Lucio Bonato

Department of Biology - University of Padova

via U.Bassi 58b I-35131 Padova (Italy) tel. 0039 049 8277487

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geophilomorph centipedes

evolution and diversity


Geophilomorph centipedes

are a group of terrestrial arthropods 

and a major component of soil ecosystems throughout the world.

These tiny predators are specialised

to carry on a secretive life inside the soil.

Knowledge on their evolution, diversity and ecology

is well behind that of many other animal groups.





Within extant geophilomorphs, mecistocephalids

comprise a very distinct lineage

retaining quite primitive features in morphology and behaviour.

Puzzling is the fact that, as a rule,

each species is invariant in the number of trunk segments

but this number frequently changed during evolution.



Geophilids comprise one of the most diverse groups

within the geophilomorphs,

with extraordinarily specialised traits 

for living among soil grains.

However, their evolutionary history still remains unclear.

Mediterranean Geophilomorphs


The Mediterranean basin

is a major hotspot of diversity for geophilomorphs.

Interesting patterns of geographical variation

and fascinating regional radiations are emerging.

However, the current taxonomy is inconsistent and unsatisfactory:

many taxa are in need of revision

...and a lot of unknown species are still to dig out. 



selected publications


Comparative analysis of diet 

in syntopic geophilomorph species (Chilopoda, Geophilomorpha) 

using a DNA-based approach 

Bortolin F., Fusco G., Bonato L. 2018.

Soil Biology and Biochemistry, 127: 223-229.


How many species of centipedes 

coexist in temperate forests

Estimating local species richness 

of Chilopoda in soil coenoses 

of the South-Eastern Prealps

Peretti E., Bonato L., 2018.
European Journal of Soil Biology

89: 25-32.



New insights into Plutonium, 

one of the largest and least known European centipedes (Chilopoda): distribution, evolution and morphology


Bonato L., Orlando M., Zapparoli M., Fusco G., Bortolin F. 2017.

Zoological Journal of the Linnean Society, 180: 887909.


Evolution of Strigamia centipedes (Chilopoda): a first molecular assessment of phylogeny and divergence times

Bonato L., Bortolin F., Drago L., Orlando M., Danyi L. 2017.
Zoologica Scripta, 46: 486-495.




an interactive identification tool 

for the geophilomorph centipedes 

of Europe


Bonato L., Minelli A., Lopresti M., Cerretti P.


Zookeys, 443: 1-9.

article  -   Chilokey

The phylogenetic position of Dinogeophilus

and a new evolutionary framework 

for the smallest epimorphic centipedes


Bonato L., Minelli A., Drago L., Pereira L.A.


Contributions to Zoology, 84: 237-253.






Phylogeny of Geophilomorpha 

inferred from 

new morphological and molecular evidence


Bonato L., Drago L., Murienne J. 2014. Cladistics, 30: 485-507.







Geophilomorph centipedes

from the Cretaceous amber of Burma


Bonato L., Edgecombe G.D., Minelli A. 2014.

Palaeontology, 57: 97-110.





 Morphology and diversity of the forcipules in Strigamia centipedes


Maruzzo D., Bonato L. 2014.

Arthropod Structure and Development, 43: 17-25








Treatise on Zoology - Anatomy, Taxonomy, Biology The Myriapoda - Volume 1


Minelli A., ed., 2011.






The centipede genus Clinopodes:

a reassessment of species diversity and distribution, with a new species from the Maritime Alps


Bonato L., Iorio E., Minelli A. 2011. Zoosystema, 33: 175-205





Pincer-like claws in centipedes:

multiple evolutionary origin

of similar form and serial pattern


Bonato L., Drago L., Minelli A. 2011. Zoomorphology, 130: 17-29





Morphology and phylogeny of Dicellophilus,

a centipede genus with highly

disjunct distribution


Bonato L., Danyi L., Minelli A. 2010. 

Zoological Journal of the Linnean Society, 158: 501-532





Geophilomorph centipedes

in the Mediterranean region:

revisiting taxonomy opens

new evolutionary vistas


Bonato L., Minelli A. 2009. 

Soil Organisms, 81: 489-503




Diversity in the maxillipede dentition

of Mecistocephalus centipedes,

with the description of a new species

with unusually elongate denticles


Bonato L., Minelli A.

2009. Contributions to Zoology, 78: 85-97.



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


Leśniewska M., Bonato L.,  Minelli A., Fusco G.


Arthropod Structure and Development, 38: 417-426






A geophilomorph centipede 

from La Buzinie amber

(Late Cretaceous: Cenomanian), SW France


Edgecombe G.D., Bonato L., Minelli A. 2009. Geodiversitas, 31: 5-15.



Morphology, taxonomy and distribution

of Diphyonyx n. gen.,

a lineage of geophilid centipedes

with unusually shaped claws


Bonato L., Zapparoli M., Minelli A. 2008. European Journal of Entomology, 105: 343354.




a hitherto unrecognized lineage

of western Palaearctic centipedes

with unusual diversity

in body size and segment number


Bonato L., Minelli A. 2008

Zoological Journal of Linnean Society, 153: 253-286.






The Mecistocephalidae

of the Japanese and Taiwanese islands


Uliana M., Bonato L., Minelli A. 2007

Zootaxa, 1396: 1-84.




The European centipedes hitherto referred

to Eurygeophilus, Mesogeophilus and Chalandea:

taxonomy, distribution

and geographic variation 

of the segment number


Bonato L., Barber A., Minelli A. 2006

Journal of Natural History, 40: 415-438.





The centipede genus Mecistocephalus

in the Indian Peninsula


Bonato L., Minelli A., 2004

Tropical Zoology, 17: 15-63.




Evolutionary trends and patterns

in centipede segment number

based on a cladistic analysis

of Mecistocephalidae


Bonato L., Foddai D., Minelli A., 2003

Systematic Entomology, 28: 539-579.



Phylogeny and systematics of the Arrupinae

with the description of a new dwarfed species


Foddai D., Bonato L., Pereira L., Minelli A., 2003

Journal of Natural History, 37: 1247-1267.





Parental care 

in Dicellophilus carniolensis:

new behavioural evidence with implications

for the higher phylogeny of centipedes


Bonato L., Minelli A., 2002

Zoologischer Anzeiger, 241: 193-198.




Increase by duplication 

and loss of invariance of segment number

in the centipede

Mecistocephalus microporus


Bonato L., Foddai D., Minelli A., 2001

Italian Journal of Zoology, 68: 345-352.



full list of publications in progress...




what do centipedes eat in the wild?

diet analysis 

by molecular identification 

of prey remains in the gut













ecological and geographical differentiation

in Strigamia in the Italian region:

hints from molecules and anatomy








new funtional morphologies

discovered in Sardinia and Cyprus:

neglected life in the Mediterranean islands