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). The Stephanocircidae, helmet fleas, are found in Australia and South America (Dunnet & Mardon 1991) and have the front of the head modified into a vertical ‘helmet’ surrounded by a conspicuous ctenidium.

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]
| |--Saurophthirus longipesGE05
| `--Saurophthiroides mongolicus Ponomarenko 1986RJ93
`--+--Tarwinia australisGE05
|--CoptopsyllaS00 [CoptopsyllidaeDM91]
| `--C. lamelliferS00
|--VermipsyllaGE05 [VermipsyllidaeDM91]
| `--V. alakurtGE05
|--Leptopsylla [Leptopsyllidae]DM91
| `--L. segnisDM91
| |--Macropsylla herculesDM91
| `--Stephanopsylla Rothschild 1911DM91, T13
| `--*S. thomasi (Rothschild 1903) [=Stephanocircus thomasi]T13
| |--CraneopsyllaGE05 [CraneopsyllinaeDM91]
| | `--C. minervaGE05
| `--StephanocircinaeDM91
| |--Coronapsylla Traub & Dunnet 1973D74
| | `--C. jarvisiDM91
| `--StephanocircusDM91
| |--S. dasyuriDM91
| `--S. pectinipesM70
|--Ischnopsyllus octatenusA71
|--Lagaropsylla meraDM91
|--Coorilla longictenaDM91
|--Serendipsylla marshalliDM91
| |--M. collinsiBVP96
| `--M. gentilisGE05
|--P. bathyllusD70
`--P. caminaeDM91

Siphonaptera incertae sedis:
Cediopsylla simplexA71
Odontopsylla multispinosusA71
|--A. bishopiA71
`--A. nuperusA71
Callopsylla waterstoniA71
Frontopsylla laetaA71
|--M. quiriniA71
|--M. turbidusA71
`--M. walkeriA71
Typhlocerus poppeiA71
Malaraeus penicilligerA71
Peromyscopsylla silvaticaA71
Nearctopsylla brooksiA71
Ornithopsylla laetitiaeA71
Myoxopsylla laveraniKK54
Euhoplopsyllus glacialisGE05
Niwratia elongataGE05
Chiastopsylla rossiSEC08
Macroscelidopsylla albertyniSEC08
Plocopsylla scotinomi Tipton & Mendez 1966S69
Glaciopsyllus antarcticusN91
Micropsylloides Ewing 1938E38
`--*M. jaonis [=Rhadinopsylla jaonis]E38
Trichopsylloides Ewing 1938E38
`--*T. oregonensis Ewing 1938E38
Citellophilus tesquorumS00
Monopsyllus ciliatusS00
|--M. c. cilatusS00
`--M. c. protinusS00

*Type species of generic name indicated


[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 2nd 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 2nd 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 2nd 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.

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