Presence of morphological integration and modularity of the forcipular apparatus in Lithobius melanops (Chilopoda: Lithobiomorpha: Lithobiidae).

The presence of morphological integration and modularity of the forcipular apparatus, despite its evolutionary significance, has not been analyzed in centipedes. This morphological structure has a crucial role in feeding and defense, thanks to its poisonous part (forcipules), which is important for catching the prey. The aims of our study were: i) to test the hypothesis of modularity of the forcipular apparatus in centipede Lithobius melanops; and ii) to investigate the influence of allometry on overall morphological integration in the aforementioned species using a geometric morphometric approach. The presence of fluctuating asymmetry was obtained by Procrustes ANOVA. Allometry was significant only for the symmetric component of the forcipular apparatus. The modularity hypothesis was not accepted, because the covariance coefficients for symmetric and asymmetric components were lower than 89.5% and 72.1% (respectively) of other RV coefficients obtained by a random contiguous partition of the forcipular apparatus. Results of the present study indicate that allometry does increase the level of morphological integration in the forcipular apparatus. According to our results, the forcipular coxosternite and forcipules could not be considered as separate modules; namely, they probably share similar developmental pathways and function in different forms of behavior and survival in L. melanops.

Morphological integration of the head capsule in the millipede Megaphyllum unilineatum (C. L. Koch, 1838) (Diplopoda: Julida): can different modules be recognized?

Covariation of multiple morphological traits and modularity have been widely studied in the field of evolutionary developmental biology. Subunits of a morphological structure can evolve separately from each other in a modular fashion. The aims of our study therefore were: i) to test the hypothesis of modularity in the dorsal part of the head capsule and the gnathochilarium separately during late postembryogenesis in the julidan millipede Megaphyllum unilineatum (C. L. Koch, 1838) using geometric morphometrics; and ii) to investigate the influence of allometry on overall morphological integration in the dorsal part of the head capsule and the gnathochilarium in the mentioned species. Individuals from different ontogenetic stadia (stadium VI - stadium XI) were included in the analyses. Significant influence of fluctuating asymmetry on the dorsal part of the head capsule shape was detected by Procrustes ANOVA. Regressions were significant for the symmetric component of both analysed morphological traits, while non-significant regression was detected for the asymmetric component of the head capsule's dorsal part. Hypotheses of modularity for the dorsal part of the head capsule and the gnathochilarium are rejected because our results indicate that a small proportion of alternate partitions has higher covariation between subsets of structure than between the hypothesized modules. Contrary to our expectations, results of the present study show that allometry does not increase the level of morphological integration in the dorsal part of the head capsule and the gnathochilarium in M. unilineatum. Based on the obtained results, we conclude that the dorsal part of the head capsule and the gnathochilarium are not composed of independent modules and that in the case of the capsule's dorsal part, developmental processes affect morphological integration in different ways at different levels of shape variation.

Sexual size and shape dimorphism in Brachydesmus troglobius Daday, 1889 (Diplopoda, Polydesmida).

Until now, morphological trait variation has been investigated in several millipede species using geometric morphometrics. The present study is the first attempt to explore sexual shape and size dimorphism (SShD and SSD) of morphological structures in Polydesmida. We here analyse antennal, head, and leg SShD and SSD in Brachydesmus troglobius Daday, 1889. Our results show that SSD exists in all of the analysed structures, while SShD is present only in the legs. In comparison with females, males possess longer and wider legs, as well as longer antennae and a shorter head. Contrary to previous findings in some Julida, in B. troglobiusSSD of the antennae and legs varies more than SShD in these morphological structures.

Life after the mother's hug: Late post-embryonic development of Cryptops parisi (Chilopoda: Scolopendromorpha: Cryptopidae).

Here we describe in detail the late post-embryonic development of the common European scolopendromorph centipede Cryptops parisi. Canonical variate analyses of two groups of external morphological characters, viz., cephalic capsule characters (head length, length of the anterior and posterior paramedian cephalic sutures) and coxopleuron surface characters (number of pores in the coxal pore-field, number of setae on the posterior coxopleuron edge, their number on the coxal pore-field, and their number posterior to the coxal pore-field) were conducted on a large sample of specimens collected from two localities in Serbia. Ten free-living stages are recognized: three pre-adult stages (adolescens I, II, and III) and seven adult stages (one maturus junior stage, four maturus, and two maturus senior stages). The fourth late post-embryonic stage is the first mature stage in both sexes. Sexual dimorphism in the aforementioned characters was not observed. Morphological variation of coxopleuron characters was more informative for the discrimination of developmental stages in Cryptops than the morphological variation of cephalic capsule characters.

Operculum shape variation in Theodoxus Montfort, 1810 (Gastropoda: Neritidae).

The operculum morphology of neritid snails of genus Theodoxus Montfort, 1810 yields important species-specific taxonomic characters. This study is the first attempt to describe morphological differences in snails based on operculum shapes using a geometric morphometric approach. We examined the variability of opercular shapes between sexes and among populations and species based on 91 opercula of adult specimens belonging to six populations of three species of Theodoxus from the central Balkan and the southern border of the Pannonian plain. There are no sex-related differences in operculum shape and size in the studied species. The presence of shape variations influenced by size (allometry) was confirmed for T. danubialis (C. Pfeiffer, 1828) indicating that allometry could be a component of the morphological variation of this species. At the intraspecific level, phenotypic plasticity of operculum shape was found in T. danubialis and T. fluviatilis (Linnaeus, 1758). Of particular interest is the lack of morphological variability in the rare and endangered T. transversalis (C. Pfeiffer, 1828). At the species level a clear morphological distinction of T. fluviatilis from T. danubialis and T. transversalis was detected, confirming that the operculum with a rib could be used as a species-specific morphological character. The main morphological differences between species are an outwardly-stretched opercular/apophysal rib and a shortened tip of the opercular plate in T. fluviatilis, compared to the same structures in T. danubialis and T. transversalis.