Anomalous Erosion Patterns on the Cheek Teeth of Babirusa (Babyrousa Perry, 1811.)
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Canadian Journal of Zoology Anomalous erosion patterns on the cheek teeth of babirusa (Babyrousa Perry, 1811.) Journal: Canadian Journal of Zoology Manuscript ID cjz-2020-0173.R1 Manuscript Type: Article Date Submitted by the 08-Sep-2020 Author: Complete List of Authors: Macdonald, Alastair; The University of Edinburgh, Royal (Dick) School of Veterinary Studies Is your manuscript invited for consideration in a Special Not applicableDraft (regular submission) Issue?: Molar teeth, premolar teeth, wear, fruit seed, molaires, prémolaires, Keyword: Musa L. © The Author(s) or their Institution(s) Page 1 of 38 Canadian Journal of Zoology 1 1 Anomalous erosion patterns on the cheek teeth of babirusa (Babyrousa Perry, 1811.) 2 Profils d’érosion irréguliers sur les dents jugales du babiroussa (Babyrousa Perry, 1811.) 3 4 5 A A Macdonald 6 7 Royal (Dick) School of Veterinary Studies 8 The University of Edinburgh 9 Easter Bush Campus 10 Midlothian EH25 9RG 11 Scotland Draft 12 13 [email protected] 14 15 16 Key words: Molar teeth, premolar teeth, wear, fruit seed, Musa L., babirusa, Babyrousa 17 Mots-clés : molaires, prémolaires, usure, graines de fruit, Musa L., babirusa, Babyrousa 18 19 20 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 2 of 38 2 21 A A Macdonald 22 Anomalous erosion patterns on the cheek teeth of babirusa (Babyrousa Perry, 1811.) 23 Profils d’érosion irréguliers sur les dents jugales du babiroussa (Babyrousa Perry, 1811.) 24 25 26 Abstract 27 28 Two types of anomalous erosive wear, designated ‘Wear’ and ‘Groove’, were found in 29 the molar teeth of adult babirusa species (Babyrousa Perry, 1811.) from Buru or the 30 Sula Islands and Sulawesi, Indonesia. A survey of 590 specimens of babirusa crania and 31 mandibles from international museumDraft and private collections revealed their presence 32 in 20 of 187 specimens from Buru or the Sula Islands, in 99 of 391 specimens from 33 Sulawesi, and in four of 31 subfossil specimens from South Sulawesi. ‘N_Wear’ 34 comprised a discrete indentation or almost circular erosion of the enamel and 35 underlying dentine in a molar tooth; ‘W_Wear’ comprised tooth indentations of 36 irregular shape larger than 7x7mm in size; ‘E_Wear’ comprised extensive tooth 37 indentation. ‘Wear’ anomalies were found on 10% of specimens from Buru or the Sula 38 Islands, and 17% of all Sulawesi crania and mandibles. It was hypothesised that tooth 39 grinding of the abrasive seeds of the wild banana Musa balbisiana Colla initiated this 40 anomaly. ‘Groove’ was a deep, oval-shaped cleft cut into the enamel and dentine 41 between two molar teeth. ‘Groove’ anomalies were evident in 3% of Buru or the Sula 42 Islands specimens and 12% of all Sulawesi specimens. No conclusion was reached 43 regarding the initiator of the ‘Groove’ anomaly. 44 45 © The Author(s) or their Institution(s) Page 3 of 38 Canadian Journal of Zoology 3 46 Draft © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 4 of 38 4 47 1. Introduction 48 49 The babirusa (Babyrousa Perry, 1811.) is a suid endemic to eastern Indonesia, 50 inhabiting the islands of Buru, the Sula Islands of Sehu, Taliabu and Mangole, the island 51 of Sulawesi and the Togian Islands (Macdonald and Pattikawa 2017; Macdonald et al. 52 2017; Macdonald 2018;). Early anatomical investigations described the appearance of 53 the teeth in the adult male babirusa (Nieuhof 1682; Ducrotay de Blainville 1839-64; 54 Owen 1840-1845; Stehlin 1899, 1900). More recently the growth pattern of the 55 maxillary canine teeth has been described (Macdonald et al. 2016; Macdonald and Shaw 56 2018), and attention drawn to the wear patterns present on those teeth (Macdonald 57 2017). Draft 58 59 Recent studies have investigated the anatomy and wear of babirusa cheek teeth 60 (Macdonald 2019). In the process a small number of anomalous wear patterns were 61 observed. Two examples of these had been noticed earlier and briefly commented upon 62 in a study of dental pathology in Sus celebensis Müller and Schlegel, 1843 from Sulawesi 63 (Kaudern and Forshufvud 1938). The conclusion reached was that the erosion pattern 64 was evidence of resolved dental decay. However, closer inspection of the teeth of a 65 larger number of skulls soon indicated that there was no evidence of bacteria-induced 66 dental caries in these instances. An alternative hypothesis was that they may have been 67 due to specific elements of dietary intake. The current study sought to examine the 68 anomalous wear patterns in more detail, and to investigate their possible causes. 69 70 2. Materials and methods 71 © The Author(s) or their Institution(s) Page 5 of 38 Canadian Journal of Zoology 5 72 A survey was made of 621 specimens of babirusa (Babyrousa spp) crania, mandibles 73 and sub-fossil teeth from museum and private collections around the world (Macdonald 74 and Shaw 2018; Frantz et al. 2018). Many of these had recorded location data 75 associated with them, but many others could only be identified to islands or island 76 groups (viz. ‘Buru or the Sula Islands’, and ‘Sulawesi’). From among these, one female 77 and 19 male specimens from Buru or the Sula Islands (of 187) (Babyrousa babyrussa 78 Linnaeus, 1758), and 99 specimens from Sulawesi comprising eight females and 91 79 males (of 391 from Sulawesi) (Babyrousa celebensis Deninger, 1909), were found to 80 have examples of anomalous wear of the mandibular and/or maxillary cheek teeth; 31 81 subfossil babirusa M2 and M3 teeth from South Sulawesi (the south-west peninsula) (B. 82 celebensis) were studied, of which fourDraft showed comparable anomalous wear (Tables 1 83 and S1). 84 85 All specimens were photographed, and where possible, particular photographic 86 attention was paid to the occlusal surfaces of those skulls showing anomalous patterns 87 of wear. To enable anatomical clarity in the description of the wear patterns, the dental 88 anatomical nomenclature of the babirusa teeth was derived from that published for 89 Suidae by van der Made (1996). The main cusps on the mandibular molars are called 90 protoconid, metaconid, hypoconid, entoconid, and pentaconid (Figure 1). A smaller 91 sixth cusp can be seen on the M3 and it is named hexaconid (Figure 1b). The main cusps 92 on the maxillary molars are called paracone, protocone, metacone, tetracone, and 93 pentacone (Figure 2). 94 95 Visits were made in 2017 and 2018 to Buru, and in 2018 to Sehu and Taliabu Islands in 96 the Sula Archipelago, during which information on the diet of babirusa was offered by © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 6 of 38 6 97 village residents (Macdonald and Pattikawa 2017; Macdonald et al. 2017). Photographs 98 and samples of local plant fruits and seeds were collected there for subsequent closer 99 inspection. Scanning electron microscopy was carried out on seeds of Musa L. After 100 washing in water and olive oil, the dried Musa seeds were mounted on aluminium stubs 101 with carbon tabs attached, sputter coated with 20 nm gold palladium, and viewed using 102 a Hitachi S-4700 scanning electron microscope. 103 104 3. Results 105 106 In general terms, the cheek teeth of babirusa exhibited two types of anomalous tooth 107 wear. These have been termed ‘Wear’Draft indentation and ‘Groove’ indentation. The term 108 ‘Wear’ has been further sub-categorised as narrow (N_Wear), wide (W_Wear) and 109 expanded (E_Wear). These observations are summarised in Table 1. 110 111 The first sub-category, ‘N_Wear’, comprised a discrete indentation or almost circular 112 erosion of the enamel and underlying dentine in a molar tooth (Figure 3). The width of 113 these varied between 4x4mm and 7x7mm. These were found on the second and third 114 molar teeth of 13 specimens (Table 1). Four specimens from Buru or the Sula Islands 2 115 showed five examples of ‘N_Wear’ (two on M3 and three on M ), with none present 116 among the 64 specimens known to be from Buru. Nine specimens from Sulawesi 2 3 117 exhibited 20 examples of ‘N_Wear’ (one on M2, eight on M3, seven on M , and four on M 118 (Figure 4), with none on the 49 specimens and 31 subfossil teeth identified as coming 119 from central or southern peninsulae of that island (Table 1). They were also not present 120 on specimens from the Togian Islands. They were also not observed on mandibular or 121 maxillary first molar teeth. © The Author(s) or their Institution(s) Page 7 of 38 Canadian Journal of Zoology 7 122 123 The sub-category ‘W_Wear’ comprised tooth indentations of somewhat irregular shape 124 and larger than 7x7mm in size (Figure 5a). These were found on the molar teeth of 28 125 specimens, five of which were from Buru or the Sula Islands (Table 1). There were nine 126 examples on the teeth of one specimen from the Sula Islands (four on M2, three on M3, 2 127 and two on M ), and two examples on the M3 of a specimen from Buru or the Sula 128 Islands. On the 23 Sulawesi specimens there were 24 indentation examples on M2, 26 on 2 3 129 M3, 29 on M , and 11 on M . Again, there were none on the 49 specimens and 31 130 subfossil teeth identified as coming from the central regions or the southern peninsulae 131 of that island (Table 1). They were also not present on specimens from the Togian 132 Islands. Draft 133 134 The sub-category ‘E_Wear’ comprised extensive tooth indentations, larger than those of 135 ‘W_Wear’, often of somewhat irregular shape and often eroding two or more cusps 136 (Figures 5b,c).