Typified names:

Enteracantha Kluge 2010

NOMEN: Enteracantha Kluge 2010 [N.J. Kluge. Circumscriptional names of higher taxa in Hexapoda.  Bionomina, 2010, No.1: 15–55]. PDF

ORIGINAL LISTED MEMBERSHIP (Kluge 2010): Scorpiomusci (Panorpa/fg) + Nannomecoptera (Nannochorista/fg) + Metamecoptera (Bittacus/fg) + Pleuroptera [= Raphioptera (Boreus/fg) + Aphaniptera (Pulex/fg)]

TYPIFIED NAME IN BASIC FORMAT: Panorpa/fg (incl. Pulex)


MODERN STATUS:  the valid, the oldest name of a holophyletic taxon.

Classification of Enteracantha:

Enteracantha (Panorpa/fg1)
Scorpiomusci (Panorpa/fg2)
Nannomecoptera (Nannochorista/fg1)
Metamecoptera (Bittacus/fg1)
Pleuroptera (Pulex/fg1)
Raphioptera (Boreus/fg1)
Aphaniptera (Pulex/fg2)

Kluge 2010 BioNomina Dual-Nom :

According to the widely accepted idea about flea origin, the order Aphaniptera (= Siphonaptera) is a sister group to the family Boreide, which belongs to the order Mecaptera (= Mecoptera); thus Mecaptera being paraphyletic. It is also generally accepted, that taxa, whose paraphyly is revealed, should be disbanded and substituted by holophyletic taxa; however, the paraphyletic taxon Mecaptera is still in general use, while the holophyletic taxa Mecapera + Aphaniptera and Boreidae + Aphaniptera remain to be unnamed (Zrzavy 2008).

The taxon that consists of the scorpionflies and their relatives, but excludes fleas, has the circumscriptional name Mecaptera Packard 1886. Originally the name Mecaptera was applied to an order that did not include fleas (treated as a separate order, Siphonaptera). The name Mecaptera was generally accepted till the beginning of 20th century (Banks 1907); then Mecoptera Hyatt & Arms 1891, its unjustified emendation and junior circumscriptional synonym, became prevalent.
I propose a new name, Enteracantha Kluge 2010, for the taxon Mecaptera + Aphaniptera. The name refers to the acanthae present in adult proventricula, its autapomorphic character. The typified name of this taxon should be derived from Panorpa rather than Pulex, because the earliest established family-group name is Panorpatae Latreille 1802, not Pulicides Billberg 1820, and the generic name Panorpa Linnaeus 1758 (genus 221 on page 551) has page priority over Pulex Linnaeus 1758 (genus 234 on page 614).

Pleuroptera Kluge 2010 is the name I propose for the taxon consisting of Raphioptera (= Neomecoptera = Boreus/fg1) and Aphaniptera. This name refers to one of autapomorphies of this taxon — partial fusion of wing vestiges with thoracic pleurites. Male Boreus have wings transformed into sclerotized pointed hooks partly fused basally with the pleurite (and thus of limited mobility) and used to restrain the female during mating. In Aphaniptera wings are lost, but remnants of their fusion with pleurites are retained: mesothorax has a pair of separated pleural ridges, hidden under integral external pleural walls; metathorax has tergal flaps, which overlap dorsal parts of pleural ridges (Snodgrass 1946). The Pleuroptera is a part of Enteracantha. 

Classification of Enteracantha other than Pleuroptera is being disputed; above is a possible arrangement.

The taxon Scorpiomusci Kluge 2004 includes Panorpidae, Meropeidae, Choristidae, Notiothaumidae, and allies. Its holophyly is proven by (1) presence of ‘notal organ’ — a clasper formed by outgrowths of abdominal terga 3 and 4 (present in Panorpidae and Notiothaumidae, but lost in Meropeidae and Choristidae); (2) unique structure of male seminal pump; (3) unique structure of female abdominal sterna 8 and 9; (4) unique structure of female abdominal tip (Mickoleit 1975, Willmann 1981, Kluge 2004).

Nannomecoptera Hinton 1981 is group of a few closely related Notogean species. Probably, it is a young group related to Scorpiomusci; claims that the larvae possess archaic features are not sustainable (Kluge 2004).

Metamecoptera Crampton 1930 (= Raptipedia Willmann 1987) is clearly holophyletic. Its relationships with other Enteracantha are being discussed.


Billberg G.J. 1820. Enumeratio insectorum in museo Gust. Joh. Billberg. Typus Gadelianus, 1–138.

Crampton G.C. 1930. The wings of the remarkable archaic mecopteron Notiothauma reedi McLachlan with remarks on their Protoblattoid affinities. // Psyche, 37 (1), 83–103.

Hinton H.E. 1981. Biology of insect eggs. Pergamon Press, vol. 1, 1–473; vol. 2, 475–778; vol. 3, 779–1125.

Hyatt A. & Arms J.M. 1890. Ephemeroptera. In: Guides for Science Teaching, No.VIII. Insecta. Boston Society of Natural History, 69–72.

Kluge N.J. 2004. Larval leg structure of Nannochorista and characteristics of Mecoptera. // Russian Entomological Journal, (2003), 12 (4), 349–354.

Latreille P.A. 1802–1805. Histoire naturelle, générale et particulière des Crustacés et des Insectes. Paris, T.1–14.

Linnaeus C. 1758. Systema Naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum Characteribus, Differentiis, Synonymis, Locis. T.I. [A photographic facsimile of the first volume of the tenth edition. London, British Museum of Natural History, 1956, 1–824].

Mickoleit G. .1975. Die Genital- und Postgenitalsegmente der Mecoptera-Weibchen (Insecta, Holometabola). I. Das Exoskelet. // Zeitschrift für Morphologie der Tiere, 80 (2), 97–135.

Packard A.S. 1886. A new arrangement of the orders of insects. // The American Naturalist, 20 (9), 808.

Snodgrass R.E. 1946. The skeletal anatomy of fleas (Siphonaptera). // Smithsonian Miscellaneous Collections, 104 (18), 1–89, Pl. 1–21.

Willmann R. 1981. Das Exoskelett der männlichen Genitalien der Mecoptera (Insecta). // Zeitschrift für zoologische Systematik und Evolution, 19 (2), 96–150; (3), 153–174.

Willmann R. 1987. The phylogenetic system of the Mecoptera. // Systematic Entomology, 12 (4), 519–524.

Zrzavy J. 2008. Four chapters about the monophyly of insect ‘orders’: A review of recent phylogenetic contributions. // Acta Entomologica Musei Nationalis Pragae, 48 (2), 217–232.