Holmesina cryptae Moura, Góis, Galliari & Fernandes, 2019

Moura, Jorge Felipe, Góis, Flávio, Galliari, Fernando Carlos & Fernandes, Marcelo Adorna, 2019, A new and most complete pampathere (Mammalia, Xenarthra, Cingulata) from the Quaternary of Bahia, Brazil, Zootaxa 4661 (3), pp. 401-444 : 405-406

Holmesina cryptae Moura, Góis, Galliari & Fernandes sp. nov.

Figs. 2–23

Type locality. Two specimens (holotype and paratypes) associated to three juvenile individuals of Glyptodontinae were found in the inferior level of the cave Gruta da Lapinha in Iramaia Municipality, Bahia state, Brazil.

Etimology. “ Gruta ” (cave/grotto in Portuguese) comes from the Latin “ crypta ” (crypt). At the entrance of Lapinha Cave, there is an altar that is used by Catholic believers, and the skeletons of the pampatheres were found in a chamber below which is analogous to a crypt, reinforcing the choice of this name.

Holotype. LPP-PV-001: Skull. Maxillary teeth (15 – right and left: Mf3, Mf4, Mf5, Mf6, Mf7, Mf8 and Mf9; and left: Mf2). Mandibles (fragmented). Hyoid bones (right stilohyal and epihyal). Cervical vertebrae (C1–C7). Thoracic vertebrae (T1–T10). Lumbar vertebra (L1). Synsacral vertebrae (S1–S11). Caudal vertebrae (Cd1–Cd22). Chevron bones (5). Scapulae (incomplete). Clavicles (right and left). Manubrium. Ribs (10 – vertebral portion). Humeri (right and left). Radii (right and left). Ulnae (right and left). Right trapezoid bone. Hamate bones (right and left), Triquetral bones (right and left). Lunate bones (right and left). Pisiform bones (right and left). Anterior distal phalanges (right and left: I, II, III, IV, V). Anterior medial phalanges ( II, III and IV). Anterior proximal phalanges (right: I, II, III, IV and V; left: I, II, III, IV and V). Metacarpi (right and left: I, II, III and IV). Sesamoid bones in right manus (3), and right and left palmar bones. Pelvis. Femora (right and left). Patellae (right and left). Tibiofibulas (right one is complete; distal and proximal epiphyses of the left one). Calcanei (right and left). Astragali (right and left). Naviculars (right and left). Cuboids (right and left). Metatarsi (right and left: II, III, IV and V). Posterior distal phalanges (right: I, II, IV and V; left: II, III, IV and V). Posterior medial phalanges (right: II, IV and V; left: II, III, IV and V). Posterior proximal phalanges (right: III; left: I, II, III, IV and V). Right and left plantar bones. Osteoderms (more than 1000 and fragments).

Paratype. LPP-PV-002: Skull almost complete. Maxilary teeth (right and left M1–M9). Mandible (complete in right side with nine teeth; the left one is fragmented). Hyoid bones (basithyrohyal). Cervical vertebrae (C1–C7). Thoracic vertebrae (T1–T10). Lumbar vertebra (L1). Synsacral vertebrae (S1–S11). Caudal vertebrae (Cd1–Cd4). Chevron bones (4). Manubrium. Ribs (fragments). Incomplete humeri. Radii (right and left). Ulnae (right and left). Pelvis. Femora (distal and proximal epiphyses). Patellae (right and left). Tibiofibulas (fragments). Calcaneus (left). Astragalus (left). Cuboids. Anterior distal phalanges (right and left I, II, III, IV and V). Osteoderms (more than 200 and fragments) .

Age: SALMA Lujanian (Late Pleistocene).

Diagnosis. Holmesina cryptae sp. nov. differs from other species of pampatheres in having: a less elevated frontal crests in frontal bones; a less robust, thin and low sagittal crest; a nuchal crest narrower in the lateral borders; a robust and pointed mastoid process in the petrosal bones; a C-shaped notch anteriorly in lateral view of premaxillas; a larger infraorbital foramen in maxillas; very robust postorbital apophyses (postorbital process of jugal). The shape of the H. cryptae sp. nov. skull is more triangular than the other species and possesses robust zygomatic arches with an orthogonal process zygomatic of squamosal. The skull also shows the most robust parietal region, and the widest and more robust snout. The molariforms are farthest from the sagittal line than any of the compared species. Holmesina cryptae sp. nov. presents osteodermal ornamentations less marked than in Scirrotherium carinatum , Kraglievichia , H. paulacoutoi and H. occidentalis , however more defined and punctured than in Pampatherium . Anteriorly to the first lower molariform (mf1) in the dentary, there is a toothless pointed region longer than in any other species of Holmesina , however it is less developed than in P. typum and P. humboldtii . The first three anterior upper molariforms (Mf1–Mf3) show an oval morphology as in the rest of the species. The morphology of the other molariforms is more similar to other Holmesina species (short and wide, eight-shaped), with reduced interdental spaces, in contrast to Pampatherium . The mandible is much less robust than in P. humboldtii , horizontal and ascending rami narrower than in P. humboldtii . In the ulna, the lateral and medial fossae are superficial and the styloid apophysis is rounded. The femur shows a deeper and more delimited trochanteric fossa than other species of Holmesina . The patella has a longer and straighter apex in comparison to H. floridana .

Figures

FIGURE 2. Exterior and interior of Lapinha Cave. A, External view; in the right below of the picture, notice the cave entrance closed by a gate. B–C, Specimens of Holmesina cryptae sp. nov. buried in the cave, B, LPP-PV-001; and C, LPP-PV-002 at the moment of the excavation.

FIGURE 3. Complete skeleton of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Scale bar = 300 mm.

FIGURE 4. Osteoderms of Holmesina cryptae sp. nov. A–D, LPP-PV-001, A, osteoderm of cephalic shield. B, fixed osteoderm of scapular buckler in dorsal and lateral views. C, fixed osteoderm of pelvic buckler. D, movable osteoderm in dorsal and right lateral views. E, (LPP-PV-002) semimovable osteoderm of pelvic buckler, F, (LPP-PV-001) semimovable osteoderm of scapular buckler. G, (LPP-PV-002) caudal osteoderm (proximal portion of the tail). H–I, LPP-PV-001, H, caudal osteoderm (median portion), and I, caudal osteoderm (distal portion). Scale bar = 20 mm.

FIGURE 5. Skull of Holmesina cryptae sp. nov. (holotype, LPP-PV-001), dorsal and ventral views. A, dorsal; and B, ventral views. Scale bar = 50 mm.

FIGURE 6. Skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), dorsal and ventral views. A, dorsal; and B, ventral views. Scale bar = 50 mm.

FIGURE 7. Skull of Holmesina cryptae sp. nov. (holotype, LPP-PV-001), lateral and posterior views. A, lateral; and B, and occipital views. Scale bar = 50 mm.

FIGURE 8. Skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), lateral and posterior views. A, lateral; and B, and occipital views. Scale bar = 50 mm.

FIGURE 9. Ventrolateral view of the skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002) focusing the left auditory region. aptt, anteroventral process of tegmen tympany; as, alisphenoid; bb, bony bridge between tympanohyal and crista interfenestralis; bcc, basicochlear commissure; bo, basioccipital; bs, basisphenoid; cf, carotid foramen; cp, crista parotica; eam, external auditory meatus; eo, exoccipital; f, frontal; fo, foramen ovale; fr/sof, fused foramen rotundum and sphenorbital fissure; fm, foramen magnum; fs, facial sulcus; gf, glenoid fossa; hf, hypoglossal foramen; itc, infratemporal crest; Mf9, ninth upper molariform; mp, mastoid process (pop, paroccipital process of petrosal of Gaudin, Lyon (2017)); mx, maxilla; nc, nuchal crest; occ, occipital foramen; of, optic foramen; pcp, paracondylar process of exoccipital (= paroccipital process); pe, petrosal; pf, piriform fenestra; pgf, postglenoid foramen; pgp, postglenoid process; pr, promontorium of petrosal; pt, pterygoid; pt/pl, fused pterygoid and palatine; spf/cpf, fused sphenopalatine and caudal palatine foramina; smf, suprameatal foramen; stf, stylomastoid foramen; sq, squamosal; th, tympanohyal; ttf, tensor tympani fossa on epitympanic wing of petrosal; zp, zygomatic process of squamosal. Perspective figure, dimensions must be compared to Figs. 6 and 8. Based on Gaudin & Lyon (2017).

FIGURE 10. Ventral view of frontal plane of skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), computed tomographic scan image, –15 mm from the palate surface. Abbreviations: ju, jugal bone; ma, maxilla; Mf1, first molariform; Mf2, second molariform; nc, nasal cavity; pm, premaxilla; pt, pterygoid; zpm, zygomatic process of maxilla. Scale bar = 35 mm

FIGURE 11. Schematic representation of the right upper tooth rows of South American Quaternary pampatheres.A, Holmesina cryptae sp. nov. (paratype, LPP-PV-002). B, H. rondoniensis (holotype, MERO-P-002). C, H. occidentalis (ROM 3881). D, H. paulacoutoi (holotype, MCL-501/01). E, H. septentrionalis (HMNS 173). F, Pampatherium typum (MACN Pv 11543). G, P. humboldtii (MHD-P-28). In black, absent molariforms (shape inferred from the alveolus); in some cases, the occlusal surface is totally or partially destroyed, thus some illustrations were completed from the left tooth row of the same exemplar. Scale bar = 30 mm.

FIGURE 12. Sagittal planes of skull of Holmesina cryptae sp. nov. (paratype, LPP-PV-002), computed tomographic scan images. A, 1 mm; B, 2 mm; and C, 4 mm from the median sagittal plane.Abbreviations: ec, endocranial cavity; ma, maxilla; Mf1, first molariform; Mf2, second molariform; nc, nasal cavity; pm, premaxilla. Scale bars = 35 mm.

FIGURE 13. Right hemimandible of Holmesina cryptae sp. nov. (paratype, LPP-PV-002).A, lateral; B, medial; and C, occlusal views. Scale bar = 50 mm.

FIGURE 14. Hyoid apparatus of Holmesina cryptae sp. nov. A–B, lateral view (holotype, LPP-PV-001); A, stylohyal; B, epihyal; and C, (paratype, LPP-PV-002) ventral view of the basithyrohyal co-ossification (= v-bone). Scale bar = 30 mm.

FIGURE 15. Vertebral column of Holmesina cryptae sp. nov. A-E, presynsacral vertebrae of the holotype (LPP-PV-001). A, dorsal view of the articulated presynsacral vertebrae. B–C, mesocervical bone in B, dorsal; and C, lateral views. D–E, postcervical bone in D, dorsal; and E, lateral views. F, xenarthrous vertebrae of the paratype (LPP-PV-002). Scale bars = 50 mm. G, caudal vertebrae of the holotype (LPP-PV-001); Scale bar = 100 mm.Abbreviations: C1–C7, cervical vertebrae; L1, first lumbar vertebra; T4–T10, thoracic vertebrae; m, mesocervical bone (C2+C3+C4+C5); p, postcervical bone (T1+T2+T3).

FIGURE 16. Chevrons of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Scale bar = 30 mm.

FIGURE 17. Vertebral portions of ribs of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Second to tenth right ribs. Scale bar = 30 mm.

FIGURE 18. Thoracic bones and clavicle of Holmesina cryptae sp. nov. (holotype, LPP-PV-001).A, right first anterior rib; B, manubrium; and C, right clavicle. Abbreviations: ace, acromial end; ssn, suprasternal notch; ste, sternal end; vee, vertebral end. Scale bar = 30 mm.

FIGURE 19. Right scapula of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). Scale bar = 30 mm.

FIGURE 20. Pelvis of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A, dorsal; and B, lateral views. Abbreviations: ac, acetabulum; lsc, lateral synsacral crest; of, obturator foramen; sif, synsacroischial foramen; ssa, synsacral ala. Scale bar = 50 mm.

FIGURE 21. Right forelimb bones of Holmesina cryptae sp. nov. (holotype, LPP-PV-001).A–B, humerus in A, anterior; and B, posterior views. C–D, radius in C, anterior; and D, posterior views. E–H, ulna in E, lateral; F, anterior; G, posterior; and H, medial views. Abbreviations: hsp, hook-shaped projection; scf, supracondylar foramen; tn, trochlear notch. Scale bar = 50 mm.

FIGURE 22. Right hind limb bones of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A–B, femur in A, anterior; and B, posterior views. C–D, patella in C, anterior; and D, posterior views. E–F, tibiofibula in E, anterior; and F, posterior views. Scale bar = 50 mm.

FIGURE 23. Right astragalus and calcaneus of Holmesina cryptae sp. nov. (holotype, LPP-PV-001). A–B, astragalus in A, dorsal; and B, ventral views. C–D, calcaneus in C, dorsal; and D, ventral views. Scale bar = 50 mm.