons, penguins, cormorants, darters) are significantly apneumatic compared to their relatives, and provide an example of how phylogenetic information can increase the statistical power to detect taxa that depart from established trait correlations. These findings provide the strongest quantitative comparative support yet for classical hypotheses regarding the evolution of avian skeletal pneumaticity.     

The patterns and modes of the evolution of disparity in Mesozoic birds

The origin of birds from non-avian theropod dinosaurs is one of the greatest transitions in evolution. Shortly after diverging from other theropods in the Late Jurassic, Mesozoic birds diversified into two major clades—the Enantiornithes and Ornithuromorpha—acquiring many features previously considered unique to the crown group along the way. Here, we present a comparative phylogenetic study of the patterns and modes of Mesozoic bird skeletal morphology and limb proportions. Our results show that the major Mesozoic avian groups are distinctive in discrete character space, but constrained in a morphospace defined by limb proportions. The Enantiornithines, despite being the most speciose group of Mesozoic birds, are much less morphologically disparate than their sister clade, the Ornithuromorpha—the clade that gave rise to living birds, showing disparity and diversity were decoupled in avian history. This relatively low disparity suggests that diversification of enantiornithines was characterized in exhausting fine morphologies, whereas ornithuromorphs continuously explored a broader array of morphologies and ecological opportunities. We suggest this clade-specific evolutionary versatility contributed to their sole survival of the end-Cretaceous mass extinction.     

Decoupled evolution of display morphology and display behaviour in phrynosomatid lizards

A widespread trend in animals is the evolution of morphological ornaments and behaviours that are involved in aggressive and courtship displays. These display traits are important from the standpoint of communication, sexual selection, and speciation. Previous authors have suggested that the evolution of display morphology and display behaviour should be closely linked. In this study, I tested for this association with behavioural and morphological data for 59 taxa of phrynosomatid lizards using phylogenetic comparative methods (Mad-dison's concentrated changes test and Felsenstein's independent contrasts). The results showed little significant association between features of display morphology and behaviour, suggesting that the evolution of these traits is not tightly coupled. This decoupling is particularly evident in the genus Sceloporus , in which several species have lost the display coloration but retain unmodified display behaviour. The results also suggest that display morphology is more evolutionarily labile than display behaviour in this group.     

Using the whole body as a sucker: Combining respiration and feeding with an attached lifestyle in hill stream loaches (Balitoridae,Cypriniformes)

Small fishes living in fast‐flowing rivers face a harsh environment as they can easily be swept away by the rapid currents. To survive such circumstances, teleosts evolved a wide variety of attachment mechanisms, based on friction, negative pressure or both. Balitorinae (Balitoridae, Cypriniformes) are exceptional in using their whole body as an adhesive apparatus. We investigated the morphological adaptations of Balitorinae by studying the osteology and myology of four species (Beaufortia leveretti, Sewellia lineolata, Pseudogastromyzon myersi, and Gastromyzon punctulatus) using clearing and staining, serial cross‐sections and CT‐scanning. A kinematic analysis was performed to study the respiration and feeding mechanisms and to identify key structures in these mechanisms. Our research showed that the whole body of Balitorinae acts as a suction disc, with friction‐enhancing structures (unculi) on the thickened anterior rays of the paired fins. The abruptly rising head profile, supported by the extremely enlarged lacrimal bone and the flat ventral body surface facilitate effective substrate attachment. During attachment, the pelvic girdle is pulled anterodorsally, suggesting the formation of a negative pressure underneath the body. Detachment by water inflow underneath the body is prevented by three mechanisms. 1) Barbels control the water inflow by detachment and reattachment to the substrate. 2) Most water present underneath the body is removed during inspiration. 3) Excess water is regularly removed by movements of the posterior pectoral fin rays. The balitorine body is thus modified as such that it allows effective attachment, while not impairing respiration. Comparison with other teleosts living in similar environments shows that most species use more locally concentrated modifications of the paired fins and/or the mouth for attachment. The high diversity in teleostean adhesive apparatuses and associated myological modifications suggest a substantial functional convergent evolution, without necessarily highly convergent anatomical adaptations. J. Morphol. 275:1066–1079, 2014. © 2014 Wiley Periodicals, Inc.     

Cotylea (Polycladida): a cladistic analysis of morphology

Abstract. Polyclad flatworms are acoelomate bilaterians found in benthic communities worldwide, predominantly in marine environments. Current polyclad systematics is unstable, with two non-concordant classification schemes resulting in a poor understanding of within-group relationships. Here we present the first phylogenetic framework for the suborder Cotylea using a morphological matrix. Representatives of 34 genera distributed among all cotylean families (except four, excluded due to their dubious taxonomic status) were investigated. The number of families included ranges from a conservative eight to a revisionary 11. Outgroup analysis indicated that the suborder is monophyletic and defined by the presence of a ventral adhesive structure, a short posteriorly positioned vagina, and cement glands. Of the eight to 11 families included, we confirmed that three were monophyletic: Boniniidae, Prosthiostomidae, and Pseudocerotidae. Boniniidae was consistently recovered as the sister group to other Cotylea, based on the retention of the plesiomorphic Lang's vesicle. The clade consisting of Anonymus, Marcusia , and Pericelis is sister to the Boniniidae and the rest of the Cotylea. Above this clade there is little resolution at the base of the sister group. The Euryleptidae are found to be paraphyletic and give rise to the Pseudocerotidae. Neither classification scheme received unequivocal support. The intrafamilial relationships of the diverse Pseudocerotidae and Euryleptidae were examined. Color pattern characters (used for species identification) were highly homoplasious but increased cladogram resolution within genera. The monophyly of seven genera within the Pseudocerotidae and Euryleptidae was not supported and many genera showed no autapomorphies, highlighting the need for taxonomic revision of these families.     

New data on the humerotriceps of penguins and its implications in the evolution of the fossa tricipitalis

A paddle-shaped wing, the general morphology of the humerus, and the muscles involved in wing movement are among the most characteristic adaptations to diving in penguins. Particularly, the humeral fossa tricipitalis and the musculus humerotriceps are clear examples of muscular rearrangement accompanying skeletal changes. In extant Spheniscidae, we were able to identify two heads of this muscle attaching within a different compartment of the bipartite fossa. Since the partition of the fossa appeared as a novelty during the Miocene, we propose that this might have had implications for underwater flight contributing to wing-propelled diving efficiency.     

Notograptidae, sister to Acanthoplesiops Regan (Teleostei: Plesiopidae: Acanthoclininae), with comments on biogeography, diet and morphological convergence with Congrogadinae (Teleostei: Pseudochromidae)

The Notograptidae contains one genus, Notograptus Günther, and five nominal species from northern Australia and southern New Guinea. Morphological evidence places Notograptus among acanthoclinine plesiopids (continuous free margin of lower lip; head naked; dorsal and anal fins with many spines and few segmented rays; no extensor proprius; reduced number of caudal-fin rays) and supports a sister relationship with Acanthoplesiops (symphyseal flap on lower lip; reduced hypural 5; reduced hypurapophysis). This hypothesis resolves the relationships within Acanthoplesiops , clarifying the polarity of autogenous middle radials of dorsal- and anal-fin pterygiophores. The proposed relationships among acanthoclinines are: Acanthoclinus ( Belonepterygion ( Beliops ( Notograptus ( Acanthoplesiops hiatti ( A. indicus ( A. psilogaster ( A. echinatus ))))))). The distribution of Notograptus compliments that of its proposed sister clade in that Acanthoplesiops is unknown from northern Australia or southern New Guinea. There are repeated geographical patterns among several groups suggesting that Australia is a basal area to a broader Indo-Pacific region. Similarity between the Congrogadinae (Pseudochromidae) and Notograptus has long been noted, both having a loosely connected suspensorium and elongate body which were mistakenly considered indicators of relationship; we add reduced branchial arches, straight, tube-like gut and highly expandable anus. We examine these similarities as an indication of a shared specialized feeding habit. Notograptus is an alpheid shrimp predator, able to swallow its large prey whole. Most species of congrogadines eat whole, large crustaceans. This is probably an example of convergent adaptation to a particular selective regime.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 179–205.     

Body size evolution in Mesozoic birds: little evidence for Cope's rule

Cope's rule, the tendency towards evolutionary increases in body size, is a long-standing macroevolutionary generalization that has the potential to provide insights into directionality in evolution; however, both the definition and identification of Cope's rule are controversial and problematic. A recent study [J. Evol. Biol. 21 (2008) 618] examined body size evolution in Mesozoic birds, and claimed to have identified evidence of Cope's rule occurring as a result of among-lineage species sorting. We here reassess the results of this study, and additionally carry out novel analyses testing for within-lineage patterns in body size evolution in Mesozoic birds. We demonstrate that the nonphylogenetic methods used by this previous study cannot distinguish between among- and within-lineage processes, and that statistical support for their results and conclusions is extremely weak. Our ancestor-descendant within-lineage analyses explicitly incorporate recent phylogenetic hypotheses and find little compelling evidence for Cope's rule. Cope's rule is not supported in Mesozoic birds by the available data, and body size evolution currently provides no insights into avian survivorship through the Cretaceous-Paleogene mass extinction.