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Phalangeal Formula of Lepidosauria
<p>Evolutionary tempo and mode summarize ancient and controversial subjects of theoretical biology such as gradualism, convergence, contingence, trends, and entrenchment. We employed an integrative methodological approach to explore the evolutionary tempo and mode of Lepidosaurian Phalangeal Formulae (PFs). This approach involves quantifying the frequencies of morphological changes along an evolutionary trajectory (ET). The five meristic characters encoded by PFs are particularly valuable in revealing evolutionary patterns, owing to their discrete nature and extensive documentation in the literature. Based on a pre-existing dataset of PFs from 649 taxa (35 Lepidosauria families, including fossils), from which there exists a unique repertoire of 53 formulations, our approach simultaneously considers phenetic and phylogenetic data. This culminates in a diagram accounting for the phylogenetic dynamic of evolution traversing across different regions of morphospace. The method involves enumerating phenotypical options, reconstructing phenotypes across the phylogeny, projecting phenotypes onto a morphospace, and constructing a flow network from the frequency of evolutionary transitions between unique phenotypic conditions. This approach links Markovian chains and evolutionary trajectories to formally define parameters that describe the underlying transitions of morphological change. Among other results, we found that: (i) PF evolution exhibits a clear trend toward reduction in the phalangeal count, and that (ii) evolutionary change tends to occur significantly between morphologically similar PFs. Notwithstanding, although minor but not trivial, transitions between distant formulas jumps occur. Our results support a pluralistic view including stasis, gradualism, and saltationism discriminating their prevalence in a target character evolution.</p>
Рис. 4. Поперечные срезы фаΛанг паΛьцев особей Bufo sachalinensis максимаΛьного возраста: A — шестиΛетний самец (L = 69.0 мм); B — семиΛетняя самка (L = 90.6 мм) Fig. 4. Cross-section image of phalanges of Bufo sachalinensis individuals of maximum age: A — six year old male (SVL = 69.0 mm); B — seven year old female (SVL = 90.6 mm) in Age structure and sexual dimorphism of the Far Eastern toad, Bufo sachalinensis Nikolsky, 1905 in the Ussurisky Nature Reserve
Рис. 4. Поперечные срезы фаΛанг паΛьцев особей Bufo sachalinensis максимаΛьного возраста: A — шестиΛетний самец (L = 69.0 мм); B — семиΛетняя самка (L = 90.6 мм) Fig. 4. Cross-section image of phalanges of Bufo sachalinensis individuals of maximum age: A — six year old male (SVL = 69.0 mm); B — seven year old female (SVL = 90.6 mm)
DT A 'rw YTTT Unix... Au .. Ornithomimus nazis, Lainbe, phalanges of right pes, external view; natural size. Page 50. in New genera and species from the Belly River Series (mid-Cretaceous)
DT A 'rw YTTT Unix... Au .. Ornithomimus nazis, Lainbe, phalanges of right pes, external view; natural size. Page 50.
Figure 2. Cross-sections from A. danfordi phalanges. A in Age and growth in two populations of Danford's lizard, Anatololacerta danfordi (Günther, 1876), from the eastern Mediterranean
Figure 2. Cross-sections from A. danfordi phalanges. A) Female lizard with 3 hibernations and B) male lizard with 7 hibernations. e.b.: Endosteal bone, e.r.: endosteal resorption, m.c.: medullary cavity, p.: peripheral, white arrows: LAGs.
Fig. 4 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 4. Photo (A) and drawing (B) of the right pedal ungual II of the camarasaurid sauropod SMA 0002 (Upper Jurassic Morrison Formation, HoweStephens Quarry, Wyoming, USA) in proximal view, showing the broken surface indicating overgrowth attachment. Modified from Tschopp et al. (2015).
Fig. 8. Overgrowth type 3 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 8. Overgrowth type 3 (arrows in A, B) on the distal articular surface of the left manual phalanx IV-1 of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA; in anterior/dorsal (A) and distal (B) views, and CT scan of frontal slice (C), note the probable healed fracture (arrows). Photos taken by Rosemarie Roth (University of Zurich, Switzerland).
Fig. 3. Overgrowth type 1 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 3. Overgrowth type 1 (arrows) in right pedal unguals I (A) and III (B) of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA. Overgrowth projects proximally from the proximal articular surface. Note the medial to mediodorsal position of the overgrowths on the proximal articular surfaces. Unguals shown in dorsal (A1, B1), medial (A2, B2), proximal (A3, B3), and lateral (A4, B4) views. Photos taken by Esther Premru (Mönchaltorf, Switzerland) and modified from Tschopp et al. (2015).
Fig. 2. Overgrowth type 1 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 2. Overgrowth type 1 (arrow) in left pedal ungual I of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, HoweStephens Quarry, Wyoming, USA; in lateral (A), dorsal (B), and proximodorsal (C) views. Note the dorsal position of the overgrowth on the proximal articular surface, and how it fits in the notch in the distal articular surface of php I-1 (C, slightly displaced taphonomically). Abbreviations: mt, metatarsal; php, pedal phalanx. Not to scale, the proximodistal length of php I-1 is 45 mm.
Fig. 9. Overgrowth type 4 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 9. Overgrowth type 4 (arrow) on the laterodistal corner of the left pedal phalanx IV-1 of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA; in anterior/dorsal view. Abbreviation: php, phalanx of pedal digit.
Fig. 1 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 1. Schematic drawing of the left and right manual (A) and pedal (B) phalanges of the camarasaurid sauropod SMA 0002, marking the elements affected by pathologies described herein. Rectangles are normal phalanges, triangles are unguals. Dashed lines indicate lacking elements. Numbers correspond to the overgrowth types as defined in the text; e, possible eburnation; p, deep pit.
Fig. 11 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 11. Comparison of the normal development of articular surfaces of phalanges of diplodocid sauropod SMA 0087 (php II-1, A) with the pathological elements of camarasaurid sauropod SMA 0002 (php III-1, B); both from the Upper Jurassic Morrison Formation, Howe Ranch, Wyoming, USA. Note the considerable extension of the lateral spurs in SMA 0002.
Fig. 10 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 10. Deep pit (arrow) in the proximal articular surface of the left pedal phalanx II-1 of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA; in anteroproximal view. The proximal width of the phalanx is 72 mm. Abbreviation: mt, metatarsal.
Fig. 7 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 7. Smoothened area (a possible case of eburnation) on the distal articular surface of the left manual phalanx I-1 of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA. The distal dorsopalmar height is 55 mm.
Fig. 6. Overgrowth type 2 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 6. Overgrowth type 2 exemplified in a drawing (A, modified from Tschopp et al. 2015) and CT scan (B) of the left manual phalanx II-1 of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA. CT scan shows variable bone densities in the osteophyte, indicated by the different gray scales. Abbreviations: phm, phalanx of manual digit; mc, metacarpal.
Fig. 5 in Pathological phalanges in a camarasaurid sauropod dinosaur and implications on behaviour
Fig. 5. Thin section of the overgrowth type 1 from the right pedal ungual I of the camarasaurid sauropod SMA 0002 from Upper Jurassic Morrison Formation, Howe-Stephens Quarry, Wyoming, USA (A). Detail (B), showing coarse-grained structure bone in mainly avascular bone (arrowheads).
Explanation of Plate I. Figure 1.—Left tibia of Ornithomimus velox, Marsh; A, front view; b, distal end; c, transverse section. Figure 2.—Left metatarsals of same specimen; A, front view; b, proximal ends; c, transverse section; d, distal ends. Figure 3.—Phalanges of second digit of same foot; front view, a, first phalange; b, second phalange; c, third, or terminal phalange. Figure 4.—Left metacarpals of same species, perhaps of smaller individual; front view. Figure 5.—Left tibia of young Ostrich (Struthio camelus, Linn.); a, front view; b, distal end. The separate calcaneum was first observed by the writer's assistant, Dr. G-. Baur, who prepared the specimen. Figure 6.—Left metatarsals of young turkey (Meleagris gallipavo, Linn.); a, front view; b, proximal ends. a, astragalus; as, ascending process of astragalus; c, calcaneum; f, fibula; f' face for fibula; II, second metatarsal; III, third metatarsal; iv, fourth metatarsal. Figures 1-4 are one-third natural size, and figures 5 and 6, one-half natural size. in Description of new dinosaurian reptiles
Explanation of Plate I. Figure 1.—Left tibia of Ornithomimus velox, Marsh; A, front view; b, distal end; c, transverse section. Figure 2.—Left metatarsals of same specimen; A, front view; b, proximal ends; c, transverse section; d, distal ends. Figure 3.—Phalanges of second digit of same foot; front view, a, first phalange; b, second phalange; c, third, or terminal phalange. Figure 4.—Left metacarpals of same species, perhaps of smaller individual; front view. Figure 5.—Left tibia of young Ostrich (Struthio camelus, Linn.); a, front view; b, distal end. The separate calcaneum was first observed by the writer's assistant, Dr. G-. Baur, who prepared the specimen. Figure 6.—Left metatarsals of young turkey (Meleagris gallipavo, Linn.); a, front view; b, proximal ends. a, astragalus; as, ascending process of astragalus; c, calcaneum; f, fibula; f' face for fibula; II, second metatarsal; III, third metatarsal; iv, fourth metatarsal. Figures 1-4 are one-third natural size, and figures 5 and 6, one-half natural size.
Figure 7. A, left digit II phalanges 2 and 3 in On the manual morphology of Compsognathus longipes and its bearing on the diagnosis of Compsognathidae
Figure 7. A, left digit II phalanges 2 and 3 (ungual) shown in near articulation. Arrow shows subcircular articular condyles typical of theropod II-2. B, right digit II, phalanx 2 lateral impression.
Phalangeal Formula of Lepidosauria
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Plate XIII Ornithomimus altus, Lambe, phalanges of right pes, external view; natural size. Page 50. in New genera and species from the Belly River Series (mid-Cretaceous)
Plate XIII Ornithomimus altus, Lambe, phalanges of right pes, external view; natural size. Page 50.
Metatarsal Phalangeal Joint Deformity Progression - R01
ClinicalTrials.gov study NCT02616263. IPD Sharing: NO. Countries: 1. Publications: 14.
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