http://www.collembola.org/taxa/collembo.htm - Last updated on 2014.11.30 by Frans Janssens
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Peter F. Bellinger (), Department of Biology, California State University, Northridge, CA 91330, USA
Kenneth A. Christiansen, Department of Biology, Grinnell College, PO Box V3, Grinnell, IA 50112-0806, USA
Frans Janssens, Department of Biology, University of Antwerp, Antwerp, B-2020, Belgium

Fig.1. Collembola habitus
(photographs 2000-2004 © Hopkin, S. (1);
2002 © Baquero, E. & Jordana, R. (2);
2004,2007 © Henderickx, H. (4);
2003 © Pettersson, B. (5);
2003 © Schoenherr, J. (6);
2004 © Baquero, E. (7);
2004 © Gielen, K. (8);
2004 © Vuijlsteke, M. (9);
2005 © Domene, X. (10);
2005 © Cheung, D. & Schmidt, J. (11);
2005 © Hall, K. (12);
2006-2007 © Stevens, M. (13);
2006 © Maddison, D.R. (14);
2008 © Ng, M. (15);
2008 © Baas, A.H. (16). )
Citation suggested: Bellinger, P.F., Christiansen, K.A. & Janssens, F. 1996-2014.
Checklist of the Collembola of the World. http://www.collembola.org

Introduction Collembola are small ([min. 0.12] 1-5 [max. 17] mm), entognathous (mouthparts, such as mandibulae and maxillae, located within a 'gnathal pouch'), wingless hexapods with antennae always present. Most but not all Collembola may be recognised by a posterior ventral forked abdominal appendage, the furca. The presence of antennae and absence of cerci distinguishes them from the other entognathous hexapods, the Protura (with antennae and cerci absent) and the Diplura (with antennae and cerci or pincers present).
There are ca 8000 described species worldwide. Collembolan fossils from the Devonian (ca 400 million years ago) are among the oldest known records of terrestrial animals. These organisms are virtually ubiquitous in terrestrial systems, ancient and thus, one of the more successful arthropod lineages.

Etymology: Lubbock (1870) proposed for the division of the Thysanura comprised in the Linnaean genus Podura the term Collembola, "as indicating the existence of a projection or mammalia enabling the creature to attach or glue itself to the body on which it stands" (Lubbock, 1873:36) (from colla (Latin), from kolla (Greek): glue; from embolon (Greek): that what has been thrown into something, e.g. a wedge, a ram, a plug; from emballein (Greek): to throw into, to insert).
This ventral projection, the ventral tube or collophore, plays an extremely important role in the fluid and electrolyte balance. The eversible vesicles of the ventral tube may also be used as a source of 'grooming' fluid and for adhering to smooth surfaces (after Hopkin, 1997:48-49). In Anurida, that do not have a furca, the eversible vesicles of the ventral tube may adhere to the surface waterfilm on which they can walk and deform it in such a way that it is springloaded; when the waterfilm is released the animal is launched upwards into the air (Bush & Hu, 2006:351). <!--<a href="../collembo.wav">kol-UM-bo-la</a>.--> <!--<bgsound src="../collembo.wav" loop="1">-->
Fig.2. Schematic diagram of the poduromorph body structure
(lateral view).
Modified after Potapov, M. in Babenko, A. (1988).
External anatomy and morphology: The body of Collembola basically comprises three tagmata, a head capsule, a thorax with three segments, and an abdomen with five segments and a terminal periproct. Thoracic and abdominal segments may be indistinct and may give the body a more globular appearance. antenna mouthparts postantennal organ eye patch profootcomplex protibiotarsus profemur protrochanter procoxa prosubcoxa proepicoxa ventral tube retinaculum manubrium dens mucro mesofootcomplex mesotibiotarsus mesofemur mesotrochanter mesocoxa mesosubcoxa mesoepicoxa metafootcomplex metatibiotarsus metafemur metatrochanter metacoxa metasubcoxa metaepicoxa anal spine protergite mesotergite metatergite periproct head prothorax mesothorax metathorax first abdominal segment second abdominal segment third abdominal segment fourth abdominal segment fifth abdominal segment The head bears two antennae, two optional postantennal organs, two optional composed eyes and the mouthparts. The antennae principally consist of four articulations. Antennal articulations may be subdivided or annulated. Each composed eye consists of maximum eight ommatidia. The mouthparts comprise the labrum, a pair of mandibulae, a pair of maxillae, the hypopharynx and the bipartite labium. The frontal labrum, the ventral labium and two lateral oral folds enclose the other mouthparts in the buccal cavity (entognathy). Each thoracic segment bears ventrally a pair of walking limbs. Each limb is made up of an epicoxa, subcoxa, coxa, trochanter, femur, tibia and footcomplex, the latter comprising the distal part of the tibia having a large outer lamellate unguis and bearing a small inner unguicular tubercle with optional unguiculus or empodium (Janssens, 1999-2010). The anterior abdominal segment bears a ventral tube having two eversible vesicles. The third abdominal segment ventrally bears the retinaculum. The fourth abdominal segment ventrally bears the furca. The furca comprises the basal manubrium, bearing two arms, each of them comprising a dens and a mucro. The genital orifice opens at the ventral side of the fifth abdominal segment. The anus opens terminally at the posterior abdominal periproct. The linea ventralis is a linear cuticular ventral groove that runs between the base of the labium and the collophore (Hopkin, 1997:60).
Some discussion on morphological issues:
Tagmatisation in arthropodans is not monophyletic. Assuming a primitive marine benthic crustacean discovering the potentials of terrestrial habitats, tagmatisation is almost a natural process, since its main effect is the localisation and specialisation of the locomotory system. Improving the locomotory system is imperative for successfully invading terrestrial habitats. On the other hand, cephalisation is just as important. Both processes led to tripartite body division in arthropods living in terrestrial habitats. In marine or freshwater habitats, tripartite tagmatisation has no special advantadge.
Fig.2a. Dicyrtomina saundersi
Left composed eye
2007 © Krebs, C.
Fig.2a1. Sminthurinus sp.
Projected maxillar heads
2012 © Murray, A.
The collembolan composed eye, with maximum 8 single eyes designated as A to H (fig.2a), is derived from the compound eye of early crustaceans (Paulus, 1972).
Entognathy in Hexapoda s.l. is not monophyletic. The collembolan entognathy might be developed as an adaptation to terrestrial habitats. Entognathy in crustaceans is rare. There are some indications of an entognathic tendence in terrestrial Amphipoda. This is interesting, because it might show that entognathy is an evolutionary advantadge during the process of invading terrestrial systems. Amphipoda are a more recent type of crustaceans: oldest fossils are from the Eocene. So, they might be still in the phase where early collembolan ancestors were in the preDevonian times.
The postantennal organs are the remnants of the 2nd pair of antennae of its ancestral crustacean. The postantennal organs might be the specialised sensory organ of the original 2nd antennal apex that remained while the 2nd antenna shaft itself reduced (Lawrence, 1999).

To be completed.

Internal anatomy: introduction.

Fig.2aa. Kiss of death. Entomobrya muscorum predated by Hybotidae. From Czechia. 2007.08.11 © Krásenský, P.
Fig.2ab. Orchesella sp. Spermatophores. From France. 2010.11.22 © Lebeaux, P.
Biology: Development is direct with adults differing from juveniles in proportion, size, pigment (usually juveniles are paler), and the absence of a genital opening (Christiansen in Dindal, 1990:967). In some genera a diapause occurs which may be associated with regressive modification of mouthparts and digestive system and even striking external modification of cuticle and the development of spines (ecomorphosis) (Christiansen in Dindal, 1990:967-968). Collembola moult throughout life with instars ranging from four to more than 50 (Christiansen in Dindal, 1990:968). Collembola are polyphagous, in general; some species are saprophagous (decomposed plants), coprophagous (excrements), necrophagous (cadavers), mycetophagous (fungi), bacteriophagous (soil micro-organisms) (Thibaud, 1970:103) or pollinophagous (pollen). Some are predacious. In Sinella coeca and Sinella pouadensis, the adults eat their own eggs, even when there is enough food supply (Thibaud, 1970:132).
Collembola have separate sexes and indirect sperm transfer (Hopkin 1997:134). Spermatophores (fig.2ab) are deposited by the males on the substrate (Christiansen in Dindal, 1990:968), or placed directly on the female genital opening (Hopkin 1997:134). A variety of mechanisms have evolved to ensure successful 'capture' of this spermatophore by the female (Christiansen in Dindal, 1990:968; Hopkin 1997:134).
Predators: are represented by species of Chilopoda, Opilionidae, Japygidae, Arachnida such as Acari (Erythraeoidea from Lithuania, Bdellidae from the UK, Undefined mite from the UK), Aranea (e.g. Salticidae such as Hentzia palmarum or Gen. spec. from the UK or Gen. spec. from the USA, or Ballus chalybeius from the UK, and Linyphiidae), Pseudoscorpiones (after Thibaud, 1970:105); furthermore by Insecta such as Hemiptera, Coleoptera larvae, Coleoptera: Pselaphinae, Coleoptera: Staphylinidae, Coleoptera: Rhyzobius litura, Dolichopodidae ( from the UK, from Sweden, from the USA:South Carolina, from USA:Alaska), Hybotidae, and Formicidae.

Physiology: introduction.

Fig.3b. Podura aquatica and Sminthurides aquaticus, typically found on the surface of stagnant freshwaters. 2007.04.01 © Cornwall, N.J.
Ecology: Collembola are soil and litter dwelling, preferring wet or damp surroundings. Collembolans inhabit soil and leaf litter, although some species move actively over the surfaces of bark and flowers in daylight. They may be found in moss, under stones, in caves, in ant nests and termite nests but also in the intertidal zone on the coast, on the surfaces of lakes and ponds or snow fields of gletjers. Collembolans are major components of terrestrial ecosystems (and particularly significant members of the soil communities), constituting a significant proportion of the animal biomass and are thus frequently and easily found. In forest soils they can reach densities of 200 to 1800 individuals per dm3, densities only surpassed by the acarian soil population (Handschin, 1955).
Abiotic factors: In Hypogastruridae, the development is impacted as follows: 1. the lethal temperatures are -4°C and 28°C, 2. the optimum temperature range is 9°C to 12°C, 3. the hygrometric optimum is 98-100% relative humidity; 4. the lethal hygrometric minimun is 93% relative humidity (Thibaud, 1970:161-173).

Paleontology: introduction.

Anthropological perspective: Collembola can be pests principally by virtue of their presence in the home. But in many cases, the Collembola are just annoying 'guests', a nuisance, rather than infestations causing a disease. The infestations are classified as domestic infestations (Collembola found in houses), incidental human infestations (infestations through pot plants in the bedroom, infestations by malfunctioning pooter), human infestations not associated with dermatitis and human infestations associated with dermatitis. In addition, one can also consider the delusional infestations (psychotic infestations) and the infestations due to 'sample contamination' (clinical errors, laboratory errors).

Phylogeny: Handlirsch (1908) considers Collembola as a more or less recent group of insects with an extreme specialisation. He considers them as forms with a retrograde development reaching maturity while in a larval state. (cited from Handschin, 1955:41,45).
Based on the discovery of the ca 400 million years old Devonian fossil collembolan Rhyniella praecursor, and the striking resemblance it shows with extant collembolan species, Tillyard (1928) concludes that Collembola are primary, ancestral, and archaic terrestrial arthropodans (cited from Handschin, 1955:41,49).
Gullan & Cranston (1994:192-194) consider Collembola as the sistergroup of Insecta + Diplura, grouped with Protura into Hexapoda.
Janssens & Lawrence (2002-2012) propose that Collembola are highly specialised terrestrial Crustacea, that have reached their evolutionary climax already in the Devonian, when they dominated most terrestrial habitats. The terrestrial competition between Collembola and early Insecta might have triggered the latter to develop wings to become 'masters in the sky' in the Carboniferous.
A phylogeny, applying the principle of total evidence, using molecular and morphological characters, strongly supports the monophyly of Pancrustacea (= Crustacea & Hexapoda) (Giribet, Edgecombe & Wheeler, 2001:160).
Molecular phylogeny of the arthropods provide support for a monophyletic Hexapoda/Branchiopoda clade (Regier & Shultz, 1997:902,911). Based on mitochondrial data, Lavrov et al. (2004) recover an (Insecta, (Branchiopoda, Malacostraca)) clade and a (Collembola, Maxillopoda) clade, which is confirmed by Cook et al. (2005) (Cook, Yue & Akam, 2005:1301).
Physiological data show that Collembola evolved directly from marine ancestors: haemolymph with high osmotic pressures and mainly composed of inorganic salts (Little, 1983, 1990 cited from D'Haese, 2003:583). So early crustaceans must have been adapted from marine habitats in the Cambrium to terrestrial soil habitats in the Devonian. Possibly, Collembola are derived from a benthic marine maxillopod that explored the potentials of terrestrial soil habitats.

Methods: introduction.

Systematics: The taxonomic hierarchy is mainly based on Bretfeld (1994, 1999), D'Haese (2002:1148), and Deharveng (2004:427). The systematics of the higher taxa that is presented here is in line with some of the more 'recent' opinions. Collembola are not considered as being Insecta but as a taxonomic group with the same rank (class). Note that also Protura and Diplura are currently classified as separate classes.
In an attempt to organise a combination in kind of harmony between two by definition incompatible classification schools - the Linnean school that uses a static, hierarchical system with emphasis on the ranking of taxa and the cladistic school that uses a dynamic, evolutionary system with emphasis on the relationship between the taxa - the classification used here tries to map the more recent cladistic system onto the conventional Linnean classification and ranking system. Note that it will never be possible to combine both systems in a 100% compatible way. In other words: different opinions and thus classifications will continue to popup in the papers...
Hexapoda Blainville, 1816. The finding of the reciprocal paraphyly of Hexapoda and Crustacea suggests an evolutionary scenario in which the acquisition of the hexapod condition may have occurred several times independently in lineages descending from different crustacean-like ancestors, possibly as a consequence of the process of terrestrialisation (Carapelli, Liò, Nardi, van der Wath & Frati, 2007). Although found paraphyletic based on recent molecular studies, Hexapoda is conveniantly maintained in the current taxonomic hierarchy untill the disagreements between molecular and morphological analyses have been resolved.
Apterygota Lang, 1889 (= Archaeognatha, Zygentoma, Diplura, Collembola and Protura) is considered as being an artificial assemblage of paraphyletic taxa (Moen & Ellis, 1984) and therefore not accepted anymore as a valid formal taxon by the cladistic school of systematists (Hopkin, 1997:19) (Bach de Roca, Gaju-Ricart & Compte-Sart, 1999:393).
Ellipura Börner, 1910 (= Collembola and Protura) is not a monophyletic group (Bach de Roca, Gaju-Ricart & Compte-Sart, 1999:393) and therefore not accepted in this classification.

Superregnum Eucarya Woese, Kandler & Wheelis, 1990
Regnum Animalia Linnæus, 1758
Subregnum Eumetazoa Butschli, 1910
Superphylum Ecdysozoa Aguinaldo AMA, Turbeville JM, Lindford LS, Rivera MC, Garey JR, Raff RA & Lake JA, 1997
Phylum Arthropoda Latreille, 1829
Subphylum Pancrustacea Zrzavy & Stys, 1997
Superclassis Hexapoda Blainville, 1816
Classis Collembola Lubbock J, 1870:295 key  key  key  key  key  key habitus lateral
Ceratophysella meets Dicyrtomina
from the UK
2008.01.05 © Valentine., B
Isotomurus palustris and Orchesella villosa
from the UK
2008.01.12 © Tonsbeek, M.
Aggregation of Ceratophysella,
Hypogastrura and Proisotoma from the USA
2008.03.09 © Boeddeker, M.

Ordo Poduromorpha Börner, 1913, sensu D'Haese CA, 2002:1148 keyhabitus lateral
Superfamilia Neanuroidea Massoud Z, 1967:58, sensu D'Haese CA, 2002:1148 key
Familia Neanuridae Börner, 1901, sensu Deharveng L, 2004:424 key
Neanuridae from New Zealand
2006 Minor, M., & Robertson, A. © SoilBugs
Neanuridae from the USA
2009.06.22 © Light, K.

Subfamilia Caputanurininae Lee, 1983 key
Subfamilia Frieseinae Massoud, 1967
Subfamilia Morulininae Börner, 1906
Morulina multatuberculata from the USA
With large morula-like post antennal organ
2009.03.03 © Roffler, D.
Morulina sp. from the USA
2013.07.05 © Brown, M.H.

Subfamilia Neanurinae Börner C, 1901:33, sensu Cassagnau, 1989 key
Neanurinae from Singapore
2007 © Anker, A.
Neanurinae from Croatia
2008.05.25 © Keresztes, G.
Neanurinae from China
2012.10.18 © Deart, Y.

Subfamilia Pseudachorutinae Börner, 1906
Pseudachorutinae from Singapore
On cerianthid in tidal zone
2008.07.06 © Ng, M.
Pseudachorutinae from Japan
2012.04.18 © Miyazaki, Y.
Pseudachorutinae from Japan
2012.04.18 © Miyazaki, Y.

Subfamilia Uchidanurinae Salmon, 1964
Holacanthella paucispinosa from New Zealand
2006 © Stevens, M.
Acanthanura sp. from Tasmania
2007 © Henderickx, H.

Familia Brachystomellidae Stach, 1949 key
Brachystomella parvula from the UK
2012.06.11 © Murray, A.
Brachystomellidae from Tasmania
2014.04.11 © Murray, A.

Superfamilia Poduroidea sensu Palacios-Vargas, 1994:409
Familia Poduridae Latreille, 1804, i.s.
Podura aquatica from Belgium
2001 © Hopkin, S.P.
Podura aquatica from Belgium
2006.03.26 © De Wilde, A.

Superfamilia Hypogastruroidea Salmon JT, 1964:103, sensu Deharveng L, 2004:427 key
Familia Hypogastruridae Börner, 1906 key
Hypogastrura sp. from the USA
2008.02.12 © Boeddeker, M.
Hypogastruridae from Peru
2009.01-03.dd © Damiano Palomino., B
Hypogastrura sp. from the USA
2009.02.26 © Cowen, R.
Hypogastruridae from the USA
Short furcula with broad manubrium
2012.03.21 © Kouri, J.

Familia Paleotullbergiidae Deharveng L, 2004:427

Superfamilia Gulgastruroidea
Familia Gulgastruridae Lee B-H & Thibaud J-M, 1998:453

Superfamilia Onychiuroidea sensu D'Haese CA, 2002-2003 key
Familia Onychiuridae Lubbock, 1867 key
Subfamilia Onychiurinae Börner, 1901 key
Onychiurinae from China
Pigment and eyes absent
2008.07.17 © NCode, A.
Onychiurinae from Canada
Pigment and eyes absent
2009.01.09 © Weeta, W.
Onychiurinae from Greece
Ventral furca absent
2012.11.14 © Henderickx, H.
Onychiurinae from France
Cave species
2014.05.07 © Alonso, C.

Subfamilia Tetrodontophorinae Stach, 1954 key
Tetrodontophora bielanensis from Germany
2004 © Hopkin, S.P.

Subfamilia Lophognathellinae Stach, 1954 key

Familia Tullbergiidae Bagnall RS, 1935:238 key  key
Tullbergiidae from the USA
2012.02.16 © Kouri, J.
Tullbergiidae from the UK
2012.10.02 © Murray, A.

Familia Isotogastruridae Thibaud J-M & Najt J, 1992:545 i.s.
Isotogastruridae from South Africa
2011 © Deharveng, L.

Familia Pachytullbergiidae Stach, 1954
Familia Odontellidae Massoud, 1967 key
Odontellidae from the USA
2006 © Bernard, E.
Odontella sp. from the USA
2006 © Bernard, E.

Ordo Entomobryomorpha Börner, 1913, sensu Soto-Adames FN et al., 2008:501 keyhabitus lateral
Superfamilia Tomoceroidea Szeptycki A, 1979:112 key
Tomocerus minor from the UK
Large rounded body scales
2009.09.22 © Robertson, A.

Familia Oncopoduridae Carl J & Lebedinsky J, 1905:565 key
Oncopodura sp. nov. from Belgium
2005 © Janssens, F.
Oncopodura crassicornis from the UK
Abd.4 subequal to abd.3
2013.11.30 © Murray, A.

Familia Tomoceridae Schäffer, 1896 key
Pogonognathellus sp. from the UK
Body covered with iridescent scales
2007.12.19 © Campbell, A.
 
Tomocerus minor from the UK
3rd abdominal segment superequal to 4th
2008.09.24 © Robertson, A.

Superfamilia Isotomoidea Szeptycki, 1979:112, sensu Soto-Adames FN et al., 2008:504 key
Familia Isotomidae Schäffer, 1896 key
Isotomidae from Portugal
2012.10.16 © Murray, A.

Subfamilia Proisotominae Stach, 1947
Proisotominae from Tasmania
2014.04.08 © Murray, A.
Proisotominae from Tasmania
2014.04.16 © Murray, A.

Subfamilia Anurophorinae Börner C, 1901:42
Subfamilia Isotominae Schäffer, 1896
Isotominae from Belgium
2006 © Vuijlsteke, M.
Isotoma viridis from the USA
2009.03.10 © Cowen, R.

Subfamilia Pachyotominae Potapov MB, 2001:18

Familia Actaletidae Börner, 1902, sensu Soto-Adames FN et al., 2008:506 key
Actaletidae from Mexico
2002 © Palacios-Vargas, J.G.

Familia Protentomobryidae Folsom, 1937, -

Superfamilia Entomobryoidea Womersley, 1934, sensu Soto-Adames FN et al., 2008:502 key
Familia Microfalculidae Massoud & Betsch, 1966
Familia Praentomobryidae Christiansen, KA et Nascimbene, P, 2006:354,- key
Familia Entomobryidae Schäffer, 1896 key  keyhabitus lateral habitus lateral (head)
Orchesella cincta & Lepidocyrtus curvicollis from France
2012.06.25 © Quintin, C.

Subfamilia Capbryinae Soto-Adames FN, Barra J-A, Christiansen K & Jordana R, 2008:508
Subfamilia Orchesellinae Börner C, 1906:162, sensu Szeptycki A, 1979:115
Orchesella sp. from Estonia
2006.10.06 © Tartes, U.
Orchesella cincta from the UK
With subdivided 2 basal antennomeres
2008.09.21 © Robertson, A.

Subfamilia Entomobryinae Schäffer, 1896, sensu Szeptycki A, 1979:115
Entomobryinae from the UK
Entomobrya nicoleti, E. intermedia, E. multifasciata
2005 © Brocklehurst, K.
Entomobryinae from the USA
Entomobrya griseoolivata
E. assuta, E. cf. nigrocincta
2006 © McClarin, J.

Subfamilia Lepidocyrtinae Wahlgren E, 1906:67, sensu Szeptycki A, 1979:115
Lepidocyrtinae from Sweden
2006.03.25 © Hall, K.
Lepidocyrtinae from Singapore
2014.01.22 © Bay, N.

Subfamilia Seirinae Yosii R, 1961, sensu Szeptycki A, 1979:115
Seirinae from Vietnam
2014.01.17 © Bertner, P.
Seirinae from Malaysia
With extended furca; left dens lost
2014.02.08 © Bertner, P.

Subfamilia Willowsiinae Yoshii R & Suhardjono YR, 1989:35, sensu Janssens F, 2008 key  key  key
Willowsia buski from France
2007.06.09 © Lebeaux, P.
Willowsia platani from the UK
2007.06.30 © Cornwall, N.J.

Familia Paronellidae Börner, 1913, sensu Soto-Adames FN et al., 2008:507 key
Subfamilia Paronellinae Börner, 1913, sensu Soto-Adames FN et al., 2008:507 key  key
Paronellinae from China
2008.07.18 © NCode, A.
Paronellinae from Taiwan
2009.02.04 © Wu, S.
Paronellinae from Taiwan
2010.06.23 © Chien, H.-C.
Paronellinae from Australia
2012.06.23 © Kathy & Marissa.

Subfamilia Cyphoderinae Börner, 1913, sensu Soto-Adames FN et al., 2008:507 key
Cyphoderus albinus from the UK
With Myrmica rubra
2007 © Cornwall, N.J.
Cyphoderus albinus from France
With Pheidole pallidula
2008.03.15 © Lebeaux, P.

Familia Oncobryidae Christiansen, KA et Pike, E, 2002:167,-
Superfamilia Coenaletoidea Soto-Adames FN et al., 2008:506
Familia Coenaletidae Bellinger PF, 1985:117
Coenaletes caribaeus
2006 © Palacios-Vargas, J.G.

Ordo Neelipleona Massoud Z, 1971:198
Familia Neelidae Folsom JW, 1896:391
Neelus? sp.
2005 © Cheung, D. & Schmidt, J.
Megalothorax? sp. nov. from the UK
2006 © Brocklehurst, K.
Neelidae from the UK
2008.04.13 © Kilford., B
Neelidae from Hungary
2010.03.09 © Pfliegler, W.

Ordo Symphypleona Börner, 1901, sensu Massoud, 1971habitus lateral
Symphypleona from Holland
Sminthurinus aureus & Dicyrtoma fusca
2013.03.03 © van Duinen, J.

Superfamilia Sminthuridoidea sensu Fjellberg A, 1989:133
Familia Mackenziellidae Yosii, 1961
Mackenziella psocoides ♂ from Tenerife
After Fjellberg, A, 1989 Fig.15
Mackenziella psocoides ♀ from Scandinavia
After Fjellberg, A, 2006 Fig.2

Familia Sminthurididae Börner, 1906, sensu Betsch J-M & Massoud Z, 1970:199
Sminthurides aquaticus from Belgium
2000 © Hopkin, S.P.
Sphaeridia serrata from the USA
2006 © Maddison, D.R.

Superfamilia Katiannoidea Bretfeld, 1994
Familia Katiannidae Börner, 1913, sensu Bretfeld G, 1999:13 key
Two new Katiannidae from the UK
2009.06.06 © Ardron, P.A.
Katiannina macgillivrayi from the USA
2009.11.26 © Justis, S.

Familia Spinothecidae Delamare Deboutteville, 1961, sensu Bretfeld, 1994 key
Adelphoderia regina from Tasmania
2014.03.10 © Murray, A.
Adelphoderia regina from Tasmania
With neck organs
2014.03.10 © Murray, A.

Familia Arrhopalitidae Stach, 1956, sensu Bretfeld G, 1999:13 key
Arrhopalites hirtus from the USA
2005 © Cheung, D. & Schmidt, J.
Arrhopalites sp. from the USA
2006 © Bernard, E.

Familia Collophoridae Bretfeld G, 1999:13

Superfamilia Sturmioidea Bretfeld, 1994
Familia Sturmiidae Bretfeld, 1994
Sturmius sp. nov. from Panama
2009.03.16 © Palacios-Vargas, J.G.
Sturmius sp. nov. from Panama
2009.03.16 © Palacios-Vargas, J.G.

Superfamilia Sminthuroidea Bretfeld, 1994
Familia Sminthuridae Lubbock, 1862, sensu Deharveng, L, 2004:427habitus lateral
Subfamilia Sminthurinae Lubbock, 1862, sensu Deharveng, L, 2004:427
Sminthurinae from Russia
2006 © Macroclub.ru.
Sminthurus sp. nov. from the USA
2007.04.30 © Cowen, R.

Subfamilia Sphyrothecinae Betsch J-M, 1980:149
Neosminthurus richardsi from the USA
2008.01.26 © Gross, J.

Familia Bourletiellidae Börner, 1912, sensu Bretfeld, 1994
Deuterosminthurus sp. from France
Courtship dance
2010.06.04 © Elpachato.
Bourletiella sp. from Australia
2012.07.03 © Hort, J.

Superfamilia Dicyrtomoidea Bretfeld, 1994
Familia Dicyrtomidae Börner, 1906, sensu Deharveng, L, 2004:427
Dicyrtomina minuta f. ornata from the USA
Abdominal tubular wax excretions
2008.11.15 © Justis, S.
Dicyrtomina minuta f. ornata & Ptenothrix renateae from the USA
2009.12.07 © Bradford, A.

Subfamilia Ptenothricinae Richards, 1968
Ptenothrix sp. from the USA
Two unpaired facial setae
2010.01.17 © Ten Eyck, C.
Ptenothrix sp. from the USA
Apical part of 3rd antennal segment annulated
2010.01.17 © Ten Eyck, C.

Subfamilia Dicyrtominae Richards, 1968
Calvatomina nr superba from the UK
More than 2 unpaired facial setae
2010.01.23 © Robertson, A.
Dicyrtominae from the UK
Dicyrtomina ornata and Dicyrtoma fusca
2012.01.21 © Barton, T.
Dicyrtomina sp. from New Zealand
2014.02.25 © Murray, A.
Dicyrtominae juv. from the UK
Dorsally positioned gut; note dark gut contents
2014.08.15 © Phillips, E.


Familia ludens

Familia Fuzzballidae Janssens, 2006
Genus Pareidolia Janssens, 2008
Species ramosi Janssens, 2008
Pareidolia ramosi from the USA
2006.02.06 © Ramos, K.

Species bloombergi Janssens, 2013
Pareidolia bloombergi from the USA
2013.04.15 © Bloomberg, M.

Discussion of Ordinal Phylogenetic Relationships

Fig.4. Tentative ordinal phylogenetic relationships
	  +----------- "protoCollembola" (+)
     +----+
     |    +------------ Poduromorpha
<-pC-+
     |    +------------ Tomoceroidea
     |    |
     +-Nc-+    +------- Neelipleona
	  +-Pe-+
	       |  +---- "Entomobryomorpha" 
	       +1-+
		  |  +- Coenaletidae
		  +2-+
		     +- Symphypleona

This ordinal tree (Fig. 4) is a compromise compilation of the views of relationships among orders of Collembola based on phylogenies proposed by Cassagnau (1971), Massoud (1971, 1976) Moen & Ellis (1984), Bretfeld (1986), Fjellberg (1994), Soto-Adames (1996), D'Haese (2002, 2003), Park (2002), Deharveng (2004), Gao & al. (2008), Xiong & al. (2008), and Schneider & al. (2011). Traditionally, the Collembola have been divided into five groups (Poduromorpha, Metaxypleona, Neelipleona, Entomobryomorpha, and Symphypleona) which different authors have considered to represent orders, sections or every category in between these two. D'Haese (2002) and Xiong & al. (2008) considered Entomobryomorpha as paraphyletic and D'Haese (2002:1148) proposed Tomoceromorpha (=Tomoceroidea) as a new basic group of Collembola. Schneider & al. (2011) confirmed 1. the monophyly of Neelipleona and 2. that Neelipleona are not closely related to Symphypleona. We consider Neelipleona as the basal apomorph sistergroup of the remaining Entomobryomorpha + Symphypleona. We propose Coenaletidae (= Entomobryomorpha with grasping antennae) as extant representatives of transient forms between Entomobryomorpha and Symphypleona (via Sminthuridoidea = Mackenziellidae + Sminthurididae (= Symphypleona with grasping antennae)) and consider Coenaletidae as the sistergroup of Symphypleona.

Acknowledgements

We would like to thank Louis Deharveng, Steve Hopkin and Toby Barton for their constructive comments.

References