Revision of Drusinae subfamily (Trichoptera, Limnephilidae): divergence by paraproct and paramere: speciation in isolation by integration

Abstract

In the last few years we have described over 70 new incipient sibling limnephild species applying the discovered Trichoptera speciation traits of the paraproct and paramere for species recognition and delimitation. In this revision on Drusinae subfamily, comprising 177 species, we have applied these subtle, but rapid and stable speciation traits and described 49 new sibling species from the “well studied” European mountain ranges. Discussing the theoretical background we have elaborated and adapted a new character state ranking system of phenomics to revise the long-neglected taxonomy of the Drusinae subfamily and synonymised the Cryptothrix, Monocentra, Metanoea, Leptodrusus, Anomalopterygella, Hadimina genera with the Drusus genus. These old genera of artificial constructs were established exclusively by divergences of secondary sexual traits known already to have only species level ranking value. According to our new character ranking system in the Drusinae subfamily, beside the Drusus genus, only the Ecclisopteryx genus has been retained having robust generic level divegences of paraproct loss and ancestral duplication of spine organising centre on the paramere pattern. Speciation trait function of the peg-packed surface on the paraproct head in Drusus genus moved to the gonopod apices and integrated into variously shaped stimulatory organ in the Ecclisopteryx genus. In the Drusus genus the ancestral divergence of the single spine organising centre has integrated 11 species groups with remarkably stable paramere spine pattern. Based upon ancestral divergences in the paraproct architecture we have differenciated 28 species complexes inside the 11 species groups. The delineation of the 163 mostly incipient siblings species, inside the 28 species complexes with 44 new Drusus species, was based primarily on the divergences of speciation trait, that is in the stimulatory head shape of the apical arms on the dorsal branches of the paraproct. In the Ecclisopteryx genus with 14 species we have established two independent lineages both with a single species, as well as two species complexes with five new species applying the speciation trait of the genus, that is the shape divergence of the stimulatory organ on the dorsoapical surface of the gonopods. Based on the Darwinian natural selection, we do not understand how the discovered 70+49 new European incipient phylogenetic species of limnephilid caddisflies have been evolved in the isolated sky island habitats of high mountain ranges. This isolation induced speciation represents a challenge to the mechanistic reductionist concept of the natural selection. Our first trial to extract information from various disciplines to answer this question is presented in a brief theoretical discourse: (1) rethinking the status of natural selection towards postdarwinism; (2) teleology or teleonomy; (3) limits and potentials in understanding reality; (4) organisation of universe by integration; (5) what are and how the organising forces are powered to work in the emerging energy mechanisms; (6) divergence by integration; (7) divergence in isolation; (8) reproductive isolation by sexual selection; (9) shape divergence; (10) speciation traits; (11) generic ranking characters. Assessing the limits and potentials, the humility and hybris attitudes towards understanding reality, we hypothesise an integrative power of organisation, instead of simplistic natural selection, that works both in isolated and sympatric populations to maintain the integer state of the integrated autonomic entities against mutations, permutations, transmutations and perturbations. The permanent flux of these adverse, neutral or advantegous external and internal effects generates and gives rise to a responsive and balancing integrative mechanisms with comparative magnitude and multitude. This permanently balancing organisation process develops and drives a large number of interactions along various patterns of supervenient emergent mechanisms in order to integrate them into an oscillating autonomy. In isolation of the sky-islands on high altitudes, the divergences of new taxa are the adaptive autonomic byproducts of this integrative organisation. Integration and not the selection is the real force mechanism that harmonises the concerted flux of stochastic processes into reproductive isolation by sexual selection. The primacy of integration is clearly demonstrated by empirical evidences in the causes and consequences of the integrated speciation traits. Any kind of selection, artificial, natural, social or sexual, is only an emergent perturbating mechanism forced to integrate into the autonomy of an entity. This is how the entanglement of the quantum world and any supervenient integer work together through energeticism of the interactive realism. This is a must in the ontic, epistemic and semantic structural realism.

Keywords

• Speciation trait
• incipient sibling species
• character ranking
• Drusinae
• new species
• humility and hybris
• speciation in sky islands
• organisation versus evolution
• integration versus selection