principles of systematics and nomenclature general system and phylogeny of insects systematics of Ephemeroptera



What is cladoendesis

Cladoendesis (κλάδος + ἔν-δεσις = "branch-linking") is a method of phylogeny analysis by means of creating a classification, in which each phylogenetic branch is represented by a holophyletic taxon, each non-resolved area of phylogenetic tree is represented by a plesiomorphon, and all taxa (holophyletic and plesiomorphons) are supplied with own names; diagnosis of each taxon is supplied with references to diagnoses of higher taxa, so that characters of all taxa are formulated as hiearachically connected (linked) ones.

Cladoendesis is apposed to matrix methods of phylogeny analysis, in which characters are at first inputted into a rectangular matrix and than calculated by these or that mathematical programs. The matrix methods contradict the modern evolutionary theory (in spite of the fact that this theory is wordily recognized by all apologists of the matrix methods), being not based on any scientific theories.

Cladoendesis is a direct continuation of that traditional method of phylogeny analysis, which had been created much before the evolutionary theory and which had generated the evolutionary theory and the concept of phylogeny. This method is that all data are recorded in a form of classification, in which all characters are attributed to hierarchically subordinated taxa. New data are inputted into the already existent classification, which is expanded and worked out while new information is accumulated, and is corrected while errors are revealed. In order to fulfil this function, classification should be able to hold all newly obtained facts and allow to find them easily. Principles of classification elaborated by Linnaeus and Latreille have partly served solution of this problem: thanks to them, accumulation of data have reached such a high level, which allowed to create the evolutionary theory and to reconstruct that part of phylogeny which we recently known. However, the principles of systematics and nomenclature existent before 1999, limited the classification by a set of artificial absolute ranks, and because of this did not allow to input data about all that hierarchically subordinated phylogenetic branches, which had been revealed. For solution of this problem, I had elaborated a rank-free nomenclature, more exactly a system of two nomenclatures ("dual nomenclature system"), which, being non-contradicting both to the generally accepted International Code of Zoological Nomenclature and to the traditions in usage of names of higher taxa, allows to supply with monosemantic names an unlimited number of taxa in unlimited number of hierarchical levels ( 1999 , Kluge 1999 , 2000, Kluge 2009a, 2010 ).

For more details see Kluge 2012

First usages of the term "cladoendesis"

While cladoendesis is used since 1999, this term appeared for the first time in 2011 in the paper by Kluge N.J. & Novikova E.A. "Systematics of the mayfly taxon Acentrella (Ephemeroptera, Baetidae), with description of new Asian and African species." // Russian Entomological Journal 20 (1): 156, in the followong text on the page 2:

The most comprehensive analysis of mayfly phylogeny is given in the book "The phylogenetic system of Ephemeroptera" (Kluge 2004). Updated electronic version of this monograph is available from "". The method of phylogenetic analysis used there can be named cladoendesis i.e., "branch-coupling" (from Greek κλάδος + ἔν-δεσις). Cladoendesis is based on a specially elaborated rational text layout and a powerful, rank-free, dual nomenclature system. This allows to arrange all characters among taxa, make clear links between characters of subordinated and higher taxa, clarify phylogenetic significance of characters, adequately use in phylogenetic analysis all know apomorphies, and find ways to discover new apomorphies (Kluge 2000, 2004). Cladoendesis is opposed to a non-scientific approach of counting characters and calculating parsimony (which is not compatible with any modern evolutionary theory).

Special explanation of this term is published in the paper by Kluge N.J. 2012. "Cladoendesis and a new look at the evolution of insect metamorphosis". // Entomological Review, 92 (6): 622632 (thranslated from Entomologicheskoe Obozrenie 91(1): 63-78).

Cladoendesis already allowed to do the following:

(non-published achievements are not listed here)

Discover nature of complete metamorphosis.

Kluge N. 2005. Larval/pupal leg transformation and a new diagnosis for the taxon Metabola Burmeister, 1832 = Oligoneoptera Martynov, 1923. // Russian Entomological Journal (2004) 13(4): 189-229. KMK

Kluge N. Cladoendesis and new look to evolution of insect metamorphosis. (in Russian). // Entomologicheskoe Obozrenie, in press. 

Discover metamorphosis of coccid feeding stages.

Kluge N. 2010. Paradoxical molting process in Orthezia urticae and other coccids (Arthroidignatha, Gallinsecta). // Zoosystematica Rossica 19(2): 246-271.

Kluge N. 2012. Cladoendesis and new look to evolution of insect metamorphosis. (in Russian). // Entomologicheskoe Obozrenie 91(1): 63-78.  

Discover homology of paired piercing-sucking mouth apparatus of Birostrata.

Kluge N. 2005. Metamorphosis and homology of mouthparts in Neuropteroidea (Hexapoda: Metabola), with remarks on systematics and nomenclature. // Russian Entomological Journal 14(2): 87-100.

Discover homology of piercing-sucking mouth apparatus of fleas.

Kluge N. 2002. Homology of the mouthparts in fleas (Aphaniptera). // Entomologicheskoe Obozrenie [Revue d'Entomologie de l'URSS] 81(4): 808-816 (in Russian).

Find error in description of Nannomecoptera and discover autapomorpy of Panzygothoraca.

Kluge N. 2004. Larval leg structure of Nannochorista and characteristics of Mecoptera. // Russian Entomological Journal 12(4): 349-354.

Reach some progress on understanding homology of insect male genitals.

Kluge N. 2003. About evolution and homology of genital appendages of insects. // Trudy Russkogo Entomologicheskogo Obshestva / Proceedings of the Russian Entomological Society, 74: 3-16 (in Russian).

Discover synapomorphy of Embioptera and Notoptera and create a new system of Neoptera.

Kluge N.J. 2012. General system of Neoptera with description of a new species of Embioptera. // Russian Entomological Journal 21(4): 371-384.

Kluge N.J. 2013. Additions to the system of neopterous insects (Neoptera). // Proceedings of the Russian Entomological Society. St. Petersburg 84 (2): 5360.

Describe apomorphies of the taxa Calyptroptera and Enteracantha, comprising mecopterans and fleas.

Kluge N.J. 2013. Additions to the system of neopterous insects (Neoptera). // Proceedings of the Russian Entomological Society. St. Petersburg 84 (2): 5360.

Describe apomorphies and elaborate phylogenetic system of Saltatoria, discovere origin of their ovipositors.

Kluge N.J. 2016. Structure of ovipositors and cladoendesis of Saltatoria, or Orchesopia. // Entomological Review 96(8): 10151040.

Describe apomorphies and elaborate phylogenetic system of Parametabola, explaine function of copeognathan mouth apparatus and disprove paleontological erriors.

Kluge N.J. 2019. Cladoendesis of Parametabola and systematic position of the extinct taxon Tetrastigmoptera taxon nov. // Entomological Review 99(3): 362383.

Elaborate the Phylogenetic System of Ephemeroptera.

Kluge N. 2004. The phylogenetic system of Ephemeroptera. // Kluwer Academic Publishers. p. i-xiii + 1-442   Google books

Kluge N. 2007 and next publications - see Publications by N.J. Kluge on Ephemeroptera

Create the first permanent and objective data base Ephemeroptera of the World.

Kluge N. 2009. New version of the database Ephemeroptera of the World as the first experience of a permanent and objective web catalogue in biology. // pp 167-180 In: A.H. Staniczek (ed.): International Perspectives in Mayfly and Stonefly Research (Proceedings of the 12th International Conference on Ephemeroptera and the 16th International Symposium on Plecoptera, Stuttgart 2008) // Aquatic Insects, Volume 31, Supplement 1, p. 1-739. online version

Create the first database on non-typified names of arthropods and higher taxa Nomina Circumscribentia Insectorum.


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