Register      Login
Australian Mammalogy Australian Mammalogy Society
Journal of the Australian Mammal Society
RESEARCH ARTICLE (Open Access)

Mammal craniodental remains in owl pellets provide new tooth row measurements for the threatened Julia Creek dunnart (Sminthopsis douglasi)

A. M. Baker A B * , C. L. Charley https://orcid.org/0009-0004-5402-5294 A , D. Lockhart A and E. L. Gray https://orcid.org/0000-0001-7556-0908 A
+ Author Affiliations
- Author Affiliations

A School of Biology and Environmental Science, Queensland University of Technology, Brisbane, Qld, Australia.

B Biodiversity and Geosciences Program, Queensland Museum, South Brisbane, Qld, Australia.

* Correspondence to: am.baker@qut.edu.au

Handling Editor: Mark Eldridge

Australian Mammalogy 47, AM25004 https://doi.org/10.1071/AM25004
Submitted: 22 January 2025  Accepted: 19 February 2025  Published: 12 March 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing on behalf of the Australian Mammal Society. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License (CC BY-NC-ND)

Abstract

Archer’s 1981 morphological revision of Sminthopsis included only a few individuals of the newly described Sminthopsis douglasi. Here, we used craniodental material sourced from owl pellets collected in north-west Queensland, and additional skeletal material from Queensland Museum specimens, to compile a detailed series of tooth row measurements for S. douglasi (85 specimens, 14 locations). These were compared with existing measurements for co-occurring and geographically proximate congeners, namely, S. macroura, S. crassicaudata, S. murina and S. virginiae. Overall, Julia Creek dunnarts had consistently longer tooth-row measurements than those of the other dunnart species examined. This research will be useful for those seeking to confidently identify S. douglasi by using limited craniodental remains.

Keywords: barn owl, carnivorous marsupial, Dasyuridae, mandible, skull, species identification.

Introduction

When Archer conducted his landmark morphological revision of the dasyurid genus Sminthopsis in 1981, he was able to include only three specimens of the Julia Creek dunnart (Sminthopsis douglasi), which had been described just 2 years earlier by the same author (Archer 1979, 1981). Thereafter, the species was presumed extinct until new records were found in Queensland in the early 1990s (Woolley 1992). Today, Sminthopsis douglasi is listed as Vulnerable under the Environment Protection and Biodiversity Conservation Act (EPBC Act 1999). The species has proven elusive, but it has been sporadically detected using live (Elliott) trapping (Mifsud 1999; Bakker et al. 2024a), camera traps (Bakker et al. 2024b) and owl pellets (Woolley 2010; Lockhart 2024; Charley et al. 2025).

Woolley (2010) examined eastern barn owl (Tyto javanica delicatula) pellets collected in north-western Queensland between 1994 and 2001, and S. douglasi was identified from 22 (of 28) locations, usually with fewer than 10 individuals per site. More recently, Charley et al. (2025) assessed owl pellets collected from 2023 to 2024 at a single location where S. douglasi occurs, namely, Toorak station in north-western Queensland. Sminthopsis douglasi comprised up to 31% of the vertebrate material found in the pellets (including >220 individuals). However, neither study published measurements of S. douglasi tooth rows. This means that the main reference source to aid S. douglasi identification is still based on the comparative measurements of only three individuals from Archer’s earlier work (Archer 1981). Such small sample sizes could over- or under-estimate the mean, variance and range of species diagnostics because of the higher weighting attributed to each sample and the increased likelihood that (by chance) one or more samples may not be representative of a ‘typical’ member of the population.

Here, we aimed to fill this knowledge gap by making a series of (upper and lower tooth row) measurements for S. douglasi craniodental remains sourced from owl pellets used by Charley et al. (2025), other unpublished material, and registered specimens held at the Queensland Museum. We then compared our measurements with those previously taken of S. douglasi (and co-occurring and geographically proximate congeners, including S. macroura, S. crassicaudata, S. murina and S. virginiae) by Archer (1981) and synthesised the data to provide a more comprehensive reference source for identifying Sminthopsis from Queensland.

Materials and methods

Sminthopsis douglasi specimens

Owl pellets were collected from four sites in north-western Queensland as part of parallel monitoring (Charley et al. 2025) and distribution (Lockhart 2024) studies of S. douglasi (Table 1, Fig. 1). The craniodental material of S. douglasi retrieved from these pellets was retained and forms the bulk of specimens used in the present study. For full details of owl pellet dissection and processing, see Charley et al. (2025).

Table 1.S. douglasi specimens and locations.

Location (site symbol; N individuals)LatitudeLongitudeCollectorDate collectedSource
Toorak Woolshed, 40 km south of Julia Creek. Collection 1 (T; 48)21°01′58″S141°47′59″EGray, Mifsud, Baker13 Apr. 2023Owl pellets QUT
Toorak Woolshed, 40 km south of Julia Creek. Collection 2 (T; 8)21°01′58″S141°47′59″EGray18 July 2023Owl pellets QUT
Toorak Woolshed, 40 km south of Julia Creek. Collection 3 (T; 7)21°01′58″S141°47′59″EGray2 Oct. 2023Owl pellets QUT
Toorak Woolshed, 40 km south of Julia Creek. Collection 4 (T; 9)21°01′58″S141°47′59″EGray21 Apr. 2024Owl pellets QUT
Stamford Racecourse, Stamford Collection 1 (S; 2)21°14′56″S143°48′53″EGray, Mifsud, Baker12 Apr. 2023Owl pellets QUT
Stamford Racecourse, Stamford Collection 2 (S; 1)21°14′56″S143°48′53″EGray6 June 2023Owl pellets QUT
Woodsberry, via Stamford (W; 1)21°34′38″S143°53′03″EGray3 Oct. 2023Owl pellets QUT
Nelia, 48 km E of Julia Creek (N; 1)20°39′16″S142°12′18″EGray, Mifsud, Baker14 Apr. 2023Owl pellets QUT
Garomna, Julia Creek (G; 1)20°42′S141°53′EM Browne24 May 1933J5459 Male Paratype QM
Eureka, via Julia Creek (E1; 1)20°50′S141°48′EP Woolley24 Apr. 1992JM9805 Male QM
Bladensburg National Park, via Winton (B; 1)22°31′58″S143°02′46″ER Hobson2 Oct. 2001JM14810 Male QM
Ashton Station (A; 1)21°09′59″S144°44′59″EA Kutt25 Sep. 2000JM14336 QM
Edith Downs Station, south-west of Julia Creek (E3; 1)20°56′S142°12′EP Woolley17 Apr. 1992JM18108 Male QM
Lyrian Station, via Cloncurry (L1; 3)19°21′S141°11′EP WoolleyApr. 1991JM18139 Owl Pellet QM
Lyrian Station, via Cloncurry (L2; 1)19°28′S141°12′EP Woolley27 Apr. 1992JM18112 Male QM
Lyrian Station, via Cloncurry (L2; 1)19°28′S141°12′EP Woolley28 Apr. 1992JM18114 Male QM
Near Edith Downs, via Julia Creek (E2; 1)20°53′S142°12′EC TrittonOct. 1991JM18122 Male QM
Proa, via Nelia (P; 1)20°54′S142°08′EG Mifsud3 June 1998JM18601 Male QM
Proa, via Nelia (P; 1)20°54′S142°08′ED FyshJan. 1999JM18602 QM
Proa, via Nelia (P; 1)20°54′S142°08′EG Mifsud15 Nov. 1997JM18603 Male QM
North-east of Cloncurry (C; 1)20°26′54″S140°46′42″EN Godwin28 Aug. 2012JM19866 QM

Four collections were made from Toorak and two collections from Stamford Racecourse. The majority of specimens were taken from Toorak Collection 1, which represented a broader age range of individuals. Representative samples were taken from each of the subsequent three Toorak collections, each of which represented a maximum age of 6 months. For the three other owl pellet sites, there was a limited amount of material available, yielding representative samples at each location. Site symbols in column 1 match those in Fig. 1. QM indicates Queensland Museum (registration number is provided). Note: J5173, the holotype, could not be used in the analysis because it is a juvenile female.

Fig. 1.

Map of Queensland, Australia (on right), and the Mitchell Grass Downs Bioregion (on left), with Julia Creek dunnart (Sminthopsis douglasi) collection locations highlighted in yellow. Figure on the left is labelled as follows: L1, Lyrian Station; L2, Lyrian Station; C, north-east of Cloncurry; G, Goromna; N, Nelia; E1, Eureka; P, Proa; E2, near Edith Downs Station; T, Toorak; E3, Edith Downs Station; B, Bladensburg National Park; S, Stamford Racecourse; W, Woodsberry; and A, Ashton Station. Site symbols match those in column 1 of Table 1. Red and orange regions display the Government habitat distribution model for S. douglasi (Department of Climate Change, Energy, the Environment and Water 2024).


AM25004_F1.gif

Comparative measurements from all craniodental material of S. douglasi registered in the Queensland Museum were also included in this study, to improve the geographic coverage of specimens (Table 1, Fig. 1).

Other Sminthopsis species included

Two dunnart species are known to co-occur with S. douglasi in central and north-western Queensland grasslands, namely, S. macroura (stripe-faced dunnart) and S. crassicaudata (fat-tailed dunnart). Sminthopsis douglasi, which can reach 70 g in weight, is notably larger than either congener (which rarely exceed 25 g), and the size differences are also apparent in their craniodentaries (Archer 1981; Van Dyck et al. 2013). Two other congeners possess distributions geographically proximate to that of S. douglasi. Sminthopsis murina is found in varied wooded habitats of eastern Australia, extending inland into semiarid areas, but it is also small, rarely exceeding 28 g (Baker and Gynther 2023). However, another large dunnart, S. virginiae, is found in savanna woodland habitat to the north and east (Baker and Gynther 2023), and it is uncertain how far the S. douglasi range extends in this direction (Kutt 2003).

Thus, measurements for S. macroura, S. crassicaudata, S. murina, S. virginiae and comparative measurements for S. douglasi were taken from Archer (1981). We made no attempt to remeasure the craniodentaries used in Archer (1981) for either S. douglasi or congeners. An additional two registered Qld Museum S. virginiae specimens (JM4015; JM18722) were measured to increase the sample size for this species.

Measurements

Tooth row measurements were taken from both the left and right side of the craniodental material. Multiple t-tests comparing left and right tooth row measurements for S. douglasi and corrected using the Holm–Bonferroni method (Holm 1979) indicated that there was no significant difference between the sides for any measurement (see Appendix 1). However, only one side of each specimen was used in the analysis, as per Archer (1981); the left side was nominally chosen, because this side had more complete samples.

Five craniodental measurements (three skull and two mandible) were made to the nearest 0.1 mm with Mititoyo digimatic calipers.

Superscripts designate tooth number in the skull (e.g. M3), subscripts define the tooth number in the dentary (e.g. M3), and neither subscript nor superscript indicates a reference to both upper and lower teeth (e.g. M3), as per Archer (1981).

Skull: C1–M4; M1–M4; M1–M3 (see Fig. 2). These matched Archer (1981): e, f, g respectively.

  • C1–M4: length of maxillary tooth row: from anterior edge of C1 root to posterior edge of M4 root.

  • M1–M4: length of upper molar row: from anterior side of M1 paracone root to posterior side of M4 posterior root.

  • M1–M3: length of first three upper molars: from anterior side of M1 paracone root to posterior side of M3 metacone root.

Fig. 2.

Photograph of a S. douglasi skull from a Toorak owl pellet, showing tooth numbers and the position that the C1–M4; M1–M4; M1–M3 measurements (described in text) should be taken.


AM25004_F2.gif

Mandible: M1–M4; M1–M3 (see Fig. 3). These matched Archer (1981): m, n respectively.

  • M1–M4: length of lower molar row: from anterior face of the anterior root of M1 to the posterior edge of M4 talonid root.

  • M1–M3: length of the first three lower molars: from anterior face of the anterior root of M1 to the posterior edge of the M3 posterior root.

Fig. 3.

Photograph of a S. douglasi mandible from a Toorak owl pellet, showing tooth numbers and the position that the M1–M4; M1–M3 measurements (described in text) should be taken.


AM25004_F3.gif

Age of specimens

Only adult specimens were used because some measurements may be affected by bone lengths, which grow during adolescence (although fully emerged teeth do not grow and M1–M3 measurements are least affected by age). As per Archer (1981), adult specimens were designated by a fully descended P3 (described as P4 by Archer), such that P3 > P2 > P1 in crown height.

Analyses

The raw data from Archer (1981) were not available for all comparative dunnart species. Therefore, sample mean, standard error of the mean, standard deviation, coefficient of variation and measurement range were calculated to match Archer (1981), but no further analyses were undertaken.

Ethics

The project was conducted under the auspices of Queensland Department of Environment, Science and Innovation (DESI) Permit P-PTUKI-100171210 and QUT Research Ethics Permit 5154.

Results and discussion

We aimed to assess variation in craniodental (tooth-row) measures of a threatened dasyurid, the Julia Creek dunnart. We measured up to 85 S. douglasi individuals across 14 locations (Table 2, Fig. 1). As expected, the increased number of individual S. douglasi assessed meant that compared with Archer (1981), both minimum and maximum ranges of all tooth-row measures were extended (Table 2).

Table 2.Absolute measurements (in mm), including both those taken in the present study [denoted: Present Study Sminthopsis douglasi and Sminthopsis virginiae] and those adapted from Archer (1981) (Sminthopsis douglasi, Sminthopsis virginiae, Sminthopsis macroura, Sminthopsis crassicaudata and Sminthopsis murina).

Itemns.e.m.RangeσCV
Sminthopsis douglasi (Present Study)
 Skulle (C1–M4)3212.50.0811.8–13.30.430.03
f (M1–M4)857.00.026.6–7.50.180.03
g (M1–M3)856.10.025.7–6.40.170.03
 Mandiblesm (M1–M4)817.80.027.5–8.20.200.03
n (M1–M3)786.00.025.6–6.20.140.02
Sminthopsis douglasi (Archer 1981)
 Skulle (C1–M4)212.40.611.8–13.00.856.84
f (M1–M4)27.00.166.8–7.10.223.19
g (M1–M3)36.10.15.9–6.20.172.84
 Mandiblesm (M1–M4)38.00.127.8–8.20.202.50
n (M1–M3)36.10.046.0–6.10.071.15
Sminthopsis virginiae (Present Study)
 Skulle (C1–M4)211.10.4510.6–11.50.640.06
f (M1–M4)26.20.196.0–6.40.260.04
g (M1–M3)25.30.025.3–5.30.040.01
 Mandiblesm (M1–M4)16.5
n (M1–M3)15.0
Sminthopsis virginiae (Archer 1981)
 Skulle (C1–M4)2110.70.19.9–11.80.464.29
f (M1–M4)235.80.055.0–6.00.244.06
g (M1–M3)255.10.054.5–5.40.234.51
 Mandiblesm (M1–M4)246.50.046.0–6.90.203.08
n (M1–M3)264.90.054.4–5.00.190.91
Sminthopsis macroura (Archer 1981)
 Skulle (C1–M4)859.40.057.9–11.30.495.23
f (M1–M4)845.20.034.7–5.80.234.48
g (M1–M3)854.60.024.2–5.10.194.21
 Mandiblesm (M1–M4)845.90.035.6–6.40.244.14
n (M1–M3)864.40.033.9–4.80.327.28
Sminthopsis crassicaudata (Archer 1981)
 Skulle (C1–M4)709.30.048.7–10.00.303.24
f (M1–M4)715.00.024.7–5.50.183.60
g (M1–M3)734.50.024.2–4.90.163.48
 Mandiblesm (M1–M4)715.70.025.1–6.20.213.60
n (M1–M3)874.30.043.8–4.70.347.80
Sminthopsis murina (Archer 1981)
 Skulle (C1–M4)1149.80.058.7–11.30.495.03
f (M1–M4)1165.20.024.7–5.80.193.72
g (M1–M3)1214.60.024.3–5.30.194.12
 Mandiblesm (M1–M4)1205.80.025.2–6.50.213.54
n (M1–M3)1234.40.023.9–4.80.204.46

Abbreviations: n, sample size; , sample mean; s.e.m., standard error of the mean; σ, standard deviation; and CV, coefficient of variation. e, f, g, m, n are equivalent measures from Archer (1981).

However, despite the increased sample size, S. douglasi was still largest in size, and there was no overlap in ranges in any of the comparative measurements made from three co-occurring or geographically proximate congeners (S. macroura, S. crassicaudata and S. murina) (Fig. 4). Of measured congeners, in terms of size, S. virginiae most closely approached S. douglasi, with the former being only slightly smaller in several measures and marginally overlapping in one (C1–M4) (Fig. 4).

Fig. 4.

The absolute measurement range (mm) for each measured craniodental feature split across assessed species. Coloured bars represent the range of each species’ measurement, and light grey bars represent the mean ± standard deviation for the respective species. Black dots represent the mean of each measurement.


AM25004_F4.gif

In short, a dunnart specimen from Queensland is likely to be an S. douglasi if in the skull: M1–M3 ≥ 5.6 mm and C1–M4 is ≥ 11.8 mm; and in the dentary: M1–M4 ≥ 7.3 mm. If the specimen is a juvenile, the M4 may not be fully emergent and is typically inflected in relation to the rest of the dental row. In this case, M1–M3 measures should be used, because they are least likely to vary between age classes, and even in juveniles, the molars of S. douglasi are distinctively large.

Thus, the clear craniodental size differentiation of S. douglasi will aid confident identification of the species if present in owl pellet material collected in the grasslands of central and north-western Queensland, even if the skull and dentary are damaged, incomplete or the teeth are loose. See Appendix 2 for a more detailed description of the condition of the craniodental material sourced from owl pellets and some suggested preservation methods.

Variation in S. virginiae

Although we were able to measure only two further S. virginiae skulls, one of them (JM18722) fell outside the reported range for the species in Archer (1981) for one measurement, M1–M4 – 6.4 mm for JM18722, versus the maximum in upper range of 6.0 mm reported in Archer (1981). Despite the increase in maximum range, this was still less than the lower range for the comparative measurement for S. douglasi of 6.6 mm (Table 2).

Implications

This research has added significantly to the limited craniodental measurements available for S. douglasi. It should provide a useful resource for future studies assessing craniodental material to help inform the distributional range of this threatened species. The easiest and most accurate measurements proved to be C1–M4 and M1–M3/M4, and at least one of these (usually the latter) is likely to be possible for dunnarts found in eastern barn owl pellets. Researchers should refer to Archer (1981) when further details of comparative craniodental characters among dunnart species are required.

Supplementary material

Supplementary material is available online.

Data availability

The data that support this study are available in the article and accompanying online supplementary material. Raw data is available via Zenodo: 10.5281/zenodo.14708185.

Conflicts of interest

The authors declare that they have no conflicts of interest in this work.

Declaration of funding

This research was principally funded by Multicom Resources as part of an Offset Management Plan conceptualised by Multicom Resources and Epic Environmental and approved under the Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act) for the Saint Elmo Vanadium project (EPBC 2017/8007). Additional funding and resources were provided by QUT.

Acknowledgements

We are grateful to Heather Janetzki of the Queensland Museum for useful discussion and providing access to specimens held in the collection. We thank members of the Museum Vertebrate Zoology Laboratory for hosting AMB during December of 2024. We thank Mike Archer for providing access to a subset of the raw datasheets used in his 1981 systematic revision of Sminthopsis and Cameron Dodd for scanning them. We thank the various field assistants and landholders who collected owl pellets, and members of the Baker Mammal Ecology Lab for additional guidance and suggestions regarding the project. Comments from two anonymous reviewers helped us to improve the paper.

References

Archer, M. (1979). Two new species of Sminthopsis Thomas (Dasyuridae: Marsupalia) from northern Australia, S. butleri and S. douglasi. Supplement to The Australian Zoologist 20(2), 327-364.
| Google Scholar |

Archer, M. (1981). Results of the Archbold Expeditions. No. 104. Systematic revision of the marsupial Dasyurid genus Sminthopsis Thomas. Bulletin of the American Museum of Natural History 168(2), 61-224.
| Google Scholar |

Baker, A. M., and Gynther, I. C. (Eds) (2023). ‘Strahan’s Mammals of Australia.’ 4th edn. (Reed New Holland: Sydney, NSW, Australia.)

Bakker, A. H., Patterson, C. R., Mifsud, G., Reside, A. E., Fuller, S., and Baker, A. M. (2024a). Density of a cryptic Australian small mammal: the threatened Julia Creek dunnart (Sminthopsis douglasi). Ecology and Evolution 14(7), e11674.
| Crossref | Google Scholar |

Bakker, A. H., Schoenefuss, P., Mifsud, G., Fuller, S., and Baker, A. M. (2024b). Comparing methods of detecting an elusive dasyurid marsupial, the threatened Julia Creek dunnart (Sminthopsis douglasi), in central western Queensland, Australia. Ecology and Evolution 14, e70507.
| Crossref | Google Scholar |

Charley, C. L., Gray, E. L., and Baker, A. M. (2025). Owl pellet content analysis proves an effective technique to monitor a population of threatened Julia Creek dunnarts (Sminthopsis douglasi) throughout a native rodent plague. Ecology and Evolution 15, e70922.
| Crossref | Google Scholar |

Department of Climate Change, Energy, the Environment and Water (2024). Species of National Environmental Significance and selected marine and cetacean species – GDB. Available at https://fed.dcceew.gov.au/datasets/9d313bb078b9421ebebc835b3a69c470/about [accessed 18 September 2024].

EPBC Act (1999). ‘Environment Protection and Biodiversity Conservation Act.’ Available at https://www.legislation.gov.au/C2004 A00485/latest/versions

Holm, S. (1979). A Simple Sequentially Rejective Multiple Test Procedure. Scandinavian Journal of Statistics 6(2), 65-70. Available at http://www.jstor.org/stable/4615733

Kutt, A. S. (2003). New records of the Julia Creek dunnart Sminthopsis douglasi in central-north Queensland. Australian Zoologist 32(2), 257-260.
| Crossref | Google Scholar |

Lockhart, D. (2024). The hidden diet: determining the distribution of the threatened Julia Creek dunnart (Sminthopsis douglasi) using eastern barn owl (Tyto javanica delicatula) pellets. BSc Honours thesis, Queensland University of Technology, Australia

Mifsud, G. (1999). Ecology of the Julia Creek dunnart, Sminthopsis douglasi, (Marsupialia: Dasyuridae). MSc thesis, La Trobe University, Victoria, Australia.

Van Dyck, S., Gynther, I., and Baker, A. (Eds) (2013) ‘Field Companion to the Mammals of Australia’. (New Holland Publishers: Australia)

Woolley, P. A. (1992). New Records of the Julia Creek Dunnart, Sminthopsis douglasi (Marsupialia: Dasyuridae). Wildlife Research 19, 779-783.
| Crossref | Google Scholar |

Woolley, P. A. (2010). The Julia Creek dunnart and other prey of the barn owl in Mitchell grass downs of north-western Queensland. Memoirs of the Queensland Museum 55(1), 107-117.
| Google Scholar |

Appendix 1.t-tests comparing left and right tooth row measurements for Sminthopsis douglasi (corrected using the Holm–Bonferroni method; Holm 1979).


Testtd.f.P-valueadj P-value
e (C1–M4)−0.1455.120.891
f (M1–M4)0.23163.40.821
g (M1–M3)−0.12166.610.911
m (M1–M4)0.11159.570.921
n (M1–M3)1.38154.930.170.85

Appendix 2.Preservation of the dunnart skull and dentary sourced from eastern barn owl pellets.

Skull. The cranium was typically crushed and fragmented. Much of the remainder of the skull was also otherwise fragmented. However, usually the separated left and right maxilla and molar/premolar teeth were preserved. Occasionally, all teeth in the maxilla from C1 to M4 were preserved intact, although most of the time the C1 was missing. The premaxilla was usually present as a fragment in the pellet, although the incisor teeth were typically lost or fell out during sorting.

Dentary. the separated left and right mandibles and associated teeth were often preserved close to intact, although the incisor teeth, and often the C1, were typically lost or fell out during sorting. The tip of the dentary, especially at I1, was often damaged. At the posterior of the mandible, the tip of the ascending ramus, the articular condyle and the angular process were sometimes snapped or lost. The molar and premolar teeth of each mandible were usually intact, although the lower molar teeth were more easily displaced than those in the skull.

Note: loose teeth were secured using a dab of woodworking glue applied via a pin tip; the glue dries clear in about 10 min and will ensure the most important teeth for identification (the molars) are kept in place. If teeth in jaws removed from owl pellets are not secured by glue, at best, they will rock in their socket if pressed (at worst, they will fall out). In the former case, care needs to be taken when measuring to ensure the calipers just touch the outside ranges of each tooth and do not compress them, which would otherwise underestimate length measurements.