INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A CATKIN WITH JUGLANDACEOUS AFFINITIES

Three specimens of one type of fossil catkin from the Middle Eocene of Tennessee are excellently preserved and have been investigated morphologically. The flowers on these catkins are subtended by elongate, three-lobed bracts, are exclusively staminate, and have three conspicuous, obovate, perianth parts that bear large peltate scales. The stamens are well preserved and contain triporate pollen grains that are equivalent to the dispersed pollen genus Momipites. Floral morphology, cuticular features, and pollen indicate close affinities with the extant genera Engelhardia, Oreomunnea, and Alfaroa of the Juglandaceae; but because the fossil catkins are distinct and are a dispersed plant organ, they are placed in a new form genus: Eokachyra. These fossil flowers represent a rare opportunity to correlate the micro- and macrofossil record and to compare the relative rates of evolution of these features. The fossil catkins also demonstrate that much structural information may be gained from the study of fossil angiosperm flowers. The similarities between the staminate flowers of the fossil catkins and the staminate flowers of Engelhardia, Oreomunnea, and Alfaroa confirm the idea that this complex has had a long evolutionary history and suggest that the pollination system of certain extant genera was well developed during Middle Eocene times.     

Numerical taxonomy of Palaeocarya (Juglandaceae) from the Mangbang Formation of West Yunnan, China

Specimens of Palaeocarya (Juglandaceae) were collected from the upper Pliocene of western Yunnan Province, which is a new geographical and age range record of engelhardioid winged fruits in China. A numerical taxonomic investigation based on 14 morphological characters of fossil and extant specimens of engelhardioid winged fruits in China was conducted using multivariate methods. Phenetic clustering shows that there are four morphotypical groups which we assign to four species. Two groups are assigned to the previously described fossil species, Palaeocarya guangxiensis Li Hao-Min et Chen Yun-Fa, 2003 and Palaeocarya koreanica (Oishi) Manchester, 1987. Two new species are recognized, Palaeocarya yunnanensis and Palaeocarya longialata. These Neogene Engelhardieae fruits provide fossil data for understanding the origin and evolution of the Engelhardia species that now inhabit South China.     

New fossil Ochteridae (Hemiptera: Heteroptera: Ochteroidea) from the Upper Mesozoic of north‐eastern China,with phylogeny of the family

Specimens of the velvety shore bugs (Hemiptera: Ochtheridae) occur in the Early Cretaceous Yixian Formation in north‐eastern China. We describe two new genera and three new species –Pristinochterus ovatus Yao, Zhang & Ren sp. nov., Floricaudus multilocellus Yao, Ren & Shih gen.n. et sp.n. and Angulochterus quatrimaculatus Yao, Zhang & Ren gen.n. et sp.n. – from this deposit. A cladistic analysis based on a combination of fossil and extant taxa, and their morphological characters, clarifies the phylogenetic status of the new fossils and allows the reconstruction of relationships within the family Ochteridae. Two main clades within Ochteridae are recognized from the cladistic analysis: Pristinochterus Yao Cai & Ren and Floricaudus Yao, Ren & Shih gen.n. form a monophyletic lineage; and Angulochterus Yao, Zhang & Ren gen.n. emerges as a sister group to all extant velvety shore bugs. A key to all fossil and extant genera of Ochteridae is provided.     

Phylogenetic Distribution and Identification of Fin-winged Fruits

Fin-winged fruits have two or more wings aligned with the longitudinal axis like the feathers of an arrow, as exemplified by Combretum, Halesia, and Ptelea. Such fruits vary in dispersal mode from those in which the fruit itself is the ultimate disseminule, to schizocarps dispersing two or more mericarps, to capsules releasing multiple seeds. At least 45 families and more than 140 genera are known to possess fin-winged fruits. We present an inventory of these taxa and describe their morphological characters as an aid for the identification and phylogenetic assessment of fossil and extant genera. Such fruits are most prevalent among Eudicots, but occur occasionally in Magnoliids (Hernandiaceae: Illigera) and Monocots (Burmannia, Dioscorea, Herreria). Although convergent in general form, fin-winged fruits of different genera can be distinguished by details of the wing number, texture, shape and venation, along with characters of persistent floral parts and dehiscence mode. Families having genera with fin-winged fruits and epigynous perianth include Aizoaceae, Apiaceae, Araliaceae, Asteraceae, Begoniaceae, Burmanniaceae, Combretaceae, Cucurbitaceae, Dioscoreaceae, Haloragaceae, Lecythidiaceae, Lophopyxidaceae, Loranthaceae, and Styracaceae. Families with genera having fin-winged fruits and hypogynous perianth include Achariaceae, Brassicaceae, Burseraceae, Celastraceae, Cunoniaceae, Cyrillaceae, Fabaceae, Malvaceae, Melianthaceae, Nyctaginaceae, Pedaliaceae, Polygalaceae, Phyllanthaceae, Polygonaceae, Rhamnaceae, Salicaceae sl, Sapindaceae, Simaroubaceae, Trigoniaceae, and Zygophyllaceae. This survey has facilitated the identification of fossil winged fruits such as Combretaceae and Araliaceae in the late Cretaceous of western North America and provides additional evidence toward the identification of various Cenozoic fossils including Brassicaceae, Fabaceae, Polygonaceae, Rutaceae, and Sapindaceae.     

A new species of winged fruit from the Miocene of Ecuador: Tipuana ecuatoriana (Leguminosae)

Tipuana ecuatoriana sp. nov. (Leguminosae; Dalbergieae), is described from winged fruits collected in Miocene sediments of the Loja and Nabon Basins of Ecuador. Extensive comparisons with families that include genera with similar samaroid fruits or disperal units led to recognition of legume affinities for the fossil. Based upon the texture of the fruit, the position of the style remnant, and the venation of the fruit wing, the fossil conforms to the extant monotypic genus, Tipuana, native to subtropical Bolivia and northwest Argentina, but widely cultivated in the subtropics. Tipuana, and thus the Dalbergieae as presently recognized, were present in South America prior to the establishment of permanent land connection between North and South America.     

CHATTAWAYA (STERCULIACEAE): A NEW GENUS OF WOOD FROM THE EOCENE OF OREGON AND ITS IMPLICATIONS FOR XYLEM EVOLUTION OF THE EXTANT GENUS PTEROSPERMUM

A new sterculiaceous wood, Chattawaya paliformis gen. et sp. n., was collected from a Middle Eocene locality in the type area of the Clamo Formation of north central Oregon. The wood differs from all known extant woods in having very large and irregularly shaped tile cells. However, other features of the xylem indicate a close natural relationship between Chattawaya and the extant paleotropical genus, Pterospermum. It is suggested that the xylem organization of the Eocene fossil represents a more primitive condition than that observed in Pterospermum. Comparative xylotomy indicates three evolutionary trends within the complex to which Chattawaya and Pterospermum belong: development of storied structure, increased uniformity in shape and size of tile cells and loss of multiseriate rays. Chattawaya is part of a growing body of paleobotanical evidence indicating that the Sterculiaceae was well differentiated by the early Tertiary, but that the lineages leading to extant genera have undergone an appreciable amount of evolutionary change since the Eocene.     

EVOLUTION OF COTYLEDONARY AND NODAL VASCULATURE IN THE JUGLANDACEAE

Cotyledonary nodal patterns of the Juglandaceae range from 1-gap, 2-trace to multi-gap, multi-trace. The development of increased nodal complexity is associated with at least two independent evolutionary shifts from epigeal to hypogeal germination. The taxa with epigeal germination such as Engelhardia sect. Engelhardia, Engelhardia sect. Psilocarpeae, Platycarya, and Pterocarya all have 1-gap, 2-trace nodes. The change to hypogeal germination in Engelhardia sect. Oreomunnea and Alfaroa is correlated with the development of 1-gap, 3-trace cotyledonary nodes. The second line has led to large, heavy-fruited members with hypogeal germination and complex cotyledonary nodes ranging from 2–6 gaps. The diversity of nodal patterns is the result of variation on a common theme; five basic vascular strands in the cotyledon, undergoing variations in dichotomy, fusion, and separation, are associated with one to many gaps. Presumably the complex development of the cotyledonary node is a response to increased functional demands of hypogeous cotyledons.     

Fossil flowers and fruits of the Actinidiaceae from the Campanian (Late Cretaceous) of Georgia

A new genus and species of Actinidiaceae (Parasaurauia allonensis gen. et sp. nov.) are established for fossil flowers and fruits from the early Campanian (Late Cretaceous) Buffalo Creek Member of the Gaillard Formation in central Georgia, USA. The fossil flowers, which are exquisitely preserved as charcoal, have five imbricate, quincuncially arranged sepals and petals. The androecium consists of ten stamens with anthers that are deeply sagittate proximally. The gynoecium is tricarpellate, syncarpous, and has three free styles that emerge from an apical depression in the ovary. The fruit is trilocular and contains numerous ovules on intruded axile placentae. The structure of mature fruits is unknown. Comparisons with extant taxa clearly demonstrate that the affinities of Parasaurauia allonensis are with the Ericales, and particularly with the Actinidiaceae, which have been placed among the Ericales in recent cladistic analyses. Because Parasaurauia allonensis is not identical to any one genus of Actinidiaceae, or other member of the Ericales, phylogenetic relationships of the fossil were evaluated through a cladistic analysis using morphological and anatomical characters. Results of this analysis place Parasaurauia allonensis within the Actinidiaceae as sister to the extant genera Saurauia and Actinidia. Parasaurauia allonensis differs from extant Saurauia only in having ten rather than numerous stamens.     

TRIPLOCHITIOXYLON (STERCULIACEAE): A NEW GENUS OF WOOD FROM THE EOCENE OF OREGON AND ITS BEARING ON XYLEM EVOLUTION IN THE EXTANT GENUS TRIPLOCHITON

A new sterculiaceous wood, Triplochitioxylon oregonensis gen. et sp. n., was collected from a Middle Eocene locality in the Clarno Formation of Oregon. Anatomical data indicate a close natural relationship between T. oregonensis and the living species of Triplochiton, a genus endemic to tropical Africa. The fossil is believed to represent a population of the group or complex from which Triplochiton evolved. The basic differences in the xylem organizations of the two genera are explained by a significant reduction of fusiform initial length and by a complete suppression of post-cambial parenchyma strand elongation in the extant genus. Paleobotanical and biogeographical evidence suggest that the xylem evolution has been strongly influenced by the increasing aridity of the African continent.     

Morphological conservatism in the foreleg structure of cicada hatchlings,Burmacicada protera n. gen., n. sp. in Burmese amber,Dominicicada youngi n. gen., n. sp. in Dominican amber and the extant Magicicada septendecim (L.) (Hemiptera: Cicadidae)

Two cicada hatchlings (Hemiptera: Cicadidae) in Burmese and Dominican amber are described as Burmacicada protera n. gen., n. sp. and Dominicicada youngi n. gen., n. sp., respectively. Although very similar in appearance, the two species can be separated by body contour, the nature of the process on the terminal antennomere and the shape and size of protrusions, teeth and spines on the forelegs. A comparison of the forelegs of the fossil hatchlings with those of an extant hatchling of the periodical cicada, Magicicada septendecim (L.), reveals a remarkable degree of morphological conservatism over 100 million years. A brief review of fossil cicadas is presented.     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

Reproductive structures of Rhamnaceae from the Cerro del Pueblo (Late Cretaceous, Coahuila) and Coatzingo (Oligocene, Puebla) Formations, Mexico

Recently discovered fossil flowers from the Cretaceous Cerro del Pueblo and flowers and fruits from the Oligocene Coatzingo Formations are assigned to the Rhamnaceae. The Cretaceous flower, Coahuilanthus belindae Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov., is actinomorphic with fused perianth parts forming a slightly campanulate to cupulate floral cup, with sepals slightly keeled and spatulate clawed petals. The Oligocene fossils include Nahinda axamilpensis Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov. (characterized by its campanulate bisexual flower with stamens opposite, adnate to and enfolded by petals; and with the ovary ripening into a drupe), and a winged fruit assigned to Ventilago engoto Calvillo-Canadell and Cevallos-Ferriz, sp. nov. The flowers and drupe features indicate closer affinity to Zizipheae and/or Rhamneae, while the single samaroid fruit suggests the presence of Ventilagineae. However, the unique character combination in the fossil flowers precludes placing them in extant genera. Nevertheless, the history of the family is long and can be traced back to the Campanian. A detailed phylogenetic revision of the group that uses morphological characters from both extant and fossil plants is needed to better understand the significance of these records as well as other important fossils of the family.     

New genera and new species of leaf beetles (Coleoptera: Polyphaga: Chrysomelidae) from Lowermost Eocene French amber

Three new monotypical genera of leaf beetles are described: Aoriopsis eocenicus n. gen, n. sp. and Acolaspoides longipes n. gen., n. sp. from the subfamily Eumolpinae, and also Crepidocnema yantarica n. gen., n. sp. from the subfamily Alticinae. Short reviews of known fossil records of the mentioned subfamilies are given and position of the new genera discussed.     

FRUITS AND SEEDS OF TRIBE GORDONIEAE (THEACEAE) FROM THE EOCENE OF NORTH AMERICA

A new genus and species Gordoniopsis polysperma and two new species of Gordonia (Gordonieae, Camellioideae, Theaceae) are described based on fossil fruit and seed remains. These specimens are part of a large flora consisting of various plant organs from the middle Eocene Claiborne Formation in western Kentucky and Tennessee. Gordoniopsis is a five-valved loculicidally dehiscent capsule similar to capsules of Gordonia but differing in having unwinged seeds and a greater number of seeds per locule. The two Gordonia species are among the earliest unequivocal records of the genus and two of only four fossil Gordonia species known with in situ seeds. Two extinct genera, Gordoniopsis and Andrewsiocarpon, and the extant genus Gordonia in the tribe Gordonieae are known from the middle Eocene Claiborne flora, suggesting an early radiation within the tribe. Based on a survey of Recent fruits and seeds we concur with Keng's proposal to merge Laplacea with Gordonia.     

A Total Evidence Phylogeny of the Arctoidea (Carnivora: Mammalia): Relationships Among Basal Taxa

A total evidence phylogenetic analysis was performed for 14 extant and 18 fossil caniform genera using a data matrix of 5.6 kbp of concatenated sequence data from six independent loci and 80 morphological characters from the cranium and dentition. Maximum parsimony analysis recovered a single most parsimonious cladogram (MPC). The topology of the extant taxa in the MPC agreed with previous molecular phylogenies. Phylogenetic positions for fossil taxa indicate that several taxa previously described as early members of extant families (e.g., Bathygale and Plesictis) are likely stem taxa at the base of the Arctoidea. Taxa in the “Paleomustelidae” were found to be paraphyletic, but a monophyletic Oligobuninae was recovered within this set of taxa. This clade was closely related to the extant genera Gulo and Martes, therefore, nested within the extant radiation of the family Mustelidae. This analysis provides a resolution to several discrepancies between phylogenies considering either fossil taxa or extant taxa separately, and provides a framework for incorporating fossil and extant taxa into comprehensive combined evidence analyses.     

NEW TAXODIACEOUS SEED CONES FROM THE UPPER CRETACEOUS OF NEW JERSEY

Nephrostrobus cliffwoodensis gen. et sp. nov., Nephrostrobus bifurcatus sp. nov., and Rhombostrobus cliffwoodensis gen. et sp. nov. are described based on anatomical studies of seed cone fragments from the Upper Cretaceous Magothy Formation of New Jersey. These species belong to the Taxodiaceae. As in Sequoia, Sequoiadendron. and Metasequoia, the vascular strands supplying the scale and bract in Nephrostrobus are about equal. These vascular strands are arranged in a reniform pattern resembling that found in Metasequoia, but the orientation differs by 180 degress. Nephrostrobus cliffwoodensis and Nephrostrobus bifurcatus differ from each other in the branching of the complex trace and associated resin canals. Rhombostrobus cliffwoodensis resembles Cunninghamia in the arrangement of vascular strands and associated resin canals in its bract-scale complexes. However, the relative amount of bract and ovuliferous scale making up the complex is more like that in Sequoia. This combination of cone features does not occur in any of the extant genera. Nephrostrobus and Rhombostrobus are not considered to be ancestral to any of the extant taxodiaceous genera, but are extinct members of an ancestral complex from which the extant genera were derived.     

Small but not ephemeral: newly discovered species of Aphelinidae and Trichogrammatidae (Insecta: Hymenoptera: Chalcidoidea) from Eocene amber

New species of fossil Aphelinidae and Trichogrammatidae are described from middle Eocene (Lutetian) Baltic amber (41.3–47.8 Ma). A new subfamily, two new genera and three new species of Aphelinidae are described, with comments on their placement: Phtuaria fimbriae gen.n. , sp.n. in Phtuariinae subf.n. , Glaesaphytis interregni gen.n. , sp.n. and Centrodora brevispinae sp.n. These represent the first described true fossil Aphelinidae. Four new species of Trichogrammatidae are described: Mirufens illusionis sp.n. , Palaeogramma eos gen.n. , sp.n. , Pterandrophysalis plasmans sp.n. and Szelenyia terebrae sp.n. , thus expanding our knowledge of fossil Trichogrammatidae beyond the single previously described species. The presence of recognizable extant genera of Aphelinidae and Trichogrammatidae in the Eocene suggests that the morphology of these genera has been relatively invariant despite highly variable conditions during and since the Eocene. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:E9AD60B6‐3D56‐4E74‐AA54‐F7B91F4FDC79 .     

Plio-Pleistocene cercopithecids from Kanam East, western Kenya

This paper describes cercopithecid craniodental and postcranial fossils recovered by L. S. B. Leakey at Kanam East, Kenya during the early 1930s. These fossil monkeys have been generally assumed to have been derived from early Pliocene horizons, but their exact geographical and stratigraphical provenience is unknown. Although the question of the evolutionary significance of these specimens must await the recovery of more securely dated material from Kanam East, some general conclusions can be drawn concerning their taxonomic affinities and paleobiology. Based on comparative studies of the craniodental material, at least three extant genera are represented—Colobus,Lophocebus, andCercopithecus. The postcranial fossils include a number of hindlimb specimens, as well as the manubrium of a sternum and a caudal vertebra. Identification of the postcranial remains to particular genera is not possible, but they are similar in morphology to modern arboreal and semiterrestrial cercopithecid monkeys of small to medium size. It is evident that Kanam East had a diverse cercopithecid community, similar to those found today in forested and woodland habitats, and this may be of some significance in reconstructing the paleoecology of the site. Because the fossil record of most extant cercopithecid genera is rather sparse at Plio-Pleistocene sites in Africa, Kanam East represents one of only a few sites that has yielded material that can be assigned toColobus,Lophocebus, orCercopithecus. The fossil monkeys from the site, therefore, provide additional evidence to help reconstruct the paleobiology, as well as the patterns of species diversity and community structure that characterized the cercopithecid radiation during the Plio-Pleistocene.     

Past and present diversity and distribution in the parasitic wasp family Megalyridae (Hymenoptera)

The phylogenetic relationships of extant and extinct Megalyridae are analysed at the genus level. The dataset comprises seven outgroup taxa, all eight extant genera and a number of extinct taxa that have been associated with Megalyridae, including two genera from Maimetshidae, whose affinity with Megalyridae is uncertain. Analytical results are unstable because some of the fossil taxa have many missing entries. The most stable results are produced when the maimetshid taxa and Cretodinapsis are excluded. When included, these taxa fall outside crown‐group Megalyridae, the maimetshid taxa being the sister of Orthogonalys (Trigonalidae). Based on the results of our analyses, we synonymize the fossil genera Rubes Perrichot n.syn . and Ukrainosa Perrichot & Perkovsky n.syn . with Prodinapsis, creating the new combinations Prodinapsis bruesi n.comb . and Prodinapsis prolata n.comb . When comparing past and present distributions of Megalyridae with the results of the phylogenetic analyses, it is evident that the genera radiated in the Mesozoic, and that the family as a whole was much more widespread then. The present‐day distribution is essentially relictual, with range contraction since the early Tertiary probably being the result of climate deterioration, which caused the disappearance of tropical forests throughout the Palaearctic.     

Expanding the molecular and morphological diversity of Apusomonadida,a deep-branching group of gliding bacterivorous protists

Apusomonads are cosmopolitan bacterivorous biflagellate protists usually gliding on freshwater and marine sediment or wet soils. These nanoflagellates form a sister lineage to opisthokonts and may have retained ancestral features helpful to understanding the early evolution of this large supergroup. Although molecular environmental analyses indicate that apusomonads are genetically diverse, few species have been described. Here, we morphologically characterize 11 new apusomonad strains. Based on molecular phylogenetic analyses of the rRNA gene operon, we describe four new strains of the known species Multimonas media, Podomonas capensis, Apusomonas proboscidea, and Apusomonas australiensis, and rename Thecamonas oxoniensis as Mylnikovia oxoniensis n. gen., n. comb. Additionally, we describe four new genera and six new species: Catacumbia lutetiensis n. gen. n. sp., Cavaliersmithia chaoae n. gen. n. sp., Singekia montserratensis n. gen. n. sp., Singekia franciliensis n. gen. n. sp., Karpovia croatica n. gen. n. sp., and Chelonemonas dolani n. sp. Our comparative analysis suggests that apusomonad ancestor was a fusiform biflagellate with a dorsal pellicle, a plastic ventral surface, and a sleeve covering the anterior flagellum, that thrived in marine, possibly oxygen-poor, environments. It likely had a complex cell cycle with dormant and multiple fission stages, and sex. Our results extend known apusomonad diversity, allow updating their taxonomy, and provide elements to understand early eukaryotic evolution.