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Millipedes faced with drought: the life cycle of a Mediterranean population... 115
Millipedes faced with drought: the life cycle of a
Mediterranean population of Ommatoiulus sabulosus
(Linnaeus) (Diplopoda, Julida, Julidae)
Jean-François David1, Mathieu Coulis1
1 Centre d’Ecologie Fonctionnelle & Evolutive, UMR 5175, CNRS–Université de Montpellier, 1919 route de
Mende, F–34170 Montpellier cedex 5, France
Corresponding author: Jean-François David (jean-francois.david@cefe.cnrs.fr)
Academic editor: Ivan H. Tuf| Received 27 October 2014| Accepted 4 May 2015| Published 30 June 2015
http://zoobank.org/B7BBC82C-84CB-47C8-BBF4-AD8D8C3C31E9
Citation: David J-F, Coulis M (2015) Millipedes faced with drought: the life cycle of a Mediterranean population of
Ommatoiulus sabulosus (Linnaeus) (Diplopoda, Julida, Julidae). In: Tuf IH, Tajovský K (Eds) Proceedings of the 16th
International
Congress of Myriapodology, Olomouc, Czech Republic. ZooKeys 510: 115–124. doi: 10.3897/zookeys.510.8838
Abstract
Growth, development and life-cycle duration of the millipede Ommatoiulus sabulosus (f. aimatopodus)
were studied in a Mediterranean shrubland of southern France and compared with previous data from
northwest Europe. Changes in the proportions of stadia during the course of the year were analysed in
several generations. e results show that stadia VII and VIII are consistently reached after the rst year
of growth, and stadia IX and X after the second year. First reproduction may occur at the age of two years
in males reaching maturity at stadium X, but not until the age of three in those reaching maturity at sta-
dia XI and XII. Reproduction cannot occur until at least the age of three in females, which carry mature
eggs from stadium XI onwards. In comparison with more northern populations, life-cycle duration is not
shorter in the Mediterranean population but there are marked dierences in its phenology: the breeding
period is in autumn, so that juveniles of stadia II to VI are never faced with the summer drought, and
larger individuals are mostly inactive in summer; moreover, all individuals moult once every winter. e
results illustrate how julid millipedes of humid temperate regions could respond to higher temperatures
and drier summer conditions in the context of climate change.
Keywords
Millipedes, life cycle, phenology, climate change
ZooKeys 510: 115–124 (2015)
doi: 10.3897/zookeys.510.8838
http://zookeys.pensoft.net
Copyright Jean-François David, Mathieu Coulis. This is an open access article distributed under the terms of the Creative Commons Attribution License
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Jean-François David & Mathieu Coulis / ZooKeys 510: 115–124 (2015)
116
Introduction
In many organisms, ongoing climate change aects the timing of life-cycle events such
as activity, growth and reproduction (Parmesan 2006). When no long-term data sets
are available to analyse trends in local populations, potential phenological responses
to climate change can be studied by examining intraspecic variation in widespread
species that live in a wide range of conditions.
In millipedes (Diplopoda), geographic variation in life-cycle characteristics has
been documented for some species of European julids (Fairhurst 1974, David 1982),
but there is little precise information for populations living in the Mediterranean re-
gion. A number of widespread species such as Cylindroiulus caeruleocinctus (Wood),
Cylindroiulus punctatus (Leach), Leptoiulus belgicus (Latzel), Ommatoiulus rutilans (C.
L. Koch) and Ommatoiulus sabulosus (Linnaeus) have populations in this area, which
is typied by cool winters and hot, dry summers. e study of phenological character-
istics in Mediterranean populations of these species is particularly interesting, because
climate change scenarios predict warmer and drier summer conditions over large parts
of western Europe for the end of the 21st century (IPCC 2013). e purpose of the
present study is to describe the life cycle of a Mediterranean population of O. sabulosus
and to compare the results with those previously obtained further north in Europe,
so as to highlight dierences between populations from the two climatic zones. e
species has a wide distribution, from Finland and Scotland to Albania and Spain, in
contrast to most Ommatoiulus species that are conned to the Iberian peninsula and
north Africa (Akkari and Engho 2012). Two forms occur in southern France: that
in which adults have two orange-yellow bands on the back, and a typically Mediter-
ranean form, the so-called O. sabulosus aimatopodus (Risso), in which adults are black
dorsally. e latter form is the most common in southern France and often occurs at
high population densities in shrubland ecosystems on limestone (Coulis et al. 2013).
e post-embryonic growth and development of O. sabulosus were described in
detail by Halkka (1958) and Sahli (1969). e life cycle, i.e. the calendar of events be-
tween birth and reproduction, was studied under eld conditions by Halkka (1958) in
Finland, Sahli (1968) in Germany, Biernaux (1972) in Belgium and Fairhurst (1974)
in Great Britain. Reproduction occurs in late spring and summer in all these regions.
Biernaux (1972) tentatively suggested that both males and females mature in two years
in Belgium, but Fairhurst (1974) concluded that males take two or three years, and
females three or four years, to reach maturity in Great Britain. As regards Mediterra-
nean populations, Sahli (1991a, 1992) studied in detail the timing of male maturation
and adult–intercalary male successions (periodomorphosis) in O. sabulosus aimatopo-
dus from the Alpes-Maritimes and Provence, southern France. is author mentioned
that egg-laying occurs in late summer–early autumn (Sahli 1991a), but provided very
limited information on the growth, development and age at reproduction of females.
Sahli (1991b) suggested that females could breed only once before dying in Mediterra-
nean populations (semelparity), in contrast to Biernaux (1972) who concluded, based
on his own study of egg development in populations from Belgium, that females can
Millipedes faced with drought: the life cycle of a Mediterranean population... 117
breed in successive years (iteroparity). e presence of an abundant population of O.
sabulosus aimatopodus in a garrigue ecosystem of Provence provided the opportunity to
clarify some aspects of the species’ biology in the eld, with particular attention to how
this julid adjusts its phenology under warmer and drier conditions.
Methods
is study was conducted at the Massif de l’Etoile near Marseille, southern France
(5°25'E; 43°22'N), in a shrubland dominated by rockrose (Cistus albidus L.), kermes
oak (Quercus coccifera L.), rosemary (Rosmarinus ocinalis L.) and gorse (Ulex parviorus
Pourr.). e soil is shallow rendzina on limestone, in which rock fragments and stones
represent about 60% of the soil volume in the top 20 cm. e mean annual temperature
in the area is 15.1 °C, mean monthly temperatures ranging from 7.1 °C in January to
24.1°C in July, and the mean annual rainfall is 555 mm (Marseille 1981–2010 climate
normals). e driest months are June, July and August, during which the soil becomes
very dry. e millipede community, heavily dominated by O. sabulosus aimatopodus, also
comprises an abundant population of Polyxenus lagurus (Linnaeus) (Polyxenidae) and
rare specimens of Leptoiulus sp. (Julidae) and Trichoblaniulus sp. (Trichoblaniulidae).
Collections of millipedes were made using dierent methods. (1) Twenty three pit-
fall traps were set on the site in late March 2010 (8 days) and late April 2010 (10 days).
(2) Leaf litter and topsoil samples were taken within 25 × 25 cm quadrats in May 2010
(31 sampling units), November 2010 (12 s.u.), May 2012 (31 s.u.), October 2013 (15
s.u.), November 2013 (11 s.u.), March 2014 (12 s.u.), April 2014 (31 s.u.) and Septem-
ber 2014 (13 s.u.). Millipedes were extracted using Tullgren funnels. (3) Large individu-
als were also collected by hand in leaf litter to determine their reproductive status.
Individuals were assigned to a stadium by counting the rows of ocelli (R.O.) on
each side of the head (1 R.O. = stadium II, 2 R.O. = stadium III, etc.) (Engho et al.
1993). e method, however, was often dicult to apply from stadium XI onwards.
e numbers of podous rings (including the collum) and apodous rings (excluding
the telson) were counted. Intercalary males were distinguished from other males by a
much smaller rst pair of legs than in immature males, but not modied into hooks
as in copulatory males (Halkka 1958). Forty-two females of stadia X and higher were
dissected to determine whether mature eggs (i.e. brownish, subspherical eggs about 0.6
mm long) were present in the ovitube.
e growth of several cohorts in the eld was studied by examining changes in the
proportions of stadia in successive samples (Blower 1970). In addition, 40 individuals
of various stadia were reared in the laboratory for periods ranging from a few months
to two years. ey were kept in transparent plastic boxes containing sieved soil and
moist leaf litter. e boxes were placed in incubators tted with a glass door, in which
temperature followed the long-term monthly mean temperatures of Marseille, with a
daily thermoperiod of low amplitude. All millipedes received a pinch of powder yeast
every month and, occasionally, rabbit faeces as supplementary food.
Jean-François David & Mathieu Coulis / ZooKeys 510: 115–124 (2015)
118
Results
Post-embryonic growth and development in the eld
e stadia identied using the numbers of R.O., and the numbers of body rings count-
ed in each stadium, are indicated in Table 1. In terms of ring numbers, growth was
slightly dierent from that reported in more northern populations, with an extra apo-
dous ring in stadia III and IV. is was conrmed by the higher numbers of podous
rings from stadium IV onwards. e maximum number of stadia is uncertain because
it was generally impossible to decipher the exact number of R.O. in the largest indi-
viduals. However, a few males had at least 12 R.O. (stadium XIII) and a female with
57 podous rings had at least 13 R.O. (stadium XIV).
Sexual dimorphism was apparent at stadium VI. Although two males reared in
the laboratory reached maturity at stadium IX, the smallest adult males found in the
eld were in stadium X (Table 1). Immature males were numerous up to and includ-
ing stadium XI, indicating that many males mature for the rst time in stadia XI or
XII. Intercalary males were found from stadium XI onwards. Dissection of females in
late summer–early autumn, just before the breeding period (see below), showed that
ovigerous females carrying mature eggs were present in any stadium from stadium XI
onwards. None of the stadium X females that were dissected (n = 6) were ovigerous.
Phenology
Juveniles were active in leaf litter in late October as stadium II, in mid-November as sta-
dia II and III (Fig. 1b), and in mid-December as stadia III and IV. Also, a female kept for
Table 1. Growth and development of O. sabulosus in Provence. e number of rows of ocelli (R.O.), the
range of podous rings (collum included) and the numbers of apodous rings (telson excluded) are given for
each stadium. Male stages: Im. = Immature; Ad. = adult; Int. = Intercalary.
Stadium R.O. Podous rings / Apodous rings Male development
Juveniles Females Males
II 1 6 / 5
III 2 11 / 5,6
IV 3 16–17 / 6,7
V 4 22–24 / 6,7
VI 5 29–32 / 6,7,8 29–32 / 6,7 Im.
VII 6 35–38 / 5,6 36–39 / 5,6 Im.
VIII 7 42–45 / 3,4 41–45 / 2,3,4 Im.
IX 8 45–49 / 1,2,3 44–49 / 1,2,3 Im.
X 9 47–50 / 1,2 47–50 / 1,2 Im., Ad.
XI 10 48–53 / 1 48–50 / 1 Im., Ad., Int.
XII+ ≥ 11 49–57 / 0,1 50–55 / 0,1 Ad., Int.
Millipedes faced with drought: the life cycle of a Mediterranean population... 119
months in the laboratory produced stadium II juveniles in late October. Samples taken
in late March showed that the new generation was mainly in stadium V by the end of
winter (Figs 1c, 2a), which implies moulting during the winter. is result was conrmed
in laboratory rearings, in which juveniles that had hatched in October emerged from the
soil as stadium V in March. e rearings further showed that the new generation contin-
ued to grow rapidly in spring, from stadium V in March (rearing temperature: 10 ± 2°C)
to stadium VI in April (13 ± 2 °C) and to stadium VII in May (17 ± 2 °C), exactly as in
the eld (Fig. 2b). Sexual dierentiation at stadium VI thus occurs at the age of about
6 months. e pace of growth slowed markedly around the summer. Litter and topsoil
samples taken in early October showed that the smallest individuals, born in autumn of
the preceding year, were in stadia VII and VIII (Fig. 1a), indicating that only one moult
had occurred since May. All those one-year old millipedes were immature in both sexes.
During the second year of growth, no moult occurred from October to mid-No-
vember (Fig. 1b). At this time of year, the population becomes progressively inactive
in the soil, both in the eld and in the laboratory. One-year old millipedes moulted
once during the winter and emerged from the soil in late March as stadia VIII and
IX (Figs 1c, 2a). is was conrmed in laboratory rearings, in which six males and
females of stadium VIII collected in the eld in October burrowed into the soil in
October–November and emerged as stadium IX in March. Field samples taken in
2010 (Fig. 2) showed that the generation that was in stadia VIII and IX at the end of
winter remained in these stadia until May, moulted in mid-May (as shown by the large
Figure 1. Phenology of O. sabulosus in Provence (October 2013–April 2014). Stadia are indicated on the
horizontal axis and those of three identiable generations (G 2011 without any individuals, G 2012 and
G 2013) are grouped together. White bars = undierentiated juveniles and females; grey bars = immature
and intercalary males; black bars = adult males. Abundant (+) or very abundant (++) juveniles of stadia II
and III were not included in the calculation of percentages.
Jean-François David & Mathieu Coulis / ZooKeys 510: 115–124 (2015)
120
proportion of millipedes that were moulting at the time of sampling), and was still in
stadia IX and X in mid-November, at the age of two. e complete absence of stadia
IX and X in the autumn of 2013 (Fig. 1a, b) indirectly conrms that these stadia are
reached in two years, since, for unexplained reasons, there was no recruitment in the
autumn of 2011. Similarly, samples taken in the autumn of 2010 (Fig. 2c) suggest
there was no or little recruitment in that year, which was conrmed in subsequent
samples (data not shown).
e number of moults during the third year of growth cannot be deduced from
the eld data. Assuming that there are two further moults — one in winter and one
in spring, as in the second year of growth — most individuals of stadia XI and XII
found in the autumn of 2013 (Fig. 1a) would be three years old. However, this would
be inconsistent with the lack of juvenile recruitment in 2010, and it is likely that most
individuals in stadia XI and XII collected in the autumn of 2013 were actually born
before 2010.
Life-cycle duration
By combining the results on individual development and phenology, one may infer
that a small proportion of males that reach maturity at stadium X reproduce at the age
of two. However, many males that mature for the rst time at stadia XI and XII cannot
reproduce until the age of three. ere is no evidence that some females breed at the
age of two, since no ovigerous females were found in stadia IX and X in late summer.
Females need at least three years to reach stadium XI and lay eggs in early autumn.
Figure 2. Phenology of O. sabulosus in Provence (March 2010–November 2010). See explanations in
the legend to Fig. 1.
Millipedes faced with drought: the life cycle of a Mediterranean population... 121
e continuation of the life cycle was observed in a few adults reared in the labora-
tory. ree adult males collected in autumn moulted during the winter and emerged
from the soil in March as intercalary males. ey became mature again after a further
moult in spring and remained in the adult stage until the following autumn. Two large
females collected in autumn also moulted during the winter but, in contrast to males,
they did not moult again in spring or summer. One of these females bred in October,
overwintered a second time in the laboratory, and survived until the following Sep-
tember but without moulting. Post-mortem inspection showed that this female had no
apodous ring and contained no eggs.
Discussion
e present study provides the rst estimate of life-cycle duration for O. sabulosus in
southern France. e interpretation of our eld data was made easy by the generally
high abundance of juveniles and also by gaps between successive generations, possibly
due to reproduction failures and/or high juvenile mortality rates in some years. In the
population studied near Marseille, stadia VII and VIII are consistently reached after
the rst year of growth, and stadia IX and X after the second year. is pattern was ob-
served in three generations born between 2008 and 2012, despite some variation from
one year to another (e.g. the cohort born in 2008 was mainly in stadium VIII in late
March 2010, while that born in 2012 was mainly in stadium IX in late March 2014).
Our results dier from those of Sahli (1992), who assumed that stadia X and XI were
reached at the age of three in the region of Provence.
As adult males were found from stadium X onwards in our samples, some males
may reproduce at the age of two. However, males that reach maturity in stadia XI or
XII cannot reproduce until the age of three at the earliest. Also, ovigerous females,
which were found from stadium XI onwards in our samples, cannot breed until the
age of three at the earliest. Moreover, the presence of some stadium XI females without
any mature eggs in early autumn suggests they may start breeding at the age of four.
erefore, the duration of the life cycle, which corresponds to the age of females at rst
reproduction, is three or possibly four years in this Mediterranean population, i.e. the
same as in populations studied by Fairhurst (1974) in Great Britain.
It remains unclear whether each female breeds only once during its lifetime (semel-
parity) or can breed over several years (iteroparity). Sahli (1991a) assumed that O. sabu-
losus females might be semelparous in Mediterranean populations, reproduction being
spread over dierent stadia and dierent years in each generation. In the present study,
the single female that bred in the laboratory survived for a further year but died without
breeding again, so that there is still no direct evidence for iteroparity. Moreover, we did
not nd clusters of small oocytes at the same time as mature eggs in ovigerous females,
which Biernaux (1972) mentioned as evidence for iteroparity in O. sabulosus. On the
other hand, dissection of females in late summer–early autumn revealed that the pro-
portion of those not carrying mature eggs in stadia XI and higher was rather low (22%),
Jean-François David & Mathieu Coulis / ZooKeys 510: 115–124 (2015)
122
and the question is whether this is suciently high to be consistent with semelparity.
Semelparity would imply that many females in stadia XI, XII and even XIII postpone
reproduction until the next year(s), which should result in a substantial proportion of
females without eggs in early autumn. is topic requires further research.
Although Mediterranean conditions do not modify the length of the life cycle in
O. sabulosus, several phenological characteristics are very dierent between the popula-
tion of Marseille and more northern populations. First, there is a shift of the breeding
season. In northwestern Europe, the species generally breeds in summer (Sahli 1968,
Biernaux 1972, Fairhurst 1974). Under milder climate conditions, as on the island of
Jersey, the species tends to breed earlier (Fairhurst 1974). However, our study con-
rms that, in the Mediterranean region, the breeding period of O. sabulosus is delayed
until the autumn (Halkka 1958, Sahli 1991a). Juveniles of stadia II to IV were col-
lected only in this season. ey grow rapidly from autumn to the following spring
and the rst part of the life cycle is similar to that of Ommatoiulus moreleti (Lucas) in
southern Portugal, which breeds in late autumn–early winter (Baker 1984). In both
species, the earliest active stadia (stadia II to VI) are never faced with the hot and dry
conditions of the summer, which may be an adaptation of the Mediterranean popula-
tions of Ommatoiulus. It should be noted that, in other millipedes, the youngest stadia
are by far the least resistant to desiccation (David and Vannier 2001).
e seasonal patterns of activity and growth also dier between the two climatic
areas. In northern populations, there is generally a single period of activity and growth
from spring to autumn and the species is active in summer (Halkka 1958, Fairhurst
1974). e duration of the active season clearly increases with increasing temperatures
in areas where the risk of summer drought is low (cf. Halkka 1958, Fairhurst 1974,
Meyer 1985). In the Mediterranean population, however, activity stops during the
summer months, and our study has shown that there is at most one moult between
May and September. Similarly, in O. moreleti living at low altitudes in Madeira, Read
(1985) reported that growth slows down during the summer, presumably due to dry
conditions. e presence of two long periods of inactivity, in summer as well as winter,
largely explains why the life cycle of O. sabulosus is not shorter under Mediterranean
conditions, as would have been expected for millipedes living in a warmer climate
(David and Handa 2010).
Conclusion
e life cycle of O. sabulosus in the Mediterranean region appears to be inuenced
mainly by the summer drought. e dry season especially impacts phenology, i.e. the
timing of activity, growth and reproduction. Contrary to many organisms that breed
earlier in spring under warmer conditions, this julid breeds in autumn under Medi-
terranean conditions, so that juveniles are unlikely to be exposed to severe drought.
Moreover, larger stadia become inactive in summer and the total duration of activity
over a year is roughly the same as in northern populations. As a result, the life cycle is
Millipedes faced with drought: the life cycle of a Mediterranean population... 123
not shorter in the Mediterranean region than in Great Britain. Although it is too soon
to generalize, the life cycle of O. sabulosus in southern France is quite similar to that of
O. moreleti in southern Portugal, suggesting ways in which a number of julids could
respond to drier summer conditions in the context of climate change.
Acknowledgements
We thank Anais Rancon, Anne Gorgeon, Mathieu Santonja, and all the members of
the Bioux team who assisted in eld sampling. is study was conducted as part of
the CLIMED project funded by the French National Agency for Research (ANR-09-
CEP-007).
References
Akkari N, Engho H (2012) Review of the genus Ommatoiulus in Andalusia, Spain (Diplopoda:
Julida) with description of ten new species and notes on a remarkable gonopod structure,
the fovea. Zootaxa 3538: 1–53.
Baker GH (1984) Distribution, morphology and life history of the millipede Ommatoiulus
moreletii (Diplopoda: Iulidae) in Portugal and comparisons with Australian populations.
Australian Journal of Zoology 32: 811–822. doi: 10.1071/ZO9840811
Biernaux J (1972) Chorologie et étude biologique comparée de deux familles de Myriapodes
Diplopodes belges: les Blaniulidae et les Iulidae. PhD esis, Faculté des Sciences Agrono-
miques de l’Etat, Gembloux, Belgium.
Blower JG (1970) e millipedes of a Cheshire wood. Journal of Zoology (London) 160:
455–496. doi: 10.1111/j.1469-7998.1970.tb03092.x
Coulis M, Hättenschwiler S, Fromin N, David JF (2013) Macroarthropod–microorganism
interactions during the decomposition of Mediterranean shrub litter at dierent moisture
levels. Soil Biology & Biochemistry 64: 114–121. doi: 10.1016/j.soilbio.2013.04.012
David JF (1982) Variabilité dans l’espace et dans le temps des cycles de vie de deux populations
de Cylindroiulus nitidus (Verhoe) (Iulida). Revue d’Ecologie et de Biologie du Sol 19:
411–425.
David JF, Handa IT (2010) e ecology of saprophagous macroarthropods (millipedes, wood-
lice) in the context of global change. Biological Reviews 85: 881–895. doi: 10.1111/j.1469-
185X.2010.00138.x
David JF, Vannier G (2001) Changes in desiccation resistance during development in the mil-
lipede Polydesmus angustus. Physiological Entomology 26: 135–141. doi: 10.1046/j.1365-
3032.2001.00226.x
Engho H, Dohle W, Blower JG (1993) Anamorphosis in millipedes (Diplopoda) — the pre-
sent state of knowledge with some developmental and phylogenetic considerations. Zoo-
logical Journal of the Linnean Society 109: 103–234. doi: 10.1111/j.1096-3642.1993.
tb00305.x
Jean-François David & Mathieu Coulis / ZooKeys 510: 115–124 (2015)
124
Fairhurst C (1974) e adaptive signicance of variations in the life cycles of Schizophylline
millipedes. Symposia of the Zoological Society of London 32: 575–587.
Halkka R (1958) Life-history of Schizophyllum sabulosum (L.) (Diplopoda, Iulidae). Annales
Zoologici Societatis Zoologicae-Botanicae Fennicae Vanamo 19: 1–71.
IPCC (2013) Climate Change 2013: e Physical Science Basis. Cambridge University Press,
Cambridge, 1535 pp.
Meyer E (1985) Distribution, activity, life-history and standing crop of Julidae (Diplopoda,
Myriapoda) in the Central High Alps (Tyrol, Austria). Holarctic Ecology 8: 141–150. doi:
10.1111/j.1600-0587.1985.tb01164.x
Parmesan C (2006) Ecological and evolutionary responses to recent climate change. Annual
Review of Ecology Evolution and Systematics 37: 637–669. doi: 10.1146/annurev.ecol-
sys.37.091305.110100
Read HJ (1985) Stadial distributions of Ommatoiulus moreleti at dierent altitudes in Madeira
with reference to life history phenomena (Diplopoda, Julidae). Bijdragen tot de Dierkunde
55: 177–180.
Sahli F (1968) Observations sur la biologie et la périodomorphose chez le Diplopode Schizo-
phyllum sabulosum (L.) en Allemagne. Bulletin Scientique de Bourgogne 25: 333–346.
Sahli F (1969) Contribution à l’étude du développement post-embryonnaire des Diplopodes
Iulides. Annales Universitatis Saraviensis, Mathematisch-Naturwissenschaftliche Fakultät
7: 1–154.
Sahli F (1991a) Recherches sur le cycle des mâles d’Ommatoiulus sabulosus (L.) (Myriapoda,
Diplopoda, Julidae) dans les Alpes-Maritimes et en Provence. Bulletin Scientique de
Bourgogne 44: 33–39.
Sahli F (1991b) Fréquence de la périodomorphose chez le Diplopode Julide Ommatoiulus sa-
bulosus (L.) dans les Alpes-Maritimes et en Provence II. Passages males intercalaires-males
adultes. Bulletin Scientique de Bourgogne 44: 41–47.
Sahli F (1992) On male reproduction strategies in Ommatoiulus sabulosus (L.) in the Maritime
Alps and Provence (France): juvenile to adult maturation moults. Berichte des Naturwis-
senschaftlich-Medizinischen Vereins in Innsbruck Supplementum 10: 167–176.
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