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Fig. 9 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 9. Muscle scars on internal mould of the rostroconch Ribeiria Sharpe, 1853, in lateral view. Both the anterior and posterior median muscle scars lie across the median dorsal plane of symmetry, the former attached to the transverse pegma preserved as a deep cleft on the internal mould. The posterior scar is a uniform attachment area, often ornamented with transverse growth lines, unlike the multiple small scars of Eotebenna (based on Pojeta and Runnegar 1976; Polechova 2015).

Fig. 7 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 7. Muscle scars on internal moulds of Palaeozoic bivalves. A. Left valve of Babinka Barrande, 1881, from the Lower Ordovician of Öland, Sweden (after Soot-Ryen 1969, length of specimen about 20 mm). B. Left valve of Palaeoneilo musculosa (Knod, 1908) from the Devonian of Bolivia (after Babin and Farjat 1994, length of specimens about 20 mm). C, D. Sketches in apical view showing asymmetry between pedal muscle scars (black) on internal molds of Palaeoneilo musculosa between left and right valves, and variation in pattern of pedal muscle between specimens (after Babin and Farjat 1994). Abbreviation: am, anterior adductor muscle scar.

Fig. 8 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 8. Muscle scars on internal moulds of helcionelloids. All sketches oriented in lateral view with the apex to the right. A. Vendrascospira frykmani Peel and Kouchinsky, 2022 (after Peel 2023). B. Anhuiconus microtuberus Zhou and Xiao, 1984 (after Parkhaev 2002). C. Hensoniconus siku (Peel and Kouchinsky, 2022) (after Peel 2023). D. Bemella communis Parkhaev, 2001 showing three pairs of muscle scars (black, after Parkhaev 2014b); Li et al. 2021) considered the two pairs of scars on the supra-apical surface (joined by grey) to be traces of a single pair of scars. E. Yochelcionella (based on outline of Yochelcionella ostentata Runnegar and Jell, 1974) showing pair of apical muscle scars described by Vendrasco et al. (2010) in Yochelcionella snorkorum Vendrasco, Porter, Kouchinsky, Li, and Fernandez, 2010. F. Eotebenna danica sp. nov., with multiple scars forming a muscle attachment area at the apex.

Fig. 6 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 6. Surface textures on internal mould of a helcionelloid mollusc Vendrascospira frykmani Peel and Kouchinsky, 2022, PMU 39208 from GGU sample 271492, Henson Gletscher Formation, Løndal, Peary Land, North Greenland, Miaolingian, middle Cambrian. A1. Lateral view with one muscle attachment scar from each of the two pairs of muscle scars (arrows). A2. Dorsal view showing pair of symmetrically placed muscle scars (arrows) on the supra-apical surface (right side of A1). The muscle scars lie on each side of the median area with botryoidal surface texture. A3. Detail of muscle scar (left scar in A1). A4. Detail of finely imbricate shell structure from the median area of A2.

Fig. 3. Helcionelloid mollusc Eotebenna viviannae Peel, 1991a in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 3. Helcionelloid mollusc Eotebenna viviannae Peel, 1991a, internal moulds, Andrarum Limestone, Bornholm, Denmark, Guzhangian, Miaolingian, middle Cambrian. A. MGUH 19565, paratype, lateral view (A1) with detail of apex (A2). B. MGUH 34273, lateral (B1) and apico-lateral views showing impression of comarginal ornamentation and cylindrical form of median sub-apical area (B2). C. Specimen lost, lateral view (C1) with detail of apex (C2), arrow locates detail of shell structure (C3). D. MGUH 34274, lateral view with detail of radial fibrous structure and overlying imbricated lamellae (D2), and patch of ornamented outer shell (D1, arrow). E. MGUH 34275, lateral view with detail of apex (E2) with muscle scars; arrows locate possible muscle scar. F. MGUH 34276, lateral view (F1) with detail of possible muscle scar (F2) located by arrows.

Fig. 2. Helcionelloid mollusc Eotebenna viviannae Peel, 1991a, MGUH 19564 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 2. Helcionelloid mollusc Eotebenna viviannae Peel, 1991a, MGUH 19564, holotype, internal mould, Andrarum Limestone, Bornholm, Denmark, Guzhangian, Miaolingian, middle Cambrian. A1. Oblique lateral view showing margin of shell (arrow) along the narrow slit joining the sub-apical and supra-apical apertures. A2. Oblique apico-lateral view. A3. Lateral view. A4. Oblique view showing inverted teardrop-shaped sub-apical aperture and irregular area (arrow) of possible muscle scar. A5. Lateral view of apex. A6. Oblique lateral view of apex showing radial fibrous structure beneath smooth outer layer.

Fig. 1. Geological and geographical background. A in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 1. Geological and geographical background. A. Cambrian stratigraphy of southern Bornholm, Denmark (based on Nielsen and Schovsbo 2007). B. Map of the Baltic area showing location of Bornholm, with location of studied locality (asterisk) on the rivulet Øleå (C), and the Lake Vänern area in southern Sweden (D), with collection locality on the western slopes of the hill Kinnekulle (asterisk).

Fig. 4 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 4. Helcionelloid mollusc Eotebenna danica sp. nov., internal moulds, Andrarum Limestone, Bornholm, Denmark, Guzhangian, Miaolingian, middle Cambrian. A. MGUH 34277, lateral view (A1) with rectangles indicating location of Fig. 5B1, B2, and B3. Muscle scars at apex illustrated in different orientations (A2–A6), with arrow in A2 indicating slight diagenetic compression or deformation along edge of muscle field. Arrow in A3 indicating imbricated lamellar structure shown in detail in A4. B. MGUH 34278, apex in lateral view. C. MGUH 34279, lateral view. D. MGUH 34280, holotype, lateral (D1) and apico-lateral (D2) views, the latter showing the laterally compressed shell form, with detail of apical muscle scars (D3). E. MGUH 34281, lateral view. F. MGUH 34282, apex in lateral view. pontifex Runnegar and Jell, 1976, from the Currant Bush southern Freuchen Land, North Greenland is more strongly Limestone (Miaolingian) of Queensland, Australia, is also coiled than Eotebenna danica, with a convex supra-apical much more elongate than Eotebenna danica but the massive surface, in lateral view, and the apex strongly overhanging snorkel is circular in cross-section compared to the inverted the sub-apical surface (Peel 1989, 1991b). teardrop-shape in the two Bornholm species. Eotebenna arctica Peel, 1989, from the Henson Gletscher Stratigraphic and geographic range.—Drumian of Sweden Formation (uppermost Series 2, Stage 4, lower Cambrian) of and Guzhangian of Denmark (both middle Cambrian).

Fig. 5 in Muscle attachment scars in helcionelloids from Denmark cast light on mollusc evolution in the Cambrian

Fig. 5. Helcionelloid mollusc Eotebenna danica sp. nov. from Miaolingian, middle Cambrian A. MGUH 34283, internal mould with traces of comarginal ornamentation and rugae (arrow), western slopes of Kinnekulle, southern Sweden, Drumian. B. MGUH 34277, details of shell structure, Andrarum Limestone, Bornholm, Denmark, Guzhangian (general view of the specimen in Fig. 4A). Surface of internal mould (B1) digitally inverted and mirrored here (B2) to depict shell structure on the interior surface of the shell. Detail of imbricate lamellae on internal mould (B3).

Fig. 7 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 7. Caste syndromes defined as in fig. 6, optimized on cladogram produced by using caste syndrome as a character in combination with the matrix of Wenzel and Carpenter.

Fig. 3 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 3. Caste discrimination found in some epiponines based on size and shape. Individuals identified by dissection to be ''queens'' are represented as squares, while those identified as ''workers'' are solid dots. Separate regression lines are drawn for illustration only; see text for discussion.

Fig. 6 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 6. Caste syndromes defined by data of table 1 and ovarian dissection, optimized on cladogram of Wenzel and Carpenter. Where genera are polymorphic for caste syndrome, terminals are coded according to parsimonious optimization.

Fig. 5 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 5. Plots of Wilks' lambda values versus queens correctly classified using discriminant function analysis for some epiponines. See text for discussion.

Fig. 4 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 4. Comparison of the level of caste distinction between epiponines (1–59, white circles indicate workers with ovarian development, black diamonds indicate workers without ovarian development) and a Vespinae (60, black square) based on wing length queen–worker ratio versus length of tergite I (except 14, 16, 17, 33–35, 37: hamuli number and 56: width of tergite I). 1. Agelaia areata, 2. A. fulvofasciata, 3. A. lobipleura, 4. A. multipicta, 5. A. pallipes, 6. A. vicina, 7. A. yepocapa, 8. Angiopolybia pallens, 9. Apoica flavissima, 10. A. gelida, 11. A. pallens, 12. Asteloeca ujhelyii, 13. Brachygastra augusti, 14. B. bilineolata, 15. B. lecheguana, 16. B. moebiana, 17. B. scutellaris, 18. Chartergellus communis, 19. Charterginus fulvus, 20. Chartergus chartarius, 21. C. globiventris, 22. C. metanotalis, 23. Clypearia sulcata, 24. Epipona tatua, 25. E. guerini, 26. Leipomeles dorsata, 27. Metapolybia aztecoides, 28. M. docilis, 29. Nectarinella championi, 30. Parachartergus colobopterus, 31. Pa. fraternus, 32. P. smithii, 33. Polybia bicytarella, 34. Po. bistriata, 35. Po. catillifex, 36. Po. dimidiata, 37. Po. emaciata, 38. Po. erythrothorax, 39. Po. jurinei, 40. P. liliacea, 41. P. occidentalis, 42. P. parvulina, 43. P. platycepha sylvestris, 44. P. quadricincta, 45. Po. rejecta, 46. Po. ruficeps, 47. Po. scutellaris, 48. Po. singularis, 49. Po. spinifex, 50. Po. striata, 51. Protonectarina sylveirae, 52. Protopolybia exigua, 53. Pr. minutissima, 54. Pr. pumilla, 55. Pr. sedula, 56. Pseudopolybia difficilis, 57. Ps. vespiceps, 58. Synoeca cyanea, 59. S. surinama, 60. Vespula squamosa. For references for the species, see table 1.

Fig. 2 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 2. Slight caste discrimination found in some epiponines based mostly on shape (when applicable). Individuals identified by dissection to be ''queens'' are represented as squares, while those identified as ''workers'' are solid dots. Separate regression lines are drawn for illustration only; see text for discussion.

Fig. 1 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 1. Representative measures for morphometric analyses of this paper: head width (HW), minimum interorbital distance (IDm), gena width (GW), width of mesoscutum (MSW), alitrunk length (AL), length of gastral tergite I (T1L), basal height of T1 (T1BH), basal widths of tergite II (T2BW), and partial length of the forewing (WL).

Fig. 8 in Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini)

Fig. 8. Wheeler's (1991) model of the evolution of caste, stepwise in graphical terms, representing morphometric variation measured within a colony. See text for explanation.