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Table 4 Selection of groundplan characters and apomorphies of Holometabola and of those holometabolan subgroups whose phylogenetic relationships were addressed in this study and whose monophyly was confirmed

From: The evolutionary history of holometabolous insects inferred from transcriptome-based phylogeny and comprehensive morphological data

Taxon Characters
Holometabola •* Larval head orthognathous
•* Larval compound eyes simplified but present
•* Ocelli absent in larvae
•* Larval tentorium X-shaped
•* Retractile larval abdominal prolegs absent
• Larval cerci absent (possible reversal in Strepsiptera [homology uncertain])
•* Adult head orthognathous
• Meso- and metasternum invaginated
• Meso- and metacoxae closely adjacent medially
• Appearance of fully developed compound eyes including external apparatus in the pupal stage (reversal in Strepsiptera)
• External wing buds absent in larval stages (partial reversal in Strepsiptera)
Aparaglossata (Holometabola excluding Hymenoptera) • Larval head prognathous
• Well-developed larval stemmata
• Larval tentorium H-shaped
• Paraglossae vestigial or absent, without muscles
• Ventral sclerites of segment VIII (gonocoxae and gonapophyses) indistinct (reversals within Neuropterida)
Neuropteroidea § (Neuropterida and Coleopterida) • Adult head prognathous or slightly inclined (reversal in Neuroptera)
Megaloptera § • Sensorium on antepenultimate larval antennomere
• Larval salivary duct strongly narrowed, without recognizable lumen
• Setiferous lateral abdominal gills present in larvae
Neuroptera + Megaloptera • Mesothoracic prealare present (also in Amphiesmenoptera)
• Muscular connection between metafurcal arm and epimeral apophysis
• Aquatic larvae (with reversal)
Coleopterida (Coleoptera and Strepsiptera) • Antenna with 9 flagellomeres or less
• Pronotum and propleuron partly or completely connected (also in Diptera)
• Metathorax enlarged, hind wings used as flight organs (posteromotorism)
• Membranous area between mesoscutellum and mesopostnotum present
Mecopterida (Antliophora and Amphiesmenoptera) • Larval dorsal tentorial arm strongly reduced or absent
• Less than 3 larval antennomeres (reversal to 3 in some groups)
• Larval galea and lacinia extensively or completely fused (also missing as separate structures in Neuroptera and Strepsiptera)
• Larval Musculus craniodististipitalis present
Siphonaptera + Mecoptera § • Muscle connecting profurcal arms (Musculus profurca-spinalis) present
• Acanthae of proventriculus close-set, prominently elongated
  1. Plesiomorphic groundplan characters are marked with an asterisk *. For a full list and for apomorphies found for additional subgroups see Additional file 4, Chapter 5. Characters apply to adults if not mentioned otherwise. For groups marked with § behind taxon name, no selection but rather all obtained apomorphies are listed. Groundplan characters and apomorphies were inferred from the morphological datamatrix of Beutel et al. [15] and the interordinal topology of the ML tree of dataset 1 by formal character mapping in Mesquite [33].