INFLUENCE OF SMALL- AND LARGE-SCALE
ECOLOGICAL FACTORS ON THE CENTIPEDE
(CHILOPODA) ASSEMBLAGES OF ARMORICAN
FORESTS (NW FRANCE)

E. Iorio, Julien Pétillon

► To cite this version:

E. Iorio, Julien Pétillon. INFLUENCE OF SMALL- AND LARGE-SCALE ECOLOGICAL FACTORS ON THE CENTIPEDE (CHILOPODA) ASSEMBLAGES OF ARMORICAN FORESTS (NW FRANCE). Vie et Milieu / Life & Environment, 2020, 70 (1), pp.79-87. 10.57890/kcxxgc45. hal-03095721

HAL Id: hal-03095721

https://hal.science/hal-03095721v1

Submitted on 26 Jun 2025

**HAL** is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire **HAL**, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
# INFLUENCE OF SMALL- AND LARGE-SCALE ECOLOGICAL FACTORS ON THE CENTIPEDE (CHILOPODA) ASSEMBLAGES OF ARMORICAN FORESTS (NW FRANCE)

E. IORIO<fnref n="1" />*, J. PÉTILLON<fnref n="2" />

MYRIAPODA
LITTER
SOIL
QUADRAT SAMPLING
COMPETITION

ABSTRACT. – Despite their high functional role as abundant predators, the drivers of centipede assemblages remain poorly known in forests, especially in France. Using replicated depletion quadrats, we sampled five forests distributed over the Armorican Massif. A total of 5209 individuals belonging to 20 species was collected between 2015 and 2017. Assemblage composition was surprisingly constant across parcels, with only different dominant species in the oldest parcel (Bercé, Pays-de-Loire) and in the forest of Cerisy (Normandy). The latter hosted two exclusive species, likely because of both biogeographic and abiotic conditions, *Geophilus truncorum* and *Lithobius aeruginosus*. Species richness was overall close among sites and parcels, and therefore little influenced by tree age or dominant species. Individual densities varied between site pair-matched parcels, probably because of differences in temporal forest continuity rather than differences in tree ages. In quadrats, the presence of numerous ants strongly affected the number of centipedes, with global densities divided by an average factor of 7. Also at small spatial scale, we did not find a strong vertical stratification of centipede species in terms of presence-absence, with only 2 species exclusive of soil and none for litter, but with important differences in the relative abundance of dominant species. This study suggests that centipedes are important components of the forest soil and litter fauna, should be more frequently studied in French forests and have a potential of bio-indicators of local conditions in these habitats, as well as of temporal forest continuity.

## INTRODUCTION

Centipedes are predators, mainly feeding upon other invertebrates living in or on the soil, or under above-ground shelters (Demange 1981, Iorio 2014). There are more than 550 centipede species known from Europe (Zapparoli 2003, Bonato & Minelli 2014, Simaiakis & Strona 2015), including 149 currently reported from metropolitan France (Iorio 2014, Iorio & Geoffroy 2019). Behind Italy, France is the second country with the highest species diversity in Europe (Zapparoli & Minelli 2006, Simaiakis & Strona 2015). Since twenty years, and even more during the last 10 years, basic knowledge on centipedes is highly increasing in France because of numerous qualitative inventories in various regions and ecosystems (Iorio 2019). But on the other hand, these arthropods remain highly neglected in broad ecological studies in France, especially in forests. Some forest studies have sometimes taken considered centipedes together with various other invertebrate groups (e.g. Geoffroy et al. 1981, Blandin et al. 1985, Auclerc et al. 2012, De Smedt et al. 2019), but fully dedicated studies to forest centipedes or myriapods are very rare (Geoffroy 1979).

The ecological importance of centipedes in forest ecosystems is yet widely recognized elsewhere in Europe. They represent indeed one of the most abundant groups

of soil predators in terms of density of individuals in temperate forests (Albert 1979, Poser 1988, Scheu et al. 2003), with many species restricted to forest habitats (Voigtländer 2009, 2011). In several countries, authors have investigated variation in centipede assemblage and diversity in forests (e.g. Leśniewska 2000, Wytwer 2000, Lock et al. 2001, Kula & Lazorík 2015, Peretti & Bonato 2018). More precisely, the effects of reforestation (Baini & Zapparoli 2015), fragmentation (Fründ et al. 1997, Grinvald 2011, Distribution of Myriapods in Forest Mosaic. Master Sci Nature Protection, Dept Ecol Environ Sci Fac Sci Palacky), succession (Grgič & Kos 2003, 2005, Leśniewska et al. 2005, Lock et al. 2005) and the influence of various environmental factors (e.g. moisture, temperature, exposition, litter mass, openings caused by a windstorm) on the distribution of centipedes were studied over the last decades, showing the potential importance of this group in French forests (Poser 1990, Tuf 2002, Jabin 2008, Leśniewska & Skwierczyński 2018). Finally, some studies focused on the efficiency of sampling methods, showing that the pitfall trapping is not the most reliable for assessing centipede diversity and number of individuals (Gerlach et al. 2009, Tuf 2015). Despite this recent literature, centipedes are still considered less known than other groups of predators (e.g. spiders and carabid beetles: Bonato et al. 2018).

<footnote n="1">¹ EI - Entomologie & Myriapodologie, 522 chemin Saunier, F-13690 Graveson, France</footnote>

<footnote n="2">² UMR CNRS 6553 Ecobio, Université de Rennes, 263 Avenue du Gal Leclerc, CS 74205, 35042 Rennes cedex, France</footnote>

<footnote>* Corresponding author: cingulata@hotmail.fr</footnote>
To increase our knowledge of forest centipedes in France, we characterized both litter and soil assemblages of Chilopoda over five forests of the Armorican Massif, a wide homogenous geologic area in North-Western France. More precisely, we aimed at assessing how assemblages (in terms of numbers of species and individuals, species composition) changed at different spatial scales, i.e. between sites, habitats and micro-habitats. Thanks to our sampling design, we especially could test for the effects of (i) biogeographic conditions (forest scale), (ii) dominant tree species and age (parcel scale, although they were sometimes confounded) and (iii) the vertical distribution and the presence of ants (quadrat scale).

## MATERIAL AND METHODS

*Sampling sites and their characteristics:* Five forests were selected in the Armorican Massif area or its margins in western France: four in the Pays de la Loire region and one in the west of the Normandy region (Fig. 1). In each forest, two distinct and geographically close parcels, from 300 m to 5 km between almost all paired parcels (only those of Bercé are more distant: 16 km), were chosen (Table I). The following environmental variables were taken:

- The dominant tree species and the age of forest stands (data from "Office National des Forêts" and private owners, or from our personal observations: Table I). The forest of Cerisy was the only forest mainly dominated by beech (*Fagus sylvatica*).

Dominant tree species and climatic factors such as annual temperature were closely linked (Table I).

- The temporal forest continuity (data from the French National Geographic Institute: https://www.geoportail.gouv.fr; https://remonterletemps.ign.fr), which shows that one of the parcels (BE2) was clear-cut in 1972, when all the others were ancient.

- The mean annual rainfall (mm) and mean annual temperature (°C), with quotation of the mean summer temperature (i.e. the season with the highest differences between the forests), data from Météo-France (2012), Artélia (2013) and Cantat *et al.* (2015) (Table I).

*Field sampling:* In each parcel, we collected ten leaf-litter and soil samplings in a 0.25 m² metallic square from 10 cm of depth; each sampling being spaced with a minimum of three metres to the other, but without criterion of choice. Leaf-litter and soil were systematically separated on the field in each 0.25 m² square: collection of the leaf-litter horizons L and F in one bag; collection of below horizons (H and A) in a second bag; both being numbered with indelible felt. The specimens trying to escape were immediately captured and placed in the corresponding bag. All the contents of the bags were placed in Berlèse apparatus to extract centipedes. The litter and soil sampling has been used with success by various authors to study this group (e.g. Geoffroy 1979, Leśniewska *et al.* 2005, Auclerc 2012). Because quadrats with many (> 50) ants had few centipedes (1 in BE1, BE2 and VI1, and 2 in VI2), five quadrats were excluded from analyses, except when testing the effects of ants.

<figure><img src="image_2.png" /><figcaption>Fig. 1. – Map showing the studied forest parcels in western France (base: IGN FranceRaster).</figcaption></figure>
<table><caption>Table I. – Main characteristics of the five studied forests. * Oak = *Quercus* sp. (both *Q. petraea* and *Q. robur* are present; almost only *Q. petraea* in Grande Charnie and BE1).</caption><thead><tr><th>Name of the forest</th><th>Bercé</th><th>Grande Charnie</th><th>Mervent</th><th>Vibraye</th><th>Cerisy</th></tr></thead><tbody><tr><td>Parcel</td><td>BE1</td><td>BE2</td><td>GR1</td><td>GR2</td><td>ME1</td></tr><tr><td>Altitude</td><td>143</td><td>150</td><td>212</td><td>197</td><td>90</td></tr><tr><td>Dominant tree species*</td><td>Oak</td><td>Oak</td><td>Oak</td><td>Oak</td><td>Oak</td></tr><tr><td>Age of the forest stand</td><td>> 300 years</td><td>> 100 years</td><td>> 120 years</td><td>> 120 years</td><td>> 180 years</td></tr><tr><td>Mean annual rainfall</td><td>700-800</td><td>700-800</td><td>800-900</td><td>800-900</td><td>900-1000</td></tr><tr><td>Mean annual temperature (mean summer temperature)</td><td>11-12 (18-19)</td><td>11-12 (18-19)</td><td>11-12 (18-19)</td><td>11-12 (18-19)</td><td>12-13 (19-20)</td></tr></tbody></table><table><thead><tr><th>Mervent</th><th>Vibraye</th><th>Cerisy</th></tr></thead><tbody><tr><td>ME2</td><td>VI1</td><td>CE1</td></tr><tr><td>78</td><td>179</td><td>97</td></tr><tr><td>Oak</td><td>Oak</td><td>Oak</td></tr><tr><td>40 years</td><td>> 120 years</td><td>> 80 years</td></tr><tr><td>900-1000</td><td>700-800</td><td>800-900</td></tr><tr><td>900-1000</td><td>700-800</td><td>800-900</td></tr><tr><td>12-13</td><td>11-12</td><td>10-11</td></tr><tr><td>(19-20)</td><td>(18-19)</td><td>(15-17)</td></tr></tbody></table>

Most forests were sampled in spring (Mervent: 05/05/2015; Grande Charnie: 05/04/2016; Vibraye: 01/06/2016; Bercé: 06/04/2017), except for Cerisy that sampled in autumn (03/11/2017). Because of time and logistic constraints, sampling was carried over different years, but assemblages of forest centipedes have few inter-annual variations (Albert 1979, Geoffroy 1979). In temperate forests, centipedes have their highest activity in spring, but also are very active in autumn (Geoffroy 1979, Voigtländer 2009, 2011). In North-Western France and adjacent regions, all species included in the present paper are much found in both seasons (e.g. Iorio 2005a, b, 2007, 2014, Racine & Iorio 2017, Iorio & Racine 2018, Tillier 2018) as well as in United Kingdom for those which are also present in this country (Barber & Keay 1988). Dominant species are consequently similar between spring and autumn (Geoffroy 1979). We consequently argue that assemblages of Cerisy can still be compared with the others despite a difference in sampling periods.

*Data analysis:* Compositions were compared between parcels and forests using an analysis of similarity based on Pearson correlations and average links on the occurrence matrix.

To test whether numbers of individuals and species differed among spatially pair-matched parcels (for testing the effects of dominant tree species and parcel age) Wilcoxon tests were used.

Results were analyzed with PAST and Mintab softwares. We have taken into account the data of the centipede atlas of the Pays de la Loire region in progress (6707 data and 20976 centipedes identified in this region at the end of May 2018) (Iorio & Racine, 2018), in addition of other references, to support some parts of our discussion.

## RESULTS

A total of 5209 specimens belonging to 20 species was collected (Table SI), including 166 immature individuals of the genus *Lithobius*.

### Forest-scale analysis

Species richness ranged between 13 to 16 species between forests (Table SI). Assemblages were constant

<figure><img src="image_3.png" /><figcaption>Fig. 2. – Dendrogram based on correlation coefficients of centipede assemblages (complete link).</figcaption></figure>

between sites with high degrees of similarity in species composition, but also between parcels of a given forest (Fig. 2). Only the forest of Cerisy strongly differed from the other sites, and that was mostly explained by the high abundance of *Geophilus truncorum* and the low abundance of *Schendyla nemorensis*, while the oldest parcel (Bercé) was quite different from the other parcels and sites due to the high abundance of *Cryptops hortensis* (Fig. 3). *Geophilus truncorum* was an exclusive species from Cerisy, with a relative abundance of 48.8 % in this forest. On the other way, the density of *Schendyla nemorensis* was lowest in Cerisy (28 % of total numbers) when in all the other forests it dominated assemblages (from 56.5 % until 86.7 % in Bercé and Grande Charnie, respectively).

Because of this high similarity, we did not check for differences in species composition between pair-matched parcels, but only in numbers of species and individuals (both total and for dominant species).

### Parcel-scale analysis

The mean densities of centipedes ranged from 118.4 specimens/m² to 350.4 specimens/m² per parcel, with extreme densities from the same forest (forest of Mervent, parcels ME2 and ME1, respectively) (Fig. 4). The species richness was rather constant between parcels,
<figure><img src="image_4.png" /><figcaption>Fig. 4. – Mean density (± standard deviation) of centipedes (all species confounded) per forest parcel.</figcaption></figure>

<figure><img src="image_5.png" /></figure>

between 6 and 14 species, with some important variations (e.g. Bercé).

In Cerisy, between CE1 and CE2 parcels having different age and dominant tree species, the difference in centipede densities was not significant (Wilcoxon & Mann-Whitney test, Qobs = 35, P = 0.272), and the same goes for the densities of dominant species *Geophilus truncorum* (Qobs = 58, P = 0.570). *Lithobius aeruginosus* was an exclusive species to CE1, the younger parcel dominated by the alder (*Alnus glutinosa*). On the other way around, *Strigamia acuminata* was more abundant in CE2, the older parcel dominated by the beech (56.5 % of its total numbers).

Comparisons of pair-matched oak parcels with different ages (ME1 vs ME2; BE1 vs BE2) show a significant difference in centipede densities in one case (ME1 vs ME2) with more individuals in the older parcel, but not in the other (BE1 and BE2) (Wilcoxon & Mann-Whitney tests, Qobs = 100, P < 0.001 and Qobs = 40.5, P = 1, respectively).

### Quadrat-scale analysis

1915 specimens were collected in the litter vs 3294 individuals in the upper soil samples. Two species were exclusive from the soil samples: *Geophilus electricus* and *Stigmatogaster subterraneus*, while all litter species were found in the soil samples. However, *G. electricus* and *S. subterraneus* were present by few specimens (n = 11 and 8 respectively). Most of other species were better represented in the upper soil than in the litter, but for some species the opposite pattern was found, like e.g. *Strigamia acuminata* (80.4% of relative abundance in the litter), *Geophilus easoni* (71.4 %) or *Lithobius piceus* (64.1 %) (Fig. 5).

Centipede were on an average 7 times less abundant when ants are numerous (> 50 individuals per quadrat) compared to quadrats where they are not, or few, present (Table II).
<figure><img src="image_6.png" /><figcaption>Fig. 5. – Relative abundance (%) of centipede species in litter vs soil samples.</figcaption></figure>

<table><caption>Table II. – Proportion of mean centipede densities per parcel without or with few (< 10) ants vs with many (> 50) ants (± standard deviation, number of quadrats is given between brackets).</caption><thead><tr><th>Parcel</th><th>Square samplings<br>without ants</th><th>Square samplings<br>with numerous ants</th><th>Proportion</th></tr></thead><tbody><tr><td>BE1</td><td>42.3 ± 12.2 (9)</td><td>6 (1)</td><td>7.05</td></tr><tr><td>BE2</td><td>44.2 ± 13.5 (9)</td><td>6 (1)</td><td>7.37</td></tr><tr><td>VI1</td><td>71.8 ± 21.8 (9)</td><td>10 (1)</td><td>7.18</td></tr><tr><td>VI2</td><td>74.75 ± 19.3 (8)</td><td>9.75 ± 4.9 (2)</td><td>7.67</td></tr></tbody></table>

# DISCUSSION

## Biogeographic conditions

The forest of Cerisy strongly differs of those of Pays de la Loire because of the presence and dominance of *Geophilus truncorum*. This is a North-Western European species found in Normandy and elsewhere in forests of North-Western France, but that becomes very rare south-ern, as in Pays de la Loire region (Iorio 2014, Racine & Iorio 2017, Iorio & Racine 2018). In this region, it is almost only present in the most northern parts of the northern departments (Sarthe and Mayenne departments) (Iorio & Racine 2018). We also found in Cerisy *Lithobius aeruginosus*, a Central European species which reaches its western limit of distribution in very few localities of the Armorican region. It was found in the coldest forest of the five forests studied: mean annual and summer temper-atures differences between Pays de la Loire forests only vary of 1 °C at most (Mervent vs others), but the mean annual and summer temperatures of Cerisy are respec-tively lower of 1 °C and 2.5 °C than three of the Pays de la Loire forests, and of 2 °C and 3.5 °C of the fourth. The mean temperatures of Cerisy seem low enough to lead to a different species composition, getting closer to the assem-blages of more northern or continental European forests (Fründ et al. 1997, Lock et al. 2001, Iorio 2014).

In terms of species richness, the studied forests were slightly richer (by a difference from 1 to 4 species) than a Francilian forest like the forest of Commanderie (south of Fontainebleau, Ile-de-France: n = 12 species); this last being however considered as fairly poor because of its suburban location (Geoffroy 1979). The forest of Com-manderie has had a much higher pressure of sampling than ours, thus the difference is potentially higher in favour of our cases. Species richness is yet overall simi-lar to other Northern European forests, but the Lithobio-morpha are more numerous in the latter when continen-tal (e.g. Leśniewska 2000, Lock et al. 2001, Leśniewska et al. 2005, 2011, Jabin 2008, Voigtländer 2009). The Southern European or Alpine forests are obviously richer than Armorican forests, until two times (e.g. Grgič & Kos 2005, Kula & Lazorík 2015, Peretti & Bonato 2018).

The superdominant centipede species of Commanderie and of Pays de la Loire forests is the same (i.e. *Schendyla nemorensis*), but it is different from more continental for-ests. *Lithobius mutabilis* and *Strigamia acuminata* are indeed over-dominating the latter, yet sometimes co-dom-inating with *S. nemorensis* and *G. truncorum* (Leśniewska 2000, Jabin 2008, Voigtländer 2009, Grinvald 2011). Dif-ferences in species composition are getting larger when leaving French Atlantic for the continental area, with e.g. *Arctogeophilus inopinatus* and *Geophilus easoni* in the first area, and *Lithobius dentatus* and *Strigamia*
*transsilvanica* in the second (Spelda 1999, Wytwer 2000, Jabin 2008, Voigtländer 2009, Grinvald 2011, Iorio 2014, Racine & Iorio 2017, Iorio & Racine 2018). The centipede assemblages of Atlantic forests thus show some clear biogeographic particularities compared to other regions, as well as inside its area according to finer climatic conditions (northern location of Cerisy).

## *Dominant tree species and stand age*

Seemingly, tree species, all deciduous, only had a weak influence on centipede densities and assemblages in the Armorican Massif. The densities of both Cerisy parcels, respectively dominated by the alder (CE1) and the beech (CE2), are in the mid-range of those observed in oak parcels. Also, there was no significant difference between alder and beech parcels in terms of centipede numbers, as well as for the superdominant species *Geophilus truncorum*. Although *Lithobius aeruginosus* was exclusive to CE1, we cannot assume that its presence is linked to the alder because this species is also reported from forest parcels dominated by other deciduous trees in France and elsewhere (Spelda 1999, Lock 2000, Jabin 2008, Iorio 2014). In the parcel CE1, the dominance of the alder reflect of the more humid soil of this parcel. It must be underlined that the three other known Armorican localities of *L. aeruginosus* are from parcels dominated by other tree species (beech, oak) but also under particular abiotic conditions, e.g. fairly high altitude for this area (from 200 m a.s.l. to > 300 m a.s.l.) or in shaded and humid valleys with small streams and mesohydrophile to hygrophile flora species like *Paris quadrifolia* and *Chryosplenium* sp. (Racine & Iorio 2017, Iorio & Racine 2018, Franck Noël pers. com.). The assumption of Racine & Iorio (2017) and Iorio & Racine (2018) that *L. aeruginosus* only inhabits cold and humid forest habitats in the Armorican Massif tends to be confirmed here. Also, as already reported for several centipede species, both humidity and temperature are important factors shaping population dynamics (Tuf 2002, Jabin 2008, Voigtländer 2009, 2011). The presence of *L. aeruginosus* in Cerisy is thus likely due to both biogeographic and parcel abiotic conditions.

In this study only *Strigamia acuminata* seemed to favour a parcel dominated by a specific tree species, with differences between the beech-dominated parcel (CE2) and the alder-dominated parcel (CE1) in Cerisy. This tendency is broadly in accordance with the knowledge in other areas (Iorio 2014, Kula & Lazorík 2015).

The age of parcels was not acting significantly on centipede densities in both Bercé & Cerisy, but was highly significant in Mervent. Not only ME2 was younger than ME1 but, to the difference of other young parcels, was clear-cut in 1972 (https://remonterletemps.ign.fr). The parcel ME1 has never been clear-cut (as far back as we can go in time); nor the other studied forest parcels. The

lower density of centipede in ME2 could therefore be the result of interrupted forest continuity (by a clear-cutting of the parcel). Openings in forest habitats, even if resulting of a natural phenomenon like windstorm, are indeed known to have a strong impact on centipede assemblages (Leśniewska & Skwierczyński 2018). Lock et al. (2005) showed that, in forest extensions on former pastures or agricultural lands, it takes more than 23 years to have some typical woodland species, yet not reaching the population density of old forests for these species. Pontégnie et al. (2005) also stated that large clearings in forests have a negative effect on centipedes. Jabin (2008) also demonstrated that centipede density was much higher in primeval forests than in recently or historically managed forests of Central Europe (> 400 individuals/m² vs rarely more than 200 individuals/m²). Abundance of forest species such as *L. mutabilis* were also reduced according to the same author. Baini & Zapparoli (2015) finally showed that centipede assemblages were different between native forests and reforestations in Central Italy. Thus the factor which had an importance here was more probably the temporal forest continuity (i.e. interrupted or not by a clear-cutting) than age tree, at least for trees a few decades old.

Other important factors not studied here, like small-scale heterogeneity in forest stands, can also play an important role in driving local densities, species richness and to support populations of specialised species. It has indeed been shown that too homogeneous even-aged parcels contain poorer centipede assemblages than uneven-aged parcels (Grgič & Kos 2003, 2005). This could be another reason why the very regular and even-aged parcel of BE1 had much lower and more variable densities than the second oldest forest stand of ME1, also lacking some forest-specialized species like *Arctogeophilus inopinatus* and conversely the surprising overdominance of the eurytopic *Cryptops hortensis* in the first (Table SI).

## *Small-scale distribution patterns*

Our observations on the vertical distribution of centipedes are in accordance with other studies in European forests. The majority of the Geophilomorpha is indeed considered having a preference for the upper horizons of soil, thus euedaphic; but several species such as *Strigamia acuminata* and several *Lithobius* spp. prefer the litter layers (F and L-layers) (Geoffroy 1979, Poser 1990, Voigtländer 2009, 2011). It is interesting to note that *Geophilus easoni* had its highest relative abundance in the litter and not in the upper soil, contrary to other studied *Geophilus* spp. on this aspect.

Ants are known to feed upon centipedes as well as on many other ground- and litter-dwelling animals (Cerdá & Dejean 2011), but their interspecific relations with centipedes in Western Europe are poorly known. The slow-moving geophilomorphs have glandular ventral secre-
tions to defend against predators; however, even a large European species like *Henia vesuviana* cannot defend itself against swarms of ants (Hopkin & Anger 1992). The very reduced centipede densities in quadrats with numerous ants tends to confirm that the presence of ants (probably near their nest) is antagonistic with most centipedes.

As a conclusion, this first work on forest centipedes on the Armorican Massif, and more widely on Western France, show that, even if species richness is quite low, centipedes are important components of the soil and litter fauna of local deciduous forests, especially in terms of density. Thus this group should be more studied in French forests. Centipedes probably have the potential of being accurate bio-indicator at a large scale to study the effects of temporal forest continuity, and at a finer scale to study the effect of local abiotic like temperature and local humidity and biotic factors like interspecific competition factors.

ACKNOWLEDGMENTS. – We are very grateful to the Groupe d'Etude des Invertébrés Armoricains (GRETIA), F Herbrecht (GRETIA) and S Étienne (Office National des Forêts (ONF), Cerisy). We also thank Mrs P d'Harcourt and A Radenac (private forest of Vibraye), as well as Mr H Kemlin (private forest of Grande Charnie) for providing information on their forests; C Courtial for relevant advice. This study benefited financial support from the Région of Pays de la Loire, the ONF, the Sarthe department and the Direction Régionale de l'Environnement, de l'Aménagement et du Logement (DREAL). Finally, we thank the Editors of *Vie et Milieu* and the anonymous referees for their useful reading and suggestions to improve our manuscript.

REFERENCES

Albert AM 1979. Chilopoda as part of the predatory macroarthropod fauna in forests: abundance, lifecycle, biomass and metabolism. *In* Camatini M ed, Myriapod Biology. Academic Press, London: 215-231.

Artélia 2013. L'adaptation aux effets du changement climatique en Haute et Basse-Normandie. Analyse du climat du futur en Haute et Basse-Normandie. Artelia Eau & Environnement, Unité Climat Énergie.

Auclerc A 2012. Effets d'amendements calco-magnésiens sur la diversité des macro-invertébrés de sols forestiers et sur certains processus fonctionnels associés. Cas du massif vosgien (nord-est, France). Thèse de doctorat en vue de l'obtention du grade de Docteur de l'Université de Lorraine, mention Écotoxicité, Biodiversité, Écosystèmes.

Auclerc A, Nahmani J, Aran D, Baldy V, Callot H, Gers C, Iorio É, Lapied E, Lassauce A, Pasquet A, Spelda J, Rossi JP, Guerold F, 2012. Changes in soil macroinvertebrate communities following liming of acidified forested catchments in the Vosges Mountains (North-eastern France). *Ecol Eng* 42: 260-269.

Baini F, Zapparoli M 2015. Centipede assemblages in semi-natural and reforested sites: comparing data from a Mediterranean area (Myriapoda: Chilopoda). *Vie Milieu* 65: 157-168.

Barber AD, Keay AN 1988. Provisional atlas of the centipedes of the Bristish Isles. Biological Records Centre, Natural Environment Research Council.

Blandin P, Flogaitis E, Arbeille J, Geoffroy JJ 1985. Les variations interannuelles des macroarthropodes édaphiques dans une forêt tempérée. *Bull Ecol* 16: 273-283.

Bonato L, Minelli A 2014. Chilopoda Geophilomorpha of Europe: a revised list of species, with taxonomic and nomenclatorial notes. *Zootaxa* 3770 (1): 1-136.

Bonato L, Minelli A, Zapparoli M 2018. Centipede communities (Chilopoda) of forest soils across Europe: abundance, species richness and species composition. *Atti Accad Naz Ital Entomol* 65: 113-120.

Cantat O, Bühler MA, Héricher S, Caillebotte E, Jambu E, Barbot JF, Bosser K, Gresselin F 2015. Profil environnemental de Basse-Normandie : le climat. Report of the DREAL Basse-Normandie.

Cerdá X, Dejean A 2011. Predation by ants on arthropods and other animals. *In* Polidori C ed, Predation in the Hymenoptera: An Evolutionary Perspective. Transworld Research Network: 39-78.

Demange JM 1981. Les Mille-pattes Myriapodes. Généralités, Morphologie, Écologie, Éthologie. Détermination des Espèces de France. Éditions Boubée, Paris.

De Smedt P, Baeten L, Proesmans W, Van de Poel S, Van Keer J, Giffard B, Martin L, Vanhulle R, Brunet J, Cousins SAO, Decocq G, Deconchat M, Diekmann M, Gallet-Moron E, Le Roux V, Liira J, Valdés A, Wulf M, Andrieu E, Hermy M, Bonte D, Verheyen K 2019. Strength of forest edge effects on litter-dwelling macroarthropods across Europe is influenced by forest age and edge properties. *Divers Distrib* 25: 963-974.

Fründ HC, Balkenhol B, Ruszkowski B 1997. Chilopoda in forest habitat-islands in north-west Westphalia, Germany. *Ent Scand* 51: 107-114.

Gerlach A, Voigtländer K, Heidger CM 2009. Influences of the behaviour of epigeic arthropods (Diplopoda, Chilopoda, Carabidae) on the efficiency of pitfall trapping. *Soil Org* 81(3): 773-790.

Geoffroy JJ 1979. Les peuplements de Chilopodes et de Diplopodes d'une Chênaie-Charmaie (Station Biologique de Foljuif, Seine & Marne). Thèse de doctorat 3e Cycle. UPMC Paris VI.

Geoffroy JJ, Christophe T, Moltefas S, Blandin P 1981. Étude d'un écosystème forestier mixte. III. Traits généraux du peuplement de macroarthropodes édaphiques. *Rev Ecol Biol Sol* 18: 39-58.

Grgič T, Kos I 2003. Centipede diversity in patches of different development phases in an unevenly-aged beech forest stand in Slovenia. *Afr Invertebr* 44(1): 237-252.

Grgič T, Kos I 2005. Centipede diversity in differently structured forests in Slovenia. *Peckiana* 4: 49-56.

Hopkin SP, Anger HS 1992. On the structure and function of the glue-secreting glands of *Henia vesuviana* (Newport 1845) (Chilopoda: Geophilomorpha). *In* Meyer E, Thaler K & Schedl W eds, Proceedings of the 8th International Congress of Myriapodology, Berichte des Naturwissenschaftenlich Medizinischen Vereins in Innsbruck, Supplementum 10: 71-79.

Iorio É 2005a. A propos de quelques Chilopodes récoltés en Île-de-France, en Picardie et dans le Centre (Myriapoda, Chilopoda). *Bull Arthropoda* 23: 42-45.

Iorio É 2005b. Contribution à la connaissance des Chilopodes de Bretagne (Myriapoda, Chilopoda). *Bull Soc Linn Bordeaux* Tome 140 NS 33(2): 149-156.

Iorio É 2007. A new contribution to the knowledge of centipedes of eastern France (Chilopoda). *Schubartiana* 2: 1-12.
Iorio É 2014. Catalogue biogéographique et taxonomique des Chilopodes (Chilopoda) de France métropolitaine. *Mém Soc Linn Bordeaux* 15: 1-372.

Iorio É 2019. Les Chilopodes de France : état des connaissances. Présentation dans le cadre du 1er colloque des myriapodologistes français de Brunoy des 22 et 23 mars 2019.

Iorio É, Geoffroy JJ 2019. Étude des Chilopodes de Païolive (Ardèche, France) et description d'une nouvelle espèce du genre *Lithobius* Leach, 1814 (Myriapoda, Chilopoda). *Bull Soc Entomol Fr* 124(2): 109-126.

Iorio É, Racine A 2018. Projet d'atlas des Chilopodes (Chilopoda) des Pays de la Loire : bilan de la 4e année. GRETIA report.

Jabin M 2008. Influence of Environmental Factors on the Distribution Pattern of Centipedes (Chilopoda) and other Soil Arthropods in Temperate Deciduous Forests. Göttingen, Cuvillier Verlag.

Kula E, Lazorík M 2015. Comparison of Myriapoda in beech and spruce forests. *J For Sci* 61(7): 306-3014.

Leśniewska M 2000. Centipede (Chilopoda) communities of three beech forests in Poland. *Fragm Faun* 43: 343-349.

Leśniewska M, Skwierczyński F 2018. Impact of windstorm on a community of centipedes (Chilopoda) in a beech forest in Western Poland. *Biologia* 73(2): 165-173.

Leśniewska M, Koralewska-Batura E, Błoszyk J 2005. Centipede communities in oak-hornbeam forests of different ages and exploitation in Wielkopolska (Poland). *Peckiana* 4: 67-77.

Leśniewska M, Mock A, Kania G, Kania G 2011. Centipede (Chilopoda) Diversity in Forest Habitats of Ojcow National Park. *Pol J Environ Stud* 20(3): 581-590.

Lock K 2000. Preliminary Atlas of the centipedes of Belgium (Myriapoda, Chilopoda). I.N. and K.B.I.N., Report Institute of Nature Conservation 2000/19, Brussels.

Lock K, De Bakker D, De Vos B 2001. Centipede communities in the forest of Flanders. *Pedobiologia* 45: 27-35.

Lock K, Dekoninck W, Grootaert P 2005. Effect of forest developmental stage on centipede communities. *Bull Inst R. Sci Nat Belg Entomol* 75: 267-270.

Météo-France 2012. Atlas climatologique régional des Pays-de-la-Loire. DREAL Pays de la Loire.

Peretti E, Bonato L 2018. How many species of centipedes coexist in temperate forests? Estimating local species richness of Chilopoda in soil coenoses of the South-Eastern Alps. *Eur J Soil Biol* 89: 25-32.

Pontégnie M, Bus de Warnaffe G, Lebrun P 2005. Impacts of silvicultural practices on the structure of hemi-edaphic macrofauna community. *Pedobiologia* 49: 199-210.

Poser T 1988. Chilopoden als Prädatoren in einem Laubwald. *Pedobiologia* 31: 261-281.

Poser T 1990. The Influence of litter manipulation on the centipedes of a beech wood. In Minelli A ed, Proceedings of the Seventh International Congress on Myriapods. Brill, Leiden: 235-245.

Racine A, Iorio É 2017. Contribution à la connaissance des Chilopodes du Finistère et des Côtes d'Armor (Myriapoda, Chilopoda). *Invertébr Armoricains* 16: 3-28.

Scheu S, Albers D, Alphei J, Buryn R, Klages U, Migge S, Platter C, Salamon JA, 2003. The soil fauna community in pure and mixed stands of beech and spruce of different age: trophic structure and structuring forces. *Oikos* 101: 225-238.

Simaiakis S, Strona G 2015. Patterns and processes in the distribution of European centipedes (Chilopoda). *J Biogeogr* 42(6): 1018-1028.

Spelda J 1999. Verbreitungsmuster und Taxonomie der Chilopoda und Diplopoda Südwestdeutschlands. Diskriminanzanalytische Verfahren zur Trennung von Arten und Unterarten am Beispiel des Gattung *Rhymogona* Cook, 1896 (Diplopoda, Chordeumatida, Craspedosomatidae). Ph D Thesis, University of Ulm.

Tillier P 2018. Contribution à la connaissance des Chilopodes d'Île-de-France : première mention de *Lithobius (Lithobius) dentatus* C. L. Koch, 1844 et synthèse des connaissances régionales (Myriapoda Chilopoda). *Entomologiste* 74(2): 115-128.

Tuf IH 2002. Contribution to the knowledge of vertical distribution of soil macrofauna (Chilopoda, Oniscidea) in floodplain forest. In Tajovský K, Balik V, Pižl V eds, Studies on Soil Fauna in Central Europe. ISB AS CR, Ceské Budejovice: 241-246.

Tuf IH 2015. Different collecting methods reveal different ecological groups of centipedes (Chilopoda). *Zoologia* 32: 345-350.

Voigtländer K 2009. Ökologie. In Rosenberg J ed, Die Hundertfusser. Westarp Wissenschaften, Hohenwarsleben: 385-409.

Voigtländer K 2011. Chilopoda – Ecology. In Minelli A ed, Treatise on Zoology – Anatomy, Taxonomy, Biology. The Myriapoda, volume I. Brill: 309-325.

Wytwer J 2000. Centipede (Chilopoda) communities of some forest habitats of Puszcza Bialowieska in Poland. *Fragm Faun* 43: 333-342.

Zapparoli M 2003. The present knowledge on the European fauna of Lithobiomorpha (Chilopoda). *Bull Brit Myr Grp* 19: 20-41.

Zapparoli M, Minelli A 2006. Chilopoda. In Ruffo S, Stoch F eds, Checklist and Distribution of the Italian Fauna. *Mem Mus Civ Storia Nat Verona*, 2 Ser: 123-125 (with distribution maps on CD-Rom).

Received on February 14, 2020

Accepted on March 31, 2020

Associate editor: C Battisti
Appendix

<table><caption>Table SI. – Taxonomic list of centipede species, and total numbers per parcel.</caption><thead><tr><th></th><th>BE1</th><th>BE2</th><th>GR1</th><th>GR2</th><th>ME1</th><th>ME2</th><th>VI1</th><th>VI2</th><th>CE1</th><th>CE2</th><th>Total</th></tr></thead><tbody><tr><td>Geophilomorpha</td><td>136</td><td>352</td><td>361</td><td>562</td><td>829</td><td>260</td><td>542</td><td>542</td><td>384</td><td>347</td><td>4315</td></tr><tr><td>Arctogeophilus inopinatus</td><td>0</td><td>21</td><td>0</td><td>0</td><td>33</td><td>4</td><td>11</td><td>6</td><td>31</td><td>12</td><td>118</td></tr><tr><td>Geophilus easoni</td><td>0</td><td>7</td><td>0</td><td>1</td><td>7</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Geophilus electricus</td><td>0</td><td>0</td><td>7</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Geophilus gavoyi</td><td>2</td><td>0</td><td>0</td><td>0</td><td>7</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Geophilus osquidatum</td><td>0</td><td>0</td><td>2</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>8</td></tr><tr><td>Geophilus truncorum</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>226</td><td>186</td><td>412</td></tr><tr><td>Schendyla nemorensis</td><td>123</td><td>324</td><td>347</td><td>547</td><td>775</td><td>247</td><td>529</td><td>535</td><td>121</td><td>119</td><td>3667</td></tr><tr><td>Strigamia acuminata</td><td>9</td><td>0</td><td>1</td><td>3</td><td>1</td><td>0</td><td>1</td><td>1</td><td>4</td><td>26</td><td>46</td></tr><tr><td>Strigamia crassipes</td><td>2</td><td>0</td><td>3</td><td>1</td><td>1</td><td>0</td><td>1</td><td>0</td><td>2</td><td>3</td><td>13</td></tr><tr><td>Stigmatogaster subterraneus</td><td>0</td><td>0</td><td>1</td><td>1</td><td>5</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Lithobiomorpha</td><td>50</td><td>39</td><td>31</td><td>69</td><td>45</td><td>29</td><td>85</td><td>58</td><td>22</td><td>29</td><td>457</td></tr><tr><td>Lithobius aeruginosus</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td><td>0</td><td>6</td></tr><tr><td>Lithobius agilis</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>0</td><td>3</td><td>1</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lithobius calcaratus</td><td>3</td><td>8</td><td>4</td><td>1</td><td>0</td><td>3</td><td>6</td><td>14</td><td>2</td><td>1</td><td>42</td></tr><tr><td>Lithobius forficatus</td><td>1</td><td>0</td><td>0</td><td>0</td><td>2</td><td>4</td><td>2</td><td>2</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Lithobius macilentus</td><td>7</td><td>0</td><td>0</td><td>1</td><td>6</td><td>2</td><td>0</td><td>2</td><td>0</td><td>0</td><td>18</td></tr><tr><td>Lithobius microps</td><td>9</td><td>0</td><td>3</td><td>8</td><td>30</td><td>11</td><td>0</td><td>4</td><td>3</td><td>2</td><td>70</td></tr><tr><td>Lithobius muticus</td><td>0</td><td>0</td><td>5</td><td>10</td><td>0</td><td>1</td><td>0</td><td>1</td><td>9</td><td>0</td><td>26</td></tr><tr><td>Lithobius piceus</td><td>3</td><td>0</td><td>4</td><td>7</td><td>0</td><td>2</td><td>3</td><td>1</td><td>2</td><td>17</td><td>39</td></tr><tr><td>Lithobius sp.</td><td>13</td><td>11</td><td>15</td><td>33</td><td>4</td><td>3</td><td>52</td><td>29</td><td>0</td><td>6</td><td>166</td></tr><tr><td>Lithobius tricuspidis</td><td>14</td><td>20</td><td>0</td><td>9</td><td>2</td><td>3</td><td>19</td><td>4</td><td>0</td><td>3</td><td>74</td></tr><tr><td>Scolopendromorpha</td><td>201</td><td>13</td><td>4</td><td>7</td><td>2</td><td>7</td><td>91</td><td>37</td><td>1</td><td>74</td><td>437</td></tr><tr><td>Cryptops hortensis</td><td>201</td><td>13</td><td>4</td><td>7</td><td>2</td><td>7</td><td>91</td><td>37</td><td>1</td><td>74</td><td>437</td></tr><tr><td>Total number of individuals (parcel)</td><td>387</td><td>404</td><td>396</td><td>638</td><td>876</td><td>296</td><td>718</td><td>637</td><td>407</td><td>450</td><td>5209</td></tr><tr><td>Number of species (parcel)</td><td>11</td><td>6</td><td>11</td><td>14</td><td>13</td><td>13</td><td>10</td><td>12</td><td>11</td><td>12</td><td>20</td></tr><tr><td>Total number of individuals (forest)</td><td>791</td><td></td><td>1034</td><td></td><td>1172</td><td></td><td>1355</td><td></td><td>857</td><td></td><td>5209</td></tr><tr><td>Number of species (forest)</td><td>13</td><td></td><td>14</td><td></td><td>16</td><td></td><td>13</td><td></td><td>13</td><td></td><td>20</td></tr></tbody></table>