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Diptera) in Tertiary: Eocene Ambers

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Diptera) in Tertiary: Eocene Ambers 1 Review and phylogenetic placement of Asilidae (Diptera) in Tertiary: Eocene ambers Torsten Dikow 0000-0003-4816-2909 @TDikow #asiloidflies Smithsonian PDF doi: 10.6084/m9.figshare.1243382 National Museum of Natural History Tertiary: Eocene amber – Baltic Amber 2 ◊ well-known amber from the Bay of Gdansk (Poland) of the southern Baltic Sea ◊ 45 – 50 Million years old (myo) ◊ very rich in insect inclusions particularly Diptera › Hoffeins & Hoffeins 2003. Studia dipterologica 10(2): 381 – 392 ◊ Baltic, Bitterfeld, and Rovno ambers most probably same fauna › but see Penney 2010. Biodiversity of fossils in Amber from the major world deposits. Siri Scientific Press, Manchester ◊ Willi Hennig studied Diptera inclusions extensively › Acalyptratae, Acroceridae, Bombyliidae, Lower Brachycera, Therevidae › recorded several acalyptrate families believed to be Sweden Southern Hemisphere endemics Lithuania Russia Belarus Germany Poland Ukraine SimpleMappr link Asilidae – “assassin flies” 3 ◊ adult flies and larvae predatory ◊ size = 5 – 60 mm ◊ > 7,500 extant species in 541 genera ◊ 58 fossil species in 32 genera catalog URL ◊ oldest definitive assassin fly – †Araripogon axelrodi › Cretaceous Crato Formation of Brazil, ≈ 112 myo †Araripogon axelrodi / › Grimaldi 1990 open-access link ◊ rare in any amber due to life history › 2x Burmese Amber (100 myo) – †Burmapogon bruckschi › 1x Raritan Amber (92 myo) – †Cretagaster raritanensis › Dikow & Grimaldi 2014 open-access link › 18x Dominican Amber (20 – 25 myo) – Dikow & Fisher in prep. †Burmapogon bruckschi ?– Morphbank #832135 Asilidae in Baltic Amber – history of discovery 4 ◊ Loew 1850 BHL link › †Asilus angustifrons, †Asilus trichurus, †Dasypogon (Holopogon) pilipes › species nomina nuda ◊ Meunier 1899 BHL link, 1908 BHL link › †Asilus trichurus Meunier, 1899 › †Asilus (Lophonotus) klebsi Meunier, 1908 ◊ Hennig 1967 › unable to locate any of the above type specimens ◊ Schumann 1984 › Bitterfeld Amber › †Protoloewinella keilbachi ◊ Geller-Grimm 1998 › recorded †Protoloewinella keilbachi from Baltic Amber material and methods 5 ◊ 25 amber pieces with assassin-fly inclusions ◊ primarily from Coll. Hoffeins (Hamburg, Germany) and AMNH › amber beautifully enhanced through cutting and embedding in “modern” resin ◊ photography › Zeiss SteREO Discovery.V12, PlanApo S 0.63x or 1x lens (5x to 10x magnification) › mixed LED lighting: Bright Field, Dark Field, and Transillumination › Olympus micro Four-Thirds mirrorless E-PL 5, RAW image › manual “slicing” – rendered with HeliconFocus ◊ morphological matrix TreeBase PURL from Dikow 2009 open-access link › TNT 1.1 (February 2014) Goloboff et al. 2008 doi link › exhaustive search of tree space (MacPro 3.5 GHz 6-core Intel Xeon E5, 64 GB RAM) › Bremer support Bremer 1988 doi link › character analysis Mesquite (3.01) and WinClada/Asado (1.08) †Protoloewinella keilbachi 6 ◊ Laphriinae: Atomosiini ◊ now known from Baltic, Bitterfeld, and Rovno ambers ◊ most abundantly preserved assassin fly(10 pieces with 13 enclosed specimens) †Protoloewinella keilbachi / coll. Hoffeins 938-3 left †Protoloewinella keilbachi / coll. Hoffeins 938-3 right †Protoloewinella keilbachi 7 ◊ Atomosiini › tree trunks or vegetation as perching sites › relatively small and “weak” assassin flies Atomosia puella © M. Thomas 3x †Protoloewinella keilbachi coll. Hoffeins 1038-2 dorsal newly discovered taxa 8 ◊ Atomosiini: morphologically similar, but distinct from †Protoloewinella keilbachi › antennal stylus longer, abdominal tergites less microrugose †Atomosiini ? gen. nov. sp. nov. / coll. Hoffeins 1392-1 dorsal †Atomosiini ? gen. nov. sp. nov. / coll. Hoffeins 1392-1 ventral †Atomosiini ? gen. nov. sp. nov. / coll. Hoffeins 1392-1 left †Atomosiini ? gen. nov. sp. nov. / coll. Hoffeins 1392-1 right newly discovered taxa 9 ◊ †Asilinae gen. nov. A sp. nov. ◊ †Asilinae gen. nov. B sp. nov. › 1 female specimen › 6 specimens (4 males, 2 females) › similarities with Astochia, Neoitamus, some Promachus › similarities with Neomochtherus †Asilinae gen. nov. B sp. nov. ? coll. Hoffeins 938-1 left †Asilinae gen. nov. A sp. nov./ †Asilinae gen. nov. A sp. nov./ coll. Hoffeins 1453-1 right coll. Hoffeins 1453-1 left †Asilinae gen. nov. B sp. nov. ? coll. Hoffeins 938-1 right newly discovered taxa 10 ◊ †Laphriinae gen. nov. sp. nov. ◊ †”Stenopogoninae” gen. nov. sp. nov. › 1 male specimen › 1 male and 1 female › similarities with Nusa †Laphriinae gen. nov. sp. nov. ? coll. Hoffeins 1305-1 dorsal †”Stenopogoninae” gen. nov. sp. nov. / coll. Hoffeins 632-2 dorsal †Laphriinae gen. nov. sp. nov. ? coll. Hoffeins 1305-1 ventral †”Stenopogoninae” gen. nov. sp. nov. / coll. Hoffeins 632-2 ventral newly discovered taxa 11 ◊ †Leptogastrinae gen. nov. sp. nov. › 1 male specimen › similarities with Leptogaster †Leptogastrinae gen. nov. sp. nov. ? coll. Hoffeins 1570 dorsal †Leptogastrinae gen. nov. sp. nov. ? coll. Hoffeins 1570 ventral phylogenetic placement 12 Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca Laphystia tolandi included fossil taxa % missing characters Hoplistomerus nobilis Trichardis testacea Laxenecera albicincta Nusa infumata Smeryngolaphria numitor Orthogonis scapularis Aphestia annulipes Atomosia puella (of 220 characters) Cerotainia albipilosa Laphriinae Goneccalypsis argenteoviridis Loewinella nigripes +Protoloewinella keilbachi +1392 Maira aenea Lampria clavipes Choerades bella †Asilinae A 43 Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris †Asilinae B 28 Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Ommatiinae Emphysomera conopsoides †Protoloewinella 36 Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii +1453-1 Asilus crabroniformis Asilus sericeus Asilidae Megaphorus pulchrus Promachus amastrus †Atomosiini 36 Apoclea algira Philodicus tenuipes Neolophonotus bimaculatus Pogonioefferia pogonias Proctacanthus philadelphicus Lochmorhynchus albicans Eichoichemus pyrrhomystax Myaptex brachyptera Asilinae † Leptogastrinae 48 Dasophrys crenulatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Neomochtherus pallipes +938 Afromochtherus mendax †”Stenopogoninae” 30 Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Tolmerus atricapillus Clephydroneura sundaica Clephydroneura sp. (Kepong) Heligmonevra laevis Zosteria rosevillensis Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Carebaricus rionegrensis Creolestes nigribarbis Prolepsis tristis Plesiomma sp. (Guanacaste) Tillobroma punctipennis Coleomyia setigera Ablautus coquilletti Oligopogon sp. (Cape Recife) Connomyia varipennis +632 Bathypogon nigrinus Bathypogon sp. (Badgingarra) Microstylum sp. (Karkloof) Daspletis stenoura Bathypogoninae Phellus olgae Obelophorus terebratus Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Phellinae Ancylorhynchus fulvicollis Scylaticus costalis Hypenetes stigmatias Lycostommyia albifacies Willistonina bilineata Trichoura sp. (Tierberg) Stenopogon sabaudus Stenopogon rufibarbis Ospriocerus aeacus Scleropogon subulatus Gonioscelis ventralis Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Trigonomiminae Holcocephala calva Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata +1570 Leptogaster arida Leptogastrinae Leptogaster cylindrica Lasiocnemus lugens Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctriinae Dioctria hyalipennis Dioctria rufipes Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Dasypogoninae Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogoninae Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris strict consensus topology of 348 most parsimonious trees of length 2833 Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Bremer ≥ 5 Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Brachyrhopalinae Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis phylogenetic placement 13 Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca Laphystia tolandi Hoplistomerus nobilis Laphriinae Trichardis testacea Laxenecera albicincta Nusa infumata Smeryngolaphria numitor Orthogonis scapularis Aphestia annulipes Atomosia puella Cerotainia albipilosa Goneccalypsis argenteoviridis Loewinella nigripes › relationships among taxa less-resolved +Protoloewinella keilbachi +1392 Maira aenea Lampria clavipes Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris › Asilinae resolution improved Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus
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