Historical Biology

An International Journal of Paleobiology

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Human tooth development, tooth length and

eruption; a study of British archaeological
dentitions

Helen M. Liversidge & Theya Molleson

To cite this article: Helen M. Liversidge & Theya Molleson (2017): Human tooth development,
tooth length and eruption; a study of British archaeological dentitions, Historical Biology, DOI:
10.1080/08912963.2017.1305375

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Historical Biology, 2017
[Link]

Human tooth development, tooth length and eruption; a study of British

archaeological dentitions
Helen M. Liversidgea and Theya Mollesonb
a
Queen Mary University of London, Dental Institute, London, UK; bDepartment of Earth Sciences (Palaeontology), Palaeontology, Natural History
Museum, London, UK

ABSTRACT ARTICLE HISTORY

Human teeth erupt during root growth but few studies report the relationship between fractions of root Received 8 March 2017
development and eruption levels. The aim of this study was to assess root stages of deciduous and early Accepted 8 March 2017
erupting permanent teeth (maxillary and mandibular central incisors and first molars) at eruption levels
KEYWORDS
and relate this to root fraction and tooth length. The sample consisted of 620 modern human skeletal Human; tooth eruption;
remains with developing teeth. Tooth stage (based on Moorrees crown and root stages) and eruption root formation stage; tooth
levels of all developing teeth were assessed where possible. Tooth length of isolated teeth was measured. length
The distribution of root stage at eruption levels was calculated. Results showed that root stage at alveolar
eruption was less variable than at partial eruption. Most teeth (72% of 138) at alveolar eruption were at
root a quarter (R¼) whereas teeth at partial eruption were at R¼ or root half (R½) (38 and 50% of 128
respectively). These findings suggest that the active phase of eruption is probably a rapid process and
occurs during the first half of root growth.

Human teeth develop within the alveolar jaw bones. The crown in this study. Teeth assessed were all deciduous teeth and perma-
is the first part of the tooth to form and once initial root forma- nent central incisors and first molars both maxillary and man-
tion occurs, the separate process of eruption begins. The active dibular. The sample consisted of fragmentary remains of 620
phase of eruption is characterised by an increase in root length individuals from Abingdon (Allen 1989), Cannington (Rahtz
as well as bodily movement of the tooth towards the oral cavity. et al. 2000), Newark Bay (Brothwell et al. 1986), Poundbury
The incisal edge or occlusal surface of the tooth moves through (Farwell and Molleson 1993), Spitalfields (Molleson and Cox
the alveolar bone margin, penetrates the oral mucosa and comes 1993) and Winchester (Biddle and Quirk 1962; Molleson et al.
into contact with the opposing tooth to be in a functional posi- 2016) detailed in Table 1. These sites range in time from Roman-
tion. This continuous process can be divided into discrete stages British to mid-nineteenth century.
relative to the alveolar bone margin and occlusal levels. There
is a vast literature documenting the age of clinical eruption of
Method
individual teeth, however, few studies report on the variation of
root stage at alveolar or clinical eruption and an understanding of All teeth were assessed for eruption level, tooth stage, and tooth
this will help to recognise what is normal and to predict eruption length was measured where possible. Radiographs of some spec-
level of isolated teeth. Such information can throw light on life imens were available and some developing teeth that were not
history events of past populations. Root length and chronological loose and able to inspected visually, were assessed from radi-
age of permanent molars at clinical eruption differs in modern ographs. Eruption level of each tooth was assessed where the
humans, early hominins and chimpanzees. Root length per se in alveolar margin was undamaged. Teeth with cusp tip(s) or incisal
an evolutionary context is a tantalising issue that needs an estab- edge below the alveolar margin were classified as un-erupted. If
lished base reference and our study focuses on root formation the cusp tip(s) or incisal edge was at or just through the bone
stage and eruption levels. viewed from the lateral aspect, the tooth was classified as at
The aim of this study was to describe the associations between alveolar eruption (AE). If cusp tip(s) or incisal edge was at the
root stages, tooth length and eruption levels of deciduous and maximum bulbosity of adjacent tooth (if present) the tooth was
permanent central incisors and first molars in a skeletal remains. classified as partial eruption (PE). Teeth in occlusion were con-
sidered fully erupted. Examples of eruption levels are illustrated
in Figure 1.
Sample
If the tooth was isolated or the developing edge was visi-
Skeletal remains with developing teeth, from the British Isles ble, crown or root stage was assessed and tooth length (from
housed at the Natural History Museum, London were included incisal edge/cusp tips to forming edge of crown or root or apex)

CONTACT  Helen M. Liversidge  [Link]@[Link]

© 2017 Informa UK Limited, trading as Taylor & Francis Group
2   H. M. LIVERSIDGE AND T. MOLLESON

Table 1. The number of individuals (N), the sites and approximate date of the sam-
ple of human skeletal remains used in this study.
Site Approximate date N
Cannington Early post-Roman 23
Newark Bay Late Roman 86
Poundbury Late Roman 195
Winchester Green Medieval 47
Abingdon Medieval seventeenth century 115
Spitalfields 1729–1853 154
Total 620

Table 2. Crown and root stage descriptions and abbreviations.

Stage Abbreviation Description
Crown three quarters C¾ Crown height > half the unit
Crown complete Cc Presence of mature enamel and
initial root visible on both
approximal margins
Initial root Ri Anatomically crown complete with
mature enamel margin and root
present less than half the unit
Root cleft Rcl Root cleft visible
Root quarter R¼ Root length at least 3 units
Root half R½ Root length at least 4 units
Root three quarters R¾ Root length at least 5 units
Root length complete Rc Root edges beginning to con-
verge/open apex
Apex half closed A½ Small apical foramen
Apex closed Ac Very small or no apical foramen

Notes: Fractions are relative to the unit of half the crown height on the approximal
surface (see Figure 1). Root cleft stage is only appropriate for molars. Figure 2. Crown and root fractions of maxillary permanent incisor and first molar
(above) and deciduous incisor and first molar (below).
Notes: Arrow on left of crown indicates crown height complete (Cc) = 2 units (labelled 2), arrow
measured in the long axis using Vernier callipers. Crown and on right of tooth indicates root half (R½) = 4 units (labelled 4) when crown height equals root
length. Tooth length measured from incisal edge to root apex/apices.
root stages were modified from radiographic tooth stages of
Moorrees et al. (1963) and are listed in Table 2. Data for crown
three quarter formed (C¾) to apex closed (Ac) are presented in

Figure 1.  Examples of alveolar eruption of mandibular deciduous central incisors (a, b, c), first molar (d), partial eruption of mandibular central incisor and alveolar
eruption of lateral incisor (e), partial eruption of mandibular incisors and first molar, canine un-erupted (f) and canine partially erupted (arrowed) with lateral incisor and
first molar fully erupted (g).
 HISTORICAL BIOLOGY   3

Table 3. Numbers of teeth assessed for eruption and tooth length. measure of rater agreement suitable for categorical data) was cal-
Eruption Tooth length culated (Cohen 1960) for eruption level and tooth stage reliability.
Tooth Maxillary Mandibular Maxillary Mandibular
Reliability of tooth length measurement was assessed by compar-
ing the mean difference between readings using a paired t-test.
Deciduous
The number of individuals at each tooth stage and eruption
 Central incisor 123 150 156 165
 Lateral incisor 123 164 142 176 level was calculated for each tooth type. Distribution of tooth
 Canine 108 145 123 157 stages at eruption level was calculated and deciduous and perma-
  First molar 129 161 156 162 nent dentitions compared. Distribution of eruption level at R¼
 Second molar 136 160 155 139
Total 619 780 732 799 and R½ was calculated and deciduous and permanent dentitions
Permanent compared. Tooth length of teeth at alveolar eruption and partial
 Central incisor  67  67  69  61 eruption was calculated for each tooth type. Tooth length at each
  First molar  96  91  98  85 tooth stage was calculated for each tooth type.
Total 163 158 167 146 Comparisons between archaeological sites could not be made
due to the fragmentary nature of the material and small number
of actively erupting teeth.
this study. The unit used to define tooth stages was half the crown
height, assessed from the incisal edge/cusp tip(s) to the enamel
margin on the approximal surface (Figure 2). Root fractions were Results
defined in terms of half the crown height, but several stages were Reliability of tooth stage and eruption levels for 28 teeth was
defined morphologically with reference to the crown margin or Kappa 0.771 and 0.892 respectively showing good agreement.
root apical edges and apex width. Crown complete (Cc) stage was The mean difference between duplicate tooth length measures
defined as the presence of mature enamel and initial root visible was −0.058 (SD 0.36) and this is not statistically significant from
on the approximal margins. Half of this approximal crown height zero. The total number of teeth assessed for eruption level and
(C½) was used as a unit of length to define subsequent stages. tooth stage was 1720 and 1844 teeth were measured (Table 3).
Initial root stage (Ri) was defined as anatomically crown com- Only a small proportion (16%) of teeth in this sample were
plete (mature enamel margin on all four aspects of crown) with actively erupting (138 at AE and 128 at PE).
root present less than half the unit. Three units was root quarter Results of root stage at eruption level are presented in terms
(R¼), four units root half (R½) and 5 units root three quarters of the eruption level and root stage. Firstly, the distribution of
(R¾). Once the root edges begin to converge, a tooth was defined root stage at active eruption levels (AE and PE) is shown in
as root length complete (Rc). A small apical foramen easily seen Figure 3. At alveolar eruption the modal root formation stage
with the naked eye was defined as apex half closed (A½) and a was R¼ (91/121 deciduous teeth and 9/17 permanent teeth). At
very small or no foramen visible was defined as apex closed (Ac). partial eruption the modal root formation stage was R½. This
Tooth length was measured using Vernier callipers in the long pattern was similar in deciduous and permanent teeth and data
axis of the tooth from the incisal edge/cusp tip(s) to the crown were pooled. Root stage at eruption level for individual teeth
or root developing edge. A number of mature erupted teeth were is detailed in Table 4. Data for one tooth type (the mandibular
also measured including some with small occlusal wear facets. deciduous first molar) are illustrated in Figure 4, showing that
Reliability of eruption level, tooth stage and tooth length was early root stages were observed at all eruption levels.
assessed from duplicate readings of 28 teeth. Cohen’s Kappa (a

Figure 3. Distribution of root stage at alveolar eruption (AE) and partial eruption (PE) for all teeth combined.
Note: Root stages Ri – initial root, Rcl – root cleft, R¼ – root quarter, R½ – root half, R¾ – root three quarters, Rc – root complete.
4   H. M. LIVERSIDGE AND T. MOLLESON

Table 4. Counts of tooth formation stage for each category of eruption level (de-
ciduous incisors, canines, deciduous molars, permanent central incisor and first
molar). Table 4. (Continued)
Tooth Eruption levels Eruption levels Tooth Eruption levels Eruption levels
Stage UE AE PE FE N UE AE PE FE N Stage UE AE PE FE N UE AE PE FE N
Maxillary deciduous central Mandibular deciduous central Maxillary permanent first molar Mandibular permanent first molar
incisor incisor C¾ 25 25 21 21
C¾ 19 19 14 14 Cc 11 11 8 8
Cc 10 10 9 9 Ri 8 8 3 3
Ri 7 7 5 1 6 Rcl 1 1 2 1 3
R¼ 3 4 2 9 3 10 6 19 R¼ 7 1 1 9 4 3 3 1 11
R½ 1 2 3 6 3 5 5 13 R½ 2 3 2 7 2 2 3 7
R¾ 13 13 1 6 7 R¾ 1 12 13 2 12 14
Rc 7 7 16 16 Rc 2 2 5 5
A½ 12 12 9 9 A½ 4 4 5 5
Ac 40 40 57 57 Ac 16 16 14 14
Total 39 5 4 75 123 31 14 12 93 150 Total 52 3 5 36 96 38 6 7 40 91
Maxillary deciduous lateral incisor Mandibular deciduous lateral Abbreviations: Eruption levels UE – un-erupted, AE – alveolar eruption, PE – partial
incisor eruption, FE – fully erupted, tooth stages see Table 1. Counts of tooth formation
C¾ 22 22 22 22 stage for each category of eruption level.
Cc 8 8 10 10
Ri 2 2 8 3 11
R¼ 5 4 1 10 9 8 3 20
R½ 2 2 4 8 1 3 9 4 17
R¾ 17 17 3 16 19
Rc 10 10 11 11
A½ 11 11 9 9
Ac 35 35 45 45
Total 37 6 3 77 123 50 14 15 85 164
Maxillary deciduous canine Mandibular deciduous canine
C¾ 15 15 20 20
Cc 3 3 3 3
Ri 6 6 5 2 7
R¼ 1 12 1 1 15 6 14 5 25
R½ 1 6 2 9 1 9 11 21
R¾ 2 18 20 23 23
Rc 11 11 16 16
A½ 2 2 5 5
Ac 27 27 25 25
Total 25 13 9 61 108 34 17 14 80 145
Maxillary deciduous first molar Mandibular deciduous first molar
C¾ 16 16 18 18
Cc 8 8 9 9
Ri 6 6 9 9
Rcl 3 3 4 3 7
R¼ 2 5 11 18 4 11 9 2 26
R½ 5 9 14 2 7 16 25
R¾ 2 11 13 13 13
Rc 14 14 15 15 Figure 4. Frequency of crown and root stages of the mandibular deciduous first
A½ 12 12 8 8 molar at eruption levels (N = 161).
Ac 25 25 31 31 Notes: UE – un-erupted, AE – alveolar eruption, PE – partial eruption, FE – full eruption. Tooth
stages C¾ – crown three quarters, Cc – crown complete, Ri – initial root, Rcl – root cleft, R¼
Total 35 5 18 71 129 44 16 16 85 161
– root quarter, R½ – root half, R¾ – root three quarters, Rc – root complete, A½ – apex half,
Mandibular deciduous second Ac – apex closed.
Maxillary deciduous second molar molar
C¾ 22 22 29 29
Cc 13 13 14 14
Ri 9 9 9 9 Secondly, results expressed by tooth stage show that almost all
Rcl 4 2 6 3 1 4 teeth in late crown stages and initial root stage were un-erupted.
R¼ 4 10 2 16 4 13 4 3 24
R½ 2 6 7 15 3 5 10 18 At R¼, the distribution of eruption level differed between decid-
R¾ 2 19 21 1 21 22 uous and permanent teeth (see Figure 5), however, the sample
Rc 10 10 1 12 13 size for permanent teeth was small. The modal eruption level at
A½ 4 4 9 9
Ac 20 20 18 18 R¼ for deciduous teeth was alveolar eruption and for the perma-
Total 52 14 10 60 136 59 17 11 73 160 nent teeth was un-erupted. At R½, the modal eruption level was
Maxillary permanent central Mandibular permanent central fully erupted and by R¾ most teeth had fully erupted.
incisor incisor
C¾ 19 19 14 14 Tooth length at alveolar and partial eruption levels are shown
Cc 6 6 6 6 in Table 5 and tooth length at each tooth stage is detailed in
Ri 4 4 4 4 Table 6. Figure 5 shows length, stage and level of eruption for
R¼ 4 3 7 2 2 1 5
R½ 1 1 2 4 2 2 3 7
the mandibular deciduous canine and permanent first molar.
R¾ 1 1 1 3 4
Rc 1 1 4 4
A½ 7 7 3 3 Discussion
Ac 18 18 20 20
Total 34 3 1 29 67 26 5 3 33 67 Tooth eruption is a complex process involving the tooth organ
(Continued) and the surrounding alveolar bone. Root formation and eruption
 eruption.
permanent teeth at root quarter (middle) and all teeth at root half.

are thought to be independent processes involving an increase in

Figure 5. Distribution of deciduous teeth at root quarter at eruption levels (top),

remodelling (Kardos 1996) although the presence of the dental

root length, cementum formation, formation of the periodontal
ligament and resorption of the alveolar bone allowing a pathway

alveolar eruption of some teeth (Garn et al. 1958; Haavikko 1970;

follicle, the ligament and bone are thought to be more important
to the oral cavity. Theories of tooth eruption include root growth,
vasculature, the developing periodontal ligament and bone

than the developing tooth. Several studies report on the age of

Note: Eruption levels UE – un-erupted, AE – alveolar eruption, PE – partial eruption, FE – full
Table 5. Tooth length in mm at alveolar eruption and partial eruption levels.
Tooth N Mean SD Min Max N Mean SD Min Max
Maxillary Alveolar eruption Partial eruption
 di1 5 8.60 1.14 7.40 10.10 4 9.64 0.77 8.90 10.70
 di2 6 9.01 1.16 7.20 10.30 2 8.55 0.78 8.00 9.10
 dc 10 9.61 0.73 8.90 11.15 9 12.41 1.53 9.90 14.80
 dm1 5 8.76 0.15 8.60 9.00 18 9.77 0.95 8.27 12.50
 dm2 12 9.92 0.88 7.77 11.06 10 11.61 0.90 10.50 13.70
Mandibular
 di1 13 8.49 0.70 7.40 9.30 12 9.26 1.35 7.30 9.26
 di2 12 9.09 1.27 7.00 10.70 15 10.67 1.37 8.67 12.92
 dc 16 8.89 0.88 7.90 11.10 13 11.14 1.33 8.53 12.32
 dm1 13 8.53 0.66 7.60 9.60 14 9.48 0.60 8.30 10.67
 dm2 15 9.94 1.19 7.54 12.81 10 11.16 1.24 9.63 13.30
Maxillary
 I1 3 16.41 0.99 15.27 17.10 1 16.48
 M1 3 14.07 1.87 11.91 15.29 5 14.76 2.41 11.29 17.60
Mandibular
 I1 4 15.53 1.47 13.60 16.98 3 16.40 0.52 16.06 17.00
 M1 6 13.07 1.41 10.62 14.44 7 13.31 1.28 11.40 14.69
HISTORICAL BIOLOGY 

Abbreviations: N – number of teeth, SD – standard deviation, Min – minimum, Max – maximum, di1 – deciduous central incisor, di2 – deciduous lateral incisor, dc – deciduous canine, dm1 – deciduous first molar, dm2 – deciduous
second molar, I1 – permanent central incisor, M1 – permanent first molar.
 5
6   H. M. LIVERSIDGE AND T. MOLLESON

Table 6. Tooth length in mm at each tooth stage for deciduous incisors, canines, deciduous molars, permanent central incisor and first molar.
Stage N Mean SD Min Max N Mean SD Min Max
Maxillary deciduous central incisor Mandibular deciduous central incisor
C¾ 17 6.15 0.61 5.00 7.30 17 5.27 0.39 4.50 5.80
Cc 11 6.79 0.33 6.40 7.40 9 5.84 0.36 5.20 6.40
Ri 9 7.93 0.65 6.90 8.90 7 7.32 1.04 6.30 9.40
R¼ 22 8.79 0.77 7.40 9.90 22 8.12 0.82 6.30 9.30
R½ 13 11.06 1.06 9.30 12.40 17 10.16 0.94 8.57 11.90
R¾ 19 13.77 1.00 12.36 16.00 7 11.31 0.48 10.65 12.10
Rc 10 14.31 0.93 13.17 15.51 18 12.29 0.62 11.06 13.23
A½ 13 15.74 1.23 14.20 18.70 11 12.89 0.92 11.41 14.88
Ac 42 15.84 1.40 13.54 21.68 57 13.25 0.93 10.26 15.11
Maxillary deciduous lateral incisor Mandibular deciduous lateral incisor
C¾ 20 5.50 0.50 4.80 6.70 20 5.41 0.65 4.10 6.30
Cc 11 6.37 0.68 5.20 7.51 9 6.67 0.72 5.00 7.40
Ri 5 7.22 0.42 6.80 7.80 13 7.28 0.59 6.40 8.50
R¼ 16 8.21 0.77 7.20 9.80 23 8.67 0.98 7.29 10.46
R½ 14 10.64 0.78 9.10 11.86 19 10.76 0.83 9.10 12.68
R¾ 24 12.81 1.23 10.60 15.10 22 12.90 0.95 11.04 14.90
Rc 10 14.40 1.34 12.70 16.58 12 14.69 1.14 13.10 16.28
A½ 10 15.80 0.91 14.20 17.38 10 15.00 0.59 14.04 15.90
Ac 32 15.42 1.06 13.70 17.34 48 15.21 1.35 10.60 17.84
Maxillary deciduous canine Mandibular deciduous canine
C¾ 19 6.77 0.68 4.40 7.73 21 6.56 0.68 4.10 7.30
Cc 2 7.85 0.64 7.40 8.30 5 7.79 0.52 6.92 8.22
Ri 7 8.09 0.51 7.30 8.72 8 8.25 1.11 7.00 10.60
R¼ 22 9.41 0.85 8.28 11.45 28 9.10 0.79 8.10 11.18
R½ 11 12.10 1.42 9.90 14.90 22 12.40 1.20 9.50 15.50
R¾ 20 15.41 1.00 13.50 16.91 25 14.84 1.08 12.56 16.93
Rc 12 16.94 1.36 14.80 18.68 17 16.41 1.10 14.40 18.58
A½ 3 17.49 0.32 17.13 17.74 5 16.19 1.04 14.60 17.34
Ac 27 16.96 1.25 14.90 19.35 26 16.45 1.19 14.51 19.09
Maxillary deciduous first molar Mandibular deciduous first molar
C¾ 19 5.37 0.52 4.40 6.49 19 5.35 0.66 4.50 6.70
Cc 12 6.20 0.50 5.60 7.50 11 6.32 0.40 5.90 7.02
Ri 13 6.91 0.67 6.10 8.59 11 6.72 0.53 5.60 7.46
Rcl 6 7.64 0.85 6.70 8.71 8 7.73 0.78 6.30 8.66
R¼ 22 8.95 0.72 7.10 10.30 25 8.86 0.75 7.20 10.02
R½ 20 10.49 1.15 8.62 13.10 24 10.94 1.25 9.29 13.90
R¾ 12 11.98 0.88 10.60 13.40 13 12.38 0.70 11.47 13.50
Rc 13 12.94 1.03 11.50 14.99 16 13.16 0.87 11.83 14.91
A½ 12 13.14 0.58 12.10 13.90 6 13.55 0.62 12.80 14.40
Ac 27 13.96 0.89 12.50 15.90 29 14.30 1.08 12.70 17.18
Maxillary deciduous second molar Mandibular deciduous second molar
C¾ 25 5.75 0.62 4.50 6.79 22 5.59 0.45 4.70 6.50
Cc 15 6.75 0.72 5.37 7.70 15 6.49 0.38 5.80 7.10
Ri 13 7.35 0.82 6.20 8.94 12 7.77 0.75 6.20 8.69
Rcl 8 8.05 0.80 7.03 9.40 4 7.73 0.71 7.20 8.78
R¼ 20 9.48 1.04 7.60 10.91 21 9.88 1.02 7.90 11.70
R½ 15 11.63 0.67 10.48 12.57 14 11.54 0.64 10.58 12.81
R¾ 19 12.91 1.03 11.31 15.00 19 13.03 0.83 11.22 14.31
Rc 10 13.64 0.69 12.70 14.76 10 13.84 0.62 13.00 14.78
A½ 5 14.73 1.34 13.60 16.99 7 14.75 1.16 12.80 16.30
Ac 25 15.46 1.21 13.18 18.16 15 15.27 0.99 13.47 16.80
Maxillary permanent central incisor Mandibular permanent central incisor
C¾ 16 9.30 0.97 7.80 11.12 13 7.87 0.96 6.50 10.29
Cc 7 11.00 0.65 9.94 11.97 7 9.82 0.84 8.00 10.45
Ri 5 12.39 1.29 11.20 13.87 3 11.79 1.19 10.42 12.46
R¼ 11 15.01 1.39 12.95 17.10 6 14.22 1.16 13.11 16.14
R½ 3 17.63 1.26 16.48 18.97 7 16.61 0.93 15.24 18.22
R¾ 3 20.48 0.70 19.72 21.10 4 18.40 1.10 16.98 19.50
Rc 2 23.88 0.49 23.54 24.23 3 19.83 0.79 19.34 20.74
A½ 7 23.80 1.65 21.95 25.89 3 20.15 1.24 19.05 21.50
Ac 15 22.85 1.54 20.48 24.84 15 20.77 1.56 18.81 24.81
Maxillary permanent first molar Mandibular permanent first molar
C¾ 24 7.09 0.54 5.96 8.05 19 6.63 0.61 5.20 7.58
Cc 9 7.97 0.69 7.05 9.48 6 7.79 0.86 6.88 9.40
Ri 12 9.33 1.01 7.20 11.10 3 9.04 0.33 8.73 9.39
Rcl 3 12.48 1.42 11.40 14.10 4 10.65 0.97 9.20 11.30
R¼ 14 11.64 1.91 9.33 16.00 14 12.40 1.36 9.87 14.30
R½ 10 15.19 0.88 13.67 16.73 6 14.46 1.19 12.87 16.55
R¾ 11 17.43 1.63 14.30 20.73 13 16.57 1.61 14.52 20.35
Rc 2 20.28 1.67 19.10 21.45 5 18.70 1.52 17.55 20.72
A½ 4 20.13 0.91 19.02 21.15 7 19.30 0.73 18.14 20.11
Ac 9 19.51 1.42 17.76 22.23 8 19.38 1.43 17.18 21.59

Abbreviations: see Table 1. N – number of teeth, SD – standard deviation, Min – minimum, Max – maximum, tooth stages see Table 1.
 HISTORICAL BIOLOGY   7

Figure 6. Tooth length in mm of the mandibular deciduous canine (left) and mandibular permanent first molar (right) plotted against tooth stage showing eruption levels
(triangle apex up un-erupted, grey diamond alveolar eruption, circle partial eruption, triangle apex down full eruption).
Notes: Tooth stages C¾ – crown three quarters, Cc – crown complete, Ri – initial root, Rcl – root cleft, R¼ – root quarter, R½ – root half, R¾ – root three quarters, Rc – root complete, A½ – apex
half, Ac – apex closed.

Liversidge and Molleson 2004) and there is a vast literature on Our findings show differences between the deciduous and
the age of clinical tooth eruption into the oral cavity. Only one early erupting permanent teeth. Deciduous root formation is
study describes root stage at clinical tooth eruption (Grøn 1962). probably uniformly rapid (Schour and Massler 1940; Stack 1967;
This study is part of an ongoing project on root stage and Deutsch et al. 1985) but root formation in permanent teeth is
eruption by the first author. Results from this study show that more complex with a root spurt. A recent study has shown that
there is variation in root stage at both alveolar eruption and par- the peak root extension rate is earlier than the time of clinical
tial eruption levels. These two eruption levels represent active eruption in some permanent teeth with the exception of the early
eruption when most teeth were at root quarter or root half stage, erupting central incisor (Dean 2007; Dean and Cole 2013). Early
suggesting that active eruption begins during the first quarter of erupting permanent teeth erupt in to the oral cavity with less root
root growth and teeth are fully erupted by the third quarter of than later erupting teeth (Grøn 1962). Our sample of actively
root formation. This suggest that the active phase of eruption erupting permanent teeth was small and showed some variation
might be a rapid process. in root stage at both alveolar and partial eruption levels.
Teeth vary in length and dividing tooth formation into dis-
crete stages of unequal duration has limitations. Figure 6 shows
Conclusions
that tooth length varies somewhat between stages and there is
overlap in tooth length at R¼ and R½ and alveolar and partial Knowing the eruption level of a developing tooth tells us about
eruption levels. Despite this, it is possible that once a critical the establishment of the masticatory system, as well as cranio-fa-
tooth length is reached the tooth can withstand the forces of cial growth and development. The timing of tooth eruption is
chewing and erupt. The critical tooth length (minimum tooth correlated to life history variables and being able to predict if an
length sufficient to withstand masticatory forces) for the man- isolated tooth is functional can be valuable to infer life history
dibular deciduous canine and mandibular permanent first molar variables. This study shows an association between root stage,
might be around 9 and 13 mm respectively (see Figure 6). root length and eruption level. Active eruption occurs during the
Another limitation of root fractions is that discriminating first half of root growth and most teeth at alveolar and partial
between tooth stages is difficult when a tooth is deemed to be eruption levels were at R¼ and R½ respectively, although some
between stages. A tooth may have just entered the stage while variation in tooth formation stage exists at both eruption levels.
others have been in the stage for some time. The duration of
tooth stages is not well documented and cannot be assessed from
Acknowledgements
a cross-sectional study such as this. Teeth with roots between
root fractions or eruption levels were categorised into the earlier We thank the editors of this special volume to celebrate Percy Butler for
stage. Our definition of root fractions is an attempt to reduce the inviting us to contribute. We are also grateful to Robert Kruszynski for help
accessing the collections.
subjectivity of fraction estimation.
8   H. M. LIVERSIDGE AND T. MOLLESON

Disclosure statement Garn SM, Lewis AB, Koski K, Polacheck DL. 1958. The sex difference in
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