Phylogeny of Carpinus and subfamily Coryloideae (Betulaceae) based on chloroplast and nuclear ribosomal sequence data

Phylogenetic studies were conducted for Carpinus and the subfamily Coryloideae (Betulaceae) using sequences of the chloroplast matK gene, the trnL-trnF region (trnL intron, and trnL [UAA] 3' exon-trnF [GAA] intergenic spacer) and the psbA-trnH intergenic spacer, and the nuclear ribosomal ITS regions. The combined analyses of the three chloroplast regions suggest that Coryloideae is monophyletic; Ostryopsis is sister to the Carpinus - Ostrya clade; Corylus is monophyletic and sister to the Ostrya - Carpinus - Ostryopsis clade; Ostrya is paraphyletic; and within Carpinus, species of sect. Carpinus from eastern Asia form a monophyletic group, whereas the positions of C. betulus from Europe and C. caroliniana from eastern North America are unresolved within the Carpinus clade. The cpDNA tree generated in this study is largely congruent with the previously published ITS results, but the ITS tree places Carpinus sect. Distegocarpus as sister to the Ostrya - Carpinus sect. Carpinus clade. Future work is needed to examine the relationships within the Ostrya - Carpinus clade, evaluate the generic status of Ostrya, and test the phylogenetic position of Ostryopsis.     

Phylogeny and evolution of the Betulaceae as inferred from DNA sequences,morphology, and paleobotany

Phylogeny of the Betulaceae is assessed on the basis of rbcL, ITS, and morphological data. Based upon 26 rbcL sequences representing most “higher” hamamelid families, the Betulaceae are monophyletic, with Casuarinaceae as its sister group, regardless of whether the outgroup is Cunoniaceae, Cercidiphyllaceae, Hamamelidaceae, or Nothofagus. Within the Betulaceae, two sister clades are evident, corresponding to the subfamilies Betuloideae and Coryloideae. However, with only 13 phylogenetically informative sites, the rbcL sequences provide limited intra-subfamilial resolution. Internal transcribed spacer (ITS) sequences provided 96 phylogenetically informative sites from 491 aligned sites resulting in a single most parsimonious tree of 374 steps (consistency index = 0.791) with two major lineages corresponding to the two traditional subfamilies: Betuloideae (Alnus, Betula) and Coryloideae (Corylus, Ostryopsis, Carpinus, Ostrya). This arrangement is mostly consistent with those from rbcL and morphology and is greatly reinforced by analyses with the three data sets combined. In the Coryloideae, the Ostryopsis–Carpinus–Ostrya clade is well supported, with Corylus as its sister group. The sister-group relationship between Ostryopsis and the Carpinus–Ostrya clade is well supported by ITS, rbcL, and morphological data. Phylogenetic relationships among the extant genera deduced by these analyses are compatible with inferences from ecological evolution and the extensive fossil record.     

Comparative Flower and Inflorescence Organogenesis among Genera of Betulaceae: Implications for Phylogenetic Relationships

Betulaceae have simple flowers but complex inflorescences. Recent phylogenetic analyses using molecular data have produced robust phylogenetic trees of Betulaceae. In this study, we evaluated the phylogenetic value of comparative organogenetic data of reproductive organs in the context of molecular phylogenies. Flower and inflorescence developmental processes of 21 species from all six genera in Betulaceae were documented with scanning electron microscopy. In each pistillate cyme, there are one primary bract, two secondary bracts, and two or three flowers in the six genera; only in Alnus are there two tertiary bracts on the abaxial side. The pistillate flower of all genera but Alnus has tepal primordia. Two tepals stop developing early on, resulting in the lack of tepals in mature flowers of Betula; while the tepals are initiated from a common circular primordium at the base of pistil in Corylus, Ostryopsis, Carpinus, and Ostrya, and the developed tepals with irregular shape and unstable number of lobes are adnate to the top of the pistil. In staminate organogenesis, each cyme has one primary bract and three flowers in all genera; two secondary bracts are only present in Alnus, Betula, and Corylus. Staminate flowers have no tepals except in Alnus and Betula, and exhibit high variation in number of stamens among genera. The number of secondary and tertiary bracts in each pistillate and staminate cyme, as well as the presence of tepals in pistillate and staminate flowers was clarified in all genera. Micro-morphological characters were used to infer the phylogenetic relationships of genera and sections of Betulaceae. Our analyses support the division of two subfamilies: Betuloideae (Alnus and Betula) and Coryloideae (Corylus, Carpinus, Ostrya, and Ostryopsis), and three tribes: Betuleae (Alnus and Betula), Coryleae (Corylus), and Carpineae (Carpinus, Ostrya, and Ostryopsis). The results agree with those from molecular phylogenetic studies, and suggest that micro-morphological characters are phylogenetically informative in Betulaceae, and reproductive organs of Betulaceae have evolved in the direction of reduction in bracts and tepals.     

Leaf Survival and Evolution in Betulaceae

Studies on the changes in leaf number of 12 species of Betulaceaewere carried out by repeated observations during several years.The inner structure of the winter buds of these species wasalso examined. The periods of leaf emergence were long in Alnusand Betula, intermediate in Corylus and short in Ostrya andCarpinus. A heavy summer leaf fall is characteristic only ofthe Gymnothyrsus subgenus of Alnus. One lamina and two stipulesis the basic unit constituting the winter buds. Only the budsof Gymnothyrsus are composed of several of these units. In theAlnaster subgenus of Alnus, Betula and Corylus, there are oneor two scales that seem to have originated from the two stipulesremaining after reduction of the lamina. Eight and 24 scalesof similar origin were found in Ostrya and Carpinus respectively.A common ancestral species having the following primitive charactersis proposed: shoot constituted only of the equal-sized units,period of leaf emergence long, and leaf fall usually occurs.Alnus (Gymnothyrsus) is assumed to be the most primitive typeas it has many characters similar to those of the proposed ancestralspecies. Carpinus is assumed to be the most advanced group.The leaf survivorship curve is assumed to have changed fromthe primitive bell-shaped to an advanced trapezoid concurrentwith an increase in the number of bud scales. Betulaceae, Alnus, Betula, Carpinus, Corylus, Ostrya, leaf survival, leaf emergence, stipules, winter buds, evolution     

Fossil wood of <Emphasis Type="Italic">Betula erkovetskiensis</Emphasis> sp. nov. (Betulaceae) from the Miocene of the Erkovetskii Brown Coal Field,Amur Region,Russia

A new species, Betula erkovetskiensis Blokhina et O.V. Bondarenko (Betulaceae), from the deposits of the Sazanka Formation (upper Middle?Upper Miocene) of the Erkovetskii Brown Coal Field (Amur Region, Russia) is described based on anatomical features of fossil wood. The new species shows some wood anatomical characters of the extant birch subgenus Betula, B. davurica, B. nigra (section Dahuricae), and B. papyrifera (section Betula). Fossil wood of Betula is found in the Amur Region for the first time.     

The Miocene floras of Iceland and their significance for late Cainozoic North Atlantic biogeography

A large number of plant macrofossils from several Middle to Upper Miocene localities from Iceland have been studied. The fossil material includes four ferns and fern allies, seven conifers, and about 40 species of flowering plants. Betula islandica and Salix gruberi are described as new species. Coniferous twigs previously ascribed to the genus Sequoia are shown to belong to Cryptomeria based on macro‐morphological and epidermal features. Fossil plants from Iceland are compared with coeval fossil taxa from Europe and North America and with living plants. The main finding is that the Miocene flora of Iceland belongs to a widespread Neogene northern hemispheric floral type including plants whose representatives are restricted to East Asia, North America and to western Eurasia at the present time. Previously inferred conspicuous similarities to North American modern equivalents appear to be misleading. The type of vegetation in four plant‐bearing sedimentary formations from the late Mid Miocene to Late Miocene, the 12 Ma Brjánslækur‐Seljá Formation, the 10 Ma Tröllatunga‐Gautshamar Formation, the 9–8 Ma Skarðsströnd‐Mókollsdalur Formation, and the 7–6 Ma Hreðavatn‐Stafholt Formation, corresponds to a humid temperate broadleaved (deciduous)–coniferous mixed forest dominated by Betulaceae, Fagaceae and Acer. Changes in species composition in the sedimentary formations reflect a shift from warm temperate (Cfa climate) to cool temperate (Cfb climate) conditions from the late Mid Miocene to the latest Miocene. This shift was connected to repeated phases of extinction and colonization. Specifically, one set of thermophilic taxa including Magnolia, Liriodendron, Sassafras and Comptonia went extinct between 12 and 10 Ma, and appears to have been replaced by another set of thermophilic taxa in the 10 Ma formation (Juglandaceae aff. Pterocarya/Cyclocarya, Rhododendron ponticum type). The 9–8 and 7–6 Ma formations are characterized by taxa that migrated to Iceland from Europe, such as Fagus gussonii, Betula cristata and Pterocarya fraxinifolia type. Although there is convincing evidence that plants colonized Iceland both from North America and Europe until 12 Ma, migration in the younger formations (9–8, 7–6 Ma) is suggested to have occurred mainly from Europe. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149 , 369–417.     

Modelling late Quaternary changes in plant distribution, vegetation and climate using pollen data from Georgia, Caucasus

Aim To use pollen data, numerical analysis and modelling to reconstruct late Quaternary vegetation and climate in a complex, mountainous environment. Location Georgia (Caucasus region). Methods Pollen data were assembled from various sources and used to map: (1) changing frequencies of individual taxa; (2) vegetation changes; and (3) reconstructed climatic parameters for the past 14,000 years. Numerical analyses were performed using two‐way indicator species analysis (twinspan ), detrended correspondence analysis (DCA), the modern analogue technique (MAT) and weighted averaging (WA). Results Mapping of pollen taxa showed that Chenopodiaceae, Artemisia and Ephedra were most abundant in the study area during the late‐glacial. Betula and Corylus expanded during the early Holocene, yielding to Abies, Carpinus, Fagus, Quercus and Castanea. Picea, Pinus, Juglans and Ostrya‐type expanded during the late Holocene. Mixed forests grew in the moist, Black Sea refugium throughout the late Quaternary. Elsewhere in Georgia, the Pleistocene–Holocene transition is recorded as a shift from desert‐steppes to oak‐xerophyte communities and mixed forests. This kind of vegetation remained relatively stable until the mid–late Holocene, when coniferous forests and mountain grasslands advanced. DCA showed that rainfall was most strongly correlated with pollen composition in the study area (r² = 0.55). No temperature signal was detected. A weighted‐averaging transfer function linking pollen percentages to annual precipitation was selected over a MAT model as it performed better when applied to a validation data set. Rainfall reconstructions indicate widespread aridity at the terminal Pleistocene, followed by a gradual increase in precipitation, peaking during the mid Holocene (7000–4000 cal. yr bp ) and generally decreasing thereafter. Main conclusions On a regional scale, the results confirm those from previous studies of palaeovegetation and palaeoclimate in Western Asia. On a local scale, reconstructions from individual sites often diverge from the regional trend because of edaphic changes, ecological succession, human impacts and other disturbances. Some of these factors are probably responsible for the increasing heterogeneity of Georgia’s vegetation in the latter half of the Holocene.     

Phylogeographic diversification of antelope squirrels (Ammospermophilus) across North American deserts

We investigated the biogeographic history of antelope squirrels, genus Ammospermophilus, which are widely distributed across the deserts and other arid lands of western North America. We combined range‐wide sampling of all currently recognized species of Ammospermophilus with a multilocus data set to infer phylogenetic relationships. We then estimated divergence times within identified clades of Ammospermophilus using fossil‐calibrated and rate‐calibrated molecular clocks. Lastly, we explored generalized distributional changes of Ammospermophilus since the last glacial maximum using species distribution models, and assessed responses to Quaternary climate change by generating demographic parameter estimates for the three wide‐ranging clades of A. leucurus. From our phylogenetic estimates we inferred strong phylogeographic structure within Ammospermophilus and the presence of three well‐supported major clades. Initial patterns of historical divergence were coincident with dynamic alterations in the landscape of western North America, and the formation of regional deserts during the Late Miocene and Pliocene. Species distribution models and demographic parameter estimates revealed patterns of recent population expansion in response to glacial retreat. When combined with evidence from co‐distributed taxa, the historical biogeography of Ammospermophilus provides additional insight into the mechanisms that impacted diversification of arid‐adapted taxa across the arid lands of western North America. We propose species recognition of populations of the southern Baja California peninsula to best represent our current understanding of evolutionary relationships among genetic units of Ammospermophilus. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 949–967.     

Fossil involucres of Ostrya (Betulaceae) from the early Oligocene of Yunnan and their biogeographic implications

A new fossil occurrence of Ostrya (Betulaceae) is reported based on 14 involucre impressions from the lower Oligocene of Löhe Basin, Yunnan Province, Southwest China. They are characterized by their bladder-like shape with longitudinal veins and perpendicular or branched intercostal veins that form a reticulate venation. The discovery of these fossil involucres represents the earliest unequivocal fossil record of Ostrya in East Asia and the record at the lowest latitude. Its fossil history suggests that the modern distribution pattern of Ostrya might have been established since the early Oligocene, and that this genus has inhabited low altitude areas since then.     

Dioscoreaceae fossils from the late Oligocene and early Miocene of Ethiopia

Dioscorea section Lasiophyton leaflets from the late Oligocene (27.23 Ma) and Tacca leaves from the early Miocene (21.73 Ma) of north‐western Ethiopia greatly expand the known fossil record of Dioscoreaceae and represent the earliest and only known records of the Afro‐Asian trifoliate, palmately veined yams (Dioscorea) and bat flowers (Tacca). Both fossils occur in volcaniclastic and clastic sediments associated with a high water table, and the palaeofloral assemblages are indicative of tropical moist forest formations. These fossils provide insight into the evolutionary history of the family in Africa during the mid‐Cenozoic and provide well‐dated taxa that can assist in phylogenetic analyses and evolutionary divergence studies for Dioscoreales and Dioscoreaceae. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 17–28.     

First fossil Molinaranea Mello‐Leitão, 1940 (Araneae: Araneidae), from middle Miocene Dominican amber,with a phylogenetic and palaeobiogeographical analysis of the genus

The first fossil Molinaranea is described, from middle Miocene Dominican amber. This record extends the known range of the genus back 16 million years; it also extends the geographical range of the genus through time, with extant species known only from Chile, Argentina, the Falkland Islands, and Juan Fernandez Island. A parsimony‐based phylogenetic analysis was performed, which indicates that the fossil species, Molinaranea mitnickii sp. nov. , is nested with Molinaranea magellanica Walckenaer, 1847 and Molinaranea clymene Nicolet, 1849 . A modified Brooks parsimony analysis was conducted in order to examine the biogeography and origins of the fossil species in the Dominican Republic; the analysis suggests that M. mitnickii sp. nov. arrived in Hispaniola from South America as a result of a chance dispersal event. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 711–725.     

Gliridae (Rodentia,Mammalia) with a simple dental pattern: a new genus and new species from the European Early and Middle Miocene

A new genus of Gliridae, Simplomys gen. nov. is proposed. It contains glirids with a simplified dental pattern from the European Early and Middle Miocene. Simplomys gen. nov. includes several species originally described as Pseudodryomys such as Simplomys simplicidens, Simplomys robustus, Simplomys julii, and Simplomys aljaphi. In addition, a new species, Simplomys meulenorum sp. nov. , is proposed from the Spanish Miocene. The species of this genus share not only a very reduced and simplified dental morphology, but also unique dental proportions that clearly separate them from any other genera of Gliridae. Simplomys gen. nov. is recorded in most of the fossil faunas from the Early and Middle Miocene of the Iberian Peninsula, and shows the maximum diversity in this area during Mammal Neogene Zones MN 3 and MN 4. The genus has been also recorded in other European countries such as France, Germany, and Switzerland, conferring to this very characteristic taxon an important role for biochronological correlations within the European continent. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 622–652.     

Species delimitation of Chinese hop‐hornbeams based on molecular and morphological evidence

Species delimitation through which infers species boundaries is emerging as a major work in modern systematics. Hop‐hornbeam species in Ostrya (Betulaceae) are well known for their hard and heavy woods. Five species were described in China and their interspecific delimitations remain unclear. In this study, we firstly explored their distributions in all recorded field sites distributed in China. We then selected 110 samples from 22 natural populations of five species from this genus and one type specimen of O. yunnanensis, for molecular barcoding analyses. We sequenced four chloroplast (cp) DNA fragments (trnH–psbA, trnL–trnF, rps16, and trnG) and the nuclear internal transcribed spacer (ITS) region for all samples. Sequence variations of Ostrya from four cpDNA fragments identified three groups that showed no correspondence to any morphological delimitation because of the incomplete lineage sorting and/or possible interspecific introgression in the history. However, phylogenetic analyses of ITS sequence variations discerned four species, O. japonica, O. rehderiana, O. trichocarpa, and O. multinervis while O. yunnanensis nested within O. multinervis. Morphological clustering also discerned four species and showed the complete consistency with molecular evidence. Moreover, our phylogenetic analyses‐based ITS sequence variations suggested that O. trichocarpa comprised an isolated lineage different from the other Eurasian ones. Based on these results, hop‐hornbeams in China should be treated as four separate species. Our results further highlight the importance of ITS sequence variations in delimitating and discerning the closely related species in plants.     

Provannid and provannid‐like gastropods from the Late Cretaceous cold seeps of Hokkaido (Japan) and the fossil record of the Provannidae (Gastropoda: Abyssochrysoidea)

A fauna of provannid and provannid‐like shells is described from Upper Cretaceous seep carbonates in Hokkaido, Japan. We describe two new provannid species, Provanna tappuensis sp. nov. and Desbruyeresia kanajirisawensis sp. nov. , with preserved protoconchs of unquestionable provannid type with decollate apex. This material confirms the occurrence of Provannidae as early as the Middle Cenomanian. We also describe Hokkaidoconcha gen. nov. and a new family Hokkaidoconchidae fam. nov. , with two named species, H. hikidai sp. nov. and H. tanabei sp. nov . Hokkaidoconchidae are possibly related to the Provannidae, judging from a similar, but not decollate larval shell, although the juvenile teleoconch whorls differ in being of a general cerithimorph appearance and the details of the aperture are unknown. Furthermore, we review the published fossil record of Provannidae and Abyssochrysidae, and we consider that in those older than the Eocene, there is no evidence preserved that unequivocally supports a position there. The Jurassic Acanthostrophia acanthica from Italy seems to be the oldest known record of Abyssochrysidae, and the most reliable occurrence of the family, older than from the Miocene. Other fossil, pre‐Miocene species that have been classified in the Abyssochryssidae are provisionally referred to Hokkaidoconchidae. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 421–436.     

Molecular dating and diversification of the South American lizard genus Liolaemus (subgenus Eulaemus) based on nuclear and mitochondrial DNA sequences

The temperate South American lizard genus Liolaemus is the one of the most widely distributed and species‐rich genera of lizards on earth. The genus is divided into two subgenera, Liolaemus sensu stricto (the ‘Chilean group’) and Eulaemus (the ‘Argentino group’), a division that is supported by recent molecular and morphological data. Owing to a lack of reliable fossil data, previous studies have been forced to use either global molecular clocks, a standardized mutation rate adopted from previous studies, or the use of geological events as calibration points. However, simulations indicate that these types of assumptions may result in less accurate estimates of divergence times when clock‐like models or mutation rates are violated. We used a multilocus data set combined with a newly described fossil to provide the first calibrated phylogeny for the crown groups of the clade Eulaemus, and derive new fossil‐calibrated substitution rates (with error) of both nuclear and mtDNA gene regions for Eulaemus specifically. Divergence date estimates for each of the crown groups and appropriate rate estimates will provide the foundation for understanding rates of speciation, historical biogeography, and phylogeographical history for various clades in one of the most diverse lizard genera in the poorly studied Patagonian region. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 825–835.     

Analisi polliniche di depositi lacustri delle pendici del M. Vulture in Basilicata

Abstract

Pollen analysis of lake sediments on Mt. Vulture in Basilicata (Southern Italy).—A marshy sediment, about 7 meters thick, containing fossil pollen, has been discovered in a little lake of volcanic origin on a slope of Mount Vulture in Lucania at 656 m above sea level.

The sediment shows four distinct layers from the top to the bottom. The deepest layer, from 7 to 5.75 m, consisting of clay containing Betula and Ostrya, indicates a rather arid climate; the next layer, from 5.75 to 3.75 m, made up of mud rich in organic substance and containing pollen of Fagus and Abies, is witness of a damp and warmer climate; the following layer, from 3.75 to 2.25 m, with clay containing Betula, Ulmus, Ostrya and Corylus suggests a return to a more arid climate; lastly, the upper layer, from 2.25 m to the surface, with marshy mud rich in organic substance and pollen of plants with greater water requirements leads us to think of a return to a damper climate.     

A non‐aquatic otter (Mammalia,Carnivora, Mustelidae) from the Late Miocene (Vallesian,MN 10) of La Roma 2 (Alfambra,Teruel, Spain): systematics and functional anatomy

A new genus and species of otter‐like mustelid, Teruelictis riparius, is created on the basis of a partial skeleton from the Late Miocene (Vallesian age, MN 10) locality of La Roma 2 (Teruel, Spain), including several postcranial elements, the skull, and the mandible. The combination of a typically lutrine dentition, similar to that of other fossil otters such as Paralutra jaegeri, with a very slender postcranial skeleton, including a long back and gracile long bones and metacarpals, thus lacking any aquatic adaptations, was previously unknown in the fossil record. This mosaic of features strongly suggests the possibility that the aquatic lifestyle of otters could have appeared after the initial development of the distinctive dental morphology of this specialized group of mustelids. © 2013 The Linnean Society of London     

Growth,biomass distribution and CO2 exchange of northern hardwood seedlings in high and low light: relationships with successional status and shade tolerance

The physiology, morphology and growth of first-year Betula papyrifera Marsh., Betula alleghaniensis Britton, Ostrya virginiana (Mill.) K. Koch, Acer saccharum Marsh., and Quercus rubra L. seedlings, which differ widely in reported successional affinity and shade tolerance, were compared in a controlled high-resource environment. Relative to late-successional, shade-tolerant Acer and Ostrya species, early-successional, shade-intolerant Betula species had high relative growth rates (RGR) and high rates of photosynthesis, nitrogen uptake and respiration when grown in high light. Fire-adapted Quercus rubra had intermediate photosynthetic rates, but had the lowest RGR and leaf area ratio and the highest root weight ratio of any species. Interspecific variation in RGR in high light was positively correlated with allocation to leaves and rates of photosynthesis and respiration, and negatively related to seed mass and leaf mass per unit area. Despite higher respiration rates, early-successional Betula papyrifera lost a lower percentage of daily photosynthetic CO₂ gain to respiration than other species in high light. A subset comprised of the three Betulaceae family members was also grown in low light. As in high light, low-light grown Betula species had higher growth rates than tolerant Ostrya virainiana. The rapid growth habit of sarly-successional species in low light was associated with a higher proportion of biomass distributed to leaves, lower leaf mass per unit area, a lower proportion of biomass in roots, and a greater height per unit stem mass. Variation in these traits is discussed in terms of reported species ecologies in a resource availability context.     

Descriptions and phylogenetic relationships of two new genera and four new species of Oligo‐Miocene waterfowl (Aves: Anatidae) from Australia

The Tertiary anatid fossils (Aves: Anatidae) from Oligocene and Miocene deposits in Australia are described. Most fossils derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations, respectively, in the Lake Eyre and Lake Frome Basins of South Australia. The local faunas from these two formations contain the same suite of anatid species. Two new genera, the oxyurine Pinpanetta, with three new species (Pi. tedfordi, 18 specimens; Pi. vickersrichae, 15 specimens; Pi. fromensis, 20 specimens), and the tadornine Australotadorna, for a large new species known from eight specimens, are established. Three anatid bones from the Waite Formation (c. 8 Mya) at Alcoota, Northern Territory reveal the presence of a tadornine that is neither Australotadorna nor an extant Tadorna species, and an indeterminate duck about the size of Malacorhynchus. Phylogenetic analyses establish Pinpanetta as a basal member of an oxyurine (stiff‐tailed duck) radiation. Oxyurines are found to include the Recent Stictonetta and Malacorhynchus as basal members, along with the fossil taxa Mionetta, Manuherikia, and Dunstanetta, and the traditionally included Recent Oxyura, Biziura, Thalassornis, and Nomonyx. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 411–454.