A new genus of Mantispidae (Insecta: Neuroptera) from the Eocene of Germany, with a review of the fossil record and palaeobiogeography of the family

A new mantispid genus and species Symphrasites eocenicus gen. sp. nov. is described from the Middle Eocene of the Messel Pit fossil site, Germany. It has been placed, with confidence, in the subfamily Symphrasinae, the fossils of which were previously unknown. A review of the known fossil species of Mantispidae is provided, and their systematic placements are discussed. Liassochrysa stigmatica Ansorge & Schlüter, 1990 is assigned to Mantispidae, and therefore Liassochrysidae is a new synonym of Mantispidae; a second, more complete specimen of Mesomantispa sibirica Makarkin, 1997 is described from the Early Cretaceous Baissa locality, Russia; Mantispidiptera Grimaldi, 2000 is excluded from the family; Vectispa Lambkin, 1986 is referred to subfamily Mantispinae; and the assignment of Whalfera Engel, 2004 to Rhachiberothidae is validated. The origin and palaeobiogeography of the mantispids are briefly discussed: the concept of ousted relicts – which assumes that former widely distributed taxa were replaced with groups originating in the tropics – is assumed to best explain the present distribution of the family.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 701–716.     

Curtisia (Cornales) from the Eocene of Europe and its phytogeographical significance

Fossil remains of Curtisia Aiton (Cornales) are recognized for the first time from the Tertiary of Europe, based on early Eocene anatomically preserved fruits from the London Clay and Poole Formations of southern England. The modern distribution of this monotypic genus is limited to the cape of South Africa. Curtisia quadrilocularis (Reid & Chandler) comb. nov. fruits have globose tetralocular endocarps composed of isodiametric sclereids with a single seed per locule, a prominent axial vascular canal, apical placentation, and four germination valves. All of these characters, as well as size, correspond to extant Curtisia . Although many fossil taxa from the Eocene of Europe have been shown to have their closest extant relatives in Asia, this occurrence of Curtisia highlights Tertiary floristic exchange between Europe and Africa. The newly recognized fossil occurrences suggest a Laurasian origin for Curtisia , in conformity with the fossil record for several other genera of the Cornales. In addition, our rejection of the former assignment of this species to Leucopogon causes us to question whether Epacridaceae were present in the Tertiary of Europe.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 127–134.     

Analysis of family-level relationships in bees (Hymenoptera: Apiformes) using 28S and two previously unexplored nuclear genes: CAD and RNA polymerase II

We analyzed a combined data set of two protein-coding nuclear genes (CAD and RNA polymerase II) and a nuclear ribosomal gene (28S D2-D4 region) for 68 bee species and 11 wasp outgroups. Our taxon sampling included all seven extant bee families, 17 of 20 subfamilies, and diverse tribes. Wasp outgroups included the two families most closely related to bees: Crabronidae and Sphecidae. We analyzed the combined and single gene data sets using parsimony and Bayesian methods, which yielded largely congruent results. Our results provide reasonably strong support for family and subfamily-level relationships among bees. Our data set strongly supports the sister-group relationship of the Colletidae and Stenotritidae, and places Halictidae as sister to this clade combined. Our analyses place the Melittidae and the long-tongued (LT) bee clade (Apidae+Megachilidae) near the base of the tree with Colletidae (and Stenotritidae) in a fairly highly derived position. This topology ("Melittidae-LT basal") was obtained in previous morphological studies under certain methods of character coding. A more widely accepted tree topology that places Colletidae (and/or Stenotritidae) as sister to all other bees ("Colletidae basal") is not supported by our data. The "Melittidae-LT basal" hypothesis may better explain patterns in the bee fossil record as well as historical biogeography of certain bee groups. Our results provide new insights into higher-level bee phylogeny and indicate that CAD, RNA polymerase II, and 28S are useful data sets for resolving Cretaceous-age divergences in bees and other Hymenoptera.     

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.     

Systematics of the bryozoan genus Macropora (Cheilostomata)

Macropora is a distinctive genus of anascan cheilostomes. All species have large zooids with an extensive cryptocystal frontal shield perforated by numerous pores through which parietal muscle strands pass. Also characteristic of Macropora are the calcified opercula closing the d -shaped orifice, avicularia resembling autozooids but having slightly enlarged, typically crenulated opercula/mandibles, and large ovicells that are often costate. This mostly Australasian genus has a documented fossil record stretching back to the Eocene. Here we revise the generic diagnosis of Macropora and describe eight new species from New Zealand (three Recent: M. nodulosa sp. nov. , M. filifera sp. nov. and M. carlosi sp. nov. ; five fossil: M. septispinosa sp. nov. , M. similis sp. nov. , M. pittensis sp. nov. , M. leeae sp. nov. and M. bullata sp. nov. ) and one new species from Japan ( M. mawatariorum sp. nov. ), the only species known with certainty to occur in the northern hemisphere. New observations are given for some previously described species, several of which are illustrated using SEM for the first time. The northern hemisphere Upper Cretaceous species Monoporella exsculpta (Marsson), showing features allowing it to be placed tentatively in the stem-group of Macropora , is redescribed. Attempts to resolve the phylogenetic relationships of species within Macropora were hampered by the relatively few morphological characters compared with the number of species, and it proved impossible to obtain a phylogeny robust to slight changes in taxon and/or character inclusion. The favoured cladogram has poor bootstrap and Bremer support, and its stratigraphical and biogeographical congruences are low.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 115–146.     

Revision of the Neotropical Scleropactidae (Crustacea: Oniscidea)

The species of Scleropactidae from America are revised and redescribed. A phylogeny hypothesis based on an analysis of morphological characters is presented. The genera are redefined and the generic placement of several species is corrected. The Neotropical Scleropactidae include Colomboscia , Scleropactes , Circoniscus , Neosanfilippia , Sphaeroniscus , Richardsoniscus , Spherarmadillo , Colomboniscus , Amazoniscus and Protosphaeroniscus . The genus Chileoniscus is excluded from the Scleropactidae. Sphaerobathytropa is excluded from the Scleropactidae and its previous record from Argentina is revealed to be erroneous. Synuropus is revalidated and excluded from the Scleropactidae. The following new genera are introduced: Scleropactoides gen. nov. , Globopactes gen. nov. , Caecopactes gen. nov. , and Troglopactes gen. nov. Thirteen species are described as new: Caecopactes minimus sp. nov. , Colomboscia parva sp. nov. , Globopactes falconensis sp. nov. , Globopactes hispidus sp. nov. , Globopactes meridae sp. nov. , Scleropactes cotopaxii sp. nov. , Scleropactes ecuadoriensis sp. nov. , Scleropactes pululahua sp. nov. , Scleropactoides curvatus sp. nov. , Circoniscus hirsutus sp. nov. , Sphaeroniscus quintus sp. nov. , Spherarmadillo nebulosus sp. nov. , and Chileoniscus armadillidioides sp. nov. Identification keys to all species are provided.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 (Suppl. 1), 1–339.     

New evolutionary evidence of Grylloblattida from the Middle Jurassic of Inner Mongolia, north-east China (Insecta, Polyneoptera)

The small modern insect order Grylloblattida has an abundant fossil record during the Late Palaeozoic and the Mesozoicirca. The relationships between these fossil taxa and the modern grylloblattids remain unclear because most of them are based on isolated wings or have poorly preserved body features. Modern grylloblattids are wingless insects. The new grylloblattid family Plesioblattogryllidae fam. nov. is erected for the new genus and species Plesioblattogryllus magnificus gen. nov., sp. nov. , from the Middle Jurassic of north-eastern China. The well-preserved specimen provides further evidence that could support its close relationships with the modern grylloblattids: (1) several very similar head structures, e.g. developed laciniae with inner row of setae, maxillary palps segmented into five, labial palps segmented into three, large labrum, and morphology of antenna; (2) paired eoplantulae on tarsomeres 1–4; (3) long ovipositor and large eggs comparable with those of modern taxa. The new genus has strongly developed mandibles with sharp pointed apical teeth and strong marginal teeth, and strong hook-like fore claws with basal teeth, suggesting it was carnivorous. The major differences between the extinct and extant Grylloblattida, such as the lack of wings, the eyes and ocelli either degenerated or absent, and the thorax degenerated in the modern forms, are probably related to their adaptation to their life under rocks and rock-crawler habits.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 17–24.     

A review of the platanaceous woods from the Eocene paratropical rainforest of south-east England

Small diameter pyritized axes, commonly referred to as 'twigs', of fossil platanaceous wood are described from the Lower Eocene London Clay Formation of south-east England. These twigs are characterized by solitary vessels with scalariform perforation plates, opposite intervessel pits, and tall, multiseriate rays that dilate in the phloem region. The wood anatomy supports close relationship to members of extant Platanaceae and the material is placed in the organ genus Plataninium Unger erected for fossil woods with close anatomical similarity to Platanus L. This material supplements the fossil record of platanaceous type wood from the Eocene London Clay and documents the first record of Plataninium decipiens Brett in the twig flora.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 181–191.     

A new species of Orycteropodidae (Mammalia, Tubulidentata) in the Mio-Pliocene of northern Chad

A new species of fossil Tubulidentata has been found by the Mission Paléoanthropologique Franco-Tchadienne in Northern Chad. It is the first fossil Orycteropodidae (aardvark) from the Mio-Pliocene of Central Africa. The new taxon, Orycteropus abundulafus sp. nov. , is considered in the framework of the available Orycteropodidae fossil record. The Chadian specimen is characterized by the highest dental robustness index among all Tubulidentata, the presence of crests on the pterygoid, the triangular-shaped olecranon fossa and the reduction of the deltoid crest. All of these characters are linked to a less fossorial animal that had a tougher diet. This new African species is closer to the Eurasian O. gaudryi than to any other Tubulidentata. Together they form a clade distinct from that which includes O. afer . This is the first evidence of a relationship for aardvarks between Africa and Eurasia. An initial step is made towards revision of the phylogeny of the order.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 109–131.     

Microsatellite DNA and behavioural studies provide evidence of host-mediated speciation in Myzus persicae (Hemiptera: Aphididae)

The fossil record of mammals records a major interchange of northern and southern faunas in the New World, upon closure of the Panamanian isthmus approximately 3 Mya, termed the Great American Biotic Interchange (GABI). Due to their poor preservation in the fossil record, the degree of participation of birds in this interchange remains largely unknown. A phylogeny for wrens of the genus Campylorhynchus (Aves: Passeriformes) was reconstructed using DNA sequences from the mitochondrial control region and cytochrome b gene. This phylogeny, in combination with biogeographical inference and molecular clock methods, allows estimates of the importance of Late Pliocene interchange to the history of the group. Biogeographical reconstructions and divergence date estimates suggest that the genus began diversification in North America prior to closure of the Panamanian isthmus, consistent with a hypothesized North American origin for the family Troglodytidae. These reconstructions are consistent with pre-GABI dispersal of at most a single Campylorhynchus lineage into South America, with subsequent dispersal of additional lineages, probably across the fully formed isthmus. Increased sampling of avian taxa with widespread New World distributions will continue to clarify the timing and direction of continental interchange.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 687–702.     

Towards a revision of the genus Halectinosoma (Copepoda, Harpacticoida, Ectinosomatidae): new species from the North Atlantic and Arctic regions

This paper contributes to a revision of the genus Halectinosoma . Four new species are described, based on examination of ectinosomatid material from localities in western Europe, eastern Canada and the Arctic. Halectinosoma mandibularis sp. nov. is distinguishable from other species by the reduced setation of the mouthparts and enlarged mandibular gnathobase. Halectinosoma latisetifera sp. nov. bears an affinity with H. cooperatum but is easily distinguished by the shape of the setae on the female fifth leg. A species previously erroneously ascribed to H. finmarchicum (Scott) by several authors is described here as Halectinosoma kliei sp. nov. Halectinosoma gothiceps (Giesbrecht) is redescribed and the closely related Halectinosoma paragothiceps sp. nov. is described and distinguished from H. gothiceps . It is considered likely that some previous records of H. gothiceps are in error.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 453–475.     

Morphological perspective on the classification and evolution of Recent Pterioidea (Mollusca: Bivalvia)

The evolutionary relationships of the Recent Pterioidea are inferred from a phylogenetic analysis of representatives of all pterioidean genera based on original observations of anatomy and shell morphology, and an extensive survey of bivalve literature. The well-resolved cladogram supports monophyly for the superfamily, but renders all but one family (the monotypic Pulvinitidae) polyphyletic. In addition, these results reveal a considerable level of convergence and parallelisms through the Pterioidea. The branching order of pterioid genera in the morphological analysis is largely corroborated by the sequence of their appearance in the fossil record. The palaeontological evidence provides important information on dating lineage splitting events and transitional taxa. The proposed phylogeny integrates the cladistic analysis of the Recent Pterioidea with the fossil record and suggests that the crown-group pterioideans probably originated in the Triassic from the Bakevelliidae, an extinct paraphyletic stem group from which the Ostreoidea are also ultimately derived.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 253–312.     

A comprehensive phylogeny of the bumble bees (Bombus)

Bumble bees ( Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear to be similar in morphology throughout their range, suggesting that behavioural adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspecific variation in colour pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographical distribution and the evolution of behavioural and morphological adaptations, including colour pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data, we report the first nearly complete species phylogeny of bumble bees, including most of the 250 known species from the 38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1α, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology-based subgenera. Most subgenera fall into two distinct clades ( short-faced and long-faced ) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one-quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a firm foundation for reclassification and for evaluating character evolution in the bumble bees.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 161–188.     

Comparative biology of pollination systems in the African-Malagasy genus Brownleea (Brownleeinae: Orchidaceae)

The role of pollinators in the evolution of the African-Malagasy orchid genus Brownleea (Brownleeinae) was investigated. Taxa show specialization for pollination by bees ( B. parviflora , B. recurvata ), short-proboscid flies ( B. galpinii ), and long-proboscid flies ( B. macroceras , B. coerulea ). All species in the genus produce nectar, but some ( B. coerulea , B. galpinii ) appear to mimic flowers of abundant sympatric species as an additional strategy to attract pollinators. Species investigated in terms of their breeding systems ( B. coerulea , B. parviflora , B. macroceras , B. galpinii ) are reliant on pollinator visits for seed production. Self-pollination results in strong inbreeding depression during embryo formation. A phylogeny of the genus, constructed using data from the ITS1-5.8S-ITS2 region and morphology, indicates that fly pollination is likely to be basal in the genus, and that there has been a single shift to bee pollination.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 65–78.     

Curved fossil bee cells as tools for reconstructing the evolutionary history and palaeogeographical distribution of Diphaglossinae (Apoidea,Colletidae)

The new ichnospecies Celliforma curvata is described to include curved fossil bee cells from Argentina, Uruguay and the USA. The upper part of the cell (neck) of the new ichnospecies is curved, and accordingly, it can be attributed to bees of the subfamily Diphaglossinae (Colletidae). The oldest record of C. curvata, from the early Eocene of North America (52–49 Ma), provides a minimum age for the appearance of this subfamily, in accordance with an already proposed calibrated phylogeny. It is also proposed that these fossil cells could be used for future calibrations of molecular clocks. C. curvata indicates that Diphaglossinae had a widespread distribution, from southern Utah to extra‐Andean Patagonia at 42°S. In contrast, extant representatives reach only 38°S in this region. Diphaglossinae were more extended southwards in the past thanks to better environmental conditions in extra‐Andean Patagonia.     

A new trirachodontid cynodont from the lower levels of the Burgersdorp Formation (Lower Triassic) of the Beaufort Group, South Africa and the cladistic relationships of Gondwanan gomphodonts

We describe a new trirachodontid cynodont from the base of the Burgersdorp Formation (Subzone A fauna of the Cynognathus Assemblage Zone), of the South African Karoo Basin. Langbergia modisei gen. et sp. nov. is characterized by circular to ovoid in outline upper postcanines and the absence of a maxillary platform lateral to the postcanine series. Apart from the new taxon, we recognize two other species of this family in the Cynognathus Assemblage Zone: Trirachodon berryi and Cricodon metabolus , the latter also represented in the Tanzanian Manda Formation. A phylogenetic analysis of gomphodont cynodonts was conducted using a data matrix of 43 craniodental characters and 18 terminals. Trirachodontidae appears as a monophyletic assemblage, with Langbergia appearing as the sister taxon of Cricodon . The monophyly of trirachodontids is weakly supported, however, with one extra step breaking it. Traversodontid cynodont relationships were also inspected and compared with a recent phylogeny proposed for this group. Considering the resulting phylogeny and the sudden appearance and diversification of Cynognathia representatives, the origin of basal Cynognathia (i.e. Cynognathus , Diademodon , and trirachodontids) is suggested to predate their first appearance in the fossil record.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 383–413.     

The family Alestidae (Ostariophysi, Characiformes): a phylogenetic analysis of a trans-Atlantic clade

The overall most parsimonious hypothesis of relationships based on 200 characters indicates that the Alestidae is the closest relative of Chalceus , a genus previously assigned to the Neotropical Characidae. Chalceus is shifted into the Alestidae, which becomes the only trans-Atlantic family level group within the Characiformes. Various previously proposed suprageneric assemblages within the Alestidae (e.g. Petersiini) failed to delimit monophyletic groups under the intrafamilial phylogenetic analysis. The evaluation of fossil alestids within the context of the phylogeny indicates that the ancestors of Alestes , Arnoldichthys , Brycinus , Bryconaethiops and Hydrocynus evolved prior to the early Eocene (Cuisian of Upper Ypresian), 49–54.8 million years ago, with the fossil Alestoides most closely related to Alestes. The phylogenetic information further indicates a minimum age of 90–112 million years for the Alestidae. Contrary to previous hypotheses, the fossil African Sindacharax was found to be most similar to the clade including the alestid genus Bryconaethiops rather than most closely related to the South American subfamily Serrasalminae. Evaluation of the fossil Mahengecharax carrolli fails to support its hypothesized placement as the sister group to all Recent members of the Alestidae. Two separate episodes of miniaturization and one episode of gigantism occurred within the evolution of the Alestidae.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 1−144. No claim to original US government works.     

Phylogeny and biogeographical history of Trogoniformes, a pantropical bird order

With highly conserved morphology throughout the family, a tropical distribution, and no close living relatives, the trogons (Aves: Trogonidae) pose a difficult problem for systematists. Disjunct tropical distributions are often attributed to Gondwanan vicariance, but the fossil record for trogons is mostly from the Tertiary of Europe. This study examined support for the basal relationships among trogons using a combination of nuclear (RAG-1) and mitochondrial (ND2) DNA sequence data. Although some nodes could not be resolved with significant support, there is strong support for the basal position of three New World genera ( Pharomachrus , Euptilotis , and Priotelus ). This phylogenetic hypothesis differs markedly from previous studies of trogon relationships and taxonomic treatments. Biogeographically, it implies an origin and early vicariance events for the crown clade in the New World. Molecular divergence estimates place all of the basal nodes of the trogon phylogeny in the Oligocene, precluding a Gondwanan origin for modern trogons.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 725–738.     

Taxonomic revision, phylogenetic analysis, and biogeography of the bee genus Tropidopedia (Hymenoptera, Apidae, Tapinotaspidini)

The species of the bee genus Tropidopedia stat. nov. are revised. Thirteen new species are proposed: T. carinata sp. nov. , T. caracicola sp. nov. , T. danunciae sp. nov. , T. duckeana sp. nov. , T. eliasi sp. nov. , T. flavolineata sp. nov. , T. friesei sp. nov. , T. japuraensis sp. nov. , T. nigrita sp. nov. , T. nigrocarinata sp. nov. , T. ornata sp. nov. , T. peruana sp. nov. and T. venezuelana sp. nov. Taxonomic notes are provided for another four species: Tropidopedia punctifrons (Smith, 1879) comb. nov. , T. pallidipennis (Friese, 1899) comb. nov. , T. seabrai (Michener & Moure, 1957) and T. arcuatilis (Vachal, 1909) comb. nov. , including designation of a lectotype for Tetrapedia arcuatilis Vachal. An identification key, illustrations for main diagnostic characters, and distributions maps for all species are provided. A phylogenetic analysis was carried out to evaluate the monophyly of Tropidopedia and its main species groups, as well as to position it among the related genera. Biogeographical patterns are discussed.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 511–554.