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Figure 223 in Taxonomy of Phanaeus revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)

Figure 223. Approximate geographic distributions of the Phanaeus (P.) hermes, beltianus and amethystinus species groups.

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Figure 1 in Deltochilini and Phanaeini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in introduced and native ecosystems of Brazil

Figure 1. Monthly precipitation, average monthly temperature and abundance and richness of Deltochilini and Phanaeini dung beetles (Coleoptera, Scarabaeinae) sampled in pasturelands with exotic grass (Brachiaria spp.) and a patch of vegetation native (Brazilian savanna) (Aquidauana, Mato Grosso do Sul, Brazil) using pitfall traps baited with carrion and human faeces, from January to December 2011. Bars on columns represent standard errors.

Figure 15 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 15. Predicted area of environmental suitability for Bolbites onitoides in South America (to the left) and the realized geographical range of the species (to the right).

Figure 13 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 13. Predicted areas of environmental suitability (temperature and precipitation) for the red (upper row) and blue (lower row) variants of Bolbites onitoides within the range of the species. Note that the area of suitable environmental conditions for each colour variant within the range of B. onitoides, particularly when precipitation is taken into account, corresponds largely to the actual range of each variant. Furthermore, these areas largely do not overlap with each other. Confirming our initial hypothesis, this result indicates that the differential distribution of the red and blue variants across the populations of B. onitoides is likely a response to varying environmental conditions.

Figure 14 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 14. Predicted areas of environmental suitability (temperature and precipitation) for the bicoloured (upper row) and coppery (lower row) variants of Bolbites onitoides within the range of the species. Note that the area of environmental suitability for each of these variants matches their observed geographical ranges and is largely corresponded to the range of the species as a whole (particularly the bicoloured one). They broadly overlap with each other and with the suitable areas for the red and blue variants (Fig. 13).

Figure 12 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 12. Distribution of each colour variant showed in separate maps for an easier discrimination. See legend for Figure 10 for more details.

Figure 11 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 11. Geographical range of B. onitoides showing the differential distribution of the colour variants. It is indisputable that there is a strong correlation between colour and geographical provenance in the species: blue individuals are abundant at the western-half of the distribution, while red specimens are dominant at the eastern-half. Nevertheless, two kinds of intermediates – the discrete bicoloured and the gradual colour-blended – are found throughout the range of B. onitoides, but nowhere are they the dominant variant. Note that there is a zone of overlap between the distribution of red and blue variants in Paraguay and Argentina, and that three localities recorded red, blue and bicoloured individuals and a fourth recorded all four variants.

Figure 10 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 10. Phylogenetic placement of Bolbites onitoides among the Phanaeini and the evolution of iridescent colouration in this clade. A growing body of evidence has accumulated over the past two decades confirming that the horneless Bolbites is a phanaeine lineage that is more closely associated to horned Phanaeina than to the primitively hornless Gromphadina (see references in the text). If this is correct, the evolution of iridescence from the plesiomorphic opaque condition preceded that of the typically male cephalic and pronotal ornamentation found widely among the Phanaeina. As a consequence, the selective pressure that led to the evolution of the two features cannot have been the same. A more likely scenario is that iridescence first evolved serving another function in the common ancestors of the Phanaeini – e.g. thermoregulation or aposematism – and was inherited serving this function by the the Gromphadina and Phanaeina alike. Then, iridescence was putatively co-opted independently during the parallel evolution of cephalic horns by the Phanaeina less Bolbites and the gromphadine clade Gromphas aeruginosa + G. lemoinei to serve a new role in the evaluation of male horns during sexual interactions. Sexual selection thus co-opted for a new use a feature that had originally evolved under natural selection serving another function. This characterizes iridescence as an exaptation for horn evaluation, not an adaptation for this function. Species illustrated (from top to bottom): Eucranium cyclosoma Burmeister, 1861, Oruscatus davus (Erichson, 1847), Gromphas lemoinei Waterhouse, 1891, G. dichroa Blanchard, 1846, Bolbites onitoides Harold, 1868, a red Coprophanaeus saphirinus (Sturm, 1826), a blue C. saphirinus, and Sulcophanaeus menelas (Castelnau, 1840).

Figure 9 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 9. Male genitalia: Endophallus of B. onitoides. A, B, general view of the extracted endophallus. A, dorsal view. B, ventral view. C–G, endophallites: C, axial + subaxial endophallite complex (dissected). D, fronto-lateral peripheral endophallite (ventral view). E, medial endophallite (dorsal view). F–H, superior right peripheral endophallite from different views: F, G, ventral view: F, seahorse-shaped; G, F-shaped. H, lateral view. A = axial endophallite; SA = subaxial endophallite; MS = medial endophallite; FLP = fronto-lateral peripheral endophallite; SRP = superior right peripheral endophallite.

Figure 8 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 8. Genitalia of both sexes of B. onitoides. A, female: spermatheca. B–D, male: B, genital segment (i.e. nineth abdominal segment). C, D, tegmen. C, dorsal view. D, lateral view. LSP = lateral sclerotized plate; MSP = medial sclerotized plate.

Figure 6 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 6. Sexual dimorphism on the hind legs of B. onitoides. A, B, metacoxae: A, male; B, female. Note the plate-like projections and the long spines directed posteriorly in the male and which are lacking in the female. C, D, metafemora and metatibiae: C, male; D, female. Observe the ridge on the posterior edge of the metafemora and the sharp ridge on the external and internal edges of the metatibiae of the males, all of which are absent in females.

Figure 7 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 7. Sexual dimorphism on the pygidium and at the apex of the sutural margin of the elytra of B. onitoides: A, male; B, female. Orange arrow shows the short spines of the apex of the sutural margin of the elytra, whereas red arrow indicates the long spines raising from the basolateral angles of the pygidium of males. Both structures are absent in females.

Figure 5. Legs. A–E in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 5. Legs. A–E, sexual dimorphism on the front legs of B. onitoides. A, B, profemora: A, male; B, female. Note that the male profemur is claviform and has a strong emargination on its posterior edge (indicated by the red arrow), while that of the female is shorter and has a simply rounded aspect. C–E, protibiae: C, D, dorsal view of male (C) and female (D). Observe that the male protibia is much longer and slenderer than that of the female and has a well-delimited tuft of long setae at its internal apical angle. E, lateral view of the male protibia. In contrast to females, males have a series of sharp, uneven teeth along the ventral protibial carina. Red arrow indicates the central tooth, which is longer than the others. F, G, meso- and metatarsi of B. onitoides. Unlike all other Phanaeini, which are clawless, B. onitoides has a pair of well-developed claws on the apical meso- and metatarsomeres. A small prolongation raises from underneath the claws and project itself anteriorly (indicated by the red arrow in F); this prolongation seems to be homologous to the tapered apex of the apical meso- and metatarsomeres of the other Phanaeini and is probably a synapomorphy of the clade. H, metanepisternum and its long tab typical of the Phanaeina.

Figure 2 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 2. Ventral sexual dimorphism in Bolbites onitoides. A, lateral view of a male. B, C, ventral view: B, male: C, female. Note the conspicuous ventral sexual dimorphism in this species, including the presence of several teeth and spines all over the body of the male, the differences in the overall shape of the protibiae, the pilosity of the metacoxae, and colour of the metaventrite.

Figure 3 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 3. Dorsal sexual monomorphism in Bolbites onitoides. A, frontal view of a male. B, C, head and pronotum: B, male; C, female. Note how little the sexes differ dorsally. Unlike most groups of Phanaeina, males of B. onitoides lack any cephalic or pronotal horns and have a pronotal ridge identical to that of the females. This dorsal monomorphism is in striking contrast with the sexual dimorphism found on the venter. Note, however, the sexually dimorphic extension of iridescence over the clypeus, males having at least its basal half metallic, while females are entirely (or almost entirely) dull black.

Figure 4 in The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem

Figure 4. Meso- and metaventrite in Bolbites and Gromphadina. A, Bolbites onitoides. B, Gromphas inermis. C, D, Oruscatus davus: C, male; D, female. The carina of the mesofurcal brace constitutes the posterior branch of the mesoventral ridge and gives it the shape of a cross (orange arrow). This cross-shaped ridge is typical of the Gromphadina and is also present in at least some Phanaeina. Note that, while in the Gromphadina the posterior branch connects the mesoventral ridge to the anterior edge of the metaventrite, no such connection exists in most Bolbites and and the other Phanaeina whose mesoventral ridge is T-shaped. In Oruscatus, although present in both sexes, the posterior branch of the mesoventral ridge is much more clearly delimited in males than in females; in females, the posterior branch is usually incomplete and interrupted, while in males it is always sharp and continuous. Note also other differences between Bolbites and the Gromphadina relating to the shape of the metaventral anterior prominence and its integument and lateral pilosity.

Figure 4 in A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)

Figure 4. Male paratype of Gromphas jardim sp. nov. and its labels (braces '{' joining two sides of a same label). We believe that the French zoologist and explorer Alcide d'Orbigny was the collector of this specimen in 1832 (see the text).

Figure 3 in A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)

Figure 3. Lateral view of pronotum. (A) Gromphas jardim sp. nov. (B) Gromphas amazonica. Note that the granulation extends much more posteriorly in G. jardim, reaching the pronotal posterior margin, than in G. amazonica, wherein the granulation is absent or rudimentary in posterolateral region after lateral fossa.

Figure 6 in A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)

Figure 6. Phylogeny of Gromphas. Above: Cladogram showing the phylogenetic relationship of the species of Gromphas and the evolution of their characters. Black circles indicate uncontroverted synapormorphies; white circles with black margin, controverted synapomorphies; red circles, ambiguous uncontroverted synapomorphies; and white circles with red margin, ambiguous controverted synapomorphies. Below: Support values given for each clade are decay index (above branch) and bootstrap (below branch).

Figure 1 in A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)

Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype).