Lucio Bonato

A common terminology for the external anatomy of centipedes (Chilopoda)

Abstract A common terminology for the external morphological characters of centipedes (Chilopoda) is proposed. Terms are selected from the alternatives used in the English literature, preferring those most frequently used or those that have been introduced explicitly. A total of 330 terms are defined and illustrated, and another ca. 500 alternatives are listed.

Trunk anomalies in the centipede Stigmatogaster subterranea provide insight into late-embryonic segmentation

We describe and analyze naturally occurring anomalies in the segmental structures of the trunk in an isolated population of the geophilomorph centipede Stigmatogaster subterranea. Recorded anomalies include mispaired tergites, shrunk segments, variously deformed sclerites, bifurcated trunk, and defects of spiracles and sternal pore areas. One specimen has a perfect segmentally patterned trunk, but with an even number of leg-bearing segments, representing the first record of such a phenotype in adult centipedes. We interpret these anomalies as the effects of perturbation of specific morphogenetic processes in trunk segmentation, occurring at different embryonic stages. The variety of segmental anomalies found in this population provides insights into the developmental process of segmentation and its evolution in geophilomorph centipedes. Variation in dorsal mispairing anomalies demonstrates that segments, as traditionally defined in arthropod morphology, are not the effective developmental units throughout embryogenesis.

Inferring the degree of incipient speciation in secondary contact zones of closely related lineages of Palearctic green toads ( Bufo viridis subgroup)

Reproductive isolation between lineages is expected to accumulate with divergence time, but the time taken to speciate may strongly vary between different groups of organisms. In anuran amphibians, laboratory crosses can still produce viable hybrid offspring >20 My after separation, but the speed of speciation in closely related anuran lineages under natural conditions is poorly studied. Palearctic green toads (Bufo viridis subgroup) offer an excellent system to address this question, comprising several lineages that arose at different times and form secondary contact zones. Using mitochondrial and nuclear markers, we previously demonstrated that in Sicily, B. siculus and B. balearicus developed advanced reproductive isolation after Plio-Pleistocene divergence (2.6 My, 3.3–1.9), with limited historic mtDNA introgression, scarce nuclear admixture, but low, if any, current gene flow. Here, we study genetic interactions between younger lineages of early Pleistocene divergence (1.9 My, 2.5–1.3) in northeastern Italy (B. balearicus, B. viridis). We find significantly more, asymmetric nuclear and wider, differential mtDNA introgression. The population structure seems to be molded by geographic distance and barriers (rivers), much more than by intrinsic genomic incompatibilities. These differences of hybridization between zones may be partly explained by differences in the duration of previous isolation. Scattered research on other anurans suggests that wide hybrid zones with strong introgression may develop when secondary contacts occur <2 My after divergence, whereas narrower zones with restricted gene flow form when divergence exceeds 3 My. Our study strengthens support for this rule of thumb by comparing lineages with different divergence times within the same radiation.

Phylogeny of Geophilomorpha (Chilopoda) inferred from new morphological and molecular evidence

To address the phylogenetic relationships of the centipede order Geophilomorpha (more than 1000 species), we have reinterpreted and expanded the knowledge on their morphological disparity, and have doubled the amount of molecular data available. We performed maximum parsimony and maximum likelihood analyses, using 195 phylogenetically informative morphological characters for 80 species, and DNA sequences of 28S, 18S, 16S rRNA and COI for up to 48 species. We found strong support for the monophyly of Geophilomorpha, the basal dichotomy between Adesmata and Placodesmata = Mecistocephalidae, and the basal dichotomy within Adesmata between two clades that are recognized here as superfamilies Himantarioidea and Geophiloidea. With respect to the families currently in use, Himantarioidea comprises three well supported clades corresponding to (i) Oryidae, (ii) Himantariidae, and (iii) Schendylidae s.l. including Ballophilidae; Geophiloidea comprises another three supported clades corresponding to (iv) a new family Zelanophilidae, (v) Gonibregmatidae s.l. including Eriphantidae and Neogeophilidae, and (vi) Geophilidae s.l. including Aphilodontidae, Dignathodontidae, Linotaeniidae, and Macronicophilidae.

Phylogeny and systematics of the Arrupinae (Chilopoda Geophilomorpha Mecistocephalidae) with the description of a new dwarfed species

We performed a cladistic analysis of the subfamily Arrupinae, based on morphological data (23 characters) from all the 16 species we recognize in the group. Based on this analysis, the following taxonomic and nomenclatural novelties are introduced: Nannarrup hoffmani n. g. n. sp. is described from introduced specimens of unknown origin found living in New York, USA; Agnostrup n. g. (type species Krateraspis striganovae Titova, 1975) is created to include A. striatus (Takakuwa, 1949) (from Taiwanella), A. paucipes (Miyosi, 1955) (from Taiwanella) and A. striganovae (Titova, 1975) (from Krateraspis), all new combinations; Nodocephalus areolatus Shinohara, 1957, N. asiaticus Titova, 1975 and Prolamnonyx mamaevi Titova, 1975 are moved to the genus Arrup (three more new combinations); Partygarrupius Verhoeff, 1939, formerly a subgenus of Tygarrup Chamberlin, 1914, is raised to the status of genus. The peculiar morphology of Nannarrup hoffmani is analysed as a possible instance of miniaturization.

Parental Care in Dicellophilus carniolensis (C. L. Koch, 1847): New Behavioural Evidence with Implications for the Higher Phylogeny of Centipedes (Chilopoda)

Abstract Parental care in Dicellophilus carniolensis (Chilopoda Geophilomorpha Mecistocephalidae) is described for the first time: the female coils around the brood with the dorsal surface outwards, as do the females of other centipede orders (Craterostigmomorpha and Scolopendromorpha) but at variance with the remaining geophilomorphs, whose brooding females remain coiled with the ventral surface outwards. The available information on this behaviour in centipedes is reviewed and interpreted in the context of the phylogeny of the group, based on morphological and molecular characters, and also in relation to the evolution of the sternal glands, a feature common to most Geophilomorpha. The evolutionary transition from brooding female with dorsal side outwards (plesiomorphic state) to brooding female with ventral side outwards (apomorphic state) was most likely associated with the evolution of defensive glands on the ventral surface of the female. The behaviour observed in D. carniolensis is probably common to the whole of Mecistocephalidae, thus supporting the basal position of this clade within the Geophilomorpha.

Evolution of the melanistic colour in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian Prealps

Abstract In order to reconstruct the evolution of the body colour within the Salamandra atra group, a phylogenetic analysis of the mitochondrial DNA D‐loop sequence was performed on seven populations of this group, including all three chromatically different forms and all known nominal subspecies. Also included were four outgroup species. Three major mitochondrial lineages were found within S. atra: (i) the invariably yellow‐patched S. a. auro‐rae lineage from the Altopiano dei Sette Comuni (Venetian Prealps); (ii) a genetically distinct lineage represented by a population from the Pasubio massif (Venetian Prealps), in which the yellow patches were either limited or completely lacking; (iii) a lineage comprising all fully melanistic populations from the Alps to the Dinarides, which were assigned to S. a. atra and also included the nominal S. a. prenjensis. Different phylogenetical reconstruction methods and statistical tests supported the hypothesis that the lineage of S. a. aurorae was the sister group to both remaining lineages of S. atra. Within these latter lineages, S. a. pa‐subiensis emerged as the sister group to all populations of the S. a. atra lineage. This phylogenetic reconstruction suggests that the yellow‐patched colour is the ancestral condition in S. atra and the evolutionary transition to fully melanistic colour occurred probably through an intermediate reduction in the extension of the patches. Therefore, the fully melanistic colour of S. lanzai, which is probably not strictly related to S. atra, has an independent origin. The population from the Pasubio massif is here described as Salamandra atra pasubiensis n. subsp.

A geophilomorph centipede (Chilopoda) from La Buzinie amber (Late Cretaceous, Cenomanian), SW France

ABSTRACT The first geophilomorph centipede to be documented from Mesozoic amber and the second Mesozoic member of the order is described as Buziniphilus antiquus n. gen., n. sp. It is represented by a single, probably immature specimen from Early Cenomanian amber at La Buzinie, Champniers, Charentes, France. Buziniphilus n. gen. is most probably a member of either Schendylidae or Geophilidae, though documentation of the labrum and mandibles is required to make a definitive familial assignment. Referral of Buziniphilus n. gen. to the crown-group Adesmata, together with a reinterpretation of the structure of the forcipulae in the Jurassic Eogeophilus Schweigen & Dietl, 1997, reinforces the modern aspect of Mesozoic chilopods that had been indicated by Cretaceous scutigeromorph and scolopendromorph fossils.

The centipede genus Clinopodes C. L. Koch, 1847 (Chilopoda, Geophilomorpha, Geophilidae): reassessment of species diversity and distribution, with a new species from the Maritime Alps (France)

Bonato L., lorio É. & Minelli A. 2011. — The centipede genus Clinopodes C. L. Koch, 1847 (Chilopoda, Geophilomorpha, Geophilidae): reassessment of species diversity and distribution, with a new species from the Maritime Alps (France). Zoosystema 33 (2): 175–205. ABSTRACT Diagnosis, circumscription, species-level taxonomy and geographical occurrence of the genus Clinopodes C. L. Koch, 1847 (Chilopoda, Geophilomorpha, Geophilidae) are revised by integrating published information with new evidence on its morphological diversity. Clinopodes is clearly distinguishable from all other geophilid lineages by a combination of features mainly in the forcipular segment, the trunk sternites and the ultimate legs. It is widespread in south-eastern Europe, from the Alps to the Caucasus. A total of 10 species are recognized, mainly differing in minor features of the forcipular coxosternite and the arrangement of sternal and coxal pores: C. carinthiacus (Latzel, 1880) n. stat.; C. caucasicus (Selivanov, 1884) n. comb. formerly Geophilus caucasicus; C. escherichii (Verhoeff, 1896); C. flavidus C. L. Koch, 1847; C. intermedius Dărăbanţu & Matic, 1969; C. latisternus (Attems, 1947) n. comb. from Pleurogeophilus latisternus; C. rodnaensis (Verhoeff, 1938); C. skopljensis (Verhoeff, 1938); C. verhoeffi n. nom. (for Geophilus flavidus porosus Verhoeff, 1934); C. vesubiensis n. sp., Geophilus lindbergi (Loksa, 1971), n. comb. formerly Clinopodes lindbergi, is demonstrated to have been classified erroneously under Clinopodes. Clinopodes vesubiensis n. sp. is described from a limited area in the southern Maritime Alps, at the western border of the entire range of the genus, and disjunct from the morphologically closest species C. carinthiacus, suggesting that the history of differentiation and colonization of the genus within the Alps has been more complex than previously thought.

The European centipedes hitherto referred to Eurygeophilus, Mesogeophilus, and Chalandea (Chilopoda, Geophilomorpha): taxonomy, distribution, and geographical variation in segment number

We have carried out a comprehensive revision of the European centipede species currently assigned to Eurygeophilus, Mesogeophilus, and Chalandea. This was based on comparative morphological study of specimens from throughout the range of the group and representative of the different nominal taxa, as well as a critical evaluation of all relevant literature. We consider this group to be represented by a single genus Eurygeophilus Verhoeff, 1899 [ = Geophilus (Mesogeophilus) Verhoeff, 1901, n. syn.; = Chalandea Brölemann, 1909, n. syn.] including two morphologically clearly distinguished species, Eurygeophilus multistiliger (Verhoeff, 1899) [ = Eurygeophilus multistiliger velmanyensis Brolemann, 1926, n. syn.] and Eurygeophilus pinguis (Brölemann, 1898) [ = Geophilus (Mesogeophilus) baldensis Verhoeff, 1901, n. syn.; = Chalandea cottiana Verhoeff, 1938; = Chalandea cottiana var. castrensis Manfredi, 1948; = Chalandea scheerpeltzi Attems, 1952, n. syn.]. On the basis of analysis of both published and new records these two species appear to be geographically vicariant: E. multistiliger occurs mainly in Mediterranean woodlands of southern regions (the western part of the Iberian Peninsula, eastern Pyrenees and Sardinia), whereas E. pinguis occurs mainly in temperate deciduous woodlands in montane and more northern regions (a limited area in Great Britain, the Cantabrian Mountains, most of the western and central Pyrenees, Corsica and the entire Alps). Despite morphological uniformity throughout its range, E. pinguis shows a consistent geographical pattern in variation of the number of segments, the modal values being different between the three major areas: (1) Pyrenees, Cantabrian Mountains and Great Britain, (2) western and central Alps, and (3) eastern Alps.