Anomalous Erosion Patterns on the Cheek Teeth of Babirusa (Babyrousa Perry, 1811.)

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.

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.)

5 A A Macdonald

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]

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

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.)

26 Abstract

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.

Draft

47 1. Introduction

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.

70 2. Materials and methods

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).

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).

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

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.

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). They were found on 10 specimens from Buru or the Sula Islands and on

137 46 Sulawesi specimens (Table 1). Owing to the variability between specimens in the

138 relative amounts of ‘E_Wear’ it was not possible to quantify its occurrence in the same

139 way as that for ‘N_Wear’ and ‘W_Wear’. Instead, solely the number of molar teeth

140 showing ‘E_Wear‘ was counted. ‘E_Wear ‘was shown by all upper and lower M2 and M3

3 141 teeth of the Buru specimens, in the Sula specimens by one M2, two M3 and two M , and

2 3 142 in the Buru or Sula specimens by 15 M2, ten M3, 15 M and eight M teeth. ‘E_Wear’

2 3 143 among the Sulawesi specimens was shown by 54 M2, 52 M3, 53 M and 38 M teeth.

144

145 The second type of tooth anomaly found, ‘Groove’, is a deep, roughly oval-shaped cleft

146 cut into the enamel and underlying dentine between two molar teeth (Figure 6). The

147 general presence of ‘Groove’ is summarised in Tables 1 and S1. The extent and depth of

148 these localised inter-tooth erosions were variable (Figure 7a,b,c). These anomalies were

149 present in five of the specimens from Buru or the Sula Islands on 11 occasions; bi-

150 laterally between the M1 and M2 in two specimens, bi-laterally between the M2 and M3

151 in three specimens and in one left mandible. They were also present in 51 of the

152 specimens from Sulawesi on 144 occasions; bi-laterally between the M1 and M2 in nine

153 specimens, in the left mandible of three specimens and one right mandible; bi-laterally

154 between the M2 and M3 in 20 skulls, on the left mandible four times and on the right

155 mandible four times; bi-laterally between the M1 and M2 in nine specimens, on the left

156 maxilla once, and on the right maxilla twice; and bi-laterally between the M2 and M3 in 157 25 specimens, on the left maxilla twice,Draft and on the right maxilla once. 158

3 159 ‘Groove’ anomalies were evident in one M3 and three M subfossil babirusa teeth from

160 South Sulawesi (Figure 8). There was no sign of similar ‘Groove’ anomalies on the teeth

161 of babirusa from East-central or West-central Sulawesi, South-east Sulawesi or the

162 Togian Islands (Table 1).

163

164 ‘Groove’ anomalies did not always occur where ‘Wear’ anomalies were present (Table

165 S1). Among those from the Buru or the Sula Islands, two specimens had ‘Grooves’ but no

166 ‘Wear’ anomalies, and 15 had ‘Wear’ anomalies with no ‘Grooves’ present. Both

167 anomalies were present on only three specimens. Likewise, among those specimens

168 from Sulawesi, 36 specimens had ‘Grooves’ but no ‘Wear’ anomalies, and 52 had ‘Wear’

169 anomalies with no ‘Grooves’ present. Both anomalies were present on 15 specimens.

170

171 4. Discussion

172

173 The relative rarity of the anomalous tooth wear present in the current study was note-

174 worthy, but had barely been mentioned in the Babyrousa literature (Kaudern and

175 Forshufvud 1938). Its scarcity in any one museum collection may have been a factor.

176 Only 18 examples of ‘Wear’ were found among the 187 specimens from Buru or the Sula

177 Islands, representing 10% of the specimens available (Table 1). However, the larger

178 number, 67 examples of ‘Wear’, found among the 391 specimens from Sulawesi,

179 represented a greater incidence, i.e. 17%, of specimens available from that island (Table

180 1). Both groups exhibited patterns of cheek tooth wear distinctly different from those

181 reported earlier (Macdonald 2019). The focal nature of the tooth erosion in the current 182 study was in contrast to the more generalDraft wear of the occlusal surface of the teeth seen 183 before. Additional details of the geographical distribution of ‘Wear’ were also evident.

184 Only a single specimen was found among the 64 specimens known to have come from

185 Buru (2%), with a slightly larger number (4 specimens) representing a larger

186 proportion (12%) of the specimens (33) from the Sula Islands. No instances were found

187 among the 12 specimens from the Togian islands, nor were there any among the 14

188 specimens from South-east Sulawesi (Table 1).

189

190 Owing to small sample sizes for these latter groups, the results presented here should

191 be treated with caution. The northern peninsula of Sulawesi (North-east Sulawesi and

192 North-west Sulawesi) as well East-central Sulawesi exhibited larger numbers and

193 percentages of ‘Wear’ (17%, 23% and 25% respectively) than other parts of Sulawesi

194 (Table 1). Interestingly, the percentage of those babirusa specimens from unknown

195 regions of Sulawesi (19%) was similar to those found in the north and east of the island.

196 Although the ‘Groove’ anomaly occurred less frequently, the pattern of occurrence was

197 somewhat similar. It was only present in about 3% of the 187 specimens from Buru or

198 the Sula Islands. This compared to about 13% of the 422 specimens from Sulawesi,

199 represented largely by those from the northern peninsula and the subfossils from South

200 Sulawesi (Table 1).

201

202 There was no identifiable difference between males and the small number of female

203 specimens with respect to anomalous tooth wear patterns (Table S1).

204

205 The narrow ‘N_Wear’ was found in skulls of all ages as evaluated according to maxillary

206 canine tooth length (Macdonald and Shaw 2018). Male skulls designated as being older 207 had larger numbers of ‘W_Wear’ andDraft ‘E_Wear’ anomalies than younger skulls (Table S1). 208 In older babirusa it often appeared that adjacent ‘Wear’ and ‘Groove’ anomalies had

209 coalesced to form elongated and widened depressions in the dentine of the tooth with

210 branching into the surrounding enamel (Figures 5b, 5c, 7c).

211

212 The location of the ‘Wear’ on the tooth was as indicated in Figures 3-6. ‘N_Wear’ was

213 usually first located in the furrows and fossae between cusps, e.g. on maxillary teeth

214 between the protocone and the tetracone (Figures 3 and 4) and on mandibular teeth

215 between the protoconid and metaconid (Figure 5a). The patterns of ‘Wear’ suggested

216 that they were initially localised, rounded erosions of some depth (Figure 4a),

217 subsequently becoming widened (Figure 4b), and expanding over other parts of the

218 occlusal surface of the tooth (Figures 4c and 5).

219

220 ‘Wear’ of M3 teeth was consistently less frequent than ‘Wear’ of the other upper and

221 lower M2 and M3 cheek teeth; of the Buru or Sula Island specimens there was ‘Wear’ of

3 222 some sort on 10 M teeth compared to ‘Wear’ on 19 M3 teeth; of the Sulawesi specimens

3 223 there was ‘Wear’ of some sort on 42 M teeth compared to ‘Wear’ on 72 M3 teeth. The

2 224 number of specimens from Buru and the Sula Islands with ‘Wear’ on M and M2 was 22

2 225 and 20 respectively. The number of specimens from Sulawesi with ‘Wear’ on M and M2

226 was 81 and 66 respectively.

227

228 The specimen differences in the occurrence of ‘Wear’ and ‘Groove’ anomalies was

229 noteworthy (Table S1). Among those from the Buru or the Sula Islands, 75% had ‘Wear’

230 anomalies with no ‘Grooves’ present. Only 15% had both anomalies. Among the

231 specimens from Sulawesi, 53% had ‘Wear’ anomalies with no ‘Grooves’ present. Again, 232 only 15% had both anomalies. So a largerDraft proportion on Sulawesi (36%) had only 233 ‘Groove’ anomalies compared to Buru or Sula Islands (10%).

234 The evidence indicates that the factors causing these two anomalous wear patterns are

235 different. Although the babirusa from the northern peninsula of Sulawesi are known to

236 ingest the mud and waters from hot springs (Patry et al. 1995; Clayton and MacDonald

237 1999), this inorganic material is more likely to induce general wear of the occlusal

238 surfaces of the cheek teeth rather than the anomalous ‘Wear’ seen. The ‘Groove’

239 anomaly bore some similarities to human dental wear, specifically incisal edge notching,

240 due to long-term consumption of sunflower seeds (Helianthus annuus L.); the shell of

241 the sunflower seed is cracked by keeping it vertical between the upper and lower

242 incisors and applying some amount of force to crack it (Rath et al. 2017). Human

243 consumption of dried roasted watermelon seeds (Citrullus lanatus (Thunb.) Matsum. and

244 Nakai) is also known to have significant abrasive effects on the levels of tooth surface

245 loss in central incisors and canine teeth (Riyad et al. 2011). The relationships between

246 mammalian tooth wear and diet have been reviewed (Ungar 2015). Large hard food

247 particles, such as seeds from fruits, fracture (brittle) enamel. Some investigative work

248 on the fracture behaviour of pig (Sus scrofa Linnaeus, 1758) molar cusps has been

249 published (Popowics et al. 2001).

250 The babirusa is recognised to be an omnivore, but with preference for fruits and seeds

251 (Leus et al. 2001). Babirusa do not dig into hard ground for roots; they lack the large

252 pre-nasal bone of S. scrofa and S. celebensis that enables the nose of those pigs to act as

253 strong digging instruments (Macdonald 2018). Babirusa tend to harvest leaves and fruit

254 from above the ground (Leus 1996; Tjiu and Macdonald 2016). Recent studies in

255 zoological collections demonstrate that they will even wash food that has fallen onto the

256 ground (Ito et al. 2017). Draft 257 Although a full understanding of the natural diet of babirusa awaits detailed

258 investigation, several studies have provided some insight. The following genera/species

259 of fruit and seeds have been identified in the stomach contents of babirusa from the

260 northern peninsula of Sulawesi: Arenga pinnata (Wurmb) Merr., Artocarpus spp.,

261 Calophyllum soulattri Burm. f., Dillenia serrata Thunb., Dracontomelon dao (Blanco)

262 Merr. and Rolfe, Ficus Röding, Pangium edule Reinw. and Pothoidium lobbianum Schott

263 (Clayton 1996). Recent studies by means of interviews and trail cameras have extended

264 these data to Buru and the Sula archipelago (Tjiu and Macdonald 2016; Macdonald and

265 Pattikawa 2017; Macdonald et al. 2017). The fruits of the following were identified on

266 Buru: Calamus zollingeri Becc., Canarium indicum L., Daemonorops robusta Warb. Ex

267 Becc., Ficus spp., Hornstedtia rumphii (Sm) Valeton, Passiflora foetida L., Rubus

268 fraxinifolius Poir. and Shorea Roxb. ex C.F. Gaertn. On the Sula Islands the fruits of

269 Castanopsis buruana Miq., Ficus spp., Hornstedtia rumphii, Shorea spp. and four other

270 species, katoan, ta meking and two plants called galoba (but not scientifically identified)

271 were recorded. Among the Ficus spp. that were available to the babirusa were F.

272 deltoidei Jack., F. hispida L.f. and F. variegate Blume. Other candidate plants in the region

273 with seeds are: Adenanthera pavonine L. (red-bead tree), Anisoptera spp., Areca vestiaria

274 Giseke, Arenga Labill., Caryota L., Coix lacryma-jobi L. (Jobes tears), Dinochloa Büse,

275 Dipterocarpus C.F. Gaertn., Elaeocarpus L., Hopea Roxb., Musa spp., Pinanga Blume, 1838,

276 Prunus L., Terminalia L. (e.g. T. catappa L.) and Ulmaciae Mirb. The seeds of some of

277 these species are relatively soft, and although this does not necessarily reduce their

278 ability to cause tooth wear (O’Toole and Mullan 2018), they were thought unlikely to be

279 the primary cause of the anomalies seen.

280 281 The structure of the ‘N_Wear’ anomalyDraft suggested that the seed of a wild Musa sp. might 282 be one candidate for the formation of this ‘Wear’ anomaly. Experience of animals in

283 zoological collections had indicated that the modern edible seedless bananas (derived

284 from Musa acuminata Colla and Musa balbisiana Colla) were attractive to the babirusa

285 (Leus 1996; Leus et al. 2001). Wild Musa ssp. are giant herbs with perennial rhizomes,

286 their pseudostem being comprised of long, stiff leaf-sheaths that are rolled round one

287 another (Morton 2001). Six species of Musa are known to grow in Sulawesi: Musa

288 acuminata; M. balbisiana; M. borneensis var donggalaensis Sulis.; M. celebica Warb. Ex

289 K.Schum.; M. itinerans Cheesman and M. textilis Nèe sensu Baker (Nasution and Yamada

290 2001; Sulistyaningsih et al. 2014 a,b; Sulistyaningsih 2016; Hastuti et al. 2019). The

291 current study also found that at least two of these species of Musa can be found on Buru:

292 M. acuminata and M. balbisiana (Sulistyaningsih personal communication). An un-

293 identified Musa specimen from Buru is also in the herbarium collection of the Royal

294 Botanical Garden Edinburgh; Van Balgooy 4564).

295 Recent botanical studies have shown that three species of wild banana grow in North

296 Sulawesi: M. acuminata, M. balbisiana and M. textilis (Nasution and Yamada 2001;

297 Sulistyaningsih et al. 2014 a,b; Hastuti et al. 2019). It was in North-east Sulawesi and

298 North-west Sulawesi that ‘Wear’ tooth erosion has been found (Table 1). Of the three

299 Musa species, the seeds of M. balbisiana appear particularly noteworthy; they have been

300 described as globular, wrinkled, and warty (Sulistyaningsih et al. 2014a; McGahan

301 1961). They were also reportedly very hard due to the high silica content of the

302 epidermis (McGahan 1961); each epidermal cell contains a multi-angular crystalline

303 silica body (Graven et al. 1996). It was hypothesised that consumption by the babirusa

304 of this species of banana could have led to the ‘Wear’ observed. Babirusa can easily 305 access the fruit situated several metresDraft above the ground; local correspondents on Buru 306 said that the babirusa easily shoulder-push the banana plant over onto the ground and

307 thereby reach not only the fruit but also the soft vegetable matter in the pseudostem.

308 Each M. balbisiana banana (botanically a ‘berry’) contained parallel rows of 150-170

309 seeds, (Figure 9a), each of which measures approximately 4.7mm in diameter (Figures

310 9b-d). It seemed feasible that the repeated grinding of these small, hard, silica-crystal

311 abrasive particles in the furrows between the cusps of the molar teeth could initiate

312 ‘Wear’ formation.

313 The seeds of the other species of Musa on Buru and Sulawesi did not appear to have the

314 same shape and material properties (Nasution and Yamada 2001; Sulistyaningsih et al.

315 2014b; Hastuti et al. 2019). The low frequency with which the ‘Wear’ anomaly was

316 found may reflect one or both of the following; local availability of the causal agent and

317 individual animal preference for that agent. Increased understanding will require

318 further research. Examination of 203 mandibles collected on Sulawesi between 1917

319 and 1919 (Kaudern and Forshufvud 1938; Macdonald, unpublished) from the other

320 endemic pig species, S. celebensis, revealed one specimen with a ‘Wear’ anomaly

321 between the left M2 and M3 and one specimen with a ‘Groove’ anomaly between the

322 right M1 and M2. Both mandibles were from North Sulawesi. Other mammalian and

323 avian species are known to feed on Musa. Seed dispersal of wild bananas is said to be

324 mostly carried out by foraging rodents, but long-tongued fruit-eating bats (Macroglossus

325 F.Cuvier, 1824.), civets, primates and birds play an important role too (Ge et al. 2005;

326 Corlett 2017).

327 It was unclear which of the proposed candidate plants might initiate the ‘Groove’ wear,

328 although various genera of the Dipterocarpaceae (Dipterocarpus, Hopea, Shorea) appear 329 to have the appropriate seed morphology.Draft Also, M. acuminata can have a somewhat 330 flattened seed (Chin 1996). It would also seem possible, from the shapes of the enlarged

331 ‘Groove’ anomalies (Figures 6, and 7b,c), that the seeds of Musa may have subsequently

332 been involved there too. Close examination of the surfaces of the ‘Wear’ and ‘Groove’

333 anomalies revealed score marks on the surfaces of the dentine and tooth enamel. It is

334 conceivable that the application of 3D microwear dental texture analysis (Ungar 2015;

335 Souron et al. 2015; Lazagabaster 2019; Rannikko et al. 2020) to the enamel and dentine

336 surfaces of these anomalies, and perhaps also to the epidermal surfaces of candidate

337 plant seeds, might shed additional light on the botanical agents causing anomalous wear

338 of the molar teeth of babirusa.

339

340 The relatively low frequency of anomalous tooth wear in babirusa from Buru and the

341 Sula Isles and its higher frequency in babirusa from the northern and southern

342 peninsulae of Sulawesi, were consistent with the geographical distribution of M.

343 balbisiana fruit with its abrasive seeds. The initiation of ‘Groove’ wear remains unclear,

344 but its enlargement may also be caused by the grinding of M. balbisiana seeds. Further

345 research is required to investigate these observations and the hypotheses they

346 generated.

347

348 Acknowledgements

349

350 The author gratefully acknowledges the hospitality shown by the Utomo family in

351 Jakarta and Ibu Mey and Bapak Eki, Ibu Fien and Bapak Ulis, and Ibu Kulung and Bapak

352 Tita for hospitality in Maluku. Thanks to Steven Mitchell at the University of Edinburgh 353 for assistance with scanning electronDraft microscopy. The author is grateful to the late 354 George Argent of the Royal Botanical Gardens Edinburgh for discussions and initial

355 guidance with respect to the botany of Musa, to Dr Mark Coode and Dr Mark Newman of

356 the Royal Botanical Gardens Edinburgh for discussions and guidance with respect to the

357 seeds of Sulawesi, to Dr John Dransfield of the Royal Botanic Gardens Kew for additional

358 botanical suggestions and to Lulut Dwi Sulistyaningsih, Bogor, Indonesia for email

359 conversations and helpful encouragement to learn more about Indonesian species of

360 wild bananas and the structural morphology of their seeds.

361

362 The author gratefully acknowledges the kind hospitality and support of Friederike

363 Johansson, Göran Nilson and Bianca Ziehmer during these studies. He would also like to

364 thank the curators and staff of the following museums for access to the babirusa

365 skeletal material that form part of their collections: Zoological Museum Amsterdam, The

366 Netherlands; Naturhistorisches Museum Basel, Switzerland; Museum für Naturkunde,

367 Berlin, Germany; Private H.M. Collection, Bogor, Indonesia; The Museum of Comparative

368 Zoology, Harvard University, Cambridge, USA; The Field Museum of Natural History,

369 Chicago, USA; Museum Zoologicum Bogoriense, Cibinong, Indonesia; Senckenberg

370 Naturhistorische Sammlungen Dresden, Germany; National Museums of Scotland,

371 Edinburgh, Scotland; Naturmuseum Senckenberg, Frankfurt am Main, Germany;

372 Göteborgs naturhistoriska museum, Göteborg, Sweden; Zoological Museum, Center of

373 Natural History (CeNak), Universität Hamburg, Germany;Zoologisk Museum,

374 København, Denmark; Naturalis Biodiversity Center, Leiden, The Netherlands; Natural

375 History Museum, London, England; Malmö Museum Malmöhus Castle, Malmö, Sweden;

376 Zoologische Staatssammlung München, Germany; American Museum of Natural

377 History, New York, USA; Muséum National d'Histoire Naturelle, Paris, France; Lee Kong 378 Chian Natural History Museum, Singapore;Draft Naturhistoriska Riksmuseet, Stockholm, 379 Sweden; Anatomy Museum, Faculteit Diergeneeskunde, Universiteit Utrecht, The

380 Netherlands;

381 National Museum of Natural History, Washington, USA; Naturhistorisches Museum

382 Wien, Austria. He would like to thank Aline Brodin, Edinburgh University Library, for

383 translations into French. He is also grateful to the University of Edinburgh and the

384 Balloch Trust for financial support.

385

386

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537

538

539

Draft

540 Table

541 Table 1. A summary of the occurrence and distribution of anomalous tooth wear

542 specimens found in 621 babirusa (genus Babyrousa) specimens in museum and private

543 collections, showing Geographical location, number of specimens studied, number of

544 ‘Wear’ anomaly specimens, the percentage of ‘Wear’ anomaly specimens, N = the

545 number of narrow indentation ‘Wear’ specimens, W = the number of wide indentation

546 ‘Wear’ specimens, E = the number of extensive indentation ‘Wear’ specimens, the

547 number of Groove anomaly specimens, and the percentage of specimens with ‘Groove’

548 wear. A large number (214) of Sulawesi skulls could not be identified to specific

549 geographical locations on Sulawesi. Similarly, a large number of Moluccan babirusa 550 skulls (90) could not be identified specificallyDraft to either Buru or the Sula Islands. 551

Specimen Wear Groove Geographical location numbers Anomalies specimens % Wear N_Wear W_Wear E_Wear specimens % Groove Buru 64 1 1 2 0 0 1 0 Sula Islands 33 4 4 12 3 1 1 0 Sum of Buru and Sula Isles 97 5 5 5 3 1 2 0 Buru or Sula Islands 90 15 13 14 1 4 8 5 6 Total all Buru and/or Sula Isles 187 20 18 10 4 5 10 5 3

North-east Sulawesi 85 19 14 17 3 6 9 11 13 North-west Sulawesi 43 12 10 23 1 2 8 5 12 East-central Sulawesi 4 1 1 25 0 0 1 0 West-central Sulawesi 31 2 2 6 0 0 2 0 South-east Sulawesi 14 0 0 0 0 0 0 0 Sum of Sulawesi locations 177 34 27 13 4 8 20 16 10 Sulawesi 214 65 40 19 5 15 26 31 14 Total all Sulawesi 391 99 67 17 9 23 46 51 12

South Sulawesi (sub-fossils) 31 4 0 0 0 0 0 4 13

Togian Islands 12 0 0 0 0 0 0 0

Total crania and mandibles 590 119 85 14 13 28 56 56 9

Total number of specimens 621 123 85 14 13 28 56 60 9 552

553

554 Figures

555

556 Figure 1. Nomenclature, derived from Made (1996), of a number of the cusps of the

557 mandibular molar teeth of Buru babirusa (Babyrousa babyrussa Linnaeus,

558 1758) [Naturmuseum Senckenberg 7100]: (a) Left M2. (b) Left M3. 1 -

559 protoconid, 1c - protoendoconulid, 2 - metaconid, 3 - hypoconid, 3a -

560 hypoectoconulid, 3b - hypopreconulid, 4 - entoconid, 5 - pentaconid, 5a -

561 pentaectoconulid, 5b - pentapreconulid, 6 - hexaconid. (Teeth scaled to fit

562 column width.)

563 564 Figure 2. Nomenclature, derived fromDraft Made (1996), of the maxillary molar M3 tooth of 565 Buru babirusa (Babyrousa babyrussa Linnaeus, 1758) [Naturmuseum

566 Senckenberg 7100]: 1 - paracone, 2 - protocone, 3 - metacone, 4 - tetracone,

567 4b - tetrapreconule, 5 - pentacone, 5b - pentapreconule.

568

569 Figure 3. ‘N_Wear’ indentation of the protocone, the tetracone and the tetrapreconule

570 of the left and right maxillary M2 cheek teeth of North-east Sulawesi babirusa

571 (Babyrousa celebensis Deninger, 1909) [Göteborgs naturhistoriska museum

572 17.939]. (scale =10 mm)

573

574 Figure 4. ‘N_Wear’ indentation of the cheek teeth of Sulawesi babirusa (Babyrousa

575 celebensis Deninger, 1909): (a) right maxilla of North-west Sulawesi

576 [Göteborgs naturhistoriska museum 17.946], ‘Wear’ indentation of the

577 caudal surface of the tetracone of M2 and into the tetrapreconule between

578 the protocone and the tetracone of M3, and in the fossa between the

579 metacone and tetracone; (b) left maxilla of North-east Sulawesi [Göteborgs

580 naturhistoriska museum 17.975], ‘Wear’ indentation between the protocone

581 and the tetracone of M3; (c) left maxilla of Sulawesi [National Museums of

582 Scotland 1991.15.006] ‘Wear’ indentation of the tetrapreconule, and smaller

583 ‘Wear’ indentation between the protocone and the tetracone of M3. Note the

584 ‘Groove’ between the caudal surface of M2 and the protocone of M3. (scale = 5

585 mm).

586

587 Figure 5. ‘Wear’ indentation of the cheek teeth of Sulawesi babirusa (Babyrousa

588 celebensis Deninger, 1909): (a) right mandible of North-east Sulawesi 589 [Göteborgs naturhistoriskaDraft museum 17.964], illustrating ‘W_Wear’ of three 590 parts of M3 - the protoconid, protoendoconulid, metaconid, hypoectoconulid

591 and hypopreconulid – the hypoconid, entoconid and pentapreconule – and

592 the pentaconid and hexaconid; (b) right mandible of North-west Sulawesi

593 [National Museum of Natural History, Washington 199884b] illustrating

594 ‘E_Wear’ of the entoconid, pentaectoconulid, pentaconid and hexaconid of

595 M3; (c) left mandible North-west Sulawesi [National Museum of Natural

596 History, Washington 199885b]illustrating ‘E_Wear’ of the hypoconid,

597 entoconid, pentaconid, pentaectoconulid, pentapreconulid and hexaconid of

598 M3; note the ‘Groove’ indentation between the caudal surface of M2 and

599 rostral surface of M3. (scale = 5 mm).

600

601 Figure 6. ‘Groove’ indentation of the caudal surfaces of left and right M1 and into the

602 rostral surfaces of left and right M2, and into (both left and right) the

603 pentaconid of M2 and the metaconid of M3 of North-east Sulawesi babirusa

604 (Babyrousa celebensis Deninger, 1909) [Göteborgs naturhistoriska museum

605 17.939]. Note the ‘Wear’ indentation between the metaconid and entoconid

606 of the left M2. (scale = 5 mm)

607

608 Figure 7. ‘Groove’ indentation of the cheek teeth of Sulawesi babirusa (Babyrousa

609 celebensis Deninger, 1909): (a) left maxilla of North-east Sulawesi

610 [Senckenberg Naturhistorische Sammlungen Dresden 3070] between caudal

611 surface of M2 and rostral surface of M3; (b) right mandible of Sulawesi [Lee

612 Kong Chian Natural History Museum ZRC.4.1959]. between caudal surface of

613 M2 and rostral surface of M3; (c) right maxilla North-west Sulawesi 614 [Göteborgs naturhistoriskaDraft museum 17.947], between caudal surface of M2 615 and rostral surface of M3 and also showing widespread dental erosion

616 around this locus into adjacent cusps of both molars. (scale = 5 mm).

617

618 Figure 8. Subfossil left M3 of babirusa (Babyrousa celebensis Deninger, 1909)

619 [Göteborgs naturhistoriska museum RMNH.MAM.Hooijer40] from South

620 Sulawesi with ‘Groove’ indentation of the protocone (triangle) and ‘Wear’

621 indentation of the tooth between the protocone and the tetracone. (scale = 5

622 mm).

623

624 Figure 9. (a) Fruit of a banana (Musa balbisiana Colla) from Buru peeled to illustrate

625 the distribution of seeds in situ (scale = 10 mm); (b) photographic image of a

626 banana (Musa balbisiana) seed from Buru; (c) scanning electron micrograph

627 of banana (Musa balbisiana) seed from Buru. Both ‘b’ and ‘c’ show the

628 micropylar collar around the operculum and micropylar plug, the wrinkled

629 outer epidermis and the projecting crystalline silica bodies; (d) shows the

630 chalazal end of the seed as well as the outer epidermis and the projecting

631 crystalline silica bodies. (scale [b,c,d] = 1 mm).

Draft

Figure 1. Nomenclature, derived from Made (1996), of a number of the cusps of the mandibular molar teeth of Buru babirusa (Babyrousa babyrussa Linnaeus, 1758) [Naturmuseum Senckenberg 7100]: (a) Left M2. (b) Left M3. 1 - protoconid, 1c - protoendoconulid, 2 - metaconid, 3 - hypoconid, 3a - hypoectoconulid, 3b - hypopreconulid, 4 - entoconid, 5 - pentaconid, 5a - pentaectoconulid, 5b - pentapreconulid, 6 - hexaconid. (Teeth scaled to fit column width.)

161x219mm (300 x 300 DPI)

Draft

Figure 2. Nomenclature, derived from Made (1996), of the maxillary molar M3 tooth of Buru babirusa (Babyrousa babyrussa Linnaeus, 1758) [Naturmuseum Senckenberg 7100]: 1 - paracone, 2 - protocone, 3 - metacone, 4 - tetracone, 4b - tetrapreconule, 5 - pentacone, 5b - pentapreconule.

169x127mm (300 x 300 DPI)

Draft

Figure 3. ‘N_Wear’ indentation of the protocone, the tetracone and the tetrapreconule of the left and right maxillary M2 cheek teeth of North-east Sulawesi babirusa (Babyrousa celebensis Deninger, 1909) [Göteborgs naturhistoriska museum 17.939]. (scale =10 mm)

169x128mm (300 x 300 DPI)

Draft

Figure 4. ‘N_Wear’ indentation of the cheek teeth of Sulawesi babirusa (Babyrousa celebensis Deninger, 1909): (a) right maxilla of North-west Sulawesi [Göteborgs naturhistoriska museum 17.946], ‘Wear’ indentation of the caudal surface of the tetracone of M2 and into the tetrapreconule between the protocone and the tetracone of M3, and in the fossa between the metacone and tetracone; (b) left maxilla of North- east Sulawesi [Göteborgs naturhistoriska museum 17.975], ‘Wear’ indentation between the protocone and the tetracone of M3; (c) left maxilla of Sulawesi [National Museums of Scotland 1991.15.006] ‘Wear’ indentation of the tetrapreconule, and smaller ‘Wear’ indentation between the protocone and the tetracone of M3. Note the ‘Groove’ between the caudal surface of M2 and the protocone of M3. (scale = 5 mm).

169x246mm (300 x 300 DPI)

Draft

Figure 5. ‘Wear’ indentation of the cheek teeth of Sulawesi babirusa (Babyrousa celebensis Deninger, 1909): (a) right mandible of North-east Sulawesi [Göteborgs naturhistoriska museum 17.964], illustrating ‘W_Wear’ of three parts of M3 - the protoconid, protoendoconulid, metaconid, hypoectoconulid and hypopreconulid – the hypoconid, entoconid and pentapreconule – and the pentaconid and hexaconid; (b) right mandible of North-west Sulawesi [National Museum of Natural History, Washington 199884b] illustrating ‘E_Wear’ of the entoconid, pentaectoconulid, pentaconid and hexaconid of M3; (c) left mandible North-west Sulawesi [National Museum of Natural History, Washington 199885b] illustrating ‘E_Wear’ of the hypoconid, entoconid, pentaconid, pentaectoconulid, pentapreconulid and hexaconid of M3; note the ‘Groove’ indentation between the caudal surface of M2 and rostral surface of M3. (scale = 5 mm).

169x208mm (300 x 300 DPI)

Draft

Figure 6. ‘Groove’ indentation of the caudal surfaces of left and right M1 and into the rostral surfaces of left and right M2, and into (both left and right) the pentaconid of M2 and the metaconid of M3 of North-east Sulawesi babirusa (Babyrousa celebensis Deninger, 1909) [Göteborgs naturhistoriska museum 17.939]. Note the ‘Wear’ indentation between the metaconid and entoconid of the left M2. (scale = 5 mm).

169x159mm (300 x 300 DPI)

Draft

Figure 7. ‘Groove’ indentation of the cheek teeth of Sulawesi babirusa (Babyrousa celebensis Deninger, 1909): (a) left maxilla of North-east Sulawesi [Senckenberg Naturhistorische Sammlungen Dresden 3070] between caudal surface of M2 and rostral surface of M3; (b) right mandible of Sulawesi [Lee Kong Chian Natural History Museum ZRC.4.1959]. between caudal surface of M2 and rostral surface of M3; (c) right maxilla North-west Sulawesi [Göteborgs naturhistoriska museum 17.947], between caudal surface of M2 and rostral surface of M3 and also showing widespread dental erosion around this locus into adjacent cusps of both molars. (scale = 5 mm).

169x221mm (300 x 300 DPI)

Draft

Figure 8. Subfossil left M3 of babirusa (Babyrousa celebensis Deninger, 1909) [Göteborgs naturhistoriska museum RMNH.MAM.Hooijer40] from South Sulawesi with ‘Groove’ indentation of the protocone (triangle) and ‘Wear’ indentation of the tooth between the protocone and the tetracone. (scale = 5 mm).

169x123mm (300 x 300 DPI)

Draft

Figure 9. (a) Fruit of a banana (Musa balbisiana Colla) from Buru peeled to illustrate the distribution of seeds in situ (scale = 10 mm); (b) photographic image of a banana (Musa balbisiana) seed from Buru; (c) scanning electron micrograph of banana (Musa balbisiana) seed from Buru. Both ‘b’ and ‘c’ show the micropylar collar around the operculum and micropylar plug, the wrinkled outer epidermis and the projecting crystalline silica bodies; (d) shows the chalazal end of the seed as well as the outer epidermis and the projecting crystalline silica bodies. (scale [b,c,d] = 1 mm).

221x169mm (300 x 300 DPI)