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Fig. 21. Wings. A‒E. Elytra. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 21. Wings. A‒E. Elytra. A. Lateral view of elytra of Streblopus opatroides van Lansberge, 1874. Note the very wide pseudepiplera and narrow epipleura. B‒C. Pseudepipleural carina (indicated by red arrows). B. S. opatroides. C. S. punctatus (Balthasar, 1938). Observe that the pseudepipleural carina in B is completely interrupted by umbilicate punctures, whereas the pseudepipleural carina in C is almost completely smooth, with very tiny and isolated punctures. D‒E. Lateral view of apex of elytra. D. S. opatroides. E. S. punctatus. Note how the distal calluses (shown by blue arrows) of S. punctatus are much more strongly projecting than those of S. opatroides. F. Right hind wing of Streblopus opatroides. Red arrow shows the notch of the anal fold. Abbreviations: AA = anal anterior; AP = anal posterior; Cu = cubitus; J = jugal; MP = media posterior; MPa = accessory MP; r4 = cross-vein r4 (cross-vein connecting veins RA3+4 and RP2); RA = radius anterior; RP = radius posterior; ScA = subcosta anterior.

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Fig. 20. Posterior legs. A‒D. Metatibiae. A‒B. Streblopus opatroides van Lansberge, 1874. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 20. Posterior legs. A‒D. Metatibiae. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. Note the strong sexual dimorphism and the many differences between the species. E‒F. Metatarsi and apex of metatibia of S. opatroides. E. ♂. F. ♀. Observe how much shorter the male metatibial spur is in comparison to that of the female.

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Fig. 23. Genital capsule. A‒D. Streblopus opatroides van Lansberge, 1874. A. Dorsal view. B‒C in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 23. Genital capsule. A‒D. Streblopus opatroides van Lansberge, 1874. A. Dorsal view. B‒C. Lateral view from slightly different angles (red arrow shows the ventral surface and its keel). D. Ventral view. E‒H. S. punctatus. (Balthasar, 1938). E. Dorsal view. F‒G. Lateral view from slightly different angles (red arrow indicates ventral surface). H. Ventral view. Observe the differences in the shape of the parameres, in particular how the apical hook is much more prominent in S. punctatus.

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Fig. 22 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 22. Abdomen of Streblopus opatroides van Lansberge, 1874. A‒B. Lateral view. A. ♂. B. ♀. Note that the female abdomen has a straighter surface than that of the male, which is concave (red arrow shows this concavity). C. Pygidium and prepygidium.

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Fig. 28 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 28. Current distribution of Streblopus van Lansberge, 1874 plotted on a Middle Miocene palaeomap (~ 15 million years ago). Presently, both species of Streblopus are largely isolated from one another by means of the South America Dry Diagonal that separates the humid forests of the Amazon Basin from the Atlantic Forest on the South American eastern coast. This disjunct distribution is probably a result of the fragmentation of an once much more widely-distributed genus, which crossed the Dry Diagonal from the Atlantic Forest to southwestern Amazonia through forest corridors connecting those two biomes during periods of warmer climate over the past 25 million years. When the climate cooled and the tropical forests retreated, the open and drier environments of the Dry Diagonal advanced, and the populations of Streblopus in the Atlantic Forest and in the Amazon became isolated. It is known that the Middle Miocene experienced the highest temperatures of the last 20 million years (Zachos et al. 2001), and perhaps it was during its ʻclimate optimumʼ about 15 million years ago that the Amazonian and the Atlantic Forest populations of Streblopus were connected for the last time. Palaeomap modified from Scotese (2016).

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Fig. 26. Genital segment. A. Streblopus opatroides van Lansberge, 1874. B. S in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 26. Genital segment. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note that the medial sclerotized plate is larger and more sclerotized in S. punctatus than in S. opatroides; other differences refer to their lateral arches (indicated by blue arrows), which are concave in S. opatroides and more or less straight in S. punctatus, and the posterior arms of the MSP (indicated by red arrows), which are well developed and separated by a deep ʻU-shapeʼ in S. punctatus and only slightly indicated in S. opatroides. Abbreviations: LSP = lateral sclerotized plate; MSP = medial sclerotized plate.

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Fig. 18. Mesotibiae. A‒B. Streblopus opatroides van Lansberge, 1874. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 18. Mesotibiae. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. Observe that females of S. opatroides differ from those of S. punctatus by having a deep emargination at the apical edge of the mesotibiae (red arrow shows emargination).

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Fig. 24 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 24. Internal sac of Streblopus opatroides van Lansberge, 1874. A‒C. Different views of the undissected internal sac and sclerites in resting position. D. Superior right peripheral sclerite. Arrow indicates a triangular area of a more sclerotized tegument that is probably homologous to a similar triangular area on the SRP of S. punctatus (Balthasar, 1938). E‒F. Apical and subapical sclerites. E. Resting position. F. Sclerites after separation. Abbreviations: A = axial sclerite; Rp = raspule; SA = subaxial sclerite; SRP = superior right peripheral sclerite; T = temones.

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Fig. 17. Mesofemora. A‒B. Streblopus opatroides van Lansberge, 1874. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 17. Mesofemora. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. Note the sexual dimorphism in the presence of a long row of setae on the posterior edge of the male mesofemur. C‒F. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. E‒F. Detail of posterior region of metafemora. E. ♂. F. ♀. Note the strong alveolar microsculpture present near the posterior edge in males and the high concentration of umbilicate punctures in the same region in females.

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Fig. 16. Protibiae. A‒B. Lateral view. A. Streblopus opatroides van Lansberge, 1874. B. S in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 16. Protibiae. A‒B. Lateral view. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note the how the protibia of S. opatroides is much more curved than that of S. punctatus. C. Frontal view of the protibia of S. punctatus. Note the presence of a protibial spur.

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Fig. 19. Posterior legs. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 19. Posterior legs. A. Metacoxa of Streblopus opatroides van Lansberge, 1874. Arrow shows the lateral spur present in both sexes of both species. Note also that the spur covers the epipleura and possibly maintains the elytra pressed against the body, thus avoiding loss of water or the entry of sediment beneath the elytra in a similar way to that hypothesized for the metanepisternal tab of the Phanaeini and other tunneller groups (Edmonds 1972; Cupello & Vaz-de-Mello 2016). B‒E. Metafemora and metatrochanters. B‒C. S. opatroides. B. ♂. C. ♀. D‒E. S. punctatus (Balthasar, 1938). D. ♂. E. ♀. Note the strong sexual dimorphism and the many differences between the species.

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Fig. 13. Venter. A‒B. Streblopus opatroides van Lansberge, 1874. A. Hypomeron. B in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 13. Venter. A‒B. Streblopus opatroides van Lansberge, 1874. A. Hypomeron. B. Mesoventrite and anterior region of metaventrite. C‒F. Metaventrite and abdomen. C‒D. S. opatroides. C. ♂. D. ♀. E‒F. S. punctatus (Balthasar, 1938). E. ♂. F. ♀. Note in particular the distinction between the species and sexes in the shape of the concavity of the posterior region of the metaventrite.

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Fig. 15. Protibiae. A‒D. Dorsal view. A‒B. Streblopus opatroides van Lansberge, 1874. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 15. Protibiae. A‒D. Dorsal view. A‒B. Streblopus opatroides van Lansberge, 1874. A. ♂. B. ♀. C‒D. S. punctatus (Balthasar, 1938). C. ♂. D. ♀. E‒H. Ventral view. E‒F. S. opatroides. E. ♂. F. ♀. G‒H. S. punctatus. G. ♂. H. ♀. Notice the many differences between the species and sexes, in particular the presence of a protibial spur in males of S. punctatus (and its absence in S. opatroides) and the different shapes of the spurs and of the lateral teeth in females of the two species.

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Fig. 11 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 11. Labium of Streblopus opatroides van Lansberge, 1874 (left palpus removed). A. Ventral view (aboral surface). B. Dorsal view (oral surface). C. Lateral view. D. Removed left palpus. Note the unusual shape of the palpus, with the basal palpomere remarkably enlarged. Abbreviations: gl = glossa; glf = glossal flap; lbpl = labial palp; lshpx = lateral hypopharyngeal sclerite; mt = mentum; pmts = premental sclerite; shp = suspensorium of hypopharynx; vapmts = ventral angles of premental sclerites.

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Fig. 4 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 4. Distribution of Streblopus van Lansberge, 1874. Note how the two species are widely separated by the vast South American Dry Diagonal stretching between Amazonia and the Atlantic Forest. During several different times over the Neogene, forest corridors connected the two biomes and the populations of Streblopus could move from one to the other. When the forest retreated, the distribution of the genus became relict.

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Fig. 10 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 10. Maxillae of Streblopus opatroides van Lansberge, 1874. A. Dorsal view with extended lacinia. B. Dorsal view. C. Ventral view. Abbreviations: car = cardo; darsgal = dorsal articular sclerite of galea; gal = galea; lac = lacinia; mxpl = maxillary palpus; prcar = articular process of cardo; ss1–4 = stipal sclerite 1–4; varsgal = ventral articular sclerite of galea.

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Fig. 6. Head. A‒B. Clypeal process. A. Streblopus opatroides van Lansberge, 1874. B. S in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 6. Head. A‒B. Clypeal process. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). C. Ventral view of head, S. opatroides.

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Fig. 8. Labrum. A‒B. Streblopus opatroides van Lansberge, 1874. A in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 8. Labrum. A‒B. Streblopus opatroides van Lansberge, 1874. A. Aboral surface (dorsal view). B. Oral surface (ventral view; epipharynx). C‒D. S. punctatus (Balthasar, 1938). C. Aboral surface. D. Oral surface. Note the many differences between the two species, particularly the much denser and shorter setation of the median area in S. punctatus, the much longer median brush of S. opatroides and the dissimilarities in the shape of the anterior epypharingeal margin. The black arrow shows the anterior margin of the labrum, while the red arrow indicates the anterolateral region. Abbreviations: afr = apical fringe of labrum; ampr = antero-median process of labral suspensorium; lf = lateral files; mbr = medial brush of labrum; pmpr = postero-median process of labral suspensorium; ppm = proplegmatium; ssma = socketed setae of median area; topr = tormal process of labral suspensorium.

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Fig. 9 in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 9. Mandibles of Streblopus opatroides van Lansberge, 1874. A‒B. Ventral view. A. Left mandible. B. Right mandible. C. Lateral view of right mandible. Abbreviations: apmm = apodemes of mandibular muscles; cmb = comb of incisor lobe of mandible; conj = conjunctivus of mandible; inlb = incisor lobe of mandible; mac = mandibular acetabulum; molb = molar lobe of mandible.

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Fig. 7. Antennae. A. Streblopus opatroides van Lansberge, 1874. B. S in Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae)

Fig. 7. Antennae. A. Streblopus opatroides van Lansberge, 1874. B. S. punctatus (Balthasar, 1938). Note the differences between the two species concerning the colour of the apical flagellomeres and the number and shape of the articles. In S. punctatus, the suppression of arthrogenesis between ancestral articles III and IV has resulted in a reduction of the total number of antennomeres from 9 to 8.

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International Brain Laboratory public data

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