Desiomorphs in Amber

American Entomologist • Winter 2012 214 George Poinar, Jr. Desiomorphs in Amber Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 ossils from various epochs often do not easily fit into the present system of classification because they have morphological characters found in two or more extant or extinct groups, from the generic to the ordinal level. Without genetic analysis, it is difficult to determine their phylogenetic affinity and how they should be classified. We have previously referred to them as chimeras (Poinar and Poinar 2008) in the literary sense, which implies a creature combining features of two or more separate animals, such as the Greek “she-monster” with a lion’s head, goat’s body, and serpent’s tail; the part-horse, part-human centaur; or the minotaur with the head of a bull and body of a man. However, chimera now has a medical definition that makes its use confusing. Transitional fossils imply a phylogenetic relationship (like the older terms “missing links” and “intermediate forms”) between the fossil in question and similar lineages and are often regarded as stem groups. However, morphological similarities do not always imply phylogenetic relationships because a fossil can represent a separate, unique lineage or similarities can be the result of convergence. A case in point is Archaeopteryx, a Jurassic vertebrate considered a “transitional fossil.” Over the years, it has been considered a winged reptile, a primitive bird, a winged dinosaur, and more recently, a link between birds and dinosaurs. However, there is no hard evidence to say it does not represent a completely separate lineage that is only distantly related to either birds or dinosaurs. Because of the uncertainty of determining whether a fossil is truly transitional and phylogenetically intermediate between two or more separate lineages, I propose the term “desiomorph” (from the Greek desis, “a binding together” and morphe, “form or shape”) to describe fossil individuals that possess morphological characters found in two or more groups, whether fossil, extant, or a combination of both. Desiomorphs are characterized by morphology alone, without any direct implication of phylogenetic relationships. A recent vertebrate desiomorph is the 50 mya Eocene bat, Onychonycteris finneyi Simmons, Seymour, Habersetzer and Gunnell (2008), which has characters of modern bats as well as those of ground-dwelling mammals that used their claws to ascend trees, but is not phylogenetically transitional between these two groups. The present work provides accounts of desiomorphs, mostly insects, encountered in amber. Even though there are rare accounts of extant insects surviving since the middle Tertiary (Hörnschemeyer et al. 2009) and some modern insect genera detected as far back as the Early Cretaceous (Poinar and Milki 2001), the great majority of insect species exist only for several million years. As one goes back in time, desiomorphs become more common and the differences between them and extant lineages become more pronounced, thus widening their systematic separation. Determining the level of difference and systematic position of desiomorphs is generally a combination of objective and subjective judgments. The amber-embedded Tertiary and Cretaceous desiomorphs discussed below have morphological characters connecting them to two or more extant or extinct genera, subfamilies, families, or orders. F three of the five known New World genera. Paleoeuglossa melissiflora Poinar (1998) shared its metallic sheen with members of the extant genera Euglossa and Eufresea and its scutellar tuft with Euglossa and Eulaema. Its protruding clypeus and genal projections (Fig. 1) partly account for its placement in a new genus. Both specimens were females, and finding a pair together in amber suggests that they were collecting resin for nest construction when entrapped, a habit similar to the behavior of stingless bees that are commonly found in Dominican amber. Orchid bees have since been extirpated from Hispaniola. Desiomorphs at the Subfamily Level The batfly Enischnomyia stegosoma in 20-30 mya Dominican amber. Bat flies (Streblidae and Nycteribiidae) feed on the blood A Desiomorph at the Generic Level The orchid bee Paleoeuglossa melissiflora in 20-30 mya Dominican amber. New World orchid bees have attracted the attention of entomologists and botanists for their pollination activities, often being the sole pollinators for tropical orchid species. These bees are normally fairly large and often brilliantly colored with metallic markings. A pair of orchid bees in Dominican amber had characters of American Entomologist • Volume 58, Number 4 Fig. 2. The batfly Enischnomyia stegosoma in Dominican amber. 215 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 Fig. 1. The orchid bee Paleoeuglossa melissiflora in Dominican amber. Brown 2006), but the additional segment in the front legs was the really astonishing aspect (Fig. 4). All extant insects possess legs with 5 basic segments: the coxa, trochanter, femur, tibia, and tarsus. The presence of a “patella” at the apical half of the tibia with a bend at its joint showed that this is a true dicondylic joint (allowing movement in only one direction). While myriapods, mites, and spiders possess patellae, one has to travel back to the Late Paleozoic to find insect groups with extra segments, such as members of the Paleodictyoptera, Monura, and Thysanura. Since extant scydmaenids use their forelegs to manipulate mite prey during feeding, the extra foreleg segment on Hapsomela probably served as a maneuvering device for securing mites, an example of functional morphology. It is unknown whether Hapsomela’s front legs were a resurgent ancestral character from Palaeozoic times or the result of a spontaneous mutation that occurred in this particular lineage. Desiomorphs at the Family Level The flat bug Brevisensoria incrustata in 20-30 mya Dominican amber. Termite bugs (Termitaphididae: Hemiptera) are small, flattened insects that live in the nests of termites (Fig. 5). In such a hazardous environment, intruders require good defenses. Thus, the head, thorax, and abdomen of termite bugs are merged into a hardened, smooth dorsum, analogous to the shell of a turtle. The geniculate antennae and shortened legs can be withdrawn under this protective outer case. Living in a cavern-like environment for millions of years has left termite bugs eyeless and wingless. Scientists feel that termite bugs are closely related to flat bugs (Aradidae: Hemiptera) that have distinct heads and eyes and exposed antennae. Brevisensoria incrustata in Dominican amber has characters of both flat bugs and termite bugs, as well as a few unique features of its own. This fossil lacks eyes and wings and the dorsum forms a continuous body covering similar to termite bugs; however, the exposed antennae, longer legs, and incrusted dorsum align it with the flat bugs (Fig. 6). Its own unique characters include an unusual body shape and antennal structure. It is impossible to determine the original retreat of Brevisensoria. The absence of eyes, incrusted surface, and fused dorsum suggest that it inhabited a dark region Fig. 3. The strange ant-like stone beetle Hapsomela burmitis in Burmese amber. Arrows show joints between patella and tibia in front legs. Fig. 4. Foreleg of Hapsomela burmitis with extra segment. F=femur; P=patella; T=tibia. Arrow shows joint between patella and tibia. 216 Fig. 5. The blind, wingless termite bug Termitaradus dominicanus in Dominican amber. Fig. 6. The blind flat bug Brevisensoria incrustata in Dominican amber. American Entomologist • Winter 2012 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 of bats and spend the greater part of their lives on their hosts, exiting only when searching for mates. A streblid bat fly, Enischnomyia stegosoma Poinar and Brown (2012) in Dominican amber possessed characters of two extant subfamilies (Fig. 2). The strongly compressed body and laterally flattened profemora are characteristic of the Nycterophilinae, while the fairly complete wing venation, antennal pedicel extending above the flagellum, and elongate proboscis are characteristic of the Trichobiinae. A biological character that allies the fossil with members of the Nycteribiidae pertains to its role as a vector of bat malaria. Oocysts and sporozoites of bat malaria, Vetufebrus ovatus Poinar (2011), were found in the gut wall and salivary glands of the fossil fly. Presently, there are no records of streblids transmitting malaria; however, representatives of at least four genera of Nycteribiidae are known vectors of bat malaria. The ant-like stone beetle Hapsomela burmitis in 97-110 mya Burmese amber. A most unique desiomorph was a little beetle with its front legs equipped with 6 instead of the normal 5 segments found in all living insects (Fig. 3). While Hapsomela burmitis was considered an ant-like stone beetle (Coleoptera: Scydmaenidae), aside from this extra foreleg segment, it contained a combination of characters found in two extant subfamilies (Scydmaeninae and Mastiginae). These characters, along with other rather unique features, were why it was placed in its own subfamily, the Hapsomelinae (Poinar and Fig. 7. Holotype (designated as worker no. 1) of Sphecomyrma freyi in New Jersey amber (photo by the late Frank Carpenter: specimen later accidently destroyed at the AMNH). Fig. 8. Fore and hind wing typical of a “modern” aphid in Dominican amber. Note smaller membranous hind wing. American Entomologist • Volume 58, Number 4 consisting of large forewings and smaller hind wings (Fig. 8). However, some 100 million years ago, there were aphid clades that only had functional forewings, while the hind wings were reduced to stubs or halteres (Fig. 9). The hind wings of the Cretaceous Burmese amber Parvaverrucosa annulata Poinar and Brown (2005) resemble the hind pair (metathoracic) of wings of male coccids. Male coccids have functional forewings, but the hind wings are reduced to rod-like structures with terminal hooks called hamulohalters. Aphids and coccids belong to separate families in the suborder Sternorrhyncha of the order Hemiptera. The tips of Parvaverrucosa annulata’s stubby hind wings bear small hooks, thus making the reduced hind wings almost identical to those of coccids (Poinar and Brown 2006a) (Fig. 10). Although it has been placed with the aphids, the annulated antennae of P. annulata are a unique character that separates it from all fossil and extant aphids. The unicorn fly Cascoplecia insolitis in 97-110 mya Burmese amber. Bibionid flies (Diptera: Bibionidae), commonly known as march flies or love bugs, are noted for suddenly appearing in large mating swarms in the spring. The adults normally don’t feed and the larvae mostly scavenge on plant debris. The Early Cretaceous Burmese amber Cascoplecia insolitis (Diptera: Cascopleciidae) is a desiomorph between the Bibionidae and the extinct Mesozoic family Protopleciidae. Analyses of the fossil were based on wing venation Fig. 9. The two-winged aphid Parvaverrucosa annulata in Dominican amber. Arrows show reduced haltere-like hind wings. Fig. 10. Detail of hooks (arrow) on the end of the haltere of Parvaverrucosa annulata in Dominican amber. 217 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 where camouflage and a defensive posture were critical, such as an animal nest (Poinar and Heiss 2011). The primitive ant Sphecomyrma freyi in 90 mya New Jersey amber. One of the first amber desiomorphs reported was Sphecomyrma freyi Wilson, Carpenter, and Brown (1967), an Upper Cretaceous New Jersey amber fossil considered a “near-perfect link between certain nonsocial tiphiid wasps and the most primitive myrmecioid wasps” by possessing “a mosaic of wasp-like and antlike character states” (Wilson et al. 1967)(Fig. 7). The short, bidentate mandibles, antennal funiculi, and alitrunk (thorax and propodium) are more similar to characters of aculeate wasps, while the elongated scapes, single-segmented petiole, paired tibial spurs, and toothed tarsal claws are more like those of ants. The presence of metapleural glands, which are peculiar to most modern ants, is one of the main reasons Sphecomyrma was placed in the Formicidae. The aphid Parvaverrucosa annulata in 97-110 mya Burmese amber. Aphids or plant lice are a very successful group, as most gardeners would agree. Aphids have a complex life cycle, and during the summer, the large wingless parthenogenetic females produce large numbers of offspring that damage plants by removing sap and transmitting plant viruses. At some point in their development, usually at the end of the growing season, winged sexual stages appear. All extant and Tertiary winged aphids possess two pairs of wings tarsi with divaricate claws, and leveled and subequal abdominal ventrites are characters of the Caridae. The 11-segmented, loosely clubbed antennae, well-developed eyes, and lateral carina of the pronotum are characters of the Late Jurassic family Eobelidae. This above combination of characters, as well as the loose antennal club, vertical mandibles, and deep insertion of the unguitractor plate into the pretarsus (resulting in a lobed fifth segment), justified placing this weevil in its own family (Poinar and Brown 2009). Using functional morphology to determine the biology of Anchineus, the bi-lobed tarsi and robust claws indicate an arboreal existence, while the long rostrum and vertical mandibles suggest it fed on and oviposited in stems, buds, or soft fruits (Fig. 12). The primitive crane fly Dacochile microsoma in 97-110 mya Burmese amber. The curious Dacochile microsoma possesses character states found in both the Psychodidae (Bruchomyiinae) and the Tanyderidae (Poinar and Brown 2006b). While the proportion of the palpal segments, immaculate wings lacking an anal lobe, and a short Rs vein align Dacochile with the Bruchomyiinae, the presence of an m-m cross vein, serrated mandibles, setae between the ommatidia, and two segmented cerci and the absence of flagellar ascoids align it with the Tanyderidae (Fig. 13). A row of heavily sclerotized, thorn-shaped processes (spines/ spurs) on the inside of the hind tibiae and first two tarsal segments of Dacochile suggest it may have hung from vegetation by its hind legs to wait for passing prey. This behavior of hanging from plants has been noted in the New Zealand Tanyderus forcipatus. The serrated mandibles of Dacochile (Fig.14) are another tany- Fig. 11. Head of the unicorn fly Cascoplecia insolitis in Burmese amber. Fig. 13. The primitive crane fly Dacochile microsoma in Burmese amber. Arrow shows m-m crossvein, a key character of the Tanyderidae. Fig. 12. The primitive weevil Anchineus dolichobothris in Burmese amber. 218 Fig. 14. Serrated mandibles (arrow) of Dacochile microsoma in Burmese amber. American Entomologist • Winter 2012 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 comparisons because most Mesozoic Diptera are known only from isolated wings. Wing venation within the Bibionomorpha is fairly conserved, so using venational patterns to assign fossils to extant or extinct clades (pterotaxonomy) is generally dependable. The wing venation of Cascoplecia insolitis fell between the extant genus Plecia (Bibionidae) and the Mesozoic genus Protoplecia (Protopleciidae). Since Cascoplecia was complete, body characters could also be used in determining its systematic placement. The strange S-shaped antennomeres, elongate palps, and head protuberance bearing ocelli placed Cascoplecia apart from any extant or extinct member of the Bibionomorpha (Poinar 2009) (Fig. 11). The weevil Anchineus dolichobothris in 97-110 mya Burmese amber. A small but significant weevil in Burmese amber, Anchineus dolichobothris, had a combination of character traits of three extant and one extinct weevil families. The 7-jointed funicle and bifid claws resemble members of the Apionidae, while the geniculate antennae with loosely three-segmented antennal clubs are characters of the Nanophyidae. The antennae inserted on the sides of the rostrum, Fig. 15. An intertidal shore bug Saldula laticollis along the Oregon coast. 16. The shore bug Palaeoleptus burmanicus in Burmese amber. American Entomologist • Volume 58, Number 4 17. The mantidfly, Doratomantispa burmanica in Burmese amber. Note the white, spherical bodies in the abdomen, which indicates a rickettsial infection. 18. Trichosors (arrows) on the wing margin of Doratomantispa burmanica in Burmese amber. While such infections are not known in extant mantispids or thorny lacewings, they have been reported in chrysopid neuropterans. Female heterostigmatid mites (Khaustov and Poinar 2011) adjacent to the wings and body of Doratomantispa presented another interesting aspect of this fossil. Phoretic mites have not been reported on any extant Neuroptera. The primitive bee Melittosphex burmensis in 97-110 mya Burmese amber. Melittosphex burmensis (Apoidea: Melittosphecidae) represents an extinct lineage of bees with some wasp-like characters. It provides important insights into the transition from predatory crabronid wasps to pollen-collecting bees. Its mixture of bee- and wasp-like features makes it an ideal desiomorph and explains why it was placed in a new sister family to the modern bees. In general shape, Melittosphex resembles a small andrenid or halictid bee, and like modern bees, it is covered with branched hairs over most of its body (Fig. 19). While its wing venation is more beelike than wasp-like and the branched hairs (Fig. 20) separate it from all apoid wasps, Melittosphex possesses two mid-tibial spurs and a slender hind basitarsus, which are basic features of apoid wasps. Autapomorphic features are large, clearly tridentate mandibles, slender hind legs, and prominent tubercles on the propodeum (Danforth 219 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 derid character and give some indication of the feeding habits of the fossil. A piercing-sucking habit and preying on invertebrates or vertebrates may have provided its major means of obtaining nourishment. The shore bug Palaeoleptus burmanicus in 97-110 mya Burmese amber. Shore bugs (Hemiptera: Saldidae) represent a family of small insects that favor damp or wet habitats, such as streams, lakebeds, ocean beaches, or bays. An intertidal shore bug that is common along the Pacific coast is Saldula laticollis (Fig. 15), a predator capable of surviving short periods of immersion in seawater. A shore bug in Early Cretaceous Burmese amber (Fig. 16) had some characters found in modern saldids as well as in the related family, Leptopodidae. While the body shape, rostrum length and antennal structure of Palaeoleptus burmanicus are saldid features, the position of the ocelli, rostral spines, and profemur armature are leptopodid characters. Aside from these shared characters, Palaeoleptus has such a unique forewing venation that it was placed in its own family (Poinar and Buckley 2009). The asymmetrical genital plate directed to the left side of the body suggests that Palaeoleptus practiced side-by-side mating, similar to the condition in saldids. The mantidfly Doratomantispa burmanica in 97-110 mya Burmese amber. Mantidflies (Mantispidae: Neuroptera) are strange neuropterans with large raptorial forelegs similar in form and function to those of the unrelated praying mantises. A mantispid, Doratomantispa burmanica, in Early Cretaceous Burmese amber (Fig. 17) not only possessed characters found in all four extant subfamilies (Calomantispinae, Symphrasinae, Drepanicinae, and Mantispinae) of mantidflies, but also exhibited a wing character found in the related thorny lacewings (Rhachiberothidae) (Poinar and Buckley 2010). The character shared with the thorny lacewings was the arrangement of short supporting rods (trichosors) along the wing margin. Most mantispids lack trichosors, or if they are present, they are usually reduced and only occur along the wing tips. Thorny lacewings not only have numerous trichosors along most of the wing margin, but there is only a single trichosor between the veinlets. This is the exact condition found in Doratomantispa (Fig. 18). An ecological feature of Doratomantispa that could function as a desiomorphic character is the presence of spherical-subspherical bodies of a possible rickettsial pathogen in its abdomen (Fig.17). 20. Branched hairs on body of Melittosphex burmensis. and Poinar 2011). The small size of Melittosphex correlates with the small flowers that have been recovered from Burmese amber. These Early Cretaceous flowers, which represent diverse plant families, only range from 1-6 mm in diameter (Chambers et al. 2010) (Fig. 21). The eugregarine Primigregarina burmanica in 97-110 mya Burmese amber. Gregarines are extremely diverse protozoan parasites that infect invertebrates, especially annelids and arthropods. Many parasitize insects, especially cockroaches, grasshoppers, and other members of the Dictyoptera. The first fossil gregarine, Primigregarina burmanica, occurred adjacent to a cockroach in Burmese amber that had been attacked by a predator and had its midgut ripped open, thus exposing the contents (Poinar 2010). This action revealed the epimerate and gametocyst stages of the gregarine and showed that cockroaches harbored these parasites in the Early Cretaceous (Fig. 22). The fossil shares characters with both of the extant families Monoductidae and Gregarinidae. While the single sporoduct on the gametocyst (Fig. 23) aligns the fossil with members of the Monoductidae, the large spherical gametocyst and non-pronged epimerate are characters of the Gregarinidae. The short inverted neck of the epimerite is a unique character of Primigregarina. Evidence of spore expulsion from the gametocyst in the gut of the cockroach host is another unusual character. Normally, gametocysts dehisce after being passed from the body of the host and the time between capture and becoming embedded in the resin would seem to be too short for the formation of a spore duct with spore release. 220 22. Part of a predated cockroach showing a gametocyst of the eugregarine protozoan Primigregarina burmanica (arrow) in Burmese amber. 23. Sporoduct (arrow) emerging from the gametocyst of Primigregarina burmanica in Burmese amber. American Entomologist • Winter 2012 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 19. The primitive bee Melittosphex burmensis in Burmese amber. 21. Flower of Tropidogyne pikei in Burmese amber. This is one of the small flowers that females of Melittosphex could have visited. The flower diameter is slightly over 5 mm. Desiomorphs at the Ordinal Level 26. The hard tick Cornupalpatum burmanicum in Burmese amber. 27. Anterior end of Cornupalpatum burmanicum in Burmese amber. Arrow shows claws on tips of palps. related to mating behavior. They were probably used to assist the tick in attaching to a host. Summary 24. The Dominican amber onychophoran Tertiapatus dominicanus. 25. The Baltic amber onychophoran Succinipatopsis balticus. American Entomologist • Volume 58, Number 4 While these desiomorphs reveal the long geological past of certain invertebrate clades, it is difficult to determine how many are stem groups and represent transitional fossils and how many are separate, unique lineages that share characters as the result of convergence. As aforementioned, morphological characters can be misleading in determining whether a particular organism is a true transitional fossil. Darwin cited the absence of “innumerable transitional forms” as a major problem with his theory of phyletic evolution. While the number of fossils has increased greatly since Darwin’s time, his comment on the “extreme imperfection of the fossil record” still applies in large part today. One of the difficulties in deciding if desiomorphs represent actual transition fossils is our lack of knowledge of the biology of the fossils, especially if there are no characters that indicate functional morphology. If closely related to an extant clade, one can deduce the biology of a fossil by using the principal of behavior fixity, which infers that the behavior, ecology, and climatic preferences of fossil organisms will be similar to those of their present- day descendants at the generic, and in some cases, the family level (Boucot and Poinar 2010). 221 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 Velvet worms in 20-30 mya Dominican amber and 40-50 mya Baltic amber. Velvet worms of the Subphylum Onychophora are enigmatic invertebrates with a unique accumulation of characters. At first appearance, they resemble annelids with legs, but their affinity (which is still somewhat of a mystery) seems to lie with the arthropods. As predators, their singular method of prey capture employs the use of a slimy secretion propelled toward the victim from a pair of head papillae. The slime ensnares the prey, which is then consumed. All extant velvet worms have an apical non-retractable foot portion at the base of their legs. This foot portion bears a terminal pair of claws and well-defined pads with associated papillae. However, the foot portion is absent on a pair of velvet worms found in Baltic and Dominican amber (Poinar 2000). Even though the Dominican amber Tertiapatus dominicanus (Fig. 24) and Baltic amber Succinipatopsis balticus (Fig. 25) are terrestrial, possess a ventral mouth, and have numerous simple legs and slime-secreting head papillae, they both lack clawed footpads. Simple legs lacking footpads with claws are also characteristic of Paleozoic onychophorans. The hard tick Cornupalpatum burmanicum in 97-110 mya Burmese amber. Ticks are rare as fossils, but both hard (Ixodidae) and soft (Argasidae) ticks have been found in amber deposits. The oldest records of hard ticks are in Early Cretaceous Burmese amber, and one of these has features of a desiomorph (Poinar and Brown 2003). While resembling modern ticks in most characters (Fig. 26), the first instar larva of Cornupalpatum burmanicum possesses palpal claws (Fig. 27). Such armature is lacking on all extinct and extant ticks, but terminal or subterminal palpal claws occur in members of the mite groups Opilioacarida, Holothyrida, and Mesostigmata. The function of the palpal claws on Cornupalpatum is unknown, but because the fossil is immature, it is doubtful the claws would be Desiomorphs are among the most intriguing topics in paleontology, and questions about their lifestyle, disappearance, and classification will occupy us for years to come. They show the inadequacy of our present system of classification in dealing with ancient invertebrates and give us a time zone for the appearance and extinction of ancient lineages. Acknowledgements The author thanks Art Boucot and Roberta Poinar for comments on earlier versions of this work. References Cited Dr. George Poinar, who received his higher education from Cornell University, has been studying life forms in amber since 1975, shortly after he joined the Entomology Department at the University of California, Berkeley. Travels to the Baltic area, Dominican Republic, Mexico, Alberta, Canada and New Zealand gave him an opportunity to collect amber firsthand from various depositional sites. His early retirement at Berkeley offered an opportunity to transfer to Oregon State University where he continues to study amber inclusions from around the world. Call for Contributions Annals of the Entomological Society of America announces a new emphasis on integrative and comparative insect biology. Authors are encouraged to submit manuscripts that contribute to the development of trans-disciplinary theory that would be of interest to entomologists from different areas of specialization. We also solicit collections of articles from groups of scientists who are able to present up-to-date review and current research in a field of common interest (for an example, please see the Special Feature in Vol. 104 number 6, 2011, pp. 1029–1148). Ideas for group submissions may be submitted directly to the Editorin-Chief, Lawrence Hurd: hurdl@wlu.edu. All manuscripts are subject to peer review. Accepted papers for Special Features are eligible for waiver of page charges and free open access online. 222 American Entomologist • Winter 2012 Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 Boucot, A. and G. O. Poinar, Jr. 2011. Fossil Behavior Compendium, CRC Press, Boca Raton. Chambers, K. L., G. Poinar, Jr. and R. Buckley. 2010. Tropidogyne, a new genus of Early Cretaceous Eudicots (Angiospermae) from Burmese amber. Novon, 20:23–29. Danforth, B. D. and G. O. Poinar, Jr. 2011. Morphology, Classification, and Antiquity of Melittosphex burmensis (Apoiodea: Melittosphecidae) and implications for early bee evolution. J. Paleontol. 85: 882-891. Hörnschemeyer, T., S. Wedmann and G. Poinar. 2010. How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insects: Coleoptera). Zoological Journal of the Linnean Society 158: 300-311. Khaustov, A. A. and G. O. Poinar, Jr. 2011. Protoresinacarus brevipedis gen. n., sp. n. from Early Cretaceous Burmese amber: the first fossil record of mites of the Family Resinacaridae (Acari: Heterostigmata: Pyemotoidea). Historical Biol. 23: 219-222. Poinar, Jr., G. O. 1998. Paleoeuglossa melissiflora gen. n., sp. n. (Euglossinae: Apidae), fossil orchid bees in Dominican amber. J. Kansas Entomol. Soc. 71: 29-34. Poinar, Jr., G. O. 2000. Fossil onychophorans from Dominican and Baltic amber: Tertiapatus dominicanus n. g., n. sp. (Tertiapatidae n. fam) and Succinipatopsis balticus n.g., n. sp. (Succinipatopsidae n. fam.) with a proposed classification of the subphylum Onychophora. Invertebr. Biol. 119: 104-109. Poinar, Jr., G. 2009. Cascoplecia insolitis (Diptera: Cascopleciidae), a new family, genus, and species of flower-visiting unicorn fly (Bibionomorpha) in Early Cretaceous Burmese amber. Cretaceous Res. 31: 71-76. Poinar, Jr., G. O. 2010. Primigregarina burmanica n. gen., n. sp., an Early Cretaceous gregarine (Apicomplexa: Eugregarinorida) parasite of a cockroach (Insecta: Blattodea). pp. 54-56 in: Fossil Behavior Compendium, Boucot, A.J. and G. O. Poinar, Jr. (Eds.), CRC Press, Boca Raton. Poinar, Jr., G. O. 2011. Vetufebrus ovatus n. gen., n. sp. (Harmospororida: Plasmodiidae) vectored by a streblid bat fly (Diptera: Streblidae) in Dominican amber. Parasites & Vectors 2011 4: 229. Poinar, Jr., G. and A. Brown. 2003. A new genus of hard ticks in Cretaceous Burmese amber (Acari: Ixodida: Ixodidae). Systemat. Parasitol. 54: 199-205. Poinar, Jr., G. and A. E. Brown. 2004. A new subfamily of Cretaceous antlike stone beetles (Coleoptera: Scydmaenidae: Hapsomelinae) with an extra leg segment. Proc. Entomol. Soc. Washington 106: 789-796. Poinar, Jr., G. O. and A. E. Brown. 2005. New Aphidoidea (Hemiptera: Sternorrhyncha) in Burmese amber. Proc. Entomol. Soc. Washington 107: 835-845. Poinar, Jr., G. and A. E. Brown. 2006a. Remarks on Parvaverrucosa annulata (= Verrucosa annulata Poinar & Brown 2005) (Hemiptera: Sternorrhyncha: Aphidoidea) Proc. Entomol. Soc.Washington 108: 734-735. Poinar, Jr., G. O. and A. E. Brown. 2006b. The enigmatic Dacochile microsoma Poinar & Brown: Tanyderidae or Bruchomyiinae? Zootaxa 1162: 19-31. Poinar, Jr., G. O. and A. E. Brown. 2009. Anchineus dolichobothris, a new genus and species of Early Cretaceous weevils (Curculionoidea: Coleoptera) in Burmese amber. Proc. Entomol. Soc. Washington 111: 263-270. Poinar, G. & A. Brown. 2012. The first fossil streblid bat fly, Enischomyia stegosoma n. g., n. sp. (Diptera: Hippoboscoidea: Streblidae). Systematic Parasitol. 81: 79-86. Poinar, Jr., G. O. and R. Buckley. 2009. Palaeoleptus burmanicus n.g., n.sp., an Early Cretaceous shore bug (Hemiptera: Palaeoleptidae n. fam.) in Burmese amber. Cretaceous Res. 30: 1000-1004. Poinar, Jr., G. O. and R. Buckley. 2010. Doratomantispa burmanica n. gen.,n. sp. (Neuroptera: Mantispidae), a new genus of mantidflies in Burmese amber. Historical Biology 23:169-176. Poinar, Jr., G. and E. Heiss. 2011. New Termitaphididae and Aradidae (Hemiptera) in Mexican and Dominican amber. Palaeodiversity 4: 41-52. Poinar, Jr., G.O. and R. Milki. 2001. Lebanese amber: the oldest insect ecosystem in fossilized resin. Oregon State University Press, Corvallis. Simmons, N. B., K. L. Seymour, J. Habersetzer and G. F. Gunnell. 2008. Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation. Nature 451: 818-821. Wilson, E. O., F. M. Carpenter and W. L. Brown, Jr. 1967. The first Mesozoic ants, with a description of a new subfamily. Psyche 74: 1-19. Just Published and Available Now! Handbook of Turfgrass Insects Handbook of Turfgrass Insects SECOND SECOND EDITION EDITION Second Edition EDITED BY Rick L. Brandenburg Callie P. Freeman This highly-anticipated second edition of the Handbook of Turfgrass Insects contains the most current information covering all areas of turfgrass insect management. The handbook provides a comprehensive, yet easy-to-use guide for students, practitioners, extension staff, Master Gardeners, teachers, and others. American Entomologist • Volume 58, Number 4 Order by February 11, 2013 from APS at a special introductory price: $62.96 ESA Member Price $69.95 Nonmember Price $79.95 Regular Price Order at: www.apsnet.org/esaturf MOLOGIC NTO SOCIET Y OF A AL RICA ME 144 pages, softcover, 2012 ISBN 978-0-9776209-4-4 ENTOMOLOGICAL SOCIETY OF AMERICA E The past ten years have seen the spread of many longtime pests and the occurrence of several new ones, so proper identification and knowledge of the most current pest biology and ecology is critical. The book covers all major pests of warm- and cool-season turfgrass in the United States, with each section written by one or more experts on each pest. Numerous color photos of various insect stages and damage are included as well as illustrations of life stages in their actual size, life cycle charts, and distribution maps. There are important chapters on the principles of integrated turfgrass pest management, microbial control, use of insecticides, insecticide resistance management, and beneficial and innocuous invertebrates in turf. A glossary, index, and sources of local information are also included. Downloaded from https://academic.oup.com/ae/article-abstract/58/4/214/6617 by guest on 20 May 2020 Edited by Rick L. Brandenburg and Callie P. Freeman Published by the Entomological Society of America in cooperation with the American Phytopathological Society. 223