Journal of Orofacial Orthopedics

https://doi.org/10.1007/s00056-024-00532-3

ORIGINAL ARTICLE

Frequency and variability of nonmetric dental crown traits of primary

and permanent molars in a group of orthodontic patients
Ariane Beatriz Blancato1 · Eva Paddenberg-Schubert2 · Peter Proff2 · Maria Angélica Hueb de
Menezes-Oliveira1 · Svenja Beisel-Memmert3 · Flares Baratto-Filho4,5 · Carsten Lippold6 · Christian Kirschneck3 ·
Erika Calvano Küchler3 · Cesar Penazzo Lepri1

Received: 18 October 2023 / Accepted: 18 March 2024

© The Author(s) 2024, corrected publication 2024

Abstract
Background The present study aimed to assess the frequency and variation of 13 nonmetric dental crown traits (NDCT)
in permanent and primary molars in German orthodontic patients.
Methods Dental records from orthodontic patients were screened and evaluated. First and second permanent and primary
upper and lower molars (from left and right sides) were assessed. Teeth with cavitated dental caries, occlusal wear,
restorations and obvious dental deformities were not evaluated. The NDCT for permanent molars were identified and scored
according to the odontoscopic system developed by Arizona State University Dental Anthropology System (ASUDAS).
The NDCT for primary molars were identified and scored according to ASUDAS, Hanihara’s method and Sciulli’s method.
The χ2 test was used to investigate side preference and sexual dimorphism at a significance level of p ≤ 0.050.
Results A total of 163 orthodontic patients (82 males and 81 females) aged 8–14 years were included. A sexual dimorphism
was observed for the hypocone in first upper permanent molar (p = 0.041). The protostylid was observed in lower permanent
molars (range 2.1–10%). Males presented more hypoconulid than females (p = 0.019). Only females presented the distal
trigonid crest in lower first permanent molars (p = 0.002). The most common groove pattern in primary molars was Y; male
presented more Y grade than females in the lower second primary molar (p = 0.039). Asymmetry was observed in some
traits, ranging from 0 to 100%.
Conclusion The present study showed the frequency of NDCT of molars in German orthodontic patients and demonstrated
that some traits present sexual dimorphism.

Keywords Anatomy · Dentition · Tooth crown · Deciduous tooth · Orthodontics

Häufigkeit und Variabilität nicht metrischer Kronenmerkmale von primären und permanenten
Molaren in einer kieferorthopädischen Patientengruppe

Zusammenfassung
Hintergrund Ziel der vorliegenden Studie war es, die Variation und Häufigkeit 13 nicht metrischer Kronenmerkmale
(„nonmetric traits“, NDCT) von Milchmolaren und bleibenden Molaren innerhalb einer deutschen kieferorthopädischen
Patientengruppe zu untersuchen.

 Erika Calvano Küchler

[email protected]

1 4
Department of Biomaterials, University of Uberaba – School of Dentistry, Tuiuti University from Paraná,
UNIUBE, Uberaba, Minas Gerais, Brazil Curitiba, Paraná, Brazil
2 5
Department of Orthodontics, University Medical Centre of Department of Dentistry, Univille – University from the
Regensburg, Regensburg, Germany Joinville Region, Joinville, Santa Catarina, Brazil
3 6
Department of Orthodontics, University of Bonn, Private Orthodontic Practice, Synagogenstr. 1, Ibbenbüren,
Welschnonnenstr. 17, 53111 Bonn, Germany Germany

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 A. Blancato et al.

Material und Methoden Anhand diagnostischer Patientenunterlagen wurden die ersten und zweiten Molaren jedes Qua-
dranten der lactealen und permanenten Dentition untersucht. Zähne mit einer kavitierten Karies, Restauration, okklusalen
Abnutzung oder offensichtlichen Malformation wurden nicht mit einbezogen. Die NDCT der bleibenden Molaren wurden
nach dem odontoskopischen Arizona State University Dental Anthropology System (ASUDAS) bewertet. Die Analyse der
NDCT von Milchmolaren erfolgte nach ASUDAS sowie nach der Hanihara- und der Sciulli-Methode. Seitenpräferenz
sowie Geschlechtsdimorphismus wurden mittels χ2-Test auf einem Signifikanzniveau von p ≤ 0,050 ermittelt.
Ergebnisse Die Untersuchungsgruppe umfasste 163 kieferorthopädische Patienten (davon 82 männliche, 81 weibliche) im
Alter zwischen 8 und 14 Jahren. Für den Hypokonus des ersten oberen bleibenden Molaren ergab sich ein Geschlechtsdi-
morphismus (p = 0,041). Das Protostylid wurde bei den unteren bleibenden Molaren identifiziert (zwischen 2,1 und 10 %).
Das Hypoconulid konnte innerhalb der männlichen Untersuchungsgruppe häufiger nachgewiesen werden als in der weib-
lichen (p = 0,019). Ausschließlich weibliche Patienten wiesen einen distalen Trigonidkamm der unteren ersten bleibenden
Molaren auf (p = 0,002). Das häufigste Fissurenmuster der Milchmolaren war Y-förmig, bei den unteren zweiten Milch-
molaren männlicher Patienten war es stärker ausgeprägt als bei weiblichen Patienten (p = 0,039). Einige Merkmale zeigten
Asymmetrien von 0–100 %.
Schlussfolgerung Die Untersuchungen zeigen die Häufigkeit der NDCT von Milch- und permanenten Molaren in einer
deutschen kieferorthopädischen Patientengruppe und deuten auf einen Geschlechtsdimorphismus einiger Merkmale hin.

Schlüsselwörter Anatomie · Dentition · Zahnkrone · Dens deciduus · Kieferorthopädie

Introduction they produce data that express the variation tendency of all
NDCT present on the molar crown [8, 10].
Molars are the largest teeth in the oral cavity and due to The analysis of dental morphology aims to explore the
their position and eruption time, a central pillars of the de- frequency, the existence of sexual dimorphism and bilat-
velopment of the occlusion. Human molars are found in the eral symmetry of NDCT in permanent and primary teeth
permanent and primary dentition and present a morphologi- [11], evaluating the degrees of expression of these traits
cal variation that ranges in size and shape [1, 2]. The human [12, 13] and allowing the comparison among different pop-
dental morphology, particularly the molars morphology, has ulations. However, dental morphology is not only of interest
been described as presenting several parameters with a va- from an anthropological point of view as the shape of the
riety of morphological traits [3, 4]. The study of crown occlusal relief also has an effect on the development of
morphology, including identification of the molars’ traits, the dental arches. It has been shown in an animal model
is based on evaluation of the occlusal surfaces [5]. Morpho- that interdigitation of molars and canines contributes to the
logical categories have been used over the past century to development of the dental arches [14]. In a recent study,
describe nonmetric dental crown traits (NDCT) in occlusal NDCT were associated with the development of different
surfaces of the mandibular and maxillary molars [6–9]. types of malocclusions in a human sample from India [15].
The first study of dental morphological characteristics in This highlights the importance to understand the frequency
humans was reported by Hrdlička in 1920 [6]. This author of different NDCTs in a wider sample of populations and
described different expressions of a shovel shape on upper how they are connected to the development of different
permanent incisors, in which the trait ranged from minimal malocclusions. There are few studies that investigated mo-
to maximal expression [6, 9]. In 1956, Dahlberg [7] intro- lars anatomy in humans [16–19], and the investigation of
duced scales with grades of trait expressions and a series of these traits in different populations are necessary [12, 13].
standardized plaques to study teeth’s morphological varia- Therefore, this study aimed to investigate the frequency and
tions, such as Carabelli trait, protostylid and hypocone. In variation of 13 NDCT in permanent and primary molars
1961, Hanihara [8] published an important study describing in German orthodontic patients. The presence of bilateral
new classifications of morphological traits of the molars on symmetry and sexual dimorphism was also investigated.
primary dentition, as well as the crown pattern and distal
trigonid crest. Turner et al. [9] developed a dental system
for permanent molars called the ASUDAS (Arizona State Method
University Dental Anthropology System) with an extensive
series of crown and root trait classifications, which is cur- Population and sampling
rently the most widely used system for scoring dental mor-
phology. The use of these systems in dental and anthropo- This descriptive cross-sectional study was approved by the
logical research allows replicability among observers, and institutional review board from the University of Regens-

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Molars’ non-metric crown traits

Table 1 Classification used to assess the morphology of the permanent upper and lower molars
Tab. 1 Klassifikation zur Beurteilung der Morphologie der oberen und unteren bleibenden Molaren
Trait Features Grade
Carabelli’s It occurs on the lingual surface of the protocone (mesiolingual 0. Smooth mesiolingual surface
trait(UM) cusp) and is expressed from complete absence, groove or pit 1. Vertical groove present
(regarded a negative expression) to a large cusp (positive
2. Pit present
expression)
3. Small Y-shaped depression
4. Large Y-shaped depression
5. Small cusp without free apex
6. Medium cusp with free apex making contact with
the medial lingual groove
7. Large free cusp
Metacone(UM) The upper molar has 3 major main cusps regarding the metacone 0. Absent
cusp. It is named the distobuccal cusp or cusp 3 1. There is a ridge but no free apex
2. Faint cuspule with a free apex
3. Weak cusp
3.5. Intermediate-sized cusp that falls between
grades 3 and 4
4. Large cusp
5. Very large cusp, equal in size to a large hypocone
Hypocone(UM) The distolingual cusp is derived from the cingulum and attached to 0. Smooth surface
the distolingual surface of the trigon (protocone or cusp 1, 1. Faint ridge
paracone or cusp 2 and metacone or cusp 3)
2. Faint cuspule
3. Small cusp
3.5. Moderate-sized cusp
4. Large cusp
5. Very large cusp
Metaconule(UM) A 5th cusp on the distal border between the metacone (cusp 3) and 0. Trait is absent
hypocone (cusp 4). The cusp shows two parallel vertical grooves 1. Faint cuspule
2. Trace cuspule
3. Small cuspule
4. Small cusp;
5. Medium-sized cusp
Parastyle(UM) It is most commonly expressed on the paracone (mesiobuccal cusp 0. Smooth buccal surface
or cusp 2). In some instances, it occurs on the metacone (cusp 3) 1. A small pit near the buccal groove
2. Small cusp without free apex
3. Medium cusp with free apex
4. Large cusp with free apex
5. Very large cusp that may extend onto the surfaces
of both cusps 2 and 3
6. Peg-shaped crown attached to root of second or
third molar
Groove Lower molars can have 5 cusps: cusp 1 (protoconid), cusp 2 Y. Cusps 2 and 3 are in contact
pattern(LM) (metaconid), cusp 3 (hypoconid), cusp 4 (entoconid) and cusp 5 +. Cusps 1–4 are in contact
(hypoconulid). The cusps can make contacts creating patterns
X. Cusps 1 and 4 are in contact
Hypoconulid(LM) The cusp 5 occurs on the distal occlusal surface and it associated 0. Absent. The molar has only 4 cusps
with cusp 3 (hypoconid) 1. Very small
2. Small
3. Medium-sized
4. Large
5. Very large

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Table 1 (Continued)
Tab. 1 (Fortsetzung)
Trait Features Grade
Cusp 6(LM) This cusp is expressed on the distal portion, but it is associated 0. Absent
with the cusp 4 (entoconid). It is important to note that cusp 5 1. Much smaller than cusp 5
(hypoconulid) has to be present
2. Smaller than cusp 5
3. Equal in size to cusp 5
4. Large than cusp 5
5. Much larger than cusp 5
Cusp 7(LM) It is a wedge-shaped accessory cusp expressed in the lingual 0. Absent
groove between cusps 2 (metaconid) and 4 (entoconid). Cusp 7 is 1. Faint cusp, two weak lingual grooves
never considered in determining cusp number
1A. Faint tipless displaced on the lingual surface of
cusp 2
2. Small
3. Medium-sized
4. Large
Protostylid(LM) This trait is a cingular derivate on the buccal surface associated 0. Smooth surface
with the buccal groove, particularly separating cusp 1 (protoconid) 1. Pit present
and cusp 3 (hypoconid)
2. Buccal groove curve distal
3. Faint groove extending mesial from the buccal
groove
4. Groove more pronounced
5. Groove stronger
6. Groove extend across the buccal surface
7. Free cusp
Anterior This trait is expressed on the mesial occlusal surface. It involves 0. Absent
Fovea(LM) distinct essential ridges on cusp 1 (protoconid) and cusp 2 1. Trace, with a weak ridge connecting the mesial
(metaconid) that meet close to the centre of the trigonid, and aspects
a mesial marginal ridge that is expressed to varying degrees
2. Essential ridges on trigonid better developed and
resulting groove deeper than in grade 1
3. Essential ridges pronounced and marginal ridge
well developed, producing a distinctive fovea on the
anterior portion of the trigonid
4. The mesial ridge is robust and the marginal ridge
produces a well-defined fovea with a very long groove
Deflecting The form of manifestation of the essential medial ridge on cusp 2 0. Absent
Wrinkle(LM) (metaconid) 1. The essential ridge is straight and shows a midpoint
constriction
2. The essential ridge is deflected distally. There is no
contact with cusp 4 (entoconid)
3. The essential ridge shows strong deflection at the
midpoint forming an L-shaped ridge. There is contact
with cusp 4 (entoconid)
Distal The major mesial cusps of the trigonid (protoconid or cusps 1, and 0. Absent
Trigonid metaconid or cusp 2) express distal accessory ridges that are 1. Present: distal borders are connected by a ridge
Crest(LM) directly connected along the distal portion of the cusps. They can
be continuous or discontinuous

ASUDAS method was used [9]

(UM)permanent upper molar, (LM)permanent lower molar

burg (approval number ID: 19-1549-101). All included sub- Statement of Strengthening the Reporting of Observational
jects and/or their legal guardians signed the informed con- Studies in Epidemiology (STROBE) [20].
sent prior to the inclusion in the study. Age-appropriate Dental casts (orthodontic diagnostic casts) from or-
assent documents were also used for individuals younger thodontic patients (children older than 8 years old and
than 14 years. This project was performed according to the teenagers) undergoing orthodontic treatment were con-
Helsinki Declaration. This study was reported following the secutively selected by convenience at the University of

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Molars’ non-metric crown traits

Regensburg and private orthodontic practices in Regens- The NDCT for primary molars were identified and
burg, Germany. Dental cast from patients with syndromes, scored according to the ASUDAS [9], Hanihara [8] and
oral cleft, congenital alterations including tooth agenesis Sciulli methods [21]. The evaluated traits and the variability
(except for third molar agenesis), and severe bruxism with of their expressions are shown in Table 2 and illustrated in
dental tissue loss were excluded to prevent distortion of the Fig. 2. The name and position of each cusp are illustrated
data. To maximize data interpretability, only patients with in Fig. 2.
a Middle-European ancestry (at maximum one grandparent Examples of NDCT for primary molars (Supplementary
not from Middle Europe) were included. Included individ- Fig. 1) and for permanent molars (Supplementary Fig. 2)
uals should have at least one set (maxilla and mandible) of are shown in the supplementary figures.
dental casts. Some individuals presented more than one set To describe the jaw, letters were used: UM for perma-
of dental casts (at different ages and from different stages nent upper molar, um for primary upper molar, and LM for
of the orthodontic treatment). In these cases, both dental permanent lower molar, lm for a primary lower molar. To
casts were assessed in order to evaluate the majority of describe the tooth position, numbers were used: 1 for the
molars possible (primary and permanent), however, each first molar, and 2 for the second molar [22].
tooth was evaluated only once. All analyses were performed by one single trained and
calibrated examiner. The kappa (κ) test for intraobserver
Morphological analysis reliability was performed, in which the assessments were
carried out twice by the same examiner within a 2-week
All dental casts were scanned and processed into the soft- interval. The κ values for agreement ranged from 0.71 to
ware using the three-dimensional (3D) measuring OrthoXS- 1.00.
can 2.8 (Dentaurum, Ispringen, Germany). The software
OnyxCeph3™ (version 3.2.52, Image Instruments GmbH, Statistical analysis
Chemnitz, Germany) was used to take images of the mo-
lars from the virtual 3D casts. First and second permanent The prevalence of each trait as well as each variability ex-
and primary maxillary and mandibular molars (from left pression were described. Left–right symmetry/asymmetry
and right side) were assessed. Teeth with cavitated dental were evaluated only in subjects that presented the molar
caries, occlusal wear, restorations and obvious dental de- and its contralateral available for analysis. Also, symme-
formities were not evaluated. try and asymmetry were only investigated when the trait
The NDCT for permanent molars were identified and (regardless the expression) was present.
scored according to the odontoscopic system developed The χ2 test was used to investigate whether there is a side
from Arizona State University Dental Anthropology Sys- (laterality) and/or gender (sexual dimorphism) preference.
tem (ASUDAS) [9]. The ASUDAS uses standard recording The odds ratio (OR) and 95% confidence interval (CI)
forms to evaluate traits and the variability of their expres- were also calculated. Data were analysed using the soft-
sions, which are shown in Table 1 and illustrated in Fig. 1. ware PRISM 9 (GraphPad Software, 9.0, San Diego, CA,
For the hypocone trait, grades 1 and 2, and grades 3 and 4 USA) at a significance level of p ≤ 0.050.
were grouped and assessed as having the same degree of
expression. The name and position of each cusp are also
illustrated in Fig. 1.

Fig. 1 Permanent maxillary and

mandibular right first molar
showing cusps indicated by
numbers according to their
location. UM1 permanent upper
molar 1, LM1 permanent lower
molar 1
Abb. 1 Bleibender oberer und
unterer rechter erster Molar. Die
Höcker sind entsprechend ihrer
Lage nummeriert. UM1 bleiben-
der oberer Molar 1, LM1 blei-
bender unterer Molar 1

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Table 2 Classification used to assess the morphology of the primary upper and lower molars
Tab. 2 Klassifizierung zur Beurteilung der Morphologie der primären oberen und unteren Molaren
Trait Features Grade
Carabelli’s It occurs on the lingual surface of the protocone (mesiolingual 0. Smooth surface
trait3 (um) cusp). This tuberculum projection is expressed from complete 1. Pit, groove
absence, groove or pit to a large cusp
2. Two grooves are roughly parallel
3. The area between grooves raised, apex not free
4. The expression is similar to type 3 but there is a free apex
Crown The pattern is based on the development of the crown cusps of 2. Protocone and paracone
pattern the upper 1st deciduous molars 3M. Protocone, paracone and metacone
1st molar2 (um)
3H. Protocone, paracone and hypocone
4–. All four cusps but hypocone reduced
4. All four cusps but hypocone not reduce
Crown The pattern is based on the development of the crown cusps of 3. Protocone, paracone, metacone, and a small hypocone.
pattern the upper 2nd deciduous molars The distal marginal ridge can be interrupted by a groove
2nd molar2 (um) 4–. Protocone, paracone, metacone, and a small hypocone.
The distal marginal ridge follows its course without interrup-
tion
4. Protocone, paracone, metacone, and large hypocone
Meta- A 5th cusp on the distal border between the metacone and 0. Absent
conule3 (um) hypocone. The cusp shows two parallel vertical grooves P. Present
Parastyle1 (um) It is most common expressed on the paracone (mesiobuccal 0. Smooth buccal surface
cusp). In some instances, it occurs on the metacone 1. A small pit near the buccal groove
2. Small cusp without free apex
3. Medium cusp with free apex
4. Large cusp with free apex
5. Very large cusp that may extend onto the surfaces of both
cusps 2 (paracone) and 3 (metacone)
6. Peg-shaped crown attached to root of second or third
molar
Groove Lower molars can have 5 cusps: cusp 1 (protoconid), cusp 2 Y. Cusps 2 and 3 are in contact
pattern1 (lm) (metaconid), cusp 3 (hypoconid), cusp 4 (entoconid) and cusp 5 +. Cusps 1–4 are in contact
(hypoconulid). The cusps can make contacts creating patterns
X. Cusps 1 and 4 are in contact
Hypo- Cusp 5 occurs on the distal occlusal surface and is associated 0. Absent
conulid3 (lm) with cusp 3 (hypoconid) P. Present
Cusp 61 (lm) This cusp is expressed on the distal portion, but it is associated 0. Absent
with the cusp 4 (entoconid). It is important to note that cusp 5 1. Much smaller than cusp 5
(hypoconulid) has to be present
2. Smaller than cusp 5
3. Equal in size to cusp 5
4. Large than cusp 5
5. Much larger than cusp 5
Cusp 72 (lm) It is a wedge-shaped accessory cusp expressed in the lingual 0. Absent
groove between cusps 2 (metaconid) and 4 (entoconid). Cusp 7 1. Only a very weak short groove extends downward from
is never considered in determining cusp number the lingual ridge of cusp 3 (hypoconid). There is no cusp
2. There are two grooves on the lingual surface and a small
cusp is present
3. The cusp is well developed
Proto- This trait is a cingular derivate on the buccal surface associated 0. Smooth surface
stylid1 (lm) with the buccal groove, particularly separating 1. Pit present
cusp 1(protoconid) and cusp 3 (hypoconid)
2. Buccal groove curve distal
3. Faint groove extending mesial from the buccal groove
4. Groove more pronounced
5. Groove stronger
6. Groove extend across the buccal surface
7. Free cusp

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Molars’ non-metric crown traits

Table 2 (Continued)
Tab. 2 (Fortsetzung)
Trait Features Grade
Anterior This trait is expressed on the mesial occlusal surface. It involves 0. Absent
fovea1 (lm) distinct essential ridges on cusp 1 (protoconid) and cusp 2 1. Trace, with a weak ridge connecting the mesial aspects
(metaconid) that meet close to the centre of the trigonid, and
2. Essential ridges on trigonid better developed and resulting
a mesial marginal ridge that is expressed to varying degrees
groove deeper than in grade 1
3. Essential ridges pronounced and marginal ridge well de-
veloped, producing a distinctive fovea on the anterior por-
tion of the trigonid
4. The Mesial ridge is robust and the marginal ridge pro-
duces a well-defined fovea with a very long groove
Central The form of manifestation of the essential medial ridge on 1. The essential ridge of the cusp 2 is expressed similar in
ridge of the cusp 2 (metaconid). This trait is the same as Deflecting Wrinkle size and prominence as that of the other cusps
metaconid2 (lm) on permanent dentition 2. The essential ridge is very well developed in its thickness
on cusp 2 and also expands its width in the trigonid basin.
The ridge sometimes seems to curve distally at its inner end
Distal The major mesial cusps of the trigonid (cusps 1 and 2) express 0. Absent
trigonid distal accessory ridges that are directly connected along the 1. Present: distal borders are connected by a ridge
crest2 (lm) distal portion of the cusps. They can be continuous or
discontinuous
1
ASUDAS methods [9], 2Hanihara methods (1961), 3Sciulli methods (1998), (um)deciduous upper molar, (lm)deciduous lower molar

Fig. 2 Primary maxillary and

mandibular right first and second
molars showing cusps indicated
by numbers according to their
location. um1/um2 primary up-
per molar 1/2, lm1/lm2 primary
lower molar 1/2, asterisk Cara-
belli’s trait
Abb. 2 Erste und zweite Milch-
molaren des Ober- und Unter-
kiefers rechts. Die Höcker sind
entsprechend ihrer Lage num-
meriert. um1/um2 lactealer obe-
rer Molar 1/2, lm1/lm2 lactealer
unterer Molar 1/2, Asterisk Ca-
rabelli-Zeichen

Results frequency in UM. The parastyle cusp was highly uncom-

mon; only 2 teeth were affected. A sexual dimorphism was
Dental records from 163 orthodontic patients (82 males and observed for the hypocone in UM1, in which only females
81 females) were included, in primary molars, a total of presented reduced expression of the traits (p = 0.041; the
40 um1 (r), 39 um1 (l), 65 um2 (r), 68 um2 (l), 32 lm1 (r), whole dataset is available in Supplementary Table 1).
37 lm1 (l), 59 lm2 (r), and 64 lm2 (l). In permanent mo- The prevalence of each trait in LM is presented in
lars, a total of 162 UM1 (r), 162 UM1 (l), 160 UM2 (r), Table 4. In LM1 the Y groove pattern was the most com-
158 UM2 (l), 160 LM1 (r), 160 LM1 (l), 149 LM2 (r), monly observed. The hypoconulid was highly prevalent in
and 143 LM2 (l) were investigated. Their age ranged from LM1. Cusp 6 was an uncommon trait and appeared only in
8–14 years, and they were in mixed or permanent dentition. LM1. A similar pattern was observed in cusp 7; however,
The prevalence of each trait in UM is presented in the 4 teeth in LM2 presented the trait. The protostylid (a pit
Table 3. The most common cusp was the metacone, in feature) was observed in LM1 and LM2 (LM1r = 10.0%;
which none of the UM showed the absence of metacone LM1l = 8.7%; LM2r = 2.1%; and LM2l = 2.1%). Anterior
(grade 0). The grade 4/5 of hypocone was highly preva- fovea was observed in about half of the teeth. Deflecting
lent in UM1. In Carabelli trait negative expression traits wrinkles were highly frequent in LM1. The distal trigo-
(grades 1–4) were more frequent compared to positive ex- nid crest was not a common trait. Males presented more
pressions (grades 5–7). The metaconule trait presented low hypoconulids than females, when the analysis is performed

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Table 3 Traits prevalence in the upper permanent molars

Tab. 3 Prävalenz von Merkmalen der oberen bleibenden Molaren
Trait/Grade First molars Second χ2 test
molars (laterality) (dimorphism)
UM1 (r) UM1 (l) UM2 (r) UM2 (l) UM1 UM2 UM1 UM2
CARABELLI TRAIT, n (%)
0. Smooth surface 86 (54.1) 88 (55.0) 14 (91.2) 147 (93.0) 0.985 0.910 0.637 0.279
1. Vertical groove 17 (10.7) 12 (7.5) 5 (3.1) 4 (2.5)
2. Pit 6 (3.8) 7 (4.4) 4 (2.5) 1 (0.6)
3. Small Y-shaped 14 (8.8) 16 (10.0) 2 (1.3) 3 (1.9)
4. Large Y-shaped 7 (4.4) 9 (5.6) 1 (0.6) 1 (0.6)
5. Small cusp 9 (5.7) 8 (5.0) 1 (0.6) 1 (0.6)
6. Medium cusp 17 (10.7) 17 (10.6) 1 (0.6) 1 (0.6)
7. Large free cusp 3 (1.9) 3 (1.9) – –
METACONE, n (%)
2. Faint cuspule – – – 1 (0.7) 0.850 0.713 0.791 0.698
3. Weak cusp – – 8 (5.3) 10 (6.8)
4. Large cusp 20 (12.6) 21 (13.3) 85 (56.3) 82 (55.8)
5. Very large cusp 139 (87.4) 137 (86.2) 58 (38.4) 54 (36.7)
HYPOCONE, n (%)
0. Smooth surface – – 23 (16.5) 17 (12.7) 0.999 0.818 0.041* 0.578
1/2. Faint ridge and Faint cuspule 1 (0.6) 1 (0.6) 34 (24.5) 35 (26.1)
3. Small cusp 2 (1.3) 2 (1.3) 52 (37.4) 54 (40.3)
4/5. Large and very large cusp 157 (98.1) 157 (98.1) 30 (21.6) 28 (20.9)
METACONULE, n (%)
0. Trait is absent 142 (89.9) 145 (92.4) 116 (87.2) 115 (91.3) 0.950 0.962 0.054 0.617
1. Faint cuspule 6 (3.8) 3 (1.9) 4 (3.1) 4 (3.2)
2. Trace cuspule 5 (3.2) 4 (2.5) 7 (5.4) 5 (3.9)
3. Small cuspule 1 (0.6) 1 (0.6) 2 (1.5) 1 (0.8)
4. Small cusp 1 (0.6) 1 (0.6) 1 (0.75) 1 (0.8)
5. Medium-sized cusp 3 (1.9) 3 (1.9) – –
PARASTYLE, n (%)
0. Smooth surface 162 (100.0) 162 (100.0) 158 (99.4) 154 (99.4) – 0.999 – 0.150
4. Large cusp – – 1 (0.6) 1 (0.6)
U upper, M molar, 1 first, 2 second, (r) right, (l) left
*Statistical difference (p ≤ 0.05)

grouping grades 1–5 compared with grade 0, males (11.3%) trait was absent in the sample. There was no gender or side
presented statistically more hypoconulid traits than females preference observed in primary molars (the whole dataset
(2.8%; p = 0.019; OR = 4.2, CI 95% = 1.3–14.3). Gender is available in Supplementary Table 3).
differences were also observed for cusp 7, in which only The prevalence of each trait for primary lower molar (lm)
males presented the trait in LM2 (p = 0.050). On the other is presented in Table 6. The most common groove pattern
hand, the distal trigonid crest was more common in fe- was Y (in which cusps 2 and 3 are in contact). Hypoconulid
males. In the LM1 only females (6.2%) presented the distal appeared in all lm2. Cusp 6 and cusp 7 were uncommon
trigonid crest (p = 0.002). In the LM2 the prevalence of dis- traits and were observed only in lm2. Protostylid was an
tal trigonid crest was 8.2% in males, but 24.1% in females uncommon trait in lm1, but a common trait in lm2. The
(p = 0.0003; OR = 3.6, CI95% =1.7–7.0; the whole dataset presence of the different expressions of anterior fovea was
is available in Supplementary Table 2). prevalent in lm2. The central ridge metaconid showed that
The prevalence of each trait for upper primary molars this trait described as the cusp 2 ridge is similar to the other
(um) is presented in Table 5. The Carabelli trait was ob- cusp is more prevalent in lm1 and lm2. The distal trigo-
served only in um2. The most common crown pattern in nid crest was an uncommon trait in lm1 and lm2. Gender
um1 was the protocone and paracone; and large hypocone difference was present for groove pattern, in which males
in um2. Metaconule cusp was a rare found, and parastyle presented more Y grade and females more + grade in lm2

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Table 4 Trait prevalence in the lower permanent molars

Tab. 4 Prävalenz von Merkmalen der unteren bleibenden Molaren
Trait / Grade First Second χ2 test
molars molars (laterality) (dimorphism)
LM1 (r) LM1(l) LM2 (r) LM2 (l) LM1 LM2 LM1 LM2
GROOVE PATTERN, n (%)
Y. Cusps 2 and 3 are in contact 94(74.0) 97(77.6) 38(27.3) 37(27.8) 0.623 0.685 0.247 0.067
+. Cusps 1–4 are in contact 22(17.3) 21(16.8) 57(41.0) 60(45.1)
X. Cusps 1 and 4 are in contact 11(8.7) 7(5.6) 44(31.7) 36(27.1)
HYPOCONULID, n (%)
0. Absent 22(13.8) 19(11.9) 104(91.2) 89(94.7) 0.994 0.692 0.204 0.070
1. Very small 11(6.9) 13(8.2) 1(0.9) 1(1.1)
2. Small 34(21.4) 33(20.8) 6(5.3) 2(2.1)
3. Medium-sized 72(45.3) 74(46.5) 3(2.6) 2(2.1)
4. Large 17(10.7) 17(10.7) – –
5. Very large 3(1.9) 3(1.9) – –
CUSP 6, n (%)
0. Absent 150(94.3) 151(94.9) 107(100.0) 87(100.0) 0.908 – 0.168 –
1. Much smaller than cusp 5 4(2.5) 3(1.9) – –
2. Smaller than cusp 5 3(1.9) 2(1.3) – –
3. Equal in size to cusp 5 2(1.3) 3(1.9) – –
CUSP 7, n (%)
0. Absent 155(96.9) 157(98.1) 147(98.7) 141(98.6) 0.733 0.967 0.091 0.050*
1. Faint cusp 1(0.6) 1(0.6) 2(1.3) 2(1.4)
2. Small 1(0.6) – – –
3. Medium-sized 2(1.3) 2(1.3) – –
4. Large 1(0.6) – – –
PROTOSTYLID, n (%)
0. Smooth surface 144(90.0) 146(91.3) 141(97.9) 143(97.9) 0.701 0.986 0.701 0.999
1. Pit present 16(10.0) 14(8.7) 3(2.1) 3(2.1)
ANTERIOR FOVEA, n (%)
0. Absent 77(55.4) 76(53.1) 66(46.1) 56(39.4) 0.983 0.708 0.591 0.184
1. Trace with a weak ridge 14(10.1) 15(10.5) 18(12.6) 19(13.4)
2. Essential ridges on trigonid 43(30.9) 47(32.9) 55(38.5) 63(44.4)
developed
3. Essential ridges pronounced 5(3.6) 5(3.5) 4(2.8) 4(2.8)
DEFLECTING WRINKLE, n (%)
0. Absent 45(33.6) 44(31.9) 111(77.1) 111(77.6) 0.871 0.795 0.437 0.932
1. Midpoint constriction 45(33.6) 49(35.5) 19(13.2) 21(14.7)
2. Deflected distally 37(27.6) 35(25.4) 14(9.7) 11(7.7)
3. L-shaped ridge 7(5.2) 10(7.2) – –
DISTAL TRIGONID CREST, n (%)
0. Absent 130(97.0) 132(97.1) 123(87.2) 113(81.3) 0.983 0.172 0.002* 0.0003*
1. Present 4(2.9) 4(2.9) 18(12.8) 26(18.7)
L lower, M molar, 1 first, 2 second, (r) right, (l) left
*Statistical difference (p ≤ 0.05)

(p = 0.039). Side preferences were not observed (the whole hypoconulid in LM1 (7.1%); LM2 (57.1%), lm1 (25%);
dataset is available in Supplementary Table 4). protostylid in lm2 (6.9%); anterior fovea in LM1 (4.7%),
Asymmetry was observed in some traits in low fre- LM2 (10.9%), lm2 (0.03%); deflecting wrinkle in LM1
quency, as follows: for Carabelli’s trait in um2 (12.5%), (17.2%); LM2 (11.8%); and central ridge of the metaconid
UM1 (21.1%); hypocone in UM2 (17.5%); groove pattern in lm2 (5%). Other traits presented high prevalence of
in LM1(4.2%), LM2 (8.4%), lm1 (5.6%), lm2 (3.2%); asymmetry: Carabelli’s trait in UM2 (40.0%); metaconule

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Table 5 Trait prevalence in the upper primary molars

Tab. 5 Merkmalsprävalenz der oberen Milchmolaren
Trait/Grade First Second χ2 test
molars molars (laterality) (dimorphism)
um1 (r) um1 (l) um2 (r) um2 (l) um1 um2 um1 um2
CARABELLI TRAIT, n (%)
0. Smooth mesiolingual surface 40(100.0) 39(100.0) 18(28.6) 21(31.8) – 0.957 – 0.383
1. Pit, groove – – 26(41.3) 24(36.4)
2. Two grooves are parallel – – 6(9.5) 7(10.6)
3. The area between grooves raised – – 8(12.7) 10(15.1)
4. Free apex – – 5(7.9) 4(6.1)
CROWN PATTERN (first molars), n (%)
2. Protocone and paracone 28(73.7) 27(77.1) – – 0.905 – 0.133 –
3M. Protocone, paracone and metacone 3(7.9) 3(8.6) – –
3H. Protocone, paracone and hypocone 3(7.9) 3(8.6) – –
4–. Four cusps present but hypocone 4(10.5) 2(5.7) – –
reduced
CROWN PATTERN (second molars), n (%)
3. The distal marginal ridge has – – 7(10.9) 8(12.1) – 0.976 – 0.469
a groove
4–. Distal marginal ridge without inter- – – 15(23.4) 15(22.7)
ruption
4. Large hypocone – – 42(65.2) 43(65.1)
METACONULE, n (%)
0. Absent 38(100.0) 35(100.0) 61(98.4) 64(98.5) – 0.973 – 0.227
1. Present – – 1(1.6) 1(1.5)
PARASTYLE, n (%)
0. Absent 40(100.0) 39(100.0) 65(100.0) 68(100.0) – – – –
u upper, m molar, 1 first, 2 second, (r) right, (l) left

in UM1 (40.0%), UM2 (66.7%), um2 (100%); cusp 6 in we investigated an orthodontic sample of German children
LM1 (40.0%), lm2 (100%); cusp 7 in LM1 (85.7%), lm2 and teenagers in order to access molar morphology in a pop-
(53.8%); protostylid in LM1 (33.3%), LM2 (50%) and ulation with a middle European ancestry.
distal trigonid crest in LM2 (55.2%). Trait asymmetry was One important aspect to be emphasized is that dental
not present in metacone, crown pattern and parastyle in size and morphology are studied from an interdisciplinary
UM2. viewpoint, such as anthropology, paleopathology, archeol-
ogy forensic science, and dentistry, especially orthodontics.
To investigate dental traits requires comprehensive knowl-
Discussion edge of morphology, comparative anatomy, function and
occlusion [4, 24]. Dental anthropology involves the study
The present study investigated the frequency and variability of the origin and variations of the human dentition, includ-
of non-metric permanent and primary molars crown traits ing the identification of structures such as cusp size, number
in German orthodontic patients. Additionally, the presence and location of cusps, occlusal pattern, root configuration,
of laterality and sexual dimorphism were also investigated. number and position of teeth, and individual measurements
Although several studies have been focusing on specific [12, 16, 25]. In our study, we used similar methods used in
traits, such as Carabelli’s trait [23], so far, only few studies dental anthropology research to investigate the frequency,
have been converging many aspects of dental morphology the sexual dimorphism and the left–right symmetry of mo-
in living humans, and these studies investigated only few lar traits in orthodontic patients.
populations and ethnicities, such as Venkatesh et al. [18] One of the most well-known methods for assessing the
and Sujitha et al. [19] who evaluated populations from In- morphology of the permanent dentition is the Arizona State
dia, Felemban and Manjunatha [17] who examined a sample University Dental Anthropology System (ASUDAS) [9]
from Saudi Arabian, and Aguirre et al. [16] who studied that describes the dental root and crown. A tooth is split
a sample from Colombia. Therefore, in the present study, into two main parts: the root and the crown, and cusp is

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Molars’ non-metric crown traits

Table 6 Trait prevalence in the lower primary molars

Tab. 6 Merkmalsprävalenz der unteren Milchmolaren
Trait/Grade First Second χ2 test
molars molars (laterality) (dimorphism)
lm1 (r) lm1(l) lm2 (r) lm2 (l) lm1 lm2 lm1 lm2
GROOVE PATTERN, n (%)
Y. Cusps 2 and 3 are in contact 20 (95.2) 21 (95.5) 28 (77.8) 34 (89.5) 0.973 0.290 0.701 0.039
+. Cusps 1–4 are in contact – – 4 (11.1) 3 (7.9)
X. Cusps 1 and 4 are in contact 1 (4.8) 1 (4.5) 4 (11.1) 1 (2.6)
HYPOCONULID, n (%)
0. Absent 25 (89.3) 29 (85.3) – – 0.640 – 0.549 –
1. Present 3 (10.7) 5 (14.7) 57 (100.0) 63(100.0)
CUSP 6, n (%)
0. Absent 28 (100.0) 34 (100.0) 57 (100.0) 62 (98.4) – 0.339 – 0.275
2. Cusp 6 smaller than cusp 5 – – – 1 (1.6)
CUSP 7, n (%)
0. Absent 29 (100.0) 35 (100.0) 48 (84.2) 53 (82.8) – 0.968 – 0.422
1. Weak short groove of cusp 3 * – – 8 (14.0) 10 (15.6)
2. Small cusp – – 1 (1.7) 1 (1.6)
PROTOSTYLID, n (%)
0. Smooth surface 31 (96.9) 36 (97.3) 29 (49.1) 31 (48.4) 0.917 0.936 0.101 0.089
1. Pit present 1 (3.1) 1 (2.7) 30 (50.9) 33 (51.6)
ANTERIOR FOVEA, n (%)
0. Absent 24 (100.0) 29 (93.6) 12 (30.0) 7 (16.7) 0.447 0.396 0.603 0.136
1. Trace with a weak ridge – 1 (3.2) 6 (15.0) 11 (26.2)
2. Essential ridges on trigonid devel- – 1 (3.2) 21 (52.5) 21 (50.0)
oped
3. Essential ridges pronounced – – 1 (2.5) 2 (4.8)
4. Well-defined fovea – – – 1 (2.4)
CENTRAL RIDGE OF METACONID, n (%)
1. Cusp 2 ridge is similar to the other 24 (92.3) 31(91.2) 40 (88.9) 46 (92.0) 0.875 0.605 0.065 0.934
cusps *
2. Ridge is well developed and expands 2 (7.7) 3 (8.8) 5 (11.1) 4 (8.0)
to trigonid ***
DISTAL TRIGONID CREST, n (%)
0. Absent 27(93.1) 31 (91.2) 35 (94.6) 35 (92.1) 0.777 0.665 0.087 0.080
1. Distal borders are connected by 2 (6.9) 3 (8.8) 2 (5.4) 3 (7.9)
a ridge
l lower, m molar, 1 first, 2 second, (r) right, and (l) left
*Weak short groove on the lingual ridge of cusp 3
**The essential ridge of the cusp 2 is expressed similar in size and prominence as that of the other cusps
***The essential ridge is very well developed in its thickness on cusp 2 and also expands its width in the trigonid basin

the singular part constituting the crown in addition to es- lars [8, 21]. The NDCT for primary molars were identified
sential lobes and ridges (the NDCT) [4, 16], which plays and scored according to ASUDAS [9]; Hanihara method [8]
an important role in the establishment of dental occlusion. was used for the crown pattern of 1st and 2nd upper mo-
In our study, we used ASUDAS to access and describe lar, cusp 7, central ridge of the metaconid and distal trigo-
13 NDCT of the permanent dentition, including Carabelli’s nid crest. For Carabelli’s trait, metaconule and hypoconulid
trait, metacone, hypocone, metaconule, parastyle, groove classification, the Sciulli method [21] was used, and for
pattern, hypoconulid, cusp 6, cusp 7, protostylid, anterior parastyle, groove pattern, cusp 6, protostylid and anterior
fovea, deflecting wrinkle and distal trigonid crest. For pri- fovea the ASDUDAS system [9] was used. The three stud-
mary dentition, different methods have been proposed to ies established systems for classification allowing measure-
classify primary molars. Thus, in addition to ASUDAS [9], ment of minimal and maximal trait expressions and degrees
we also used other methods to access NDCT in primary mo- between these two points [22]. It is important to highlight

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the importance of using the same method of classification Cusp 6, or the entoconulid, is a supernumerary cusp po-
for the permanent and primary dentition [19] for some traits sitioned on the distal portion and associated with the en-
observed in both dentitions. toconid (cusp 4) [22]. Similar to that found in Europeans
According to Scott and Pilloud [10], over 90% of the (5–15%) [22] and in agreement with Kirthiga et al. [33], our
published manuscripts on human dental morphology fo- study showed a frequency of 5% on LM1. Primary dentition
cus on permanent teeth, despite primary teeth offering an- had only one case, which was different from the previous
other perspective on morphological variation. It has been studies that showed a frequency higher than 5% [19, 33];
described that primary teeth hold more primitive traits than this difference may be explained by the population differ-
permanent teeth [4]. Classic studies from the past century ence or by the sample size that was small for primary teeth
stated that teeth are independent of each other in variation in our study.
and evolution [26]. Butler [26] and Dahlberg [27] published A second supernumerary cusp of the lower molar is
studies describing the role of cusp variations as a field ef- cusp 7, or metaconulid, expressed between metaconid
fect. The most distal member on the tooth crown is the most (cusps 2) and entoconid (cusp 4). Cusp 7 is relatively rare
variable element. This gives us an indication of how NDCTs ranging from 3 to 8% worldwide [22], which is in agree-
might be associated with the development of malocclusions ment with the prevalence observed in our study. In contrast
as recently described by a research team from India [15]. to the permanent dentition and similar to previously studies,
As some NDCTs, for example the size of the hypocone, the the frequency on lm2 was higher ranging from 17.2% to
hypoconulid, cusp 6 and cusp 7, influence mesiodistal width 15.8% [11, 16]. Different methods to classify the perma-
of teeth and therefore the amount of space they require in nent and primary dentition were used [8, 9], although both
the dental arch. As for deciduous molars, their mesiodistal considered similar features from cusp 7.
width correlates directly with the amount of space available How these NDCTs impact mesiodistal tooth width ex-
for the second dentition. But also in the second dentition, actly and how they are connected to malocclusions as
the mesiodistal width of molars influences the amount of crowding or the amount of space available for the eruption
posterior crowding and is therefore associated with tooth of the second dentition is an important question and shall
retention. Our study aimed to characterize the frequency of be assessed in future studies. Some NDCTs as Carabelli’s
different NDCTs. trait or protostylid trait, are not directly connected to the
Upper molars have 3 major main cusps and one of them mesiodistal width of the tooth but are also associated with
is the metacone (cusp 3) [30]. The last major cusp added tooth size. This is of interest to the orthodontist, as or-
during primate evolution is the hypocone (cusp 4) [22]. thodontic bands are used especially on the first molars in
The metaconule (cusp 5) is an occlusal cusp on the dis- the permanent dentition or on the second deciduous mo-
tal border [31]. In our study we observed mesial cusps, lars. The degree to which these features are pronounced
such as metacone (cusp 3), demonstrating less variation as can influence the fit of the orthodontic bands, as these are
compared to distal cusps, such as hypocone (cusp 4) and usually aligned to the most common tooth shape with-
metaconule (cusp 5). Metacone (cusp 3) was prevalent in out the mentioned NDCTs. It is therefore advisable for
grades 4 and 5. Yadav et al. [32] showed similar frequencies orthodontists, if the expression of the traits is especially
for these traits in Indians. In the primary dentition, our re- pronounced, to consider the necessity of an oral attachment
sults showed low prevalence of metacone (cusp 3) in um1 and, sometimes, to forgo the placement of a band and bond
and higher prevalence of the hypocone (cusp 4) in um2. a bracket instead.
Sujitha et al. [19] also investigated Indians and reported In our sample the Carabelli’s trait was especially frequent
that metacone (cusp 3) had a high frequency in um1. The in first molars of the permanent dentition. Carabelli’s trait
parastyle is an accessory cusp on the mesiobuccal surface is a singular derivate expressed on the lingual surface of the
of upper molars and sometimes linked to Bolk’s paramolar protocone and researchers assumed for decades that this is
tubercle [9]. The frequency of this morphological trait was a feature of European-descendant dentition; however, this
below 10% [22], and indeed, in our study only one case latter trait was identified in other populations [22, 28]. In
was observed in UM2. our sample, the prevalence of Carabelli’s trait was high in
Lower molar cusp number depends on the presence of UM1. In the primary dentition, Carabelli’s trait is observed
cusp 5, or the hypoconulid. Each cusp is named and num- only in um2. A recent systematic review and meta-analysis
bered. The last major cusp added during primate evolution from Bhavyaa et al. [23] observed a similar prevalence.
is the hypoconulid (cusp 5) as a distal cusp integrated more For um2 the authors reported an estimated prevalence of
closely with the hypoconid (cusp 3) than entoconid (cusp 4) 72%; likewise the overall prevalence was 59% for UM1
[22]. In the present study, the frequency of five-cusped mo- and 8% for UM2. The subgroup analysis also showed that
lars was predominant on LM1 and lm2. Previous studies the European continent reported the highest prevalence of
support the same result in other populations [11, 19, 34].

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Molars’ non-metric crown traits

Carabelli’s trait [23]. Interestingly, Neanderthals are char- and 7.9%). A possible explanation for these result differ-
acterized by the presence of a larger Carabelli trait [29]. ences may be the variability in the methods used.
The protostylid trait occurs on the buccal surface of the The anterior fovea is a polymorphic trait present in the
mesiobuccal cusp or protoconid (cusp 1) and is a cingu- mesial ridge’s aspect on the protoconid (cusp 1) and meta-
lar derivative as Carabelli’s trait [22]. The present study conid (cusp 2) [6]. Data for this feature have not been tab-
reported in lm2 negative expression. Díaz et al. [11] indi- ulated on a world scale because Turner et al. [9] did not
cated a high prevalence of this trait on lm2, whereas Su- consider the anterior fovea as one of Turner’s key 29 traits.
jitha et al. [19] showed a low prevalence. The protostylid Our study had prevalent results from presence of different
was more common in Australopithecines than in modern degrees on lm2, LM1 and LM2. It has been reported that
humans [40]. Europeans have higher frequencies of this trait in lower
While some NDCTs influence tooth size, others affect molars [22].
the occlusal relief of the molars and therefore are likely to Left–right asymmetry is observed in many conditions,
have an effect on occlusal interdigitation. These traits are of including tooth agenesis and tooth morphological alter-
interest as involvement in the development of malocclusion ations, such as microdontia [36]. Deviation from perfect
has also been suggested [15]. It is our task as orthodontists symmetry expressed by changes in structure, as seen in
to examine the influence of these characteristics on occlu- dental morphology studies, is called fluctuating asymme-
sion and to identify possible disruptive factors in dental try [4, 16, 37]. In our study, the Carabelli trait, metaconule
occlusion. In the current study, the aim was to determine (cusp 5), hypoconulid (cusp 5), cusp 6, cusp 7, protostylid
the frequency of the various characteristics in the first place, and distal trigonid crest were very often observed to be
but studies are to follow that will investigate the involve- asymmetrical. However, a statistically significant difference
ment of NDCTs in the development of malocclusions. was not observed between left and right teeth, showing no
The major mesial cusps of the lower molars, protoconid side preference.
(cusp 1) and metaconid (cusp 2), form the trigonid and The investigation of the difference in morphological
they can exhibit connected ridges [22]. When the distal characteristics of teeth between men and women cor-
accessory ridges run a direct path along the distal portion of roborates the elucidation of sexual dimorphism. Sexual
the cusps (protoconid and metaconid) and come in contact dimorphism is known to be more significant in areas of
at a point close to the central occlusal sulcus, the distal the dental crown that have a later development [4], which
trigonid crest is present [8]. This trait is not common in explains our results that observed a male–female differ-
modern humans and was found by Weidenreich [41] on ence only in permanent molars. Dental development is
lm2. The noteworthy reduction in trigonid crest prevalence influenced by genes located on X and Y chromosomes.
is one hallmark of the modern human dentition [22]. In the The X chromosome is associated with enamel thickness,
present study, the prevalence was 2.9% (LM1) and 18.7% while the Y chromosome promotes growth of enamel and
(LM2) in permanent dentition, and 5.4% (lm2) and 8.8% dentin thickness [4, 38, 39]. The morphological structure,
(lm1) in the primary dentition. Hanihara [8] reported results in terms of size and shape of the cusps, is influenced by
similar to our study, whereas King, Tongkoom and Wong the sex chromosomes in the formation of their phenotype,
[35] (in a Chinese population) and Sujitha et al. [19] (in although they may not be equally influenced in the forma-
an Indian population) reported higher prevalence (33.6%, tion of enamel and dentin [39, 42, 43]. In our study, gender
65.25% and 93.06%, respectively). difference was observed for hypocone, groove pattern, dis-
Deflector wrinkle and central ridge of the metaconid are tal trigonid crest and cusp 7, which suggests that sexual
different terms, but all relate to a common feature on lower chromosomes are carrying markers for these traits.
molars [8, 9]. This trait is expressed on the occlusal surface
of metaconid (cusp 2) and is considered a manifestation of
the essential ridge. In most instances, this ridge runs from Conclusion
the cusp tip to the central occlusal fossa. The deflecting
wrinkles present a wide result variation of the results rang- The present study showed the frequency of nonmetric den-
ing from lower and high prevalence on a global level [22]. tal crown traits (NDCT) of primary and permanent mo-
Our study showed more prevalence of presence degrees on lars in German orthodontic patients and demonstrated that
LM1 (66.4 and 68.1%). King, Tongkoom and Wong [35] some traits present sexual dimorphism in the permanent and
and Sujitha et al. [19] also evaluated the primary dentition primary teeth. Knowledge regarding the interplay between
using the NDCT classification for the permanent dentition dental anatomy, physiology, and occlusion is important for
[9]. In their results, deflecting wrinkle was higher on lm2 clinical practice and should be explored in future studies.
(63.0 and 87.4% respectively) differing from our study (5.4

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Supplementary Information The online version of this article (https:// 4. Kondo S, Morita W, Ohshima H (2022) The biological significance
doi.org/10.1007/s00056-024-00532-3) contains supplementary mate- of tooth identification based on developmental and evolutional
rial, which is available to authorized users. viewpoints. J Oral Biosci 64:287–302. https://doi.org/10.1016/j.
job.2022.05.004
Acknowledgements The authors thank Alexander von Humboldt 5. Aoki S (2022) A cognitive psychological approach for tooth iden-
Foundation and Coordenação de Aperfeiçoamento de Pessoa de Nível tification based on brain event-related potentials. J Oral Biosci
Superior for scientific scholarships. 64:303–311. https://doi.org/10.1016/j.job.2022.04.002
6. Hrdlička, A., 1920. Shovel-shaped teeth. Am J Phys Anthrop.
Funding This study was financed in part by the Coordenação de Aper- 3:429–465.
feiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance 7. Dahlberg AA (1956) Materials for the Establishment of Standards
Code 001 and by the Alexander von Humboldt Foundation, Germany. for Classification of Tooth Characteristics, Attributes, and Tech-
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Menezes-Oliveira, S. Beisel-Memmert, F. Baratto-Filho, C. Lippold, anthropology system. In: Turner II, C.G. Nichol C: (R)
C. Kirschneck, E. Calvano Küchler and C.P. Lepri declare that they 10. Scott GR, Pilloud M, Navega D, Coelho J, Cunha E, Irish J (2018)
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from tooth morphology. Forens. Anthrop, vol 1, pp 18–31 https://
Ethical standards This research project was approved by the insti- doi.org/10.5744/fa.2018.0003
tutional review board from the University of Regensburg (approval 11. Díaz E, García L, Hernández M, Palacio L, Ruiz D, Velandia N,
number ID: 19-1549-101) and is a part of a dissertation from the Uber- Villavicencio J, Moreno F (2014) Frequency and variability of den-
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2428). This project was performed according to the Helsinki Dec- digenous group in the municipality of Morales, Cauca, Colombia.
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signed the informed consent prior to the inclusion in the study. Age- 13. Scott GR, Pilloud M (2018). Dental Morphology. In: Katzenberg,
appropriate assent documents were also used for individuals younger M. A., Grauer, A.L. Biological Anthropology of the Human Skele-
than 14 years. ton. John Wiley & Sons, pp.257–292.
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tion, distribution and reproduction in any medium or format, as long as https://doi.org/10.1093/ejo/19.5.531
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otherwise in a credit line to the material. If material is not included pp 364–380 https://doi.org/10.15644/asc57/4/7
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