Maximum Likelihood tree Maximum Likelihood Bootstrap tree Phylogeny of Clambus Chauliognathus Euspilotus Clambus Micromalthus Creophilus Tetraphalerus Dascillus Priacma Euspilotus Cupes Lucanus Prolixocupes Nosodendon Hydroscapha Nycteus Optioservus Derodontus Delevea 9 7 Laricobius Torridincola Cyphon Declinia Declinia David Maddison, Kojun Kanda - Oregon State University, Corvallis Cyphon 100 Elodes Elodes Pytho Nycteus Thymalus Kipling Will - University of California, Berkeley Derodontus 9 8 Tribolium Laricobius Crioceris Chauliognathus Hydroscapha Optioservus Lepicerus Wendy Moore - University of Arizona, Tucson Dascillus Sphaerius Lucanus Delevea Nosodendon 9 5 100 Torridincola Alex Wild - University of Texas, Austin Creophilus Micromalthus Thymalus 100 Tetraphalerus Kelly Miller - University of New Mexico Crioceris 100 Priacma Pytho Cupes Tribolium 100 Prolixocupes Peltodytes callosus Meru Peltodytes dispersus Peltodytes callosus Brychius 100 Peltodytes dispersus Haliplus eremicus 9 9 Brychius Apterliplus Apterliplus Haliplus "Argentina" 9 8 Haliplus "Argentina" Meru 100 Haliplus eremicus Notomicrus Notomicrus Noterus Noterus Tonerus 100 Suphisellus 4 Suphisellus 100 Tonerus Hydrocanthus 9 6 Hydrocanthus Mesonoterus 100 Mesonoterus Spanglerogyrus Gyrinus Amphizoa insolens 100 Aulonogyrus carinipennis Amphizoa lecontei 9 8 100 Aulonogyrus striatus OVERVIEW Lancetes Macrogyrus Coptotomus 100 Porrorhynchus Australphilus Dineutus CHIMERA 1 Dineutus CHIMERA 2 Neptosternus 100 9 8 Laccophilus Gyretes Philodytes Orectocheilus discifer Orectocheilus andamanicus Hydrotrupes 100 100 As part of the NSF-funded Tree of Life (BToL) Agabus Orectogyrus masculinus Batrachomatus 100 Orectogyrus posticalis Matus Aspidytes Carabdytes Hygrobia project, we are re-examining the phylogeny of major Rhantus calidus Amphizoa insolens 3 Rhantus CHIMERA 100 Amphizoa lecontei Hoperius Agabus Meladema Batrachomatus lineages of the beetle suborder Adephaga, using six genes. Neoscutopterus Hydrotrupes Dytiscus Lancetes Hyderodes Matus Eretes Coptotomus Our results confirm some results from previous studies Acilius Agaporomorphus Thermonectus 9 8 Copelatus Guignotites Dytiscus (i.e., trachypachines within Geadephaga). Two of the more Hydaticus 9 9 9 3 Hyderodes Methles Australphilus Sternhydrus 3 Neptosternus Cybister marginicollis 100 Laccophilus notable results are the continued presence of a Cybister tripunctatus 100 Philodytes Megadytes carcharias Sternhydrus Megadytes glaucus Cybister marginicollis Agaporomorphus 100 Cybister tripunctatus cicindeline-rhysodine-paussine-scaritine clade, even with Copelatus 100 Megadytes carcharias Celina Megadytes glaucus Hydrovatus Acilius additional gene sampling, and clear refutation of the Hyphydrus CHIMERA Eretes Hyphydrus elegans 9 1 Thermonectus 1 Neoporus 100 9 0 Guignotites Hydaticus Liodessus 100 proposal that rhysodines are related to Salcediina. Anodocheilus 9 7 Methles Bidessodes Rhantus CHIMERA Yola Carabdytes Allodessus 9 9 Rhantus calidus Uvarus 100 Hoperius Bidessus Meladema Clypeodytes 9 7 Neoscutopterus Stictonectes Amarodytes Coelambus Barretthydrus Hygrotus acaroides Celina Herophydrus Coelambus Hygrotus CHIMERA Deronectes METHODS Antiporus Hydrocolus Chostonectes Hydroporus Vatellus Hydrovatus Barretthydrus Hygrotus acaroides Necterosoma Necterosoma Deronectes Neoporus San lippodytes Peschetius Maximum likelihood searches of a concatenated matrix of six Amarodytes San lippodytes Peschetius 100 Stictonectes Hydrocolus Vatellus Hydroporus Antiporus gene fragments (18S, 28S, argK, wg, CAD2 and CAD4) and Spanglerogyrus 100 Chostonectes Herophydrus Gyrinus 100 Aulonogyrus carinipennis 9 8 Hygrotus CHIMERA Aulonogyrus striatus 1 Hyphydrus CHIMERA 291 terminal taxa were performed to infer Adephaga phylogeny 5 Macrogyrus 100 Hyphydrus elegans Porrorhynchus Allodessus Dineutus CHIMERA 1 Bidessus Dineutus CHIMERA 2 Clypeodytes using RAxML. Here we present the preferred tree with branch Uvarus Gyretes 100 Orectocheilus discifer Yola Orectocheilus andamanicus Liodessus lengths and the result of our bootstrap analysis (1000 reps). Anodocheilus Orectogyrus masculinus 100 Orectogyrus posticalis 100 Bidessodes Cychrus Cicindis Scaphinotus Eucamaragnathus Calosoma Gehringia Carabus Laccocenus 2 Ceroglossus Migadops Pamborus Nomius Notiophilus aquaticus Promecognathus Notiophilus ru pes Psydrus Opisthius Apotomus 1158 Paropisthius 100 Apotomus ru thorax Pelophila Cymbionotum pictulum 100 Cymbionotum semelederi Nippononebria Loricera foveata Blethisa 100 Loricera pilicornis RESULTS Diacheila Omophron (Phrator) Elaphrus californicus 100 Omophron obliteratum Elaphrus clairvillei Siagona europea Systolosoma breve 100 Siagona jenissoni Higher-level relationships Systolosoma lateritium Akephorus Trachypachus slevini 100 Dyschirioides Trachypachus gibbsii Platidiolus 1 6 Diplous CHIMERA • Adephaga monophyletic Trachypachus holmbergi 100 Cicindis 100 Patrobus Loricera foveata Blethisa 2 Loricera pilicornis Diacheila • Geadephaga monophyletic Migadops 100 Elaphrus californicus Amarotypus 100 Elaphrus clairvillei Monolobus Amarotypus Stichonotus Monolobus Antarctonomus 100 Stichonotus Antarctonomus Migadopidius 9 8 Nomius 100 Migadopidius Psydrus Relationships within Hydradephaga Mecodema 100 Oregus 100 3 Axonya 6 100 • Dysticidae monophyletic, in clade with Amphizoidae, Creobius 100 Promecoderus Cychrus Zacotus 100 Scaphinotus Broscosoma Calosoma Hygrobiidae, and Aspidytidae 100 Miscodera 9 9 Carabus Eucamaragnathus 100 Ceroglossus Laccocenus 100 Pamborus 4 Promecognathus • Meru sister to Noteridae but weakly supported Omophron (Phrator) Mouhoutia Omophron obliteratum Platidiolus 100 5 Diplous CHIMERA 8 1788 • Hydradephaga excluding Gyrinidae monophyletic in this Patrobus 100 Scaritinae 1789 Trechodes Notiophilus aquaticus Paratrechodes 100 Notiophilus ru pes analysis, but the placement of Gyrinidae is unclear Trechimorphus Pelophila Trechus 100 Nebria Typhlocharis 9 4 100 Nippononebria Geocharidius Opisthius Serranillus 100 Paropisthius Asaphidion Amblycheila Bembidion Mantichora Relationships within Geadephaga Sloaneana 100 Omus Merizodus Ctenostoma Oopterus 100 Megacephala 6 Diplochaetus 100 Cicindela • Trachypachines are placed as an early branching group of Sirdenus 100 Oxycheila Pericompsis sp. Elaphropus 100 Clinidium baldu Tachys 9 9 Clinidium calcaratum Geadephaga but exact placement varies depending upon the Erwiniana 100 100 Dhysores Mioptachys Akephorus 2 100 Grouvellina Dyschirioides 9 5 Leoglymmius analysis. In analyses where nuclear protein coding genes or ArgK 7 Salcedia 1785 Clivinini 1785 8 Aspidoglossa Salcedia are removed they are placed as sister to rest of Geadephaga. 100 Schizogenius 8 Semiardistomis Camptodontus Camptodontus 100 Clivina However, this placement not strongly supported. Apotomus 1158 Mecodema Apotomus ru thorax 100 Oregus Siagona europea Axonya 7 Siagona jenissoni 9 3 Creobius • Rhysodines are implicated in the CRPS quartet (see 9 below) Promecoderus Gehringia 100 100 Cymbionotum pictulum Zacotus Broscosoma Cymbionotum semelederi 100 and there is no support for a relationships to Salcedia, which is Mantichora 9 8 Miscodera Amblycheila Metrius contractus Omus 100 Metrius explodens Cicindela Mystropomus regularis confidently placed as a clivinine Oxycheila 100 Mystropomus subcostatus Ctenostoma 100 Microzaena Megacephala Pseudozaena 8 Clinidium sp. 9 1 100 Sphaerostylus • Scaritinae is not monophyletic Clinidium baldu Goniotropis Clinidium calcaratum 100 Platycerozaena Dhysores 100 100 Pachyteles 9 9 7 Omoglymmius 100 Physea • CRPS quartet (Cicindelinae, Rhysodinae, Paussinae, and Grouvellina Arthropterus Leoglymmius Homopterus Paussus Carenum 100 Mouhoutia Carabidomemnus Scaritini) is found but weakly supported in this analysis, and is Pasimachus 100 Cerapterus 8 Scaritinae 1788 Paratrechodes Scarites Trechimorphus not found in all analyses Scaritinae 1789 100 Trechodes Metrius contractus Trechus Metrius explodens Asaphidion 10 Mystropomus regularis 100 Bembidion • The relationship of (Moriomorphini (Brachininae, Harpalinae)) Diplochaetus Mystropomus subcostatus 9 7 Arthropterus 100 Sirdenus Carabidomemnus Geocharidius Cerapterus Serranillus is well supported in this ML analysis, but only weakly supported Homopterus 100 Typhlocharis Paussus 9 9 Sloaneana Merizodus Pseudozaena 100 in other analyses Sphaerostylus 100 Oopterus Microzaena Pericompsis Goniotropis Elaphropus 11 Tachys Platycerozaena 100 9 9 • Brachinini monophyly and its sister-group relationship to Pachyteles Erwiniana Physea 100 Mioptachys Meonis Meonis Melisodera Melisodera Harpalinae are strongly supported. Sitaphe 100 100 Sitaphe Amblytelus 100 Amblytelus Mecyclothorax 12 Mecyclothorax 9 5 • Harpalinae is monophyletic with strong support Tropopterus 9 7 Tropopterus Crepidogaster ambreana Mastax Crepidogaster atrata Crepidogaster ambreana Mastax 100 Crepidogaster atrata Pheropsophus 0528 Pheropsophus 0528 Pheropsophus aequinoctialis 100 Pheropsophus aequinoctialis Brachinus hirsutus 10 Brachinus hirsutus 9 9 Aptinomimus Aptinomimus 100 100 Brachinus armiger 9 5 Brachinus armiger Aptinus alpinus 10 Aptinus alpinus Aptinus displosor 100 Aptinus displosor Styphlodromus 100 Styphlodromus Styphlomerus 100 Styphlomerus Catapiesis Abacetus 11 Eripus Amara Pelecium Caelostomus Diploharpus Catapiesis Chlaenius Cnemalobus Stenocrepis 100 Dicaelus Galerita 11 Diploharpus 12 Morion Hybothecus Moriosomus Metius Caelostomus Pterostichus Abacetus Synuchus Agonum Agonum Metacolpodes 100 Metacolpodes Amblygnathus Amblygnathus Pelmatellus 100 Pelmatellus Cnemalobus Ega This research was supported by a grant from the National Sci- Amara 100 100 Lachnophorus Dicaelus 12 Chlaenius Hybothecus 100 Stenocrepis ence Foundation, EF-0531754 Metius Eripus Pterostichus 100 Pelecium Synuchus Galerita Cymindis 9 2 Morion Lebia 100 Moriosomus Anaulacus Anaulacus Tetragonoderus Cymindis Ega 100 Lebia Lachnophorus Tetragonoderus Pseudomorpha Cryptocephalomorpha Sphallomorpha Pseudomorpha Adelotopus Sphallomorpha Cainogenion 100 Adelotopus Cryptocephalomorpha 100 Cainogenion Orthogonius sp. 1 Orthogonius sp. 1 Orthogonius sp. 2 100 Orthogonius sp. 2