Fig. 4 in The Pupation Chamber of Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae)

Fig. 4. Unambiguous optimization for the character "pupation chamber wall" on an adapted Scarabaeinae phylogenetic tree from Monaghan et al. (2007). Gray circles indicate pupation chamber wall character states: 0 – without visible pellets or pellets without a particular arrangement; 1 – row of pellets arranged in concentric rings; 2 – imbricate pellets grouped in slices oriented longitudinally; 3 – helicoid composed of whorls showing transverse rows of imbricate pellets. Dotted lines represent ambiguous optimization of the character. Node with * indicates when clusters A and B (Scarabaeinae) diverged, and probably when pellets arranged in rows in the construction of pupation chamber wall appeared. Black arrow shows a minimum age of approximately 54 million years for clade B. This is only for illustrative purpose. Branch lengths do not reflect any relative age for the groups. Principal genera within different polyphyletic tribes are: Canthonini 1: Panelus; Canthonini 2 (Ca 2): Temnoplectrom + Lepanus + Cephalodesmius + Saphabius; Canthonini 3 (Ca 3): Circellium; Canthonini 4 (Ca 4): Anachalcos; Neotropical Canthonini: Canthon + Megathoposoma + Malagoniella + Deltochilum; Coprini 1: Copris + Microcopris; Coprini 2 (Co 2): Coptodactyla; Coprini 3 (Co 3): Catharsius + Metacatharsius; Dichotomiini 1: Heliocopris; Dichotomiini 2 (Di 2): Uroxys + Bdelyropsis + Demarziella; Dichotomiini 3 (Di 3): Ateuchus; Dichotomiini 4 (Di 4): Ontherus + Canthidium; Onthophagini 1: Onthophagus + Euonthophagus + Hyalonthophagus + Caccobius + Milichus + Cleptocaccobius; Onthophagini 2: Proagoderus + Onthophagus + Digitonthophagus.