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Primera cita de la Familia Polyzoniidae Newport, 1844 (Diplopoda, Polyzoniida) en la Península Ibérica

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Los milpiés (Diplopoda) son un grupo de artrópodos terrestres con una alta diversidad específica a escala global. En el mundo están citadas unas 12,000 especies distribuidas en 16 órdenes y 145 familias. La familia Polyzoniidae está representada en Europa por tres especies, siendo Polyzonium germanicum Brandt, 1837 la que presenta una distribución más amplia. Hasta la fecha no existen citas en la Península Ibérica para esta familia. Polyzonium germanicum habita fundamentalmente en suelos de bosques húmedos o muy húmedos, ya sean de coníferas, de frondosas o bosques mixtos. La presente nota describe una población de P. germanicum localizada en suelos de un hayedo del norte de la provincia de Barcelona (España). Este registro representa la primera cita de la especie P. germanicum y de la familia Polyzoniidae para la fauna ibérica.
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Graellsia, 77(1): e129
enero-junio 2021
ISSN-L: 0367-5041
https://doi.org/10.3989/graellsia.2021.v77.279
ABSTRACT
Millipedes (Diplopoda) are a group of terrestrial arthropods with high specific diversity at a global scale. There
are 12,000 species worldwide, distributed in 16 orders and 145 families. The family Polyzoniidae is represented
in Europe by three species, being Polyzonium germanicum Brandt, 1837 the one with the widest distribution. To
date there are no records in the Iberian Peninsula for this family. Polyzonium germanicum mainly inhabits humid
or very humid soils, either in coniferous, deciduous or mixed forests. This note describes a population of P. ger-
manicum in soils of a beech forest located in the north of Barcelona province (Spain). This report represents the
first record of P. germanicum and the family Polyzoniidae in the Iberian Peninsula.
Keywords: Millipedes, Polyzonium germanicum, Cadí Moixeró Natural Park, Gresolet forest, beech forest.
RESUMEN
Primera cita de la Familia Polyzoniidae Newport, 1844 (Diplopoda, Polyzoniida) en la Península
Ibérica
Los milpiés (Diplopoda) son un grupo de artrópodos terrestres con una alta diversidad específica a escala
global. En el mundo están citadas unas 12,000 especies distribuidas en 16 órdenes y 145 familias. La familia
Polyzoniidae está representada en Europa por tres especies, siendo Polyzonium germanicum Brandt, 1837 la
que presenta una distribución más amplia. Hasta la fecha no existen citas en la Península Ibérica para esta familia.
Polyzonium germanicum habita fundamentalmente en suelos de bosques húmedos o muy húmedos, ya sean de
coníferas, de frondosas o bosques mixtos. La presente nota describe una población de P. germanicum localizada
en suelos de un hayedo del norte de la provincia de Barcelona (España). Este registro representa la primera cita
de la especie P. germanicum y de la familia Polyzoniidae para la fauna ibérica.
Palabras clave: Milpiés, Polyzonium germanicum, Parc Natural del Cadí Moixeró, bosc de Gresolet, hayedo.
Recibido/Received: 5/05/2020; Aceptado/Accepted: 13/11/2020; Publicado en línea/Published online: 21/05/2021
Cómo citar este artículo/Citation: Mateos, E. 2021. First record of the Family Polyzoniidae Newport, 1844 (Diplopoda,
Polyzoniida) in the Iberian Peninsula. Graellsia, 77(1): e129. https://doi.org/10.3989/graellsia.2021.v77.279
Copyright: © 2021 SAM & CSIC. This is an open-access article distributed under the terms of the Creative Commons
Attribution 4.0 International (CC BY 4.0) License.
Notas / Notes
First record of the Family Polyzoniidae Newport, 1844 (Diplopoda, Polyzoniida)
in the Iberian Peninsula
Eduardo Mateos
Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals; Facultat de Biologia; Universitat de Barcelona; Avinguda Diagonal 643,
08028 Barcelona; Spain.
Email: emateos@ub.edu – ORCID iD: https://orcid.org/0000-0001-9741-5744
et al., 2012), distributed in 16 orders and 145 fami-
lies (Shelley, 2007). Of this astonishing diversity
9 orders, 48 families and well over 1,500 species are
currently described from Europe (Kime & Enghoff,
2017). The family Polyzoniidae Newport, 1844,
Millipedes (Class Diplopoda) are terrestrial ar-
thropods that belong to the subphylum Myriapoda.
There are probably as many as 80,000 millipede
species worldwide (Sierwald & Bond, 2007), but
around 12,000 have been described so far (Brewer
2 Notas / Notes
Graellsia, 77(1), junio 2021, e129 — ISSN-L: 0367-5041 – https://doi.org/10.3989/graellsia.2021.v77.279
in the beech forest at 1,300 m above sea level (posi-
tion 42°15’32.5” N, 1°42’47.2” E). The ve young
specimens are preserved in absolute ethanol and
deposited at the E. Mateos collection (Departament
de Biologia Evolutiva, Ecologia i Ciències Ambien-
tals, Facultat de Biologia, Universitat de Barcelona).
The ve adult specimens (2  and 3 ) are pre-
served in 70% ethanol and deposited at the Centre
de Recursos de Biodiversitat Animal (Facultat de
Biologia, Universitat de Barcelona) with collection
codes CRBA-92751 to CRBA-92755. The Gresolet
beech forest represents the southernmost locality of
the western European population of the species and
the rst record for the Iberian Peninsula. The forest is
crossed by the Mulleres stream and is distributed in
a range of altitudes between 1,000 and 2,000 m. The
area is characterized by a continental climate with
Mediterranean inuence, with relatively high pre-
cipitation and soft temperatures. The Gresolet forest
host a mosaic of plant communities including oak
grove (Quercus pubescens Willd.), beech forest (of
Fagus sylvatica L.), r forest (of Abies alba Mill.)
and pine forests (of Pinus sylvestris L. and Pinus
mugo Turra).
Polyzonium germanicum is a distinctive rather
slug-like millipede, some 5 to 18 mm in length, with a
yellowish to orange brown body composed of 35–60
segments and a very small and characteristic triangu-
lar head (Fig. 2A); when curled into a spiral it is said
to resemble beech bud scales (British Myriapod &
Isopod Group, 2020; Enghoff et al., 2015). In P. ger-
manicum (as in all Polyzoniida) males and females
are externally very similar. Only the transformation of
legs-9 and legs-10 into gonopods in males (in stadium
IV) allows differentiating the sexes (Couret & David,
1985). Polyzonium germanicum is a liquid feeder spe-
cies (Couret, 1985) and very hygrophilous in its habi-
tat requirements (David, 1983). Forests with thick hu-
mus layers and high humidity levels around the year is
the preferred habitat of the species (David & Vannier,
1995). These forests may be coniferous, mixed or de-
ciduous, with dense, open or clearing structure, but al-
ways with wet (or even waterlogged) soils (see Kime
& Enghoff, 2011). Characteristically P. germanicum
females brood the eggs (Enghoff, 1984). They remain
curled up around them until hatching, while secret-
ing a repellent substance that probably also acts as a
biocide preventing the rotting of the eggs (Minelli,
2015). A couple of specimens collected on 14.v.2009
were observed during mating activities (Fig. 2B), and
one female deposited an egg mass on 16.vii.2009
(Figs. 2C–D).
Acknowledgements
I would like to thank Dr. Henrik Enghoff for the identica-
tion of the specimens. José D. Gilgado and Per Djursvoll provided
valuable comments that improved the manuscript.
with 6 genera and 20 described species worldwide
(Enghoff et al., 2015), is represented in Europe by
three species of the genus Polyzonium Brandt, 1837,
P. eburneum Verhoef, 1907, P. transsilvanicum Ve r -
hoeff, 1898 and P. germanicum Brandt, 1837. The
rst two mentioned species have a restricted distri-
bution in central and Eastern Europe, respectively,
while P. germanicum is widespread in most of the
continent (Kime & Enghoff, 2011). Two main Euro-
pean populations (Fig. 1), one in the eastern half of
Europe and one in the western half (including France
and SE Britain), have been described for P. germani-
cum (see Kime & Enghoff, 2011). An area of central
Europe, with only a few records of the species, seems
to separate these two populations (Kime, 2001), but
no morphological (Schubart, 1934) nor ecologi-
cal (Kime & Enghoff, 2011) differences have been
found between them. To date P. germanicum was not
known in the Iberian Peninsula.
This report describes a population that has been
found in the Gresolet forest (Fig. 1), located into the
Cadí-Moixeró Natural Parc (Saldes municipality,
Barcelona province, Spain). Ten specimens of P. ger-
manicum were collected by the author (with collec-
tion permit inside the natural park) in two dates (14.
xi.2008, 5 young specimens, and 14.v.2009, 2 
and 3 ) from litter layer and under logs and stones
Fig. 1.— Polyzonium germanicum distribution map. Data from
Enghoff & Kime (2009) and Kime & Enghoff (2011). Dark gray:
western population; light gray: eastern population; green: area
located between eastern and western populations with few
records for the species; white: no records for the species;
red dot: forest population of Gresolet (Spain) described in the
present report.
Fig. 1.— Mapa de distribución de Polyzonium germanicum.
Datos de Enghoff & Kime (2009) y Kime & Enghoff (2011). Gris
oscuro: población occidental; gris claro: población oriental;
verde: área ubicada entre las poblaciones oriental y occidental,
con pocos registros de la especie; blanco: sin registros para la
especie; punto rojo: población forestal de Gresolet (España)
descrita en el presente informe.
Notas / Notes 3
Graellsia, 77(1), junio 2021, e129 — ISSN-L: 0367-5041 – https://doi.org/10.3989/graellsia.2021.v77.279
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tic analysis. Journal of Zoological Systematics and
Evolutionary Research, 22 (1): 8–26. https://doi.
org/10.1111/j.1439-0469.1984.tb00559.x
Enghoff, H., Golovatch, S., Short, M., Stoev, P. & Wesener
T., 2015. Diplopoda – Taxonomic overview. In: Minelli
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ology. The Myriapoda, Vol. 2. Brill. Leiden: 363–447.
Enghoff H. & Kime R.D., 2009. Fauna Europaea: Diplop-
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www.faunaeur.org [accessed 05 May 2020].
Kime, R.D., 2001. The continental distribution of British
and Irish millipedes, part 2. Bulletin of the British Myri-
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specimen) pairing for sperm transfer, scale bar = 5 mm. C. Female and eggs, scale bar = 5 mm. D. Eggs, scale bar = 1 mm.
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C. Hembra y huevos, barra de escala = 5 mm. D. Huevos, barra de escala = 1 mm.
4 Notas / Notes
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... A pesar de su interés, los milpiés son un grupo poco estudiado en la península ibérica en comparación con otras regiones de Europa (Kime y Enghoff 2011, 2021. No obstante, en la última década ha habido un resurgir de la diplopodología en esta región (Akkari y Enghoff 2012Mauriès 2012Mauriès , 2013Mauriès , 2014aMauriès , 2014bMauriès , 2015Mauriès , 2018Enghoff y Reboleira 2013a, 2013bReboleira y Enghoff 2013, 2014a, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aGilgado et al. , 2020bSerra y Mauriès 2015, 2018Akkari et al. 2018;Djursvoll 2019;Recuero y Rodríguez-Flores 2020;Mateos 2021), y una buena parte de este estudio se ha centrado en especies del medio subterráneo (Enghoff y Reboleira 2013a, 2013bMauriès 2013Mauriès , 2014aReboleira y Enghoff 2013, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aPérez Fernández y Mauriès 2015;Akkari et al. 2018;Djursvoll 2019;Mauriès y Pérez Fernández 2019). ...
... A pesar de su interés, los milpiés son un grupo poco estudiado en la península ibérica en comparación con otras regiones de Europa (Kime y Enghoff 2011, 2021. No obstante, en la última década ha habido un resurgir de la diplopodología en esta región (Akkari y Enghoff 2012Mauriès 2012Mauriès , 2013Mauriès , 2014aMauriès , 2014bMauriès , 2015Mauriès , 2018Enghoff y Reboleira 2013a, 2013bReboleira y Enghoff 2013, 2014a, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aGilgado et al. , 2020bSerra y Mauriès 2015, 2018Akkari et al. 2018;Djursvoll 2019;Recuero y Rodríguez-Flores 2020;Mateos 2021), y una buena parte de este estudio se ha centrado en especies del medio subterráneo (Enghoff y Reboleira 2013a, 2013bMauriès 2013Mauriès , 2014aReboleira y Enghoff 2013, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aPérez Fernández y Mauriès 2015;Akkari et al. 2018;Djursvoll 2019;Mauriès y Pérez Fernández 2019). Pese a este reciente impulso, el conocimiento sobre una gran parte de las especies ibéricas, incluyendo las troglobias, se reduce a su morfología y su localidad típica, y apenas nada sobre su biología, ecología o estado de conservación. ...
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Los milpiés (Myriapoda: Diplopoda) del medio subterráneo en la península ibérica e islas baleares. Ecosistemas 33(2): 2458. https://doi.org/10.7818/ECOS.2458 Los milpiés (Myriapoda: Diplopoda) del medio subterráneo en la península ibérica e islas baleares Resumen: Los milpiés son un grupo de artrópodos muy diverso, pero relativamente poco estudiado, con casi 80 especies conocidas adaptadas al medio subterráneo en el ámbito ibero balear. En el presente trabajo se hace una revisión del conocimiento de los milpiés citados en el medio subterráneo en la península ibérica e islas baleares. Se hace una breve introducción, una mención al desarrollo histórico, y a las adaptaciones de los milpiés subterráneos. Se enumeran todas las especies troglobias encontradas en cuevas y el MSS de cada uno de los sistemas montañosos de la península, excluyendo la vertiente francesa de los pirineos. También se discuten las diferencias más relevantes en la diversidad de los diferentes sistemas, y se mencionan algunas características de interés de algunas de las especies más reseñables. Además, se proporciona un mapa con todas las cuadrículas UTM de 10x10 km en las que se han citado milpiés subterráneos. Palabras clave: artrópodos; cuevas; distribución; MSS; sistemas montañosos; troglobios Millipedes (Myriapoda: Diplopoda) of the subterranean environment in the Iberian Peninsula and Balearic Islands Abstract: Millipedes constitute a very diverse, and yet understudied, group of arthropods, with almost 80 subterranean species adapted to live in the subterranean environment in the Ibero-Balearic region. In the present work, a review of the knowledge on Ibero-Balearic subterranean millipedes is made. A brief introduction, history, and comments on the adaptations of subterranean millipedes are presented. All subterranean millipede species recorded in caves or MSS in each main mountain range of the Ibero-Balearic region, excluding the French Pyrenees, are enumerated. The main differences in subterranean millipede diversity among regions are discussed, and some remarks on relevant species are presented. Additionally, a map with all the 10x10 km UTM squares with subterranean millipede records is given.
... A pesar de su interés, los milpiés son un grupo poco estudiado en la península ibérica en comparación con otras regiones de Europa (Kime y Enghoff 2011, 2021. No obstante, en la última década ha habido un resurgir de la diplopodología en esta región (Akkari y Enghoff 2012Mauriès 2012Mauriès , 2013Mauriès , 2014aMauriès , 2014bMauriès , 2015Mauriès , 2018Enghoff y Reboleira 2013a, 2013bReboleira y Enghoff 2013, 2014a, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aGilgado et al. , 2020bSerra y Mauriès 2015, 2018Akkari et al. 2018;Djursvoll 2019;Recuero y Rodríguez-Flores 2020;Mateos 2021), y una buena parte de este estudio se ha centrado en especies del medio subterráneo (Enghoff y Reboleira 2013a, 2013bMauriès 2013Mauriès , 2014aReboleira y Enghoff 2013, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aPérez Fernández y Mauriès 2015;Akkari et al. 2018;Djursvoll 2019;Mauriès y Pérez Fernández 2019). ...
... A pesar de su interés, los milpiés son un grupo poco estudiado en la península ibérica en comparación con otras regiones de Europa (Kime y Enghoff 2011, 2021. No obstante, en la última década ha habido un resurgir de la diplopodología en esta región (Akkari y Enghoff 2012Mauriès 2012Mauriès , 2013Mauriès , 2014aMauriès , 2014bMauriès , 2015Mauriès , 2018Enghoff y Reboleira 2013a, 2013bReboleira y Enghoff 2013, 2014a, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aGilgado et al. , 2020bSerra y Mauriès 2015, 2018Akkari et al. 2018;Djursvoll 2019;Recuero y Rodríguez-Flores 2020;Mateos 2021), y una buena parte de este estudio se ha centrado en especies del medio subterráneo (Enghoff y Reboleira 2013a, 2013bMauriès 2013Mauriès , 2014aReboleira y Enghoff 2013, 2014b, 2018Gilgado et al. 2015aGilgado et al. , 2015bGilgado et al. , 2015cGilgado et al. , 2017Gilgado et al. , 2020aPérez Fernández y Mauriès 2015;Akkari et al. 2018;Djursvoll 2019;Mauriès y Pérez Fernández 2019). Pese a este reciente impulso, el conocimiento sobre una gran parte de las especies ibéricas, incluyendo las troglobias, se reduce a su morfología y su localidad típica, y apenas nada sobre su biología, ecología o estado de conservación. ...
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Los milpiés son un grupo de artrópodos muy diverso, pero relativamente poco estudiado, con casi 80 especies conocidas adaptadas al medio subterráneo en el ámbito ibero balear. En el presente trabajo se hace una revisión del conocimiento de los milpiés citados en el medio subterráneo en la península ibérica e islas baleares. Se hace una breve introducción, una mención al desarrollo histórico, y a las adaptaciones de los milpiés subterráneos. Se enumeran todas las especies troglobias encontradas en cuevas y el MSS de cada uno de los sistemas montañosos de la península, excluyendo la vertiente francesa de los pirineos. También se discuten las diferencias más relevantes en la diversidad de los diferentes sistemas, y se mencionan algunas características de interés de algunas de las especies más reseñables. Además, se proporciona un mapa con todas las cuadrículas UTM de 10x10 km en las que se han citado milpiés subterráneos.
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The modern era in diplopod taxonomy began in 1971 with publication of the Nomenclator Generum et Familiarum Diplopodorum. Along with the internet and electronic communications, this pioneering work catalyzed an increased output of synthetic publications beyond mere species descriptions, thereby leading to accelerating advancements. Significant taxonomic and/or cataloging studies have since occurred in 14 of the 16 established orders, all except Stemmiulida and Siphoniulida. Important faunistic works have appeared on Canada, Mexico, Asian Russia, China, Taiwan, Vietnam, Thailand, Iran, Turkey, central Asia in general, Belgium, Scandinavia, Great Britain & Ireland, Madagascar, and the Comoros, Canary, Cape Verde, and Galapagos Islands. The complete known ranges of four orders, four superfamilies, and 47 families have been mapped along with partial, regional mappings for seven other orders and seven families. With only one resident taxonomist, continental regions of the Neotropics warrant concentrated attention in future years, and emphasis on Chinese millipeds will surely continue because of the discovery of a substantial southeast Asian callipodidan fauna. Taxonomic emphases are needed in the Spirobolidea (Spirobolida), to assess affinities between the 10 component families, and Spirostreptida, to determine taxonomic positions of the suborders Cambalidea & Epinannolenidea. Resolution of the latter may derive from development of the "8 th gonopod clade" concept, involving the orders Chordeumatida, Callipodida, Stemmiulida, Polydesmida, & Siphoniulida, and the suborder Spirostreptidea (Spirostreptida). Attributes of the global diplopod community are discussed.
Article
Two hundred and seventy-nine Polyzonium germanicum individuals of various stadia were collected seasonally in winter, late spring-early summer (the breeding period) and early autumn. Fresh weight, dry weight and defatted dry weight were determined for each specimen. Seasonal changes in water content and fat content, expressed as percentages of defatted dry weight, were analysed in relation to body size and sex. Changes in water content were more pronounced in the smallest individuals (juveniles), whereas changes in fat content were more pronounced in the largest (adults). Seasonal fluctuations in water and fat content were similar in both sexes despite the sexual dimorphism of body weight. The results suggest the following relationships within the species life-cycle: (i) water content increases during the warm months (up to 295% for juveniles in early autumn), which coincides with the moulting period, (ii) Fat content increases subsequently to moulting; overall maxima are reached in early autumn (population mean = 62–5%), as well as in early summer for the young, (iii) Both water and fat content decrease (to about 150% and 50%, respectively) in winter; however, there is no evidence of starvation–at least in ovigerous females which increase in dry weight during that season, (iv) Fat content strongly decreases (below 30% in the largest specimens) in adults of both sexes during the breeding period.
Article
Application of the principles of cladistics, or phylogenetic systematics, to the class Diplopoda indicates monophyly (in the strict sense) of the taxa Penicillata, Pentazonia, Colobognatha (minus Siphoniulus?), Nemathophora, Merocheta, and Juliformia. The result agrees with traditional classifications but stands in contrast to the classification by Hoffman (R. L. Hoffman, 1979: Classification of the Diplopoda. 237 pp. Muséum d'Histoire naturelle, Genève).At higher levels, the analysis indicates strict monophyly of the taxa Diplopoda, Chilognatha, Helminthomorpha, and Eugnatha. The following sister-group relationships are proposed:Penicillata – ChilognathaPentazonia – HelminthomorphaColobognatha (minus Siphoniulus?) – EugnathaThe interrelationships of the eugnathan groups: Nematophora, Merocheta and Juliformia, remain unsettled, as does the position of Siphoniulus.A formal classification expressing the results is presented.ZusammenfassungPhylogenie der Tausendfüßer- eine kladistische AnalyseDie Anwendung von den Prinzipien der kladistischen – oder phylogenetischen – Systematik an der Klasse Diplopoda zeigt die Monophylie (sensu strictu) der Taxa Penicillata, Pentazonia, Colobognatha (ohne Siphoniulida?), Nematophora, Merocheta and Juliformia. Diese Ergebnisse stimmen mit traditionellen Klassifikationen überein, stehen aber in Widerspruch zur Klassifikation von R. L. Hoffman N (1979). Auf höheren taxonomischen Niveaus zeigt die Analyse Monophylie (sensu strictu) für die Taxa Diplopoda, Chilognatha, Helminthomorpha and Eugnatha. Die folgenden Schwester-Gruppen-Verhaltnisse werden vorgeschlagen:Penicillata – ChilognathaPentazonia – HehninthomorphaColobognatha (ohne Siphoniulida?) – Eugnatha.Die verwandschaftlichen Beziehungen der eugnathen Gruppen: Nematophora, Merocheta and Juliformia sind nosh unsicher, ebenso die Stellung von Siphoniulida.Die Ergebnisse werden in foreteller Klassifikation dargelegt.
Fauna Europaea: Diplopoda. Fauna Europaea version 2.0
  • H Enghoff
  • R D Kime
Enghoff H. & Kime R.D., 2009. Fauna Europaea: Diplopoda. Fauna Europaea version 2.0. Available from http:// www.faunaeur.org [accessed 05 May 2020].
Atlas of European Millipedes
  • R D Kime
  • H Enghoff
  • Glomerida
  • Platydesmida
  • Siphonocryptida
  • Polyzoniida
  • Polydsmida Callipodida
Kime, R.D. & Enghoff, H., 2011. Atlas of European Millipedes (Class Diplopoda). Vol.1 Orders Polyxenida, Glomerida, Platydesmida, Siphonocryptida, Polyzoniida, Callipodida, Polydsmida. Pensoft Publishers, Sofia-Moscow & European Invertebrate Survey, Leiden, 282 pp.
Recherches sur la biologie et l'ecologie de deux populations de Diplopodes en forêt d'Orléans: Polyzonium germanicum Brandt, 1831 et Polydesmus angustus Latzel, 1884. PhD
  • T Couret
Couret, T., 1985. Recherches sur la biologie et l'ecologie de deux populations de Diplopodes en forêt d'Orléans: Polyzonium germanicum Brandt, 1831 et Polydesmus angustus Latzel, 1884. PhD, University of Orleans.