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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).
Text-fig. 5. Mytoconula vonkai sp. n. The holotype NM L 31983, internal mould. A – apical, B – right apical and right antero-lateral views, both with visible series of muscle scars in two raised zones, ×......;. Note the two long divergent scar zones in A. Dobrotivá F., Mýto near Holoubkov. in Patelliconus Horný, 1961 And Mytoconula Gen. N. (Mollusca, Tergomya) From The Ordovician Of Perunica
Text-fig. 5. Mytoconula vonkai sp. n. The holotype NM L 31983, internal mould. A – apical, B – right apical and right antero-lateral views, both with visible series of muscle scars in two raised zones, ×......;. Note the two long divergent scar zones in A. Dobrotivá F., Mýto near Holoubkov.
Fig. 5 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 5. External view of 23 species examined of Loxoconcha: A — L. shanhaiensis (Ơ, LV); B — Loxoconcha japonica (Ơ, RV); C — Loxoconcha sp. 9 (LV); D — Loxoconcha sp. 10 (LV); E — Loxoconcha sp. 8 (LV); F — Loxoconcha sp. 1 (♀, RV); G — Loxoconcha sp. 30 (LV); H — L. mutsuensis (Ơ, LV); I — Loxoconcha sp. 13 (LV); J — L. tosaensis (LV); K — L. harimensis (LV); L — L. damensis (Ơ, LV); M — L. pulchra (LV); N — L. uranouchiensis (Ơ, LV); O — L. noharai (Ơ, LV); P — L. santosi (Ơ, LV); Q — Loxoconcha sp. 5 (LV); R — Loxoconcha sp. 4 (RV); S — Loxoconcha sp. 26 (♀, RV); T — L. kosugii (Ơ, LV); U — L. sesokoensis (Ơ, RV); V — L. yoshidai (Ơ, RV); W — Loxoconcha sp. 3 (Ơ, LV). Scale 200 µm. Abbreviations: LV, left valve; RV, right valve.
Fig. 3 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 3. Study sites in Japan. Details of occurrence are shown in Appendix 1. Star shapes (Sampling and using ready specimens), solid circles (Only using ready specimens).
Fig. 2 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 2. Loxoconcha elliptica (adult Ơ), seen from the left side with left valve removed, to show the general arrangement of the appendages (only one of each pair of appendages shown for clarity) (Athersuch et al., 1989).
Fig. 6 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 6. External view of 18 species examined of Xestoleberis: A — Xestoleberis hanaii (Ơ, LV); B — Xestoleberi sp. 1 (Ơ, LV); C — Xestoleberi sp. 2 (♀, LV); D — Xestoleberi sp. 5 (♀, LV); E — Xestoleberi sp. 6 (Ơ, RV); F — Xestoleberis sp. 7 (Ơ, LV); G — X. vietnamensis (Ơ, LV); H — X. munensis (Ơ, LV); I — X. magutiensis (♀, LV); J — X. kamiya (Ơ, RV); K — X. ikeya (LV); L — X. planuventer (♀, LV); M — X. ryukyuensis (♀, LV); N — X. sesokoensis (Ơ, LV); O — X. setouchiensis (♀, RV); P — X. kuroshio (Ơ, LV); Q — X. magnoculus (♀, LV); R — X. notoensis (♀, LV). Scale 200 µm. Abbreviations: LV, left valve; RV, right valve.
Fig. 1 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 1. Sketching of male right valve in internal lateral view of carapace of Loxoconcha damensis (adult Ơ) indicating the adductor scars, mandibular scars and frontal scar. Scale 200 µm.
Fig. 10 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 10. Adult mandible in 17 species of the genus Xestoleberis: A — X. hanaii (Ơ); B — Xestoleberis sp. 1 (♀); C — Xestoleberis sp. 6 (Ơ); D — Xestoleberis sp. 7 (Ơ); E — X. magutiensis (♀); F — X. kamiya (Ơ); G — X. vietnamensis (Ơ); H — X. ikeya (Ơ); I — X. planuventer (Ơ); J — X. ryukyuensis (Ơ); K — X. sesokoensis (Ơ); L — X. setouchiensis (Ơ); M — X. Kuroshio (Ơ); N — X. magnoculus (Ơ); O — X. notoensis (Ơ); P — X. sagamiensis (Ơ); Q — X. munensis (Ơ). Note: E, F, after Hirosaki (2013); L, after Okubo (1979); O, P, after Sato & Kamiya (2007). Scale 100 µm.
Fig. 7 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 7. Muscle scars of 23 species of the genus Loxoconcha: A — L. shanhaiensis (Ơ, LV); B — Loxoconcha sp. 1 (♀, LV); C — L. japonica (Ơ, LV); D — Loxoconcha sp. 9 (LV); E — Loxoconcha sp. 10 (LV); F — Loxoconcha sp. 30 (RV); G — L. mutsuensis (Ơ, RV); H — L. modesta (Ơ, RV); I — L. tosaensis (Ơ, RV); J — L. harimensis (♀, LV); K — Loxoconcha sp. 8 (♀, LV); L — L. damensis (Ơ, LV); M — L. pulchra (Ơ, LV); N — L. kosugii (Ơ, LV); O — L. uranouchiensis (Ơ, LV); P — L. noharai (Ơ, RV); Q — L. santosi (♀, RV); R — Loxoconcha sp. 5 (♀, RV); S — Loxoconcha sp. 4 (LV); T — Loxoconcha sp. 26 (Ơ, LV); U — L. sesokoensis (Ơ, RV); V — L. yoshidai (Ơ, LV); W — Loxoconcha sp. 3 (Ơ, LV). Scale 100 µm. Abbreviations: LV, left valve; RV, right valve.
Fig. 4 in Added Morphological Characteristics Of Muscle Scars, Maxillula And Mandible Of Two Ostracod Genera Loxoconcha (Loxoconchidae) And Xestoleberis (Xestoleberididae)
Fig. 4. Map of Vietnam showing four surveyed areas with solid circles, Phu Quoc Island, Nha Trang Bay, Ha Long Bay and Cat Ba Island.
Text-fig. 1. Schizocrania incola (PERNER, 1903). Drawings of visceral area and muscle scars. in Schizocrania (Brachiopoda, Discinoidea): Taxonomy, Occurrence, Ecology And History Of The Earliest Epizoan Lingulate Brachiopod
Text-fig. 1. Schizocrania incola (PERNER, 1903). Drawings of visceral area and muscle scars.
Data from: Gut evolution in early Cambrian trilobites and the origin of predation on infaunal macroinvertebrates: evidence from muscle scars in Mesolenellus.
Trilobites are particularly common Cambrian fossils, but their trophic impact on the rapidly evolving marine ecosystems of that time is difficult to assess, due to uncertainties on how diverse their feeding habits truly were. Gut anatomy might help to constrain inferences on trilobite feeding ecology, but preservation of digestive organs is exceedingly rare. Muscle scars on the glabella, known as 'frontal auxiliary impressions' (FAIs), have been interpreted as evidence of the evolution of a pouch-like organ with powerful extrinsic muscles (i.e. a crop) in some trilobites. Here we describe FAIs in Mesolenellus hyperboreus from Cambrian Stage 4 strata of North Greenland, which represents the oldest example of such structures and their first report in the Suborder Olenellina. Mesolenellus FAIs suggest that the crop in trilobites was clearly differentiated from the rest of the digestive tract, and essentially located under a hypertrophied glabellar frontal lobe. Reviews of the digestive anatomy of trilobite sister-taxa and the glabellar morphology of the oldest-known trilobites suggest that the gut of the trilobite ancestor was an essentially simple tract (i.e. no well-differentiated crop) flanked laterally by numerous midgut glands. A crop first evolved in the Cambrian in groups like olenelloids and (later) paradoxidoids. Using ichnological evidence, we hypothesize that the emergence of olenelloids yields evidence for the evolution of predatory inclinations in a group of arthropods originally dominated by surface-deposit-feeders. By allowing the exploitation of a rapidly developing food source, infaunal animals, the diversification of feeding strategies in trilobites might partially explain their unparalleled evolutionary success.
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