What are the most accurate categories for mammal tarsus arrangement? A review with attention to South American Notoungulata and Litopterna - PubMed (original) (raw)

Review

. 2019 Dec;235(6):1024-1035.

doi: 10.1111/joa.13065. Epub 2019 Aug 2.

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Review

What are the most accurate categories for mammal tarsus arrangement? A review with attention to South American Notoungulata and Litopterna

Malena Lorente. J Anat. 2019 Dec.

Abstract

The arrangement of the tarsus has been used to differentiate afrotherian and laurasiatherian ungulates for more than a century, and it is often present in morphological matrices that include appendicular features. Traditionally, it has two states: (i) an alternating tarsus, where proximal elements are interlocked with central and distal elements positioned like the bricks of a wall; and (ii) a serial tarsus, where elements are not interlocked. Over the years, these states became synonymous with the presence or absence of an astragalocuboid contact. Within the South American order Notoungulata, a third disposition was recognized: the reversed alternating tarsus, associated with a calcaneonavicular contact. This state was considered to be a synapomorphy of 'advanced' Toxodontia families (Notohippidae, Leontiniidae and Toxodontidae), but a further inspection of its distribution shows that it occurs throughout Mammalia. Additionally, it overlaps the serial tarsus condition as originally defined, and it probably has no functional or phylogenetic significance. Calcaneonavicular and astragalocuboid contacts are non-exclusive, and their presence within a species, genus or family is not constant. Serial and alternating imply movements of the articulations of the mid-tarsus in the transverse axis, while reverse alternating refers to a small calcaneonavicular contact that sometimes occurs in a serial condition or to a significant displacement of the tarsal articulations in a different (proximodistal) axis. The proximodistal arrangement of the joints could be functionally significant. Two new states are observed and defined: (i) 'flipped serial', present in Macropodidae, in which the calcaneocuboid articulation is medially displaced and significantly larger than the astragalonavicular contact, but the relationships between proximal and central elements are one to one; and (ii) 'distal cuboid', an extreme proximodistal displacement of the astragalonavicular joint. Serial and alternating, as originally defined (i.e. without any reference to which bone contacts which), seem to be the best states for classifying tarsal arrangement though as the disposition of distal or central bones in relationship to proximal bones.

Keywords: Mammalia; Meridiungulata; morphology; phylogeny; tarsus.

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Figures

Figure 1

Figure 1

Outline of serial, alternating tarsal and previously proposed reverse alternating tarsal arrangements.

Figure 2

Figure 2

(A) Plantar view of right astragalus of Tremarctos ornatus (

MLP

1.I.03.62); shading in the surface of contact of cuboid bone. (B) Plantar view of left astragalus of Notostylops sp. (

LIEB

‐

PV

4016); shading in the space between navicular and sustentacular facets. (C) Plantar view of left astragalus of: (1) a juvenile of perissodactyl Tapirus terrestris (

MLP

1); (2) an adult Tapirus terretris (

MLP

1070). In yellow shading, ectal facet; in green shading, sustentacular facet; in pink shading, anterior astragalocalcaneal facet. Observe the disconnection between sustentacular and anterior astragalocalcaneal facet in the adult. (D) Left astragalus of Lagostomus maximus (

MLP

1683). Plantar view, slightly oblique to better observe the facets. Facets as previous image; in soft dark blue shading, the facet for the sesamoid. Scale bar: 10 mm.

Figure 3

Figure 3

Proximal view of the left navicular of proterotheriid Eoauchenia (

MLP

48‐

XII

‐16‐1). In light blue shading, sesamoid facet; in green shading, calcaneal facet; in red shading, astragalar facet. Scale bar: 10 mm.

Figure 4

Figure 4

Presence of facets mapped over: (A) strict consensus cladogram from the analysis of Billet (2011, fig. 9); and (B) Bayesian consensus tree of

COL

1 protein sequence data, with chicken (Gallus) as outgroup from the analysis of Welker et al. (2015).

Figure 5

Figure 5

Displacement of joints in the proximodistal axis mapped over: (A) strict consensus cladogram from the analysis of Billet (2011, fig. 9); and (B) Bayesian consensus tree of

COL

1 protein sequence data, with chicken (Gallus) as outgroup from the analysis of Welker et al. (2015).

Figure 6

Figure 6

Displacement of joints in the transversal axis mapped over: (A) strict consensus cladogram from the analysis of Billet (2011, fig. 9); and (B) Bayesian consensus tree of

COL

1 protein sequence data, with chicken (Gallus) as outgroup from the analysis of Welker et al. (2015).

Figure 7

Figure 7

(A) Dorsal view of right tarsus of Eutypotherium lehmannnistchei (

MLP

12‐1701). (B) Dorsal view of right tarsus of Theosodon sp. (

MLP

700). In blue shading, astragalus; in green shading, navicular; in red shading, calcaneus; in yellow shading, cuboid. Scale bar: 10 mm.

Figure 8

Figure 8

(A) Dorsal view of right tarsus of Macropodidae indet. (

MLP

951). (B) Dorsal view of right tarsus of Loxodonta africana (

MLP

1123; specimen in exposition). In blue shading, astragalus; in green shading, navicular; in red shading, calcaneus; in orange shading, cuboid.

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