Tiphiidae

Female Tiphia femorata, copyright Hectonichus.

Belongs within: Euaculeata.

The Tiphiidae are a group of small wasps that develop as parasites of subterranean beetle larvae (Naumann 1991).

Characters (from Naumann 1991): Inner margins of compound eyes not strongly emarginate; pronotum freely articulating with mesothorax, mid-dorsally usually long, posterolaterally reaching tegula, ventrally rounded; mesostemum usually with plate-like extensions over mid coxae; propodeum not tripartite; fore wing without pseudovenation. Larvae with four-toothed mandibles and second pair of thoracic spiracles reduced or absent. Cocoon present.

<==Tiphiidae [Tiphidae, Tiphioidea]
    |  i. s.: Colocistis HR11
    |           |--C. crassa HH07
    |           |--C. eremi BD17
    |           `--C. sulcatus HR11
    |         Architiphia rasnitsyni Darling in Darling & Sharkey 1990 RJ93
    |         Pterombrus RD77
    |--Anthobosca Guérin de Méneville 1838 BL17 [incl. Adontothynnus Cameron 1906 T08; Anthoboscinae GE05]
    |--Diamma [Diamminae] K04
    |    `--D. bicolor N91
    |--Brachycistidinae K04
    |    |--Brachycistis timberlakei JB13
    |    `--Glyptometopa K04
    `--Tiphia [Tiphiinae] N91
         |  i. s.: T. annandalei Turner 1908 AJ30
         |         T. auripennis Bingham 1897 AJ30
         |         T. borneana Cameron 1907 AJ30
         |         T. brevipennis Lucas 1848 E12
         |         T. carbonaria Smith 1861 AJ30
         |         T. clypealis Cameron 1896–1897 AJ30
         |         T. consueta Smith 1879 AJ30
         |         T. flavipennis Smith 1858 AJ30
         |         T. fumipennis Smith 1858 AJ30
         |         T. fuscinervis Cameron 1896–1897 AJ30
         |         T. higoensis S94
         |         T. ifri S94
         |         T. implicata Cameron 1896–1897 AJ30
         |         T. incisa Cameron 1896–1897 AJ30
         |         T. intrudens Smith 1864 AJ30
         |         T. khasiana Cameron 1902 AJ30
         |         T. kisukei S94
         |         T. levisternalis S94
         |         T. maculata L02
         |         T. magnoliae S94
         |         T. nipponica S94
         |         T. ordinaria Smith 1873 AJ30
         |         T. oswini Turner 1911 AJ30
         |         T. punctata Smith 1873 AJ30
         |         T. punctifrons Cameron 1909 AJ30
         |         T. robusta Cameron 1904 AJ30
         |         T. rufipes Smith 1855 AJ30
         |         T. rufotibiata S94
         |         T. (Tengyra) sanvitali G20
         |         T. spinosa Cameron 1902 AJ30
         |         T. stigma Smith 1858 AJ30
         |         T. teranishii S94
         |         T. tibetana Turner 1908 AJ30
         |         T. transversa RD77
         |         T. vulgata S94
         |--T. (Tiphia) femorata G20
         |    |--T. f. femorata S94
         |    `--T. f. biseculata Allen & Jaynes 1930 S94, AJ30
         |--T. clauseni Allen & Jaynes 1930 AJ30
         |--T. latistriata Allen & Jaynes 1930 AJ30
         |--T. longitegulata Allen & Jaynes 1930 AJ30
         |--T. matura Allen & Jaynes 1930 AJ30
         |--T. minutopunctata Allen & Jaynes 1930 AJ30
         |--T. nana Allen & Jaynes 1930 AJ30
         |--T. pigmentata Allen & Jaynes 1930 AJ30
         |--T. pullivora Allen & Jaynes 1930 AJ30
         |--T. totopunctata Allen & Jaynes 1930 AJ30
         |--T. vernalis Rohwer 1924 AJ30
         |--T. agilis group AJ30
         |    |--T. agilis Smith 1873 AJ30
         |    `--T. asericae Allen & Jaynes 1930 AJ30
         |--T. bicarinata group AJ30
         |    |--T. bicarinata Cameron 1902 AJ30
         |    |--T. brevilineata Allen & Jaynes 1930 AJ30
         |    |--T. cilicincta Allen & Jaynes 1930 AJ30
         |    `--T. fukiensis Allen & Jaynes 1930 AJ30
         |--T. capillata group AJ30
         |    |--T. capillata Allen & Jaynes 1930 AJ30
         |    `--T. levipunctata Allen & Jaynes 1930 AJ30
         |--T. koreana group AJ30
         |    |--T. antigenata Allen & Jaynes 1930 AJ30
         |    |--T. assamensis Allen & Jaynes 1930 AJ30
         |    |--T. autumnalis Rohwer 1924 AJ30
         |    |--T. communis Allen & Jaynes 1930 AJ30
         |    |--T. fossata Allen & Jaynes 1930 AJ30
         |    |--T. koreana Rohwer 1927 AJ30
         |    |--T. ovidorsalis Allen & Jaynes 1930 AJ30
         |    `--T. tegitiplaga Allen & Jaynes 1930 AJ30
         |--T. malayana group AJ30
         |    |--T. brevistigma Allen & Jaynes 1930 AJ30
         |    |--T. compressa Smith 1855 AJ30
         |    `--T. malayana Cameron 1910 AJ30
         |--T. popilliavora group AJ30
         |    |--T. inconspicua Allen & Jaynes 1930 AJ30
         |    |--T. nervidirecta Allen & Jaynes 1930 AJ30
         |    |--T. ovinigris Allen & Jaynes 1930 AJ30
         |    |--T. phyllophagae Allen & Jaynes 1930 AJ30
         |    `--T. popilliavora Rohwer 1924 AJ30
         `--T. rufomandibulata group AJ30
              |--T. brevicarinata Allen & Jaynes 1930 AJ30
              |--T. lyrata Magretti 1892 AJ30
              |--T. notopolita Allen & Jaynes 1930 [incl. T. notopolita var. intermedia Allen & Jaynes 1930] AJ30
              |--T. rufomandibulata Smith 1873 AJ30
              |--T. singularis Allen & Jaynes 1930 AJ30
              |--T. sternocarinata Allen & Jaynes 1930 AJ30
              `--T. sternodentata Allen & Jaynes 1930 AJ30

*Type species of generic name indicated

REFERENCES

[AJ30] Allen, H. W., & H. A. Jaynes. 1930. Contribution to the taxonomy of Asiatic wasps of the genus Tiphia (Scoliidae). Proceedings of the United States National Museum 76 (17): 1–105.

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[HH07] Hines, H. M., J. H. Hunt, T. K. O'Connor, J. J. Gillespie & S. A. Cameron. 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences of the USA 104 (9): 3295–3299.

[JB13] Johnson, B. R., M. L. Borowiec, J. C. Chiu, E. K. Lee, J. Atallah & P. S. Ward. 2013. Phylogenomics resolves evolutionary relationships among ants, bees, and wasps. Current Biology 23: 2058–2062.

[K04] Kimsey, L. S. 2004. Illustrated keys to genera of the male wasps in the subfamily Thynninae (Hymenoptera: Tiphiidae). Proceedings of the Entomological Society of Washington 106 (3): 571–585.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S94] Shimizu, A. 1994. Phylogeny and classification of the family Pompilidae (Hymenoptera). Tokyo Metropolitan University Bulletin of Natural History 2: 1–142.

[T08] Turner, R. E. 1908. A revision of the Thynnidae of Australia. [Hymenoptera]. Part II. Proceedings of the Linnean Society of New South Wales 33: 70–256.

Euaculeata

Female Methocha japonica, from here.

Belongs within: Aculeata.
Contains: Apoidea, Formicidae, Scoliidae, Pompiloidea, Tiphiidae, Thynninae, Vespidae.

The Euaculeata are a major clade of stinging wasps, united by sexually dimorphic antennae with thirteen antennomeres in males and twelve in males, and a hidden and largely desclerotised seventh metasomal tergum in females (Grimaldi & Engel 2005).

<==Euaculeata [Formicina, Heterogyna, Melliniores, Mutillariae, Praedones, Scolietae, Sphegimae] GE05
    |  i. s.: Falsiformicidae GE05
    |           |--Falsiformica cretacea Rasnitsyn 1975 P92
    |           `--Taimyrisphex pristinus Evans 1973 P92
    |--+--+--Apoidea BD17
    |  |  `--+--Formicidae BD17
    |  |     `--Armania [Armaniidae] GE05
    |  |          `--A. robusta GE05
    |  `--Scolioidea BD17
    |       |--Scoliidae JB13
    |       `--Bradynobaenidae JB13
    |            |--Typhoctes [Typhoctinae] GE05
    |            `--Apterogyna [Apterogyninae] JB13
    |                 `--A. olivieri [=Mutilla (Apterogyna) olivieri] G20
    `--+--+--Sierolomorpha GE05 [Sierolomorphidae, Sierolomorphoidea BD17]
       |  |    |--S. bicolor B11
       |  |    `--S. canadensis B11
       |  `--+--Pompiloidea PK17
       |     `--+--Tiphiidae JB13
       |        `--Thynnoidea BD17
       |             |--Chyphotes JB13 [Chyphotidae BD17, Chyphotinae]
       |             |    `--C. mellipes RBV08
       |             `--Thynnidae BD17
       |                  |--Myzinum [Myzininae] BD17
       |                  |    |--M. frontale S94
       |                  |    `--M. quinquecinctum (Fabricius 1775) [=Tiphia quinquecincta, Sphex quinquecinctus] BM76
       |                  `--+--Thynninae BD17
       |                     `--Methocha Latreille 1804 BD17, LB08 [Methochinae N91, Methocidae]
       |                          |--M. articulata PK17
       |                          |--M. formicaria (Linnaeus 1758) G20, L58 [=Mutilla formicaria G20]
       |                          |--M. ichneumonides A71
       |                          |--M. japonica S94
       |                          `--M. yasumatsui S94
       `--Vespoidea BD17
             |--Vespidae JB13
             `--+--Rhopalosomatidae JB13
                |    |--Rhopalosoma nearcticum HH07
                |    |--Mesorhopalosoma cearae Darling in Darling & Sharkey 1990 RJ93
                |    |--Propalosoma GE05
                |    |--Olixon [incl. Psyllosphex Arnold 1935] S94
                |    `--Harpagocryptus australiae R70
                `--Scolebythidae HR11
                     |  i. s.: Libanobythus Prentice & Poinar in Prentice, Poinar & Milki 1996 PPM96
                     |           `--*L. milkii Prentice & Poinar in Prentice, Poinar & Milki 1996 PPM96
                     |         Dominibythus Prentice & Poinar in Prentice, Poinar & Milki 1996 PPM96
                     |           `--*D. inopinatus Prentice & Poinar in Prentice, Poinar & Milki 1996 PPM96
                     |         Boreobythus Engel & Grimaldi 2007 B11
                     |         Cretabythus sibiricus Evans 1973 P92
                     |--+--Clystopsenella Kieffer 1911 B11
                     |  |    `--C. longiventris PPM96
                     |  `--Scolebythus Evans 1963 B11
                     |       `--S. madecassus HR11
                     `--+--Ycaploca Nagy 1975 B11
                        |    `--Y. evansi GE05
                        `--Pristapenesia Brues 1933 B11
                             |--P. inopinata B11
                             `--P. primaeva B11

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

[B11] Brothers, D. J. 2011. A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata). ZooKeys 130: 515–542.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[HH07] Hines, H. M., J. H. Hunt, T. K. O'Connor, J. J. Gillespie & S. A. Cameron. 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences of the USA 104 (9): 3295–3299.

[JB13] Johnson, B. R., M. L. Borowiec, J. C. Chiu, E. K. Lee, J. Atallah & P. S. Ward. 2013. Phylogenomics resolves evolutionary relationships among ants, bees, and wasps. Current Biology 23: 2058–2062.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[PPM96] Prentice, M. A., G. O. Poinar, Jr & R. Milki. 1996. Fossil scolebythids (Hymenoptera: Scolebythidae) from Lebanese and Dominican amber. Proceedings of the Entomological Society of Washington 98 (4): 801–811.

[RBV08] Rabeling, C., J. M. Brown & M. Verhaagh. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences of the USA 105 (39): 14913–14917.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S94] Shimizu, A. 1994. Phylogeny and classification of the family Pompilidae (Hymenoptera). Tokyo Metropolitan University Bulletin of Natural History 2: 1–142.

Pompilidae

Minagenia osoria, copyright Centre for Biodiversity Genomics.

Belongs within: Pompiloidea.
Contains: Ceropales, Ctenocerinae, Pompilinae, Leptodialepis, Calopompilus, Hemipepsis, Pepsini, Psoropempula, Ageniellini, Priocnemis, Dipogon.

The Pompilidae, spider hawks, are a cosmopolitan group of wasps that develop as predators or parasites of spiders.

Characters (from Shimizu 1994): Antenna usually with 13 segments in male, 12 segments in female. Maxillary galea usually with two separate sclerites, exogalea-1 and -2; maxillary palpus 6-segmented; postmentum present; labial palpus four-segmented. Pronotum freely articulating with mesothorax; anterior "collar" usually well developed; posterolateral lobes (pronotal lobes) reaching tegula and covering the first thoracic spiracle. Propleura posteriorly contiguous mesally. Prosternum sunken between bases of fore coxae except for its anterior section. Large area of intersegmental membrane present ventrally between pro- and mesothoraces. Prepectus discontinuous midventrally, the halves widely separated, extending the height of mesepisternum, being fused to it with the line of fusion forming a sulcus. Mesepisternum with a median straight oblique sulcus (posterior oblique sulcus). Mesepimeron much reduced posterodorsally on mesopleuron. Mesosternum with weak ridges or dentate projections anteromesal to midcoxal cavities. Meso- and metapleura not mutually movable although not fused. Metapostnotum often well developed, visible medially. Metepimeron slightly expanded anteroventral to endophragmal pit so that the pit is a short distance from anterior margin of metapleuron and pleural sulcus angulate. Metasternum entirely depressed except for slight ridges anterior to hind coxae, well differentiated from mesosternum, not anteriorly produced. Metapleural-propodeal sulcus entirely visible. Propodeal spiracle oblique. Forewing usually with ten closed cells. Hindwing with two closed cells and a jugal lobe; vein C+Sc+R+Rs fused basally, and C reduced to a pigmented relict vein after separating from Sc+R+Rs; free section of vein 1A absent, of vein Cu present. Basal hamuli strongly concentrated into a basal cluster at or basad to separation of vein Sc+R+S from margin. Claval furrow barely indicated. Legs usually slender and elongated. Each pair of coxae (fore, mid and hind) large and usually contiguous. Fore tibia with single calcar almost straight, not highly modified. Mid tibia with two spurs slightly dorsally flattened. Hind tibia with two spurs slightly dorsally flattened, the inner one modified as a calcar by development of basal brush of setae dorsally. Dense brush of short, erect setae on inner side of hind tibia usually well developed for its entire length. Each tarsal segment with plantula posteroventrally; arolia well developed. Female metasoma with six visible segments and male with seven segments. Spiracles exposed on the first and second terga but those on terga 3-6 or -7 often concealed by the preceding tergum. Each tergum overlapping its corresponding sternum laterally. Each sternum posteriorly overlapping front edge of the succeeding one; first sternum not depressed posteriorly. Female seventh tergum entirely hidden, but continuously sclerotized anteriorly; section 1 of gonocoxite 9 not completely divided; no lamellate valve dorsally on gonapophysis 8 (=lancet or valvula 1). Male sixth sternum with a pair of strongly sclerotized sharp hooks posteriorly; seventh sternum reduced, withdrawn beneath sixth and overlapping the apodeme-like central process of subgenital plate (eighth sternum); eighth sternum essentially simple, not reduced nor hidden; composite tergum 9-10 (abdominal tergum 10-11) with a cercus on its each side; genitalia with elongate parapenial lobes from section 2 of gonocoxite 9 (basiparamere) and often with basal hooklets on lamina volsellaris; gonapophysis 9 (aedeagus) with notal fusion well developed.

Pompilidae [Pompilii, Psammocharidae]
    |--Ceropalinae [Ceratopalinae, Ceropalini] EG06
    |    |--Ceropales E07
    |    `--Irenangelus Schulz 1906 [incl. Xanthampulex Schulz 1906] E07
    |         |--‘Agenia’ reversa Smith 1873 [incl. *Irenangelus hornus Schulz 1906] E07
    |         |--I. eberhardi S94
    |         |--I. ichneumonoides S94
    |         |--‘Xanthampulex’ luzonensis S94
    |         |--I pernix (Bingham 1896) [=Ceropales pernis, Xanthampulex pernix] S94
    |         |--I. tenuatus (Turner 1910) [=Ceropales tenuatus] E07
    |         `--‘*Xanthampulex’ trifur Schulz 1906 E07
    `--+--+--Ctenocerinae EG06
       |  `--Pompilinae EG06
       `--Pepsinae [Cryptocheilinae, Macromerinae] EG06
            |  i. s.: Cryptosalius H04
            |         Austrosalius Turner 1917 E07
            |           `--*A. artemis (Turner 1910) [=Ferreolomorpha artemis] E07
            |         Derochorses Banks 1941 E07
            |           `--*D. seminiger Banks 1941 E07
            |         Java Pate 1946 [=Dichelonyx Haupt 1938 non Harris 1826] E07
            |           |--*J. concolor (Taschenberg 1869) [=Priocnemis (Cyphononyx) concolor, *Dichelonyx concolor] E07
            |           `--J. nigrita (Fabricius 1781) E07
            |                |--J. n. nigrita E07
            |                `--J. n. aspasia (Smith 1859) (see below for synonymy) E07
            |         Leptodialepis E07
            |         Paleogenia wahisi PK17
            |         Diplonyx Saussure 1887 E07 [incl. Monodontonyx Haupt 1929 B38, E07]
            |           |--‘Monodontonyx’ javanus B38
            |           |--‘Monodontonyx’ plutonis Banks 1938 B38
            |           `--‘Monodontonyx’ sericosoma B38
            |         Taiwania Tsuneki 1989 T89
            |           |--*T. formosana Tsuneki 1989 T89
            |           `--T. rufotibialis Tsuneki 1989 T89
            |         Poecilagenia Haupt 1927 T89
            |           |--P. rubricans S94
            |           `--P. sculpturata S94
            |         Malloscelis Haupt 1936 S94
            |           `--*M. formosus (Morawitz 1889) S94, T89 [=Salius formosus S94]
            |         Clistoderes Banks 1934 T89
            |           |  i. s.: C. onoi (Ishikawa 1955) [=Priocnemis onoi] T89
            |           |         C. taiwanus Tsuneki 1989 T89
            |           |         C. thione T89
            |           |--C. (Clistoderes) iwatai S94
            |           `--C. (Paraclistoderes Ishikawa 1962) S94
            |                |--C. (P.) futabae S94
            |                |--C. (P.) impunctatus S94
            |                |--C. (P.) nakanei S94
            |                `--C. (P.) shirozui S94
            |         Lissocnemis T89
            |           |--*L. irrasus T89
            |           |--L. apollo (Banks 1934) [=Dinagenia apollo] T89
            |           |--L. niger Tsuneki 1989 T89
            |           `--L. nigricoxis Haupt 1941 T89
            |         Taiwagenia Tsuneki 1989 T89
            |           `--*T. taiwana Tsuneki 1989 T89
            |--Chirodamus Townes 1957 S94, E07 [Chirodamini EG06]
            |--Calopompilus S94
            |--Hemipepsis S94
            |--Pepsini S94
            |--Plagicurgus Roig Alsina 1982 S94
            |    `--P. metallicus S94
            |--+--Pompilocalus Roig Alsina 1989 S94
            |  |    `--P. fraternus S94
            |  `--+--Aimatocare Roig Alsina 1989 S94
            |     `--Anacyphonyx Banks 1946 S94
            |--Cyphononyx Dahlbom 1845 S94
            |    |--C. bipartitus (Lepeletier 1845) [=Salius bipartitus, Leptodialepis bipartitus] T89
            |    |--C. bretoni S94
            |    |--C. cameroni Banks 1938 B38
            |    |--C. dorsalis (Lepeletier 1745) (see below for synonymy) T89
            |    |--C. fugidipennis B38
            |    |--C. gowdeyi (Turner 1918) E07
            |    `--C. iridipennis (Smith 1859) S94, T90 (see below for synonymy)
            `--+--Psoropempula S94
               |--Ageniellini S94
               |--+--Platydialepis Haupt 1941 S94
               |  |    |--P. ryoheii (Ishikawa 1957) [=Malloscelis ryoheii] S94
               |  |    `--P. taiwanianus (Tsuneki 1989) [=Malloscelis taiwanianus] S94
               |  `--Priocnessus Banks 1925 S94
               |       |--P. hurdi S94
               |       |--P. nebulosus S94
               |       `--P. nuperus S94
               `--+--Priocnemis S94
                  |--Dipogon S94
                  |--Ctenopriocnemis Ishikawa 1962 S94
                  |    `--C. filicornis S94
                  |--Eopompilus Gussakovskij 1933 S94
                  |    |--E. internalis S94
                  |    `--E. minor S94
                  |--Caliadurgus Pate 1946 S94
                  |    |--C. bituberculatus S94
                  |    |--C. fasciatellus S94
                  |    |--C. kurosawai S94
                  |    `--C. ussuriensis (Gussakovskij 1933) (see below for synonymy) T89
                  |--Sphictostethus Kohl 1884 S94, E07 (see below for synonymy)
                  |    |--*S. gravesii (Haliday 1836) [=Pompilus gravesii] E07
                  |    |--S. aliciae (Turner 1914) [=Calopompilus aliciae, Priocnemis aliciae] E07
                  |    |--‘*Haploneura’ apogona Kohl 1884 [=*Haploneurion apogonum] E07
                  |    |--S. calvus H05
                  |    |--‘Pompilus’ flavipes Guérin-Méneville 1836 [=*Anapriocnemis flavipes] E07
                  |    |--S. fugax [=Pompilus fugax] E07
                  |    |--S. nitidus (Fabricius 1775) E07 (see below for synonymy)
                  |    `--S. xanthochrous (Turner 1915) [=Calopompilus xanthochrous, Priocnemis xanthochrous] E07
                  `--Minagenia Banks 1934 [Minageniini] S94
                       |--M. alticola Tsuneki 1989 T89
                       |--M. brevicornis T89
                       |--M. granulosa Tsuneki 1989 T89
                       |--M. julia S94
                       |--M. montisdorsa S94
                       |--M. osoria S94
                       |--M. pempuchiensis Tsuneki 1989 T89
                       `--M. taiwana Tsuneki 1989 T89

Pompilidae incertae sedis:
  Priocnemioides Rad. 1888 H37
    `--P. unifasciatus H37
  Ceropalites infelix Cockerell 1906 EG06
  Planiceps Latreille 1825 S94 [Planicepinae R70]
    |--P. aureovestitus Turner 1910 E07
    `--P. fulviventris R13
  Atelostegus Haupt 1929 S94
  Pygmachus Haupt 1930 S94
  Dromochares Haupt 1930 S94
  Gonaporus Ashmead 1902 S94
  Taeniaporus Haupt 1930 S94
  Aeluropetrus Arnold 1936 S94
  Rhynchopompilus Arnold 1934 S94
  Euplaniceps Haupt 1930 S94
  Aspidaporus Bradley 1944 S94
  Odontaporus Bradley 1944 S94
  Cosmiaporus Bradley 1944 S94
  Drepanaporus Bradley 1944 S94
  Rhabdaporus Bradley 1944 S94
  Tupiaporus Arlé 1947 S94
  Abripepsis Banks 1946 S94
  Hatanomus Tsuneki 1990 T90
    `--*H. granulosus Tsuneki 1990 T90

Caliadurgus ussuriensis (Gussakovskij 1933) [=Caricurgus ussuriensis; incl. Car. shirakii Yasumatsu 1943] T89

Cyphononyx dorsalis (Lepeletier 1745) [=Calicurgus dorsalis, Pompilus dorsalis, Priocnemis dorsalis, Salius dorsalis; incl. Salius flavus Bingham 1897, Cyphononyx flavus, Cryptoceilus flavus, Pallosoma fulvognathus Rohwer 1911, Cyphononyx fulvognathus] T89

Cyphononyx iridipennis (Smith 1859) S94, T90 [=Cryptochilus (Cyphononyx) iridipennis T90, Dichelonyx iridipennis T90, Macromeris iridipennis T90, Salius iridipennis T90; incl. Mignimia pulchripennis Smith 1875 T90, Cyphononyx pulchripennis T90]

Java nigrita aspasia (Smith 1859) [=Mygnimia aspasia, Cryptocheilus aspasia, Cyphononyx aspasia, Dichelonyx aspasia, Salius aspasia] E07

Sphictostethus Kohl 1884 S94, E07 [=Sphictonotus (l. c.) E07; incl. Anapriocnemis Haupt 1959 E07, Chrysocurgus Haupt 1937 H37, Haploneura Kohl 1884 non Loew 1850 E07, Haploneurion Kohl 1884 E07]

Sphictostethus nitidus (Fabricius 1775) E07 [=Sphex nitida E07, *Chrysocurgus nitidus E07; incl. Priocnemis wakefieldi H37]

*Type species of generic name indicated

REFERENCES

[B38] Banks, N. 1938. Some Psammocharidae from Singapore. Proceedings of the Entomological Society of Washington 40 (8): 236–249.

[E07] Elliott, M. G. 2007. Annotated catalogue of the Pompilidae (Hymenoptera) of Australia. Zootaxa 1428: 1–83.

[EG06] Engel, M. S., & D. A. Grimaldi. 2006. The first Cretaceous spider wasp (Hymenoptera: Pompilidae). Journal of the Kansas Entomological Society 79 (4): 359–368.

[H04] Harris, A. C. 2004. Priocnemis (Trichochurgus) monachus (Smith) (Hymenoptera: Pompilidae) preying on Misgolas napua Forster (Mygalomorphae: Idiophidae) after opening the lids of the prey’s burrows, and notes of the affinities of Trichocurgus Haupt. Weta 27: 13–14.

[H05] Harris, A. C. 2005. House sparrows (Passer domesticus) repeatedly robbing red spider wasps (Sphictostethus nitidus) (Hymenoptera: Pompilidae) of their paralysed spider prey (Neoramia otagoa) (Araneae: Agelenidae). Weta 29: 21–23.

[H37] Haupt, H. 1937. Beitrag zur Kenntnis der Psammochariden-Fauna Australiens (Spinnenfangende Wegwespen). Subf. Claveliinae Hpt. Trib. Calicurgini nov. Zeitschrift für Naturwissenschaften 91: 121–136.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

[S94] Shimizu, A. 1994. Phylogeny and classification of the family Pompilidae (Hymenoptera). Tokyo Metropolitan University Bulletin of Natural History 2: 1–142.

[T89] Tsuneki, K. 1989. A study on the Pompilidae of Taiwan (Hymenoptera). Special Publications of the Japan Hymenopterists Association 35: 1–180.

[T90] Tsuneki, K. 1990. The Pompilidae of the Ryukyus (Hymenoptera). Special Publications of the Japan Hymenopterists Association 36: 1–65.

Mutillidae

Pseudophotopsis komarovii, copyright Centre for Biodiversity Genomics.

Belongs within: Pompiloidea.
Contains: Kudakrumiini, Myrmosini, Ticoplinae, Rhopalomutillinae, Sphaeropthalminae, Dasylabrinae, Myrmillinae, Mutillinae.

The Mutillidae, velvet ants, are hairy wasps that usually show significant dimorphism with females being wingless and have the sclerites of the mesosoma fused whereas males are mostly winged with articulated mesosomal plates. In the basal subfamily Myrmosinae, found in Eurasia and North America, females retain an articulated suture between the pro- and mesonotum (Brothers & Lelej 2017).

Synapomorphies (from Brothers & Lelej 2017): Tergum II and sternum I in both sexes, tergum II overlying lateral extremities of sternum I; scape in female with flangelike expansion above radicle, angled; wings absent in female; metacoxal cavities closed in female; tergum II with stridulitrum in both sexes (lost in some subgroups); sternum I in both sexes depressed posteriorly, constricted and abutting sternum II; hind wing in male with jugal lobe reduced; penis valve (gonapophysis IX) with apex dorsally produced, ventral tooth about midway (further modified in most subgroups).

Mutillidae [Cretavidae, Sphaeropthalmina]
    |  i. s.: Smicromutilla Mickel 1964 LB08
    |           `-*S. powelli Mickel 1964 LB08
    |         Cretavus Sharov 1957 LB08
    |           `--*C. sibiricus Sharov 1957 LB08
    |         Scaptopoda Lynch Arribálzaga 1878 (n. d.) LB08
    |           `--*S. pusilla Lynch Arribálzaga 1878 LB08
    |--Myrmosinae [Kudakrumiinae, Myrmosidae] BL17
    |    |--Kudakrumiini BL17
    |    `--Myrmosini BL17
    `--+--Pseudophotopsis André 1896 BL17, LB08 (see below for synonymy)
       |    |--*P. komarovii (Radoszkowski 1885) [=Agama komarovii] LB08
       |    |--P. asiatica Suárez 1965 [=Ephutomma asiatica] LB08
       |    |--P. binghami Bischoff 1920 BL17
       |    |--P. continua (Fabricius 1804) BL17 [=Mutilla continua LB08, *Sphinctomutilla continua LB08]
       |    |--P. irana (Skorikov 1935) BL17
       |    |--‘Agama’ kokpetica Radoszkowski 1885 [=Agama kotepetica (l. c.), *Alloneurion kokpeticum] LB08
       |    `--P. schachruda (Skorikov 1935) BL17
       `--+--Ticoplinae BL17
          `--+--Rhopalomutillinae BL17
             `--+--Sphaeropthalminae BL17
                `--+--Dasylabrinae BL17
                   `--+--Myrmillinae BL17
                      `--Mutillinae BL17

Pseudophotopsis André 1896 BL17, LB08 [incl. Alloneurion Ashmead 1899 LB08, Sphinctomutilla André 1899 LB08; Pseudophotopsidinae]

*Type species of generic name indicated

REFERENCES

[BL17] Brothers, D. J., & A. S. Lelej. 2017. Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research 60: 1–97.

[LB08] Lelej, A. S., & D. J. Brothers. 2008. The genus-group names of Mutillidae (Hymenoptera) and their type species, with a new genus, new name, new synonymies, new combinations and lectotypifications. Zootaxa 1889: 1–79.

Last updated: 11 March 2021.

Apocrita

Ceraphron sp., copyright Pierre Bornand.

Belongs within: Unicalcarida.
Contains: Platygastridae, Cynipoidea, Proctotrupoidea, Ichneumonoidea, Aculeata, Megalyridae, Evanioidea.

The Apocrita are a clade of wasps characterised by the development of a narrow waist between the first and second abdominal segments with the first abdominal segment being incorporated into the mesosoma as the propodeum (Grimaldi & Engel 2005). Molecular analysis supports recognition of a major clade of parasitoid forms, including the Proctotrupomorpha, Ichneumonoidea and Ceraphronoidea, as the sister taxon of the remaining members (Peters et al. 2017). The Ceraphronoidea are minute wasps characterised by two protibial spurs, an enlarged second metasomal segment, and fore wings with the costal and subcostal + radial veins fused into a bar running along the anterior margin. The Stigmaphronidae, a Mesozoic family possibly related to the Ceraphronoidea, have similarly fused anterior wing veins but have a single protibial vein and the second metasomal segment is not enlarged (Grimaldi & Engel 2005). The Proctotrupomorpha are a diverse clade of small, often minute, wasps united by the lack of a median mesoscutal sulcus and a number of characters relating to reduction in wing venation (Rasnitsyn 2002).

Other members of the Apocrita include the Trigonalidae, hyperparasitoid wasps within caterpillars or sawfly larvae whose females lay their eggs within incisions in leaves where they are ingested by the primary host when feeding. The Stephanidae are slender wasps with a mobile, globular head bearing a crown of rasp-like teeth that parasitise larvae of wood-boring insects (Naumann 1991).

<==Apocrita [Clistogastra, Ichneumonides, Proctotrupii, Terebrantes] PK17
    |--Parasitoida PK17
    |    |--Proctotrupomorpha [Cinipserae, Cynipsera, Diplolepariae] PK17
    |    |    |  i. s.: Proctorenyxidae GE05
    |    |    |         Neotelenomus G17
    |    |    |           |--N. anthereae Dodd 1913 GM79
    |    |    |           `--N. javae Girault 1917 G17
    |    |    |         Eulagynodes Girault 1917 G17
    |    |    |           `--*E. bicolor Girault 1917 G17
    |    |    |         Acoloides G17
    |    |    |           |--A. arachnophagus Girault 1917 GM79
    |    |    |           |--A. fasciatipennis Girault 1917 G17
    |    |    |           |--A. javensis Girault 1917 G17
    |    |    |           `--A. trispinosus Girault 1917 G17
    |    |    |         Mesoserphus GE05 [Mesoserphidae R02, Mesoserphinae]
    |    |    |           `--M. karatavicus GE05
    |    |    |--Platygastridae MH11
    |    |    `--+--Cynipoidea MH11
    |    |       `--Proctotrupoidea MH11
    |    `--+--Ichneumonoidea PK17
    |       `--+--Stigmaphronidae GE05
    |          |    |--Aphrostigmon vitimense Rasnitsyn 1991 RJ93
    |          |    |--Stigmaphron Kozlov 1975 P92
    |          |    |    `--*S. orphne Kozlov 1975 P92
    |          |    |--Elasmomorpha Kozlov 1975 P92
    |          |    |    `--*E. melpomene Kozlov 1975 P92
    |          |    `--Hippocoon Kozlov 1975 P92
    |          |         `--*H. evadne Kozlov 1975 P92
    |          `--Ceraphronoidea PK17
    |               |  i. s.: Libanophron astarte PK17
    |               |--Megaspilidae GE05
    |               |    |--Lagynodes Forster 1841 GM79 [Lagynodinae N91]
    |               |    `--Megaspilinae N91
    |               |         |--Megaspilus fuscipennis MH11
    |               |         |--Dendrocerus carpenteri N91, M83
    |               |         `--Conostigmus N91
    |               |              |--C. dolicharthrus P92
    |               |              `--C. lazaros PP19
    |               `--Ceraphronidae [Calliceratidae] GE05
    |                    |--Lygocerus dubitatus Brues 1937 P92
    |                    |--Aphanogmus N91
    |                    |--Allocotidus bruesi Muesebeck 1963 P92
    |                    |--Prolagynodes penniger P92
    |                    `--Ceraphron Jurine 1807 GM79 [incl. Calliceras P92]
    |                         |--C. achilles Dodd 1914 GM79
    |                         |--C. bispinosus MH11
    |                         |--C. javensis Girault 1917 G17
    |                         `--C. sulcatum [=Sparasion (Ceraphron) sulcatum] G20
    `--+--+--Aculeata PK17
       |  `--+--Megalyridae HR11
       |     `--Trigonalidae [Archiglossata, Ichneumomimidae, Trigonaloidea, Trigonalyidae] HR11
       |          |--Mimelogonalos N91
       |          |--Orthogonalys pulchella HR11
       |          |--Pseudogonalos hahnii PK17
       |          |--Darbigonalus capitatus Rasnitsyn 1986 RJ93
       |          |--Lycogaster pullata [incl. L. pullata nevadensis] S96
       |          |--Bareogonalos canadensis S96
       |          |--Poecilogonalos thwaitesii RD77
       |          |--Cretogonalys taimyricus Rasnitsyn 1977 P92
       |          |--Trigonalys B11
       |          |    |--T. maculatus RD77
       |          |    |--T. micanticeps B11
       |          |    `--T. pervetus Cockerell 1917 P92
       |          `--Taeniogonalos HR11
       |               |--T. gundlachii [incl. Poecilogonalos costalis] S96
       |               |--T. maculata B11
       |               `--T. venatoria R70
       `--+--Evanioidea PK17
          `--Stephanoidea H02
               |--Eostephanites Hong 2002 [Eostephanitidae] H02
               |    `--*E. tenuis Hong 2002 H02
               `--Stephanidae HR11
                    |  i. s.: Megischus HR11
                    |           |--M. basalis Aguiar 2005 A05
                    |           `--M. furcatus (Lepeletier & Serville 1825) [incl. M. annulator Brullé 1846] A05
                    |         Protostephanus ashmeadi A05
                    |         Electrostephanus A05
                    |           |--E. brevicornis A05
                    |           `--E. neovenatus A05
                    |         Hemistephanus A05
                    |--Stephanus PK17 [Stephaninae GE05]
                    |    |--S. coronator B35 (see below for synonymy)
                    |    `--S. serrator PK17
                    `--Schlettererinae GE05
                         |--Archaeostephanus corae GE05
                         `--Schlettererius cinctipes N91

Stephanus coronator B35 [=Bracon (Stephanus) coronatus G20, Ichneumon coronatus L02, Pimpla coronator B35]

*Type species of generic name indicated

REFERENCES

[A05] Aguiar, A. P. 2005. A new and unusual species of Stephanidae (Hymenoptera), with a discussion on its phylogenetic implications. Journal of Hymenoptera Research 14 (1): 1–6.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[B11] Brothers, D. J. 2011. A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata). ZooKeys 130: 515–542.

[G17] Girault, A. A. 1917. New Javanese Hymenoptera. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 59–71).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[MH11] Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One 6 (11): e27023.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[PP19] Palencia, L., E. Peñalver, C. E. Prieto & F. J. Poyato-Ariza. 2019. First fossil harvestmen (Arachnida: Opiliones) from Spain and notes on the fossil record of Opiliones. Palaeontologica Electronica 22.1.5A: 1–18.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[R02] Rasnitsyn, A. P. 2002. Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linné, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S96] Smith, D. R. 1996. Trigonalyidae (Hymenoptera) in the eastern United States: seasonal flight activity, distribution, hosts. Proceedings of the Entomological Society of Washington 98 (1): 109–118.

Last updated: 20 March 2021.

Aulaciformes

Aulacus striatus, copyright Jesper Lund.

Belongs within: Evanioidea.
Contains: Gasteruption.

The Aulaciformes are a clade of parasitoid or larval predatory wasps characterised by the absence of the jugal lobe in the hind wing, fusion of the first and second metasomal terga, sexually dimorphic antennal segmentation, and elongate propleuron, and rigid attachment of the pronotum and mesepisternum (Grimaldi & Engel 2005). Members of the Aulacidae have the metasoma separated from the metanotum by more than the anterior width of the first metasomal tergite whereas Gasteruptiidae have the metasoma and metanotum closer together (Naumann 1991).

Aulaciformes GE05
    |--Manlaya R02 [Baissidae GE05, Manlayinae]
    |    `--M. laevinota Rasnitsyn 1986 RJ93
    `--+--Aulacidae GE05
       |    |--Pristaulacus S96
       |    |    |--P. bilobatus S96
       |    |    |--P. canadensis S96
       |    |    |--P. fasciatus S96
       |    |    |--P. flavicrurus S96
       |    |    |--P. niger S96
       |    |    |--P. resutorivorus S96
       |    |    |--P. rufitarsis S96
       |    |    |--P. stigmaterus S96
       |    |    |--P. strangaliae HR11
       |    |    `--P. violaceus S96
       |    |--Aulacostethus apicalis R70
       |    |--Xuaulacites Hong 2002 H02
       |    |    `--*X. fushunensis Hong 2002 H02
       |    `--Aulacus N91
       |         |--A. aneurus S96
       |         |--A. brevicaudus S96
       |         |--A. burquei PK17
       |         |--A. digitalis S96
       |         |--A. dispilis S96
       |         |--A. fusicornis R70
       |         |--A. impolitus Smith 1991 S96
       |         |--A. lovei S96
       |         |--A. pallipes S96
       |         |--A. schiffi Smith 1996 S96
       |         `--A. striatus A71
       `--Gasteruptiidae [Gasteruptionidae, Kotujellidae] GE05
            |  i. s.: Tillywhimia R02
            |         Odontofoenus R70
            |--Kojutellitinae GE05
            |--Gasteruption N91
            `--Hyptiogastrinae GE05
                 |--Hyptiogaster thoracicus (Guérin-Méneville 1844) [=Foenus thoracicus] P57
                 |--Crassifoenus N91
                 |--Aulacofoenus N91
                 `--Eufoenus N91

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[P57] Pasteels, J. J. 1957. Revision du genre Gasteruption (Hymenoptera, Evanoidea, Gasteruptionidae). Espèces australiennes. Mémoires, Institut Royal des Sciences Naturelles de Belgique, sér. 2, 56: 1–125.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[R02] Rasnitsyn, A. P. 2002. Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linné, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S96] Smith, D. R. 1996. Aulacidae (Hymenoptera) in the mid-Atlantic states, with a key to species of eastern North America. Proceedings of the Entomological Society of Washington 98 (2): 274–291.

Last updated: 18 March 2021.

Crabronidae

Dinetus pictus, copyright Sarah Gregg.

Belongs within: Apoidea.
Contains: Sericophorus, Bothynostethini, Solierella, Miscophus, Oxybelini, Crabronini, Palarus, Plenoculus, Paranysson, Larrini, Lyroda, Sphodrotes, Trypoxylini, Nitela.

The Crabronidae are a group of apoid wasps restricted by Sann et al. (2018) on the basis of molecular data to a clade including members of the genus Dinetus together with the subfamilies Crabroninae and Larrinae as recognised by Bohart & Menke (1976). Members of this clade usually have antennal sockets low on the face, often contiguous with the frontoclypeal suture, and have a relatively short jugal lobe on the hind wing. Dinetus is a Palaearctic genus of small, colourful wasps in which the males have distinctively rolled and twisted antennae (Bohart & Menke 1976).

Crabronidae
    |--Dinetus Panzer 1806 PK17, BM76 [Dinetinae, Dinetini]
    |    |--*D. pictus (Fabricius 1793) BM76 (see below for synonymy)
    |    |--D. arenarius Kazenas 1973 BM76
    |    |--D. cereolus Morice 1897 BM76
    |    |    |--D. c. cereolus BM76
    |    |    `--D. c. politus Turner 1917 BM76
    |    |--D. dentipes Saunders 1910 [incl. D. gracilis Giner Marí 1945] BM76
    |    |--D. nabataeus Beaumont 1960 BM76
    |    |--D. pulawskii Beaumont 1960 BM76
    |    |--D. simplicipes Saunders 1910 [incl. D. perezi Ferton 1914] BM76
    |    `--D. venustus Beaumont 1956 BM76
    `--Crabroninae [Larridae, Larrinae, Miscophini] SN18
         |  i. s.: Sericophorini [Sericophorinae] LP10
         |           |--Sericophorus LP10
         |           `--Larrisson Menke 1967 LP10
         |                |--*L. abnormis (Turner 1914) C85 [=Sericophorus abnormis LP10]
         |                |--L. azyx Menke 1979 C85
         |                |--L. nedymus Menke 1979 C85
         |                `--L. rieki Menke 1979 C85
         |         Aha Menke 1977 LP10, C85
         |           |--*A. ha Menke 1977 C85
         |           `--A. evansi Menke 1977 C85
         |         0--Bothynostethini BM76
         |         `--Scapheutini BM76
         |              |--Bohartella Menke 1968 BM76
         |              |    `--*B. scapheutoides Menke 1968 BM76
         |              `--Scapheutes Handlirsch 1887 BM76
         |                   |--*S. mocsaryi Handlirsch 1887 BM76
         |                   |--S. brasilianus Handlirsch 1895 BM76
         |                   |--S. flavopictus (Smith 1860) [=Pison flavopictus] BM76
         |                   |--S. friburgensis Brèthes 1913 BM76
         |                   `--S. laetus (Smith 1860) [=Pison laetus] BM76
         |         0--Solierella BM76
         |         `--+--+--Miscophus BM76
         |            |  `--Saliostethus Brauns in Kohl 1896 [incl. Mutillonitela Bridwell 1920] BM76
         |            |       |--*S. lentifrons Braun 1896 BM76
         |            |       |--S. capicola Arnold 1923 BM76
         |            |       |--S. lounsburyi (Bridwell 1920) [=Mutillonitela lounsburyi] BM76
         |            |       |--S. mimicus (Bridwell 1920) [=*Mutillonitela mimica] BM76
         |            |       `--S. rhodesianus Arnold 1929 BM76
         |            `--+--Saliostethoides Arnold 1924 BM76
         |               |    `--*S. saltator Arnold 1924 (see below for synonymy) BM76
         |               `--Miscophoides Brauns in Kohl 1896 BM76
         |                    |--*M. (Miscophoides) handlirschii Brauns 1896 BM76
         |                    `--M. (Miscophoidellus Menke in Bohart & Menke 1976) BM76
         |                         |--M. (*M.) formosus Arnold 1924 BM76
         |                         `--M. (M.) minutus Arnold 1952 BM76
         |--+--Oxybelini PK17
         |  `--Crabronini PK17
         `--+--+--Palarus SN18
            |  `--+--Mesopalarus Brauns 1899 BM76
            |     |    |--*M. mayri Brauns 1899 BM76
            |     |    `--M. turneri Arnold 1931 BM76
            |     `--+--Plenoculus BM76
            |        `--Paranysson SN18
            `--+--+--Larrini SN18
               |  `--+--Lyroda SN18
               |     `--Sphodrotes SN18
               `--+--Trypoxylini SN18
                  `--+--Nitela SN18
                     `--Auchenophorus Turner 1907 BM76
                          |--*A. coruscans Turner 1907 C85
                          |--A. aeneus Turner 1907 C85
                          `--A. fulvicornis Turner 1907 C85

Crabronidae incertae sedis:
  ‘Vespa’ argyrostoma Gmelin 1790 [=V. argurostoma (l. c.)] BM76
  ‘Vespa’ braccata Gmelin 1790 BM76
  ‘Vespa’ canescens Gmelin 1790 BM76
  ‘Vespa’ chrysogona Gmelin 1790 BM76
  ‘Vespa’ chrysopus Gmelin 1790 BM76
  ‘Vespa’ chrysostoma Gmelin 1790 BM76
  ‘Vespa’ clavata Gmelin 1790 BM76
  ‘Vespa’ cylindrica Gmelin 1790 BM76
  ‘Vespa’ dichroa Gmelin 1790 BM76
  ‘Vespa’ ferruginea Gmelin 1790 BM76
  ‘Vespa’ flavicans Gmelin 1790 BM76
  ‘Vespa’ flavipes Gmelin 1790 BM76
  ‘Vespa’ immaculata Gmelin 1790 [=V. immacula (l. c.)] BM76
  ‘Vespa’ leskii Gmelin 1790 BM76
  ‘Vespa’ lutea Gmelin 1790 BM76
  ‘Vespa’ marginata Gmelin 1790 BM76
  ‘Vespa’ melanostoma Gmelin 1790 [=V. melanops (l. c.)] BM76
  ‘Vespa’ nigerrima Gmelin 1790 BM76
  ‘Vespa’ ochropus Gmelin 1790 BM76
  ‘Vespa’ punctulata Gmelin 1790 BM76
  ‘Vespa’ tibialis Gmelin 1790 BM76
  ‘Vespa’ unifasciata Gmelin 1790 BM76

*Dinetus pictus (Fabricius 1793) BM76 [=Crabro pictus BM76, Larra (Dinetus) picta G20; incl. C. ceraunius Rossi 1794 BM76, Sphex guttatus Fabricius 1793 non Gmelin 1790 BM76]

*Saliostethoides saltator Arnold 1924 [=Saliostethus saltator; incl. Miscophus reptans Arnold 1962] BM76

*Type species of generic name indicated

REFERENCES

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[C85] Cardale, J. C. 1985. Vespoidea and Sphecoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 150–303. Australian Government Publishing Service: Canberra.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[LP10] Lomholdt, O., & W. J. Pulawski. 2010. A revision of the wasp genus Sericophorus F. Smith, 1851 (Hymenoptera: Apoidea: Crabronidae). Proceedings of the California Academy of Sciences, series 4, 61 (1): 1–234.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[SN18] Sann, M., O. Niehuis, R. S. Peters, C. Mayer, A. Kozlov, L. Podsiadlowski, S. Bank, K. Meusemann, B. Misof, C. Bleidorn & M. Ohl. 2018. Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. BMC Evolutionary Biology 18: 71.

Last updated: 27 August 2020.