New species of Primozygodactylus from Messel and the ecomorphology and evolutionary significance of early Eocene zygodactylid birds (Aves,Zygodactylidae)

Zygodactylids (Zygodactylidae) are the extinct sister taxon of passerines (Passeriformes) and among the more abundant small arboreal birds in the early Eocene German fossil site Messel. Four species of the taxon Primozygodactylus have previously been identified and here two new species are described. In addition, new fossils of the poorly known P. eunjooae are reported. The fossils corroborate the presence of two very long central tail feathers for Primozygodactylus, and the feathering of the taxon corresponds with that of extant birds foraging in scrub and undergrowth. Species diversity of zygodactylids falls short of that of passerines, the most species-rich extant avian clade, and in bill shapes they also do not reach the diversity seen in extant Passeriformes. Because zygodactylids closely resemble passerines in skeletal morphology, the evolutionary success of passerines is unlikely to be due to particular skeletal features. Passerines and zygodactylids coexisted in Europe from the early Oligocene to the middle Miocene, and both groups probably differed in ecological characteristics. The zygodactyl foot of zygodactylids may have represented an adaptation for clinging to tree trunks, and if nesting in tree cavities, zygodactylids may have succumbed to competition for safe nesting places with avian or mammalian competitors.

http://zoobank.org/urn:lsid:zoobank.org:pub:7F142141-7E1F-4568-89BF-E2363D128C36     

On the osteology and phylogenetic affinities of the Pseudasturidae – Lower Eocene stem-group representatives of parrots (Aves, Psittaciformes)

The osteology of the early Eocene (about 50 mya) avian taxon Pseudasturidae Mayr, 1998 is revised and its phylogenetic affinities are analysed. Members of the Pseudasturidae are known from abundant and excellently preserved skeletal material, both complete skeletons on slabs as well as isolated, three-dimensional bones. Although this taxon is thus among the best represented of all small early Tertiary birds, its systematic affinities were unknown so far. Derived osteological characters which are visible in newly recognized specimens from the Lower Eocene London Clay of England most convincingly support classification of the Pseudasturidae into the Psittaciformes (parrots). Both, in overall morphology and in terms of derived characters, the tarsometatarsus of the Pseudasturidae closely resembles that of the Eocene Quercypsittidae, which were assigned to the Psittaciformes by Mourer-Chauviré (1992 ). The Pseudasturidae are considered to be stem-group representatives of the Psittaciformes and the sister taxon of all other known psittaciform birds. The Eocene taxon lacks the specialized bill morphology of crown-group Psittaciformes of the Psittacidae. Several other osteological differences between the Pseudasturidae and the Psittacidae probably are also functionally correlated with the specialized feeding technique of the latter.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 715–729.     

A new psittaciform bird from the London Clay (Lower Eocene) of England

A new psittaciform bird from the Lower Eocene (Ypresian) London Clay of England is described. This taxon, Pulchrapollia gracilis gen. et sp. nov., is assigned to the order Psittaciformes (parrots) on the basis of several distinctive structures of the tarsometatarsus, namely the trochlea for metatarsal III (trochlea metatarsi III) bearing a tubercle on its lateral side and the trochlea for metatarsal IV (trochlea metatarsi IV) completely retroverted (fully zygodactyl foot). Comparisons with other fossil and Recent taxa further support this conclusion. Cladistic analysis shows that Pulchrapollia is the sister-taxon of the single extant family within Psittaciformes, the Psittacidae. Palaeopsittacus georgei, a taxon previously described from the London Clay, is most likely based on some unassociated material and is regarded here as incertae sedis.     

The phylogenetic relationships of the early Tertiary Primoscenidae and Sylphornithidae and the sister taxon of crown group piciform birds

The phylogenetic relationships of the early Tertiary Primoscenidae and Sylphornithidae are, for the first time, evaluated in a cladistic context. Both taxa include small arboreal birds with a permanently (Primoscenidae) or facultatively (Sylphornithidae) retroverted fourth toe. Primoscenidae were hitherto considered to be most closely related to either woodpeckers and allies (Piciformes) or to songbirds (Passeriformes), whereas the Sylphornithidae were classified into the roller-kingfisher-hornbill assemblage (Coraciiformes). Analysis of 56 morphological characters supports monophyly of a clade including Sylphornithidae and crown group Piciformes and results in sister group relationship between Passeriformes and a clade including Primoscenidae and the early Miocene Zygodactylidae. However, an analysis in which the search was constrained to trees supporting piciform affinities of the Primoscenidae resulted in trees that were only five steps longer than those from the primary analysis. The character evidence for each hypothesis is discussed. The systematic position of the Primoscenidae appears to be connected to the identity of the sister taxon of crown group Piciformes, as the primary search indicated Upupiformes (hoopoes and wood-hoopoes) and Bucerotiformes (hornbills) as sister taxa of Piciformes, whereas the constrained search resulted in sister group relationship between Coliiformes (mousebirds) and Piciformes. Songbirds do not show the slightest indication of a zygodactyl foot but in these birds the hindtoe is greatly elongated, an alternative strategy to increase the grasping capabilities of the foot. If Passeriformes are indeed the sister group of the clade (Primoscenidae + Zygodactylidae), these birds would be an example that, in closely related taxa, selection towards the same functional demands can result in entirely different morphological specializations.Communicated by F. Bairlein     

Pumiliornis tessellatus Mayr, 1999 revisited — new data on the osteology and possible phylogenetic affinities of an enigmatic Middle Eocene bird

Although the avian speciesPumiliornis tessellatus Mayr, 1999 is known from two skeletons from the Middle Eocene of Messel in Germany, its phylogenetic affinities remained enigmatic. The new osteological data presented in this study document thatP. tessellatus had an at least semizygodactyl foot, with a very wide basal phalanx of the fourth toe, and lacked an ossified pons supratendineus on the distal tibiotarsus. Compared to the known zygodactyl and semizygodactyl birds, this tiny Middle Eocene species resembles the late Eocene/early Oligocene taxonEocuculus Chandler, 1999. Anew, tentatively referred wing ofEocuculus from the early Oligocene of France is described and compared withPumiliornis.      

New specimens of the early Eocene bird Vastanavis and the interrelationships of stem group Psittaciformes

We describe new skeletal elements of Vastanavis from the early Eocene Cambay Shale Formation of western India, including a small coracoid that represents an unnamed new species, and comment on the relationships between this avian taxon and the recently described Avolatavis from the early Eocene Green River Formation in North America. Like the previously described ones, the new Vastanavis bones resemble those of the late Eocene Quercypsittidae, thus strengthening psittaciform affinities of the Indian taxon. Vastanavis differs from Avolatavis in the presence of a crista medianoplantaris on the tarsometatarsus and in claw morphology, but a fossil from the early Eocene London Clay, which was previously assigned to Vastanavidae, closely resembles Avolatavis in these features and all other osteological aspects. We show that most branches in a recent phylogeny of stem group Psittaciformes collapse after modification of a single erroneous character scoring for Vastanavis. We further describe a morphologically distinctive distal humerus of a small bird resembling the stem group nyctibiid Paraprefica, which was discovered in the most recent excavation in Vastan Lignite Mine.     

The developmental origin of zygodactyl feet and its possible loss in the evolution of Passeriformes

The zygodactyl orientation of toes (digits II and III pointing forwards, digits I and IV pointing backwards) evolved independently in different extant bird taxa. To understand the origin of this trait in modern birds, we investigated the development of the zygodactyl foot of the budgerigar (Psittaciformes). We compared its muscular development with that of the anisodactyl quail (Galliformes) and show that while the musculus abductor digiti IV (ABDIV) becomes strongly developed at HH36 in both species, the musculus extensor brevis digiti IV (EBDIV) degenerates and almost disappears only in the budgerigar. The asymmetric action of those muscles early in the development of the budgerigar foot causes retroversion of digit IV (dIV). Paralysed budgerigar embryos do not revert dIV and are anisodactyl. Both molecular phylogenetic analysis and palaeontological information suggest that the ancestor of passerines could have been zygodactyl. We followed the development of the zebra finch (Passeriformes) foot muscles and found that in this species, both the primordia of the ABDIV and of the EBDIV fail to develop. These data suggest that loss of asymmetric forces of muscular activity exerted on dIV, caused by the absence of the ABDIV, could have resulted in secondary anisodactyly in Passeriformes.     

Eocene fossil is earliest evidence of flower-visiting by birds

Birds are important pollinators, but the evolutionary history of ornithophily (bird pollination) is poorly known. Here, we report a skeleton of the avian taxon Pumiliornis from the middle Eocene of Messel in Germany with preserved stomach contents containing numerous pollen grains of an eudicotyledonous angiosperm. The skeletal morphology of Pumiliornis is in agreement with this bird having been a, presumably nectarivorous, flower-visitor. It represents the earliest and first direct fossil evidence of flower-visiting by birds and indicates a minimum age of 47 million years for the origin of bird–flower interactions. As Pumiliornis does not belong to any of the modern groups of flower-visiting birds, the origin of ornithophily in some angiosperm lineages may have predated that of their extant avian pollinators.     

Passerine diversity in the late Oligocene of Germany: earliest evidence for the sympatric coexistence of Suboscines and Oscines

A passerine avifauna from the late Oligocene ( c. 26–25 mya) of Germany was characterized by a high diversity of conspicuously small birds ranging in size from the smallest known Oscines to moderately small forms. The avifauna comprised both Oscines and Suboscines. Other passerine fragments showed such an unexpected mosaic of characters that it was impossible to assign them with certainty to any subordinate clade within the Passeriformes. The isolated remains of oscine passerines are the earliest evidence of this taxon in the Northern Hemisphere. Coexistence of oscine and suboscine passerines during the late Oligocene is also documented for the first time in the Northern Hemisphere. These finds reduce a major gap in the passerine fossil record and allow new insights into the composition and natural history of ancient avifaunas.     

Ultrastructure of the spermatozoon of Apus apus (Linnaeus 1758), the common swift (Aves; Apodiformes; Apodidae), with phylogenetic implications

The spermatozoon of Apus apus is typical of non‐passerines in many respects. Features shared with palaeognaths and the Galloanserae are the conical acrosome, shorter than the nucleus; the presence of a proximal as well as distal centriole; the elongate midpiece with mitochondria grouped around an elongate distal centriole; and the presence of a fibrous or amorphous sheath around the principal piece of the axoneme. The perforatorium and endonuclear canal are lost in A. apus as in some other non‐passerines. All non‐passerines differ from palaeognaths in that the latter have a transversely ribbed fibrous sheath whereas in non‐passerines it is amorphous, as in Apus, or absent. The absence of an annulus is an apomorphic but homoplastic feature of swift, psittaciform, gruiform and passerine spermatozoa. The long distal centriole, penetrating the entire midpiece, is a remarkably plesiomorphic feature of the swift spermatozoa, known elsewhere only in palaeognaths. The long centriole of Apus, if not a reversal, would be inconsistent with the former placement of the Apodiformes above the Psittaciformes from DNA–DNA hybridization. In contrast to passerines, in A. apus the microtubules in the spermatid are restricted to a transient single row encircling the cell. The form of the spermatozoon fully justifies the exclusion of swifts from the passerine family Hirundinidae.     

New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

A specimen of Eocuculus Chandler, 1999 (Aves, ? Cuculidae) from the early Oligocene of France

A postcranial skeleton of a small bird from the early Oligocene locality Pichovet in Southern France is described and identified as Eocuculus cf. cherpinae Chandler, 1999. It is the second fossil record of Eocuculus which was hitherto known from a postcranial skeleton from the late Eocene of North America only. Although Eocuculus shares some derived similarities with Cuculidae (cuckoos), it distinctly differs in a number of osteological features from crown group members of this taxon. If future, more complete skeletons prove its cuculiform affinities, Eocuculus is a stem lineage representative of this taxon and not within the crown group. Recognition of Eocuculus in the early Oligocene of France provides evidence for the presence of an extinct late Eocene/early Oligocene avian taxon with an intercontinental Northern Hemisphere distribution.     

A new kinosternoid from the Late Cretaceous Hell Creek Formation of North Dakota and Montana and the origin of the <Emphasis Type="Italic">Dermatemys mawii</Emphasis> lineage

A nearly complete turtle shell from the Late Cretaceous (Maastrichtian) Hell Creek Formation of Slope County, North Dakota, represents the most complete remains to date of a Mesozoic kinosternoid turtle and a new species, Hoplochelys clark nov. sp. The new taxon is diagnosable from other representatives of Hoplochelys by the plesiomorphic placement of the humeral/femoral sulcus behind the hyo/hypoplastral suture and the autapomorphic development of an interrupted median (neural) keel. All six previously named Paleocene (Puercan and Torrejonian) representatives of Hoplochelys lack diagnostic characters and are synonymized as Hoplochelys crassa. A phylogenetic analysis reveals that Hoplochelys spp. and Agomphus pectoralis are most parsimoniously placed within Kinosternoidea along the phylogenetic stem of the extant Mesoamerican River Turtle Dermatemys mawii, extending that taxon’s stem lineage from the early Eocene to the late Maastrichtian. The two primary crown lineages of Kinosternoidea are thus known from the Mesozoic and split prior to the late Campanian. The presence of a thickened cruciform plastron, true costiform processes, only three inframarginals, and the reduction of the medial contact of the abdominals are synapomorphies of Chelydroidea, the clade formed by Chelydridae and Kinosternoidae.     

Testing hypotheses about the sister group of the passeriformes using an independent 30-locus data set

Although many phylogenetic studies have focused on developing hypotheses about relationships, advances in data collection and computation have increased the feasibility of collecting large independent data sets to rigorously test controversial hypotheses or carefully assess artifacts that may be misleading. One such relationship in need of independent evaluation is the position of Passeriformes (perching birds) in avian phylogeny. This order comprises more than half of all extant birds, and it includes one of the most important avian model systems (the zebra finch). Recent large-scale studies using morphology, mitochondrial, and nuclear sequence data have generated very different hypotheses about the sister group of Passeriformes, and all conflict with an older hypothesis generated using DNA-DNA hybridization. We used novel data from 30 nuclear loci, primarily introns, for 28 taxa to evaluate five major a priori hypotheses regarding the phylogenetic position of Passeriformes. Although previous studies have suggested that nuclear introns are ideal for the resolution of ancient avian relationships, introns have also been criticized because of the potential for alignment ambiguities and the loss of signal due to saturation. To examine these issues, we generated multiple alignments using several alignment programs, varying alignment parameters, and using guide trees that reflected the different a priori hypotheses. Although different alignments and analyses yielded slightly different results, our analyses excluded all but one of the five a priori hypotheses. In many cases, the passerines were sister to the Psittaciformes (parrots), and taxa were members of a larger clade that includes Falconidae (falcons) and Cariamidae (seriemas). However, the position of Coliiformes (mousebirds) was highly unstable in our analyses of 30 loci, and this represented the primary source of incongruence among analyses. Mousebirds were united with passerines or parrots in some analyses, suggesting an additional hypothesis that needs to be considered in future studies. There was no clear evidence that base-compositional convergence, saturation, or long-branch attraction affected our conclusions. These results provide independent evidence excluding four major hypotheses about the position of passerines, allowing the extensive studies on this group to be placed in a more rigorous evolutionary framework.     

Phylogeny of major lineages of suboscines (Passeriformes) analysed by nuclear DNA sequence data

Phylogenetic relationships among major groups of passeriform birds were studied by analyses of nucleotide sequence data from two nuclear genes, c- myc and RAG-1. The results corroborated both the monophyly of the order Passeriformes, and the major dichotomy into oscine and suboscine passerines previously suggested based on syringeal morphology and DNA-DNA hybridizations. The representatives of the Old World suboscines (families Eurylaimidae, Philepittidae and Pittidae) formed a monophyletic clade. The New World suboscines clustered into two clades. The first contained Conopophaga (Conopophagidae), Furnarius (Furnariidae), Lepidocolaptes (Dendrocolaptidae), Thamnophilus (Formicariidae), and Rhinocrypta (Rhinocryptidae). Previously, the monophyly of this group has been inferred from their possession of a unique, "tracheophone" syrinx, and from DNA-DNA hybridisation data. The second clade of New World suboscines includes Gubernetes and Muscivora (Tyrannidae), Phytotoma (Phytotomidae), Tityra (Cotingidae) and Pipra (Pipridae). This group of families have been considered monophyletic based on morphology (although ambiguously) and DNA-DNA hybridisation. The sister group relationship of Tityra and Phytotoma supports the previously supposed cotingid affinity of Phytotoma . Nuclear DNA data also unambiguously group the lyrebirds Menura with the oscines.
The presented results from the analysis of nuclear DNA agree well with morphology and DNA-DNA hybridisation data. The precise age of the divergences studied herein are unknown but based on interpretations of the fossil record of passerine birds many of them might date back to the early Tertiary. The agreement between data from the nuclear DNA and other sources, along with the fact that neither of the studied genes showed sign of saturation, indicate the great potential of these two nuclear genes to resolve very old divergences in birds.     

Cryptosporidium proventriculi sp. n. (Apicomplexa: Cryptosporidiidae) in Psittaciformes birds

Cryptosporidiosis is a common parasitic infection in birds that is caused by more than 25 Cryptosporidium species and genotypes. Many of the genotypes that cause avian cryptosporidiosis are poorly characterized. The genetic and biological characteristics of avian genotype III are described here and these data support the establishment of a new species, Cryptosporidium proventriculi. Faecal samples from the orders Passeriformes and Psittaciformes were screened for the presence of Cryptosporidium by microscopy and sequencing, and infections were detected in 10 of 98 Passeriformes and in 27 of 402 Psittaciformes. Cryptosporidium baileyi was detected in both orders. Cryptosporidium galli and avian genotype I were found in Passeriformes, and C. avium and C. proventriculi were found in Psittaciformes. Cryptosporidium proventriculi was infectious for cockatiels under experimental conditions, with a prepatent period of six days post-infection (DPI), but not for budgerigars, chickens or SCID mice. Experimentally infected cockatiels shed oocysts more than 30 DPI, with an infection intensity ranging from 4,000 to 60,000 oocysts per gram (OPG). Naturally infected cockatiels shed oocysts with an infection intensity ranging from 2,000 to 30,000 OPG. Cryptosporidium proventriculi infects the proventriculus and ventriculus, and oocysts measure 7.4 × 5.8 μm. None of the birds infected C. proventriculi developed clinical signs.     

Identification of Potentially Human-Pathogenic Enterocytozoon bieneusi Genotypes in Various Birds

Enterocytozoon bieneusi was detected in 24 of 83 samples from birds of the orders Columbiformes, Passeriformes, and Psittaciformes. It was identical to or closely related to the Peru6 genotype, which was previously found in humans in Peru. Thus, various birds can be a significant source of environmental contamination by potentially human-pathogenic E. bieneusi.