Division of Entomology (Paleoentomology) Natural History Museum 1501 Crestline Drive - Suite 140 University of Kansas Lawrence, Kansas 66049-2811 United States
Professor, Department of Ecology & Evolutionary Biology Senior Curator & Curator-in-Charge, Division of Entomology, Natural History Museum Courtesy Curator, Division of Invertebrate Paleontology, Natural History Museum Courtesy Professor, Department of Geology Research Associate, American Museum of Natural History Fellow, Linnean Society of London
Abstract: An enigmatic new genus and species of rove beetle (Coleoptera: Staphylinidae) is described as Dolichoxenus newtoni, new genus and species, in Early Miocene (Burdigalian) amber from the Dominican Republic. The species is the first oxyteline discovered as an amber inclusion. The genus is apparently a member of the tribe Oxytelini (Staphylinidae: Oxytelinae) and may be allied to the African genus Jerozenia Herman, 2003. The genus exhibits several remarkable apomorphies for the subfamily, and in general possesses features otherwise known in the subfamily for inquilines of ants, suggesting a possible paleobiological association for the genus.
Abstract: The purpose of this application is to emend the family-group name NOMIIDAE Gozis, 1875 (Insecta, Coleoptera), a senior homonym of NOMIINAE Robertson, 1904 (Insecta, Hymenoptera) and long-considered a synonym of PSYDRINA LeConte, 1853, thereby removing the homonymy between the two names. It is proposed that the stem of the generic name Nomius Laporte, 1835, on which the beetle family-group name is based, be emended to give NOMIUSIDAE, while leaving the bee name (based on Nomia Latreille, 1804) unaltered.
Abstract: Caupolicana (Zikanapis) wileyi sp. n. from Guatemala is described. New locality records are noted for other species, and the hitherto unknown female of C. (Z.) rozenorum Michener, Engel, and Ayala from Guatemala is described. A key for the identification of Central American Caupolicana is provided.
Abstract: A giant termite is described and figured from the Late Miocene of the Styrian Basin in southeastern Austria. Gyatermes styriensis gen. n. et sp. n. is represented by a relatively complete forewing, with basal scale. The fossil approximates in size the largest of all termites today and is the largest fossil termite on record. The presence of this species in the Late Miocene fauna of Europe indicates that climatic conditions were appropriate for the persistence of species and colonies requiring relatively stable, warm conditions. The genus is primitive in overall features but shares some similarity with the dampwood termites.
Abstract: A new species of anthidiine bee (Megachilidae: Megachilinae: Anthidiini), Anthidiellum (Ranthidiellum) ignotum sp. n., is described and figured from northeastern Thailand, the furthest North this subgenus has yet been recorded. The species is distinguished from its congeners, being most similar to A. (R.) apicepilosum (Dover) in the coloration of the legs, but is otherwise quite unique for the group. A provisional key to the species of Ranthidiellum is provided.
Abstract: Aim---This study aimed to evaluate the probability of suitable habitats in the United States for two adventive orchid bee species (Eulaema polychroma (Mocsáry) and Euglossa viridissima Friese), one of which has become established in southern Florida despite the absence of its associated orchid hosts.
Location---North and Central America, northern South America, and the Caribbean.
Methods---Using positive occurrence data within the native range of both orchid bee species, Maxent species distribution modelling was employed to evaluate the probability of suitable habitats in the United States. The power of predictability for the model was tested using partitions of the data.
Results---Our results show the absence of suitable habitat in southern Arizona for E. polychroma to maintain populations there, as well as establishing the northernmost limit for the species at around 29° N in northwestern Mexico. Suitable habitat was found for E. viridissima in various locations throughout southern Florida. This species is predicted to spread to occupy roughly the southern half of the Florida Peninsula.
Main conclusions---The findings indicate that species distribution modelling is useful for evaluating records of species occurrence outside of their native range. Our results indicate that the isolated record of a male of E. polychroma from southern Arizona should not be considered representative of an established population in the absence of further males and females from the same region. Conversely, E. viridissima has successfully become established in southeastern Florida after a seemingly accidental introduction first noticed in the summer of 2003. We discuss the naturalization of E. viridissima in Florida, the probability of suitable habitat across the Caribbean (where orchid bees are otherwise natively absent today), and the absence of perfume orchids (Orchidaceae). Lastly, we discuss the implications of these results for understanding the biology and biogeography of Euglossini.
Abstract: The extinct, parasitoid wasp family Stigmaphronidae (Proctotrupomorpha: Ceraphronoidea) is reviewed and a cladistic analysis of relationships undertaken. Stigmaphronids are presently known principally in Cretaceous amber from Siberia, Alaska, Canada, New Jersey, Myanmar, and Lebanon, but also from a few compressions from the Early Cretaceous of Siberia and Mongolia. As a result of the study the following new taxa are proposed, more than doubling the size of the family: Elasmophron kurthi nov. gen. et spec. (New Jersey amber), Libanophron astarte nov. gen. et spec. (Lebanese amber), Burmaphron nov. gen., Burmaphron tridentatum nov. spec. (Burmese amber), B. prolatum nov. spec. (Burmese amber), Tagsmiphron nov. gen., Tagsmiphron muesebecki nov. spec. (New Jersey amber), T. gigas nov. spec. (New Jersey amber), T. ascalaphus nov. spec. (New Jersey amber), and T. canadense nov. spec. (Canadian amber). The genus Elasmomorpha KOZLOV is proposed as a junior synonym of Allocotidus MUESEBECK (nov. syn.) resulting in Allocotidus melpomene (KOZLOV) nov. comb. Relationships are well supported, so the lack of any stratigraphic-clade rank correlation strongly suggests poor stratigraphic sampling of what was probably a very diverse lineage.
Abstract: The fossil history of leaf beetles (Chrysomelidae) is relatively poorly documented despite an abundance of available material. Of particular interest is the origin and radiation of the diverse tortoise beetles, a derived group within Cassidinae s.l. (=Cassidinae + Hispinae) defined by the exophagous life history and specialized morphology of the immature stages. Cassidinae is also a group with relatively few fossil records that can be assigned with any degree of certainty. Here we report two of the oldest definitive tortoise beetle fossils, Eosacantha delocranioides gen.n. et sp.n. and Denaeaspis chelonopsis gen.n. et sp.n., from the Eocene Green River Formation (ca. 47 million years old) in northwestern Colorado, U.S.A. Owing to the fine level of preservation, many important features can be observed and are coded into the recent cladistic analysis for the subfamily. Phylogenetic analysis highlights that both genera have affinities with modern lineages, one restricted to the Old World and the other restricted to the Neotropics. Although Cassidinae as a whole extend into the Cretaceous, the available information suggests that the tortoise beetles perhaps originated and diversified during the Early Tertiary. As such, the morphological and biological transitions from the leaf-mining hispiforms to the distinctive tortoise-like cassidiforms, with their elaborate defensive larval shields and other unique behaviours, probably took place during the latest Paleocene or earliest Eocene. These Green River fossils are the oldest yet to document the specialized morphology associated with the transition in cassidine feeding and immature biology.
Abstract: Males of solitary bees usually spend the night in clusters on small branches of plants, cavities and flowers. The individuals usually return to the same location each evening during their life, exhibiting site fidelity to a particular plant. The present paper reports the sleeping roosts of the males of some oil-collecting bees of the genera Centris, Paratetrapedia, Lanthanomelissa, Monoeca and Tetrapedia.
Abstract: A second species of dasyleptid bristletail (Archaeognatha: Dasyleptidae) is described and figured from the Lower Permian Wellington Formation of Kansas. Dasyleptus artinskianus Engel, new species, is distinguished from other dasyleptids and a key to the species of the genus is provided.
Abstract: Like ants, termites are entirely eusocial and have profound ecological significance in the tropics. Following upon recent studies reporting more than a quarter of all known fossil termites, we present the first phylogeny of termite lineages using exemplar Cretaceous, Tertiary, and Recent taxa. Relationships among Recent families were largely unaffected by the addition of extinct taxa, but the analysis revealed extensive grades of stem-group taxa and the divergence of some modern families in the Cretaceous. Rhinotermitidae, Serritermitidae, and the âhigherâ termites (family Termitidae), which comprise 84% of the world termite species, diverged and radiated entirely in the Tertiary, corresponding with significant increase of termite individuals in the fossil record. Radiation of the higher termites may have affected the formation of terrestrial carbon reserves like oil and coal. The higher classification of Isoptera is slightly revised based on the phylogenetic results. The following new taxa are proposed: Cratomastotermitidae, new family; Euisoptera, new clade; Archotermopsidae, new family; and Neoisoptera, new clade. In addition, the families Stolotermitidae, Stylotermitidae, and Archeorhinotermitidae are newly recognized or resurrected, and the families Termopsidae and Hodotermitidae are significantly restricted in composition.
Abstract: Bees of the genus Chiasmognathus (Ammobatini) occurring in the Arabian Peninsula are revised. Three species, all new, are recognized as: Chiasmognathus nearchus n. sp. (from the UAE and Oman), C. aurantiacus n. sp. (from the UAE), and C. himyarensis n. sp. (from Yemen). Species are cleptoparasitic on the halictid bee genus Nomioides and likely also on Ceylalictus (Halictinae: Nomioidini). The host of C. nearchus n. sp. is Nomioides rotundiceps Handlirsch, 1888.
Abstract: Aside from pollen and nectar, bees of the subfamily Megachilinae are closely associated with plants as a source of materials for nest construction. Megachilines use resins, masticated leaves, trichomes and other plant materials sometimes along with mud to construct nests in cavities or in soil. Among these, the leafcutter bees (Megachile s.l.) are the most famous for their behaviour to line their brood cells with discs cut from various plants. We report on fossil records of one body fossil of a new non-leafcutting megachiline and of 12 leafcuttings from three European sitesâEckfeld and Messel, both in Germany (Eocene), and Menat, France (Paleocene). The excisions include the currently earliest record of probable Megachile activity and suggest the presence of such bees in the Paleocene European fauna. Comparison with extant leafcuttings permits the interpretation of a minimal number of species that produced these excisions. The wide range of size for the leafcuttings indirectly might suggest at least two species of Megachile for the fauna of Messel in addition to the other megachiline bee described here. The presence of several cuttings on most leaves from Eckfeld implies that the preferential foraging behaviour of extant Megachile arose early in megachiline evolution. These results demonstrate that combined investigation of body and trace fossils complement each other in understanding past biodiversity, the latter permitting the detection of taxa not otherwise directly sampled and inferences on behavioural evolution.
Abstract: The first fossil honey bee (Apini: Apis Linnaeus) from the New World is described and figured, expanding the former native range of the tribe Apini into the Western Hemisphere. Apis nearctica sp. nov., is represented by a single female worker preserved in paper shale from the middle Miocene of Stewart Valley, Nevada. The species belongs to the armbrusteri species group (= Cascapis Engel) and is most similar to the extinct species A. armbrusteri Zeuner from the Miocene of southwestern Germany. The species is described and its affinities discussed, as well as its implications for our understanding of honey bee and corbiculate bee biogeography and evolution.
Abstract: A new genus and species of the extinct Paleozoic insect order Diaphanopterodea (Palaeodictyopterida) is described and figured from the Early Permian (Artinskian) Wellington Formation of Noble County, Oklahoma. Anomalohymen dochmus, new genus and species, is most similar to Biarmohymen bardense Zalessky from the Early Permian (Kungurian) of Tshekarda in Russia. The two genera share the proximal coalescence of M and CuA along with a generally reduced venation. Rather than erect another monogeneric family, we here include Anomalohymen in the Biarmohymenidae, emending the diagnosis of the family, providing a key to the constituent genera, and extending the biogeographic and temporal range of the lineage.
Abstract: The first tiphiid wasp (Aculeata: Euaculeata: Vespoidea: Tiphiiformes) in Cretaceous amber is described and figured. Thanatotiphia nyx, new genus and species, is represented by a male entombed in mid-Cretaceous (latest Albian) amber from Myanmar. Thanatotiphia possesses remarkable apomorphies in wing venation, lacks key traits of modern subfamilies, and is thus classified in a new subfamily, Thanatotiphiinae. The fossil is further shown to be nested well within the family, indicating that major lineages of Tiphiidae diverged by the mid-Cretaceous. The new taxon is compared with modern tiphiid subfamilies and the sparse fossil history of the family briefly overviewed.
Abstract: The presence of an adventitious distal abscissa in the forewing venation of honey bees is recorded. The vein arises from the posterior quarter of the second [sic: third] submarginal cell, on the apical surface of the 2rs-m crossvein. The homology of this vein is discussed and its presence broadly compared across Hymenoptera so as to establish its correct identity. Based on the overall homology of hymenopteran wing venation it is concluded that this is a reacquisition of the vein "Rs2" and we accordingly tentatively term it "aRs2". The aRs2 was found in both Apis cerana Fabricius and A. mellifera Linnaeus at moderate frequencies, although it predominantly occurs in northeast Asian populations of the former. Species of the giant honey bees (Megapis) and dwarf honey bees (Micrapis) were found to lack the vein entirely, while the vein was found at low levels in other Asian lineages (Apis s.str.). The predilection to possess the vein appears to be a trait supporting the cavity-nesting honey bee clade (Apis s.str.).
Notes: Note an error in the abstract which refers to the "second" submarginal cell but should in fact refer to the "third" submarginal cell.
Abstract: The fossil crown wasp Electrostephanus petiolatus Brues comb. rev. (Stephanidae: Electrostephaninae) is redescribed from a single male preserved in middle Eocene Baltic Amber. The holotype was lost or destroyed around the time of World War II and subsequent interpretations of its identity have been based solely on the brief descriptive comments provided by Brues in his original account. The new specimen matches the original description and illustration provided by Brues in every detail and we hereby consider them to be conspecific, selecting the specimen as a neotype for the purpose of stabilizing the nomenclature for this fossil species. This neotype exhibits a free first metasomal tergum and sternum, contrary to the assertion of previous workers who indicated these to be fused. Accordingly, this species does indeed belong to the genus Electrostephanus Brues rather than to Denaeostephanus Engel & Grimaldi (Stephaninae). Electrostephanus petiolatus is transferred to a new subgenus, Electrostephanodes n. subgen., based on its elongate pseudo-petiole and slender gaster, but may eventually warrant generic status as the phylogenetic placement of these fossil lineages continues to be clarified. A revised key to the Baltic amber crown wasps is provided.
Abstract: New lacewings (Neuroptera) and snakeflies (Raphidioptera) are reported in Cretaceous ambers from Myanmar (Albian-Cenomanian), New Jersey (Turonian), and Canada (Campanian). Those newly reported and described in Burmese amber comprise the most species of any Cretaceous amber deposit, including a remarkable diversity of beaded lacewings (Berothidae) and the first Cretaceous amber alderfly (Megaloptera: Sialidae). The newly reported diversity includes: Jersiberotha myanmarensis sp. n. (Berothidae); Jersiberotha tauberorum sp. n. (Berothidae); Iceloberotha kachinensis gen. et sp. n. (Berothidae); Iceloberotha simulatrix sp. n. (Berothidae); Haploberotha persephone gen. et sp. n. (Berothidae); Telistoberotha libitina gen. et sp. n. (Berothidae); Systenoberotha magillae gen. et sp. n. (Berothidae); Dasyberotha eucharis gen. et sp. n. (Berothidae); Ethiroberotha elongata gen. et sp. n. (Berothidae); a beaded lacewing larva (Berothidae); Scoloberotha necatrix gen. et sp. n. (Rhachiberothidae); Litopsychopsis burmitica gen. et sp. n. (Psychopsidae); a silky lacewing larva (Psychopsidae); a putative first-instar silky lacewing larva (Psychopsidae); Elenchonymphes electrica gen. et sp. n. (Nymphidae); an osmylid lacewing larva (Osmylidae); and a fragmentary alderfly (Megaloptera: Sialidae). In New Jersey amber is described a new thorny lacewing, Rhachibermissa phenax sp. n. (Rhachiberothidae), while from Canadian amber are reported an egg and first instar of a green lacewing (Chrysopidae), the fragmentary remains of a beaded lacewing distinct from Plesiorobius but left unassigned (Berothidae), and a fragmentary larva of a mesoraphidiid snakefly (Raphidioptera: Mesoraphidiidae). The subfamily Paraberothinae is newly synonymized with Rhachiberothinae (syn. n.). The neuropterid fauna of Burmese, New Jersey, and Canadian amber is briefly summarized.
Abstract: A new fossil antlion, Choromyrmeleon aspoeckorum n.sp., is described and illustrated from the Yixian Formation (conservatively considered as Early Cretaceous or latest Jurassic) of Liaoning Province, China. This constitutes the second species of the genus Choromyrmeleon as well as the second fossil record of Myrmeleontidae from China.
Abstract: A fossil micropterigid moth (Zeugloptera: Micropterigoidea) is described and figured from a well preserved forewing of the Fur Formation [âMo-clayâ: earliest Eocene (Early Ypresian)] of Denmark. Moleropterix kalbei nov. gen. et nov. sp. is distinguished from other fossil micropterigids as well as modern counterparts.
Abstract: A fossil cimicoid bug is described and figured from a single male preserved in mid-Cretaceous (latest Albian) amber from Myanmar. Quasicimex eilapinastes n.gen., n.sp., shares many features with the bed bug family Cimicidae (Cimicomorpha: Cimicoidea), as well as a few features of primitive cimicids such as Primicimicinae, while simultaneously retaining some significant plesiomorphies relative to crown-group cimicids. The genus is tentatively retained in Cimicidae s. lato, basal to all other cimicids.
Abstract: The first formally described fossil of the beetle family Prostomidae (Tenebrionoidea) is presented. Vetuprostomis consimilis n.gen. et n.sp., is described and figured from a single individual preserved in mid-Cretaceous amber from Myanmar (Burma). The fossil is remarkably similar to modern prostomids from which it is distinguished. The only other records of fossil jugular-horned beetles are three undescribed Baltic amber inclusions in a private collection.
Abstract: Well-preserved insects are reported in pelitic sediments from the Feldbach and Paldau Formation (Late Miocene: Early Pannonian) of the Styrian Basin, Austria. Herein we provide a brief overview of the hitherto recovered fauna. The fauna includes representatives of five insect orders, representing at least 10 morphospecies of the families Phrygaenidae (Trichoptera), Tipulidae (Diptera), putative Rhagionidae (Diptera), Carabidae and Chrysomelidae (Coleoptera), Formicidae (Hymenoptera), and a giant primitive termite near Termopsidae (Isoptera). The various identifiable morphospecies recovered to date are summarized.
Abstract: A new crabronid wasp species, Pison menkei sp. n., is described and figured from a single female preserved in Early Miocene (Burdigalian) Dominican amber. The fossil is herein distinguished from extant and extinct congeners and regarded as an isolated species removed from modern Neotropical species groups. It is the seventh named fossil Pison and the second from Dominican amber.
Abstract: A new morphological structure is described and figured for orchid bees (Apinae: Euglossini). These minute features are located posterolateral to the ocelli and are termed âjuxtocellar structuresâ. The diversity of their form across Euglossini is described. Homologous structures were found in some related lineages of apid bees (e.g., Anthophorini, Centridini, Eucerini) while in others the structures were absent (e.g., Xylocopinae, Nomadinae). Most notably juxtocellar structures are apomorphically lost in all other corbiculate apine tribes (i.e., Bombini, Meliponini, Apini, Electrobombini, Electrapini, and Melikertini), a pattern which is in accordance with current phylogenetic hypotheses for the clade.
Abstract: Four new cleptoparasitic bees of the tribe Ammobatini (Apidae: Nomadinae) are described and figured from southeastern Pakistan and Sri Lanka. Ammobates maxschwarzi sp. n. (Pakistan) is known from a series of females, while A. cinnamomeus sp. n. (Pakistan), Parammobatodes craterus sp. n. (Pakistan), and Chiasmognathus taprobanicola sp. n. (Sri Lanka) are known from males and females. The species are distinguished from their congeners and comments made on potential hosts. In addition, C. gussakovskii (Popov), previously known only from Tajikistan, is newly recorded from southern Kyrgyzstan.
Abstract: A family of extinct lacewings is described and figured from the Yixian Formation [Early Cretaceous (M.S.E.) or latest Jurassic (D.R.)] of Liaoning Province, China. Aetheogrammatidae Ren and Engel, new family, is established for Aetheogramma speciosa Ren and Engel, new genus and species, and is distinguished from other myrmeleontiform families. The enigmatic kalligrammatid genus Kallihemerobius Ren and Oswald is placed in a new subfamily, Kallihemerobiinae Ren and Engel.
Abstract: The first Cretaceous fossil of the wasp family Rhopalosomatidae (Aculeata: Euaculeata: Vespoidea) is described and figured from a male preserved in Burmese amber. Eorhopalosoma gorgyra Engel, new genus and species, is the first rhopalosomatid discovered in amber and the second â but first definitive â member of the family from the Cretaceous. The new species is distinguished from its modern counterparts. The modern genus Olixon Cameron is transferred to a separate new subfamily, Olixoninae.
Abstract: A new species of fig wasp, Ceratosolen (Ceratosolen) polyodontos n. sp., is described from females captured at Los Baños, Luzon, Philippines. The species can be distinguished from its congeners by the possession of a much greater number of ventral mandibular lamellae (22â23), divided into an anterior and posterior area, and posterior metasomal structures associated with the ovipositor.
Abstract: Publishing taxonomic and systematics studies in the digital era faces major challenges and requires new approaches, many of which are currently stimulating spirited discussions among taxonomists and systematists. New amendments to the International Code of Zoological Nomenclature are expected to regulate electronic publishing of new taxa and create a standard form for their registration (ZooBank). Responding to a perceived need, this editorial announces establishment of ZooKeys - a new online and print journal in zoological taxonomy and systematics, which aims to quickly respond and adapt to the newest developments in taxonomic publishing. Open Access is accepted as mandatory for ZooKeys. The rationale for and concept of ZooKeys is discussed in detail.
Abstract: Cretopiesma suukyiae, new genus and species, is described, based on a unique female specimen in mid-Cretaceous (c. 100 myo) amber from northern Myanmar. Features of C. suukyiae unique for the small Recent family Piesmatidae include a long, protrudent clypeus, a dorsal carina of the head, lack of "jugal" lobes/appendices, widely separated coxae, very large scutellum, and the venation of the corium; some of these are plesiomorphic and shared with Aradidae. C. suukyiae possesses among the cuticular areolation and propleural cavities distinctive to Piesmatidae. Phylogenetic analysis of Recent and fossil genera of piesmatids resulted in a cladogram with Cretopiesma as sister group to the remainder of the family. Relationships of this unusual species and of Piesmatidae within Pentatomomorpha are discussed.
Abstract: A new species of the highly specialized hemipteran family of social ectosymbionts, the Termitaphididae, is reported in Miocene amber from the Dominican Republic: Termitaradus avitinquilinus, n. sp. It differs from the only other fossil termitaphidid (Termitaradus protera Poinar and Doyen), in Miocene Mexican amber, and from modern species by various distinctive features. Two of the three specimens of T. avitinquilinus are preserved in a piece of amber with its presumed host, a worker of the basal termite, Mastotermes electrodominicus Krishna and Grimaldi. The hypothesis that the family Termitaphididae is ancient and inhabited Pangaea is disputed in favor of a much more recent, Tertiary origin. It is suggested, in fact, that termitaphidids are highly modified Aradidae, possibly derived from the Mezirinae.
Abstract: A new high-elevation species of the augochlorine bee genus Augochloropsis Cockerell (Anthophila: Apoidea: Halictidae: Halictinae: Augochlorini) is described and figured. Augochloropsis (Paraugochloropsis) isabelae, new species, is distinguished from its congeners and it is noted that females of the species exhibit head size variation. The variation in head size and morphology seen between females is suggestive of social behavior in this species. Where known, species of Augochloropsis range from solitary to semisocial, and there is even a single record of primitive eusociality in the genus.
Abstract: Two new genera and species of snakeflies (Raphidioptera) are described and figured from the Jurassic Jiulongshan Formation of eastern Inner Mongolia, China. Ororaphidia megalocephala, new genus and species, and Styporaphidia magia, new genus and species, are both differentiated from other Mesozoic snakefly lineages. These constitute the oldest records of snakeflies from China.
Abstract: A new genus and species of primitive termite (Isoptera) is described and figured from the remains of an imago preserved in middle Eocene (Lutetian) Baltic amber. Idanotermes desioculus Engel, new genus and species (near to Mastotermitidae), is distinguished from related genera as well as all other termites known as inclusions in Baltic amber. In addition, the genus Heterotermes (Rhinotermitidae: Heterotermitinae) is newly recorded from Baltic amber and Heterotermes eocenicus Engel, new species, is described. A revised key to Baltic amber termites is provided so as to accommodate the newly discovered species.
Abstract: A new species of leaf-cutter bee, Megachile (Eutricharaea) ventrisi sp. n., is described and figured from western Saudi Arabia. The species is remarkably similar to M. walkeri Dalla Torre from the same region but differs strikingly in the structure of the metafemoral setae, metasomal scopa, and clypeal integument.
Abstract: A new species of the rare cleptoparasitic bee genus Chiasmognathus Engel (Ammobatini) is described and figured from a female collected in northwestern Iran. Chiasmognathus rhagae sp. n. is generally similar to C. orientanus (Warncke) (Bulgaria, Turkey, Cyprus, Crete, and the Levant) or C. pashupati Engel (from southeastern Pakistan) but can be readily recognized by coloration and integumental sculpturing. In addition, a new locality in Israel is provided for C. aegyptiacus (Warncke).
Abstract: A new genus and species of the peculiar braconid wasp subfamily Masoninae is described and figured from a female preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Anoblepsis konokeraia Engel and Bennett, new genus and species, belongs to the tribe Masonini but differs from the hitherto only known genus, Masona van Achterberg. The genus is diagnosed and compared with its modern counterparts. Anoblepsis konokeraia is the third braconid species documented from Dominican amber and the second of the subfamily Masoninae.
Abstract: The remains of a new species of zorapteran (Polyneoptera: Anartioptera: Mystroptera: Zoraptera) are described and figured from a female preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Zorotypus (Zorotypus) mnemosyne Engel sp. n. is represented by a female of the apterous, blind morph (âapteronâ) and can be readily distinguished from the only other Miocene zorapterans, Z. goeleti Engel & Grimaldi (winged morph) and Z. palaeus Poinar (apterous morph) in the terminal abdominal structure, the shape of the cerci, and the spination of the metafemora. Notes on the status of the recently described Jordanian amber (contemporaneous with Lebanese amber) zorapteran are provided and the genus Paleospinosus is newly synonymized with Octozoros Engel (new synonymy) resulting in the new combination Z. hudae (Kaddumi). In addition, the name âZ. confirmans Schevenâ is considered a nomen nudum et invalidum. However, the taxon is distinct from other Dominican amber species and is therefore validated as Z. absonus Engel sp. n. and distinguished from other fossil zorapterans.
Abstract: A new species of the extinct bee genus Ctenoplectrella Cockerell (Megachilinae: Ctenoplectrellini) is described and figured from two females preserved in a single piece of middle Eocene (Lutetian) Baltic amber. Ctenoplectrella gorskii Engel, sp. n. is distinguished from other species of the genus most notably in the venation of the forewing.
Abstract: The termite species from Brazil's Early Cretaceous (Aptian-aged) Crato (Santana) Formation are evaluated on the basis of the degree of character variation seen in modern species, using a series of 56 specimens, scanning electron microscopy of minute structures, and a bivariate plot of the proportional sizes of sclerotized body structures. Of the previously described species only the following are considered valid: Mariconitermes talicei Fontes and Vulcano, Meiatermes araripena Krishna, Cratomastotermes wolfschwenningeri Bechly, Cratokalotermes santanensis Bechly, and Cretarhinotermes novaolindense Bechly. The combination M. araripena Krishna (once placed in Cretatermes) is restored, and Cretatermes pereirai Fontes and Vulcano is proposed as a junior synonym of M. araripena. The following new species is described: Meiatermes hariolus Grimaldi, new species. The following are considered nomina dubia based on superficial and even contradictory diagnoses: Caatingatermitinae Martins-Neto et al. (likely synonymous with Hodotermitinae); Araripetermes nativa Martins-Neto et al. (nomen incorrectum, recte: nativus), Caatingatermes megacephalus Martins-Neto et al., and Nordestinatermes obesa Martins-Neto et al. (nomen incorrectum, recte: obesus). These are probably all new synonyms of the Cretaceous genus Meiatermes Lacasa-Ruiz and MartÃnez-Delclòs and of M. araripena in particular, but this assertion cannot be verified without access to the type specimens. Of the six definitive species of Crato termites, reconstructions are provided for four. All species appear to be basal taxa, either a stem group to Isoptera, to Hodotermitidae sensu lato, or to Kalotermitidae. Despite very partial preservation, Cretarhinotermes appears to be within the hodotermitid grade and not a rhinotermitid. Some specimens have yielded detailed preservation of soft internal tissues, including the midgut, which has further phylogenetic implications.
Abstract: Specimens of Paleozoic Coleoptera are quite rare, particularly so in North America where they have been hitherto represented by a single specimen. The first Permian beetle from North America was only recently described from a single specimen of a complete elytron (no counterpart). A second specimen from another locality in those same beds, comprised of the part and counterpart of a fragment of an elytron, is figured and discussed here. While representing a somewhat larger individual than the original specimen, the present material appears to represent the same species, Permocoleus wellingtonenesis Lubkin and Engel.
Abstract: The morphology of the enigmatic, Mesozoic, aquatic insect family Chresmodidae is redescribed and its phylogenetic affinities among the polyneopterous orders discussed. Study of the complete venation of both fore- and hind wings observed in some specimens from the Spanish Barremian, permit us to postulate the hypothesis that the family belongs to the Archaeorthoptera, thus to the orthopteroid lineage rather than to crown-group Phasmatodea or to the more inclusive group Holophasmatodea (sensu Grimaldi & Engel, 2005). New specimens from Spain, Lebanon, Brazil, and Germany permit a new re-description of some chresmodid body structures, with concomitant implications for the phylogenetic position of the family. Chresmoda neotropica n. sp. is described from the Aptian-Albian of the Crato Formation (northeast Brazil). The functional morphology proposed for some of their specialized structures suggest a new hypothesis of Chresmoda palaeobiology, and related to this some implications for the localized palaeoenvironment as well as global palaeoclimate.
The problematic Sternarthron spp. from the Upper Jurassic of Solnhofen were described as probable palpigrades (Arachnida: Palpigradi), based on type material originally thought to be fossil insects. The affinities of Sternarthron Haase, 1890 have been questioned. Our restudy of Haaseâs types clearly indicates that these fossils represent nymphal specimens of chresmodids. Consequently, Sternarthron has to be considered as a junior synonym of the fossil insect genera Propygolampis Weyenbergh, 1874 and Chresmoda Germar, 1839.
Abstract: The first Dominican amber fossil of the parasitoid family Orussidae (Euhymenoptera: Orussomorpha) is described and figured from a single individual preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Ophrynopus peritus Engel, new species, is the first orussid fossil described from Tertiary amber and the first species documented from the West Indies.
Abstract: The bees (Apoidea: Anthophila) first diverged from the spheciform wasps sometime in the later part of the Early Cretaceous, about 125 million years ago. The sister group of the bees is probably the Crabronidae within the paraphyletic Sphecidae, which contains about 90% of the described spheciform diversity. The palaeoentomological data for sphecids are relatively poorly understood, but support the assumption of an early Cretaceous origin of the Apoidea (bees + sphecids). Cladistic, paleobotanical, and paleclimatological evidence suggests that the bees and sphecids originated in semi-arid regions of Gondwanaland and diversified from there. By the Late Cretaceous the major lineages had been established and these persisted into the Early Tertiary, with seemingly little affect from the K-T mass extinction event. The bees appear to have experienced more significant losses during the dramatic episode of global cooling marking the Eocene-Oligocene transition (one of the famed, "lesser" mass extinction events). Indeed, at this time several lineages of eusocial bees became extinct, leaving the four distinctive survivors of the corbiculate bees (i.e., honey bees, bumble bees, orchid bees, and stingless bees). Incorporating these "missing" tribes into phylogenetic studies supports the single origin hypothesis for eusocial and advanced eusocial behavior within this group of bees. During the Oligocene the bee and sphecid wasp fauna took on a relatively modern appearance and as such can be thought of as the epoch in which the modern bee fauna was born. Bees from the Oligocene and younger deposits provide evidence of localized extinctions (e.g., losses of several genera from the West Indian islands), recent episodes of diversification (e.g., speciation events among the honey bees), and other smaller-scale, evolutionary phenomena. ---------- Zusammenfassung: Bienen (Apoidea: Anthophila) entstanden irgendwann im späteren Verlauf der Unterkreide, also vor etwa 125 Millionen Jahren, aus Grabwespen-ähnlichen Vorfahren. Die Schwestergruppe der Bienen sind wahrscheinlich die Crabronidae innerhalb der paraphyletischen Sphecidae, die etwa 90% der bekannten Grabwespendiversität umfassen. Die paläoentomologischen Daten sind bei Grabwespen nunr unzureichend untersucht, unterstützen aber die Annahme eines unterkreideteitlichen Ursprungs der Apoidea (Bienen und Grabwespen). Kladistische, paläobotanische und paläoklimatologische Belege lassen vermuten, dass Bienen und Grabwespen in semi-ariden Gebieten Gondwanas und sich von hier aus diversifizierten. Bis zur Oberkreide hatten sich alle wesentlichen Linien etabliert und überdauerten bis in der frühe Tertiär, offenbar wenig beeinträchtigt durch das Massenaussterben an der K-T-Grenze. Stattdessen erlitten insbesondere die Bienen stärkere Verluste durch während der dramatischen Phase der globalen Abkühlung, die den Ãbergang vom Eozän zum Oligozän kennzeichnet (eine der bekannten, weniger signifikanten Massenextinktionen). In dieser Zeit starben eine Reihe von eusozialen Bienentaxa aus, und nur vier Gruppen eusozialer Bienen überlebten bis heute (die Honigbienen, die Hummeln, die Prachtbienen und die stachellosen Bienen). Phylogenetische Analyse der Beziehungen innerhalb der Bienen unter Einbeziehung der ausgestorbenen eusozialen Gruppen stützen die Annahme eines einmaligen Entstehens eusozialen und des höheren eusozialen Verhaltens innerhalb dieser Bienen. Während des Oligozäns nahm die Bienen- und Grabwespenfauna kontinuierlich moderne Gestalt an, und so kann diese Epoche als Geburtsstunde der moderne Bienen- und Grabwespenfaune angesehen werden. Bienen des Oligozäns und jüngerer Fossillagerstätten belegen lokale Extinktionen (z.B. der Verlust einiger Bienengattungen von den Westindischen Inseln), relativ rezente Diversifikationsereignisse (z.B. Speziationen bei Honigbienen) sowie anderer, weniger bedeutsamer Evolutionsereignisse.
Abstract: Three new species of the parasitoid wasp superfamily Mymarommatoidea (Proctotrupomorpha: Bipetiolarida) are described and figured in Cretaceous amber from New Jersey (Turonian) and Myanmar (Albian-Cenomanian boundary). The new taxa are Archaeromma carnifex Engel and Grimaldi, new species, in New Jersey amber, A. gibsoni Engel and Grimaldi, new species, in Burmese amber (both Mymarommatidae), and Galloromma kachinensis Engel and Grimaldi, new species, in Burmese amber (Gallorommatidae).
Abstract: A new species of the bee genus Ctenoplectra Kirby (Apinae: Euceriti: Ctenoplectrini) is described and figured from a series of males and females collected in central and northeastern Sulawesi (Indonesia). Ctenoplectra elsei sp. n. is most superficially similar to C. chalybea Smith (a largely mainland species with which many metallic blue species are confused) but differs in integumental sculpturing and structure of the terminalic sclerites, among other features. The species has been collected from Momordica cochinchinensis (Lour.) Spreng. (Cucurbitaceae: Cucurbitoideae: Joliffieae: Thladianthinae). The available information on nesting biology in Ctenoplectra is briefly reviewed along with flower visitation by Southeast Asian species.
Abstract: The first Chinese fossil of the family Nymphidae (Neuroptera: Myrmeleontiformia) is briefly described and figured along with two episomyline Osmylidae, a generally plesiomorphic group which can be easily confused with nymphids when only wings are known. Four new species and three new genera are characterized from Jurassic deposits of the Jiulongshan Formation (Daohugou Biota), Inner Mongolia, China. New genera are Liminympha (Nymphidae), Enodinympha (Osmylidae), and Nilionympha (Osmylidae), while the new species are Liminympha makarkini, Enodinympha translucida, Nilionympha pulchella, and N. imperfecta.
Abstract: The species of the firefly genus Vesta occurring in Taiwan are revised. Three species are recognized: Vesta scutellonigra Olivier (nomen emendatum), V. impressicollis Fairmaire, and V. rufiventris (Motschulsky). Vesta chevrolatii var. scutello nigro Olivier is automatically emended to scutellonigra under ICZN rules. Okada described as a distinct species V. scutellonigra, employing the same name and material as Olivier, thereby automatically rendering his taxon a junior synonym (n. syn.). The name V. scutellonigro, simultaneously employed by Okada, is considered an incorrect subsequent spelling and therefore unavailable. Vesta formosana Pic (n. syn.) is a new junior synonym of V. scutellonigra. The spelling chevrolati, employed by Gemminger and Harold, is determined to be an incorrect subsequent spelling of chevrolatii and thus unavailable. Lectotypes are newly designated for V. scutellonigra Olivier, V. impressicollis Fairmaire, and V. formosana Pic. A key to the species of Taiwan is provided. Vesta flaviventris (Fairmaire) (n. comb.) from China is transferred from Lucernuta.
Abstract: The geological history of the wasp superfamily Cynipoidea is reviewed, with the description of various new taxa mostly in Late Cretaceous amber from New Jersey and Canada. The various fossil lineages are incorporated into a phylogenetic analysis of the superfamily and their implications for understanding the evolution of the group explored. The following new taxa or taxonomic changes are proposed (authorship of all taxa Liu and Engel): Protimaspidae, new family; Stolamissidae, new family; Stolamissus, new genus; Stolamissus mirabilis, new species; Proliopterinae, new subfamily; Proliopteron, new genus; Proliopteron redactus, new species; Goeraniinae, new subfamily; Goerania, new genus; Goerania petiolata, new species; Micropresbyteria, new genus; Micropresbyteria caputipressa, new species; Anteucoila, new genus; Anteucoila delicia, new species; Jerseucoila, new genus; Jerseucoila plesiosoma, new species; Syneucoila, new genus; Syneucoila magnifica, new species; Tanaoknemus, new genus; Tanaoknemus ecarinatus, new species; Kinseycynips, new genus; Kinseycynips succinea (Kinsey), new combination. The extinct family Rasnicynipidae is newly transferred to Figitidae and classified as a basal subfamily therein (Rasnicynipinae, status novus). The Gerocynipidae, its type genus Gerocynips, and the type species upon which they are founded, Gerocynips zherichini, are found to be nomenclaturally unavailable. Gerocynips zherichini is regarded as a nomen nudum; the genus newly validated as Gerocynips, new genus (with G. siberica Kovalev as type species); the family validated as Gerocynipidae, new family. The fossil records of Cynipoidea are summarized.
Abstract: A brief overview and dichotomous key is provided for those termites (Isoptera) occurring in middle Eocene (Lutetian) Baltic amber. Ten species in seven genera are presently documented from Baltic amber, these being as follows: Garmitermes succineus n. gen. n. sp., Termopsis bremii (Heer), T. ukapirmasi n. sp., Archotermopsis tornquisti Rosen, Proelectrotermes berendtii (Pictet-Baraban), Electrotermes affinis (Hagen), E. girardi (Giebel), Reticulitermes antiquus (Germar), R. minimus Snyder, and Parastylotermes robustus (Rosen). Garmitermes succineus n. gen. n. sp. is the first Mastotermes-like termite discovered in Baltic amber. Hemerobites Germar, a senior synonym of Reticulitermes Holmgren, is designated as a nomen oblitum under the ICZN rules. Maresa Giebel, another senior synonym of Reticulitermes, is tentatively suppressed pending a petition to the ICZN to preserve the latter name.
Abstract: New fossil termites (Isoptera) are described and figured from four Early Cretaceous deposits across Asia, including some of the oldest records for the order. In total seven new genera and six new species are established from these sites. A single, alate specimen is documented from the Zaza Formation (Berriasian) of Baissa, Transbaikalia (Siberia, Russia) and is described as Baissatermes lapideus n. gen. n. sp. Baissatermes lapideus n. gen. n. sp. is the oldest fossil termite presently known and the oldest known example of a social organism. Valditermes acutipennis Ponomarenko, from the Hauterivian of Mongolia, is transferred to a new genus, Khanitermes n. gen. (resulting in the new combination, Khanitermes acutipennis n. comb.). Melqartitermes myrrheus n. gen. n. sp. is described in Neocomian (Hauterivian) amber from Lebanon. The late Albian to early Cenomanian Burmese amber (Myanmar) harbors the greatest diversity of termites hitherto discovered from any Cretaceous amber locality. In total six species are documented in Burmese amber, including the following new taxa and combinations: Mylacrotermes cordatus n. gen. n. sp., Dharmatermes avernalis n. gen. n. sp., Proelectrotermes swinhoei (Cockerell) n. comb., P. holmgreni n. sp., Kachinitermes n. gen., Kachinitermes tristis (Cockerell) n. comb., Tanytermes anawrahtai n. gen. n. sp. The significance of these new taxa for understanding early termite evolution and basal relationships within Isoptera is discussed. A checklist of Cretaceous termites is provided.
Abstract: The purpose of this application, under Article 23.9.3 of the Code, is to conserve the usage of the genus-group name Reticulitermes Holmgren, 1913 (type species Termes flavipes Kollar, 1837) for an economically important and universally known group of termites, the subterranean termites responsible for most economic damage in the Northern Hemisphere. Two genus-group names have priority over Reticulitermes, both having been established for a common species of fossil termite in Baltic amber. Hemerobites Germar, 1813 (type species Hemerobites antiquus Germar, 1813) has not been used after 1899 and can be considered a nomen oblitum under Article 23.9.1 of the Code. Maresa Giebel, 1856 (type species Maresa plebeja Giebel, 1856) has been used once since 1899, in a large compendium on fossil insects, and, therefore, cannot be automatically considered a nomen oblitum. It is proposed that the genus-group name Reticulitermes be given precedence over Maresa.
Abstract: A new bee species of the genus Amegilla (Apinae: Anthophorini) is described and figured from northeastern Egypt. Amegilla argophenax spec. nova, belongs to the fasciata group and is most similar to A. deceptrix (Priesner) comb. nova which occurs in the same region. Characters are provided to distinguish the species from its congeners. Podalirius pyramidalis Kirby, from Socotra (Republic of Yemen), is resurrected from synonymy under Amegilla albigena (Lepeletier de Saint Fargeau) (as A. pyramidalis comb. nova) where it is, like A. argophenax and A. deceptrix, a member of the fasciata group.
Abstract: Those weevils belonging to the subfamily Cossoninae occurring in Early Miocene (Burdigalian) amber from the Dominican Republic are described and figured. The fauna includes 10 species (nine new): Micromimus orcus spec. nova, Caulophilus ashei Davis & Engel, C. falini spec. nova, C. swensoni spec. nova, C. bennetti spec. nova, Dryotribus amplioculus spec. nova, Paralicus abnormis spec. nova, Proeces longirostrum spec. nova, Stenotrupis breviscapus spec. nova, and Cossonus hinojosai spec. nova.
Abstract: Two new orchid bees of Euglossa subgenus Euglossella (Apinae: Euglossini) are described and figured from Peru. Both Euglossa (Euglossella) cosmodora sp. n. and E. (E.) urarina sp. n. belong to the decorata species group (i.e., the clade superficially resembling stingless bees of the genus Melipona) of Euglossella and are distinguished from their congeners as well as each other.
Abstract: An unusual new lampyrid genus, Oculogryphus Jeng, Engel, and Yang, new genus, and its type species, Oculogryphus fulvus Jeng, new species, from Vietnam, is described and figured. The genus is characterized by its filiform antennae, large and ventrally approximate compound eyes that are deeply emarginate posteriorly, largely exposed head (i.e., not covered by the pronotum), eight abdominal ventrites, rectangular abdominal tergites, absence of photogenic organs, and symmetric aedeagal sheath and genitalia in the male (the female is presently unknown). The genus does not adequately fit into any subfamily as they are currently defined, highlighting the difficulties of the subfamilial and tribal classification presently in use. Though most likely allied to Ototretinae or Luciolinae, the systematic position of Oculogryphus cannot be determined as currently there is no comprehensive phylogeny of Lampyridae and related groups. The species may rely on photic cues for purposes of mating. A key to the genera of Lampyridae (sensu Crowson) occurring in Palaearctic East Asia, Oriental, and Australian regions is provided. Some taxonomic problems of Ototretinae are commented upon.
Abstract: Two new species of augochlorine bees are described and figured from Ecuador. Caenaugochlora silvicola sp. n. is the first species of its genus with a relatively elongate malar space (although admittedly still shorter than true long-headed augochlorines) and can be readily distinguished from other Caenaugochlora by this trait as well as characters of coloration and integumental sculpturing. Caenaugochlora cuprina Moure & Hurd (= Augochlora cupriventris Friese), also from Ecuador, has proven to be a junior synonym of Megaloptilla callopis (Vachal) (syn. n.). Thus, C. silvicola sp. n. is the first true species of this genus formally documented from Ecuador. The second species does not correspond to any known group of augochlorines and is placed in a new genus herein. Rhynchochlora chlerogopsis gen. et sp. n. is most similar to Chlerogas but differs in head, antennal, and mesoscutellar structure.
Abstract: The zorapteran fauna of Fiji is reviewed based on newly collected material from the âTerrestrial Arthropod Survey of Fijiâ. A single, endemic species, Zorotypus zimmermani Gurney, is recognized from Fiji, with new records from Vanua Levu and Taveuni. The alate has hitherto not been characterized and a description of the wing venation is accordingly provided for the species.
Abstract: Cenocephalus quasiexquisitus n.sp., is described and figured from the remains of a male preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. The new species is distinguished from C. exquisitus Bright & Poinar, also from Dominican amber.
Abstract: A lateral gynandromorph of Thyreus redactulus? Cockerell is described and figured, with a particular emphasis on the genitalic sclerites. As such, the sting morphology and mechanism are described for T. ramosus (Lepeletier de Saint Fargeau) so as to provide a comparative framework for understanding the gynandromorph. The implications of the gynandromorphâs genitalic arrangements for intersexual homologies are explored.
Abstract: A new bee species of the genus Amegilla (Apinae: Anthophorini) is described and figured from western Malaysia and southern Thailand. Amegilla anekawarna Engel, new species, belongs to the zonata group of species and can be most readily con