, Volume 138, Issue 1, pp 85–96 | Cite as

Variations in body shape of mountain habitat specialist Carabus croaticus and its sister species Carabus caelatus (Coleoptera: Carabidae) populations across Dinaric Alps

  • Željka Jambrošić VladićEmail author
  • Hugo A. Benítez
  • Alja Pirnat
  • Slavčo Hristovski
  • Lucija Šerić JelaskaEmail author
Original Article


There are abundant phenotypic variations of Carabus croaticus Dejean 1826, an endemic species of the Dinaric Alps, and its sister species Carabus caelatus Fabricius 1801, resulting in taxonomic inflation at intraspecific level, synonymy and inconsistency between relevant catalogues and checklists. The main aims of this research were to explore population structure based on morphology and geographic patterns of phenotypic variability, and compare the results with taxa described by nonmetric visual comparisons of morphological traits. Our study included 224 specimens of C. croaticus and 192 specimens of C. caelatus, covering most of their distributional range. Shapes of the pronotum and head were analysed using geometric morphometrics (GMM). Principal component and canonical variate analyses were used to characterize the main features of shape variation between populations and mountain ranges. GMM delimitated interspecific morphological variations but at the intraspecific level it showed many overlaps within populations for both species. Conducting the morphological analyses for the first time on most of the described phenotypic variants of studied species, we wanted to provide a new evidence for the possible solution of the taxonomic relations within these two endemic species by measuring body shape variability, and thus to enable a better understanding of the evolutionary processes in the Dinaric Mountains.


Balkan Peninsula Ground beetles Endemic species Geometric morphometrics Speciation Morphological variability 



We would like to thank Croatian Ecological Society and Plitvice National Park for financial support (Grant: 4492/16), B. Drovenik and P. Durbešić for the permission to work on the material from their collections, to Mrs Drovenik for her hospitality during our visits to their home and during long hours working on the collection; to L. Vladić, R. Vladić, A. Jakovac and S. Jelaska for all their help in the field and J. Skejo for constructive discussion on the manuscript. HB would like to thanks to the Universidad de Tarapacá Grant DGI-UTA 9719-17. The authors are grateful to two reviewers whose comments substantially improved the manuscript, and special thanks to dr. A. Cardini for valuable comments on the MS in particular for advices on GMM analyses.

Author contributions

LSJ, ZJV and AP designed the study; ZJV, LSJ, AP and SH collected the samples, ZJV performed morphometric measurements of sampled material and data analyses, ZJV and HAB performed statistical analysis; ZJV, HAB and LSJ wrote the manuscript; all authors contributed in improving the draft of the manuscript by adding valuable comments; LSJ supervised this work from the very beginning.


This study was funded by Croatian Ecological Society and Plitvice National Park (Grant number 4492/16, LSJ project leader).

Compliance with ethical standards

Data accessibility

The datasets generated during the current study are available in the figshare repository (doi:10.6084/m9.figshare.7415351). Details with the link will be added upon acceptance of publishing.

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed.

Supplementary material

435_2018_428_MOESM1_ESM.jpg (825 kb)
Online Resource 1 Canonical variate analysis showing pronotum shape variations for all subspecies (phenotypic variants) of (A) C. caelatus (black=C. caelatus cabuljensis; grey=C. caelatus caelatus; blue=C. caelatus dalmatinus; light blue=C. caelatus macretus; red= C. caelatus malissorum; purple= C. caelatus metalkanus; pink=C. caelatus procerus; orange= C. caelatus sarajevoensis; green= C. caelatus schreiberi; light green= C. caelatus volujakianus) and (B) C. croaticus (black=C. croaticus babinjensis; grey=C. croaticus bosnicus; blue=C. croaticus bosiljevici; light blue=C. croaticus kraetschmeri; red= C. croaticus croaticus; purple=C. croaticus droveniki, light purple= C. croaticus durmitorensis; pink=C. croaticus frankenbergeri; light pink= C. croaticus kobingeri; orange= C. croaticus ljubetensis; green= C. croaticus mediterraneus; light green= C. croaticus pretneri, brown= C. croaticus travnikanus; yellow=C. croaticus zepcensis) (JPG 824 KB)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.School of Natural Sciences, Architecture and MiningVaraždinCroatia
  2. 2.Departamento de Recursos Ambientales, Facultad de Ciencias AgronómicasUniversidad de TarapacáAricaChile
  3. 3.ZVERCE, Researching Beetles and DragonfliesKamnikSlovenia
  4. 4.Institute of Biology, Faculty of Natural Sciences and MathematicsSs. Cyril and Methodius UniversitySkopjeMacedonia
  5. 5.Department of Biology, Faculty of ScienceUniversity of ZagrebZagrebCroatia

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