1

Review and phylogenetic placement of Assassin () in Tertiary: Eocene ambers

Torsten Dikow 0000-0003-4816-2909 @TDikow #asiloidflies Smithsonian PDF doi: 10.6084/m9.figshare.1466885 National Museum of Natural History Tertiary: Eocene amber – Baltic amber 2 ◊ well-known amber from the Bay of Gdansk (Poland) in the south-eastern Baltic Sea ◊ 45 – 50 Million years old (myo) ◊ very rich in inclusions particularly Diptera (midges, flies and relatives) › Hoffeins & Hoffeins 2003. Studia dipterologica 10(2): 381 – 392 ◊ Bitterfeld and Rovno amber › Penney 2010. Biodiversity of fossils in Amber from the major world deposits. Siri Scientific Press

Sweden

Lithuania Russia Belarus

Germany Poland

Ukraine

SimpleMappr link Tertiary: Eocene amber – Baltic amber 3 ◊ Hennig studied Diptera inclusions extensively › Acalyptratae, Acroceridae, Bombyliidae, Lower , Therevidae › recorded several acalyptrate families believed to be Southern Hemisphere endemics ◊ Cypselosomatidae (acalyptrate flies, Nerioidea) › Hennig 1965 BHL link › 1965 – 3 extant + 1 extinct › 2015 – 13 extant + 1 extinct species

distribution after Hennig 1965 current known distribution Assassin flies – Asilidae 4 ◊ adult flies and larvae predatory ◊ size = 5 – 60 mm ◊ > 7,500 species in 541 genera ◊ speciose world-wide › arid & semi-arid environments › tropical environments ◊ perching sites to look for flying prey duncani › primarily on ground, rocks, leaves, small vegetation, or twig tips › some species on living tree trunks › Leptogastrinae capture resting prey ◊ morphological phylogeny › Dikow 2009 open-access link

Ceraturgus fasciatus © M. Thomas Assassin flies – Asilidae 5

◊ 58 fossil species in 32 genera catalog URL ◊ oldest definitive assassin †A– raripogon axelrodi › Cretaceous Crato Formation of Brazil, ≈ 112 myo › Grimaldi 1990 open-access link ◊ rare in any amber due to life history › strong flies that might pull their legs out of sticky tree resin

◊ known diversity in amber †Araripogon axelrodi / amber age # pieces # sp. # sp. nov. Burmese 100 myo 2 1 Dikow & Grimaldi 2014 open-access link Raritan 92 myo 1 1 Dikow & Grimaldi 2014 Baltic 45–50 myo 28 4 6 Dominican 20–25 myo 18 2 4 Dikow & Fisher in prep.

†Burmapogon bruckschi ?– Morphbank #832135 Asilidae in Baltic amber – history of discovery 6

◊ Loew 1850 BHL link › †Asilus angustifrons + †Asilus trichurus (species names nomina nuda) › †Holopogon pilipes

◊ Meunier 1899 BHL link, 1908 BHL link › †Asilus trichurus Meunier, 1899 › †Asilus (Lophonotus) klebsi Meunier, 1908

◊ Hennig 1967 BHL link › unable to locate any of the above type specimens ◊ Schumann 1984 › Bitterfeld amber › †Protoloewinella keilbachi ◊ Geller-Grimm 1998 › recorded †Protoloewinella keilbachi from Baltic amber ◊ to date, 4 species from Baltic – includes 1 species also known from Bitterfeld amber material and methods 7 ◊ 27 amber pieces with assassin-fly inclusions › 25 Baltic + 2 Rovno amber ◊ 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 8 ◊ Laphriinae: Atomosiini ◊ previously known from Baltic and Bitterfeld ambers › newly recorded from Rovno amber (K-4739, Schmalhausen Institute of Zoology, Kiev) ◊ 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 9 ◊ 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 10 ◊ another unique Atomosiini species? › 3 specimens morphologically similar to †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 11 ◊ †Asilinae gen. nov. A sp. nov. ◊ †Asilinae gen. nov. B sp. nov. › 1 female specimen › 6 specimens (4 males, 2 females) › similarities to Astochia, Neoitamus, some Promachus › similarities to 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 12 ◊ †Leptogastrinae gen. nov. sp. nov. ◊ †”” gen. nov. sp. nov. › 1 male specimen › 1 male and 1 female specimens › similarities to Leptogaster

†Leptogastrinae gen. nov. sp. nov. ? coll. Hoffeins 1570 dorsal †”Stenopogoninae” gen. nov. sp. nov. / coll. Hoffeins 632-2 dorsal

†Leptogastrinae gen. nov. sp. nov. ? coll. Hoffeins 1570 ventral †”Stenopogoninae” gen. nov. sp. nov. / coll. Hoffeins 632-2 ventral newly discovered taxa 13 ◊ †Laphriinae gen. nov. sp. nov. › 1 male specimen › specimen covered by milky “fog”

†Laphriinae gen. nov. sp. nov. ? coll. Hoffeins 1305-1 dorsal

†Laphriinae gen. nov. sp. nov. ? coll. Hoffeins 1305-1 ventral newly discovered taxa 14 ◊ †Laphriinae gen. nov. sp. nov. › Zeiss Xradia micro-CT scan phylogenetic placement 15

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 tristis Plesiomma sp. (Guanacaste) Tillobroma punctipennis Coleomyia setigera coquilletti Oligopogon sp. (Cape Recife) Connomyia varipennis +632 Bathypogon nigrinus Bathypogon sp. (Badgingarra) 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) sabaudus Stenopogon rufibarbis 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 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 punctipennis phylogenetic placement 16

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 higher-level taxa less- +Protoloewinella keilbachi +1392 Maira aenea Lampria clavipes Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis resolved Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Ommatiinae Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus › 3 subfamily taxa non-monophyletic Lycomya germainii +1453-1 Asilus crabroniformis Asilus sericeus Asilidae Megaphorus pulchrus Promachus amastrus Asilinae Apoclea algira Philodicus tenuipes Neolophonotus bimaculatus › Stenopogoninae Pogonioefferia pogonias Proctacanthus philadelphicus Lochmorhynchus albicans Eichoichemus pyrrhomystax Myaptex brachyptera Dasophrys crenulatus Nomomyia murina Satanas gigas › Tillobromatinae Machimus chrysitis Machimus occidentalis Neomochtherus pallipes +938 Afromochtherus mendax Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Tolmerus atricapillus › Willistonininae 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) Bathypogoninae Daspletis stenoura Phellus olgae Obelophorus terebratus Acnephalum cylindricum Phellinae Sisyrnodytes sp. (Gamka) 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) Trigonomiminae Holcocephala abdominalis Holcocephala calva Acronyches maya Beameromyia bifida Euscelidia pulchra Leptogastrinae Tipulogaster glabrata +1570 Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Broticosia paramonovi Dicolonus simplex Echthodopa pubera Dioctriinae Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Dasypogoninae Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogoninae Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Brachyrhopalinae strict consensus topology of 348 most parsimonious trees of length 2833 Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis phylogenetic placement – †Protoloewinella 17

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 › all Atomosiini specimens represent sngle species +Protoloewinella keilbachi +1392 Maira aenea Lampria clavipes Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis › †Protoloewinella keilbachi Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Ommatiinae Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus › 13 amber pieces with 16 flies preserved Lycomya germainii +1453-1 Asilus crabroniformis Asilus sericeus Asilidae Megaphorus pulchrus Promachus amastrus Asilinae Apoclea algira Philodicus tenuipes Neolophonotus bimaculatus Pogonioefferia pogonias Proctacanthus philadelphicus Lochmorhynchus albicans Eichoichemus pyrrhomystax › sister-group Loewinella nigripes + Goneccalypsis Myaptex brachyptera Dasophrys crenulatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Neomochtherus pallipes +938 argenteoviridis Afromochtherus mendax 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) Bathypogoninae Daspletis stenoura Phellus olgae Obelophorus terebratus Acnephalum cylindricum Phellinae Sisyrnodytes sp. (Gamka) 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) Trigonomiminae Holcocephala abdominalis Holcocephala calva Acronyches maya Beameromyia bifida Euscelidia pulchra Leptogastrinae Tipulogaster glabrata +1570 Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Broticosia paramonovi Dicolonus simplex Echthodopa pubera Dioctriinae Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Dasypogoninae Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogoninae Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Brachyrhopalinae strict consensus topology of 348 most parsimonious trees of length 2833 Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis phylogenetic placement – †Asilinae 18

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 › Asilinae resolution improved +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 › in 2009 hypothesis Asilinae is unresolved Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Ommatiinae Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii +1453-1 Asilus crabroniformis Asilidae Asilus sericeus › †Asilinae A in clade with Promachus Megaphorus pulchrus Promachus amastrus Asilinae Apoclea algira Philodicus tenuipes Neolophonotus bimaculatus Pogonioefferia pogonias Proctacanthus philadelphicus Lochmorhynchus albicans Eichoichemus pyrrhomystax › addition of extant Astochia needed Myaptex brachyptera Dasophrys crenulatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Neomochtherus pallipes +938 Afromochtherus mendax Dysmachus trigonus Philonicus albiceps Congomochtherus inachus › †Asilinae B in clade with Neomochtherus 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) Bathypogoninae Daspletis stenoura Phellus olgae Obelophorus terebratus Acnephalum cylindricum Phellinae Sisyrnodytes sp. (Gamka) 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) Trigonomiminae Holcocephala abdominalis Holcocephala calva Acronyches maya Beameromyia bifida Euscelidia pulchra Leptogastrinae Tipulogaster glabrata +1570 Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Broticosia paramonovi Dicolonus simplex Echthodopa pubera Dioctriinae Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Dasypogoninae Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogoninae Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Brachyrhopalinae strict consensus topology of 348 most parsimonious trees of length 2833 Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis phylogenetic placement – †Leptogastrinae 19

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 › deeply placed within Leptogastrinae +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 › sister-group to Tipulogaster glabrata Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Ommatiinae Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii +1453-1 Asilus crabroniformis Asilus sericeus Asilidae Megaphorus pulchrus Promachus amastrus Asilinae Apoclea algira Philodicus tenuipes Neolophonotus bimaculatus Pogonioefferia pogonias Proctacanthus philadelphicus Lochmorhynchus albicans Eichoichemus pyrrhomystax Myaptex brachyptera Dasophrys crenulatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Neomochtherus pallipes +938 Afromochtherus mendax 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) Bathypogoninae Daspletis stenoura Phellus olgae Obelophorus terebratus Acnephalum cylindricum Phellinae Sisyrnodytes sp. (Gamka) 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) Trigonomiminae Holcocephala abdominalis Holcocephala calva Acronyches maya Beameromyia bifida Euscelidia pulchra Leptogastrinae Tipulogaster glabrata +1570 Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Broticosia paramonovi Dicolonus simplex Echthodopa pubera Dioctriinae Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Dasypogoninae Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogoninae Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Brachyrhopalinae strict consensus topology of 348 most parsimonious trees of length 2833 Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis phylogenetic placement – †”Stenopogoninae” 20

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 › not easily placed +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 › Stenopogoninae non-monophyletic Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Ommatiinae Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii +1453-1 Asilus crabroniformis Asilidae Asilus sericeus › female and male specimen available Megaphorus pulchrus Promachus amastrus Asilinae Apoclea algira Philodicus tenuipes Neolophonotus bimaculatus Pogonioefferia pogonias Proctacanthus philadelphicus Lochmorhynchus albicans Eichoichemus pyrrhomystax Myaptex brachyptera Dasophrys crenulatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Neomochtherus pallipes +938 Afromochtherus mendax 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) Bathypogoninae Daspletis stenoura Phellus olgae Obelophorus terebratus Acnephalum cylindricum Phellinae Sisyrnodytes sp. (Gamka) 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) Trigonomiminae Holcocephala abdominalis Holcocephala calva Acronyches maya Beameromyia bifida Euscelidia pulchra Leptogastrinae Tipulogaster glabrata +1570 Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Broticosia paramonovi Dicolonus simplex Echthodopa pubera Dioctriinae Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Dasypogoninae Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogoninae Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Brachyrhopalinae strict consensus topology of 348 most parsimonious trees of length 2833 Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis summary 21 ◊ 6 new species for a total of 10 assassin-fly species in Baltic amber ◊ 5 new genera ◊ 2 specimens await further micro-CT data analysis for placement ◊ Baltic, Bitterfeld, and Rovno amber preserved same fauna – †Protoloewinella keilbachi ◊ phylogenetic study important to evaluate morphological variation ◊ phylogenetic placement successful except for 1 taxon acknowledgements 22 ◊ Fritz Geller-Grimm ◊ Christel and Hans-Werner Hoffeins and David Grimaldi for specimens ◊ Masako Terada at Zeiss Xradia ◊ U.S. National Science Foundation REVSYS DEB-0919333

presentation PDF doi: 10.6084/m9.figshare.1466885

Smithsonian National Museum of Natural History