Bittacidae

Harpobittacus septentrionis, copyright Summerdrought.

Belongs within: Mecopteroidea.

The Bittacidae, hanging-flies, are a group of large, predatory scorpionflies that commonly hunt by hanging from vegetation with their fore legs and grasping prey with the hind legs (Byers 1991).

Characters (from Byers 1991): Single-clawed, raptorial tarsi. Wings elongate, slender basally. Compound eyes large, protruding; ocelli large, on a median prominence; rostrum slender. Basistyles of male bulbous, broadly fused ventrally; dististyles ordinarily small, inconspicuous; T9 divided into two lobes, clasper-like in appearance but not in function. Larvae with two dorsal rows of branched, fleshy processes.

<==Bittacidae
    |--Anomalobittacus N02
    |--Apterobittacus N02
    |--Archebittacus exilis N02
    |--Orobittacus N02
    |--Symbittacus L94
    |--Kalobittacus L94
    |--Nannobittacus L94
    |--Hylobittacus L94
    |--Tytthobittacus Smithers 1973 R74
    |--Austrobittacus B91
    |--Edriobittacus B91
    |--Probittacus avitus RJ93
    |--Bittacus Latreille 1802 L02
    |    |--*B. tipularius [=Panorpa tipularia] L02
    |    |--B. apterus R13
    |    |--B. eremus L94
    |    |--B. mastrillii CM07
    |    |--B. pilicornis KP19
    |    `--B. strigosus HR11
    `--Harpobittacus Gerstaecker 1885 L94
         |--+--+--H. rubricatus Riek 1954 [=H. tillyardi rubricatus] L94
         |  |  `--H. tillyardi Esben-Petersen 1915 [incl. H. nigratus Navás 1932] L94
         |  `--+--*H. australis (Klug 1838) (see below for synonymy) L94
         |     `--+--H. albatus Riek 1954 [=H. tillyardi alatus; incl. H. limnaeus Smithers 1973] L94
         |        `--H. christine Lambkin 1994 L94
         `--+--+--H. quasisimilis Lambkin 1994 L94
            |  `--H. similis Esben-Petersen 1935 L94
            `--+--+--H. nigriceps (Selys-Longchamps 1868) [=Bittacus nigriceps] L94
               |  `--H. phaeoscius Riek 1954 L94
               `--+--H. scheibeli Esben-Petersen 1935 [incl. H. brewerae Smithers 1973] L94
                  `--H. septentrionis Lambkin 1994 L94

*Harpobittacus australis (Klug 1838) [=Bittacus australis; incl. B. corethrarius Rambur 1842, B. intermedius Selys-Longchamps 1868, Harpobittacus intermedius, H. australis rubripes Riek 1954] L94

*Type species of generic name indicated

REFERENCES

[B91] Byers, G. W. 1991. Mecoptera (scorpion-flies, hanging-flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 696–704. Melbourne University Press: Carlton (Victoria).

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[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.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L94] Lambkin, K. J. 1994. Revision of the Australian scorpion-fly genus Harpobittacus (Mecoptera: Bittacidae). Invertebrate Taxonomy 8: 767–808.

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

[N02] Novokshonov, V. G. 2002. Order Panorpida Latreille, 1802. The scorpionflies (=Mecaptera Packard, 1886, =Mecoptera Comstock et Comstock, 1895, +Neomecoptera Hinton, 1958, +Paratrichoptera Tillyard, 1919, +Paramecoptera Tillyard, 1919). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 194–199. Kluwer Academic Publishers: Dordrecht.

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

[R74] Riek, E. F. 1974. Mecoptera (scorpion-flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 90. 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.

Mecopteroidea

Orthophlebia grandis part and counterpart fossil, copyright Bjoertvedt.

Belongs within: Panorpoidea.
Contains: Siphonaptera, Aneuretopsychina, Bittacidae, Boreidae, Panorpidae.

The Mecopteroidea are a clade of insects uniting the modern orders Mecoptera and Siphonaptera which share a fused clypeus and labrum together with a lining of spicule-like teeth in the proventriculus (Grimaldi & Engel 2005). The Early–Mid-Jurassic Muchoriidae, characterised by wing with a short subcosta, vein R angular at the pterostigmal base, two-branched RS, a short MA fork and four-branched MP (Novokshonov 2002) may represent the sister group of the two orders together.

The Mecoptera are commonly known as scorpion-flies due to the large bulbous terminal genitalia of males in most families, which superficially resemble a scorpion's tail. The head and mouthparts are elongate and ventrally prolonged. They are commonly regarded as among the most generalised representatives of the holometabolous insects, retaining many plesiomorphic features such as wings with an extensive venation and relatively little differentiation between fore and hind wings (Grimaldi & Engel 2005). The fossil record of Mecoptera is correspondingly extensive, going back to the Late Permian. Among modern taxa, the Tasmanian species Apteropanorpa tasmanica is entirely wingless with the mesothoracic and metathoracic sclerites reduced and fused (Byers 1991).

<==Mecopteroidea [Mecopterida] RL09
    |--Muchoria RJ93 [Muchoriidae RL09]
    |    `--M. reducta RJ93
    `--+--Siphonaptera RL09
       `--Mecoptera [Eumecoptera] RL09
            |  i. s.: Anormochorista [Anormochoristidae] RJ93
            |           `--A. oligoclada RJ93
            |         Choristopanorpa [Choristopanorpidae] RJ93
            |           |--C. bifasciata Riek 1950 F71
            |           `--C. drinnani Jell & Duncan 1986 RJ93
            |         Cimbrophlebia [Cimbrophlebiidae] RJ93
            |           `--C. bittaciformis RJ93
            |         Cyclopterina [Cyclopterinidae] RJ93
            |           `--C. autumnalis RJ93
            |         Cyclorista [Cycloristidae] RJ93
            |           `--C. convexicosta RJ93
            |         Lithopanorpa [Lithopanorpidae] RJ93
            |           |--L. kuznetskiensis RJ93
            |           `--L. pusilla RJ93
            |         Martynopanorpa [Martynopanorpidae] RJ93
            |           `--M. angustata RJ93
            |         Neorthophlebia [Neorthophlebiidae] RJ93
            |           `--N. maculipennis RJ93
            |         Neoparachorista [Neoparachoristidae] RJ93
            |           |--N. clarkae Jell & Duncan 1986 RJ93
            |           |--N. perkinsi K-P91
            |           `--N. splendida K-P91
            |         Petrochorista [Petrochoristidae] RJ93
            |           |--P. elegantula RJ93
            |           `--P. prona RJ93
            |         Protopanorpa [Protopanorpidae] RJ93
            |           `--P. permiana RJ93
            |         Tomiochorista [Tomiochoristidae] RJ93
            |           `--T. minuta RJ93
            |         Triassochorista [Triassochoristidae] RJ93
            |           `--T. nana RJ93
            |         Xenochoristidae RJ93
            |           |--Xenochoristella hillae RJ93
            |           `--Xenochorista F71
            |                |--X. sobrina Riek 1953 F71
            |                `--X. splendida Riek 1953 F71
            |--Aneuretopsychina RL09
            `--+--+--+--Bittacidae RL09
               |  |  `--Robinjohnia F71 [Robinjohniidae RL09]
               |  |       `--R. tillyardi Martynov 1948 F71
               |  `--+--Boreidae RL09
               |     `--Nannochoristidae RL09
               |          |--Itaphlebia multa N02
               |          |--Nannochorista neotropica B91, GE05
               |          `--Microchorista philpotti B91, WT11 [=Nannochorista philpotti GE05]
               `--+--+--+--Englathaumatidae N02
                  |  |  `--Dinopanorpa RJ93 [Dinopanorpidae RL09]
                  |  |       --D. megarche RJ93
                  |  `--+--Parachorista N02 [Parachoristidae RL09]
                  |     |    |--P. bairdae (Tillyard 1922) [=Archipanorpa bairdae] F71
                  |     |    |--P. comica N02
                  |     |    |--P. pincombeae Tillyard 1926 [incl. P. warnerensis Tillyard 1926] F71
                  |     |    `--P. splendida Tillyard 1926 F71
                  |     `--+--Eomeropidae [Notiothaumatidae] RL09
                  |        |    |--Notiothauma reedi GE05
                  |        |    |--Eomerope GE05
                  |        |    `--Pronotiothauma neuropteroides RJ93
                  |        `--+--Thaumatomeropidae RL09
                  |           `--Meropeidae RL09
                  |                |--Merope tuber B91
                  |                `--Austromerope poultoni B91
                  `--+--Orthophlebiidae [Austropanorpidae] GE05
                     |    |--Austropanorpa australis N02, RJ93
                     |    `--Orthophlebia GE05
                     |         |--O. curta RJ93
                     |         |--O. grandis N02
                     |         |--O. heidemariae N02
                     |         `--O. longicauda N02
                     `--+--+--Holcorpa [Holcorpidae] GE05
                        |  |    `--H. maculosa GE05
                        |  `--+--Panorpidae RL09
                        |     `--Panorpodidae GE05
                        |          |--Brachypanorpa carolinensis GE05, K-P87
                        |          `--Panorpodes brevicauda RJ93
                        `--+--Apteropanorpa B91 [Apteropanorpidae GE05]
                           |    `--A. tasmanica B91
                           `--Choristidae N02
                                |--Neochorista ruficeps B91
                                |--Taeniochorista pallida B91
                                |--Cretacochorista parva Jell & Duncan 1986 RJ93
                                `--Chorista Klug 1836 ICZN03
                                     |--*C. australis Klug 1838 ICZN03
                                     `--C. ruficeps R70

*Type species of generic name indicated

REFERENCES

[B91] Byers, G. W. 1991. Mecoptera (scorpion-flies, hanging-flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 696–704. Melbourne University Press: Carlton (Victoria).

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

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

[ICZN03] ICZN. 2003. Choristidae Verrill, 1882 (Mollusca, Gastropoda): spelling emended to Choristeidae, so removing the homonymy with Choristidae Esben-Petersen, 1915 (Insecta, Mecoptera). Bulletin of Zoological Nomenclature 60 (3): 224–225.

[K-P87] Kukalová-Peck, J. 1987. New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta). Canadian Journal of Zoology 65: 2327–2345.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[N02] Novokshonov, V. G. 2002. Order Panorpida Latreille, 1802. The scorpionflies (=Mecaptera Packard, 1886, =Mecoptera Comstock et Comstock, 1895, +Neomecoptera Hinton, 1958, +Paratrichoptera Tillyard, 1919, +Paramecoptera Tillyard, 1919). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 194–199. Kluwer Academic Publishers: Dordrecht.

[RL09] Ren, D., C. C. Labandeira, J. A. Santiago-Blay, A. Rasnitsyn, C. Shih, A. Bashkuev, M. A. V. Logan, C. L. Hotton & D. Dilcher. 2009. A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies. Science 326: 840-847.

[R70] Riek, E. F. 1970. Mecoptera (scorpion-flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 636–646. 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.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated: 24 April 2022.

Ceratophyllidae

Male hen flea Ceratophyllus gallinae, copyright Katja ZSM.

Belongs within: Siphonaptera.

The Ceratophyllidae are a Holarctic group of fleas found on both mammals and birds (Dunnet & Mardon 1991).

<==Ceratophyllidae
    |--Dasypsyllus gallinulae GE05
    |--Nosopsyllus DM91
    |    |--N. fasciatus KK54
    |    |--N. laeviceps S00
    |    `--N. londiniensis DM91
    `--Ceratophyllus [Ceratophyllinae] DM91
         |  i. s.: C. borealis A71
         |         C. fasciatus RD77
         |         C. garei A71
         |         C. rossittensis A71
         |         C. silantiewi K01
         |         C. styx A71
         |         C. tesquorum K01
         |         C. tundrensis A71
         `--C. (Ceratophyllus) S76
              |--C. (C.) fringillae (Walker 1856) S76
              |--C. (C.) gallinae S76
              `--C. (C.) phrillinae Smit 1976 S76

*Type species of generic name indicated

REFERENCES

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

[DM91] Dunnet, G. M., & D. K. Mardon. 1991. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 705–716. Melbourne University Press: Carlton (Victoria).

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

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[KK54] Klemmer, K., & H. E. Krampitz. 1954. Zur Kenntnis der Säugetierfauna Siziliens. Senckenbergiana Biologica 35 (3–4): 121–135.

[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.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2^nd ed. CABI Publishing: Wallingford (UK).

[S76] Smit, F. G. A. M. 1976. A new bird-flea from Nepal (Siphonaptera). Senckenbergiana Biologica 57 (1–3): 63–66.

Malacopsylloidea

Rhopalopsyllus sp. in Dominican amber, from Poinar (2014).

Belongs within: Siphonaptera.

The Malacopsylloidea are a group of fleas mostly found on Neotropical rodents with the exception of the genus Parapsyllus found on sub-Antarctic seabirds (Dunnet & Marson 1991).

<==Malacopsylloidea DM91
    |--Malacopsyllidae GE05
    `--Rhopalopsyllidae DM91
         |  i. s.: Parapsyllus DM91
         |           |--P. australiacus DM91
         |           |--P. magellanicus A71
         |           `--P. taylori D70
         `--Rhopalopsyllinae L07
              |--Rhopalopsyllus GE05
              `--Polygenis L07
                   |--P. (Polygenis) L07
                   |    |--P. (P.) axius (Jordan & Rothschild 1923) L07
                   |    |--P. (P.) bohlsi (Wagner 1901) L07
                   |    `--P. (P.) platensis (Jordan & Rothschild 1908) L07
                   `--P. (Neopolygenis) L07
                        |--P. (N.) atopus (Jordan & Rothschild 1922) L07
                        `--P. (N.) massoiai Del Ponte 1967 L07

*Type species of generic name indicated

REFERENCES

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

[D70] Dunnet, G. M. 1970. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 647–655. Melbourne University Press.

[DM91] Dunnet, G. M., & D. K. Mardon. 1991. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 705–716. Melbourne University Press: Carlton (Victoria).

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

[L07] Lareschi, M. 2007. Seasonal occurrence of ectoparasites associated with the water rat Scapteromys aquaticus (Muridae, Sigmodontinae) from Punta Lara, Argentina. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 233–240. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

Pygiopsyllidae

Bandicoot flea Pygiopsylla hoplia, copyright R. E. Pugh.

Belongs within: Siphonaptera.

The Pygiopsyllidae are a group of fleas that is most diverse in Australasia, with further representatives in tropical Asia and South America. The majority of species feed on marsupials and rodents with a smaller number found on birds (Dunnet & Mardon 1991).

<==Pygiopsyllidae DM91
    |  i. s.: Austropsylla DM91
    |--Lycopsyllinae DM91
    |    |--Lycopsylla nova DM91
    |    |--Uropsylla tasmanica DM91
    |    |--Bradiopsylla echidnae DM91
    |    `--Choristopsylla ochi DM91
    |--Stivaliinae DM91
    |    |--Parastivalius novaeguineae DM91
    |    |--Stivalius rectus D70
    |    `--Metastivalius DM91
    |         |--M. molestus DM91
    |         `--M. rectus DM91
    `--Pygiopsyllinae DM91
         |--Notiopsylla kerguelensis DM91
         |--Geohollandia solida DM91
         |--Wurunjerria warnekei DM91
         |--Hoogstraalia vandiemeni DM91
         |--Acanthopsylla DM91
         |    |--A. jordani DM91
         |    `--A. scintilla D70
         |--Bibikovana DM91
         |    |--B. rainbowi DM91
         |    `--B. tiptoni DM91
         `--Pygiopsylla DM91
              |--P. hoplia DM91
              |--P. phiola DM91
              `--P. rainbowi D70

*Type species of generic name indicated

REFERENCES

[D70] Dunnet, G. M. 1970. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 647–655. Melbourne University Press.

[DM91] Dunnet, G. M., & D. K. Mardon. 1991. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 705–716. Melbourne University Press: Carlton (Victoria).

Pulicidae

Oriental rat flea Xenopsylla cheopis, from here.

Belongs within: Siphonaptera.

The Pulicidae is a family of fleas including some species of the most significance for humans, such as the human flea Pulex irritans, the dog and cat fleas Ctenocephalides spp., and the Oriental rat flea Xenopsylla cheopis. Pulex irritans is the only flea species known to habitually parasitise a primate. Xenopsylla cheopis is notorious as the primary vector for the plague bacterium Yersinia pestis. Females of the chigoe flea or jigger Tunga penetrans burrow into the skin of mammals including humans, leaving only the tip of the abdomen exposed for excretion, mating and laying eggs.

Characters (from Brinck-Lindroth & Smit 2007): Terga II-VII each with only one row of setae. Mesonotum without pseudosetae under collar; metanotum and terga without marginal spinelets or ctenidia. Head without anterior tentorial rod. Outer internal ridge of mid coxa absent; hind coxa with a row or group of small spiniform setae on lower part of inner side; hind tibia without an apical tooth on outside. No setae above spiracular fossa of tergum VIII.

<==Pulicidae [Pulicoidea]
    |--Tunga [Tungidae, Tunginae] GE05
    |    `--T. penetrans GE05
    `--Archaeopsyllini KP19
         |--Archaeopsylla erinacei A71
         `--Ctenocephalides KP19
              |--C. canis DM91
              `--C. felis (Bouché 1835) H82

Pulicidae incertae sedis:
  Hoplopsyllus anomalus TW05
  Typhlopsylla typhlus K01
  Spilopsyllus cuniculi DM91
  Pulex Linnaeus 1758 L58
    |--P. echidnae O86
    |--P. fasciatus Bosc 1800 B00
    |--P. irritans Linnaeus 1758 L58
    |--P. (Pulex) larimerius R02
    `--P. penetrans Linnaeus 1758 L58
  Echidnophaga Olliff 1886 O86
    |--*E. ambulans Olliff 1886 O86
    |--E. eyrei DM91
    |--E. gallinacea GE05
    |--E. murina KK54
    |--E. myrmecobii DM91
    |--E. octotricha DM91
    `--E. perilis W91
  Xenopsylla GE05
    |--X. astia A71
    |--X. bechuanae SEC08
    |--X. brasiliensis SEC08
    |--X. cheopis GE05
    |--X. cornigera SEC08
    |--X. cryptonella SEC08
    |--X. erilli SEC08
    |--X. hamula SEC08
    |--X. nubica SEC08
    |--X. philoxera SEC08
    |--X. piriei SEC08
    |--X. scopulifer SEC08
    |--X. trifaria SEC08
    |--X. versuta SEC08
    `--X. vexabilis DM91
         |--X. v. vexabilis D70
         `--X. v. meseris D70

*Type species of generic name indicated

REFERENCES

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

[B00] Bosc, C. 1800. Description d'une nouvelle espèce de puce (Pulex fasciatus). Bulletin des Sciences, par la Societé Philomathique de Paris 2 (44): 156.

[D70] Dunnet, G. M. 1970. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 647–655. Melbourne University Press.

[DM91] Dunnet, G. M., & D. K. Mardon. 1991. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 705–716. Melbourne University Press: Carlton (Victoria).

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

[H82] Helmore, D. W. 1982. Drawings of New Zealand insects. Bulletin of the Entomological Society of New Zealand 8: 1–52.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[KK54] Klemmer, K., & H. E. Krampitz. 1954. Zur Kenntnis der Säugetierfauna Siziliens. Senckenbergiana Biologica 35 (3–4): 121–135.

[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.

[O86] Olliff, A. S. 1886. Description of a new aphanipterous insect from New South Wales. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 171–172.

[R02] Rasnitsyn, A. P. 2002. Order Pulicida Billbergh, 1820. The fleas (=Aphaniptera). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 240–242. Kluwer Academic Publishers: Dordrecht.

[SEC08] Shihepo, F., S. Eiseb & P. Cunningham. 2008. Fleas (Insecta: Siphonaptera) associated with small mammals in selected areas in northern Namibia. Journal Namibia Wissenschaftliche Gesellschaft 56: 5–23.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240–257.

[W91] Waterhouse, D. F. 1991. Insects and humans in Australia. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 221–235. Melbourne University Press: Carlton (Victoria).

Last updated: 24 April 2022.

Siphonaptera

Stephanocircus sp., from here.

Belongs within: Mecopteroidea.
Contains: Ctenophthalmidae, Pulicidae, Malacopsylloidea, Ceratophyllidae, Pygiopsyllidae, Hystrichopsyllidae.

The Siphonaptera are the fleas, wingless parasites of birds and mammals. Modern fleas are laterally flattened, with strong jumping abilities. Larvae are vermiform and usually free-living in the nest or burrow of the host, feeding on organic debris. The Cretaceous Tarwinia australis is well supported as a stem flea; the Early Cretaceous Saurophthirus longipes, suggested to be a stem flea by Rasnitsyn (2002), is of more uncertain relationships. Saurophthirus longipes was a large (12 mm in length), long-legged insect that has been suggested to be an ectoparasite of pterosaurs. Tarwinia australis was more similar to modern fleas but the hind coxae were smaller; it also possessed ctenidia on the fore and hind tibiae rather than on the thorax (Giribet & Engel 2005). The classification of living fleas remains uncertain with higher taxa often defined by combinations of features rather than well defined synapomorphies (Dunnet & Mardon 1991).

Synapomorphies for Tarwinia and crown Siphonaptera (from Giribet & Engel 2005): Body laterally compressed; wings absent; thorax greatly reduced; antennal flagellum compact; pygidium with patch of sensilla.

Characters of crown Siphonaptera (from Rasnitsyn 2002): Size small. Body streamlined, compressed, apterous, usually with ctenidia (combs of backward-directed spines) on thoracic segments, sometimes also on head and some abdominal segments. Head hardly movable, with two stemmata only. Antenna 13-segmented, flagellum modified into tight club, hidden in groove, used as additional clasping device in male. Mouthparts piercing, with epipharyngeal and lacinial stylets, mandible lost, maxillary palp free, labial palp forming beak sheath. Thoracic nota subequal in length, metapleuron large due to leading role of hind leg in leaping, legs strong, not particularly long, saltatory, with 5-segmented tarsus and large clinging claws. Abdomen 10-segmented, with 10th bearing patch of specialised sensoria (sensilium), and cerci. Male with sternum 9 modified into clasping organ, with gonocoxa and gonostylus normally retained, volsella lost, external aedegus reduced and replaced by large endophallus in its intromittent function. Ovipositor lost. Digestive tract modified to digest blood (salivary pump, small proventriculus furnished with spines [acanthae] and used in disintegrating blood cells, large stomach). Four malpighian tubes. Ovaries neopanoistic. Larva worm-like, legless, 13-segmented, with head eyeless, one-segmented antenna, mandible of chewing type, weakly developed maxilla and labium, 10th abdominal sternum bearing unsegmented appendages possibly of cercal nature. Pupa adecticous, exarate, sometimes with mesothoracic outgrowths, enclosed in loose silk cocoon.

<==Siphonaptera [Aphaniptera, Pulicida]
    |--Saurophthiridae R02
    |    |--Saurophthirus longipes GE05
    |    `--Saurophthiroides mongolicus Ponomarenko 1986 RJ93
    `--+--Tarwinia australis GE05
       `--+--Pulicidae DM91
          |--Malacopsylloidea DM91
          `--Ceratophylloidea DM91
               |--Ceratophyllidae DM91
               |--Pygiopsyllidae DM91
               |--Xyphiopsyllidae DM91
               |--Ancistropsyllidae DM91
               |--Chimaeropsyllidae DM91
               |--Hystrichopsyllidae DM91
               |--Coptopsylla S00 [Coptopsyllidae DM91]
               |    `--C. lamellifer S00
               |--Vermipsylla GE05 [Vermipsyllidae DM91]
               |    `--V. alakurt GE05
               |--Leptopsylla [Leptopsyllidae] DM91
               |    `--L. segnis DM91
               |--Macropsyllidae DM91
               |    |--Macropsylla hercules DM91
               |    `--Stephanopsylla Rothschild 1911 DM91, T13
               |         `--*S. thomasi (Rothschild 1903) [=Stephanocircus thomasi] T13
               |--Stephanocircidae DM91
               |    |--Craneopsylla GE05 [Craneopsyllinae DM91]
               |    |    `--C. minerva GE05
               |    `--Stephanocircinae DM91
               |         |--Coronapsylla Traub & Dunnet 1973 D74
               |         |    `--C. jarvisi DM91
               |         `--Stephanocircus DM91
               |              |--S. dasyuri DM91
               |              `--S. pectinipes M70
               `--Ischnopsyllidae DM91
                    |--Ischnopsyllus octatenus A71
                    |--Lagaropsylla mera DM91
                    |--Coorilla longictena DM91
                    |--Serendipsylla marshalli DM91
                    |--Myodopsylla GE05
                    |    |--M. collinsi BVP96
                    |    `--M. gentilis GE05
                    `--Porribius DM91
                         |--P. bathyllus D70
                         `--P. caminae DM91

Siphonaptera incertae sedis:
  Tarsopsylla A71
  Cediopsylla simplex A71
  Odontopsylla multispinosus A71
  Atyphloceras A71
    |--A. bishopi A71
    `--A. nuperus A71
  Callopsylla waterstoni A71
  Frontopsylla laeta A71
  Megabothris A71
    |--M. quirini A71
    |--M. turbidus A71
    `--M. walkeri A71
  Typhlocerus poppei A71
  Nycteridopsylla A71
  Malaraeus penicilliger A71
  Peromyscopsylla silvatica A71
  Nearctopsylla brooksi A71
  Ornithopsylla laetitiae A71
  Myoxopsylla laverani KK54
  Euhoplopsyllus glacialis GE05
  Niwratia elongata GE05
  Chiastopsylla rossi SEC08
  Macroscelidopsylla albertyni SEC08
  Plocopsylla scotinomi Tipton & Mendez 1966 S69
  Ctenophthalmidae GE05
  Glaciopsyllus antarcticus N91
  Micropsylloides Ewing 1938 E38
    `--*M. jaonis [=Rhadinopsylla jaonis] E38
  Micropsylla E38
  Trichopsylloides Ewing 1938 E38
    `--*T. oregonensis Ewing 1938 E38
  Trichopsylla E38
  Citellophilus tesquorum S00
  Monopsyllus ciliatus S00
    |--M. c. cilatus S00
    `--M. c. protinus S00

*Type species of generic name indicated

REFERENCES

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

[BVP96] Batalla, I. B., M. Vargas & O. J. Polaco. 1996. Arthropods associated with Myotis thysanodes Müller 1897, in San Josecito Cave, Nuevo Léon, México. In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 109–111. Ohio Biological Survey: Columbus (Ohio).

[D70] Dunnet, G. M. 1970. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 647–655. Melbourne University Press.

[D74] Dunnet, G. M. 1974. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 91. Melbourne University Press.

[DM91] Dunnet, G. M., & D. K. Mardon. 1991. Siphonaptera (fleas). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 705–716. Melbourne University Press: Carlton (Victoria).

[E38] Ewing, H. E. 1938. Two new genera and one new species of Siphonaptera. Proceedings of the Entomological Society of Washington 40 (4): 93–95.

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

[KK54] Klemmer, K., & H. E. Krampitz. 1954. Zur Kenntnis der Säugetierfauna Siziliens. Senckenbergiana Biologica 35 (3–4): 121–135.

[M70] Mackerras, I. M. 1970. Evolution and classification of the insects. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 152–167. Melbourne University Press.

[N91] Norris, K. R. 1991. General biology. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 68–108. Melbourne University Press: Carlton (Victoria).

[R02] Rasnitsyn, A. P. 2002. Order Pulicida Billbergh, 1820. The fleas (=Aphaniptera). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 240–242. Kluwer Academic Publishers: Dordrecht.

[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.

[SEC08] Shihepo, F., S. Eiseb & P. Cunningham. 2008. Fleas (Insecta: Siphonaptera) associated with small mammals in selected areas in northern Namibia. Journal Namibia Wissenschaftliche Gesellschaft 56: 5–23.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2^nd ed. CABI Publishing: Wallingford (UK).

[S69] Steyskal, G. C. 1969. The mistreatment of the Latin genitive case in forming names of parasites. Systematic Zoology 18 (3): 339–342.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

Last updated: 24 April 2022.