# The effect of captivity on craniomandibular and calcaneal ontogenetic trajectories in wild boar

Dimitri Neaux, Hugo Harbers, Barbara Blanc, Katia Ortiz, Yann Locatelli, Anthony Herrel, Vincent Debat, Thomas Cucchi

► To cite this version:

Dimitri Neaux, Hugo Harbers, Barbara Blanc, Katia Ortiz, Yann Locatelli, et al. The effect of captivity on craniomandibular and calcaneal ontogenetic trajectories in wild boar. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, 2022, 338 (8), pp.575-585. 10.1002/jez.b.23130. hal-03608590

HAL Id: hal-03608590

https://hal.science/hal-03608590

Submitted on 24 Nov 2022

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>
1

The effect of captivity on craniomandibular and calcaneal ontogenetic trajectories in wild boar

2

Short running title: Captivity and ontogenetic trajectories

3

4

Dimitri Neaux¹,²*, Hugo Harbers¹, Barbara Blanc³, Katia Ortiz³,⁴, Yann Locatelli³,⁵, Anthony Herrel⁶,

5

Vincent Debat⁴, Thomas Cucchi¹*

6

7

¹ Archéozoologie, Archéobotanique: Sociétés, Pratiques et Environnements, UMR 7209, Muséum national d'Histoire naturelle CNRS, Paris, France

8

9

² Laboratoire Paléontologie Evolution Paléoecosystèmes Paléoprimatologie, UMR 7262, Université de Poitiers CNRS, Poitiers, France

10

11

³ Réserve Zoologique de la Haute-Touche, Muséum national d'Histoire naturelle, Obterre, France

12

⁴ Institut de Systématique, Evolution, Biodiversité, UMR 7205, Muséum national d'Histoire naturelle CNRS UPMC EPHE, UA, Paris, France

13

14

⁵ Physiologie de la Reproduction et des Comportements, UMR 7247, INRAE CNRS Université de Tours IFCE, Nouzilly, France

15

16

⁶ Mécanismes Adaptatifs et Evolution, UMR 7179, Muséum national d'Histoire naturelle CNRS, Paris, France

17

18

19

*Corresponding authors: Dimitri Neaux: dimitrineaux@gmail.com, Thomas Cucchi:

20

thomas.cucchi@mnhn.fr

21
22 ORCID IDs:

23 Dimitri Neaux: 0000-0001-9465-220X

24 Thomas Cucchi: ORCID ID: 0000-0001-6021-5001

25 Yann Locatelli: ORCID ID: 0000-0002-2657-9327

26 Anthony Herrel: ORCID ID: 0000-0003-0991-4434

27 Vincent Debat: ORCID ID: 0000-0003-0040-1181
28

**Abstract**

29

30

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

Deciphering the plastic (i.e. non-heritable) changes induced by human control over wild animals in the archaeological record is challenging. Previous studies detected morphological markers associated with captivity in the cranium, mandible, and calcaneus of adult wild boar (*Sus scrofa*) but the developmental trajectories leading up to these changes during ontogeny remain unknown. To assess the impact of growth in a captive environment on morphological structures during postnatal ontogeny, we used an experimental approach focusing on the same three structures and taxon. We investigated the form and size differences of captive-reared and wild-caught wild boar during growth using three-dimensional (3D) landmark-based geometric morphometrics. Our results provide evidence of an influence of captivity on the morphology of craniomandibular structures, as wild specimens are smaller than captive individuals at similar ages. The food resources inherent to anthropogenic environments may explain some of the observed differences between captive-reared and wild specimens. The calcaneus presents a different contrasted pattern of plasticity as captive and wild individuals differ in terms of form but not in terms of size. The physically more constrained nature of the calcaneus and the direct influence of mobility reduction on this bone may explain these discrepancies. These results provide new methodological perspectives for bioarchaeological approaches as they imply that the plastic mark of captivity can be observed in juvenile specimens in the same way it has been previously described in adults.

46

47

Keywords: ontogeny, growth, domestication, geometric morphometrics, phenotypic plasticity

48
49 Research Highlights

50 We showed the influence of captivity, an early step of domestication, on the morphology of
51 craniomandibular and postcranial structures during development. It underlines the plastic nature of
52 bony structures and their ability to change in a short time period.

53
<figure><img src="image_2.png" /></figure>
56 **Introduction**

Animal domestication is an ongoing process (Vigne, 2011; Zeder, 2012) associated with substantial phenotypic changes that form part of the so-called domestication syndrome (Lord, Larson, Coppinger, & Karlsson, 2020; Sánchez-Villagra, Geiger, & Schneider, 2016; Zeder, 2012). Exploring the developmental mechanisms associated with the emergence of domestic phenotypes is crucial to document the roots of animal domestication over the last 15,000 years (Zeder, 2018).

Zooarchaeologists previously considered that morphological changes observed in the archaeological record, such as bone shape and size modifications, were subsequent to the integration of animals into human society (Clutton-Brock, 1992) through adaptations to the new constraints of the anthropogenic environment (Price, 1999). Therefore, morphological markers have been deemed irrelevant to document the early processes of domestication (e.g. population control through captivity; Vigne, Carrère, Briois, & Guilaine, 2011), as they would only be detectable when genetic isolation and breeding selection are already in place (Frantz et al., 2015; Marshall, Dobney, Denham, & Capriles, 2014). Yet, a series of recent experimental studies with wild boar (*Sus scrofa*) have demonstrated that early domestication steps, such as the control of wild animals, can be detected and quantified. These studies further showed that a lifetime in captivity induces changes in the functional demands (e.g. locomotor, foraging or feeding behaviours), modifying the shape of craniomandibular (Neaux, Blanc, Ortiz, Locatelli, Laurens, et al., 2021; Neaux, Blanc, Ortiz, Locatelli, Schafberg, et al., 2021) and postcranial (Harbers, Neaux, et al., 2020; Harbers, Zanolli, et al., 2020) bony structures. More importantly, these studies showed that captivity leaves an anatomical imprint on the musculoskeletal system beyond the phenotypic variation range observed in animals in their natural habitat. These results have been confirmed by studies on reindeer (*Rangifer tarandus*) comparing wild and captive populations (Pelletier, Kotiaho, Niinimäki, & Salmi, 2020, 2021).
While previous studies detected morphological markers associated with captivity in the cranium, mandible (Neaux, Blanc, Ortiz, Locatelli, Laurens, et al., 2021; Neaux, Blanc, Ortiz, Locatelli, Schafberg, et al., 2021), and calcaneus (Harbers, Neaux, et al., 2020), and humerus (Harbers, Zanolli, et al., 2020) of adult specimens, the tempo of these changes during postnatal ontogeny remains unknown. Substantial differences in terms of shape and size between wild and captive animals have already been identified in mammals during growth but with contradictory results (O'Regan & Kitchener, 2005). While some studies found greater cranial dimensions in captive lion cubs (*Panthera leo*; Smuts, Anderson, & Austin, 1978) and captive-bred chinchillas (*Chinchilla lanigera*; Crossley & del Mar Miguélez, 2001), others showed a decrease in cranial dimensions in captive Indian rhinoceroses (*Rhinoceros unicornis*; Groves, 1982) and equids (*Equus* spp.; Groves, 1966). The morphological ontogenetic changes associated with captivity therefore remain to be understood.

To assess the impact of growth in a captive environment on morphological structures during postnatal ontogeny, we used an experimental approach focusing on the same bones and taxon on which morphological markers associated with captivity were detected, i.e. the cranium, mandible (Neaux, Blanc, Ortiz, Locatelli, Laurens, et al., 2021; Neaux, Blanc, Ortiz, Locatelli, Schafberg, et al., 2021), and calcaneus (Harbers, Neaux, et al., 2020) of wild boar. We collected weaned wild boar piglets from a genetically homogenous population and raised them in a captive anthropogenic environment from the age of 6 months. We scanned them *in vivo* at five different age classes and compared them with wild-caught wild boar populations.

To determine the influence of captivity on the growth and development of the wild boar skeleton, we compared ontogenetic changes in form (i.e. size and shape) and size between captive-reared and wild-caught specimens. We predicted that form and size differences should not differ significantly at an early age but should rather start diverging in later age classes, when the effect of captivity on morphological structures becomes more prominent. Next, we investigated differences in ontogenetic
allometry, i.e. the relationship between shape and size over the course of ontogeny at different ages (Alberch, Gould, Oster, & Wake, 1979; Klingenberg, 2016), as allometry has been shown to affect postnatal ontogenetic trajectories in domesticated clades when compared to their wild counterparts (Sánchez-Villagra et al., 2017; Wilson, 2018). We hypothesized that differences in ontogenetic allometry should be significant between captive-reared and wild-caught wild boar, indicative of a plastic effect of captivity upon developmental trajectories.

## 109 Methods

## 110 Experimental design

Captive wild boar groups (Appendix S1) consist of wild boar from the DOMEXP project: a multidisciplinary experiment aiming to assess the effect of captivity on the musculoskeletal system (http://anr-domexp.cnrs.fr/). They include the same specimens that were studied as adults by Neaux et al. (2021; 2021) and Harbers et al. (2020; 2020). We relied on a control population of wild boar living in a 100,000 m² fenced forest in Urciers (France). These specimens came from a wild boar farm, where human interactions are intentionally kept to a minimum in order to ensure that the behaviour of the wild boar remains as natural as possible. They are free to forage for food in the woods. From this population, we sampled 24 piglets that were divided into two groups of 12 specimens of equal sex ratio (6 males and 6 females). These groups were raised from the age of 6 months at the zoological reserve of La Haute-Touche (France) in two different contexts of mobility reduction: a 3,000 m² wooded pen ('enclosure' group) and an indoor stall of 100 m² ('stall' group). These space restrictions respectively represent a reduction of 97% and 99.9% of the range of the control population and do not allow the captive specimens to roam as freely as animals from natural populations of wild boar (Palencia et al., 2019; Russo, Massei, & Genov, 1997). We supplied individuals from both groups with processed dry food pellets including 15.5% of raw proteins adapted for domestic pig diet. These specimens were repeatedly scanned *in vivo* at the age of 6, 8, 11, 14, and 20 months, using a Computed Tomography
(CT) scanner with a spatial resolution of between 100 and 500 µm at the Chirurgie et Imagerie pour la
Recherche et l'Enseignement (CIRE) platform of the Institut National de Recherche pour l'Agriculture,
l'Alimentation et l'Environnement (INRAE). This experiment received ethics approval from the French
Ministère de l'Enseignement Supérieur, de la Recherche et de l'Innovation (APAFIS#5353-
20160511133847).

132 Comparative wild-caught wild boar samples

For comparison with the two captive groups, we collected free-ranging specimens ('wild-caught'
group). This group included 6 individuals from the control free-ranging population of Urciers (100,000
m²) mentioned previously. Also included in the 'wild-caught' groups are 15 free-ranging wild boar that
were sampled in the forests of Chambord and Compiègne (approximated at 54,400,000 m² and
150,000,000 m² respectively; Harbers, Neaux, et al., 2020), belonging to similar geographic and
climatic environment (i.e. temperate central France), to reduce the confounding effects of geographic
and climate-induced morphological variation known to exist in Sus scrofa (Albarella, Dobney, &
Rowley-Conwy, 2009; Groves, 2021; Iannucci, Sardella, Strani, & Mecozzi, 2020). Like most wild
boar in western Europe, these free-ranging specimens likely had an omnivorous diet consisting mostly
of plants (e.g. acorns, roots, and crops) supplemented with animal matter (e.g. insect and earthworms)
as a primary source of protein (Schley & Roper, 2003). These specimens were wild-caught either
between 8 and 10 months or between 16 and 18 months of age. We based age estimation on the
mandibular tooth eruption and wear stages in occlusal view comparing our specimens with the charts
developed by Grant (1982) and Horard-Herbin (1997). The selection of wild-caught specimens has
been performed to match at best the age variation of captive specimens. Due to the inherent difficulty
of collecting juvenile wild-caught specimens, the individuals from our study do not cover fully this
variation, leading possibly to slight over-interpretation of results.
151 **Data acquisition and analyses**

152 We used homologous landmarks and semilandmarks placed on three-dimensional (3D) surfaces to
153 describe the morphology. Digitisation and landmark definition were performed following published
154 protocols (Appendix S2; Harbers, Neaux, et al., 2020; Neaux, Blanc, Ortiz, Locatelli, Laurens, et al.,
155 2021; Neaux et al., 2020). We performed all the analyses in the R environment (R Core Team, 2019).
156 Coordinates were aligned using a generalised Procrustes superimposition (Rohlf & Slice, 1990),
157 implemented in the procSym function of the package ‘Morpho’ (Schlager & Jefferis, 2020).

158 We chose to work on form, i.e. the combination of size and shape (Dryden & Mardia, 1998), rather
159 than shape as form is a more comprehensive description of an object than shape alone in the context of
160 ontogenetic studies (Mitteroecker, Gunz, Windhager, & Schaefer, 2013). We constructed the form
161 space by augmenting the Procrustes coordinates by the logarithm (log) of centroid size (CS;
162 Mitteroecker, Gunz, Bernhard, Schaefer, & Bookstein, 2004). The exploratory approach of the major
163 directions of variation in this form space relies on a principal component analysis (PCA) on form
164 variables performed on the mean form of each age class of each of the three groups. We visualized the
165 deformations between negative and positive scores on the first two principal components as well as the
166 deformations between age classes for each of the three groups using heatmaps through the meshDist
167 function of ‘Morpho’. The lack of congruence in age class among wild-caught groups and the two
168 captive groups prevented us from directly comparing the ontogenetic trajectories (Adams & Collyer,
169 2009). We therefore tested the difference in form between captive-reared and wild-caught groups using
170 a factorial MANOVA through the procD.lm function of ‘the package ‘geomorph’ (Adams, Collyer, &
171 Kaliontzopoulou, 2019). We also tested the difference in CS and in body mass (Appendix S3) between
172 captive-reared and wild-caught groups with a pairwise test and visualized it graphically with a bivariate
173 plot. Body masses were measured on a scale before each scan *in vivo* for the captive-reared specimens
174 and on uneviscerated specimens for the wild-caught ones.
To estimate and compare ontogenetic allometries between the three groups, we used a multivariate regression (Drake & Klingenberg, 2008) between shape (Procrustes coordinates) and size, computed as log CS (Collyer, Sekora, & Adams, 2015). We displayed graphically the difference in ontogenetic trajectories among our three mobility groups using a biplot of regression shape scores against log CS and tested it with a MANCOVA through the procD.lm function.

180

## 181 Results

### 182 Form

The factorial MANOVA (Table 1) showed that the ‘8-10 month wild-caught’ cranium, mandible, and calcaneus forms are not significantly different from the 6-month and 8-month old captive forms (i.e. stall and enclosure). Those structures are different from those of all the older captive groups. The ‘16-18 month wild-caught’ group is not significantly different from the 11-month and 14-month captive groups for any of the studied structures. The mandible and calcaneus of this age class do not differ from those of the 8-month ‘enclosure’ nor from the 8-month ‘stall’ groups for the mandible. They are different from those of all the other captive groups (see Appendix S4 for the Factorial MANOVA between all groups).

For the PCA (Fig. 1), PC1 for the cranium, mandible, and calcaneus show respectively 79.8%, 86.1%, and 75.5% of the total variance. For the three studied structures, PC1 displays mostly changes associated with growth as the younger specimens have the lower PC scores and the older ones have the higher scores. The cranium form changes associated with positive scores involve (1) an anteroposteriorly longer and more concave rostrum, (2) more robust zygomatic arches, and (3) smaller orbits relative to the overall cranium size (Fig 1.a). For the mandible, the associated form changes towards positive scores include (1) a reduction of the gonial angle, (2) a longer corpus and mandibular
symphysis, and (3) a mediolateral reduction of the space between the two mandibular rami in dorsal view (Fig 1.b). For the calcaneus, the changes towards positive scores consisted in (1) a shift of the sustentaculum tali and of the calcaneal sulcus towards the distal extremity, (2) a more dorsoplantarily curved calcaneus, and (3) a more elongated epiphysis that is orientated toward the plantar side (Fig 1.c). PC2 bear less variation in form for the cranium, mandible, and calcaneus (respectively 7.4%, 5.1%, and 9.1% of the total variance) but clearly distinguishes captive and wild-caught specimens. Cranial form changes along PC2 from wild-caught (low values) to captive individuals (high values) involved (1) a narrower cranium in dorsal view, specifically in the zygomatic region, (2) a less vertical occipital region, and (3) an anteroposteriorly shorter nasal and maxillary region (Fig 1.a). The mandible form changes along PC2 are characterized by (1) a mediolateral narrowing, (2) a higher ramus, and (3) a ventrally orientated ramus (Fig 1.b). The calcaneus form changes along PC2 mainly express (1) a downward shift of the sustentaculum tali and (2) an epiphysis more orientated toward the plantar side (Fig. 1.c). From 6 months to 20 months, the main changes for captive specimens are localised on the zygomatic arches and on the occipital region for the cranium (Fig. 1.d), on the ramus and on the symphysis for the mandible (Fig. 1.e), and on the epiphysis, the sustentaculum tali, and the cuboid facet for the calcaneus (Fig. 1.f). From 8-10 months to 16-18 months, the changes for wild-caught specimens are localised on the same areas but are less prominent.

## Size

The size of the '8-10 month wild-caught' group is not significantly different from the 6-month and 8-month captive groups for the cranium, mandible, and calcaneus (Fig. 2; Table 2). It is different from all the older captive groups. The '16-18 month wild-caught' group is not significantly different from the 11- and 14-month captive groups for the cranium, the 8-, 11- and 14-month captive groups for the mandible, and all the captive groups from 8 to 20 months for the calcaneus. The body mass of the '8-10 month wild-caught' is not significantly different from the 6-, 8- and 11-month captive groups and
the '16-18 month wild-caught' group is not significantly different from the 11- and 14-month captive
groups and the '20 months - enclosure' group. All the other groups are statistically different.

224 **Ontogenetic allometry**

We found overall ontogenetic allometry across all three structures in the three groups of mobility,
which explains greater variation in calcaneus (26%) than in cranium (21%), and mandible (16%; Fig.
3). The ontogenetic allometric trajectories depicted graphically are parallel between enclosure and stall
captive groups across the three structures. They differ from the wild-caught group with a more positive
allometry, although the interaction term was only significant for the calcaneus, indicative of a
difference in allometric growth among the three groups for this structure.

231 **Discussion**

For the cranium, mandible, and calcaneus, we found that the form and size of the ‘8-10 month wild-
caught’ wild boar are similar to those of younger captive specimens aged between 6 and 8 months, but
are significantly different from the 11-month-old animals and older ones. This may suggest that the
growth in captivity of a wild ungulate impacts the development of the three studied structures (O’Regan
& Kitchener, 2005). Differences also exist later for the cranium and mandible, when the effect of the
reduction of mobility on morphological structures becomes more prominent. Indeed, the form and size
of the ‘16-18 month wild-caught’ group are similar to the 11- and 14-month-old captive specimens but
statistically different from the 20-month-old ones. The same results are observed when comparing body
mass between wild-caught and captive specimens. These results are strengthened by the fact that the
wild-caught control specimens, belonging to the same initial herd as the captive ones, follow the same
trend as the other wild-caught wild boar (see Fig. 1.a and 1.b). Unfortunately, 6-month-old wild-caught
wild boar were not available. Yet, since the ‘8-10 month wild-caught’ group is similar to the 6-month-
old captive specimens, we could speculate that 6-month-old wild-caught wild boar would be similar to
younger specimens. This suggests that captive growth in wild boar induces a morphological divergence driven by an acceleration of changes in the size and form of the skull. This developmental delay in wild-caught wild boar compared to captive ones is characterised by a retention of more juvenile traits observed in younger wild specimens as evidenced by the MANOVA. This implies an increase in the developmental rate of captive wild boar when compared to that of wild-caught specimens. These findings are in line with previous studies on postnatal growth assessing that, for a given age, captive specimens are often significantly larger than wild-caught individuals. Comparing skeletal measurements, Zihlman et al. (2007) found that the tempo of growth in wild chimpanzees (Pan troglodytes) contrasts sharply with the rate demonstrated for captive individuals that can mature as much as 3 years earlier. Cheverud et al. (1992) also found a significant increase of the growth rate in toque macaques (Macaca sinica) raised in the laboratory when compared to wild specimens. Finally, captive-raised lion cubs were reported as being nearly twice the size of wild cubs of the same age (Schaller, 1973). The more consistently available food is probably the main cause of these differences (Turner, Cramer, Nisbett, & Gray, 2016). Indeed, one of the consequences of the spatial control of wild animals by humans is the presence of more constantly available food resources. As it is the case in the context of our experimental study, animals in captivity are regularly provisioned and do not need to spend energy searching for food, hence removing intra-group feeding competition. In addition, intrasexual competition is removed in stall specimens and drastically reduced for the pen specimens. Differences in cranial size during the postnatal development were previously described between domestic pigs and wild boar, especially in the neurocranial region, surrounding and protecting cerebral structures (Evin et al., 2017). Our study stresses that this increase of the size of neurocranial structures is not necessarily a product of the long selective breeding leading to the morphology observed in domestic pigs. Indeed, we observe a similar increase as a plastic response to captivity, considered as one of the earliest domestication steps (Vigne, 2011), most likely due to an unrestricted access to food
269 leading to a faster growth (Kimura & Hamada, 1996). Furthermore, these findings are consistent with 270 recent studies suggesting that the commonly assumed reduction of brain size associated with 271 domestication should be questioned (Lord, Larson, Coppinger, et al., 2020; Lord, Larson, & Karlsson, 272 2020). Indeed, both early domestication steps, in the context of our study, and long term selective 273 processes (Evin et al., 2017) result in a size increase and not a reduction of the structures surrounding 274 the brain. Yet, although described previously (Finarelli, 2006; 2011), the presence of a clear positive 275 relationship between brain size and cranial structures size is still unclear (Logan, & Palmstrom, 2015), 276 highlighting the need for future studies directly addressing the evolution of endocranial shape and size 277 in relation to domestication.

278 The calcaneus presents a different pattern when the specimens are older, i.e. when the effect of captivity 279 becomes prominent as the ‘16-18 month wild-caught’ group is similar to the 11-, 14-, and 20-month- 280 old captive specimens in size. This result is supported by the fact that the wild-caught control specimens 281 follow the same trend as the captive ones (see Fig. 1.c). The more physically constrained nature of the 282 calcaneus, articulating with both the talus and the cuboid bones, may partly explain this difference. In 283 this sense, Hanot et al (2017) described a strong and significant morphological integration between the 284 calcaneus and talus in horses (Equus caballus). In comparison, lower level of integration were found 285 between the cranium and mandible of wild boar, specifically for captive individuals (Neaux, Blanc, 286 Ortiz, Locatelli, Schafberg, et al., 2021). This comparatively loose integration in the skull may allow 287 more size variation, and a greater and more rapid growth of these structures in captive individuals. 288 Further covariation studies of the tarsus of less specialized taxa than horses, such as wild boar, will 289 help untangle the role of morphological integration in the observed differences between the 290 craniomandibular and postcranial structures.

291 The calcaneus is subjected to high tensile, bending and compressive forces (Su, Skedros, Bachus, & 292 Bloebaum, 1999) and has often being described has a key proxy to assess terrestrial mammal locomotor
behaviours (Bassarova, Janis, & Archer, 2009; Ginot, Hautier, Marivaux, & Vianey-Liaud, 2016; 293
Panciroli, Janis, Stockdale, & Martín-Serra, 2017). In this respect, the phenotypic plasticity in shape 294
(but not the size) of the calcaneus has been shown to capture the direct influence of the anthropogenic 295
control of wild boar locomotor behaviour (Harbers, Neaux, et al., 2020). Conversely, the 296
morphological modifications associated with the mandible in captive animals are not the direct 297
consequence of mobility reduction but may rather be related to functional demands resulting from the 298
anthropogenic environment (Neaux, Blanc, Ortiz, Locatelli, Laurens, et al., 2021). They include the 299
increase of feeding activity (Turner et al., 2016), the reduction of foraging behaviour (Mason & Mendl, 300
1997), and the appearance of stereotypy (Rhodes et al., 2005), i.e. repeated sequences of movements 301
with no obvious purpose, particularly common in captive animals (Fraser & Broom, 1990). In this 302
sense, in PC2 of the PCA on the form space for the calcaneus, from negative to positive values, there 303
is a clear gradient from wild-caught, to enclosure, to stall specimens (Fig. 1.c), while for the cranium 304
(Fig. 1.a) and mandible (Fig. 1.b) captive individuals (i.e. enclosure and stall) are similar. This result 305
is consistent with the hypothesis that the calcaneus captures mainly locomotor behaviour as the habitats 306
of the wild-caught group is the larger (≥ 100,000 m²), followed by the enclosure group (3,000 m²) and 307
finally the stall group (100 m²). The PC2 for the cranium and mandible form space primarily records 308
changes between the captive specimens (i.e. enclosure and stall) under anthropogenic control on the 309
one hand and the wild-caught group on the other. This result is in line with the hypothesis that 310
craniomandibular changes are not directly associated with a reduced mobility but rather with functional 311
requirements resulting from the anthropogenic environment (e.g. feeding activity, foraging behaviour, 312
stereotypy). The difference in the way captivity affects the calcaneus (direct influence) and the cranium 313
and mandible (indirect influence) may explain this distinction and should be further explored. 314

The slope of the ontogenetic allometry for the cranium and mandible is not significantly different 315
between captive-reared and wild-caught specimens. This result is in line with previous findings that 316
317 ontogenetic allometry generally does not evolve on short evolutionary time scales and that
318 modifications of ontogenetic trajectories are usually achieved by heterochronic shifts along a shared
319 slope rather than directional changes (Voje, Hansen, Egset, Bolstad, & Pélabon, 2014; Wilson, 2018).
320 Moreover, previous results (Sánchez-Villagra et al., 2017; Wilson, 2018) found no significant
321 differences in the ontogenetic slopes between domestic pigs and wild boar contrary to dogs (Canis
322 lupus familiaris) and wolves (C. lupus), and llamas (Lama glama) and guanacos (L. guanicoe). This is
323 congruent with our results on wild boar captivity, as the control of mobility is considered one of the
324 first steps of the domestication process leading to the morphology observed in modern pigs (Vigne,
325 2011). The differences in the direction of the slope for the calcaneus shows that some level of
326 directional change can nevertheless occur on a short-time scale. The distinctions between the skull and
327 the calcaneus may once again arise from how mobility reduction influences these structures, i.e.
328 respectively indirectly and directly.
329

329 Conclusions

330 Our results provide evidence that captivity influences the morphology of craniomandibular and
331 postcranial structures of wild boar, as wild specimens are significantly smaller than captive individuals
332 of a similar age. Consistently available food resources and the reduction of stress associated with the
333 search for food and intra-group feeding competition may explain the distinction between captive-reared
334 and wild specimens. The calcaneus presents a different pattern as captive and wild individuals differ
335 in terms of form but not in terms of size. Furthermore, it is the only structure presenting differences in
336 ontogenetic allometry. The more physically constrained nature of the calcaneus and the direct influence
337 of mobility reduction on this bone may explain these specificities. These results provide new
338 methodological perspectives for bioarchaeological approaches as they imply that the plastic mark of
339 captivity can be observed in juvenile specimens as well as in adults (Cucchi et al., 2021; Harbers,
340 Neaux, et al., 2020; Neaux, Blanc, Ortiz, Locatelli, Laurens, et al., 2021; Neaux, Blanc, Ortiz, Locatelli,
341 Schafberg, et al., 2021). Further studies need to explore the morphological integration during growth

342 in captive conditions for both craniomandibular and postcranial structures in order to decipher the role

343 of developmental and functional correlates between structures in generating the differences observed

344 in our study.
345 Acknowledgements

346 We are most grateful to the staff of the Réserve Zoologique de la Haute-Touche (Christophe Audureau,
347 Jérémy Bernard, Christophe Jubert, Sandrine Laloux, Emmanuel Marechal, Régis Rabier, Patrick
348 Roux, and Colin Vion) for their help during the setup of the experimental structures, the care they
349 provided to the experimental specimens and during the data acquisition. We thank the CIRE platform
350 at INRAE (Hans Adriensen, Frédéric Elbout, Christian Moussu and Luc Perrigouard). This research
351 was funded by ANR, through the Domexp project (ANR-13-JSH3-0003-01), the LabEx ANR-10-
352 LABX-0003-BCDiv, in the programme ‘Investissements d’avenir’ ANR-11-IDEX-0004-02,
353 programme Emergence SU-19-3-EMRG-02, Muséum national d’Histoire naturelle (Paris), and the
354 CNRS INEE institute.

355

356 Conflict of Interest Statement

357 The authors declare no conflicts of interest.

358

359 Data Availability Statement

360 All analytical codes and data are freely available at: https://zenodo.org/record/5547335 (Neaux,
361 Harbers, Blanc, Ortiz, Locatelli, et al., 2021)

362
363 Adams, D. C., Collyer, M., & Kaliontzopoulou, A. (2019). geomorph: Geometric morphometric
364 analyses of 2D/3D landmark data (Version 3.1.2).

365 Adams, D. C., & Collyer, M. L. (2009). A general framework for the analysis of phenotypic
366 trajectories in evolutionary studies. *Evolution; International Journal of Organic Evolution*,
367 63(5), 1143–1154. https://doi.org/10.1111/j.1558-5646.2009.00649.x

368 Albarella, U., Dobney, K., & Rowley-Conwy, P. (2009). Size and shape of the Eurasian wild boar
369 (*Sus scrofa*), with a view to the reconstruction of its Holocene history. *Environmental
370 Archaeology*, 14(2), 103–136. https://doi.org/10.1179/146141009X12481709928283

371 Alberch, P., Gould, S. J., Oster, G. F., & Wake, D. B. (1979). Size and Shape in Ontogeny and
372 Phylogeny. *Paleobiology*, 5(3), 296–317.

373 Bassarova, M., Janis, C. M., & Archer, M. (2009). The Calcaneum—On the Heels of Marsupial
374 Locomotion. *Journal of Mammalian Evolution*, 16(1), 1–23. https://doi.org/10.1007/s10914-008-9093-7

376 Cheverud, J. M., Wilson, P., & Dittus, W. P. J. (1992). Primate population studies at Polonnaruwa.
377 III. Somatometric growth in a natural population of toque macaques (*Macaca sinica*). *Journal
378 of Human Evolution*, 23(1), 51–77. https://doi.org/10.1016/0047-2484(92)90043-9

379 Clutton-Brock, J. (1992). The process of domestication. *Mammal Review*, 22(2), 79–85.
380 https://doi.org/10.1111/j.1365-2907.1992.tb00122.x

381 Collyer, M. L., Sekora, D. J., & Adams, D. C. (2015). A method for analysis of phenotypic change
382 for phenotypes described by high-dimensional data. *Heredity*, 115(4), 357–365.
383 https://doi.org/10.1038/hdy.2014.75

384 Crossley, D. A., & del Mar Miguélez, M. (2001). Skull size and cheek-tooth length in wild-caught
385 and captive-bred chinchillas. *Archives of Oral Biology*, 46(10), 919–928.
386 https://doi.org/10.1016/S0003-9969(01)00055-3
387 Cucchi, T., Domont, A., Harbers, H., Leduc, C., Guidez, A., Bridault, A., ... Vigne, J.-D. (2021).

388 Bones geometric morphometrics illustrate 10th millennium cal. BP domestication of

389 autochthonous Cypriot wild boar (*Sus scrofa circeus* nov. *ssp*). *Scientific Reports*, *11*(1),

390 11435. https://doi.org/10.1038/s41598-021-90933-w

391 Drake, A. G., & Klingenberg, C. P. (2008). The pace of morphological change: Historical

392 transformation of skull shape in St Bernard dogs. *Proceedings of the Royal Society B:*

393 *Biological Sciences*, *275*(1630), 71–76. https://doi.org/10.1098/rspb.2007.1169

394 Dryden, I. L., & Mardia, K. V. (1998). *Statistical Shape Analysis*. New York, NY: John Wiley and

395 Sons.

396 Evin, A., Owen, J., Larson, G., Debiais-Thibaud, M., Cucchi, T., Vidarsdottir, U. S., & Dobney, K.

397 (2017). A test for paedomorphism in domestic pig cranial morphology. *Biology Letters*, *13*(8),

398 20170321. https://doi.org/10.1098/rsbl.2017.0321

399 Finarelli, J. A., (2006). Estimation of endocranial volume through the use of external skull measures in

400 the carnivora (Mammalia). *Journal of Mammalogy*, *87*(5), 1027–1036,

401 https://doi.org/10.1644/05-MAMM-A-430R1.1

402 Finarelli, J. A., (2011). Estimating endocranial volume from the outside of the skull in Artiodactyla.

403 *Journal of Mammalogy*, *92*(1), 200–212, https://doi.org/10.1644/09-MAMM-A-391.1

404 Frantz, L. A. F., Schraiber, J. G., Madsen, O., Megens, H.-J., Cagan, A., Bosse, M., ... Groenen, M.

405 A. M. (2015). Evidence of long-term gene flow and selection during domestication from

406 analyses of Eurasian wild and domestic pig genomes. *Nature Genetics*, *47*(10), 1141–1148.

407 https://doi.org/10.1038/ng.3394

408 Fraser, A. F., & Broom, D. M. (1990). *Farm Animal Behaviour and Welfare*. Wallingford, UK:

409 CABI Publishing.
410 Ginot, S., Hautier, L., Marivaux, L., & Vianey-Liaud, M. (2016). Ecomorphological analysis of the
411 astragalo-calcaneal complex in rodents and inferences of locomotor behaviours in extinct
412 rodent species. *PeerJ*, 4, e2393. https://doi.org/10.7717/peerj.2393

413 Grant, A. (1982). The use of tooth wear as a guide to the domestic ungulates. In B. Wilson, C.
414 Grigson, & S. Payne (Eds.), *Ageing and Sexing Animal Bones from Archaeological Sites*
415 (BAR British Series, pp. 991–108). UK.

416 Groves, C P. (1966). Skull-changes due to captivity in certain Equidae. *Zeitschrift Fuer
417 Saugietierkunde*, pp. 44–66.

418 Groves, C. P. (1981). *Ancestors for the Pigs, Taxonomy and Phylogeny of the Genus Sus*. Canberra,
419 Australia: Department of Prehistory Technical Bulletin, Australian National University Press.

420 Groves, C P. (1982). The skulls of Asian rhinoceroses: Wild and captive. *Zoo Biology, 1*(3), 251-
421 261. https://doi.org/10.1002/zoo.1430010309

422 Hanot, P., Herrel, A., Guintard, C., & Cornette, R. (2017). Morphological integration in the
423 appendicular skeleton of two domestic taxa: The horse and donkey. *Proceedings of the Royal
424 Society B: Biological Sciences*, 284(1864), 20171241. https://doi.org/10.1098/rspb.2017.1241

425 Harbers, H., Neaux, D., Ortiz, K., Blanc, B., Laurens, F., Baly, I., ... Cucchi, T. (2020). The mark of
426 captivity: Plastic responses in the ankle bone of a wild ungulate (*Sus scrofa*). *Royal Society
427 Open Science*, 7(3), 192039. https://doi.org/10.1098/rsos.192039

428 Harbers, H., Zanolli, C., Cazenave, M., Theil, J.-C., Ortiz, K., Blanc, B., ... Cucchi, T. (2020).
429 Investigating the impact of captivity and domestication on limb bone cortical morphology: An
430 experimental approach using a wild boar model. *Scientific Reports*, 10(1), 19070.
431 https://doi.org/10.1038/s41598-020-75496-6
432 Horard-Herbin, M.-P. (1997). *Le village celtique des Arènes à Levroux. L'élevage et les productions animales dans l'économie de la fin du second âge du Fer-Levroux 4* (Vol. 12). Fédération pour l'édition de la Revue archéologique du Centre de la France.

435 Iannuci, A., Sardella, R., Strani, F., Mecozzi, B. (2020). Size shifts in late Middle Pleistocene to Early Holocene *Sus scrofa* (Suidae, Mammalia) from Apulia (southern Italy):
ecomorphological adaptations? *Hystrix, the Italian Journal of Mammalogy*, 31(1), 10–20.
https://doi.org/10.4404/hystrix-00258-2019

439 Kimura, T., & Hamada, Y. (1996). Growth of wild and laboratory born chimpanzees. *Primates*, 37(3), 237–251. https://doi.org/10.1007/BF02381856

441 Klingenberg, C. P. (2016). Size, shape, and form: Concepts of allometry in geometric morphometrics. *Development Genes and Evolution*, 226, 113–137.
https://doi.org/10.1007/s00427-016-0539-2

444 Logan, C. J., & Palmstrom, C. R. (2015). Can endocranial volume be estimated accurately from external skull measurements in great-tailed grackles (*Quiscalus mexicanus*)? *PeerJ*, 3:e1000. https://doi.org/10.7717/peerj.1000

447 Lord, K. A., Larson, G., Coppinger, R. P., & Karlsson, E. K. (2020). The History of Farm Foxes Undermines the Animal Domestication Syndrome. *Trends in Ecology & Evolution*, 35(2), 125–136. https://doi.org/10.1016/j.tree.2019.10.011

450 Lord, K. A., Larson, G., & Karlsson, E. K. (2020). Brain Size Does Not Rescue Domestication Syndrome. *Trends in Ecology & Evolution*, 35(12), 1061–1062.
https://doi.org/10.1016/j.tree.2020.10.004

453 Marshall, F. B., Dobney, K., Denham, T., & Capriles, J. M. (2014). Evaluating the roles of directed breeding and gene flow in animal domestication. *Proceedings of the National Academy of Sciences*, 111(17), 6153–6158. https://doi.org/10.1073/pnas.1312984110
Mason, G., & Mendl, M. (1997). Do the stereotypies of pigs, chickens and mink reflect adaptive species differences in the control of foraging? *Applied Animal Behaviour Science*, 53(1), 45–58. https://doi.org/10.1016/S0168-1591(96)01150-1

Mitteroecker, P., Gunz, P., Bernhard, M., Schaefer, K., & Bookstein, F. L. (2004). Comparison of cranial ontogenetic trajectories among great apes and humans. *Journal of Human Evolution*, 46(6), 679–698. https://doi.org/10.1016/j.jhevol.2004.03.006

Mitteroecker, P., Gunz, P., Windhager, S., & Schaefer, K. (2013). A brief review of shape, form, and allometry in geometric morphometrics, with applications to human facial morphology. *Hystrix*, the Italian Journal of Mammalogy, 24(1), 59–66. https://doi.org/10.4404/hystrix-24.1-6369

Neaux, D., Blanc, B., Ortiz, K., Locatelli, Y., Laurens, F., Baly, I., ... Cucchi, T. (2021). How changes in functional demands associated with captivity affect the skull shape of a wild boar (*Sus scrofa*). *Evolutionary Biology*, 48, 27–40. https://doi.org/10.1007/s11692-020-09521-x

Neaux, D., Blanc, B., Ortiz, K., Locatelli, Y., Schafberg, R., Herrel, A., ... Cucchi, T. (2021). Constraints associated with captivity alter craniomandibular integration in wild boar. *Journal of Anatomy*, 239(2), 489–497. https://doi.org/10.1111/joa.13425

Neaux, D., Harbers, H., Blanc, B., Ortiz, K., Locatelli, Y., Herrel, A., ... Cucchi, T. (2021). *The effect of captivity on cranial and postcranial ontogenetic trajectories in wild boar* [Data set]. Zenodo. https://doi.org/10.5281/zenodo.5547335

Neaux, D., Sansalone, G., Lecompte, F., Haruda, A., Schafberg, R., & Cucchi, T. (2020). Examining the effect of feralization on craniomandibular morphology in pigs, *Sus scrofa* (Artiodactyla: Suidae). *Biological Journal of the Linnean Society*, 131(4), 870–879. https://doi.org/10.1093/biolinnean/blaa156
479 O'Regan, H. J., & Kitchener, A. C. (2005). The effects of captivity on the morphology of captive,
480 domesticated and feral mammals. *Mammal Review*, 35(3–4), 215–230.

481 https://doi.org/10.1111/j.1365-2907.2005.00070.x

482 Palencia, P., Vicente, J., Barroso, P., Barasona, J. Á., Soriguer, R. C., & Acevedo, P. (2019).

483 Estimating day range from camera-trap data: The animals' behaviour as a key parameter.
484 *Journal of Zoology*, 309(3), 182–190. https://doi.org/10.1111/jzo.12710

485 Panciroli, E., Janis, C., Stockdale, M., & Martín-Serra, A. (2017). Correlates between calcaneal
486 morphology and locomotion in extant and extinct carnivorous mammals. *Journal of
487 Morphology*, 278(10), 1333–1353. https://doi.org/10.1002/jmor.20716

488 Pelletier, M., Kotiaho, A., Niinimäki, S., & Salmi, A.-K. (2020). Identifying early stages of reindeer
489 domestication in the archaeological record: A 3D morphological investigation on forelimb
490 bones of modern populations from Fennoscandia. *Archaeological and Anthropological
491 Sciences*, 12(8), 169. https://doi.org/10.1007/s12520-020-01123-0

492 Pelletier, M., Kotiaho, A., Niinimäki, S., & Salmi, A.-K. (2021). Impact of selection and
493 domestication on hindlimb bones of modern reindeer populations: Archaeological
494 implications for early reindeer management by Sámi in Fennoscandia. *Historical Biology*,
495 *0*(0), 1–19. https://doi.org/10.1080/08912963.2021.1947268

496 Price, E. O. (1999). Behavioral development in animals undergoing domestication. *Applied Animal
497 Behaviour Science*, 65(3), 245–271. https://doi.org/10.1016/S0168-1591(99)00087-8

498 R Core Team. (2019). R: A language and environment for statistical computing (Version 3.1.2).
499 Vienna, Austria.

500 Rhodes, R. T., Appleby, M. C., Chinn, K., Douglas, L., Firkins, L. D., Houpt, K. A., ... Wills, R. W.
501 (2005). A comprehensive review of housing for pregnant sows. *Journal of the American*
502 Veterinary Medical Association, 227(10), 1580–1590.

503 https://doi.org/10.2460/javma.2005.227.1580

504 Rohlf, F. J., & Slice, D. (1990). Extensions of the Procrustes method for the optimal superimposition
505 of landmarks. *Systematic Biology*, *39*(1), 40–59. https://doi.org/10.2307/2992207

506 Russo, L., Massei, G., & Genov, P. V. (1997). Daily home range and activity of wild boar in a
507 Mediterranean area free from hunting. *Ethology Ecology & Evolution*, *9*(3), 287–294.

508 https://doi.org/10.1080/08927014.1997.9522888

509 Sánchez-Villagra, M. R., Geiger, M., & Schneider, R. A. (2016). The taming of the neural crest: A
510 developmental perspective on the origins of morphological covariation in domesticated
511 mammals. *Royal Society Open Science*, *3*(6), 160107. https://doi.org/10.1098/rsos.160107

512 Sánchez-Villagra, M. R., Segura, V., Geiger, M., Heck, L., Veitschegger, K., & Flores, D. (2017). On
513 the lack of a universal pattern associated with mammalian domestication: Differences in skull
514 growth trajectories across phylogeny. *Royal Society Open Science*, *4*(10), 170876.

515 https://doi.org/10.1098/rsos.170876

516 Schaller, G. B. (1973). *The Serengeti Lion: A Study of Predator-Prey Relations* (University of
517 Chicago). Chicago.

518 Schlager, S., & Jefferis, G. (2020). *Morpho: Calculations and visualisations related to geometric
519 morphometrics*.

520 Schley, L., & Roper, T. J. (2003). Diet of wild boar *Sus scrofa* in Western Europe, with particular
521 reference to consumption of agricultural crops. *Mammal Review*, *33*(1), 43–56.

522 https://doi.org/10.1046/j.1365-2907.2003.00010.x

523 Smuts, G. L., Anderson, J. L., & Austin, J. C. (1978). Age determination of the African lion
524 (*Panthera leo*). *Journal of Zoology*, *185*(1), 115–146. https://doi.org/10.1111/j.1469-5255.1978.tb03317.x
526 Su, S. C., Skedros, J. G., Bachus, K. N., & Bloebaum, R. D. (1999). Loading conditions and cortical bone construction of an artiodactyl calcaneus. *Journal of Experimental Biology*, 202(22), 3239–3254. https://doi.org/10.1242/jeb.202.22.3239

529 Turner, T. R., Cramer, J. D., Nisbett, A., & Gray, J. P. (2016). A comparison of adult body size between captive and wild vervet monkeys (*Chlorocebus aethiops sabaeus*) on the island of St. Kitts. *Primates; Journal of Primatology*, 57(2), 211–220. https://doi.org/10.1007/s10329-015-0509-8

533 Vigne, J.-D. (2011). The origins of animal domestication and husbandry: A major change in the history of humanity and the biosphere. *Comptes Rendus Biologies*, 334(3), 171–181. https://doi.org/10.1016/j.crvi.2010.12.009

536 Vigne, J.-D., Carrère, I., Briois, F., & Guilaine, J. (2011). The Early Process of Mammal Domestication in the Near East: New Evidence from the Pre-Neolithic and Pre-Pottery Neolithic in Cyprus. *Current Anthropology*, 52(S4), S255–S271. https://doi.org/10.1086/659306

540 Voje, K. L., Hansen, T. F., Egset, C. K., Bolstad, G. H., & Pélabon, C. (2014). Allometric Constraints and the Evolution of Allometry. *Evolution*, 68(3), 866–885. https://doi.org/10.1111/evo.12312

543 Wilson, L. A. B. (2018). The evolution of ontogenetic allometric trajectories in mammalian domestication. *Evolution*, 72(4), 867–877. https://doi.org/10.1111/evo.13464

545 Zeder, M A. (2012). Pathways to animal domestication. In P. Gepts, T. R. Famula, & R. L. Bettinger (Eds.), *Biodiversity in Agriculture: Domestication, Evolution, and Sustainability* (pp. 227–259). Cambridge, UK: Cambridge University Press.
548 Zeder, M A. (2018). Why evolutionary biology needs anthropology: Evaluating core assumptions of
549 the extended evolutionary synthesis. *Evolutionary Anthropology*, 27(6), 267–284.
550 https://doi.org/10.1002/evan.21747

551 Zihlman, A. L., Bolter, D. R., & Boesch, C. (2007). Skeletal and dental growth and development in
552 chimpanzees of the Taï National Park, Côte D’Ivoire. *Journal of Zoology*, 273(1), 63–73.
553 https://doi.org/10.1111/j.1469-7998.2007.00301.x

554
<table><caption>Table 1. MANOVA *p*-values of the form coordinates between captive (stall and enclosure) and wild-caught groups of wild boar (*Sus scrofa*) computed for the cranium, mandible, and calcaneus. Significant values (*p* < 0.05) are in bold. m.: months.</caption><thead><tr><th rowspan="3"></th><th colspan="2">Cranium</th><th colspan="2">Mandible</th><th colspan="2">Calcaneus</th></tr><tr><th>8-10 m.</th><th>16-18m.</th><th>8-10 m.</th><th>16-18 m.</th><th>8-10 m.</th><th>16-18 m.</th></tr><tr><th>wild-caught</th><th>wild-caught</th><th>wild-caught</th><th>wild-caught</th><th>wild-caught</th><th>wild-caught</th></tr></thead><tbody><tr><td>6 m. - enclosure</td><td>0.21</td><td><strong>&lt; 0.01</strong></td><td>0.55</td><td><strong>&lt; 0.01</strong></td><td>0.49</td><td><strong>&lt; 0.01</strong></td></tr><tr><td>6 m. - stall</td><td>0.27</td><td><strong>&lt; 0.01</strong></td><td>0.74</td><td><strong>&lt; 0.01</strong></td><td>0.35</td><td><strong>0.01</strong></td></tr><tr><td>8 m. - enclosure</td><td>0.11</td><td><strong>0.02</strong></td><td>0.27</td><td>0.09</td><td>0.13</td><td>0.09</td></tr><tr><td>8 m. - stall</td><td>0.22</td><td><strong>0.01</strong></td><td>0.38</td><td>0.05</td><td>0.22</td><td><strong>0.03</strong></td></tr><tr><td>11 m. - enclosure</td><td><strong>&lt; 0.01</strong></td><td>0.12</td><td><strong>0.01</strong></td><td>0.42</td><td><strong>0.01</strong></td><td>0.35</td></tr><tr><td>11 m. - stall</td><td><strong>0.01</strong></td><td>0.13</td><td><strong>0.01</strong></td><td>0.30</td><td><strong>&lt; 0.01</strong></td><td>0.20</td></tr><tr><td>14 m. - enclosure</td><td><strong>&lt; 0.01</strong></td><td>0.17</td><td><strong>&lt; 0.01</strong></td><td>0.21</td><td><strong>&lt; 0.01</strong></td><td>0.25</td></tr><tr><td>14 m. - stall</td><td><strong>&lt; 0.01</strong></td><td>0.15</td><td><strong>&lt; 0.01</strong></td><td>0.12</td><td><strong>&lt; 0.01</strong></td><td>0.09</td></tr><tr><td>20 m. - enclosure</td><td><strong>&lt; 0.01</strong></td><td><strong>0.01</strong></td><td><strong>&lt; 0.01</strong></td><td><strong>0.01</strong></td><td><strong>&lt; 0.01</strong></td><td><strong>0.02</strong></td></tr><tr><td>20 m. - stall</td><td><strong>&lt; 0.01</strong></td><td><strong>0.01</strong></td><td><strong>&lt; 0.01</strong></td><td><strong>0.01</strong></td><td><strong>&lt; 0.01</strong></td><td><strong>0.01</strong></td></tr></tbody></table>
Table 2. ANOVA *p*-values of the centroid size (CS) between captive (stall and enclosure) and wild-caught groups of wild boar (*Sus scrofa*) computed for the cranium, mandible, and calcaneus and ANOVA *p*-values of body mass. Significant values (*p* < 0.05) are in bold. . m.: months.

<table><thead><tr><th rowspan="2"></th><th colspan="2">CS Cranium</th><th colspan="2">CS Mandible</th><th colspan="2">CS Calcaneus</th><th colspan="2">Body mass</th></tr><tr><th>8 to 10<br>m.<br>wild-caught</th><th>16 to 18<br>m.<br>wild-caught</th><th>8 to 10<br>m.<br>wild-caught</th><th>16 to 18<br>m.<br>wild-caught</th><th>8 to 10<br>m.<br>wild-caught</th><th>16 to 18<br>m.<br>wild-caught</th><th>8 to 10<br>m.<br>wild-caught</th><th>16 to 18<br>m.<br>wild-caught</th></tr></thead><tbody><tr><td>6 m. – enclosure</td><td>0.75</td><td><strong>&lt; 0.01</strong></td><td>0.54</td><td><strong>&lt; 0.01</strong></td><td>0.93</td><td><strong>&lt; 0.01</strong></td><td>0.50</td><td><strong>&lt; 0.01</strong></td></tr><tr><td>6 m. – stall</td><td>0.67</td><td><strong>&lt; 0.01</strong></td><td>0.58</td><td><strong>0.01</strong></td><td>0.78</td><td><strong>0.01</strong></td><td>0.45</td><td><strong>&lt; 0.01</strong></td></tr><tr><td>8 m. – enclosure</td><td>0.37</td><td><strong>0.03</strong></td><td>0.39</td><td>0.09</td><td>0.18</td><td>0.12</td><td>0.81</td><td><strong>0.01</strong></td></tr><tr><td>8 m. – stall</td><td>0.60</td><td><strong>0.00</strong></td><td>0.61</td><td>0.05</td><td>0.32</td><td>0.06</td><td>0.64</td><td><strong>&lt; 0.02</strong></td></tr><tr><td>11 m. – enclosure</td><td><strong>0.01</strong></td><td>0.42</td><td><strong>&lt; 0.01</strong></td><td>0.79</td><td><strong>0.01</strong></td><td>0.62</td><td>0.20</td><td>0.12</td></tr><tr><td>11 m. – stall</td><td><strong>0.02</strong></td><td>0.37</td><td><strong>0.02</strong></td><td>0.81</td><td><strong>&lt; 0.01</strong></td><td>0.95</td><td>0.16</td><td>0.22</td></tr><tr><td>14 m. – enclosure</td><td><strong>&lt; 0.01</strong></td><td>0.71</td><td><strong>&lt; 0.01</strong></td><td>0.49</td><td><strong>&lt; 0.01</strong></td><td>0.63</td><td><strong>&lt; 0.02</strong></td><td>0.60</td></tr><tr><td>14 m. – stall</td><td><strong>&lt; 0.01</strong></td><td>0.60</td><td><strong>&lt; 0.01</strong></td><td>0.33</td><td><strong>&lt; 0.01</strong></td><td>0.28</td><td><strong>0.01</strong></td><td>0.99</td></tr><tr><td>20 m. – enclosure</td><td><strong>&lt; 0.01</strong></td><td><strong>0.04</strong></td><td><strong>&lt; 0.01</strong></td><td><strong>0.02</strong></td><td><strong>&lt; 0.01</strong></td><td>0.12</td><td><strong>&lt; 0.01</strong></td><td>0.06</td></tr><tr><td>20 m. – stall</td><td><strong>&lt; 0.01</strong></td><td><strong>0.02</strong></td><td><strong>&lt; 0.01</strong></td><td><strong>0.01</strong></td><td><strong>&lt; 0.01</strong></td><td>0.08</td><td><strong>&lt; 0.01</strong></td><td><strong>0.03</strong></td></tr></tbody></table>
<figure><img src="image_3.png" /></figure>

a.

<figure><img src="image_4.png" /></figure>

wild-caught 8-10 months to 16-18 months

d.

<figure><img src="image_5.png" /></figure>

b.

<figure><img src="image_6.png" /></figure>

wild-caught 8-10 months to 16-18 months

e.

<figure><img src="image_7.png" /></figure>

c.

<figure><img src="image_8.png" /></figure>

wild-caught 8-10 months to 16-18 months

f.
Figure 1. Principal component analyses for the cranium (a) mandible (b), and calcaneus (c) of wild boar (*Sus scrofa*) in the PC1-PC2 form space. Symbols represent the mean form for each group. Black dots represent the “control” specimens from the wild-caught group, i.e. the specimens belonging to the same initial population as the individuals from the stall and enclosure groups. Form changes are depicted in lateral, dorsal and frontal views for the cranium and the mandible, and in medial and plantar views for the calcaneus. Heatmap of the intensity of form variation between captive (i.e. enclosure and stall) groups (6 months and 20 months) and between wild-caught groups (8-10 months and 16-18 months) for the cranium (d), mandible (e), and calcaneus (f). Blue indicates a low intensity of variation and red indicates a high intensity of variation. Form changes are depicted in lateral, dorsal and frontal views for the cranium and the mandible, and in medial and plantar views for the calcaneus
<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /></figure>

<figure><img src="image_11.png" /></figure>

<figure><img src="image_12.png" /><figcaption>Figure 2. Boxplots of centroid size (CS) for the cranium (a), mandible (b), and calcaneus (c) and of body mass (d) of wild boar (*Sus scrofa*). The vertical lines represent all the values within 1.5 times of the interquartile range accounting for 50% of the data, from the 25th percentile to the 75th percentile. The horizontal lines are the median values.</figcaption></figure>
<figure><img src="image_13.png" /></figure>

<figure><img src="image_14.png" /></figure>

<figure><img src="image_15.png" /></figure>
Figure 3. Regression of log centroid size (CS) on shape scores for the cranium (a), mandible (b) and calcaneus (c) of wild boar (*Sus scrofa*) and effects of size, group, and interaction between size and group on the regression of log CS on shape scores. Significant values (*p* < 0.05) are in bold.

583

584

585

586

587

588
589 The effect of captivity on craniomandibular and calcaneal ontogenetic trajectories in wild boar

590 Short running title: Captivity and ontogenetic trajectories

591 Dimitri Neaux¹,²*, Hugo Harbers¹, Barbara Blanc³, Katia Ortiz³,⁴, Yann Locatelli³,⁵, Anthony Herrel⁶,

592 Vincent Debat⁴, Thomas Cucchi¹*

593

594 ¹ Archéozoologie, Archéobotanique: Sociétés, Pratiques et Environnements, UMR 7209, Muséum

595 national d'Histoire naturelle CNRS, Paris, France

596 ² Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie, UMR 7262, Université de

597 Poitiers CNRS, Poitiers, France

598 ³ Réserve Zoologique de la Haute-Touche, Muséum national d'Histoire naturelle, Obterre, France

599 ⁴ Institut de Systématique, Evolution, Biodiversité, UMR 7205, Muséum national d'Histoire naturelle

600 CNRS UPMC EPHE, UA, Paris, France

601 ⁵ Physiologie de la Reproduction et des Comportements, UMR 7247, INRAE CNRS Université de

602 Tours IFCE, Nouzilly, France

603 ⁶ Mécanismes Adaptatifs et Evolution, UMR 7179, Muséum national d'Histoire naturelle CNRS, Paris,

604 France

605

606 *Corresponding authors: Dimitri Neaux, Thomas Cucchi

607 dimitrineaux@gmail.com, thomas.cucchi@mnhn.fr

608
611 a. List of groups included in the study and number of specimens

<table><thead><tr><th>Cranium</th><th>Cranium</th><th>Mandible</th><th>Calcaneus</th></tr></thead><tbody><tr><td>Enclosure</td><td>12</td><td>12</td><td>12</td></tr><tr><td>Stall</td><td>12</td><td>12</td><td>12</td></tr><tr><td>8-10 months wild-caught</td><td>10</td><td>12</td><td>10</td></tr><tr><td>16-18 months wild-caught</td><td>8</td><td>6</td><td>8</td></tr><tr><td>TOTAL</td><td>42</td><td>42</td><td>42</td></tr></tbody></table>
b. List of specimens. M: male, F: female, nd: not determined individuals. Specimens are localised at the Muséum national d'Histoire naturelle (Paris, France)

<table><thead><tr><th>Catalogue number</th><th>Sex<sup>1</sup></th><th>Age class</th><th>Status</th><th>Location</th><th>Cranium</th><th>Mandible</th><th>Calcaneus</th></tr></thead><tbody><tr><td>2017-557</td><td>F</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>H285</td><td>M</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-560</td><td>M</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-562</td><td>M</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-555</td><td>F</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-556</td><td>F</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-569</td><td>F</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>H319</td><td>F</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-554</td><td>F</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-571</td><td>M</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-574</td><td>M</td><td></td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr></tbody></table>
<table><tr><td>2017-575</td><td>M</td><td>stall</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-558</td><td>M</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-559</td><td>F</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-561</td><td>M</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-563</td><td>M</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-564</td><td>M</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-565</td><td>F</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-566</td><td>F</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-567</td><td>F</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-568</td><td>F</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-570</td><td>F</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-572</td><td>M</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-573</td><td>M</td><td>enclosure</td><td>Réserve de la Haute-Touche</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>PRA_186</td><td>nd</td><td>8-10 months</td><td>wild-caught</td><td>Urciers</td><td>Yes</td><td>Yes</td><td>No</td></tr><tr><td>PRA_172</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Urciers</td><td>Yes</td><td>Yes</td><td>Yes</td></tr></table>
<table><tr><td>PRA_174</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Urciers</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-582</td><td>M</td><td>8-10 months</td><td>wild-caught</td><td>Urciers</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-584</td><td>M</td><td>8-10 months</td><td>wild-caught</td><td>Urciers</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>CHA_S_77</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>CHA_S_509</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>CHA_S_581</td><td>M</td><td>8-10 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>CHA_S_664</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>No</td><td>Yes</td></tr><tr><td>CHA_S_577</td><td>nd</td><td>8-10 months</td><td>wild-caught</td><td>Chambord</td><td>No</td><td>Yes</td><td>No</td></tr><tr><td>COMP_2013-1262</td><td>nd</td><td>8-10 months</td><td>wild-caught</td><td>Compiègne</td><td>Yes</td><td>Yes</td><td>No</td></tr><tr><td>COMP_2013-1247</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Compiègne</td><td>No</td><td>Yes</td><td>Yes</td></tr><tr><td>COMP_2013-1269</td><td>F</td><td>8-10 months</td><td>wild-caught</td><td>Compiègne</td><td>No</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-583</td><td>M</td><td>16-18 months</td><td>wild-caught</td><td>Urciers</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-581</td><td>F</td><td>16-18 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-577</td><td>M</td><td>16-18 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-579</td><td>F</td><td>16-18 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr></table>
<table><tr><td>2017-580</td><td>F</td><td>16-18 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>2017-578</td><td>F</td><td>16-18 months</td><td>wild-caught</td><td>Chambord</td><td>Yes</td><td>No</td><td>Yes</td></tr><tr><td>COMP_2013-1264</td><td>F</td><td>16-18 months</td><td>wild-caught</td><td>Compiègne</td><td>Yes</td><td>Yes</td><td>Yes</td></tr><tr><td>COMP_2013-1270</td><td>M</td><td>16-18 months</td><td>wild-caught</td><td>Compiègne</td><td>Yes</td><td>No</td><td>Yes</td></tr></table>

¹Sexes and ages for wild-caught specimens were estimated based on osteological observations, using respectively the morphology of canine cross section (Mayer & Brisbin, 1988) and the mandibular tooth eruption and wear stages in occlusal view (Grant, 1982).

Grant, A., 1982. The use of tooth wear as a guide to the domestic ungulates. In: Wilson, B., Grigson, C., Payne, S. (Eds.), Ageing and Sexing Animal Bones from Archaeological Sites. UK, pp. 991–108.

Mayer, J.M., & Brisbin, I.L., 1988. Sex identification of *Sus scrofa* based on canine morphology. Journal of Mammalogy 69:408–4
# Appendix S2: Digitisation and definitions of landmarks

## a. Digitisation protocol

All specimens were scanned using a Computed Tomography (CT) scanner with a spatial resolution of between 100 and 500 µm. The wild boar from Urciers were scanned as living specimens at the *Chirurgie et Imagerie pour la Recherche et l'Enseignement* (CIRE) platform of the *Institut National de Recherche pour l'Agriculture, l'Alimentation et l'Environnement* (INRAE). Other individuals were scanned as dry specimens using a CT scanner close to the collections they were housed in. We segmented the bones using the segmentation tools of the Avizo v8.0 software, and then converted the volumes into three-dimensional PLY surfaces format. We digitised the anatomical landmarks and semilandmarks using IDAV Landmark v3.0 software (Wiley et al., 2005). To remove variation related to their initial arbitrary position along the curves, the semilandmarks were slid along the tangent of the curves minimising bending energy (Gunz and Mitteroecker, 2013).

Gunz, P., Mitteroecker, P., 2013. Semilandmarks: a method for quantifying curves and surfaces. Hystrix, the Italian Journal of Mammalogy. 24, 103–109.

Wiley, D., Amenta, N., Alcantara, D., Ghosh, D., Kil, Y.J., Delson, E., Harcourt-Smith, W., Rohlf, F.J., St. John, K., Hamann, B., Motani, R., Frost, S., Rosenberger, A.L., Tallman, L., Disotell, T., O'Neill, R., 2005. Evolutionary Morphing. In: Proceedings of IEEE Visualization 2005. Presented at the VIS'05, IEEE, Minneapolis, MN, pp. 431–438.
b. Number of landmarks

<table><thead><tr><th></th><th>Homologous<br>landmarks</th><th>Sliding semi-<br>landmarks</th><th>Surface sliding semi-<br>landmarks</th></tr></thead><tbody><tr><td>Cranium</td><td>14</td><td>181</td><td>0</td></tr><tr><td>Mandible</td><td>70</td><td>28</td><td>0</td></tr><tr><td>Calcaneus</td><td>23</td><td>48</td><td>763</td></tr></tbody></table>
c. Definitions of cranial (1-70), mandibular (71-94), and calcaneus (95-108) homologous landmarks.

<table><thead><tr><td>Landmark</td><td>Definition</td></tr></thead><tbody><tr><td>1</td><td>Most anterior midline point of the nasals</td></tr><tr><td>2</td><td>Most anterior, dorsal midline point of the premaxillae</td></tr><tr><td>3, 4</td><td>Most anterior point of the nasal-premaxilla suture</td></tr><tr><td>5, 6</td><td>Most anterior, lateral point of the upper canine alveolus</td></tr><tr><td>7, 8</td><td>Suture at the meeting point of premaxilla, maxilla, and nasal</td></tr><tr><td>9, 10</td><td>Most anterior point of the infraorbital foramen</td></tr><tr><td>11, 12</td><td>Most posterior point of the infraorbital foramen</td></tr><tr><td>13, 14</td><td>Most anterior lateral point of the facial tuberosity</td></tr><tr><td>15, 16</td><td>Most ventral point of the zygomatic-maxilla suture</td></tr><tr><td>17, 18</td><td>Most anterior, lateral point of the orbit</td></tr><tr><td>19, 20</td><td>Most dorsal point of the lower lacrimal foramen</td></tr><tr><td>21, 22</td><td>Most posterior point of the supraorbital foramen</td></tr><tr><td>23, 24</td><td>Most dorsal point of the orbit</td></tr><tr><td>25, 26</td><td>Most ventral point of supraorbital process of the frontal bone</td></tr><tr><td>27, 28</td><td>Meeting point of the parietal-frontal suture and temporal line</td></tr><tr><td>29, 30</td><td>Most anterior, dorsal point of the zygomatic process of the squamosal bone</td></tr><tr><td>31, 32</td><td>Most posterior point of the zygomatic bone</td></tr><tr><td>33, 34</td><td>Most dorsal point of the zygomatic process of the squamosal bone</td></tr><tr><td>35, 36</td><td>Most anterior, lateral point of the nuchal crest</td></tr><tr><td>37, 38</td><td>Most anterior point of the palatine fissure</td></tr><tr><td>39, 40</td><td>Most posterior point of the palatine fissure</td></tr></tbody></table>
<table>
  <tr>
    <td>41, 42</td>
    <td>Most anterior point of the cheek-tooth row (excluding P1)</td>
  </tr>
  <tr>
    <td>43, 44</td>
    <td>Most posterior point of the cheek-tooth row</td>
  </tr>
  <tr>
    <td>45</td>
    <td>Most posterior point of the posterior nasal spine on the palatine bone</td>
  </tr>
  <tr>
    <td>46, 47</td>
    <td>Most ventral, lateral point of the pterygoid process of the sphenoid</td>
  </tr>
  <tr>
    <td>48, 49</td>
    <td>Most posterior point of the pterygoid hamulus</td>
  </tr>
  <tr>
    <td>50, 51</td>
    <td>Meeting point of the pterygoid process with the ridge of the lateral pterygoid plate</td>
  </tr>
  <tr>
    <td>52, 53</td>
    <td>Meeting point of the pterygoid hamulus with the ridge of the medial pterygoid plate</td>
  </tr>
  <tr>
    <td>54</td>
    <td>Most posterior point of the vomer in contact with the sphenoid</td>
  </tr>
  <tr>
    <td>55, 56</td>
    <td>Most ventral, lateral, posterior point of the sphenoid-squamosal suture</td>
  </tr>
  <tr>
    <td>57, 58</td>
    <td>Most ventral, medial, posterior point of the sphenoid-squamosal suture</td>
  </tr>
  <tr>
    <td>59, 60</td>
    <td>Most posterior, medial point of the petro-occipital fissure</td>
  </tr>
  <tr>
    <td>61, 62</td>
    <td>Most lateral point of the occipital condyle</td>
  </tr>
  <tr>
    <td>63</td>
    <td>Most anterior, ventral midline point of the premaxilla</td>
  </tr>
  <tr>
    <td>64</td>
    <td>Most posterior midline point of the nuchal crest</td>
  </tr>
  <tr>
    <td>65, 66</td>
    <td>Most posterior, lateral point of the nuchal crest</td>
  </tr>
  <tr>
    <td>67, 68</td>
    <td>Most lateral point of the foramen magnum</td>
  </tr>
  <tr>
    <td>69</td>
    <td>Most posterior, dorsal point of the foramen magnum</td>
  </tr>
  <tr>
    <td>70</td>
    <td>Most anterior point, ventral of the foramen magnum</td>
  </tr>
  <tr>
    <td>71, 72</td>
    <td>Most anterior, lateral point of the lower canine alveolus</td>
  </tr>
  <tr>
    <td>73, 74</td>
    <td>Most anterior point of the cheek-tooth row (excluding P1)</td>
  </tr>
  <tr>
    <td>75, 76</td>
    <td>Most lateral point at the maximum of curvature between the mandibular ramus and corpus</td>
  </tr>
  <tr>
    <td>77, 78</td>
    <td>Most lateral point at the maximum of curvature between the coronoid process and the mandibular ramus</td>
  </tr>
</table>
<table>
  <tr>
    <td>79, 80</td>
    <td>Most dorsal point of the coronoid process</td>
  </tr>
  <tr>
    <td>81, 82</td>
    <td>Most lateral point of the mandibular condyle</td>
  </tr>
  <tr>
    <td>83, 84</td>
    <td>Most posterior point of the mandibular condyle</td>
  </tr>
  <tr>
    <td>85, 86</td>
    <td>Point at the maximum of curvature of the mandibular angle</td>
  </tr>
  <tr>
    <td>87</td>
    <td>Most ventral, posterior point of the mandibular symphysis</td>
  </tr>
  <tr>
    <td>88</td>
    <td>Most ventral, anterior point of the mandibular symphysis</td>
  </tr>
  <tr>
    <td>89, 90</td>
    <td>Most medial point of the mandibular condyle</td>
  </tr>
  <tr>
    <td>91</td>
    <td>Most dorsal, posterior point of the mandibular symphysis</td>
  </tr>
  <tr>
    <td>92</td>
    <td>Most dorsal, anterior point of the mandibular symphysis</td>
  </tr>
  <tr>
    <td>93, 94</td>
    <td>Most anterior point of the mandibular foramen</td>
  </tr>
  <tr>
    <td>95</td>
    <td>Distal end of the cuboid facet</td>
  </tr>
  <tr>
    <td>96</td>
    <td>Proximo-plantar end of the cuboid facet</td>
  </tr>
  <tr>
    <td>97</td>
    <td>End of the beak of the coracoid process</td>
  </tr>
  <tr>
    <td>98</td>
    <td>Maximum of curvature of the plantar bulge on the plantar margin</td>
  </tr>
  <tr>
    <td>99</td>
    <td>Dorso-proximal end of the calcaneal sulcus</td>
  </tr>
  <tr>
    <td>100</td>
    <td>Planto-lateral end of sustentaculum tali</td>
  </tr>
  <tr>
    <td>101</td>
    <td>Dorsal end of the sustentaculum tali</td>
  </tr>
  <tr>
    <td>102</td>
    <td>Medial end of sustentaculum tali</td>
  </tr>
  <tr>
    <td>103</td>
    <td>Plantar end of the epiphysis</td>
  </tr>
  <tr>
    <td>104</td>
    <td>Dorso-proximal end of the bulge of the proximal part (not on the epiphysis)</td>
  </tr>
  <tr>
    <td>105</td>
    <td>Proximal end of the lateral lobe of the epiphysis (secondary lobe)</td>
  </tr>
  <tr>
    <td>106</td>
    <td>Proximal end of the medial lobe of the epiphysis (main lobe)</td>
  </tr>
  <tr>
    <td>107</td>
    <td>Dorsal end of the epiphysis</td>
  </tr>
  <tr>
    <td>108</td>
    <td>Dorso-proximal end of the lateral part of the coracoid process</td>
  </tr>
</table>
d. Definitions of cranial (I to XIV), mandibular (XV to XXII), and calcaneus (XVI to XXIX) curves

<table><thead><tr><td>Curve</td><td>Definition</td></tr></thead><tbody><tr><td>I</td><td>from LM 13 to LM31</td></tr><tr><td>II</td><td>from LM 14 to LM32</td></tr><tr><td>III</td><td>from LM 29 to LM33</td></tr><tr><td>IV</td><td>from LM 30 to LM34</td></tr><tr><td>V</td><td>from LM 27 to LM35</td></tr><tr><td>VI</td><td>from LM 28 to LM36</td></tr><tr><td>VII</td><td>from LM 1 to LM64</td></tr><tr><td>VIII</td><td>from LM 45 to LM63</td></tr><tr><td>IX</td><td>from LM 50 to LM55</td></tr><tr><td>X</td><td>from LM 51 to LM56</td></tr><tr><td>XI</td><td>from LM 52 to LM57</td></tr><tr><td>XII</td><td>from LM 53 to LM58</td></tr><tr><td>XIII</td><td>from LM 64 to LM65</td></tr><tr><td>XIV</td><td>from LM 64 to LM66</td></tr><tr><td>XV</td><td>from LM 75 to LM77</td></tr><tr><td>XVI</td><td>from LM 76 to LM78</td></tr><tr><td>XVII</td><td>from LM 83 to LM85</td></tr><tr><td>XVIII</td><td>from LM 84 to LM86</td></tr><tr><td>XIX</td><td>from LM 85 to LM87</td></tr><tr><td>XX</td><td>from LM 86 to LM87</td></tr><tr><td>XXI</td><td>from LM 87 to LM88</td></tr></tbody></table>
XXII from LM 91 to LM92

XXIII Edge of the articular surface of the cuboid facet

XXIV Medial edge of the coracoid process

XXV Edge of the articular surface of the sustentaculum tali

XXVI Lateral edge of the coracoid process

XXVII Edge of the attachment surface of the tendon on the epiphysis

XXVIII Distal delineation of the junction zone between the epiphysis and the rest of the calcaneus

XXIX Proximal delineation of the junction zone between the epiphysis and the rest of the calcaneus
<figure><img src="image_16.png" /><figcaption>e. Wild boar (*Sus scrofa*) cranium and mandible showing the homologous landmarks (red dots and Arabic numerals) and semilandmarks (blue dots and Roman numerals) used in the study.</figcaption></figure>
f. Wild boar (*Sus scrofa*) calcaneus showing the homologous landmarks (red dots and Arabic numerals) and semilandmarks (blue dots and Roman numerals) used in the study.

<figure><img src="image_17.png" /></figure>
<table><caption>Appendix S3: Body mass measurements (kg)</caption><thead><tr><th rowspan="2"></th><th colspan="5">Captive wild boar</th></tr><tr><th>6 months</th><th>8 months</th><th>11 months</th><th>14 months</th><th>20 months</th></tr></thead><tbody><tr><td>2017-557</td><td>18.5</td><td>32.5</td><td>51.0</td><td>60.5</td><td>83.0</td></tr><tr><td>H285</td><td>24.0</td><td>27.0</td><td>30.5</td><td>38.0</td><td>59.0</td></tr><tr><td>2017-560</td><td>21.5</td><td>28.5</td><td>55.5</td><td>56.5</td><td>77.0</td></tr><tr><td>2017-562</td><td>29.0</td><td>37.5</td><td>40.5</td><td>50.0</td><td>76.0</td></tr><tr><td>2017-555</td><td>15.5</td><td>21.5</td><td>31.0</td><td>49.5</td><td>77.0</td></tr><tr><td>2017-556</td><td>27.0</td><td>34.0</td><td>35.0</td><td>40.0</td><td>59.0</td></tr><tr><td>2017-569</td><td>26.0</td><td>36.5</td><td>38.5</td><td>45.0</td><td>57.2</td></tr><tr><td>H319</td><td>19.5</td><td>29.5</td><td>51.5</td><td>67.5</td><td>97.8</td></tr><tr><td>2017-554</td><td>21.5</td><td>31.5</td><td>42.5</td><td>54.0</td><td>77.0</td></tr><tr><td>2017-571</td><td>18.0</td><td>24.0</td><td>31.0</td><td>44.5</td><td>68.0</td></tr><tr><td>2017-574</td><td>33.0</td><td>37.5</td><td>50.0</td><td>67.0</td><td>95.5</td></tr><tr><td>2017-575</td><td>30.5</td><td>35.0</td><td>42.5</td><td>57.5</td><td>78.5</td></tr><tr><td>2017-558</td><td>19.0</td><td>30.5</td><td>40.0</td><td>55.5</td><td>80.5</td></tr><tr><td>2017-559</td><td>18.0</td><td>24.0</td><td>36.0</td><td>43.0</td><td>62.0</td></tr><tr><td>2017-561</td><td>26.0</td><td>39.5</td><td>47.0</td><td>59.5</td><td>83.0</td></tr><tr><td>2017-563</td><td>31.0</td><td>45.0</td><td>56.5</td><td>73.5</td><td>89.0</td></tr><tr><td>2017-564</td><td>22.0</td><td>31.2</td><td>30.5</td><td>44.5</td><td>61.0</td></tr><tr><td>2017-565</td><td>17.5</td><td>29.0</td><td>33.0</td><td>41.5</td><td>61.5</td></tr><tr><td>2017-566</td><td>21.0</td><td>32.0</td><td>38.0</td><td>46.0</td><td>66.5</td></tr><tr><td>2017-567</td><td>32.0</td><td>38.0</td><td>50.0</td><td>53.5</td><td>81.0</td></tr><tr><td>2017-568</td><td>27.0</td><td>38.5</td><td>48.5</td><td>50.5</td><td>77.0</td></tr></tbody></table>
<table><tbody><tr><td>2017-570</td><td>19.5</td><td>30.0</td><td>40.0</td><td>45.0</td><td>67.0</td></tr><tr><td>2017-572</td><td>32.0</td><td>45.5</td><td>55.0</td><td>73.0</td><td>90.0</td></tr><tr><td>2017-573</td><td>24.0</td><td>37.5</td><td>36.5</td><td>47.0</td><td>73.5</td></tr><tr><td colspan="6" style="text-align: center;">wild-caught wild boar</td></tr><tr><td></td><td>8-10 months</td><td>16-18 months</td><td></td><td></td><td></td></tr><tr><td>PRA_186</td><td>30.0</td><td></td><td></td><td></td><td></td></tr><tr><td>PRA_172</td><td>23.0</td><td></td><td></td><td></td><td></td></tr><tr><td>PRA_174</td><td>28.0</td><td></td><td></td><td></td><td></td></tr><tr><td>2017-582</td><td>35.0</td><td></td><td></td><td></td><td></td></tr><tr><td>2017-584</td><td>52.0</td><td></td><td></td><td></td><td></td></tr><tr><td>CHA_S_77</td><td>20.9</td><td></td><td></td><td></td><td></td></tr><tr><td>CHA_S_509</td><td>24.3</td><td></td><td></td><td></td><td></td></tr><tr><td>CHA_S_581</td><td>20.5</td><td></td><td></td><td></td><td></td></tr><tr><td>CHA_S_664</td><td>36.0</td><td></td><td></td><td></td><td></td></tr><tr><td>CHA_S_577</td><td>25.0</td><td></td><td></td><td></td><td></td></tr><tr><td>COMP_2013-1262</td><td>NA</td><td></td><td></td><td></td><td></td></tr><tr><td>COMP_2013-1247</td><td>23.9</td><td></td><td></td><td></td><td></td></tr><tr><td>COMP_2013-1269</td><td>39.7</td><td></td><td></td><td></td><td></td></tr><tr><td>2017-583</td><td></td><td>53.0</td><td></td><td></td><td></td></tr><tr><td>2017-581</td><td></td><td>49.8</td><td></td><td></td><td></td></tr><tr><td>2017-577</td><td></td><td>35.3</td><td></td><td></td><td></td></tr><tr><td>2017-579</td><td></td><td>31.7</td><td></td><td></td><td></td></tr><tr><td>2017-580</td><td></td><td>68.5</td><td></td><td></td><td></td></tr><tr><td>2017-578</td><td></td><td>20.0</td><td></td><td></td><td></td></tr></tbody></table>
COMP_2013-1264 60.8

COMP_2013-1270 86.0
# 1 Appendix S4: Pairwise analyses between all groups

## 2 a. MANOVA p-values of the form coordinates between all groups computed for the cranium,

## 3 mandible, and calcaneus.

<table><thead><tr><th rowspan="2">Cranium</th><th>8 to 10</th><th>16 to 18</th><th rowspan="2">6 months<br>enclosure</th><th rowspan="2">6 months<br>stall</th><th rowspan="2">8 months<br>enclosure</th><th rowspan="2">8 months<br>stall</th><th>11</th></tr><tr><th>months wild<br>caught</th><th>months wild<br>caught</th><th>months<br>enclosure</th></tr></thead><tbody><tr><td>16-18 months wild caught</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.21</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.27</td><td>0.00</td><td>0.98</td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.11</td><td>0.02</td><td>0.24</td><td>0.20</td><td></td><td></td><td></td></tr><tr><td>8 months stall</td><td>0.22</td><td>0.01</td><td>0.34</td><td>0.46</td><td>0.88</td><td></td><td></td></tr><tr><td>11 months enclosure</td><td>0.00</td><td>0.12</td><td>0.00</td><td>0.00</td><td>0.11</td><td>0.03</td><td></td></tr><tr><td>11 months stall</td><td>0.01</td><td>0.13</td><td>0.00</td><td>0.00</td><td>0.13</td><td>0.06</td><td>0.70</td></tr><tr><td>14 months enclosure</td><td>0.00</td><td>0.17</td><td>0.00</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.26</td></tr><tr><td>14 months stall</td><td>0.00</td><td>0.15</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.15</td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td></tr></tbody></table><table><thead><tr><th rowspan="2">Mandible</th><th>8 to 10</th><th>16 to 18</th><th rowspan="2">6 months<br>enclosure</th><th rowspan="2">6 months<br>stall</th><th rowspan="2">8 months<br>enclosure</th><th rowspan="2">8 months<br>stall</th><th>11</th></tr><tr><th>months wild<br>caught</th><th>months wild<br>caught</th><th>months<br>enclosure</th></tr></thead><tbody><tr><td>16-18 months wild caught</td><td>0.02</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.55</td><td>0.01</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.74</td><td>0.01</td><td>0.89</td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.27</td><td>0.09</td><td>0.18</td><td>0.22</td><td></td><td></td><td></td></tr></tbody></table>
<table><tr><td>8 months stall</td><td>0.38</td><td>0.04</td><td>0.25</td><td>0.39</td><td>0.73</td><td></td></tr><tr><td>11 months enclosure</td><td>0.00</td><td>0.42</td><td>0.00</td><td>0.00</td><td>0.07</td><td>0.03</td></tr><tr><td>11 months stall</td><td>0.00</td><td>0.30</td><td>0.00</td><td>0.00</td><td>0.04</td><td>0.02</td><td>0.95</td></tr><tr><td>14 months enclosure</td><td>0.00</td><td>0.21</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.33</td></tr><tr><td>14 months stall</td><td>0.00</td><td>0.12</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.16</td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td></tr></table>

<table><thead><tr><th>Calcaneus</th><th>8 to 10<br>months wild<br>caught</th><th>16 to 18<br>months wild<br>caught</th><th>6 months<br>enclosure</th><th>6 months<br>stall</th><th>8 months<br>enclosure</th><th>8 months<br>stall</th><th>11<br>months<br>enclosure</th></tr></thead><tbody><tr><td>16-18 months wild caught</td><td>0.01</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.49</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.35</td><td>0.01</td><td>0.89</td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.13</td><td>0.09</td><td>0.18</td><td>0.21</td><td></td><td></td><td></td></tr><tr><td>8 months stall</td><td>0.22</td><td>0.03</td><td>0.31</td><td>0.68</td><td>0.74</td><td></td><td></td></tr><tr><td>11 months enclosure</td><td>0.01</td><td>0.35</td><td>0.01</td><td>0.03</td><td>0.37</td><td>0.09</td><td></td></tr><tr><td>11 months stall</td><td>0.00</td><td>0.20</td><td>0.00</td><td>0.02</td><td>0.13</td><td>0.07</td><td>0.56</td></tr><tr><td>14 months enclosure</td><td>0.00</td><td>0.25</td><td>0.00</td><td>0.00</td><td>0.02</td><td>0.01</td><td>0.29</td></tr><tr><td>14 months stall</td><td>0.00</td><td>0.09</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.08</td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.02</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.01</td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.01</td></tr></tbody></table>
5 b. ANOVA *p*-values of the centroid size (CS) between all groups computed for the cranium,

6 mandible, and calcaneus and of body mass (kg).

<table><thead><tr><th rowspan="2">Cranium</th><th>8 to 10</th><th>16 to 18</th><th rowspan="2">6 months<br>enclosure</th><th rowspan="2">6 months<br>stall</th><th rowspan="2">8 months<br>enclosure</th><th rowspan="2">8 months<br>stall</th><th rowspan="2">11<br>months<br>enclosure</th><th rowspan="2">months<br>stall</th></tr><tr><th>months wild<br>caught</th><th>months wild<br>caught</th></tr></thead><tbody><tr><td colspan="9" style="text-align: left; font-weight: bold;">16-18 months wild</td></tr><tr><td>caught</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.75</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.67</td><td>0.00</td><td>0.86</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.37</td><td>0.03</td><td>0.22</td><td>0.14</td><td></td><td></td><td></td><td></td></tr><tr><td>8 months stall</td><td>0.60</td><td>0.00</td><td>0.37</td><td>0.30</td><td>0.70</td><td></td><td></td><td></td></tr><tr><td>11 months enclosure</td><td>0.01</td><td>0.42</td><td>0.01</td><td>0.00</td><td>0.09</td><td>0.04</td><td></td><td></td></tr><tr><td>11 months stall</td><td>0.02</td><td>0.37</td><td>0.00</td><td>0.00</td><td>0.11</td><td>0.04</td><td>0.96</td><td></td></tr><tr><td>14 months enclosure</td><td>0.00</td><td>0.71</td><td>0.00</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.20</td><td></td></tr><tr><td>14 months stall</td><td>0.00</td><td>0.60</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.14</td><td></td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.04</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td></td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.02</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td></td></tr><tr><td colspan="9" style="text-align: left; font-weight: bold;">Mandible</td></tr><tr><td> caught</td><td>0.02</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.54</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.58</td><td>0.01</td><td>0.94</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.39</td><td>0.09</td><td>0.15</td><td>0.18</td><td></td><td></td><td></td><td></td></tr></tbody></table>
<table><tr><td>8 months stall</td><td>0.61</td><td>0.05</td><td>0.25</td><td>0.29</td><td>0.74</td><td></td><td></td></tr><tr><td>11 months enclosure</td><td>0.01</td><td>0.79</td><td>0.00</td><td>0.00</td><td>0.09</td><td>0.03</td><td></td></tr><tr><td>11 months stall</td><td>0.02</td><td>0.81</td><td>0.00</td><td>0.00</td><td>0.08</td><td>0.03</td><td>0.99</td></tr><tr><td>14 months enclosure</td><td>0.00</td><td>0.49</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.25</td></tr><tr><td>14 months stall</td><td>0.00</td><td>0.33</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.15</td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.02</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td></tr></table>

<table><thead><tr><th></th><th>8 to 10<br>months wild<br>caught</th><th>16 to 18<br>months wild<br>caught</th><th>6 months<br>enclosure</th><th>6 months<br>stall</th><th>8 months<br>enclosure</th><th>8 months<br>stall</th><th>11<br>months<br>enclosure</th><th>months<br>wild<br>stall</th></tr></thead><tbody><tr><td>Calcaneus</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>16-18 months wild<br>caught</td><td>0.01</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.93</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.78</td><td>0.01</td><td>0.74</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.18</td><td>0.12</td><td>0.15</td><td>0.27</td><td></td><td></td><td></td><td></td></tr><tr><td>8 months stall</td><td>0.32</td><td>0.06</td><td>0.29</td><td>0.50</td><td>0.70</td><td></td><td></td><td></td></tr><tr><td>11 months enclosure</td><td>0.01</td><td>0.62</td><td>0.01</td><td>0.02</td><td>0.25</td><td>0.12</td><td></td><td></td></tr><tr><td>11 months stall</td><td>0.00</td><td>0.95</td><td>0.00</td><td>0.00</td><td>0.10</td><td>0.03</td><td>0.65</td><td></td></tr><tr><td>14 months enclosure</td><td>0.00</td><td>0.63</td><td>0.00</td><td>0.00</td><td>0.02</td><td>0.01</td><td>0.28</td><td></td></tr><tr><td>14 months stall</td><td>0.00</td><td>0.28</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.09</td><td></td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.12</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.02</td><td></td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.08</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.01</td><td></td></tr></tbody></table>

<table><thead><tr><th></th><th>8 to 10<br>months wild<br>caught</th><th>16 to 18<br>months wild<br>caught</th><th>6 months<br>enclosure</th><th>6 months<br>stall</th><th>8 months<br>enclosure</th><th>8 months<br>stall</th><th>11<br>months<br>enclosure</th><th>months<br>wild<br>stall</th></tr></thead><tbody><tr><td>Mass</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>
<table><tbody><tr><td>16-18 months wild</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>caught</td><td>0.01</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months enclosure</td><td>0.50</td><td>0.00</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>6 months stall</td><td>0.45</td><td>0.00</td><td>0.91</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>8 months enclosure</td><td>0.81</td><td>0.01</td><td>0.33</td><td>0.28</td><td></td><td></td><td></td><td></td></tr><tr><td>8 months stall</td><td>0.64</td><td>0.02</td><td>0.19</td><td>0.18</td><td>0.80</td><td></td><td></td><td></td></tr><tr><td>11 months enclosure</td><td>0.20</td><td>0.12</td><td>0.04</td><td>0.03</td><td>0.27</td><td>0.39</td><td></td><td></td></tr><tr><td>11 months stall</td><td>0.16</td><td>0.22</td><td>0.02</td><td>0.01</td><td>0.19</td><td>0.26</td><td>0.82</td><td></td></tr><tr><td>14 months enclosure</td><td>0.02</td><td>0.60</td><td>0.00</td><td>0.00</td><td>0.02</td><td>0.05</td><td>0.26</td><td></td></tr><tr><td>14 months stall</td><td>0.01</td><td>1.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.01</td><td>0.09</td><td></td></tr><tr><td>20 months enclosure</td><td>0.00</td><td>0.06</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td></td></tr><tr><td>20 months stall</td><td>0.00</td><td>0.01</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td>0.00</td><td></td></tr></tbody></table>

7

8

9