Sexual dimorphism in Apfelbeckia insculpta (L. Koch, 1867) (Myriapoda: Diplopoda: Callipodida)


  • Bojan S. Ilić University of Belgrade – Faculty of Biology, Institute of Zoology, Studentski Trg 16, 11000 Belgrade
  • Bojan M. Mitić University of Belgrade – Faculty of Biology, Institute of Zoology, Studentski Trg 16, 11000 Belgrade
  • Slobodan E. Makarov University of Belgrade – Faculty of Biology, Institute of Zoology, Studentski Trg 16, 11000 Belgrade


sexual dimorphism, adult body plan, Balkan Peninsula, evolutionary morphology, millipedes


Apfelbeckia insculpta (L. Koch, 1867) is one of the largest European millipedes and an endemic species of the Balkan Peninsula. We present data on sexual dimorphism in size and body proportions obtained from 179 adult specimens of this species from four caves in Serbia and one in Montenegro using univariate and multivariate morphometric techniques. Sexual dimorphism was apparent and female-biased for all measured characters, except for lengths of the antennae and the 24th leg pair (which were larger in males) and lengths of the first, second and fourth leg pairs, which exhibited small differences between sexes. Generally, females had significantly greater body size than males, while males expressed significantly greater values in traits that can be associated with mobility and copulation behavior. Also, we found significant variations in sexual size and body proportions dimorphism among analyzed populations. The influences of fecundity and sexual selection on the adult body plan in A. insculpta are discussed.

DOI: 10.2298/ABS160229060I

Received: February 29, 2016; Revised: May 5, 2016; Accepted: May 18, 2016; Published online: July 27, 2016

How to cite this article: Ilić B, Mitić B, Makarov S. Sexual dimorphism in Apfelbeckia insculpta (L. Koch, 1867) (Myriapoda: Diplopoda: Callipodida). Arch Biol Sci. 2017;69(1):23-33.


Download data is not yet available.


Teder T, Tammaru T. Sexual size dimorphism within species increases with body size in insects. Oikos. 2005;108(2):321-34.

Foellmer MW, Moya-Laraño J. Sexual size dimorphism in spiders: patterns and processes. In: Fairbairn DJ, Blanckenhorn WU, Székely T, editors. Sex, size and gender roles: evolutionary studies of sexual size dimorphism. Oxford: Oxford University Press; 2007. p. 71-81.

Stillwell RC, Blanckenhorn WU, Teder T, Davidowitz G, Fox C. Sex differences in phenotypic plasticity affect variation in sexual size dimorphism in insects: from physiology to evolution. Annu Rev Entomol. 2010;55:227-45.

Shine R. Sexual size dimorphism in snakes revisited. Copeia. 1994;1994:326-46.

Monnet JM, Cherry MI. Sexual dimorphism in anurans. Proc Biol Sci. 2002;269(1507):2301-7.

Lindenfors P, Gittleman JL, Jones KE. Sexual size dimorphism in mammals. In: Fairbairn DJ, Blanckenhorn WU, Székely T, editors. Sex, size and gender roles: evolutionary studies of sexual size dimorphism. Oxford: Oxford University Press; 2007. p. 16-26.

Székely T, Lislevand T, Figuerola J. Sexual size dimorphism in birds. In: Fairbairn DJ, Blanckenhorn WU, Székely T, editors. Sex, size and gender roles: evolutionary studies of sexual size dimorphism. Oxford: Oxford University Press; 2007. p. 27-37.

Webb TJ, Freckleton RP. Only half right: species with female-biased sexual size dimorphism consistently break Rensch’s rule. PloS One. 2007;2(9):e897.

Honek A. Intraspecific variation in body size and fecundity in insects: a general relationship. Oikos. 1993;66(3):483-92.

Preziosi RF, Fairbairn DJ, Roff DA, Brennan JM. Body size and fecundity in the water strider Aquarius remigis: a test of Darwin’s fecundity advantage hypothesis. Oecologia. 1996;108(3):424-31.

Fox CW, Czesak ME. Evolutionary ecology of progeny size in arthropods. Annu Rev Entomol. 2000;45:341-69.

Davidowitz G. Population and environmental effects on the size-fecundity relationship in a common grasshopper across an aridity gradient. J Orthoptera Res. 2008;17(2):265-71.

Salavert V, Zamora-Muñoz C, Ruiz-Rodriguez M, Soler JJ. Female-biased size dimorphism in a diapausing caddisfly, Mesophylax aspersus: effect of fecundity and natural and sexual selection. Ecol Entomol. 2011;36(3):389-95.

Navarro J, Kaliontzopoulou A, Gonzalez-Solis J. Sexual dimorphism in bill morphology and feeding ecology in Cory’s shearwater (Calonectris diomedea). Zoology. 2009;112(2):128-38.

Berns CM. The evolution of sexual dimorphism: understanding mechanisms of sexual shape differences. In: Moriyama H, editor. Sexual dimorphism. Rijeka: InTech; 2013. p. 1-16.

Pekár S, Martišová M, Bilde T. Intersexual trophic niche partitioning in an ant-eating spider (Araneae: Zodariidae). PloS One. 2011;6(1):e14603.

Roitberg ES. Variation in sexual size dimorphism within a widespread lizard species. In: Fairbairn DJ, Blanckenhorn WU, Székely T, editors. Sex, size and gender roles: evolutionary studies of sexual size dimorphism. Oxford: Oxford University Press; 2007. p. 143-53.

Stillwell RC, Morse GE, Fox CW. Geographic variation in body size and sexual size dimorphism of a seed-feeding beetle. Am Nat. 2007;170(3):358-69.

Lengkeek W, Didderen K, Côté IM, van der Zee EM, Snoek RC, Reynolds JD. Plasticity in sexual size dimorphism and Rensch’s rule in Mediterranean blennies (Blennidae). Can J Zool. 2008;86(10):1173-78.

Djordjević S, Djurakić M, Golubović A, Ajtić R, Tomović Lj, Bonnet X. Sexual size and body shape dimorphism of Testudo hermanni in central and eastern Serbia. Amphibia-Reptilia. 2011;32:445-58.

Fairbairn DJ. Allometry for sexual size dimorphism: pattern and process in the coevolution of body size in males and females. Annu Rev Ecol Syst. 1997;28:659-87.

Székely T, Freckleton RP, Reynolds JD. Sexual selection explains Rensch’s rule of size dimorphism in shorebirds. PNAS. 2004;101(3):12224-7.

Fairbairn DJ. Allometry for sexual size dimorphism: testing two hypotheses for Rensch’s rule in the water strider Aquarius remigis. Am Nat. 2005;166(S4):S69-S84.

Hopkin SP, Read HJ. The biology of millipedes. Oxford: Oxford University Press; 1992.

Wojcieszek JM, Simmons LW. Divergence in genital morphology may contribute to mechanical reproductive isolation in a millipede. Ecol Evol. 2013;3(2):334-43.

Sierwald P, Bond JE. Current status of the myriapod class Diplopoda (millipedes): Taxonomic diversity and phylogeny. Annu Rev Entomol. 2007;52:401-20.

Hoffman RL. Classification of the Diplopoda. Genève: Museum d'Histoire Naturelle; 1979.

Blower JG. Milipedes. Keys and notes for the identification of the species. 35. The Linnean Society of London and the Estuarine and Brackish-Water Sciences Association. London: E.J. Brill/Dr. W. Backhuys; 1985.

Hoffman RL, Payne JA. Diplopods as carnivores. Ecology. 1969;50:1096-8.

Stoev P, Sierwald P, Billey, A. An annotated world catalogue of the millipede order Callipodida (Arthropoda: Diplopoda). Zootaxa. 2008;1706:1-50.

Minelli A, Michlik P. Diplopoda - Reproduction. In: Minelli A, editor. The Myriapoda. Vol. 2, Treatise on zoology - Anatomy, taxonomy, biology. Leiden, Boston: Brill; 2015. p. 237-65.

VandenSpiegel D, Golovatch SI. A new millipede of the family Ammodesmidae found in central Africa (Diplopoda, Polydesmida). Zookeys. 2015;483:1-7.

Stoev P, Enghoff H. A revision of the millipede tribe Apfelbeckiini Verhoeff, 1900 (Diplopoda: Callipodida: Schizopetalidae). Steenstrupia. 2008;29(1):47-66.

Frederiksen SB, Petersen G, Enghoff H. How many species are there of Pachyiulus? A contribution to the taxonomy of Europe's largest millipedes (Diplopoda: Julida: Julidae). J Nat Hist. 2012;46(9-10):599-611.

Ilić BS, Tomić VT, Lučić LR, Mitić BM. Anamorphic development of Apfelbeckia insculpta (L. Koch, 1867) (Diplopoda: Callipodida: Schizopetalidae). Arch Biol Sci. 2016;68(2):445-50.

Berner D. Size correction in biology: how reliable are approaches based on (common) principal component analysis? Oecologia. 2011;166:961-71.

Zelditch ML, Swiderski DL, Sheets HD, Fink WL. Geometric morphometrics for biologists: a primer. San Diego: Elsevier Academic Press; 2004.

Rice WR. Analyzing tables of statistical tests. Evolution. 1989;43(1):223-5.

Rossolimo OL, Pavlinov IY. Sex-related differences in development, size, and proportions of the skull of martens Martes martes (Mammalia, Mustelidae). Bull Moscow Soc Nat Biol Ser. 1974;79:23-35. Russian.

Lovich JE, Gibbons JW. A review of techniques for quantifying sexual size dimorphism. Growth Develop Aging. 1992;56:269-81.

Willemsen RE, Hailey A. Sexual dimorphism of body size and shell shape in European tortoises. J Zool. 2003;260(4):353-65.

Jolicoeur P. The multivariate generalization of the allometry equation. Biometrics. 1963;19:497-99.

Son MH, Hughes RN. Sexual dimorphism of Nucella lapillus (Gastropoda: Muricidae) in North Wales, UK. J Moll Stud. 2000;66:489-98.

Shariffi M, Farsat H, Vaissi S. Sexual size dimorphism in Neurergus kaiseri (Caudata: Salamandridae) in south-western Zagros Mountains, Iran. Amphib Reptile Conserv. 2012;6(4):1-8.

Du W-G, Lü D. An experimental test of body volume constraint on female reproductive output. J Exp Zool. 2010;313A:123-8.

Heath J, Bocock KL, Mountford MD. The life history of the millipede Glomeris marginata (Villers) in north-west England. Symp Zool Soc Lond. 1974;32:433-62.

Baker GH. The post-embryonic development and life history of the millipede, Ommatoiulus moreletii (Diplopoda: Julidae), introduced in south-eastern Australia. J Zool. 1978;186:209-28.

Bhakat S, Bhakat A, Mukhopadhyaya MC. The reproductive biology and post-embryonic development of Streptogonopus phipsoni (Diplopoda: Polydesmoidea). Pedobiologia. 1989;33(1):37-47.

Rowe M. Copulation, mating system and sexual dimorphism in an Australian millipede, Cladethosoma clarum. Aust J Zool. 2010;58(2):127-32.

Enghoff H, Golovatch S, Short M, Stoev P, Wesener T. Diplopoda – Taxonomic overview. In: Minelli A, editor. The Myriapoda. Vol. 2. Treatise on zoology – Anatomy, taxonomy, biology. Leiden, Boston: Brill; 2015. p. 363-453.

Krause MA, Burghardt GM, Gillingham JC. Body size plasticity and local variation of relative head and body size sexual dimorphism in garter snakes (Thamnophis sirtalis). J Zool. 2003;261(4):399-407.

Chung K-H, Moon M-J. Microstructure of the adhesive pad in the millipede Orthomorphella pekuensis (Polydesmida: Paradoxosomatidae). Entomol Resarch. 2008;38(3):216-20.

Telford SR, Dangerfield JM. Mating tactics in the tropical millipede Alloporus uncinatus (Diplopoda: Spirostreptidae). Behavior. 1993;124(1):45-56.

Cooper MI, Telford S. Copulatory sequences and sexual struggles in millipedes. J Insect Behav. 2000;13(2):217-30.

Tanabe T, Sota T. Complex copulatory behavior and the proximate effect of genital and body size differences on mechanical reproductive isolation in the millipede genus Parafontaria. Am Nat. 2008;171(5):692-99.

Yasuda H, Dixon AFG. Sexual dimorphism in the two spot ladybird beetle Adalia bipunctata: developmental mechanism and its consequences for mating. Ecol Entomol. 2002;27(4):493-98.

Manton SM. The evolution of arthropodan locomotory mechanisms. Part 11. Habits, morphology and evolution of the Uniramia (Onychophora, Myriapoda, Hexapoda) and comparisons with the Arachnida, together with a functional review of uniramian musculature. Zool J Linn Soc. 1973;53:257-375.

Blanckenhorn WU, Preziosi RF, Fairbairn DJ. Time and energy constraints and the evolution of sexual size dimorphism: to eat or to mate? Evol Ecol. 1995;9(4):369-81.

Blanckenhorn WU. Behavioral causes and consequences of sexual size dimorphism. Ethology. 2005;111(11):977-1016.

Haacker U. Patterns of communication in courtship and mating behavior of millipedes (Diplopoda). Symp Zool Soc Lond. 1974;32:317-28.

Carey CJ, Bull CM. Recognition of mates in the Portuguese millipede Ommatoiulus moreletii. Aust J Zool. 1986;34(6):837-42.

Darwin C. The descent of man and selection in relation to sex. London: John Murray; 1871.

Savalli UM. Sexual selection. In: Fox CW, Roff DA, Fairbairn DJ, editors. Evolutionary ecology. Concepts and case studies. Oxford: Oxford University Press; 2001. p. 207-21.

Andersson M, Iwasa Y. Sexual selection. TREE. 1996;11(2):53-8.

Butler MA, Losos JB. Mutivariate sexual dimorphism, sexual selection, and adaptation in Greater Antillean Anolis lizards. Ecol Monogr. 2002;72(4):541-59.

Rutherford PL. Proximate mechanisms that contribute to female-biased sexual size dimorphism in an anguid lizard. Can J Zool. 2004;82(5):817-22.

Stillwell RC, Fox CW. Geographic variation in body size, sexual size dimorphism and fitness components of a seed beetle: local adaptation versus phenotypic plasticity. Oiokos. 2009;118(5):703-12.

King RB. Body size variation among island and mainland snake populations. Herpetologica. 1989;45(1):84-8.

Bronikowski AM. Experimental evidence for the adaptive evolution of growth rate in the garter snake Thamnophis elegans. Evolution. 2000;54(5):1760-7.

Badyaev AV, Hill GE. The evolution of sexual dimorphism in the house finch. I. Population divergence in morphological covariance structure. Evolution. 2000;54(5):1784-94.

Enghoff H. The size of a millipede. In: Meyer E, Thaler K, Schedl W, editors. Advances in Myriapodology. Proceedings of the 8th International Congress of Myriapodology, Innsbruck, 15-20 July 1990. Ber nat-med Verein Innsbruck, 10. Innsbruck: Universtitätsverlag Wagner; 1992. p. S47-S56.




How to Cite

Ilić BS, Mitić BM, Makarov SE. Sexual dimorphism in Apfelbeckia insculpta (L. Koch, 1867) (Myriapoda: Diplopoda: Callipodida). Arch Biol Sci [Internet]. 2016Aug.19 [cited 2024Jun.18];69(1):23-3. Available from:




Most read articles by the same author(s)

Similar Articles

You may also start an advanced similarity search for this article.