On the Middle Miocene avifauna of Maboko Island,Kenya

The Middle Miocene sediments of Maboko Island (Lake Victoria) in western Kenya yielded numerous avian bones, which remained, however, little studied. The significance of this material is shown by the recent identification of an opisthocomiform bird. In the present study, further avian remains from Maboko Island are described. Most of the specimens belong to aquatic or semi-aquatic groups, of which some are closely related to taxa known from Early and Middle Miocene European avifaunas, that is, Nectornis cormorants (N. africanus nov. sp.) and Laricola-like Laromorphae. The fossil material also includes Ciconiidae (cf. Ciconia), Pelecanidae, Phoenicopteridae (Leakeyornis aethiopicus), Musophagidae, and a species of Ardeidae, which closely resembles the taxon Pikaihao from the Early Miocene of New Zealand. Some avian remains from Maboko Island belong to higher-level taxa unknown from the Middle Miocene of Europe. The occurrence of a giant Jacanidae (?Nupharanassa mabokoensis nov. sp.) is of particular interest, because these are globally absent in extant avifaunas and were previously only known from the Late Eocene/Early Oligocene of Egypt. Further unknown from contemporaneous European sites are small representatives of Jacanidae, Bucerotidae, and Alcedinidae, with the fossils of the latter two taxa being among the earliest published records of their respective groups. Several of the taxa that are common in contemporaneous European avifaunas have not been found in Maboko, and in spite of less pronounced climatic differences, Middle Miocene Afrotropical avifaunas already appear to have been distinct from contemporaneous European ones.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.     

Systematics of early and middle Miocene Old World monkeys

New information about the early cercopithecoids Prohylobates tandyi (Wadi Moghra, Egypt) and Prohylobates sp. indet. (Buluk and Nabwal, Kenya) is presented. Comparisons are made among all major collections of Early and Middle Miocene catarrhine monkeys, and a systematic revision of the early Old World monkeys is provided. Previous work involving the systematics of early Old World monkeys (Victoriapithecidae; Cercopithecoidea) has been hampered by a number of factors, including the poor preservation of Prohylobates material from North Africa and lack of comparable anatomical parts across collections. However, it is now shown that basal cercopithecoid species from both northern and eastern Africa can be distinguished from one another on the basis of degree of lower molar bilophodonty, relative lower molar size, occlusal details, symphyseal construction, and mandibular shape. Results of particular interest include: 1) the first identification of features that unambiguously define Prohylobates relative to Victoriapithecus; 2) confirmation that P. tandyi is incompletely bilophodont; and 3) recognition of additional victoriapithecid species.     

步林在甘肃党河流域塔奔布鲁克地区的早期工作记录——经典脊椎动物化石地点与现代地层框架的解译

1931-1932年博格·步林在甘肃省西部塔奔布鲁克地区(又名党河地区,位于肃北县城西南)首次发现丰富的新生代哺乳动物化石。步林在塔奔布鲁克盆地中部燕丹图沟中找到三个小哺乳动物化石异常丰富的层位。三个层位紧靠在一起,其中产出的哺乳动物群被命名为燕丹图动物群。步林明确指出燕丹图动物群为晚渐新世。该动物群以后成为晚渐新世塔奔布鲁克哺乳动物期的典型动物群并沿用至今。     

Biogeography of living and fossil Nitellopsis (Charophyta) in relationship to new finds from Morocco

Aim To document the distribution of two closely related macrophyte species, the fossil Nitellopsis (Tectochara) merianii (A. Braun ex Unger) Grambast & Soulié‐Märsche and Nitellopsis obtusa (Desvaux) J. Groves, the single living species of genus Nitellopsis and to draw the relationships between first finds of N. merianii in North Africa and previously known records. Location South part of the High Atlas Range, Aït Kandoula basin, Morocco. Methods New finds of fossil charophyte gyrogonites were determined and illustrated. Using a large bibliographic database of both fossil and extant charophytes, we depict the biogeographical distribution N. merianii and N. obtusa and summarize the ecological requirements of N. obtusa as a modern analogue. Results The Aït Kandoula deposits, dated by magnetostratigraphy and vertebrate fossils to Upper Miocene, provided the first fossil populations of N. merianii in Africa. The distribution patterns of the Tertiary N. merianii and the living N. obtusa superimpose on a large area that extends from Spain to East Asia and characterize them as typically Eurasian taxa indicative for permanent, relatively cold and deep freshwater lakes. During the Quaternary, similar palaeolakes in the Sahara were colonized by N. obtusa. Main conclusions The presence of the Eurasian species N. merianii in North Africa is the result of dispersal by migratory waterbirds and attests to N–S migration occurring during the Upper Miocene (10–5 Myr ago). Gyrogonites imported from Southern Europe could germinate and fulfil their life cycle so as to form a new seed bank of gyrogonites because they had been deposited in a suitable environment consisting of a freshwater mountain lake.     

First macroflora data from La Val (Late Oligocene/Early Miocene), Estadilla (Huesca,Spain)

This paper reports a previously unknown leaf-flora from the Upper Oligocene/Lower Miocene of the Ebro Basin, NE Spain, a period with a relatively poor vascular-plant fossil record in Southern Europe. The presence of Acrostichum sp. is also important. This fern is extremely significant from the point of view of palaeoecology and the depositional environment. The macroflora appears to yield sufficient morphological characteristics to be identified at genus level, and sometimes at species level, although cuticles are not preserved. This article presents the first data obtained from the new outcrop at La Val; the following families have been identified: Pteridaceae, Dennstaedtiaceae, Equisetaceae, Pinaceae, Lauraceae, Hamamelidaceae, Betulaceae, Myricaceae and Salicaceae. The fossil plant assemblage is correlated with the Cadibona floristic complex (Mai, Tertiäre Vegetationsgeschichte Europas. Methoden und Ergebnisse, Gustav Fischer, Jena, 691 pp., 1995) and suggests a subtropical-to-warm temperate climate, rainy and wet, with a short dry season. The age of the assemblage is Late Oligocene/Early Miocene.     

内蒙古中部敖尔班地区的岩石及生物地层(英文)

中国北方新近纪地层中,早中新世的哺乳动物化石地点比较稀少,化石面貌也不十分清楚。在内蒙古中部,尽管通古尔台地上有研究历史近百年的新近纪经典地点,但上世纪90年代发现的嘎顺音阿得格,依然是内蒙古目前惟一产出早中新世动物群的小盆地。同内蒙古诸多其他新近纪哺乳动物群一样,嘎顺音阿得格动物群与相邻动物群彼此缺乏时空上的联系,还不易建立可靠的层序关系。位于苏尼特左旗东南约60 km的敖尔班(曾用名"奥尔班",见Liddicoat et al.,2007),红色地层大面积出露,是2004年发现的一个新的哺乳动物化石地点。敖尔班剖面总厚50余米,大、小哺乳动物化石共生,岩性特征易识别,有利层位对比,是研究生物地层学的理想地点。自2004年以来,我们连续在敖尔班地区采集化石并进行地层工作,建立了一个较完整的哺乳动物序列,时代跨越早中新世晚期到晚中新世晚期。更加难得的是,敖尔班同一剖面上含有4个哺乳动物化石层位,其上下层序关系一目了然,在内蒙古新近纪地层中仅此一例。敖尔班哺乳动物序列的建立,无疑将对整个内蒙古中部新近纪,尤其是早中新世动物群面貌的了解具有促进作用。本文着重对敖尔班岩石地层进行描述,并结合我们4年来对脊椎动物化石的积累与认识,试图进一步完善内蒙古中部地区的生物地层层序,进而建立这一地区的年代地层框架。对化石的详细描述将适时发表。根据岩性及接触关系,敖尔班剖面可分为三大段:敖尔班组、巴伦哈拉根层及必鲁图层。敖尔班组(新建组)由一套红色和绿色的泥岩及粉砂岩组成,发育有成熟的古土壤层,厚约42 m,时代属于早中新世中晚期。敖尔班组的层型剖面建立在敖尔班露头出露最厚的中部,下部未见底,顶部与上覆巴伦哈拉根层呈假整合或不整合接触。该组目前已知分布仅局限于敖尔班露头。敖尔班组可进一步划分出三段:下红泥岩段、中绿泥岩段及上红泥岩段。三段呈连续沉积。敖尔班组与巴伦哈拉根层之间的假整合所代表的沉积间断延续了中中新世的大部分时段。巴伦哈拉根层为一套橘红色砂岩、粉砂岩及底砾岩,时代大致是最晚中中新世至最早晚中新世。不整合于巴伦哈拉根层之上是必鲁图层。两层之间似乎缺失了晚中新世的大部分堆积。必鲁图层的底砾岩是一种切割与充填构造,其中所含钙质结核及相当数量的化石都可能是巴伦哈拉根层原生堆积物再沉积的结果。必鲁图层的分布还需做更多工作,其时代可能是晚中新世晚期。主要依据小哺乳动物的组合,在敖尔班剖面中可建立4个动物群。最早为敖尔班组下红泥岩段产出的下敖尔班动物群。该动物群的特征是,小哺乳动物中在渐新世十分兴旺的一些科,如Ctenodactylidae,Tachyoryctoididae,Aplodontidae和Zapodidae还相当繁荣;中新世出现的属,如Mioechinus,Keramidomys,Heterosminthus和Democricetodon等占动物群总量的半数以上;大哺乳动物中残留有Palaeogale和裂爪兽。比下敖尔班动物群稍晚的是上敖尔班动物群,产出于敖尔班组的上红泥岩段。上敖尔班动物群的特征是,小哺乳动物在渐新世中占统治地位的一些科或完全绝迹,如Ctenodactylidae,或在种类和数量上明显减退,如Aplodontidae和Zapodidae;缺少在下敖尔班动物群中还相当常见的一些古老属,如Amphechinus,Tachyo- ryctoides和Sinolagomys等;出现了下敖尔班动物群中所没有的Megacricetodon,Cricetodon和Alloptox属;大哺乳动物中出现了长鼻类和柄杯鹿(Ligeromeryx/Lagomeryx)。经过相当长的一个沉积间断,敖尔班剖面的上部出现了巴伦哈拉根动物群。该动物群中渐新世常见的小哺乳动物科进一步衰落,同时出现了亚洲古北界晚中新世以后常见的跳鼠科(Dipodidae)和鼢鼠科(Siphneidae),具有明显的中中新世晚期或晚中新世早期生物组合的特点。最后是敖尔班剖面顶部必鲁图层中的必鲁图动物群。必鲁图动物群显然带有中中新世及晚中新世的混合特征,很可能是水流作用再沉积的结果。根据其中最进步分子的成分判断,估计必鲁图动物群的年代是晚中新世的晚期,其特征是,鼠科(Muridae)动物高度分化,而野兔科(Leporidae)尚未出现;含有晚中新世宝格达乌拉动物群或最晚中新世二登图动物群中大量出现的属种,如Lophocricetus grabaui,Paralactaga suni,Dipus fraudator和Hansdebruijnia pusilla等;但二登图动物群中很繁荣的一些属在必鲁图动物群中未被发现或者发现的个体数量很少,如Prospermophilus, Paralophicricetus和Microtodon等。     

Miocene spiders in Dominican amber (Oonopidae,Mysmenidae)

Oonops seldeni sp. nov. and Mysmenopsis lissycoleyae sp. nov. are described from male specimens preserved in Miocene amber from the Dominican Republic and are the first fossil records of these extant genera. Gamasomorpha incerta Wunderlich, 1988 is transferred to Stenoonops. Two females previously described as Orchestina dominicana? Wunderlich, 1981 are transferred to Orchestina sp. indet., and one new specimen of a male of Orchestina dominicana Wunderlich, 1988 is identified. Oonopidae is reported for the first time from Cretaceous ambers of Burma, Lebanon and New Jersey. These are the oldest records of this family, extending the known range from the Rupelian (Oligocene) to the Valanginian (Lower Cretaceous). Recent StenoonopsMysmenopsis are unknown from Hispaniola. The presence of these genera in amber suggests that they will be found. The presence of Mysmenopsis in amber is further evidence of kleptoparasitic/commensal spiders in the Miocene.     

The flora of the early Miocene Brandon Lignite, Vermont, USA. VIII. Caldesia (Alismataceae)

Caldesia, a genus of aquatic monocotyledons, is represented by four living species, which are widely distributed in the temperate and tropical Old World. The genus has an extensive Oligocene through Pleistocene fossil record in Eurasia. We survey the morphology of the extant and fossil fruits of the Alismataceae, and provide a detailed review of the morphology and anatomy of living and fossil Caldesia fruits. The latter exhibit substantial similarity, making the recognition of separate species on the basis of fruit morphology difficult. We erect the new species Caldesia brandoniana from the Early Miocene Brandon Lignite of Vermont primarily on the basis of its geographic isolation; careful revision of all fossil fruiting material of Caldesia might require placement of the Brandon specimens in a more inclusive form species. Together with leaves of Caldesia from the Miocene Clarkia flora of Idaho, this occurrence indicates that Caldesia was in the New World as recently as the Early Miocene.     

New genus of leaf-mimicking katydids (Orthoptera: Tettigoniidae) from the Late Eocene–Early Oligocene of France and England

A new leaf-mimicking katydid (Orthoptera: Tettigoniidae: Pseudophyllinae), Archepseudophylla fossilis gen. et sp. nov. is described based on tegmina from the Early Oligocene of France. Lithymnetes laurenti Théobald, 1937 from the Late Eocene of France is transferred to Archepseudophylla comb. nov. The enigmatic ‘Poekilloptera’ melanospila Cockerell, 1921 from the Early Oligocene of the Isle of Wight (UK) is transferred to Archepseudophylla comb. nov. The present discoveries in the Palaeogene of the Isle of Wight, Céreste, and Marseilles Basins represent the first fossil records of Pseudophyllinae. In addition, these records support warm and humid forest conditions near the palaeolakes of the Isle of Wight, Camoins-les-Bains, and Céreste.     

Exceptionally preserved North American Paleogene metatherians: adaptations and discovery of a major gap in the opossum fossil record

A major gap in our knowledge of the evolution of marsupial mammals concerns the Paleogene of the northern continents, a critical time and place to link the early history of metatherians in Asia and North America with the more recent diversification in South America and Australia. We studied new exceptionally well-preserved partial skeletons of the Early Oligocene fossil Herpetotherium from the White River Formation in Wyoming, which allowed us to test the relationships of this taxon and examine its adaptations. Herpetotheriidae, with a fossil record extending from the Cretaceous to the Miocene, has traditionally been allied with opossums (Didelphidae) based on fragmentary material, mainly dentitions. Analysis of the new material reveals that several aspects of the cranial and postcranial anatomy, some of which suggests a terrestrial lifestyle, distinguish Herpetotherium from opossums. We found that Herpetotherium is the sister group to the crown group Marsupialia and is not a stem didelphid. Combination of the new palaeontological data with molecular divergence estimates, suggests the presence of a long undocumented gap in the fossil record of opossums extending some 45Myr from the Early Miocene to the Cretaceous.     

Oldest known avian footprints from Australia: Eumeralla Formation (Albian), Dinosaur Cove,Victoria

Two thin‐toed tridactyl tracks in a fluvial sandstone bed of the Eumeralla Formation (Albian) at Dinosaur Cove (Victoria, Australia) were likely made by avian trackmakers, making these the oldest known fossil bird tracks in Australia and the only Early Cretaceous ones from Gondwana. These tracks, which co‐occur on the same surface with a slightly larger nonavian theropod track, are distinguishable by their anisodactyl form, hallux impressions and wide digit II–IV divarication angles. A lengthy hallux impression and other deformational structures associated with one track indicate foot movement consistent with an abrupt stop, suggesting its tracemaker landed after either flight or a hop. The single nonavian theropod track is similar to other tracks described from the Eumeralla Formation at another locality. The avian footprints are larger than most Early Cretaceous avian tracks recorded worldwide, indicating sizeable enantiornithine or ornithurine species in formerly polar environments of Australia. The avian tracks further supplement scant body fossil evidence of Early Cretaceous birds in southern Australia, which includes a furcula from the Wonthaggi Formation. Because of this discovery, Dinosaur Cove, previously known for its vertebrate body fossils, is added to a growing list of Early Cretaceous vertebrate tracksites in southern Australia.     

A new coral‐associated decapod assemblage from the Upper Miocene (Messinian) Upper Coralline Limestone of Malta (Central Mediterranean)

Abstract: A rich coral‐associated decapod assemblage is recorded from the ‘Depiru Beds’ of the upper part of the Upper Coralline Limestone (Messinian, Upper Miocene), from the island of Malta. Nineteen species within 17 genera have been discovered, where 14 genera are new for Malta. Four new species are described, namely Micippa annamariae sp. nov., Pilumnus scaber sp. nov., Panopeus muelleri sp. nov. and Herbstia melitense sp. nov. Herbstia melitense sp. nov. constitutes the first record of the genus from the fossil record in the Mediterranean region. This discovery more than doubles the number of known fossil decapod species from Malta. The fossil bivalve Jouannetia (J.) semicaudata Des Moulins, 1830 and the extant decapod Maja goltziana D’Oliveira, 1888, are also recorded for the first time from Malta. Other Neogene coral‐associated decapod assemblages are investigated and correlated with the new assemblage from Malta. The migration of taxa between the Mediterranean region and the Paratethys, particularly during the Lower Badenian (Langhian), is evidenced by the strong affinity of the Maltese decapod assemblage with that of the Middle Miocene Badenian assemblages from Hungary, Poland and Ukraine. Upper Miocene, Messinian assemblages from Spain, Algeria and Morocco are also similar to that from Malta.     

'First' appearances in the Cenozoic land-mammal record of the Greater Antilles: significance and comparison with South American and Antarctic records

Aim Through analysis of fossil records, the aim of this paper is to show that fossil representatives of at least three land‐mammal clades (pitheciine atelid primates, heteropsomyine echimyid rodents, and megalonychid phyllophagan xenarthrans) that once lived in the Greater Antilles are as old as, if not older than, ‘first’ occurrences of these same groups on the South American mainland. Location Greater Antilles, South America, Antarctic Peninsula. Methods Analysis of Cenozoic land‐mammal fossil records for the three areas. Results Comparison reveals an interesting similarity to the Tertiary vertebrate palaeontological record for the Antarctic Peninsula (Seymour Island), in the sense that the latter also includes early (Eocene) representatives of some typical ‘South American’ groups (e.g. meridiungulates, sloths, certain marsupial groups). Conclusions Given how limited the Antillean and Antarctic records are in quantity and quality, it seems unlikely that these ‘first’ appearances have much bearing on real origins (basal divergences). Rather, it suggests that the fossil basis for interpreting the origin and earliest diversification of ‘South American’ clades during the latest Cretaceous/early Cenozoic is probably even scantier than generally realized. In particular, the Antillean record strengthens arguments that some crown‐group continental lineages are considerably older than fossil evidence currently allows – a point increasingly (if unevenly) supported by molecular studies of many of the same clades.     

Warm and humid Trans-Himalaya during the late Miocene: plant fossil evidence

Neogene fossil records from the Indus Basin sedimentary rocks (IBSR), deposited in the Indus Tsangpo Suture Zone (ITSZ), are very rare, but are important to understand the history of plant diversity and paleoclimate in the Himalaya. We report fossil wood ascribed to Ebenoxylon siwalicus Prakash from late Miocene sediments of the Karit Formation belonging to ITSZ. The anatomical details of the fossil wood, such as small to medium-sized vessels occluded with tyloses, scanty paratracheal to diffuse-in-aggregate axial parenchyma, 1–3 seriate homo to heterocellular rays, bordered intervessel pits with lenticular apertures and simple perforations, suggest its close affinity with Diospyros Linnaeus of the family Ebenaceae. Further anatomical details suggest a close resemblance with extant D. ehretioides Don and D. macrophylla Blume. The present fossil, along with previously known fossil records of Lagerstroemia (Lythraceae) and palms, indicate that the Trans-Himalaya was warm and humid during the late Miocene, quite different from the modern cool and dry climate in the study area.     

The origin and early evolution of birds

Birds evolved from and are phylogenetically recognized as members of the theropod dinosaurs; their first known member is the Late Jurassic Archaeopteryx, now represented by seven skeletons and a feather, and their closest known non-avian relatives are the dromaeosaurid theropods such as Deinonychus. Bird flight is widely thought to have evolved from the trees down, but Archaeopteryx and its outgroups show no obvious arboreal or tree-climbing characters, and its wing planform and wing loading do not resemble those of gliders. The ancestors of birds were bipedal, terrestrial, agile, cursorial and carnivorous or omnivorous. Apart from a perching foot and some skeletal fusions, a great many characters that are usually considered ‘avian’ (e.g. the furcula, the elongated forearm, the laterally flexing wrist and apparently feathers) evolved in non-avian theropods for reasons unrelated to birds or to flight. Soon after Archaeopteryx, avian features such as the pygostyle, fusion of the carpometacarpus, and elongated curved pedal claws with a reversed, fully descended and opposable hallux, indicate improved flying ability and arboreal habits. In the further evolution of birds, characters related to the flight apparatus phylogenetically preceded those related to the rest of the skeleton and skull. Mesozoic birds are more diverse and numerous than thought previously and the most diverse known group of Cretaceous birds, the Enantiornithes, was not even recognized until 1981. The vast majority of Mesozoic bird groups have no Tertiary records: Enantiornithes, Hesperornithiformes, Ichthyornithiformes and several other lineages disappeared by the end of the Cretaceous. By that time, a few Linnean ‘Orders’ of extant birds had appeared, but none of these taxa belongs to extant ‘families’, and it is not until the Paleocene or (in most cases) the Eocene that the majority of extant bird ‘Orders’ are known in the fossil record. There is no evidence for a major or mass extinction of birds at the end of the Cretaceous, nor for a sudden ‘bottleneck’ in diversity that fostered the early Tertiary origination of living bird ‘Orders’.     

A succession of Miocene rodent assemblages from fissure fillings in southern France: palaeoenvironmental interpretation and comparison with Spain

An Early to Late Miocene sequence of rodent assemblages from southern France has been quantitatively studied. The resulting pattern seems very similar to a contemporary sequence from central Spain (Calatayud–Teruel Basin). The fossil mammal-bearing localities are of different types: mainly karst infills in France and localities situated in sedimentary basins in Spain. In order to interpret the fossil record, a comparison has been made between southern France faunas of similar age but collected in karst infills and in basin deposits. There seems to be no difference between the two kinds of faunas and thus there is no indication that karst infills systematically give a picture of drier and more open environments. Both types of localities may give a similar relative abundance of taxa and when differences exist they can be attributed to local conditions. The comparison between southern France and the Calatayud–Teruel Basin (central Spain) shows that: (1) similar trends occurred in the two areas; (2) differences between spectra were more important during the late Early Miocene than during the Middle Miocene; (3) the shift between the late Early Miocene and the Middle Miocene environments in southern France does not seem to be correlated with a general drop in temperatures as inferred from the analysis of central Spain faunas.     

Tempo and mode of hummingbird evolution

Lack of adequate historical data has hindered understanding of the evolutionary tempo and mode of many ecologically well-characterized avian radiations. DNA hybridization distances among 28 hummingbirds (Trochilidae) were used to establish a timescale for this family's radiation into more than 330 species. Under a variety of analytical assumptions, genetic distances calibrated with a fossil divergence date corrected for incompleteness in the geologic record indicated that all extant hummingbird lineages began to diverge in the Early Miocene, approximately 40 Myr (million years) after the Paleocene date estimated for the divergence of hummingbirds and swifts. The long period prior to the radiation of living forms provides ample time for divergent evolution to produce the large morphological gap that has tended to obscure the sister-relationship of hummingbirds and swifts. The Miocene radiation of extant hummingbird lineages itself began with the divergence of the hermit and nonhermit subfamilies approximately 17 Ma (million years ago), followed by the rapid divergence of two Andean and one principally Central and North American clade at approximately 12 Ma. Younger subsidiary lineages, including ones found mainly in the Andes or in North America, date to the later Miocene-earlier Pliocene, approximately 6 Ma. The DNA hybridization-based chronology thus indicates a protracted, rather than stricdy rapid, radiation. Evidence from a broader spectrum of organisms supports the general pattern that higher taxonomic structure within many extant continental families evolved in the Miocene, suggesting that a common environmental pacemaker initiated radiation in unrelated groups. Compared to those in the Pleistocene, radiations tracing to the Miocene may have depended less on rapid climate cycling than on creation of new habitats by major geologic and climatic upheavals. For extant hummingbirds, a principal cause for their Miocene diversification probably was the ability of the ecologically generalized subfamily of nonhermits to radiate in montane areas created by the Andean and other orogenies. Similar interactions between new habitats and their exploitation by ecological generalists may explain, at least in part, the contemporaneous radiation of Passeriformes, the most diverse avian order.     

The First Occurrence in the Fossil Record of an Aquatic Avian Twig-Nest with Phoenicopteriformes Eggs: Evolutionary Implications

Background

We describe the first occurrence in the fossil record of an aquatic avian twig-nest with five eggs in situ (Early Miocene Tudela Formation, Ebro Basin, Spain). Extensive outcrops of this formation reveal autochthonous avian osteological and oological fossils that represent a single taxon identified as a basal phoenicopterid. Although the eggshell structure is definitively phoenicopterid, the characteristics of both the nest and the eggs are similar to those of modern grebes. These observations allow us to address the origin of the disparities between the sister taxa Podicipedidae and Phoenicopteridae crown clades, and traces the evolution of the nesting and reproductive environments for phoenicopteriforms.

Methodology/Principal Findings

Multi-disciplinary analyses performed on fossilized vegetation and eggshells from the eggs in the nest and its embedding sediments indicate that this new phoenicopterid thrived under a semi-arid climate in an oligohaline (seasonally mesohaline) shallow endorheic lacustine environment. High-end microcharacterizations including SEM, TEM, and EBSD techniques were pivotal to identifying these phoenicopterid eggshells. Anatomical comparisons of the fossil bones with those of Phoenicopteriformes and Podicipediformes crown clades and extinct palaelodids confirm that this avian fossil assemblage belongs to a new and basal phoenicopterid.

Conclusions/Significance

Although the Podicipediformes-Phoenicopteriformes sister group relationship is now well supported, flamingos and grebes exhibit feeding, reproductive, and nesting strategies that diverge significantly. Our multi-disciplinary study is the first to reveal that the phoenicopteriform reproductive behaviour, nesting ecology and nest characteristics derived from grebe-like type strategies to reach the extremely specialized conditions observed in modern flamingo crown groups. Furthermore, our study enables us to map ecological and reproductive characters on the Phoenicopteriformes evolutionary lineage. Our results demonstrate that the nesting paleoenvironments of flamingos were closely linked to the unique ecology of this locality, which is a direct result of special climatic (high evaporitic regime) and geological (fault system) conditions.