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. 2025 Apr 25;11(3):e70368. doi: 10.1002/vms3.70368

Geometric Morphometrics of Astragalus and Shape Variation Analysis in Capra, Ovis, and Bovidae

Yasemin Üstündağ 1, İftar Gürbüz 2,✉, Yasin Demiraslan 1, Özcan Özgel 2
PMCID: PMC12023762  PMID: 40278643

ABSTRACT

In the study, it was investigated whether taxonomic differences have an effect on shape using bovine, ovis, and capra astragalus. A total of 142 samples, 32 bovine (Holstein), 71 ovis (Merino ovis), and 39 capra (Hair goat) astragalus, were used as material. The geometric morphometry method was applied in the study. 13 homologous landmarks were marked on the photographed astragalus. Principal component analysis was performed on the new coordinates obtained as a result of Procrustes analysis. In addition, in areas where shape differences were concentrated, allometric effect and grouping features were determined. The MorphoJ program was used for all these analyses. As a result, 22 principal components were calculated for astragalus. In terms of PC1, capra samples were placed between bovine and ovis sample clusters. Most of the shape variations were formed on the medial side of the astragalus. In conclusion, ovis and bovine astragalus samples were grouped taxonomically completely (100%) separately, while capra samples were grouped largely (97.2%) separately between ovis and capras in cross‐validation scores. The results are important in terms of contributing to taxonomy, morphology, and zooarcheology studies.

Keywords: astragalus, bovine, caprine, ovine, shape

It was investigated whether taxonomic differences have an effect on shape using bovine, sheep, and capra astragalus. Shape variation of sheep, goat, and bovine astragalus was evaluated by the geometric morphometry method. Similar and different findings were obtained from the literature in terms of these features. The variation was seen at landmarks LM3, LM4, LM8, LM9, LM10, LM11, and LM13 between bovine and capras, at landmarks LM2, LM3, LM4, LM8, LM9, and LM11 between bovine and ovis, and at landmarks LM2, LM7, LM11, and LM13 in ovis and capras. It was observed that the distal end of the bovine astragalus was wider than that of the ovis and capras. The results are important in terms of contributing to taxonomy, morphology, and zooarcheology studies.

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1. Introduction

Geometric morphometry is a method that allows the determination of differences and similarities between species of biological shapes through statistical analysis (Rohlf 2000). Geometric morphometry enables the detection of shape differences that cannot be detected by the eye via landmark coordinates (Özden 2008). These landmark coordinates are anatomical points obtained from two or three‐dimensional Cartesian coordinates. It analyzes shapes in a way that preserves their integrity without allowing linear distortion (O'Higgins 2000, Slice 2007, Aytek 2017). In addition, since this method provides more comprehensive shape measurement results than alternative approaches, it is frequently used in scientific fields such as anatomy, biology, ecology, evolution, and archaeology (Adams and Otarola‐Castillo 2013).

Anatomically, the astragalus, together with the calcaneus, participates in the proximal row of the ankle joints. It articulates with the distal tibia proximally and the distal row of tarsal bones distally. A spool‐shaped articular process called the trochlea joins these joints. In ruminants, the trochlea is in two symmetrical pieces, and the direction of the trochlea extends perpendicular to the ground. The astragalus is rectangular in shape in ruminantia (Özcan and Demiraslan 2021).

Since bones have survived for a long time, their remains are also encountered in archaeological excavations. In archaeology, the distinction between ovis, goat, and bovine bones is important in terms of clarifying the basic aspects of human‐animal relationships and the purposes for which these animals were used in the past and in different parts of the world (Buitenhuis 1995; Noddle 1994). However, the information remains of archaeological bone is important in terms of providing information about faunal identification, phylogenetic relationships, and other historical processes, as well as estimating the morphological characteristics of animals. The morphological characteristics of bones are determined by methods such as macroanatomy, biometry, gene analysis, traditional morphometry, and geometric morphometry (Fernandez et al. 2001; Evin et al. 2013).

In recent years, the integration of artificial intelligence, which has been frequently used as a learning tool, into veterinary and medical education provides potential benefits for research, diagnosis, and treatment. In addition, artificial intelligence has an important place in advancing the field of veterinary anatomy (Choudhary et al. 2023; Choudhary et al. 2025). In veterinary anatomy, artificial intelligence is used in advanced diagnostic methods such as imaging analysis and shape/pattern recognition (Vickram et al. 2025). It has been reported that machine learning algorithms can be integrated into morphological descriptions (Bellin et al. 2021). Therefore, deep learning can be achieved in geometric morphometric studies and artificial intelligence‐supported shape analysis. This may be the hypothesis of a morphological study.

Morphological and morphometrical studies are powerful tools for biogeographic, phylogenetic, and systematic studies, especially in the absence of molecular studies. In morphological studies, in addition to the skull (Dayan et al. 2023; Ahani et al. 2024; Alizadeh et al. 2024; Kazemi et al. 2025), mandible (Demircioğlu et al. 2021), vertebral column (Derakhshi et al. 2024), long bones (Manuta et al. 2023), and also short bones such as astragalus (Andrew Barr 2014, Davis 2017) and calcaneus (Lloveras et al. 2022) are frequently examined. One of the bones that allows distinguishing between species is the astragalus. (Haruda 2017). There are studies that previously investigated astragalus morphologically and morphometrically (Haruda 2017; Zeder and Lapham 2010). However, limited information was provided about the variation in the shape of the astragalus of bovine, ovis, and capra species. Therefore, this study aimed to reveal the shape variations in the astragalus of ovis, capra, and bovine, which are taxonomically close.

2. Material and Methods

2.1. Samples

In the study, adult bovine, ovis, and capra astragalus (left side) were used to investigate whether taxonomic differences have an effect on shape. For this purpose, 142 samples were used, including 32 bovine (Holstein), 71 ovis (Merino), and 39 capra (Hair goat) astragalus. Samples were collected from slaughterhouses in the western Mediterranean region. Since the study used astragalus samples from sheep, goats, and cattle, which were slaughterhouse material, and no procedure was applied to live animals, according to the Regulation on “Working Procedures and Principles of Ethics Committees for Animal Experiments” No. “28914” dated “15 February 2014”, there is no need for ethics committee approval.

2.2. Imaging and Digitization

After the left astragalus was dissected, it was photographed from the dorsal side (Canon 600D 18×55 lens, Japan). The photographs were saved to the computer in the JPEG format. A “tps” file was created from the photographs using the TpsUtil (Version 1.79) (Rohlf 2019) software, and 13 homologous landmarks were marked with the TpsDig2 (Version 2.64) (Rohlf 2018) software.

Homologous landmarks (LM) marked on astragalus (Figure 1) were determined as follows; LM1: Most proximal end of lateral proximal trochlea, LM2: Lateral side of the center, LM3: Most concave point of the lateral side, LM4: Most lateral end of distal lateral trochlea, LM5: Most medial end of distal lateral trochlea, LM6: Most concave point between distal lateral and medial trochlea, LM7: Most lateral end of distal medial trochlea, LM8: Most medial end of distal medial trochlea, LM9: Distal start point of the medial protuberence, LM10: Peak point of the medial protuberence, LM11: Proximal start point of the medial protuberence, LM12: Most proximal end of medial proximal trochlea, LM13: The most proximal concave point between the lateral and medial trochlea. With the landmark marking, the x and y cartesian coordinates of the points representing the general shape of the material were determined on the analytical plane.

FIGURE 1.

FIGURE 1

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Homologous landmarks (LM) marked on left astragalus from dorsal view (from distal to proximal).

2.3. Statistical Analysis

Since there were differences in size, position, and direction in the photographs, general Procrustes analysis (superimposition) was performed separately for all bones (Slice 2007). Principal components (PCs) analysis was performed on the new coordinates obtained as a result of the Procrustes analysis. In addition, in areas where shape differences were concentrated, allometric effects and grouping features were determined. MorphoJ (Klingenberg 2011) program was used for all these analyses.

3. Results

As a result of the regression analysis conducted to determine whether the size (centroid size) has an effect on the shape, it was determined that 0.9909% (p = 0.1634) of the shape could be predicted in terms of size (according to the 95% confidence interval). Since the size did not have a statistically significant effect on the shape, other analyses were carried out.

A total of 22 PCs were calculated for the astragalus. The first PC explained 25.190% of the total shape variation, while the first four PCs (PC1+PC2+PC3+PC4) explained 61.841%. The distribution of samples according to the first four PCs is shown in the graph in Figure 2. In terms of PC1, capra samples were placed between bovine and ovis sample clusters.

FIGURE 2.

FIGURE 2

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Graphical distribution of samples according to PCs. BV, bovine; CP, capra; OV, ovis.

Graphs showing the landmark levels at which shape differences were concentrated according to PC1, PC2, PC3, and PC4 are shown in Figure 3. Significant variations were observed at LM2, LM3, LM8, LM9, LM10, LM11 for PC1 and LM3, LM6, LM7, LM9, LM10, LM13 for PC2, and LM2, LM3, LM4, LM8, LM9, LM13 for PC3, and LM2, LM4, LM5, LM6, LM9, LM10, LM11 for PC4. It was striking that these variations were concentrated, especially on the medial surface of the bone.

FIGURE 3.

FIGURE 3

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Wireframe graphical representation of shape differences with respect to PC1 (a), PC2 (b), PC3 (c), and PC4 (d). Dark blue represents the average shape with respect to the principal component.

The shape variation graphs of the astragalus between groups are given in Figure 4. Accordingly, the variation was seen at landmarks LM3, LM4, LM8, LM9, LM10, LM11, and LM13 between bovine and capras, at landmarks LM2, LM3, LM4, LM8, LM9, and LM11 between bovine and ovis, and at landmarks LM2, LM7, LM11, and LM13 in ovis and capras. It was observed that the distal end of the bovine astragalus was wider than that of the ovis and capras.

FIGURE 4.

FIGURE 4

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Graph of shape variation by groups. BV: bovine, CP: Capra, OV: ovis (change in shape is from light blue to dark blue and from BV to CP (BV—CP), and from BV to OV (BV—OV), and from CP to OV (CP—OV)).

In Figure 4, it was observed that the distal and proximal sulcus in the middle of the trochlea were deeper in ovis than capras, the proximal sulcus was deeper in bovine than capras and ovis, and the distal sulcus was deeper in bovine than capras. The protuberance at the level of LM9, LM10, and LM11 was more pronounced in capras than in ovis and bovine.

The data of the discriminant function analysis were presented in Table 1. In terms of grouping characteristics, it was observed that 89.1% of ovis and capras, 97.2% of capras and bovine, and 100% of ovis and bovine were grouped correctly. Mahalanobis and Procrustes distance were calculated as 5.0734 and 0.05262534 between bovine and capras, 5.5075 and 0.06447940 between bovine and ovis, and 3.9408 and 0.04216149 between ovis and capras, respectively. These results were statistically significant (P < 0.0001). According to the cross‐validation scores (Figure 5), while bovine and ovis were grouped completely correctly, a low level of incorrect grouping was detected in the grouping of capra samples.

TABLE 1.

Results of discriminant function analysis.

CP OV
BV MD 5.0734 5.5075
PD 0.05262534 0.06447940
p‐value <0.0001 <0.0001
Misclassification % 2.8 0
CP MD 3.9408
PD 0.04216149
p‐value <0.0001
Misclassification % 10.9
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Abbreviations: MD, Mahalanobis distance; PD, Procrustes distance; p‐value, p‐value calculated for Procrustes distance (from permutation tests, 10,000 permutation rounds); BV, bovine; CP, capra; OV, ovis.

FIGURE 5.

FIGURE 5

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Frequency graph of cross‐validation scores by groups. BV, bovine; CP, capra; OV, ovis.

4. Discussion

Taxonomic classification is important for understanding the natural world, but current methods for the assessment of species generally focus on bone morphology. Knowing the morphological variation in bones is important for determining the genetic structure, evolutionary process, and adaptation of the population (Herrel et al. 2012). Moreover, it is currently unknown how much variation there may or should be between species. Here, the shape characteristics and grouping degrees of the astragalus bones of ovis, capra, and bovine, which are phylogenetically in the same class, were investigated using the geometric morphometry method. Thus, the effect of taxonomy on astragalus shape variation and how bone shapes reflect taxonomy according to species were determined. At the same time, it was determined that the shape characteristics of the astragalus varied between groups and at which landmark level this variation occurred.

Klein et al. (2010) examined the ecomorphology of astragalus in an article on morphometric measurements of bovid metapodiums, which they examined for taxonomic identification. Klein et al. (2010) argued that the distinguishing feature in astragalus morphology is the general size and that size‐independent shape does not distinguish well between bovids in different habitat groups. However, it was stated that there may be a functional relationship between bovid astragalus morphology according to body size differences (Barr 2014). Haruda (2017) examined the astragalus morphology in ovis and capras using the geometric morphometry method. Haruda (2017) determined that ovis and capra astragalus were completely separate groups as a result of canonical variance analysis. In the study, it was statistically determined that bone size had no effect on shape in the comparison of astragalus in ovis, capra, and bovine groups. In addition, while ovis and bovine astragalus collected from the same region were clustered in completely (100%) separate groups taxonomically, capra samples were also found to be grouped separately to a large extent (97.8% and 89.1%). This situation was evaluated as the species difference may have a determining effect on the shape of the astragalus.

Taxonomic criteria that were previously defined and frequently used on the astragalus were the shape of the distal lateral articular facet and the height of the proximal trochlea (Fernandez 2001; Zeder and Lapham 2010), while other features were often unclear (Haruda 2017). Here, in line with the findings of the researchers, it was observed that the proximal trochlea showed variation according to the species, and it was wider in bovine. Unlike the researchers (Fernandez 2001; Zeder and Lapham 2010), no clear difference was observed between capra and ovis in the distal lateral trochlea. The reason for this difference may be that the photographs were taken from different directions. However, Boessneck (1969) reported that the sulcus in the middle of the trochlea is generally deeper in ovis than in capras. In the study, it was similarly observed that the sulcus in the middle of the trochlea (LM6) was deeper in ovis than in capras.

The protuberance of the medial articular ridge of the astragalus (in our study, this protuberance is at the LM10 level) has been noted as valuable for distinguishing between ovis and goats (Boessneck 1969; Fernandez 2001; Prummel and Frisch 1986; Zeder and Lapham 2010). In ovis, this protuberance is parallel to the proximal‐distal plane or has a proximal direction, while in goats, this protuberance is more strongly developed and has a distal direction. In this study, like the findings of the researchers (Boessneck 1969; Fernandez 2001; Prummel and Frisch 1986; Zeder and Lapham 2010), it was observed that in goats, this protuberance is more evident than in ovis and capras (LM10). On the other hand, it was determined that bovine was different from ovis and capras at this protuberance point. It was located more proximally in ovis and capras than in bovine. In addition, a significant difference was observed at this protuberance in PC1.

In a biometric study (Salvagno and Albarella 2017), where ovis and goat astragalus were also discussed, it was emphasized that the measurements of the ratio between the height of the central constriction the greatest depth of the lateral half, plotted against a ratio between the breadth of the distal end and the greatest length of the lateral half, were important. At the same time, as a result of the discriminant analysis evaluating these measurements, it was determined that the discrimination could be made as 89%. It is noteworthy that this variation detected by Salvagno and Albarella (2017) with linear measurements was mostly on the lateral side of the bone. In this study, there was variation on both the lateral and medial sides of the bone in ovis and capra. In accordance with the findings of the researchers (Salvango and Albarella 2017), the results of the discriminant function as 89.1% between ovis and capra.

In a study comparing goat astragalus according to gender using the geometric morphometry method (Oktay et al. 2024), it was determined that the shape variation was in the medioproximal part of the astragalus, and the proximal part of the astragalus was wider in male goats. In the study, according to the principal component analysis, it was observed that the variations detected between individuals were mostly on the medial side of the bone.

Another factor affecting anatomical shape changes in bones is the function of the joint. Schaeffer (1947) defined the astragalus as a structure that articulates with the calcaneus, distal tibia, and cubonavicular, and it rotates in a space. The movements of this joint vary among individuals. These differences can be divided into anatomical, functional, and pathological. Various anatomical elements of the joint and tissues around the joint serve as restrictors of movement in the joint. Under the influence of physical weight, joints can undergo functional and structural changes that increase or decrease their mobility (Potekhina et al. 2018). Locomotor behavior (Vermeulen et al. 2022) may affect the shape of the bone. It was thought that the findings were consistent with the literature and that the shape variation may be more related to joint movement, while the variation determined according to races may be more related to genetics and developmental processes.

The study has some limitations in terms of ruminant species from which astragalus was collected. Therefore, bovine, ovis, and capras astragalus used in the study could not be compared with other ruminant species such as gazelle, mountain goat, and deer.

5. Conclusion

As a result, in the cross‐validation results, while bovine and ovis were completely grouped correctly, capra samples showed a low level of incorrect grouping. It was evaluated that the reason for the variation determined between bovine, ovis, and capra could be due to differences in posture, vital differences, locomotor behavior, body weight, and genetic structure differences. The results are important in terms of contributing to taxonomy, morphology, and zooarchaeology studies.

Author Contributions

Conceptualization: Yasemin Üstündağ, İftar Gürbüz, Yasin Demiraslan, and Özcan Özgel. Material collation: Yasemin Üstündağ and Özcan Özgel. Methodology and statistical analysis: Yasemin Üstündağ and Yasin Demiraslan. Resources: İftar Gürbüz and Yasin Demiraslan. Writing–original draft preparation: İftar Gürbüz and Yasemin Üstündağ. Writing—review and editing: İftar Gürbüz, Yasin Demiraslan, Yasemin Üstündağ, and Özcan Özgel. All authors have read and agreed to the published version of the manuscript.

Ethics Statement

Since the study materials (sheep, goat, and bovine astragalus) are existing slaughterhouse material, and no procedure is applied to live animals, according to the “Regulation on the Working Procedures and Principles of Animal Experimentation Ethics Committees”, there is no need for Ethics Committee approval.

Conflicts of Interest

The authors declare no conflicts of interest.

Peer Review

The peer review history for this article is available at https://publons.com/publon/10.1002/vms3.70368.

Funding: The authors received no specific funding for this work.

Data Availability Statement

The data that support the findings of this study are available from the corresponding author upon reasonable request.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

The data that support the findings of this study are available from the corresponding author upon reasonable request.

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