Fonseca, Claudio Ruy Vasconcelos Da, Barbosa, Márcio Luís Leitão & Fernandez, Maria Fernanda Souza, 2011, A hypothetical evolutionary history of passalid beetles narrated by the comparative anatomy of the hindgut (Coleoptera: Passalidae), Zootaxa 3012, pp. 1-20: 12-13
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The Aulacocyclinae forms a natural group recognized by the early authors of Passalidae taxonomy ( Kaup 1868; Kuwert 1891, 1896, 1897; Gravely 1914, 1918; Dibb 1933; Reyes-Castillo 1970). In this analysis, it was possible to find three pairs of heterobatmy (levels of evolutionary development for parts of the body) to support the hypothesis of monophyly:
[Character 3] Long diverticula either not protruding (0) or protruding from the same level around the proximal region of the ileum (1). Fast optimization (ci = 100; ri = 100) ( Fig. 1View FIGURE 1 D).
This character state was synapomorphic.
[Character 4] Ileum without conspicuous segmentation (0), or with conspicuous segmentation (1). Fast optimization (ci = 100; 100).
Although segmentation also occurred in Solenocyclinae ( Figs 2View FIGURE 2 E, F), it was conspicuous only in Aulacocyclinae (i.e., there were strong, ring-like constrictions) ( Fig. 1View FIGURE 1 D). Therefore, this state found in Aulacocyclinae was considered synapomorphic.
[Character 6] Colon not striated (0); with longitudinal striate (1). Unambiguous optimization (ci = 100; ri = 100).
This character state (1) was synapomorphic for this level of hierarchy.
Branch Solenocyclinae +
The branch Solenocyclinae+ is supported by one synapomorphy:
[Character 5] Ileum with a smooth external surface (0); with a series of protuberances covering the external surface (1). Unambiguous optimization (ci = 100; ri = 100).
Character state 1 was synapomorphic to this level of hierarchy. Even though the quantity of protuberances varied among the genera that composed these two branches, the presence of protuberances was constant, indicating that this character became an evolutionary novelty in the Solenocyclinae+ ancestor ( Figs 2–9View FIGURE 2View FIGURE 3View FIGURE 4View FIGURE 5View FIGURE 6View FIGURE 7View FIGURE 8View FIGURE 9). In Solenocyclinae, the protuberances can be restricted to just half of the distal of the ileum ( Figs 2View FIGURE 2 C–F, 3 E) or, at most, covering three quarters of the surface ( Fig. 2View FIGURE 2 A), but never extending throughout the ileum as occurs in Proculinae+ ( Figs 4–8View FIGURE 4View FIGURE 5View FIGURE 6View FIGURE 7View FIGURE 8).
[Character 7] Proximal area of the ileum with more than one long diverticulum (0); with just one long diverticulum (1). Fast optimization (ci = 100; ri = 100).
In Aulaclocyclinae, the diverticula are long, forming a ring on the proximal apex ( Fig. 1View FIGURE 1). Here the diverticula were shorter and more protuberance-like, in contrast to Aulaclocyclinae, but there was always one longer diverticulum, the form and size of which differed from Solenocyclinae+ ( Figs 2–8View FIGURE 2View FIGURE 3View FIGURE 4View FIGURE 5View FIGURE 6View FIGURE 7View FIGURE 8). Interestingly, the ileum in the outgroup was smooth, presenting no diverticula.
[Character 8] Diverticula restricted to the proximal extremity of the ileum (0); covering at least the third proximal of the ileum (1). Fast optimization (ci = 100; ri = 100).
Character state 1 was synapomorphic for Solenocyclinae+.
[Character 9] Colon without wrinkles (0); with longitudinal wrinkles (1). Unambiguous optimization (ci = 100; ri = 100).
In Aulacocyclinae , the colon of the intestine has longitudinal striations ( Fig. 1View FIGURE 1), while in the other taxa it was smooth. Additionally, the presence of longitudinal wrinkles ( Figs 2–8View FIGURE 2View FIGURE 3View FIGURE 4View FIGURE 5View FIGURE 6View FIGURE 7View FIGURE 8) was an synapomorphy for this level of hierarchy.
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